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REVISION OF POCADIUS ERICHSON (COLEOPTERA: NITIDULIDAE).
A Dissertation
Submitted to the Graduate Faculty of the
Louisiana State University and
Agricultural and Mechanical College
in partial fulfillment of the
requirements for the degree of
Doctor of Philosophy
in
The Department of Entomology
by
Andrew R. Cline
B.S. University of Alabama - Huntsville, 1996
M.S. University of Missouri, 2000
December 2005
© Copyright 2005
Andrew R. Cline
All rights reserved
ii
DEDICATION
This dissertation is dedicated to my wife JoAnna. Through the many years we
have spent together you have often been my balance and point of reason. Your patience
and understanding were always a comfort and inspiration. The work produced herein
would not have been possible without you.
iii
ACKNOWLEDGEMENTS
I thank my graduate advisor Dr. Christopher Carlton for his advice throughout the
completion of this research. I thank former and current entomology department head’s
Drs. Frank Guillot and Timothy Schowalter respectively for their support of my graduate
studies. Drs. Dorothy Prowell, Meredith Blackwell, Mark Hafner and Daniel Burba
graciously served as graduate committee members. Alexey Tishechkin, a lab associate
during my tenure at LSU, provided an immeasurable amount of inspiration and
encouragement through our daily discussions and across-office chats, and not only
broadened my scientific horizons but also developed my global point of view on cultural,
socioeconomic, and historical matters. Dr. Shaun Winterton aided in the analysis of data.
I graciously thank the numerous curators and directors that generously loaned
specimens used in this study, and without whom this research would have been
impossible. In particular, I thank Al Newton and Margaret Thayer at the Field Museum;
Robert Anderson, Francois Genier, and Henry and Anne Howden at the Canadian
Museum of Nature; Alexander Kirejtshuk at the Zoological Institute in St. Petersburg,
Russia; Max Barclay and Malcolm Kerley at the National Museum of Natural History -
London; Phillip Perkins at the Museum of Comparative Zoology at Harvard University;
and Mike Thomas and Paul Skelley at the Florida State Collection of Arthropods for their
kindness during visits made by the author to their respective institutions. Paul Skelley
aided in the production of the SEM photographs. Richard Leschen of the New Zealand
Landcare; Jose Luis Navarette-Heredia of the University of Guadalajara; Steve Ashe of
the Snow Entomological Collection at the University of Kansas; Roberta Brett and Dave
Kavanaugh at the California Academy of Sciences; Nancy Adams, and Gary Hevel and
David Furth of the United States National Museum - Smithsonian Institute provided
iv
numerous specimens for this work. Robert Turnbow, Roy Morris, Jim Wappes, Bill
Warner, and Alan Gillogly all graciously donated material.
Funding for this research was provided by the Louisiana State University
Agricultural Center, an Ernst Mayr grant from the Museum of Comparative Zoology at
Harvard University, visiting systematist grants from the Field Museum and the Canadian
Museum of Nature, travel grants from the Florida Entomological Society and Louisiana
State University Graduate School, an A.W. Mellon grant from the Organization for
Tropical Studies, a Louisiana State University Sigma Xi Grant-in-Aid of Research, and a
National Science Foundation Doctoral Dissertation Improvement Grant DEB 0308764.
v
TABLE OF CONTENTS
ACKNOWLEDGEMENTS………………………………………………… iv
ABSTRACT……………...…………………………………………………. vii
CHAPTER
1. INTRODUCTION………………………………………….. 1
General Background………………………………………… 1
Taxonomic Background. I. Cucujoidea Latrielle 1802……... 2
Taxonomic Background. II. Nitidulid-Lineage ……………. 7
Taxonomic Background. III. Nitidulidae Latrielle 1802…… 8
Systematics and Classification of Nitidulidae……………… 13
Biological / Ecological Considerations…………………….. 19
Morphological / Molecular Considerations………………… 25
Pocadiini / Pocadius Background………………………….. 27
2. MATERIALS AND METHODS…………………………… 34
Taxonomic Materials / Methods……………………………. 34
Systematic Methods. I. Ingroup / Outgroup Discussion..….. 39
Systematic Methods. II. Character Discussion……………... 44
Systematic Methods. III. Phylogenetic Reconstruction……... 55
3. SYSTEMATIC TREATMENT...………………………….. 58
Generic Redescription……………………………………… 58
Key to Species……………………………………………… 62
Species Accounts............…………………………………… 72
Pocadius Phylogeny….……………………………………... 299
Checklist of Pocadius Erichson.............................................. 329
4. DISCUSSION……………………………………………….. 332
Monophyly of Pocadius and Its Placement in Nitidulinae…. 332
Origin and Distribution of Pocadius..…………….…………. 335
Host Fungal Evolution……………………………………… 340
Phenological Considerations……………………………….. 342
Future/Continuing Research...……………………………… 343
REFERENCES CITED……………………………………………………... 346
APPENDIX: DATA MATRIX…………………………………………..….. 369
VITA………………………………………………………………………… 375
vi
ABSTRACT
A revision of Pocadius Erichson (Coleoptera: Nitidulidae) was completed. A
total of 46 species were (re)described, including 25 new species, a key to species
constructed, and phylogenetic analysis performed. Taxonomic changes, including
nomenclatural emendations, reinstatement of species, and description of new taxa, within
this dissertation do not constitute formal changes as defined by the International Code of
Zoological Nomenclature. Illustrations of the genitalia, key characters, and dorsal
habitus of species are provided. The phylogeny demonstrates a monophyletic Pocadius.
The phylogeny suggests a Palearctic origin of the genus with subsequent speciation into
the Old World tropics and New World. Host specialization on gasteromycetes fungi by
Pocadius species was shown not to be obligate but rather facultative. Some sympatric
species were shown to have temporally disjunct occurrences that may provide
reproductive isolation.
vii
CHAPTER 1. INTRODUCTION
“In all things of nature there is something of the marvelous.”
Aristotle, On the Parts of Animals, circa 350B.C.
GENERAL BACKGROUND
Nitidulidae, or sap beetles, are small to minute Clavicornia taxa (e.g. beetles having
clubbed antennae) with a variety of body forms, ecologies, life history strategies, and
evolutionary peculiarities. Characterization of the family is difficult, and the literature is replete
with misplacements of members of other, sometimes unrelated families, into Nitidulidae. Until
the 20th century, Nitidulidae contained several other families or parts of families (e.g.
Cyclaxyridae, Smicripidae, Kateretidae, Protocucujidae, and Phloeostichidae). These small
convex beetles with drab coloration have had historically unclear systematic affinities thereby
making this family a quintessential “dumping ground” for small beetles with clubbed antenna.
Characterization of what a nitidulid is remains questionable as remarked upon by Murray
(1864) prior to his monograph, “this I knew to be no easy task, no journey of the Sabbath day.”
Through comparative morphology, Lawrence et al. (1999a, 1999b) and Habeck (2002a, based on
Parsons 1943 and Audisio 1993) provided the following that differentially define Nitidulidae:
antennae 11 segmented with at least the three terminal antennomeres forming a well-defined
club, labrum free and visible, maxilla with a single lobe, procoxae transverse, tarsal formula 5-5-
5, larvae with a complex mandibular prostheca and pretarsal setae present. Of these, I find two
most useful when identifying material, i.e. transverse procoxae and compact 3-segmented club.
However, only the former character as well as a single-lobed maxilla and complex larval
mandibular prostheca are synapomorphic, as evidenced by their absence in other beetle lineages.
1
To understand and appreciate the complexity of the taxonomic and subsequent systematic
condition of Nitidulidae fully, it is best first to assess the superfamily in which it is contained and
continue down the classification hierarchy (Table 1). The following sections deal with our
current understanding of nitidulids and their closest allies. Taxonomic efforts focusing on
nitidulids and their relatives, how nitidulids are subdivided, the lifestyles and evolutionary
histories some nitidulids exhibit, and studies underlying comparative morphology and molecular
biology in the family also are treated. Finally, an introduction to the focal taxa of this
dissertation, i.e. the genus Pocadius Erichson and tribe Pocadiini Seidlitz, are given.
TABLE 1. Hierarchy demonstrating the placement of Pocadius, Pocadiini, and other taxa. Taxa
in bold indicate some of the taxa considered in this study.
KINGDOM Animalia
PHYLUM Arthropoda
CLASS Insecta
ORDER Coleoptera
SUBORDER Archostemata, Myxophaga, Adephaga, Polyphaga
SERIES Staphyliniformia, Scarabaeiformia, Elateriformia,
Bostrichiformia, Cucujiformia
SUPERFAMILY Lymexyloidea, Cleroidea, Cucujoidea, Tenebrionoidea,
Chrysomeloidea
FAMILY
Sphindidae (cryptic slime mold beetles), Kateretidae
(short winged flower beetles), Nitidulidae (sap beetles),
Smicripidae (palmetto beetles), Coccinellidae (lady-bird
beetles)
SUBFAMILY
Carpophilinae, Amphicrossinae, Cillaeinae,
Maynipelinae, Calonecrinae, Meligethinae, Epuraeinae,
Nitidulinae, Cryptarchinae, Cybocephalinae
TRIBE Nitidulini, Cychramini, Cyllodini, Cychramptodini,
Lawrencerosini, Amborotubini, Pocadiini
GENUS
Atarphia, Hebasculinus, Hebascus, Hyleopocadius,
Lordyrodes, Niliodes, Physoronia, Pocadioides,
Pocadius, Pocadites, Pseudoplatychora, Taraphia,
Teichostethus
SPECIES Pocadius ferrugineus (Fabricius)
TAXONOMIC BACKGROUND. I. CUCUJOIDEA LATRIELLE 1802
Cucujoidea is one of six superfamilies (Lymexyloidea, Cleroidea, Cucujoidea,
Tenebrionoidea, Chrysomeloidea, and Curculionoidea) in the series Cucujiformia Lameere
(1938). The Cucujiformia is the most speciose series (~50% of all beetle species) containing
2
hyperdiverse families such as Cerambycidae (long-horned beetles), Chrysomelidae (leaf beetles),
and Curculionidae (weevils). Interestingly, the greatest species diversity occurs in the mainly
phytophagous lineages, whereas the greatest familial diversity occurs in the mycophagous
lineages. Characters delineating the cucujoid lineage include: cryptonephridic Malpighian
tubules, ring-type or sheath-like aedeagus, hylecoetoid metendosternite, undivided mala in the
larval maxilla, retention of larval spiracular closing apparatus in adults, and absence of spiracles
on abdominal segment 8 in adults (see Crowson 1960 and Lawrence and Newton 1982).
According to Pakaluk et al. (1995), the Cucujoidea contained 31 families, 78 subfamilies, ~1500
valid genera, and >20,000 species. However, numerous changes have occurred since this
catalogue appeared, most importantly the transfer of Languriidae into a broadly defined
Erotylidae (Leschen 2003) and erection of Cyclaxyridae. According to my tabulation (Table 2)
Cucujoidea includes 31 families, 89 subfamilies, a little more than 1200 genera, and >20,000
species. Thus, some generic resolution has occurred over the last decade in some lineages and
new subfamilies erected, but relatively few new species have been described (compare Pakaluk
et al. 1995 to Table 2). This trend shows the relative push in insect systematics to produce
higher level work while for the most part neglecting species level monographs.
The first formalization of the Cucujoidea was offered by Böving and Craighead (1931) and was
based primarily on larval forms. This approach of using immature forms to recognize
relationships is an important innovation, particularly with cucujoid beetles, as there is a long and
erroneous history of classifications based on convergent adult morphological characters.
Unfortunately, this seminal work was overlooked by many beetle systematists, and the
superfamily did not begin to receive serious attention again until Crowson, the 20th century’s
most imminent coleopterist, began studies on members of the lineage in the 1950’s (e.g. his
“Classification of the Families of British Coleoptera” series, culminating with his 1955 text and
3
1960 phylogenetic assessment of the order). Böving and Craighead’s concept of the Cucujoidea
included the Tenebrionoidea (= Heteromera, so called for the 5-5-4 tarsal formula of adults),
which was not formally divided from Cucujoidea “sensu lato” until Crowson (1954, 1960).
Thus, by 1960 we began to see a well-developed concept of the Cucujoidea.
TABLE 2. Current classification summary of the Cucujoidea. Superscript numbers refer to
published records, “?” values delineate undetermined affinities of the families.
Family Number of
Subfamilies
Number of
Genera
Number of
Species
Defined
Lineages
Undefined
Lineages
Sphindidae14 9 61 Sphindid
Protocucujidae21 1 7 Sphindid
Nitidulidae310 >200 >4000 Nitidulid
Kateretidae41 12 100 Nitidulid
Smicripidae51 1 6 Nitidulid
Cyclaxyridae* 1 1 1 ?
Monotomidae62 20 250 ?
Boganiidae72 5 11 ?
Phloeostichidae84 6 10 ?
Helotidae91 5 108 ?
Cucujidae10 1 4 40 Cucujid
Silvanidae11 2 47 470 Cucujid
Passandridae12 1 9 105 Cucujid
Laemophloeidae13 1 37 400 Cucujid
Propalticidae14 1 2 35 Cucujid
Phalacridae15 2 55 600 ?
Hobartiidae16 1 2 6 ?
Cavognathidae17 1 4 5 ?
Cryptophagidae18 3 48 600 Cryptophagid
Lamingtoniidae19 1 1 1 Cryptophagid
Erotylidae20 6 110 >3500 Cryptophagid
Byturidae21 2 7 16 ?
Biphyllidae22 1 6 200 ?
Cerylonidae23 5 52 300 Cerylonid
Bothrideridae24 4 35 300 Cerylonid
Alexiidae25 1 1 50 Cerylonid
Discolomatidae26 5 18 400 Cerylonid
Endomychidae27 12 120 1300 Cerylonid
Coccinellidae28 6 360 >6000 Cerylonid
Corylophidae29 4 35 284 Cerylonid
Latridiidae30 2 25 1050 Cerylonid
∑ 89 1238 >20216 2 3 + ?
1 McHugh 2002; 2 Slipinski 1998; 3 Kirejtshuk 1986a, 1998a, 1998b; 4 Audisio 1993, Habeck 2002; 5 Price 2002; 6
Lawrence 1982, Bousquet 2002; 7 Crowson 1990, Endrödy-Younga 1991, Klimasweski and Watt 1997, 8 Lawrence
1982 and Lawrence 1995; 9 Kirejtshuk 2000; 10 Thomas 2002a; 11 Thomas 2002b; 12 Slipinski 1987, 1989, Burkhardt
and Slipinski 1991, 1995; 13 Thomas 2002c; 14 Lawrence 1982; 15 Lawrence 1991, Steiner 2002; 16 Tomaszewska
and Slipinski 1995; 17 Lawrence 1982, 1991; 18 Leschen 1996, Leschen and Skelley 2002; 19 SenGupta and Crowson
1969; 20 Leschen 2003; 21 Goodrich 2002a; 22 Lawrence 1991, Goodrich 2002b; 23 Slipinski 1990; 24 Phillips and Ivie
(2002); 25 Lawrence 1982; 26 Lawrence 1982; 27 Tomaszewska 2000; 28 Vandenberg 2002; 29 Bowestead and
Leschen 2002; 30 Lawrence 1991, Andrews 2002.
* Cyclaxyridae is incertae sedis, being monotypic and having no clear affinities for any cucujoid lineage.
4
The basal position of Cucujoidea with respect to other Cucujiformia appears well
substantiated, in particular the Lymexyloidea and Cleroidea (Crowson 1960, 1964a, 1966, 1970,
Lawrence 1991, Lawrence and Newton 1982). Cladistic efforts by Leschen, Lawrence, and
Ślipiński (2005), however, suggested that Cucujoidea were paraphyletic with respect to the
Cleroidea, which was peripherally remarked upon by Crowson in his Cleroidea work (1964a,
1966, 1970) and initially substantiated by preliminary efforts by Beutel and Ślipiński (2001).
Multiple datasets of adults, larvae and molecules are needed to bring resolution to this confusion,
and to provide a reconstruction of the phylogenetic position and composition of this superfamily.
The initial impetus on family and higher level phylogenetic work in the Cucujoidea and
its constituent taxa by Crowson were continued by collaborative research with his student Sen
Gupta. Work by these two individuals over three decades formalized many families and
problematic taxa within them, including: Sphindidae (SenGupta and Crowson 1979),
Rhizophagidae (= Monotomidae) (SenGupta 1988), Boganiidae (SenGupta and Crowson 1966,
1969a), Phloeostichidae (SenGupta and Crowson 1966, 1969a, Crowson 1973), Propalticidae
(Crowson 1955, Crowson and Sen Gupta 1969, SenGupta and Crowson 1971), Hobartiidae
(SenGupta and Crowson 1966, 1969a), Cavognathidae (Crowson 1964b, 1973, SenGupta and
Crowson 1966, 1969a), Lamingtoniidae (SenGupta and Crowson 1969b), Languriidae (=
Erotylidae) (SenGupta 1967, 1968a and 1968b, SenGupta and Crowson 1967, 1969b,1971),
Erotylidae (SenGupta 1969), Sphaerosomatidae (= Alexiidae) (SenGupta and Crowson 1971),
Cerylonidae (SenGupta and Crowson 1973), and Merophysiidae (= Endomychidae in part)
(SenGupta 1979). The work by Crowson and Sen Gupta demonstrates the first two major
modern contributions in cucujoid research (a total of three major contributions were proposed by
Lawrence and Newton 1982, two of which include the erection and definition of endemic south
temperate families, i.e. Boganiidae, Hobartiidae, Propalticidae, Cavognathidae, and
5
Lamingtoniidae; and clarification of the Erotylidae/Languriidae whereupon the Cryptophagid-
lineage could be resolved).
The third contribution to cucujoid systematics as suggested by Lawrence and Newton
(1982) was delimitation of the cerylonid-series. The “cerylonid-series” has been recognized as
the most highly derived grouping of cucujoid beetles (Ślipiński and Pakaluk 1991). The group
was originally defined by Crowson (1955). Ślipiński and Pakaluk (1991) included the following
families in the lineage: Alexiidae, Bothrideridae, Cerylonidae, Coccinellidae, Corylophidae,
Discolomidae (= Discolomatidae), Endomychidae, and Latridiidae. The group was characterized
by: adults with tarsal formula 4-4-4 or 3-3-3 in both sexes, wing without a closed radial cell and
reduced number of anal veins, aedeagi resting on side when retracted and with highly reduced
tegmen, larvae with unisetose tarsungulus, annular spiracles, and sensory appendage of second
antennal segment not as long as third segment. Current efforts by McHugh (University of
Georgia) and his students are focused on resolving the status of the cerylonid-series with respect
to other Cucujoidea, its constituent members, and phylogenetic progress within specific families.
The families Sphindidae and Protocucujidae were thought to be sister taxa (Crowson 1954, 1955,
and SenGupta and Crowson 1979), and at a basal position within Cucujoidea (Lawrence 1991,
McHugh 1993, Chiao and McHugh 2000, Ślipiński 1998, Beutel and Ślipiński 2001).
Crowson’s assessments of the relationship between these two families was based on adult
morphological features; Ślipiński ’s, McHugh’s, and Chiao and McHugh’s research combined
both adult and larval characters, and Lawrence’s and Beutel and Ślipiński ’s work further
developed larval characters. Thus, the basal clade of Cucujoidea appears well corroborated with
both larval and adult character systems. These works constitute what I propose as the fourth
major contribution to cucujoid systematics, i.e. the establishment of a basal lineage (Sphindidae-
6
Protocucujidae) that can be used for evaluating character polarity and tree rooting in other
cucujoid lineages.
With both basal and highly derived cucujoid lineages defined, the next contributions in
cucujoid systematics must be focused on clarifying remaining familial clades, including: the
nitidulid-lineage (Nitidulidae, Kateretidae, and Smicripidae); the cucujid-lineage (Cucujidae,
Laemophloeidae, Silvanidae, Passandridae, Propalticidae); the cryptophagid-lineage
(Cryptophagidae, Erotylidae, Byturidae, and Biphyllidae); and the remainder of the unplaced
families (e.g. Monotomidae, Boganiidae, Cyclaxyridae, Phloeostichidae, Helotidae, Phalacridae,
Hobartiidae, and Cavognathidae). Thomas (1984a, 1984b, 1984c) helped solidify relationships
between some members of the cucujid-lineage, providing a basis for elevating the
Laemophloeidae to a family and clarifying issues within Silvanidae. To provide a comprehensive
phylogeny of the Cucujoidea, it will be necessary to do more than strip away basal and derived
lineages leaving a polyphyletic assemblage of groups. Research on the remainder of the
Cucujoidea as well as a complete treatment of the entire superfamily is in great need.
TAXONOMIC BACKGROUND. II. NITIDULID-LINEAGE
The Nitidulid-lineage includes three families: Nitidulidae, Kateretidae, and Smicripidae.
The limits of these three families had been mired in controversy and until recently the two latter
families were designated as subfamilies of Nitidulidae. These close affinities are illustrated in
Figure 1. Price (2002) offered a modern generic review of the family Smicripidae, and Habeck
(2002b) provided a similar review of Nearctic Kateretidae (= Habeck’s Brachypteridae) with
notes on global classification. However, neither of these reviews provided information on
constituent species nor phylogenetic problems within the respective families.
The limits of Smicripidae were first assessed by Böving and Craighead (1931) and were
based solely on larval characters. Prior to this work, LeConte (1878), Sharp (1900), Casey
7
(1916), Leng (1920), and Hetschko (1930) all placed this taxon into the Monotomidae (=
Rhizophagidae) and Horn (1879) placed them within Nitidulidae. Parsons (1943) suggested a
smicripid affinity to Cucujidae, and Arnett (1963) placed the group in Monotomidae. The
following characters, when taken in combination, define Smicripidae: antennae 11 segmented
with a loose 3-segmented club, pygidium and hypopygidium longer than preceding 4 segments
combined, two abdominal tergites visible from above, frontoclypeal suture deeply impressed and
curved, and maxilla with single lobe. No global treatment of the family exists; the last
taxonomic work on the group was completed by Casey (1916).
The Kateretidae (sensu Audisio 1994, 1995), unlike Smicripidae, have had a much more
linked history with Nitidulidae, remaining a nitidulid subfamily until recently (Kirejtshuk
1986c). However, the family was defined as having a maxillary lacina with two lobes instead of
one, though this was traditionally thought to be a subfamilial attribute and numerous workers
included them with nitidulids (Arnett 1963, Blackwelder 1945, Blatchley 1910, and Downie and
Arnett 1996). Verhoeff (1923) elevated Kateretidae to a family based on larval characters.
Audisio (1984) suggested a split of Kateretidae from Nitidulidae based on adults, which was
further elaborated and formalized by Kirejtshuk (1986c) based on genitalia and a preliminary
phylogeny. Audisio’s (1993) work remains the most comprehensive treatment of Kateretidae.
No global treatment of any genus exists at any hierarchical level, and only a few generic
revisions are available, but which are regional in scope (Parsons 1943, Audisio 1979, 1989,
Hisamatsu 1976, Jelínek 1976, 1979a, Kirejtshuk 1988b, 1989).
TAXONOMIC BACKGROUND. III. NITIDULIDAE LATRIELLE 1802
Nitidulidae, or sap beetles, are the second most diverse family of cucujoid beetles (after
Coccinellidae containing >6,000 species) with more than 4,000 described species (Lawrence
8
1, 4
2, 3, 12, 14,
17, 20 6,13
5, 7 11,
16, 19
8, 9,
10, 18
15
NITIDULIDAE
SMICRIPIDAE KATERETIDAE
Figure 1. Venn diagram representing the similarities between the three Nitidulid-lineage
families (Based on Lawrence 1999a, 1999b, and pers. obs.). 1) procoxae strongly transverse;
2) abdominal process truncate/indentate; 3) antennal club always loose; 4) prothoracic trochantin always
exposed; 5) maxilla with lacinia only; 6) maxilla with galea and lacinia; 7) tarsal formula either 4-4-4 or
5-5-5; 8) procoxal cavities always open externally; 9) postcoxal lines on metaventrite always absent; 10)
apical area of hindwing with branches RA and/or RP absent; 11) anal lobe of hindwing present; 12)
metendosternal laminae reduced or absent (except in highly evolved inquilinous forms such as
Cychramptodes and Cylindroramus from Australia); 13) tegminal paramera not fused to phallobase; 14)
5th and 6th abdominal spiracles lacking; 15) abdominal ventrite 1 not much longer than 2; 16)
mesotarsomere 1 well-developed and visible; 17) posterior edge of hindwing with long fringe of hairs;
18) posterior edge of head capsule in larvae always distinctly emarginated; 19) frontal arms on larval head
capsule lyriform; 20) frontal arms on larval head capsule u- or v-shaped
1982, updated in Table 2), although this number will likely increase several fold because there
has been little to no modern (after 1950) comprehensive work in the Neotropics, Afrotropics, or
SE Asia. Some historical studies (prior to 1950) were undertaken by Grouvelle in the Old and
New World tropics (1896, 1897, 1898, 1899a, 1899b, 1899c, 1894, 1901, 1905a, 1905b, 1906,
1908a, 1908b, 1910, 1914a, 1914b, 1915, 1916, 1919); Erichson in the Old and New World
tropics (1843); Sharp in the Neotropics (1890); Murray in the Old and New World tropics (1864,
9
1867, 1868); and Reitter in the Old and New World tropics (1873, 1874a, 1874b, 1875, 1876a,
1876b, 1876c, 1880, 1884). These authors described more than 75% of all known genera and
more than 50% of all species. The works listed above are the more salient products of their
efforts to document tropical Nitidulidae, in addition to numerous smaller publications.
Some regional treatments of the nitidulid fauna, both historic and modern, have been
completed in well-defined areas such as: Japan (Reitter 1877, 1883 (adults) and Hayashi 1978
(larvae)), Korea and the Chejudo Islands (Chujo 1994 and 1992, respectively), North America
(Parsons 1943, following Horn 1879), Europe (Audisio 1993, Jelínek 1965a, 1996, and
Kirejtshuk 1997a), and part of the Russian Far East (Kirejtshuk 1992). Kirejtshuk initiated a
modern treatment of the nitidulids of the Himalayas and northern Indochina from which the first
volume was published (Kirejtshuk 1998a). Kirejtshuk also published numerous, isolated
taxonomic papers on new taxa from Australia (Kirejtshuk 1986b, 1987, 1988a, 1990a, 1990b,
1992a, 1992b, and 1997a), although a thorough revision was never completed.
Modern research on nitidulids in the tropics has consisted of revisions or partial revisions
of genera or genus groups in an area, for example the Axyroid-group from SE Asia (Audisio and
Jelínek 1993), Ithyra and Neothalycra from Africa (Audisio and Kirejtshuk 1983), Pocadius in
the Neotropics (Jelínek 1977a), Hebascus and Teichostethus in the Neotropics (Jelínek 1975),
Epuraea in Africa (Jelínek 1977b), Anister from Africa and the Middle East (Jelínek 1981a),
Stelidota in the Australasian region (Jelínek 1984), Mystrops from the Neotropics (Gillogly
1955, 1972 and Jelínek 1969), Cryptarchinae genera in the Afrotropics (Kirejtshuk 1981),
Neopallodes from the Indo-Malayan region (Kirejtshuk 1994a), Mystropini from the Neotropics
(Kirejtshuk and Jelínek 2000), Phenolia (Lasiodites) from Africa (Kirejtshuk and Kvamme
2002), Cychramus from Japan (Hisamatsu 1958), and Meligethes from South Africa (Spornraft
and Kirejtshuk 1993). However, in the Neotropics hyperdiverse genera including Colopterus
10
(Larry Watrous completed dissertation work on this genus, however it was not published and
remains available only as unpublished data), Stelidota, Camptodes, Mystrops, Brachypeplus,
Conotelus, Cyllodes, Pallodes, and Cryptarcha have received little or no attention by
taxonomists, and no comprehensive taxonomic works of any globally distributed genera
including Carpophilus, Epuraea, Cychramus, Cyllodes, or Soronia were completed until the
work reported here on Pocadius.
The modern literature contians isolated new species or genus descriptions from the
tropics and subtropics including but not limited to: Cnipsarcha Jelínek (1982) from Chile, new
species of Vietterchnus and Ceramphosia Kirejtshuk and Kirk-Spriggs (1996) from the Indo-
Malayan region, Cryptarchopria kabokowi Kirejtshuk (1979) from Vietnam, new species of
Atarphia, Lordyrodes, Pocadites, Trimenus, and other genera from the Indo-Malayan region
(Kirejtshuk 1984a), new Cyllodes from Vietnam (Kirejtshuk 1985), new Propetes from Vietnam
and the Philippines (Kirejtshuk 1997c), new Lasiodactylus from the Neotropics (Cline and
Carlton 2004b), new Epuraea (Orthopeplus) from Mexico (Cline and Carlton 2004a), a new
Psilotus from Peru (Cline 2004b), two new Pocadius from the Neotropics (Leschen and Carlton
1994), a new Eusphaerius from the Neotropics (Leschen and Carlton 1996), and a new Pallodes
from the southern U.S. (Leschen 1988).
Several regional nitidulid checklists are available covering the Neotropics (Blackwelder
1945), the Nearctic (Poole and Gentili1996), Vietnam and Laos (Kirejtshuk 1997c), the
Anatolian, Caucasian and Middle East regions (Audisio et al. 2000), Italy (Audisio 1993), the
Philippine and Bismarck Islands (Gillogly 1969), Bhutan (Jelínek 1978), the Tokara Islands
(Nakane 1959), Great Britain (Kirk-Spriggs 1996), Hungary (Audisio 1981, 1987, 1996),
Namibia (Ferrer et al. 2000), Albania (Jelínek 1965a), Mongolia (Jelínek 1965b, 1966),
11
Afghanistan (Jelínek 1964, 1967a), Turkey (Jelínek 1967b), Saudi Arabia (Jelínek 1979b, 1988),
and Iran (Jelínek 1981b).
Keys for higher taxa in various regions include: global subfamilies of Nitidulidae
(Kirejtshuk 1986a); global subfamilies and tribes of Nitidulidae (Kirejtshuk1998a); the world
Nitidulinae genera (Gillogly 1965); the world Cyllodini (Leschen 1999); Meligethinae of Great
Britain (Kirk-Spriggs 1996); the world Physoronia-group of genera (Jelínek 1999a); the world
Aethina-complex of genera (Kirejtshuk and Lawrence 1999); the world Thalycra-complex of
genera (Kirejtshuk and Leschen 1998); Neotropical Mystropini genera (Kirejtshuk and Jelínek
2000); Australian Cychramptodini and Thalycrodes-complex of genera (Kirejtshuk 1992a,
1992b); Oriental Cryptarchinae (Jelínek 1974); Afrotropical Cryptarchinae (Kirejtshuk 1981),
and other more historical keys such as Grouvelle (1908a) for genera and species of nitidulids and
kateretids from India, and Reitter (1873) for genera and species of nitidulids in South America.
Only one comprehensive global taxonomic catalogue has been produced, Pars 56 of the
Coleopterum Catalogus (Grouvelle 1913). A recent endeavor by Jelínek and Audisio will seek
to remedy this scarcity of nitidulid catalogues. These individuals are collaborating on a Catalog
of the Palearctic Nitidulidae, of which there is already a preliminary publication addressing some
nomenclatural issues (Jelínek and Audisio 2003). During the course of research for this generic
revision, numerous advances have been made on New World Nitidulidae including generic
reviews for Epuraea (Orthopeplus) (Cline and Carlton 2004a) and Lasiodactylus (Cline and
Carlton 2004b), new distribution information for endemic western North American Pocadius
(Cline 2003a) and Thalycra (Cline 2004a), new distribution records for some eastern North
American Lobiopa (Shockley and Cline 2004), a new nitidulid to the U.S. from the Neotropics
(Cline 2003b), a new Psilotus from Peru (Cline 2004b), and nitidulids associated with the fungus
Pleurotus ostreatus Fries (Cline and Leschen 2005).
12
Very few complete monographs exist for nitidulid genera containing more than ten
species. Kirejtshuk (1994a) revised Neopallodes Reitter, which included 20 species.
Neopallodes are associated with the sporocarps and thalli of Basidiomycetes, and attain their
highest diversity in the Eastern hemisphere, especially SE Asia. Cooper (1982) revised Pria
Stephens, which includes 73 species. Pria is an Old World taxon with its highest diversity in
Africa and surrounding areas. Members of the genus are associated with various flowers and
vegetation. Cooper’s monograph represents the largest comprehensive generic revision to date.
Few taxonomists work on nitidulids, perhaps because these beetles have small,
generalized body forms and obscure habits. Nitidulids range in size from minute forms less than
1mm in length to moderate sized beetles ~15mm long. The three most abundant and speciose
taxa, Meligethes Stephens, Carpophilus Stephens and Epuraea Erichson, are 1.5-6.0 mm in
length and most require extraction of the male genitalia or DNA to confirm species identities.
The habits of these genera include phytophagy and anthophily in Meligethes and Epuraea,
frugivory in Carpophilus, and saprophagy and fungivory in both Epuraea and Carpophilus. The
life history strategies of these taxa undoubtedly are correlated to their evolutionary success and
global distribution (see biological/ecological considerations below).
SYSTEMATICS AND CLASSIFICATION OF NITIDULIDAE
Latrielle (1802, 1807) formally defined the family based on the type genus Nitidula
(sensu Fabricius 1775), which was based on Linne’s Silpha rufipes (1758). Although Latrielle
did much for delimiting Nitidula from other unrelated Coleoptera, he did not adequately
distinguish the family from related Coleoptera or provide insights into the taxa to be included
within its limits. Erichson’s (1843) treatment was the first work not only defining in detail
characters uniting Nitidulidae, but also formally defining subfamilies, new genera (which
included the removal of other described taxa from distantly related families), and numerous new
13
species. Erichson’s work was the first true beginning of nitidulid systematics, and we begin to
see comparative morphology used to formalize groupings, i.e. subfamilies, within Nitidulidae.
Nitidulids are currently divided into ten subfamilies: Calonecrinae Kirejtshuk (1982),
Carpophilinae Erichson (1843), Amphicrossinae Kirejtshuk (= Amphicrossini, 1986a),
Meligethinae Thomson (1859), Epuraeinae Kirejtshuk (= Epuraeini, 1986c), Nitidulinae Latrielle
(1802), Cillaeinae Kirejtshuk and Audisio (in Kirejtshuk 1986a), Maynipeplinae Kirejtshuk
(1998b), Cryptarchinae Thomson (1859), and Cybocephalinae Jacquelin duVal (1858).
Systematics will from here on formalize the relationships between taxa of any lineage;
this may or may not include rigorous quantitatively substantiated efforts utilizing modern
cladistic or other tree-building protocols. In point of fact, only a few such quantitative efforts
exist for nitidulids, therefore this section will be subdivided into traditional/historical endeavors
and more modern approaches using algorithm based methods. Although it should be pointed out
that these two sections are not mutually exclusive and I do not suggest that the efforts of one are
not bound by the precepts and inferences gained by the other.
Traditional/Historical Endeavors. The status and monophyly of nitidulid subfamilies has
never been rigorously tested and it is very likely that the Nitidulinae will be paraphyletic as
currently delimited, the Cybocephalinae will be separated as a distinct family (see Lawrence et
al. 1999a, 1999b for a list of larval and adult apomorphies), and the Maynipeplinae will be a
basal member of the Cillaeinae. The only systematic views of subfamilial relationships have
been reported by Kirejtshuk (1982 and 1995), and were represented by hand-drawn diagrams
based on a rudimentary table containing 19 characters including adult morphology, the fossil
record, some biological information and his intuition about the distribution of these ‘characters’
for a few exemplar taxa from some but not all of the subfamilies . The 1982 dendrogram
represents six subfamilies, one of which is Kateretinae (= Kateretidae). The same dendrogram
14
showed the Kateretinae as the most basal subfamily, Calonecrinae and Carpophilinae as ancestral
with respect to the other nitidulid subfamilies, and the Meligethinae, Nitidulinae and
Cryptarchinae as a derived polytomy. From his 1995 dendrogram, Kirejtshuk suggested two
major groups: the carpophilin- and nitidulin-lineages. His carpophilin-lineage corresponded to
the subfamilies Epuraeinae, Carpophilinae, Amphicrossinae, and Calonecrinae with the nitidulin-
lineage containing the Meligethinae, Nitidulinae, Cillaeinae, Cryptarchinae, and Cybocephalinae.
The split of Nitidulidae into two lineages was further elaborated in Kirejtshuk’s (1998a) treatise
on Epuraeinae. Interestingly, Kirejtshuk does not address Maynipeplinae in this latest treatment,
which was published the same year. From the 1995 dendrogram, the Epuraeinae, within the
basal carpophilin- lineage, were depicted as ancestral with a Carpophilinae, Amphicrossinae, and
Calonecrinae polytomy. Within the derived Nitidulin-lineage, the Cybocephalinae were most
basal, with Cillaeinae and Cryptarchinae forming a more derived grouping, and Meligethinae and
Nitidulinae appearing as a derived polytomy. Kirejtshuk’s resistance to quantitative endeavors,
i.e. phenetics and cladistics, is a philosophical one based on his view that such methods are
examples of extreme reductionism (see Kirejtshuk 1995, pg. 13).
Alhough Kirejtshuk’s contribution to modern nitidulid systematics via rigorous analytical
methods is minimal, his traditional work on classifying nitidulid taxa has been influential. Some
of his seminal works on members of the Australian (Kirejtshuk 1986b, 1987, 1988a, 1990a,
1990b, 1992a, 1992b, and 1997a), African (Kirejtshuk 1980, 1981, 1990d, 1993, 1994b, 1996,
1998b, 2001, Kirejtshuk and Audisio 1995, Kirejtshuk and Kvamme 2002), and Asian nitidulid
faunas (Kirejtshuk 1979, 1982, 1984a, 1985, 1986d, 1987, 1990c, 1994a, 1997c, Kirejtshuk and
Kirk-Spriggs 1996) cannot be overstated. Without these works progress toward a natural
classification of the family would not be possible. Four works authored by Kirejtshuk stand out
among modern systematic treatments and deserve special mention below.
15
The first paper dealt with a predaceous nitidulid form that feeds on scale insects from
Australia and the beetle’s body is modified supposedly due to pressure by ants that tend the
scales they are eating (Kirejtshuk and Lawrence 1992a). The tribe, Cychramptodini, was
described and its affinities to another derived tribe, i.e. the Lawrencerosini, were suggested. This
latter tribe was described by Kirejtshuk (1990b) and includes highly modified body forms with
long legs and heavily sculptured surfaces thought to be associated with ants (or some other social
insect). The Cychramptodini paper suggested one purported synapomorphy between the two
tribes, i.e. the elevation of the metasternum. This character is not known in any other recognized
ant associated nitidulid genus, including a remarkably similar form from Bolivia that is also
thought to be associated with social insects (Leschen and Carlton 2004). Perhaps these two
Australian tribes should be united into a more broadly defined Lawrencerosini with the elevated
metasternum as a synapomorphy.
A second paper by Kirejtshuk and Leschen (1998), made an informal attempt to define
the Thalycra-complex of genera. They suggested that the Thalycra-complex contains 12 genera
with broad distribution except the Oriental region. The complex was defined using numerous
pleisiomorphies, with no clear synapomorphies uniting the genera or delimiting them from the
purportedly closely related Pocadius-complex as stated by them; rather the authors assumed
monophyly and a sister-group relationship between these two complexes. A potential behavioral
synapomorphy linking the two complexes (in this case feeding on fungi without a hymenium)
was used to establish monophyly. This trait also was used by Leschen (1999) in a subsequent
nitidulid paper, which quantitatively tested the limits of the tribe Cyllodini (detailed below).
A third paper dealt with the Aethina-complex of genera (Kirejtshuk and Lawrence 1999).
They defined and delimited the complex, presented a key to genera, and also reviewed the
subgenera Idaethina Gemminger et Harold and Cleidorura Kirejtshuk and Lawrence. In their
16
definition of the group they mentioned the modification of the basal portion of the pygidium as
occurring in all but two south temperate genera, i.e. Brounithina Kirejtshuk from New Zealand
and Lordyra Gemminger et Harold from Argentina. In all other genera and subgenera arc-like
impressions on the pygidium of some type and number can be found.
The fourth and final paper was a preliminary review of the tribe Mystropini (Kirejtshuk
and Jelínek 2000). In this work they made a first-attempt to formalize the tribe, disentangle the
taxonomic confusion surrounding some genera (some errors extending back to Erichson 1843),
and provided keys to genera of Mystropini and species of the genera Anthocorcina Kirejtshuk
and Nitidulora Reitter. They suggested that the synapomorphy uniting members of the tribe was
a lack of antennal grooves on the ventral surface of the epicranium. This synapomorphy seems
at best tenuous, as the absence of a character is not suggestive of shared ancestry.
Modern/Algorithmic-Based Endeavors. The first true cladistic phylogeny of a nitidulid
lineage was Endrödy-Younga’s (1978) revision of some Meligethinae genera from the Ethiopian
region of Africa. He sought to explain the rift between the Meligethes- and Pria- generic
complexes by analyzing some of the smaller, lesser known genera from tropical Africa and
Madagascar. Although not robust in scope, the paper explained the methods of Hennig, applied
them to a small group of genera, and produced a cladogram that could be tested by others.
Unfortunately, a robust ingroup and outgroup selection of taxa was lacking. Endrödy-Younga
recognized this lack of taxon sampling but also knew that the work was a preliminary assessment
of the Meligethinae and not a comprehensive approach to understanding the relationships of all
taxa within the subfamily. The paper included a discussion of characters used and their character
state transformations from plesiomorphic to apomorphic condition. An insightful discussion of
polymorphic and plesiomorphic characters and their utility, although limited, in assessing
relationships between taxa was also included.
17
The second attempt at a modern systematic view of a nitidulid group was performed by
Audisio and Jelínek (1993) on what they defined as the axyroid-group of genera. This
groundbreaking paper defined character states used in the analysis, polarized characters based on
out-group analysis, and produced a cladogram based on synapomorphic characters. The authors
used five ingroup taxa and one outgroup taxon, 16 characters, and a most parsimonious
cladogram. Their axyroid-group was supported by a single synapomorphy, the presence of a
pronotosternal mycangium. Their discussion of choice of characters and character distribution in
other taxa demonstrated the authors’ willingness to have their hypothesis of common ancestry
and relatedness of the group tested. These authors and their collaborators began the modern age
of nitidulid systematics. Audisio and DeBiase (1996) revised the genus Dapsa Latrielle in the
cucujoid family Endomychidae using similar methods. Audisio and DeBiase also initiated the
molecular taxonomy movement in Nitidulidae (see below). Jelínek (1999a) contributed greatly
to the clarification of the Nitidulinae in his treatment of the Physoronia-complex of genera.
Although mainly taxonomic in nature, Jelínek also hypothesized relationships by pointing out
that this generic complex was related to certain other genera but differed from them “by the
presumably synapomorphic presence of lateral grooves on postmentum.”
The most comprehensive cladistic revision of any nitidulid lineage was completed by
Leschen (1999) on Cyllodini Everts. Leschen attempted to solidify the traditional grouping of
glabrous convex nitidulids (conventionally referred to as Strongylinae, Strongyllini, etc. by
European workers) as a formal tribe, and with the new phylogeny in hand discussed the
evolution of convexity and phallalophagy (“stinkhorn feeding”). Leschen used 19 ingroup and 8
outgroup taxa, and 63 characters (including adult and larval morphology and fungal host use) in
his analysis. Leschen modified his trees via ACCTRAN, DELTRAN, and other weighting
measures (e.g. successive approximations and others) to produce a monophyletic Cyllodini,
18
which was still not unambiguously defined. Leschen’s derived character states supporting the
monophyly of Cyllodini, e.g. procoxal rests on the mesosternum and visible tibial lines, were
ambiguous. These two characters occur in other groups including other nitidulines as well as
other subfamilies of Nitidulidae, and they are polymorphic within some Cyllodini genera and
absent in others (i.e. Camptodes). Leschen attempted to strengthen his argumentation for a
monophyletic Cyllodini with additional tree-building and weighting techniques following the
exclusion of problematic taxa or taxa for which he did not have specimens, including larvae for
additional characters. The inclusion of larval characters was problematic as there are no
comprehensive descriptions for some genera. Thus taxon sampling became an even greater
issue, because the genera Ceramphosia, Cyllodesus and Camptomorphus were not included in
the original dataset as well as likely sister-taxa in the Lawrencerosini and other members of the
Cychramptodini. The resulting trees, therefore, offered a murky interpretation of the tribe.
Although there were problems with Leschen’s analysis, his efforts provided the first rigorous
cladistic treatment of any nitidulid lineage.
BIOLOGICAL / ECOLOGICAL CONSIDERATIONS
As their common name suggests, nitidulids can often be encountered at fresh tree wounds
and sap flows (Crowson 1981), especially members of the Cryptarchinae (e.g. Cryptarcha and
Glischrochilus), Cillaeinae (Ewing, pers. comm.), and some Nitidulinae (e.g. Phenolia, Lobiopa,
Soronia and Prometopia) (Parsons 1943, Lawrence 1991). I have used a blend of molasses,
beer, yeast, and other ingredients to mimic sap flows in bait traps, and the following genera often
have been collected: Amphicorssus, Colopterus, Carpophilus, Lobiopa, Stelidota, Epuraea,
Prometopia, Glischrochilus, Cryptarcha, Psilotus, and Brachypeplus. The greatest diversity of
nitidulid genera, however, do not feed on sap but rather feed primarily on the fruiting bodies of
Basidiomycota (with some exceptions occurring with Ascomycota and Myxomycota). Besides
19
sap and fungal feeding, other notable feeding behaviors and life history strategies include:
frugivory, saprophagy, anthophily, phytophagy, predation, necrophagy, and inquilinism with
social Hymenoptera.
Nitidulid mycophagy has interested many authors, and nitidulid fungus feeding involves
most major fungal lineages (see Lawrence 1991 for a review). Within Pocadiini (sensu
Kirejtshuk and Leschen 1998) two genera display peculiar habits of being epigeous
gasteromycetes specialists (i.e. Pocadius and Physoronia). Gasteromycetes specialists are
uncommon in Coleoptera, only occurring again in Endomychidae (Lycoperdina) and Anobiidae
(Caenocara). The gasteromycetes fungi, however, are not monophyletic (see Hibbett et al. 1997
and Binder and Bresinsky 2002), and specialization on them does not connote specialization on a
monophyletic fungal group. Most Nitidulidae, in particular Nitidulinae, feed on mushrooms and
their relatives in the Agaricales. These beetles include Apsectochilus, Carinocyllodes,
Cycolcaccus, Cyllodes, Eusphaerius, Hebascus, Neopallodes, Niliodes, Oxycnemus, Pallodes,
Somatoxus, Teichostethus, Tricanus, and others. Aphyllophorales, shelf/bracket fungi and their
relatives, represent another common host for nitidulids, in particular Lobiopa, Soronia,
Platychora, Parametopia, some Ipidia, Atarphia, Pocadites, Hebasculinus, and others. Some
nitidulines feed on hypogeous fungi in the Hymenogastrales, e.g. Rhizopogon and their relatives
(Howden 1961, Kirejtshuk and Leschen 1998). Two interesting new records for Nitidulidae
include Oxycnemus fulvus Erichson feeding on the slime-coating of a Clavicipitaceae
(Ascomycota) fungus (Cline and Bischoff, unpublished data), and Phenolia grossa (F.) feeding
and reproducing on the fruiting body of Laetiporus cinnamoides (Basidiomycota) (Cline,
unpublished data). The clavicipitaceous record for O. fulvus is the first of a host outside of
Phallales (see Navarette-Heredia 2003). The record for P. grossa may represent the first beetle
reported for this fungus species, but the fungus has not often been correctly identified (M.
20
Blackwell, pers. comm.). Larvae obtained during acquisition of P. grossa represented the first
known larva/adult/host association for the species. Fungivorous habits of other nitidulids are
evidenced in the transmittal of plant fungal pathogens (Dorsey and Leach 1956, Juzwik 1986,
Bruck and Lewis 2002).
Frugivory in Nitidulidae is most commonly associated with members Carpophilus.
Carpophilus are known from hosts including citrus, pineapple, stone fruit, figs, strawberries,
corn, almond, cherries, grapes, quince, plum, peaches, apples, and more (see Connell 1956 and
1981 for historical aspects and argumentation for a revision due to its agricultural importance).
These ubiquitous beetles occur in orchards and fields, as well as canneries, granaries, and other
places where stored products are processed and housed. Other Carpophilus spp., however, are
frugivorous on non-agricultural plants including Yucca (Huth and Pellmyr 1997, Bronstein and
Ziv 1997). Other frugivorous taxa occur in Cillaeinae, in particular Colopterus and
Brachypeplus, both of which have been collected by me in large numbers on decaying banana.
Saprophagy in detritus, subcortical cavities, or soil, is common among some taxa,
especially Epuraea, Carpophilus, some Cillaeinae, and nitidulines such as Stelidota.
Associations with nitidulids and soil ecology are undoubtedly important, in particular within
Stelidota. This genus is abundant in soil litter, often being the predominant macrocoleopteran
within a sample. The genus is widely distributed except Europe, where S. geminata, a fruit pest,
has only recently been introduced. The genus has its highest diversity in SE Asia and the
Neotropics, where it is abundant year-round with montane endemics scattered across its range
(Cline unpublished data). Fungal hyphae have been retrieved from the gut of both Stelidota
octomaculata (Say) and S. ferruginea Reitter from the Nearctic, and specimens are available to
determine gut contents from species in Costa Rica, Panama and Bolivia. Revision of this genus
will help shed light on its biology and classification, as well as the potential efficacy for which
21
species can be used in biodiversity studies (see Anderson and Ashe 2000 for commentary on
using soil beetles in conservation practices and Didham et al. 1998 on soil-inhabiting beetles in
addressing biodiversity responses with respect to forest fragmentation).
Anthophily and phytophagy are combined to include feeding on the reproductive and
vegetative structures of plants. These habits are prevalent in many nitidulids (Crowson 1981).
The most studied genus of Nitidulidae, i.e. Meligethes, is a known plant associate (see Kirk-
Spriggs 1996 for a comprehensive treatment of the British fauna). The Meligethinae are known
anthophiles and herbivores, as well as most Epuraeinae, most Cillaeinae, and some Nitidulinae
including Aethina, Camptodes, Xenostrongylus, Anister, and others. Interestingly, species of
Macrostola have begun to receive attention from tropical ecologists due to their importance as a
pollinator of palms. Recently, I identified adults and larvae of Macrostola costulata Reitter from
Colombia as part of a study on the pollination biology of the palm Xanthosoma daguense
(Garcia-Robledo et al. 2004). Mystropini is one of the most important anthophilous lineages.
These beetles are numerous in the Neotropics, and have undoubtedly been misidentified in
studies as they are readily mistaken for Epuraea (see Listabarth 1996, who unfortunately even
misspelled Epuraea), and have more restricted ranges than previously thought (Scariot and
Lleras 1991). Until recently, not only were species identifications extremely suspect, but also
generic confirmations (Kirejtshuk and Jelínek 2000). These instances of misidentification are
important because the proliferation of such literature produces an erroneous account of the
biology and subsequent ecological niches these beetles occupy, and could have profound affects
on the perceived importance of some groups over others. For example, Naskrecki and Colwell
(1995) noted the occurrence of phoretic mites on species of Mystrops that feed on the pollen of
palms. Female Mystrops, which do not exhibit marked sexual dimorphism, are not easily
distinguished from other genera of Mystropini and could be easily misidentified. The need for
22
reliable taxonomic identifications of beetles in ecological, evolutionary and other studies is
imperative for our understanding of nature. Lachance et al. (2001) suggested that perhaps
anthophily and saprophagy on yeasts might be more intimately associated than once thought.
Their results showed that anthophilous/phytophagous nitidulids often transfer specific yeasts that
could be potential feeding substrata for their larvae. These and other studies focused on specific
trophic interactions require great taxonomic attention to the entities of the study system.
Cybocephalinae, a group delimited from others in the family by both adult and larval
characters, are predators. These beetles are also peculiar for their ability to roll themselves into a
ball, i.e. conglobation, which is also seen in Eusphaerius. Cybocephalus is a diverse genus (see
Kirejtshuk 1984b, Endrödy-Younga 1968, 1982, 1984, Blumberg 1973, and Blumberg and
Swirski 1982 for treatments of the Palearctic fauna, and Yu and Tian 1995 and Tian and Pang
1994 for Chinese and Taiwanese taxa respectively) that will likely increase in species diversity
by more than ten-fold with further efforts on the Neotropical fauna. These beetles are
predaceous on scale insects, particularly Diaspididae and Coccidae, and have, therefore, been of
importance in biocontrol studies (Drea and Carlson 1988, Lima 2002, Matadha et al. 2003,
Kirejtshuk et al. 1997). Another predator of scale insects is Cychramptodes murrayi Reitter,
from Australia. Other nitidulid taxa are likely facultatively predaceous on Scolytinae larvae (i.e.
some Epuraea, Pityophagus, and Glischrochilus), however more research will be needed to
substantiate their predaceous habits (Currie et al. 1996).
Necrophagy is known in three nitidulid genera, Omosita, Nitidula, and Epuraea, with
Epuraea being the only facultative necrophagous taxon. Both Nitidula and Omosita species
occur on carrion in the latter stages of decay, feeding on the dried remains of the corpse. Their
importance in the succession of carrion decomposition is well known (Payne 1965, Payne and
23
King 1970, and Shubeck et al. 1981). The evolution of necrophagy has not been thoroughly
discussed, but likely represents a unique shift from ancestral mycophagy/saprophagy.
Inquilinism, the intimate association with social insects, is a peculiar occurrence in
nitidulids and a phenomenon that deserves more attention. At least four (three associated with
ants or termites and one with honeybees) independent originations of inquilinism within the
subfamily Nitidulinae have occurred. One event occurred in Amphotis Erichson, which is found
in association with various ant genera including Pheidole (Parsons 1943) and Lasius (Hölldobler
and Wilson 1990) from which it solicits food through simple antennation behaviors. This genus
does not have an elevated metasternum or the flattened legs as seen in the two Australasian
inquilinous tribes. The Australasian tribes Cychramptodini and Lawrencerosini include some of
the most bizarre forms in the entire Nitidulidae (see Kirejtshuk and Lawrence 1992a, and
Kirejtshuk 1990b, respectively). Only the biology of Cychramptodes murrayi is known, and it is
a predator of the wattle tick scale (Homoptera: Coccidae), and thus does not directly obtain
nutrients from social Hymenoptera, but is in intimate association with them as ants are typically
found guarding this scale insect. A recently described genus, Amborotubus (Leschen and
Carlton 2004), from Bolivia resembles the Cylindroramus species from Australia with flattened
leg segments and shielded appearance with retracted head. Amborotubus, however, was not
mentioned as having an elevated metasternum in the description, a condition indicating that it is
not likely to be a close relative of either of the Australasian tribes. Its association with social
insects has only been postulated on the basis of morphology since all known examples were
collected at lights. Aethina tumida Murray, the small hive beetle, is associated with honeybees
and the adults derive nutrients from adult bees, whereas larvae occupy combs within the hive.
There appear to be four body morphologies evolved to deal with the rigors of inquilinism:
1) Amphotis-type that is denoted by a broad flat body, such that when attacked by ants the beetle
24
lies close to the ground with legs retracted, being firmly attached to the substrate via specialized
setae on the legs, and the ants are unable to flip the beetle over to expose its venter; 2) the
Cychramptodini/Amborotubus-type that is shield-like in appearance being extremely convex
dorsally and flattened ventrally with the sides of the elytra and pronotum extending ventrally to
help protect the legs, so that when attacked the beetle lies flat on the ground with legs retracted
and the attacker is unable to grasp the highly convex surface; 3) the Lawrencerosini-type
indicated by a longer-legged beetle with heavily sculptured pronotum perhaps affording the
beetles speed and chemical camoflauge (via sequestration of frass and/or other chemicals in the
pronotal fovea) against ant attack, and 4) the Aethina tumida-type that has little or no external
modifications, but relies heavily on chemical means to confuse and distract its honeybee hosts.
The small hive beetle, Aethina tumida Murray, has become a pest of apiaries in the eastern U.S.
(see Hood 2000 for a review). Most members of Aethina (s.str.) are associated with flowering
plants. The shift to inquilinism in A. tumida is not surprising. A native Aethina from Central
America has recently been discovered as an associate in honeybee colonies, and morphological
comparisons of this species to the small hive beetle are currently underway by me.
MORPHOLOGICAL / MOLECULAR CONSIDERATIONS
The majority of revisionary taxonomic work has been accomplished using larval and
adult morphology. Species-level studies traditionally have relied on the form of male and female
genitalia (see Fig. 5), antennal segments, body vestiture, body punctation, sexually dimorphic
characters of the antennae and legs, mouthpart sensory regions, pronotal shape, elytral shape,
development of the prosternal process, pygidial shape, scutellum shape, development and shape
of coxal lines, color patterns, and overall body shape (see Fig. 2 for a dorsal view of Pocadius).
Higher level revisions have also emphasized some of these characters, but typically with a
somewhat different more broadly defined resolution. Most higher-level taxonomic works
25
PLG
MLG
MLG
SCT
PXP OAN
OAP
EXP
IAS
TDA
ANC
ANF
SCP
PDL
Figure 2. Dorsal habitus of Pocadius fulvipennis Erichson, with pro-, meso-, and meta- legs
disarticulated and close-up of antenna. Antennal funicle (ANF), antennal club (ANC), terminal
depressed area (TDA), scape (SCP), pedicel (PDL), proleg (PLG), mesoleg (MLG), metaleg
(MTG), inner apical spine (IAS), outer apical process (OAP), outer apical notch (OAN), pronotal
explanation (PXP), elytral explanation (EXP), scutellum (SCT).
26
have also included descriptions of the mentum, mandibles, maxilla, labrum, clypeus (see Fig. 4),
meso- and metasterna, shape and size of the leg segments, and abdominal segments (see Fig. 3
for ventral view of Pocadius). Distinctive larval features of Nitidulidae were described by
Böving and Rozen (1962) and those of Pocadius were discussed by Hayashi (1978) and
Lawrence (1991). A treatment of larval Pocadius was not undertaken as part of this research.
Only recently have proteins and DNA been used to acquire characters for the
reconstruction of nitidulid relationships. Audisio et al. (2000) used allozyme and RAPD
analyses to clarify the relationships among members of the Meligethes viridescens species
complex. With more than 600 species in the genus Meligethes, most of them occurring in the
Palearctic in sympatry, there is difficulty in establishing specific limits of the more cosmopolitan
species such as M. viridescens. A total of 12 enzymes were used in the isozyme analysis and
eight different oligonucleotides were used as amplification primers in the RAPD analysis. The
resulting dendrograms were produced (1978) by calculating pairwise genetic distances (Nei
1978); these provided evidence for discriminating the purported five species in the complex
using M. aeneus as an outgroup. This study was a follow up to an electrophoresis study that
discerned the identity of M. exilis (Audisio et al. 1984), and laid the foundation for the continued
study of problematic species groups in Meligethes, including Audisio et al.’s (2001) paper on the
Meligethes caracinus complex, Audisio et al.’s paper (2002) and DeBiase et al.’s paper (2003)
on other sympatric Meligethes species.
POCADIINI / POCADIUS BACKGROUND
Pocadiini are members of Nitidulinae, the most diverse subfamily of Nitidulidae with
regard to the number of constituent taxa, as well as the diversity of niches occupied (Crowson
1954, Hayashi 1978, Lawrence1991). As defined by Kirejtshuk and Leschen (1998), the tribe
Pocadiini Seidlitz (or as they state, the “Pocadius-complex”) contains: Atarphia Reitter,
27
PST
MST
MTT
MAS
MTP
MSE
MPT
APC
HYP
AST 1
AST 2-4
ATF
LAF
MTS
LVP
SF
VMF
Figure 3. Ventral habitus of Pocadius fulvipennis Erichson, with metendosternite (MTS) inset.
Anterior tendons of furca (ATF), lateral portion of ventral process (LVP), lateral arms of furca
(LAF), stalk of furca (SF), ventral median flange of furca (VMF), abdominal sternite (AST),
prosternum (PST), mesosternum (MST), metasternum (MTT), abdominal process (APC),
hypopygidium (HYP), metepisternal axillary space (MAS), metepisternum (MTP), mesepimeron
(MSE), mesepisternum (MPT).
28
AT
SAT
PT
MO
ME
B.
A.
MNT
LBM
PLP
PLP
LA PGL
STP
CRD
C.
D.
LBP
MXP
LBM
MNT
ANS
MSS
E.
Figure 4. Mouthparts of Pocadius fulvipennis Erichson. A. dorsal aspect of labrum, with
median excision (ME); B. dorsal aspect of left mandible, with apical tooth (AT), subapical tooth
(SAT), prostheca (PT), and mola (MO); C. ventral aspect of right maxilla, with palpomere
(PLP), lacinia (LA), cardo (CRD), and stipes (STP); D. ventral aspect of combined mentum
(MNT) and labium (LBM), with palpomere (PLP) and paraglossae (PGL), only elevated portion
of mentum drawn; E. scanning electron micrograph of ventral aspect of head, with mouthparts
articulated, labial palpomere (LBP), maxillary paplomere (MXP), mental/submental sulcus
(MSS), antennal sulcus (ANS).
29
AFM
LRS
MCV
AFM
LF
SPC BL
BN
A. C.
B.
AO
LRS
ISS
ER
IRS IS
MF BP
SPC
E.
D. F.
H.
AGL
PRP
ASC
EAS
TGM
ML GNC
LPR
BCL
G. ATG
ISS
IGI
Figure 5. Male and female genitalia of Pocadius fulvipennis Erichson. A. ventral aspect of the
male anal sclerite (ASC), with medial concavity (MCV) and apical fimbriae (AFM); B. ventral
aspect of eighth abdominal sternite (EAS), with apical fimbriae (AFM), lateral flange (LF), and
spiculum (SPC); C. lateral aspect of tegmen (TGM), with lateral row of setae (LRS), basal notch
(BN), and basal lobe (BL); D. ventral aspect of tegmen (TGM), with lateral row of setae (LRS),
inner row of setae (IRS), and median fossa (MF); E. ventral aspect of median lobe (ML), with
apical opening (AO), internal structure (IS), and spiculum (SPC); F. ventral aspect of primary
internal sac sclerite (ISS), with ejaculatory rods (ER) and basal piece (BP); G. ventral aspect of
articulated male intermittent organ with accessory gland (ACG) present; H. female ovipositor
(OVP), with paraprocts (PRP), gonocoxite (GNC), apical tooth of gonocoxites (ATG), lateral
prominence (LPR), baculi (BCL), and intragonocoxal invagination (IGI).
30
Hebasculinus Kirejtshuk, Hebascus Erichson, Hyleopocadius Jelínek, Lordyrodes Reitter,
Niliodes Murray, Physoronia Reitter, Pocadioides Ganglbauer, Pocadius Erichson, Pocadites
Reitter, Axyra Erichson, Taraphia Audisio and Jelínek, and Teichostethus Sharp. Confusion has
surrounded the relationships among the members of Pocadiini and those of the Thalycra complex
(which included the Thalycrodes group of genera), the Axyroid group (sensu Audisio and Jelínek
1993), and the Physoronia complex (sensu Jelínek 1999a). Members of the Physoronia complex
were included in Pocadiini by Kirejtshuk and Leschen (1998). Kirejtshuk and Lawrence (1992b)
included three genera in the Thalycrodes complex: Australycra Kirejtshuk and Lawrence,
Rixerodes Kirejtshuk and Lawrence, and Thalycrodes Blackburn. Several genera have been
included in the Pocadius-, Thalycra-, Thalycrodes-, Axyroid-, and Physoronia-genus complexes
(Table 3), and there are internal inconsistencies as to what genera constitute the Pocadius,
Physoronia, and Axyroid groups. This situation is confounded further in Kirejtshuk (1997a),
which synonymized Lordyrodes and Pocadioides as subgenera of Physoronia, but which were
retained as separate genera in later (Kirejtshuk and Leschen 1998), and which Jelínek (1999a)
subsequently conserved as subgenera.
Of the Pocadiini genera, only Pocadius is globally distributed, i.e. found on all continents
except Antarctica. Pocadius is also the most speciose of these genera, currently with more than
three times as many species as any other, including the related Physoronia, Thalycra, and
Axyroid genera. Pocadius beetles have their highest diversity in the tropics and subtropics,
particularly the Australasian and Neotropical regions. These beetles are prevalent in most forest
types and grasslands from lowland rain forests to nearly xeric conditions in higher montane
elevations, wherever conditions amenable to fungal hosts occur. Pocadius species are all
obligate fungivores, feeding and reproducing primarily in members of the gasteromycetes
(Eumycota, Basidiomycota), most commonly in puffballs, especially Lycoperdon, Calvatia, and
31
TABLE 3. Historical classification of genera related to Pocadius.
Genus Thalycrodes
Complex1Axyroid
Lineage2Thalycra
Complex3Pocadius
Complex3Physoronia
Complex4
Atarphia Reitter X X
Hebasculinus Kirejtshuk X
Hebascus Erichson X
Hyleopocadius Jelinek X X
Lordyrodes Reitter X X
Niliodes Murray X
Physoronia Reitter X X
Pocadioides Ganglbauer X X
Pocadius Erichson X
Pocadites Reitter X
Pseudoplatychora Grouv. X X
Taraphia Aud. & Jel. X X
Teichostethus Sharp X
Thalycra Erichson X
Pseudothalycra Howden X
Quadrifrons Blatchley X
Thalycrodes Blackburn X X
Rixerodes Kir.& Lawr. X X
Australycra Kir. & Lawr. X X
Neothalycra Aud. & Kir. X
Thalycrinella Kirejtshuk X
Pleuroneces Olliff X
Pocadionta Lucas X
Pocadiolycra Kir. & Les. X
Tagmalycra Kir. & Les. X
Ussuriphia Kir. X
Megauchenia MacLeay X
Axyra Erichson X
Megauchenoides Aud. & Jel. X
1Kirejtshuk and Lawrence 1992b, 2Audisio and Jelinek 1993, 3Kirejtshuk and Leschen 1998, 4Jelinek 1999a
less frequently Scleroderma (Boletales) (Donisthorpe 1935, Jelínek 1977a, 1999b, Lawrence
1991, Leschen and Carlton 1994, and Kirejtshuk 1998a). This genus, like so many other groups
of small brown non-descript cucujoids has been a quintessential “dumping ground” for other
small setose forms. More than 10 species from six genera were once classified as Pocadius, and
numerous synonymies for Pocadius species have been erected (in for P. ferrugineus and P.
audstus). This genus and tribe were long overdue for critical revisionary work.
The objectives of this thesis include: 1) an assessment and argumentation for outgroups
to be used in cladistic analysis of Pocadius; 2) a complete monograph world Pocadius species
that includes an identification key, (re)description of all species, redescription of the genus, notes
32
on the biology, seasonality, distribution, phenotypic variation, and host data for each species, 3)
phylogenetic analysis using adult and larval characters, and 4) a discussion of how the
monograph and systematic revision will lay the foundation for future work and that of other
nitidulid taxonomists/systematists.
33
CHAPTER 2. MATERIALS AND METHODS
"..I confess I often feel wearied with the work, and cannot help
sometimes asking myself what is the good of spending a week or a
fortnight in ascertaining that certain just perceptible differences
blend together and constitute varieties and not species...."
Charles Darwin
TAXONOMIC MATERIALS / METHODS
From a total of >250,000 Coleoptera obtained on loan, >2500 Pocadius
specimens and >600 non-Pocadius members of Pocadiini were examined. This material
formed the basis of this revision in conjunction with analysis of type material. All
primary type specimens from previously described Pocadius species, with the exception
of P. monticolis Lechanteur, P. brevis Grouvelle, and P. ferrugineus (Fabricius) were
examined. These types were not located, however, determined specimens that matched
species descriptions and/or were identified by nitidulid specialists were available.
Primary type species of most Pocadiini genera were also studied, as well as type material
from other species from other nitidulid genera.
Males and females (when both were available) were studied, photographed, and
dissected. Genitalia, mouthparts, internal sclerites (e.g. the metendosternite and genitalic
structures), and appendages were extracted and disarticulated with minuten pin tools.
These tools were comprised of minuten pins attached to wooden dowels (an angled
probe, recurved hook, and straight probe were all constructed for micro-dissection).
Genitalia and mouthparts were either cleaned manually with the minuten pin tools or with
34
a 5-10% KOH solution, depending on the condition of the organs; heavily sclerotized
and/or organs with excessive tissue were treated with KOH, whereas less sclerotized
and/or structures with debridement already partially to mostly completed were cleaned
with minuten tools. The cleaned organs were then placed on glycerin slide mounts for
microscopic examination. The descriptors for the external adult features illustrated
follow the terminology used in Audisio’s (1993) detailed tome on the Italian Nitidulidae,
and the terminology used for the metendosternite follows that of Crowson (1954). New
terms have been added to provide better clarity for diagnostic characters and to
consolidate multiple terms for specific anatomical features.
All drawings were prepared using a camera lucida. Genitalia drawings were
made with an Olympus® BX50 compound microscope and all other anatomical features
with a WILD® Heerbrugg dissecting microscope. Measurements were made with a
calibrated ocular micrometer. Total length is defined as the distance from the labral tips
to the pygidial apex, width is defined as the distance across the elytra base, and depth the
distance from the humeri to the mesosternum when viewed laterally. Metasternal width
to length ratio is calculated as the distance between the metepisterna and the distance
between the border of the metacoxae with the first abdominal sternite and the
mesosternum. Head width is defined as the distance between the orbits, and head length
the distance from the labral tips to the occiput. Pronotal length is defined as the distance
from the middle of the anterior margin to the middle of the posterior margin, and width
the greatest distance across the structure. Tegmen length is defined as the distance along
the longitudinal midline from the apex to the base of the structure, and the width the
greatest width across the structure. Median lobe length is defined as the distance along
the longitudinal midline from the apex to base of the structure. Unless otherwise stated,
35
body length, width and depth are defined as discrete values for taxa in which only one
specimen was observed or an average of 10 randomly selected individuals from the total
number of specimens observed (including males and females).
Photographs were prepared using a JVC® digital camera (JVC KY-F75U)
attached to a Leica® dissecting scope (Leica Z16 AP0) with z-stepper system. Images
were captured via Auto Montage Pro® (Syncroscopy®, Inc.) on an IBM® PC. Any
modification of images was performed in Adobe Photoshop®. Scanning electron
micrographs were prepared at the Florida State Collection of Arthropods in Gainesville,
FL using a JEOL system (model# JSM-5510LV).
Label information from all specimens was recorded into an Excel® file. Label
data was divided into the following categories: collection locality, collector, date
collected, collecting method, loaning institution, and notes. The notes section contained
information regarding variation in the phenotype of a particular specimen, any damage or
missing parts to the specimen (in the case of type specimens), and if the specimen
belonged to the type series for a particular species. Type labels were recorded in the
exact format they were on the specimen in the (re)descriptions, but not in the Excel file.
For type information, line breaks are here denoted by a “;” and for specimens with more
than one label the label breaks are delineated by a “/”. Holotype labels of newly
described species, as well as lectotype labels, were printed on all red paper, whereas
paratype labels were printed on all yellow paper. Labels that connoted new locality
records were provided in detail. All other label information was recorded in the Excel
database, summarized, and reported as broad geographical ranges in the distribution
section, month ranges for the seasonality/habitat section, and fungal host or other
biological peculiarity in the notes section.
36
The following acronyms were used to denote the museums and institutions that
loaned specimens or type depositories for new or described species. These abbreviations
can be found at the Bishop Museum’s website, which maintains an up to date list of all
museums worldwide (www.bishopmuseum.org/research/natsci/ento/codens-inst.html).
Those entries marked with an “*” are not included in the website.
*ACM - American Coleoptera Museum, Bulverde, TX, Jim Wappes collection
AMNH – American Museum of Natural History, New York, New York USA
ANIC – Australian National Insect Collection, CSIRO, Canberra, Australia
*ACC – Andrew Cline Collection, currently housed at the California State Collection of
Arthropods, Sacramento, CA, USA.
ASUT – Frank M. Hasbrouk Insect Collection, Arizona State University, Tempe, Arizona
USA
BMNH – The Natural History Museum (formerly the British Museum of Natural
History), London, United Kingdom
CAS – California Academy of Sciences, San Francisco, California USA
CMNC – Canadian Museum of Nature Collection, Ottawa, Canada
CMNH – The Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, USA
CNC – Canadian National Collection of Inscets, Ottawa, Canada
CSCA – California State Collection of Arthropods, Sacramento, California, USA
CUIC – Cornell University Insect Collection, Ithaca, New York USA
DEI – Deutsches Entomologisches Institut, Eberswalde-Finow, Germany
EAPZ – Escuela Agrcicola Panamerica, Tegucigalpa, Honduras
EDNC – North Carolina Department of Agriculture, North Carolina State University,
Raleigh, North Carolina USA
37
EMEC – Essig Museum of Entomology, Berkeley, California USA
FMNH – Field Museum of Natural History, Chicago, Illinois USA
FSCA – Florida State Collection of Arthropods, Gainesville, Florida USA
GSNP – Great Smoky Mountains National Park, Gatlinburg, Tennessee USA
HNHM - Hungarian Natural History Museum, Budapest, Hungary
INBC – Instituto Nacional de Biodiversidad (INBio), Santo Domingo de Heredia, Costa
Rica
LACM – Los Angeles County Museum of Natural History, Los Angeles, California USA
LSAM – Louisiana State Arthropod Museum, Baton Rouge, Louisiana USA
MCZ – Museum of Comparative Zoology, Harvard University, Cambridge,
Massachusetts USA
MEMU – Mississippi Entomological Museum, Starkville, Mississippi USA
MNGC – Museo Nacional de Historia Natural, Guatemala City, Guatemala
MNHN - Muséum National d’Histoire Naturelle, Paris, France
MTEC – Montana Entomology Collection, Bozeman, Montana USA
NMPC – National Museum (Natural History), Prague, Czech Republic
NMPI – Division of Plant Industry, New Mexico State University, Las Cruces, New
Mexico USA
OSAC – Oregon State Arthropod Collection, Corvallis, Oregon USA
OSUC – Ohio State University Collection, Columbus, Ohio USA
RLC - Richard Leschen Collection, Auckland, New Zealand
RMC - Roy Morris Collection, Lakeland, Florida, USA
RMNH - Nationaal Naturhistorische Museum (“Naturalis”), Leiden, Netherlands
RTC – Robert Turnbow Collection, Fort Rucker, Alabama, USA
38
SDMC – San Diego Natural History Museum, San Diego, California USA
SEMC – Snow Entomological Museum, University of Kansas, Lawrence, Kansas USA
TAMU – Texas A&M University, College Station, Texas USA
TMSA – Transvaal Museum, Pretoria, Gauteng, South Africa
UAIC – University of Arizona Insect Collection, Tucson, Arizona USA
UASC - Museo de Historia Natural "Noel Kempff Mercado", Santa Cruz, Bolivia
UCMC – University of Colorado Museum, Boulder, Colorado USA
UCRC – University of California-Riverside Collection, Riverside, California USA
UDCC – University of Delaware Collection, Newark, Delaware USA
UGCA – University of Georgia Collection of Arthropods, Athens, Georgia USA
UGZM –Universidade de Guadalajara Zapopan Mexico, Centro de Estudios en Zoología
Jalisco a Universidad de Guadalajara, Jalisco, Mexico
UMRM – Enns Entomology Museum, University of Missouri, Columbia, Missouri USA
UMSP – University of Minnesota, St. Paul, Minnesota USA
UNSM – University of Nebraska State Museum, Lincoln, Nebraska USA
USNM – National Museum of Natural History, Smithsonian Institute, Washington, D.C.
WSUC – Maurice T. James Entomological Collection, Pullman, Washington USA
ZISP – Zoological Institute in St. Petersburg, Russian Academy of Sciences, St.
Petersburg, Russia
ZMHB - Museum für Naturkunde der Humboldt-Universitat, Berlin, Germany
ZMUC – Zoological Museum, University of Copenhagen, Denmark
SYSTEMATIC METHODS. I. INGROUP/OUTGROUP CONSIDERATIONS
The use of outgroup taxa in assessing character polarity was implied by Hennig
(see Hennig 1965, 1966); however, the outgroup comparison method for determining
39
character polarity was not broadly defined until Watrous and Wheeler (1981) depicted it
in the following manner: for a particular character with two or more states, the state
observed in related groups is assumed to be plesiomorphic. Nixon and Carpenter (1993)
further elaborated on Watrous and Wheeler’s concept, and provided a more concise
outgroup comparison method that was recently summarized by Schuh (2000) and
followed five steps: 1) define ingroup taxa on the basis of presumed synapomorphies; 2)
select outgroups based on synapomorphies at a more inclusive level and on a higher-level
cladistic analysis; 3) perform unrooted parsimony analysis; 4) root cladogram between
outgroup and ingroup; and 5) read character polarities from cladogram. Nixon and
Carpenter (1993) suggested that outgroup taxa need not comprise a monophyletic group.
Schuh (2000) not only summarized Nixon and Carpenter’s work, but also further
suggested that phylogenetic analyses are likely best conducted using multiple outgroups.
Outgroup comparison is not a perfect methodological system and Farris (1982),
Maddison et al. (1984) and Nixon and Carpenter (1996) found this to be an
oversimplification that would obtained locally parsimonious explanations (also see
Kitching et al. 2000 for a discussion of the problems associated with outgroup
comparison methods). However, with the poor state of systematic knowledge of
Nitidulidae and reliance on a morphological dataset for this analysis, the outgroup
comparative method was the only viable alternative for ascertaining character polarity.
Due to the inherent problems with nitidulids and use of morphological data, the selection
of potential outgroup taxa was critical to the successful analysis of Pocadius. A total of
nine outgroup taxa representing species from nine genera and three subfamilies were
used. The following justification is provided the outgroup taxa in this revision.
40
Due to the purported basal position of Pocadius (Kirejtshuk pers. comm., global
distribution of genus, and large numbers of symplesiomorphic characters compared to
other related genera) in the subfamily Nitidulinae, inclusion of not only members of
closely allied genera in the Nitidulinae but also genera in more ancestral subfamilies (e.g.
Carpophilinae and Epuraeinae) was necessary. Genera used as outgroups for the
Pocadius analysis include: Thalycra Erichson, Pocadionta Lucas, Lasiodactylus Perty,
Hebascus Erichson, Teichostethus Sharp, Hyleopocadius Jelínek, Epuraea (Orthopeplus)
Erichson, and Carpophilus Stephens. These genera represented each of the complexes in
Table 2 as well as a member of the basal Aethina-complex and two ancestral nitidulid
subfamilies. All Pocadius species were included as terminal ingroup taxa.
Members of Thalycra and associated genera in the Thalycra-complex (including
Pocadionta), have been thought to be sister taxa to Pocadiini (Kirejtshuk and Leschen
1998, following Reitter 1873 and Ganglbauer 1899). Two species of Thalycra, T. acuta
Howden and T. carolina (Wickham), were used as exemplars of the genus for this
analysis. I have spent much time researching the New World members of Thalycra,
visiting the Canadian Museum of Nature to study type specimens of Howden, and also
identifying several hundreds of specimens during the course of this study as evidenced by
a short publication on new distribution records for some Thalycra (Cline 2004a). The
monotypy of Pocadionta precluded the use of anything but P. dentipes Grouvelle as an
exemplar. The endemism of Pocadionta to the Andes Mountains in Chile and Argentina
has been reported (Blackwelder 1945, Cline unpublished Neotropical catalogue). These
two genera sufficiently represented the Thalycra genus complex in the expression of both
plesiomorphic and apomorphic characters.
41
Lasiodactylus Perty constituted a member of the Aethina-complex (sensu
Kirejtshuk and Lawrence 1999). To thoroughly understand the distribution and nature of
characters used to define Lasiodactylus and the generic complex, I undertook a complete
review of the genus and described three new species (Cline and Carlton 2004b). Two
new synapomorphies were defined to establish the monophyly of this taxon, and to
provide a basis for its inclusion in this study and future analysis of the Aethina-complex.
To assess the informative nature of the two synapomorphies exhibited by Lasiodactylus
species, it was necessary to perform a study of other genera of the complex. Thus, a
preliminary review of the Aethina-complex was completed and I determined not only the
distribution of apomorphic characters within the group, but also the validity of some
characters and character systems for higher-level taxa within the Nitidulinae. One
species, L. brunneus Perty, was included in this revision.
Hebascus and Teichostethus are endemic Neotropical genera that were included in
the Pocadius-complex (Kirejtshuk and Leschen 1998, following Kirejtshuk 1997 and
Kirejtshuk and Lawrence 1990, 1992). Both genera were reviewed by Jelínek (1975),
wherein they were redescribed, types designated for each, an identification key for
species of both genera given, and remarks on the taxonomy and biology of some species
provided. Jelínek’s revision enabled me to describe 8 new species of Hebascus and 7
new species of Teichostethus (Cline unpublished data). Jelínek’s review also provided
some of the original ideas for character transformation in Pocadiini through his detailed
descriptions. In particular, his drawings and remarks on specific mouthparts including
the basal segment of the lacinia (i.e. cardo), the carinate nature of the ligula, the extent
and shape of the mandibular mola, and development of the medial excision of the labrum
can all be traced to this work by Jelínek. Also, Jelínek’s drawings of the egg-laying
42
apparatus (i.e. ovipositor), in particular the shape of the extremity of the gonocoxal
sclerites, were instrumental in developing my thoughts on the evolution of the ovipositor
in mycophagous Nitidulinae. The following species were used as exemplars in the
Pocadius analysis: H. discoideus Reitter, H. erinaceous Sharp, H. traili Sharp, T. analis
(F.), T. testaceous (Erichson), and T. villosus Sharp.
Jelínek’s efforts on Hyleopocadius (Jelínek 1977) provided insights into the
comparative morphology of a likely transitional form between the Physoronia-complex
and Pocadiini. Hyleopocadius carbonarius (Erichson) was originally placed in the genus
Pocadius in Erichson’s 1843 tome, however Jelínek provided convincing evidence to
transfer this species into a new genus, and remarked that the newly erected genus is likely
more related to the Old World members of Pocadites and other Pocadiini and
Physoronia-complex members by virtue of the prosternal process with low obsolete
apical wall, shape of anterior tibia, short terminal tibial spurs, elytral punctation and
pubescence, and the entirely dorsad position of the mesosternum. Due to the monotypy
of the genus, only H. carbonarius was included in this analysis.
The placement of Pocadius within the Nitidulinae has not been unequivocally
established. The mycophagous habits, generalized body form, setose surface, and
distinctly separated coxae suggested a likely ancestral position in the subfamily. Thus, to
thoroughly root the Pocadius revision exemplars from the ancestral Carpophilin-lineage,
i.e. Carpophilinae and Epuraeinae were used. Epuraea (Orthopeplus) is an endemic New
World subgenus. As with the previous study on Lasiodactylus, a review of this subgenus
was conducted to develop a familiarity with the characters and character state
distributions, however, this particular taxon is a distant relative of Pocadius. The
monophyly of the subgenus was discerned from the synapomorphy of fused gonocoxal
43
sclerites of the ovipositor (Cline and Carlton 2004a). Two new species were described in
the manuscript, a key to the species of the subgenus provided, and a diagnosis of the
subgenus given. During the course of the study of Epuraea (Orthopeplus), I was able to
identify numerous new Epuraea species from other subgenera (Cline unpub. data)
including: seven of Epuraea (Epuraea) from the Nearctic, three of Epuraea (Haptoncus)
from the Neotropics, and four of Epuraea (Amedanyraea) from Central America (see
Kirejtshuk and Pakaluk 1996 for a discussion of New World Epuraea subgenera). One
species, Epuraea (Orthopeplus) quadricollis (Horn) was included in this revision. The
justification of Carpophilus in this revision was similar to that of Epuraea (Orthopeplus),
i.e. to provide a clearly ancestral outgroup. One cosmopolitan species, C. humeralis (F.),
was included in this revision.
SYSTEMATIC METHODS. II. CHARACTER DISCUSSION
Characters 1-15. Modifications of the cuticle surface (including punctures, lines,
depressions, pores, and sculpturing) have been used extensively for species descriptions
and diagnoses (see Audisio 1993, Kirejtshuk 1998a, and Jelínek 1975 for general use and
Jelínek 1977a for use in Pocadius). Punctures tend to be circular to elliptical shaped
depressions on the cuticular surface that may or may not have an associated seta
projecting from it. Lines are elongate structures on the exoskeleton that often demarcate
the juncture of sclerites, areas of cuticular thickening or thinning, and moderate
invaginations. Depressions are concave regions on the external body surface, i.e. in
Pocadius the depressed frontoclypeal region (#18 below). Pores tend to be deep,
typically circular, openings of the cuticle often associated with a gland or sensory
structure. Sculpturing deals with surface markings, which vary from smooth (i.e. little to
no sculpturing) to micreticulate/rugose (i.e. minute ridges or roughened areas).
44
1. Punctation of metasternal disc: (0) absent; (1) faint/scattered; (2) deep/aggregated.
When present on the metasternal disc, punctures that are scattered are always
faintly impressed and when aggregated they are always deeply impressed.
2. Punctation of scutellum: (0) evenly dispersed; (1) aggregated anteriorly. In character
state 1 some punctures may appear evenly dispersed in the posterior third of the
structure, but most punctures are located in the anterior two-thirds.
3. Punctation of pygidium: (0) dense, <1 diameter apart; (1) dispersed, >1 diameter
apart.
4. Punctation of mental-submental sulcus: (0) smooth; (1) punctate.
5. Punctation of gula: (0) present; (1) absent.
6. Punctation of submentum: (0) present; (1) absent.
7. Punctation of head vertex: (0) punctures all similar in size; (1) punctures of two
distinct types, large and small interspersed.
8. Punctation on abdominal sternites 2-4: (0) >2 serial rows; (1) 2 rows; (2) 1 row; (3)
absent.
9. Punctation of anterior metacoxal space: (0) impunctate; (1) punctate. The anterior
metacoxal space is the area between the anterior border of the coxae and the
anterior metacoxal line. The anterior metacoxal line is always well-defined and
evenly curved laterally from the medial border of the metacoxae
10. Punctation of anterior metepisternum axillary space: (0) impunctate; (1) punctate.
11. Number of small puncture rows between larger puncture rows on elytra: (0) no
distinctly larger punctures present, punctures interspersed; (1) 1; (2) >1.
45
12. Development of metasternal line: (0) complete; (1) incomplete. A complete
metasternal line extends from the posterior margin of the metasternum anteriorly
to the border between the meta- and mesosterna.
13. Development of medial vertexal depression: (0) not extending completely between
orbits; (1) extending completely between orbits.
14. Sculpturing of mental-submental ridge: (0) granulate; (1) microreticulate; 2)
punctate/reticulate.
15. Development of mental/submental lateral sulcus: (0) absent; (1) present.
Characters 16-23. Extensions of the cuticle, as with the above cuticular surface
modifications, have been heavily used in lower-level taxonomic studies. Most cuticular
extensions include setosity, in particular: the location of particular setae in specific
localities, the type and degree of pubescence exhibited, as well as development and
placement of spines. Presence/absence of setae on the dorsal surface was demonstrated
by Leschen (1999) to be important for reconstructing the phylogeny of the Cyllodini. At
the species level, pubescence has been used repeatedly to differentiate between related
taxa. Setosity was used by Erichson (1843) in his original diagnoses of Pocadius species,
and was the key external feature used by Audisio (1993) to differentiate between the two
closely related Palearctic Pocadius species. The number of tibial spines are diagnostic
for many species of Meligethes (see Kirk-Spriggs 1996 for an overview), as well as in
several nitiduline genera, most notably Thalycra (Howden 1961).
16. Pubescence of metasternum: (0) setae short, < length of scape; (1) setae elongate, >
length of scape; (2) disc glabrous.
17. Pubescence of antennal scape: (0) moderate, <15 setae; (1) setose, >15 setae.
46
18. Pubescence of lateral margin of elytra: (0) sparsely fimbriate, setae more than 2
diameters apart; (1) densely fimbriate, setae not more than 2 diameters apart.
19. Pubescence on elytra: (0) decumbent only; (1) erect/semi-erect only; (2) both
decumbent and erect/semi-erect.
20. Pubescence on elytra (0) uniform length: (1) two different lengths.
21. Pubescence on pronotal lateral margin: (0) asetose, (1) fimbriate.
22. Development of elytral setosity: (0) diffusedly setose, (1) serially setose at 1:1, (2)
serially setose at 1:2.
23. Number of spines on lateral margin of metatibia: (0) <10 spines present; (1) > 10
spines present.
Characters 24-35. Mouthparts were used to some degree in Leschen’s (1999)
reconstruction of Cyllodini, and have been of some importance in diagnosing genera (see
Gillogly’s 1965 Nitidulinae key); however, species level use of mouthpart characters
tends to be limited except in instances of sexual dimorphism (see Cline 2004b for
mandibular characters of Psilotus).
24. Shape of mental-submental sulcus: (0) narrow, ≤ diameter of pedicel, (1) produced
laterally, >diameter of pedicel.
25. Shape of anterior border of submentum: (0) truncate; (1) indentate.
26. Shape of mentum in lateral aspect: (0) flattened; (1) bulbous, convex medially.
27. Shape of raised portion of anterior mental border in ventral aspect: (0) broadly
rounded; (1) angulate. In character state 1, the anterior borders are oblique and
join together in a distinct apex, whereas in state 2 there is no distinct apex.
28. Shape of anterior margin of labrum: (0) deeply cleft; (1) indentate.
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29. Shape of raised portion of mentum: (0) hemispherical; (1) pentagonal; (2) triangular
(3) quadrate with apical emargination (4) quadrate with apical biconcavity. If
raised portion absent, the entire shape of the mentum was assessed.
30. Development of mandibular apex: (0) simple, (1) bifid; (2) oblique.
31. Length of terminal maxillary palpus: (0) ≤length of basal 2 segments; (1) >length of
basal 2 segments. In some specimens this character may be difficult to visualize
without dissection of the entire palpus.
32. Length of mental-submental sulcus: (0) absent; (1) elongate, extending past
submentum to gula; (2) shortened, not extending past submentum.
33. Diameter of basal segment of labial palpi: (0) slender, <diameter of pedicel; (1)
robust, >diameter of pedicel. In some specimens this character may be difficult to
visualize without dissection of the entire palpus.
34. Development of mentum: (0) flattened or partially elevated in posterior 0.33; (1)
distinctly elevated in posterior 0.66.
35. Shape of terminal maxillary palpus: (0) conical; (1) barrel; (2) globular.
Characters 36-48. The shape of the ventral sclerites (including, the prosternum,
mesosternum, metasternum, metepisternum, and all abdominal sternites) had been
overlooked in species level studies, except with respect to modification of the cuticular
surface. Parsons (1943) demonstrated the importance of the metepisternal axillary space
in differentiating species groups and as a diagnostic character for species. He also noted
the importance of mesosternum and metasternum characters for delimiting genera (1972).
36. Shape of abdominal process: (0) broadly rounded without acuminate apex; (1)
narrowed with acuminate apex.
48
37. Shape of lateral aspect of prosternal process: (0) not produced; (1) produced
posteriorly; (2) produced posteriorly and anteriorly.
38. Shape of posterior face of prosternal process in lateral aspect: (0) absent (1) straight;
(2) oblique; (3) oblique with concavity.
39. Shape of posterior margin of mesosternum: (0) truncate, (1) broadly concave, (2)
indentate.
40. Development of metepisternum: (0) medial margin angulate or with slight concavity;
(1) medial margin with distinct concavity.
41. Development of carina on prosternal process: (0) no carina visible; (1) short obsolete
carina in anterior 0.25.
42. Development of carina on mesosternum: (0) obsoletely carinate; (1) moderately
carinate; (2) distinctly carinate.
43. Development of anterior portion of metepisternum: (0) absent; (1) only anterior
0.125 cut-off from rest of structure; (2) at least anterior 0.20 cut-off from rest of
structure by raised line forming small axillary space.
44. Development of the antennal grooves on head: (0) convergent; (1) parallel; (2)
absent.
45. Development of the prosternal process between procoxae: (0) broad, > length of
pedicel, (1) narrow, < length of pedicel.
46. Development of the mesosternum: (0) not carinate, (1) feebly carinate, (2) carinate.
If the carina is only present in the anterior portion of the structure then it is coded
as (1), if the carina is present along the entire structure it is coded as (2).
47. Development of the procoxal cavities: (0) open, (1) closed.
48. Development of metepisternal axillary space: (0) absent; (1) present.
49
Characters 49-56. The primary sclerites of the dorsum (including the pronotum,
scutellum, elytra, and pygidium) have been used in varying degrees at higher and lower
taxonomic levels. Leschen (1999) attempted to use overall body convexity, in particular
the dorsal convexity, for higher level systematic analysis.
49. Shape of epiplurae: (0) narrow, ≤ width of mesotibia in female; (1) broad, > width of
mesotibia in female. The female mesotibia is used due to the sexually dimorphic
condition of the mesotibia in some males that can often be observed as a
thickening of the leg segment.
50. Shape of anterior margin of pronotum: (0) truncate; (1) evenly rounded; (2)
trapezoidal.
51. Shape of posterior pronotal margin: (0) truncate; (1) bisinuate; (2) undulate.
52. Shape of scutellum: (0) broadly rounded; (1) triangular with angulate sides.
53. Shape of male pygidial apex: (0) indentate; (1) truncate; (2) broadly rounded.
54. Shape of posterior pronotal angles: (0) distinct, (2) obsolete.
55. Development of dorsal convexity: (0) flattened, (1) moderately convex, (2) greatly
convex.
56. Exposure of pygidium: (0) fully exposed; (1) partially exposed; (2) concealed.
Characters 57-62. The antennae, whether in total or segments thereof, have been
used mostly for lower-level taxonomic work, specifically the shape of the antennal club
with particular reference to the terminal antennomere. Pocadius followed this general
rule, and the shape of the terminal antennomere and placement of sensillar regions on its
surface are important for systematic and taxonomic research.
57. Shape of terminal antennomeres: (0) symmetrical; (1) asymmetrical.
58. Shape of antennal club (0) ovate; (1) obovate; (2) elongate.
50
59. Length of terminal antennomeres: (0) =length of basal 2 club antennomeres
combined; (1) >length of basal 2 club antennomeres combined.
60. Length of antennal segment 4: (0) <length of 5; (1) >length of 5.
61. Development of antennal funicle: (0) antennomeres 7-8 not flattened, disc-like; (1)
antennomeres 7-8 flattened, disc-like.
62. Development of antennal club: (0) not compact, some space evident between club
segments; (1) compact, no space evident between club segments.
Characters 63-68. The legs, in particular the shape of the individual segments and
armature of tibia are important taxonomic characters. Kirejtshuk (2003) and Kirejtshuk
and Leschen (1998) used protibial characters (i.e. lateral crenulation, presence/absence of
outer apical processes) to help define generic complexes. Meligethes treatments have
included protibial morphology, in particular dentition pattern, for species-level studies.
Likewise, Howden (1961) used protibial dentition to delimit Thalycra species.
63. Development of anterior-lateral region of protibia: (0) not produced, (1) broadly
produced or expanded, (2) produced into sharp prominence.
64. Development of protibial lateral margin: (0) smooth, (1) crenulate.
65. Development of tibial anterior-medial spines: (0) absent, (1) single spine present, (2)
two equal spines, (3) two unequal spines.
66. Development of apico-lateral protibial notch: (0) absent; (1) shallow, <length of first
tarsomere; (2) deep, >length of first tarsomere.
67. Location of protibial apico-lateral tooth: (0)absent; (1) at same level as tarsomere
junction; (2) distinctly subapical.
68. Development of tarsomeres: (0) thin, length > width; (1) thickened, width > length.
51
Characters 69-91. The aedeagus, or male intermittent organ, has had a long
history in species level taxonomic work. Surprisingly, most descriptions of nitidulid taxa
do not outwardly discuss specific structures of the aedeagus. Pocadius aedeagi, in
particular the tegmen and median lobe, are structurally similar in most species; however,
the setosity of the tegmen and development of the internal sac sclerites are quite useful.
69. Shape of apex of male spiculum gastrale (= 8th abdominal sternite): (0) truncate; (1)
indentate; (2) bisinuate; (3) crenulate; (4) ridged.
70. Shape of ventral surface of male anal sclerite shape: (0) shallowly concave; (1)
deeply concave.
71. Shape of tegminal apex: (0) truncate; (1) evenly rounded; (2) indentate.
72. Shape of apex of tegminal central fossa (i.e. fossa that houses median lobe): (0)
evenly rounded; (1) sharply incised; (2) with broad protuberance
73. Shape of ventral protuberance on phallobase: (0) produced anteriorly; (1) produced
ventrally.
74. Shape of apical sclerite of ejaculatory rods (i.e. principal sclerites of the internal sac):
(0) straight; (1) laterally curved; (2) elbowed.
75. Shape of medial margin of basal sclerite of ejaculatory rods: (0) straight; (1)
concave; (2) bisinuate.
76. Setosity of male speculum gastrale: (0) ≤10 setae; (1) >10 setae.
77. Setosity of tegminal lateral margin: (0) present from apex of central fossa to apex;
(1) only in apical 1/3 of tegmen
78. Setosity of inner tegminal row: (0) aciliate; (1) not attaining apex; (2) reaching apex
in U-shaped row.
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79. Development of subapical furrow on tegmen: (0) absent; (1) small; (2) two small
furrows; (3) elongate furrow.
80. Development of aedeagal parameres: (0) not fused to each other, (1) fused basally,
(2) completely fused throughout; (3) fused with apical division.
81. Degree of fusion of apical to basal sclerites of ejaculatory rods
(0) fused; (1) separate.
82. Degree of phallobase/paramere fusion: (0) loosely fused, (1) completely fused.
83. Location of the anal sclerite: (0) ventral, (1) posterior, (2) withdrawn into abdomen.
84. Development of tegmen: (0) without well-developed median furrow; (1) with well-
developed median furrow.
85. Setosity of spiculum gastrale: (0) >1 row of apical setae; (1) 1 row of apical setae.
86. Point of attachment of spiculum gastrale strut: (0) dorsally; (1) posteriorly.
87. Degree of fusion of ejaculatory rods to each other: (0) separate; (1) separate.
88. Number of segments comprising ejaculatory rods: (0) 1; (1) 2; (2) >2.
89. Development of tegminal lateral setae: (0) attaining apex; (1) not attaining apex.
90. Development of tegminal inner row of tegminal setae: (0) multiple rows present; (1)
single row present; (2) aciliate.
91. Development of male anal sclerite: (0) produced apically; (1) evenly rounded.
Characters 92-111. The ovipositor, or egg-laying apparatus, is a simple structure
compared to the male aedeagus. However, modifications of the gonocoxite, i.e. the
primary ovipositor sclerite, can be dramatic and host specific for oviposition. Despite
these modifications, female genitalia have been used in relatively few species-level works
(see Howden 1961 and Kirk-Spriggs 1996 for the most comprehensive use in species-
level taxonomic work). Audisio et al (2001) used the setal pattern of the gonostylus, an
53
apical prolongation of the gonocoxite, as one of several characters to validate a new
species of Meligethes. Leschen (1999) used 3 ovipositor characters in his higher-level
analysis of the Cyllodini; one of these (#115) is repeated in this study.
92. Shape of lateral margin of gonostyloid: (0) angulate; (1) sinuate.
93. Shape of the medial region of the basal gonocoxal margin: (0) broadly rounded; (1)
indentate; (2) truncate.
94. Shape of apical border of paraproct: (0) evenly rounded; (1) acute.
95. Shape of lateral margin of paraproct: (0) evenly tapering from apex to base; (1)
broadly flanged; (2) concave.
96. Shape of inner margin of gonostylus: (0) straight; (1) curved; (2) bisinuate.
97. Degree of separation of ovipositor gonocoxites:(0) ≤1 gonostyloid width; (1) >1
gonostyloid width.
98. Degree of sclerotization of medial region of gonocoxites between gonostyloids: (0)
absent; (1) present.
99. Degree of fusion of paraprocts to gonocoxite base: (0) loosely fused medially; (1)
extensively fused medially and laterally.
100. Degree of sclerotization of paraprocts: (0) medial border sclerotized; (1) medial
border and apical portion of lateral border sclerotized.
101. Development of apical gonostyloid sete-bearing region: (0) no modification; (1)
apical pit; (2) apical groove.
102. Development of sclerotized ridge along basal margin of gonocoxite: (0)
approximating lateral prominence; (1) extending to lateral prominence.
103. Development of oblique baculi on gonocoxite base: (0) absent; (1) ≤1/2 length of
gonostylus; (2) >1/2 length of gonostylus.
54
104. Development of apico-lateral region of ovipositor gonostylus: (0) evenly rounded;
(1) produced laterally into “tooth”; (2) with 2 “teeth”.
105. Development of gonostyloid terminal appendage: (0) large, (1) reduced, (2) absent.
106. Setosity of gonostyloid terminus: (0) ≤4 setae; (2) >4 setae. These numbers
indicate the presence of primary setae, not microscopic secondary setae.
107. Degree of fusion of gonocoxites: (0) separate; (1) basal fusion only in 0.5 of
structure; (2) extensive fusion.
108. Number of lateral protuberances on ovipositor gonocoxite base: (0) 1; (1) 2.
109. Cross-section of gonostyloid: (0) rounded, (1) elliptical, (2) flattened.
110. Development of intragonocoxal invagination: (0) deep, extending > 0.50 length of
gonocoxal extension; (1) shallow; not extending past 0.50 of gonocoxal extension.
111. Degree of tegmen curvature: (0) no marked declivity in lateral aspect; (1) marked
declivity in lateral aspect.
Characters 112-114. Larval characters were used in Leschen’s (1999) analysis of
Cyllodini. He used three characters, however, none of these were repeated here.
112. Development of the larval epicranial stem: (0) present, (1) absent.
113. Shape of body curvature: (0) flattened, (1) recurved.
114. Degree of separation of frontal arms (i.e. frontal sutures): (0) distant at base; (1)
adjacent at base.
SYSTEMATIC METHODS. III. PHYLOGENETIC RECONSTRUCTION
Tree Building. All characters were coded as unordered and unweighted, and
multistate characters were treated as non-additive. Phylogenetic analyses were
performed in PAUP* 4.0b10 for Macintosh (Swofford 2001, Sinauer Assoc., Inc.); all
PAUP calculations were executed on a Power Macintosh 7100/80AV with a MacOS
55
operating system. A simultaneous analysis of outgroup and ingroup taxa was performed
to polarize characters (see Farris 1972 and Nixon and Carpenter 1993). The TBR
swapping algorithm was used for all tree construction. One-hundred replicates were
performed for each tree construction. A successively weighted tree was constructed
using the strict consensus tree of Pocadius and the outgroups Carpophilus and Epuraea;
a total of 4 iterations were used in the successive weighting protocol.
Tree Statistics/Analysis. Following phylogenetic reconstruction, statistical
analyses were performed to quantitatively assess the level of support of the chosen
characters based on the phylogenies produced. Bremer methods were undertaken in lieu
of Bootstrap or Jacknife methods due to likely violation of the independence of
characters criterion. Bremer, or decay, analysis was performed using the Macintosh
software program TreeRot© (Copyright 1999, Michael D. Sorenson, Department of
Biology, Boston University, Boston, MA 02215) to determine the number of characters
needed to support a particular branch. TreeRot© aided in the determination of decay, or
Bremer support, indices (Bremer 1988) by generating a command file for PAUP or
PAUP* (Swofford 1993, 2002). The command file included 1) a constraint statement for
each node in a given shortest or strict consensus tree and 2) commands to search for trees
inconsistent with each of these constraint statements in turn (see website for details and
free download: http://people.bu.edu/msoren/TreeRot.html). The default number of
replicates executed in TreeRot© was 20. Other basic statistics computed by PAUP that
were reported include: tree length (TL), consistency index (CI), retention index (RI),
rescaled consistency index (RC), and homoplasy index (HI). Character distributions and
alternative tree topologies were analyzed using MacClade 4.0PPC (Maddison and
56
Maddison 2000). The monophyly of Pocadius was evaluated in the context of the entire
set of taxa.
57
CHAPTER 3. SYSTEMATIC TREATMENT
“Progress in natural history starts from a basis of species, and until these are accurately described
so that others can arrive at a knowledge of them, no great advance is possible.”
George H. Horn
GENERIC REDESCRIPTION
Pocadius Erichson 1843
Pocadius Erichson 1843: 318 (original description); Reitter 1873: 94, 1884: 267 (new
species LSAMounts); Horn 1879: 310 (new species accounts); Parsons 1936: 114, 1943: 235
(review of North American fauna); Jelínek 1977: 29 (review of South American fauna); Hayashi
1978: 22 (larval descriptions); Kirejtshuk 1984: 182, 1992: 191 (new species accounts);
Lawrence 1991: 456 (larval description), Audisio 1993: 171 (review of Palearctic fauna);
Leschen and Carlton 1994: 209 (new species accounts).
Type Species. Nitidula ferruginea Fabricius 1775 (= Strongylus ferrugineus Stephens
1839, Cychramus ferrugineus Heer 1841, and Pocadius ferrugineus Erichson 1843) subsequent
designation by Parsons 1943.
Diagnosis. Distinct ridge and sulcus laterad to mentum and submentum, scape
asymmetrical with convexity anteriorly, protibia with crenulate outer margin, unequal tibial
spurs, pronotum with large and small punctures interspersed, elytral pubescence and elytral
punctation serial alternating from rows of smaller simple to rows of larger umbilicate punctures,
lateral margins of elytra and pronotum fimbriate, outer apical angle of protibia produced into
sharp flat tooth, metatarsi simple without thick setal pads, head transverse with medial
depression between orbits, meso- and metatibiae finely spinulose laterally, distance between
58
mesocoxae subequal to distance between metacoxae (i.e. metacoxae not widely separated), elytra
separately rounded exposing at least part of pygidium, mesosternum situated more dorsad than
metasternum sometimes, with blunt rounded medial carina, metasternal axillary space small with
narrow elaboration posteriorly, metepisternum always with anterior 0.10-0.25 cut-off by oblique
ridge and forming a metepisternal axillary space, lateral margins of pronotum and elytra
narrowly explanate, male anal sclerite (= tergite VIII) always visible from above, tegmen
compact and always ciliate, gonocoxites always fused basally, gonocoxal apices in ovipositor
divergent and lacking styli (P. femoralis n. sp. is unique amongst Pocadius in having short
abrupt gonocoxal apices that are scarcely separated).
Description of Adult. Size: 3-6mm in length. Broadly oval to somewhat elongate oval,
convex, body varying from moderately to densely pubescent. Dorsum visibly punctate, elytra
with serial rows of punctures and setae (Fig. 2). Head wider than long, prognathous, medial
depression visible in anterior region of head prior to clypeal region. Clypeus somewhat distinct
and moderately projecting forward. Labrum visible, strongly transverse, feebly bilobed with
anterior medial emargination or incision (Fig 4a). Mandibles large and simple, often blunt
apically, with additional smaller tooth subapically on median edge; prostheca often densely
setose; basal molar region often enlarged (Fig. 4b). Maxilla with cardo elongate and in most
species biconcave laterally, stipes triangular, lacinia broadly hemispherical with dense setal
brush along medial margin, palpi elongate with second segment typically broader than others and
terminal palpomere usually longer than preceding three combined (Fig. 4c). Mentum transverse;
labium extending anteriorly from above mentum, typically with dense setose brush along
anterior margin, labial palpi 3-segmented with basal segment short and narrow, second segment
large and asymmetrical, and terminal palpomere subequal to preceding two combined (Fig. 4d).
Eyes always finely faceted. No prominence above antennal insertions. Antennae 11-segmented
59
and a little longer than length of head. Scape large, convex anteriorly, asymmetrical, bearing
setae on outer edge of anterior margin. Pedicel mostly conical in shape. Segments 3-5 variable,
but becoming more transverse towards club. Segments 6-8 often flattened, transverse, disc-like,
shorter than any of the previous segments. Antennal club compact, always three-segmented,
shape variable, though the terminal antennomere is always the largest of the club segments.
Pronotum convex, not wider than elytra, lateral edges with variable (with regards to length,
number and color) fringe of setae. Elytra a little longer than wide, separately rounded to expose
majority of pygidium. Hindwing with simple venation and fine minute ciliate posterior margin.
Epiplurae moderately wide, usually flat or with slight lateral proclivity, not reaching elytral
apices. Pygidium always more densely punctate than other dorsal sclerites. Venter punctation
mostly similar or less dense than dorsum. Prosternal process extending posteriorly past procoxal
cavities, typically convex medially. Procoxal cavities completely closed. Mesosternum situated
more dorsad than either the prosternum or mesosternum with no or not well-developed median
longitudinal carina. Meso- and metacoxal lines not diverging from coxal cavities.
Metepisternum narrowed medially, with small axillary space delimited by transverse raised
ridge. Metasternum with hemispherical glabrous region anterior to metacoxal cavity, for
reception of meta-femur. Metendosternite of the common cucujoid hylecoetoid type as defined
by Crowson (1954) but with anterior tendons arising medially on furca, sclerotized lateral arm of
furca well-developed with acute apex, stalk of furca moderately long with heavily sclerotized
ventral median flange (see Fig. 3). First abdominal sternite with process extending between
metacoxae. Sternites 2-4 similar in size to each other. Hypopygidium always more densely
punctate than any other sternite, no sexual dimorphism visible. Anal sclerite of male visible
from above, with setose posterior margin. Male pygidium typically truncate, emarginate or more
broadly rounded than that of the female. Male genitalia with tegmen consisting of a fused
60
phallobase and parameres; median lobe with long basal strut; internal sac with two sets of
sclerotized “parts” including an apical set of ejaculatory rods and a basal ejaculatory rod base,
the former of which may be a single or set of two rods, and the latter of which may or may not be
fused to the former; and an eighth abdominal sternite having two lateral sclerites associated with
one another via a membranous connection and with a long basal strut that may be as long or
longer than the median lobe strut. The anal sclerite in combination with the eighth abdominal
sternite form a “tube” through which the tegmen and median lobe pass into the bursa copulatrix
of the female with a subsequent eversion of the internal sac and associated ejaculatory rods. The
female genitalia have a membranous valvifer and paraproct that may be either loosely or
extensively fused to the well-sclerotized gonocoxites, which have two apical prolongations that
are divergent, lack terminal styli, and have pits or other apical modifications bearing setae and
some species have a recurved apical “tooth” on each prolongation; oblique baculi are often
present medially on the gonocoxal bases.
Etymology. Derivation of the Greek “poca” meaning hair or wool, apparently owing to
the vestiture of setae on the habitus.
Notes. The above description is based on the works of Audisio (1993), Jelínek (1977),
and Kirejtshuk (1992) with additional elaborations of the genitalia, mouthparts, and other salient
features such as the metendosternite. Jelínek’s (1977) revision of Neotropical Pocadius, asserted
that Pocadius was closely related to the New World genus Hyleopocadius Jelínek and Old World
genera Physoronia Reitter (= Lordyrodes Reitter) and Pocadiodes Ganglbauer based on fused,
apically divergent ovipositor gonocoxites, however, this appears to be a homeoplastic character
developed for oviposition on members of the Gasteromycetes (eluded to by Jelínek 1999a).
Synapomorphies linking all Pocadius include the flattened shape of antennomeres 6-8 (though 6
is usually less flattened than 7 and 8, the latter two being disc-like), an elevated mentum in
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posterior 0.66 of structure that forms various geometric shapes, a short epicranial stem visible
dorsally on the larval head, and recurved body of mature larvae (2nd instars and further in
development).
Distribution. Pocadius are found on all continents except Antarctica, and most major
islands and archipelagos with the exception of Greenland, Iceland, New Zealand, Madagascar,
Hawaii, and most of Polynesia. Species are also notably lacking from high latitudes.
KEY TO SPECIES
1a) Species occurring in the Old World…………………………………………….……………2
1b) Species occurring in the New World………………………………………………………..17
2a) All tarsomeres greatly thickened, 1-4 wider than long and swollen; species occurring in SE
Asia and Australia………………………………………………………..……….….……3
2b) All tarsomeres thin, 1-4 longer than wide and not swollen; species occurring in SE Asia and
throughout the Old world except Australia…………………………………..……………6
3a) Elytral pubescence moderately long, ≥ length of scape; prosternal process with
posterior face oblique. Median lobe as in Fig 152. (Australia)………..P. kirejtshuki n.sp.
3b) Elytral pubescence short, < length of scape; prosternal process with posterior face
straight…………………………………………………………..…………………………4
4a) Large, >5mm in length; terminal antennomere with clearly defined depressed region
extending across entire apex with circular areas laterally. Internal sac sclerites as in Fig
193. (Thailand)…………………...……………………….……P. majusculus Kirejtshuk
4b) Smaller, <5mm in length; terminal antennomere with depressed region not well-defined or
extending across entire apex………………………………………….………..……….....5
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5a) Posterior two-thirds of elytra and pronotal disc much darker than rest of body; male
pygidium truncate at apex. Internal sac sclerites as in Fig 187. (Sulawesi)……....
……………………………………………………………………...….P. fusiformis n. sp.
5b) Pronotal disc not darker than rest of body and only posterior on-fourth to one-eighth of
elytra darker than rest of body; male pygidium slightly emarginate at apex. Internal sac
sclerites as in Fig 171. (Sulawesi)…………………...……………...……P. barclayi n. sp.
6a) Terminal antennomere distinctly asymmetrical, longer than preceding two segments
combined…………………………………………………………………….…………….7
6b) Terminal antennomere symmetrical, equal to or shorter than preceding two segments
combined………………………………………………………………………...……….12
7a) Scutellum broadly hemispherical without distinct apical angle……………………………...8
7b) Scutellum triangular with distinct apical angle………………………………………………9
8a) Scutellum, sides of elytra, and basal half of pronotum black; all femora and tibia
testaceous; elytral surface between punctures mostly smooth and shining. Median lobe as
in Fig. 138. (Vietnam)……...…………………….………………P. decoratus Kirejtshuk
8b) Scutellum, sides of elytra, and basal half of pronotum testaceous; all femora and tibia black;
elytral surface between punctures granular and dull. Ovipositor as in Fig. 219.
(Vietnam)………...…………………………………………………..…P. femoralis n. sp.
9a) Protibia with indistinct outer apical notch (as in Fig. 2), depth < length of tarsomere 1; elytra
with markings lighter than rest of body………………………………...………………..10
9b) Protibia with distinct outer apical notch, depth ≥ length of tarsomere 1; elytra
concolorous with rest of body……………………………………………...…………….11
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10a) Terminal antennomere with distinctly depressed circular sensillar region (as in Fig. 2, i.e.
TDA) midway to apex near medial margin; pronotum unicolorous. Internal sac sclerites
as in Fig 198. (Okinawa)……………...............................................P. okinawaensis n. sp.
10b) Terminal antennomere without distinct circular sensillar region; pronotum with lateral
margins lighter than disc. Ovipositor as in Fig. 232. (Zaire)…..P. monticolis Lechanteur
11a) Terminal antennomere with distinct elliptical depressed sensillar region near medial
margin midway between base and apex; metasternum with small faint punctation and
interspaces smooth. Internal sac sclerites as in Fig 203. (India)…P. testaceous Grouvelle
11b) Terminal antennomere without distinct depressed sensillar region; metasternum with faint
moderate punctation and interspaces alutaceous. Ovipositor as in Fig. 231.
(Philippines)……………………………………………………….P. martini Kirejtshuk
12a) Male anal sclerite with produced apex (Figs. 7, 10); outer protibial notch greatly reduced or
absent (Africa)……………………………………………………………………….…..13
12b) Male anal sclerite with evenly rounded apex, not acute; outer protibial notch visible, not
reduced or absent (Palearctic).…………………………..…………………………...…..14
13a) Smaller species, ≤3mm in length; prosternal process with apical border perpendicular when
viewed laterally; elytra with decumbent and semi-erect rows of setae. Internal sac
sclerites as in Fig. 168. (West Africa)………………………………..P. africanus Kraatz
13b) Larger species, ≥3.5mm in length; prosternal process with apical border oblique when
viewed laterally; elytra with semi-decumbent and adpressed rows of setae.
Internal sac sclerites as in Fig. 181. (Southern Africa)…………..….…P. endroedyi n. sp.
14a) Anterior margin of pronotum trapezoidal; elytra unicolorous……………………………..15
14b) Anterior margin of pronotum broadly concave; elytra with pale humeri………………….16
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15a) Pronotal and elytral fimbriae elongate, longer than length of antennal scape; male
pygidium truncate apically. Internal sac sclerites as in Fig. 167. (Western
Palearctic)……...……………………………………………………….P. adustus Reitter
15b) Pronotal and elytral fimbriae not elongate, subequal to length of antennal scape; male
pygidium concave apically. Internal sac sclerites as in Fig. 184. (Western
Palearctic)………….…………………………………………P. ferrugineus (Fabricius)
16a) Elytra with rows of decumbent setae only; pronotum with dark longitudinal line
medially; basal half to two-thirds of elytral lighter in color. Internal sac sclerites as in
Fig. 197. (Japan)………………..………………………………………..P. nobilis Reitter
16b) Elytra with rows of semi-erect setae; pronotum uniformly dark with only extreme lateral
margin lighter; basal one-fourth to one-third lighter in color. Ovipositor as in Fig. 242.
(China)……………………...……………………...…………..P. yunnanensis Grouvelle
17a) Terminal antennomere enlarged, longer than preceding two segments combined……….18
17b) Terminal antennomere not enlarged, equal to or shorter than preceding two segments
combined………………………………………………………………………………....36
18a) Pronotum widest in posterior third or near posterior angles……………………...……….19
18b) Pronotum widest near middle, not in posterior third or near posterior angles………….....26
19a) Metasternal disc with few minute punctures or impunctate…………………………….....20
19b) Metasternal disc densely punctate………………………………………………..………..22
20a) Male pygidium emarginate at posterior margin; apical protibial notch shallow, equal to 0.5
length of tarsomere 1; terminal antennomere not greatly enlarged, <0.25 larger than
preceding two segments combined. Internal sac sclerites as in Fig. 199. (Venezuela)…..
…………………………………………………………………….…….…..P. pecki n. sp.
65
20b) Male pygidium truncate at posterior margin; apical protibial notch deep, greater than 0.5
length of tarsomere 1; terminal antennomere greatly enlarged, >o.25 larger than
preceding two segments combined....……………………………….………………….21
21a) Body dark reddish brown in color; pronotum and elytra with short fimbriae, subequal or
equal to the length of the antennal scape; epimeron without distinct rows of large
punctures. Internal sac sclerites as in Fig. 169. (Brazil)………....P. antennuliferus n. sp.
21b) Body light gold-brown in color; pronotum and elytra with elongate fimbriae, much greater
than the length of the antennal scape; epimeron with two distinct rows of large punctures.
Internal sac sclerites as in Fig. 178. (Guyana)…………..…………..…….P. crypsis n. sp.
22a) Elytra with alternating rows of erect and decumbent/adpressed setae……………….........23
22b) Elytra with alternating rows of erect and semi-erect setae……………………………...…24
23a) Body surface shining with smooth to finely alutaceous sculpturing; elytra with apices and
part of lateral margin much darker than rest of body; metasternum W:L < 3. Internal sac
sclerites as in Fig. 186. (SE Brazil and Argentina)………………….…P. fumatus Jelínek
23b) Body surface not or only slightly shining with mostly granular to microreticulate
sculpturing; elytra concolorous with rest of body; metasternum W:L > 3. Internal sac
sclerites as in Fig. 176. (Bolivia)…………………..………..……..P. cochabambus n. sp.
24a) Elevated portion of mentum triangular in shape; metasternum W:L ≤ 3; metasternal disc
with deeply impressed large punctures. Internal sac sclerites as in Fig. 174. (Nicaragua)..
………………………………………..........................…………..………P. carltoni n. sp.
24b) Elevated portion of mentum pentagonal in shape (as in Fig. 4); metasternum W:L > 3;
metasternal disc with faintly impressed small to moderate sized punctures. Internal sac
sclerites not as above……………………………………………………………….....…25
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25a) Procoxae much lighter than rest of body and other coxae; outer protibial notch absent; head
only slightly longer than wide (W:L = 1.35:1); body dark brown to black in color.
Internal sac sclerites as in Fig. 177. (Brazil)……...……………………..….P. coxus n. sp.
25b) Procoxae concolorous with rest of coxae and body; outer protibial notch present; head
much wider than long (W:L = 1.75:1); body light golden brown to reddish brown in color
sometimes with elytral apices and lateral margin dark brown. Internal sac sclerites as in
Fig. 200. (Peru)………………………………………………....………P. peruensis n. sp.
26a) Elevated portion of mentum pentagonal, lateral margins present and perpendicular......…27
26b) Elevated portion of mentum triangular or hemispherical, not pentagonal, without distinct
lateral margins.………….…………………………...………………………...…………31
27a) Body bicolored with the pronotum different in color than the rest of the body. Ovipositor as
in Fig. 210. (Brazil)…………………………...………………....……...…P. bicolor n. sp.
27b) Body not bicolored, pronotum concolorous with rest of body…………………………….28
28a) Abdominal process narrow with acuminate apex; elytra with alternating rows of semi-erect
setae; terminal antennomere lacking well-defined sensillar region; dorsal surface almost
completely granular in sculpture. Internal sac sclerites as in Fig. 204. (Bolivia)………….
…………………………….………………………………………..…….P. wappesi n. sp.
28b) Abdominal process broad with widely rounded or truncate apex; elytra with alternating
rows of setae that are not both semi-erect; terminal antennomere with a well-defined
sensillar region; dorsal surface variously sculptured but not completely granular….…...29
29a) Terminal antennomere with two elliptical sensillar regions; male pygidium with distinctly
concave posterior margin; posterior prosternal wall perpendicular; anterior pronotal
margin nearly truncate. Internal sac sclerites as in Fig. 183. (Paraguay)…..P. falini n. sp.
67
29b) Terminal antennomere with a single sensillar region; male pygidium with a truncate
posterior margin; posterior prosternal wall oblique; anterior pronotal margin concave or
trapezoidal…………………………………………………………...………………...…30
30a) Body smaller in size, <3.5mm in length; brown in color; prosternal process with distinct
medial concavity in lateral aspect; axillary space on metepisternum small 0.10 length of
structure; pronotum with W:L > 2:1; metasternum W:L <3:1. Internal sac sclerites as in
Fig. 191. (Costa Rica, Nicaragua)……..………………..P. jelineki Leschen and Carlton
30b) Body larger in size >3.5mm in length; black in color; prosternal process with slight medial
concavity in lateral aspect; axillary space on metepisternum moderate 0.15 length of
structure; pronotum with W:L < 2:1; metasternum W:L > 3:1. Internal sac sclerites as in
Fig. 194. (Ecuador)….……………………..P. maquipucunensis Leschen and Carlton
31a) Protibia with outer apical notch absent…………………………………………………….32
31b) Protibia with outer apical notch present…………………………………………………...33
32a) Erect and semi-erect rows of elytral setae; antennomeres 4-6 trapezoidal and subequal in
length; densely deeply punctate scutellum, metasternum L:W > 1:2; antennal club ~.85
length of segments 1-8 combined; body uniformly reddish brown. Internal sac sclerites
as in Fig. 170. (Andean Bolivia)………………….………………..………..P. ashei n. sp.
32b) Erect and decumbent rows of elytral setae; antennomeres 5-6 trapezoidal and
antennomere 4 cuboid to barrel-shaped and longer than 5 or 6; scutellum obsoletely
punctate; metasternum L:W < 1:2; antennal club ~0.75 length of segments 1-8 combined;
body dark brown with elytral apices and lateral margin darker. Internal sac sclerites as in
Fig. 179. (Argentina)……………………………………………...…P. dimidiatus Jelínek
33a) Abdominal process broad between metacoxae with widely rounded to truncate apex; large
puncture rows on elytra separated by >1.75 large puncture diameters; protibial outer
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apical notch depth shallow, ≤ 0.5 length of tarsomere 1; male pygidium with posterior
margin distinctly concave……………………………………………...……………...…34
33b) Abdominal process narrow between metacoxae with acuminate apex; large puncture rows
on elytra separated by <1.75 large puncture diameters; protibial outer apical notch depth
shallow, > 0.5 length of tarsomere 1; male pygidium with posterior margin straight…...35
34a) Elytra with alternating erect and semi-erect rows of setae; terminal antennomere with
circular to elliptical shaped sensillar region; punctures on mentum large, equal to large
punctures on vertex of head; prosternal process with posterior wall concave; metasternal
disc with faint minute punctures; head L:W < 1:1.75. Internal sac sclerites as in Fig. 196.
(Paraguay)……………………………………………………...…..…P. nigerrimus n. sp.
34b) Elytra with alternating erect and decumbent rows of setae; terminal antennomere with V-
shaped sensillar region; punctures on mentum small, equal to small punctures on vertex
of head; prosternal process with posterior wall oblique; metasternal disc with deep large
punctures; head L:W > 1:1.75. Internal sac sclerites as in Fig. 202. (Mexico)……….……
…………………………………………..…………..…………….....…P. tepicensis n. sp.
35a) Body orange brown with apical 0.5-0.66 of elytra black; elytra with alternating rows of
semi-erect and decumbent setae; punctures on mentum large, equal to large punctures on
vertex of head; prosternal process with posterior wall perpendicular; antennal club 0.6
length of segments 1-8 combined; male hypopygidium with small central asetose region.
Internal sac sclerites as in Fig. 180. (Hispaniola)...…..……...………...P. dominicus n. sp.
35b) Body uniformly light brown; elytra with alternating rows of erect and semi-erect setae;
punctures on mentum small, equal to small punctures on vertex of head; prosternal
process with posterior wall oblique; antennal club 0.95 length of segments 1-8 combined
69
male hypopygidium uniformly setose. Internal sac sclerites as in Fig. 182. (Costa Rica).
……………………………………………...…………….P. ephite Leschen and Carlton
36a) Pronotum widest in posterior third or near posterior angles……………………...……….37
36b) Pronotum widest in middle, not in posterior third or near posterior angles…………….…42
37a) Antennal scape <1.8X length of antennal pedicel; elytra with alternating rows of semi-erect
setae only……………………….……………………………………..…………………38
37b) Antennal scape >1.8X length of antennal pedicel; elytra with varying orientations of setal
rows, but both not semi-erect……………………………………………………….……40
38a) Pronotum with posterior angles indistinct and broadly rounded; prosternal process with
posterior wall concave; protibial outer apical notch absent; metasternal W:L ≥ 3:1;
mentum triangular. Internal sac sclerites as in Fig. 201. (South America)…………………
…………..……………………………………………………………P. rubidus Erichson
38b) Pronotum with posterior angles distinct, not broadly rounded; prosternal process with
posterior wall oblique; protibial outer apical notch present; metasternal W:L < 3:1;
mentum not triangular………………………………...………………………………….39
39a) Pronotum with anterior margin distinctly trapezoidal; metasternal disc with large
moderately impressed punctures; abdominal process narrow between metacoxae; body
uniformly dark reddish brown. Internal sac sclerites as in Fig. 188. (Honduras)………….
…………………………………………………………..………..……P. globularis n. sp.
39b) Pronotum with anterior margin shallowly concave; metasternal disc with small faintly
impressed punctures; abdominal process broad between metacoxae; body uniformly light
golden brown. Internal sac sclerites as in Fig. 190. (Trinidad)…………P. insularis n. sp.
40a) Abdominal process narrow between metacoxae (as in Fig. 3); prosternal process with
posterior wall concave; metasternal W:L > 2.9:1; metepisternal axillary space ≥ 0.175;
70
elytra with at least one of the alternating rows bearing erect setae. Internal sac sclerites as
in Fig. 189. (E North America through Central Mexico)………..….P. helvolus Erichson
40b) Abdominal process broad between metacoxae; prosternal process with posterior wall not
concave; metasternal W:L < 2.9:1; metepisternal axillary space < 0.175; elytra with no
rows bearing erect setae…………………………………...……………………………..41
41a) Body uniformly dark brown to blackish in color; elytra with alternating rows of decumbent
setae; mentum with small shallow punctures. Internal sac sclerites as in Fig. 175.
(Mexico, Guatemala, and Honduras)………………………………........P. centralis n. sp.
41b) Body with pronotum and head always darker in color than elytra, with elytral apices
sometimes darker as well; elytra with alternating rows of semi-erect and decumbent
setae; mentum with large moderately impressed punctures. Internal sac sclerites as in Fig.
185. (W North America and NW Mexico)……………...………..P. fulvipennis Erichson
42a) Head slightly transverse, not much wider than long (W:L < 1.3:1); pronotal anterior margin
deeply concave; metepisternal axillary space small, < 0.13 length of metepisternum.
Median lobe as in Fig. 165. (Argentina)…………………………….…...P. torresi Jelínek
42b) Head transverse, wider than long (W:L > 1.3:1); pronotal anterior margin trapezoidal or
shallowly concave; metepisternal axillary space moderate to large, > 0.15 length of
metepisternum………………………………………………………………………..…43
43a) Elevated portion of mentum pentagonal; pronotal punctation dense, separated by <1
puncture diameter……………………………………………………………………….44
43b) Elevated portion of mentum not pentagonal; pronotal punctation dispersed, punctures
separated by >1 puncture diameter….………………………………...………………....45
44a) Elytra with lighter colored maculi on humeri; antennal scape elongate, >1.9X length of
pedicel; protibia with outer apical notch deep, equal to length of tarsomere 1 and part of 2
71
combined; abdominal process narrow between metacoxae. Internal sac sclerites as in Fig.
195. (SW United States)………….………………………………………P. niger Parsons
44b) Elytra concolorous with rest of body; antennal scape not elongate, <1.9X length of
pedicel; protibia with outer apical notch absent; abdominal process broad between
metacoxae. Internal sac sclerites as in Fig. 173. (Cuba)………………P. brevis Grouvelle
45a) Head and pronotum concolorous, dark piceous to black with elytral humeri sometimes
lighter than rest of body; posterior prosternal wall oblique; metasternum and mentum
with large punctures, equal to or larger than larger ones on vertex; elytra with alternating
rows of semi-erect setae. Internal sac sclerites as in Fig. 172. (SW United States)………..
……………………………………………………………...…...…….P. basalis Schaeffer
45b) Midline of pronotum and elytra apices and suture darker than rest of body; posterior
prosternal wall concave; metasternum and mentum with small punctures, equal to smaller
ones on vertex; elytra with alternating rows of setae variable; protibial notch deep and
distinct. Internal sac sclerites as in Fig. 192. (Nayarit State, Mexico)……….....................
...……………………………………………………………...............P. luisalfredoi n. sp.
SPECIES ACCOUNTS
Pocadius adustus Reitter
(Figs. 6, 46, 85, 126, 167, 205)
Type Material Examined. HOLOTYPE (HNHM): Caucus. Occ.; Circassien; Leder
Reitter / Holotypus 1888; Pocadius ferrugin.; v. adustus; Reitter / coll. Reitter.
Non-Type Material Examined. >350 specimens from throughout the Palearctic,
ranging from the Iberian peninsula north through the British Isles and eastward through northern
Italy and eastern Europe into parts of Turkey and with eastern examples from Sweden, Belarus,
72
western parts of Russia, and the Ukraine. Kirejtshuk (1992) includes P. adustus in his treatment
of the Russian Far East, stating their presence in Siberia, the Caucuses, and Malaysia. The latter
of these countries should be confirmed to provide the most detailed distribution of this
widespread species.
Diagnosis. Most closely allied to P. ferrugineus, with which it is sympatric throughout
most of its known range. Pocadius adustus can be distinguished from this species by the broadly
concave apex of the male pygidium, more elongate hairs on the meso- and metatibia, the overall
longer body pubescence, more pronounced apico-lateral spine on meso-metatibia of males, the
prosternal process in lateral aspect with its highest convexity more posterior, and body
punctation more widely spaced throughout habitus. Differs from P. africanus by having much
longer body vestiture, particularly the elytral and pronotal fimbriae which are ~2-3 times longer,
terminal antennomere more symmetrical, interspaces between dorsal habitus punctures more
rugulose and serial elytral punctures more closely spaced. Ejaculatory rods of male distinctive,
in particular the basal piece which is more robust and resembles recurved spines.
Redescription. Length 4.2mm, Width 2.8mm, Depth 1.7mm. Body convex, somewhat
shining, light to dark reddish brown, pubescence long pale grey to golden, pronotal and elytral
fimbriae distinctly elongate (> length of scape). Pygidium and hypopygidium densely pubescent.
Head surface densely irregularly punctate, puncture surface deeply rugulose, interspaces
smooth to alutaceous, ~0.25 puncture width. Punctures not decidedly larger near orbits or
occiput. Labrum either with minute punctures and rugulose interspaces or completely rugulose.
Pronotal surface with large and small punctures interspersed throughout. Large punctures
~1.33X width of head punctures, smaller punctures ~0.33-0.5 width of head punctures;
interspaces 0.5 to 1 large puncture diameters apart, smooth to alutaceous. Each puncture bearing
an elongate semierect seta. Scutellar surface densely to moderately punctuate, punctures similar
73
in size to smaller ones on pronotum; interspaces alutaceous to rugose and separated by 0.25-0.5
puncture diameter. Eytral surface with large punctures ~2X diameter of punctures on head, each
giving rise to a decumbent seta. Small punctures ~0.25 diameter of large punctures, each giving
rise to an elongate semi-erect seta; interspaces alutaceous to rugose, larger punctures separated
from each other by 0.5-0.75 diameter, smaller punctures separated from each other by 2 small
puncture diameters. Large and small puncture rows separated from each other by ~2 small
puncture diameters. Pygidium densely irregularly punctate, punctures equal in size to small
punctures on elytra; interspaces alutaceous to rugose with some microreticulation, punctures
separated by 0.5-1 puncture diameter. Each puncture giving rise to a shorter decumbent to erect
seta, setae ~0.5 length of those on elytra.
Venter with somewhat shorter pubescence (~0.75 length of dorsal vestiture) than dorsum
with the exception of that on the head and prosternal process. Mentum with scattered punctures
similar in size to the small punctures on the pronotum, interspaces alutaceous with
microreticulation, punctures 1 diameter apart. Each puncture giving rise to a moderately
elongate seta. Submentum and gula with similar sized punctures as mentum, each giving rise to
a short seta, interspaces alutaceous to rugose. Prosternum and epimeron faintly to moderately
impressed with large punctures, punctures ~2 times larger than those on mentum, interspaces
alutaceous to rugose with microreticulation present, ~1-1 0.5 diameters apart. Mesosternum with
punctures only near posterior margin with metasternum, interspaces and anterior portion of
mesosternum rugulose. Metasternal disc faintly punctate with moderate sized punctures equal to
those on mentum. Metasternum becoming more densely punctate laterally. Interspaces
alutaceous to faintly rugose. Abdominal sternite 1 with faint punctures, equal in size to those on
metasternum, interspaces rugose, separated by ~1-2 puncture diameters, abdominal process
mostly impunctate and alutaceous. Abdominal segments 2-4 with punctures scattered throughout
74
with some punctures aligned in an irregular lateral row near the posterior border. Serial
punctures ~0.75 diameter of scattered punctures, interspaces smooth to alutaceous, serial
punctures closely spaced, ~0.25 diameter apart. Hypopygidium densely shallowly punctate,
punctures equal to those on metasternum, interspaces alutaceous with faint microreticulation,
punctures separated by 0.25-0.5 diameter of puncture.
Head wider than long (W:L = 1.5:1), fronotclypeal region well pronounced. Vertex with
deep broad concavity extending between each orbit from the posterior region of the frons to the
clypeal region. Labrum transverse with small medial incision on anterior margin. Antennal club
compact, large, obovate, asymmetrical with the last antennomere subequal to previous two
segments combined, entire club equal in length to 1.3X’s the antennal stem. Antennomeres 6-8
flattened, segment 6 less flattened than 7 and 8, their combined length slightly longer than length
of antennomere 9. Antennal scape asymmetrical, broadly hemispherical, 2 times as long as
pedicel, ≤12 setae originating from it. Pedicel more or less barrel-shaped. Antennal segment 3
equal in length to pedicel, narrowed proximally. Segments 4 and 5 globular, 0.75 the length of
segment 3. Antennal grooves deep and somewhat curved posteriomedially. Prosternal process
narrowed slightly between coxae, evenly rounded with an acute apex. Mesosternum extending
to midway between mesocoxae with a concave apex. Metasternum wider than long (W:L =
2.4:1). Metepisternum not concave medially, anterior third strongly produced anteriomedially
and impunctate with surface granular with microreticulation. First abdominal sternite with
broadly rounded process with a small point between metacoxae, ~2.3X’s longer than second
sternite. Sternites 2-4 subequal in length. Hypopygidium subequal in length to first abdominal
sternite.
Protibia with apical tooth as long as tarsomeres 1 and half of 2 combined. Outer apical
notch with no distinct angle or notch but oriented in oblique manner. Longer inner apical spine
75
subequal in length to first and second tarsomere combined. Apical border of protibia smooth, no
armature present. Mesotibia armored with a row slender spines along entire lateral edge, spines
~0.5 length of lateral setae on mesotibia. Apical border armored with 2-3 short spines, 0.33-0.5
length of lateral spines. Outer apical process elongate and robust, subequal in length to inner
apical spine. Inner apical spine equal in length to tarsomeres 1-2 combined. Metatibia more
heavily armored than mesotibia, lateral slender spines more elongate and numerous. Spines of
varying lengths, but mostly longer than those on the mesotibia. Apical border armored with a
few short spines, as in mesotibia, but more apparent. Outer apical process elongate and robust,
slightly longer than inner apical spine. Inner apical spine equal in length to tarsomeres 1-2
combined.
Male genitalia well sclerotized. Anal sclerite with large broadly curved membranous
region anteriodorsally (Fig. 6), apex densely fimbriate with elongate setae; ventrally with a deep
medial concavity approaching apex. Spiculum gastrale more lightly sclerotized than anal sclerite
and tegmen (Fig. 46), lateral region somewhat rounded and moderately explanate, apical border
undulate with >10 setae, spiculum attached posterio-medially to sclerites. Tegmen evenly
broadly rounded apically (Fig. 85), longer than wide (w:l ~ 2.6:1), lateral row of setae visible
anterior to the median fossa around the apex, inner row of setae absent, basal notch slightly
projecting forward, basal margin slightly concave. Median lobe large and robust, 0.66 the length
of the tegmen, apical opening well-developed extending posteriorly to ~ 0.5 the length of the
median lobe (Fig. 126). Ejaculatory rods not fused to each other, evenly concave, basal piece
with recurved apices and with proximally projecting medial region (Fig. 167).
Ovipositor moderately sclerotized. Gonocoxites with sclerotized basal border with one
lateral prominence, basal border giving rise to two medial oblique sclerotized baculi extending
apicolaterally, ridges long ~0.33 length of basal border. Gonocoxal apices moderately separated
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with intragonocoxal invagination evenly rounded at base, gonocoxal tips rounded at apex, each
apex with a small lateral recurved “tooth” giving rise to 5-7 short seta (Fig. 205). Valvifers
membranous with some sclerotization along medial border, evenly tapering to lateral apex.
Variation. Color varies from light red brown to dark red brown with some individuals
having dark brown elytral sutures, lateral elytral margins, and sometimes a dark scutellum. The
number of terminal setae on the gonocoxal apices varies from 5-7.
Seasonality/Habitat. Specimens studied were collected from March - October.
Distribution. Found throughout Europe eastward into Turkey and the Western Caucasus
in Russia (Audisio et al. 2000, Kirejtshuk 1992).
Notes. Pocadius adustus has been collected from the following fungal genera:
Lycoperdon, Bovista, Calvatia, and Langermannia. This species was recently redescribed by
Audisio (1993) in his treatise on Italian Nitidulidae.
Pocadius africanus Kraatz
(Figs. 7, 47, 86, 127, 168, 206)
Type Material Examined. HOLOTYPE (DEI): Togo / ♂ / Pocadius; africanus; Kraatz
95 / Coll. Kraatz; Grouvelle det. / Holotypus 1895-1968; Pocadius; africanus Kr.; Dr. Endrödy-
Younga / Coll. DEI; Eberswalde
Non-Type Material Examined. 24 specimens with the following label data: (LSAM)
GHANA: Bobiri Forest Reserve; Near FIT #1; 23-VII-2001; V. Moseley & C. Carlton collr.; in
earthstar fungi. (LSAM) GHANA: Ashanti Province; Bobiri Forest Preserve; 6˚42’N 1˚20’W;
23-31-July-2001, FIT# 3; V. Moseley and C. Carlton collr. GHANA: Ahanti Reg.; Bobiri Forest
Reserve; 6˚42’N 1˚20’W; 15-22-III-2002; E. Opuni-Frimpong collr. FIT. GHANA: Ahanti
Reg.; Bobiri Forest Reserve; 6˚42’N 1˚20’W; III-V-2002; E. Opuni-Frimpong collr. FIT.
77
Diagnosis. Differs from the Palearctic and other African fauna by its diminutive size,
lack of recurved gonocoxal apices, acute pointed apex of anal sclerite in males, and the single
fused ventrally curved ejaculatory rod in males.
Redescription. Length 2.9mm, Width 1.1mm, Depth 0.9mm. Body convex, moderately
shining, uniformly light to reddish brown, long golden pubescence throughout dorsum, setae on
head longer than that on rest of body. Pronotum and elytra narrowly explanate with margins
fimbriate. Pygidium and hypopygidium densely pubescent with posterior margins densely
fimbriate.
Head surface deeply distinctly irregularly punctate, punctures larger near orbits and
occiput becoming smaller near clypeal region with some small punctures interspersed with larger
ones on vertex. Smaller punctures ~5 times larger than eye facets, larger punctures between 2-3
times diameter of smaller punctures. Each puncture giving rise to a single elongate apically
curved seta. Interspaces narrow between lateral punctures, about 0.25 puncture diameter,
becoming wider between medial punctures about 1 puncture diameter apart; alutaceous to
smooth in texture. Labrum bearing no punctures, completely alutaceous. Pronotal surface with
large and small punctures interspersed throughout. Large and small punctures similar in size to
respective large and small punctures on head. Interspaces 0.5 to 1 puncture diameter apart,
smooth to alutaceous. Each puncture bearing a straight semidecumbent seta. Scutellar surface
densely punctate anteriorly with large punctures similar in size to the large punctures on the
head, impunctate at apex, interspaces alutaceous and 0.25 diameter of puncture in basal half.
Eytral surface with serial rows of alternating large and small shallow punctures. Large punctures
3-4 times diameter of the large punctures on head, each giving rise to a decumbent seta. Small
punctures 0.20 diameter of large punctures, each giving rise to a semi-erect seta. Interspaces
alutaceous to smooth between all punctures, larger punctures separated from each other by 0.5
78
puncture diameter, smaller punctures separated from each other by 1-2 puncture diameters.
Large and small puncture rows separated from each other by ~1 small puncture diameter.
Pygidium densely finely punctate, punctures equal in size to small punctures on elytra,
interspaces alutaceous with some minute microreticulation, punctures separated by 0.20 puncture
diameter. Each puncture giving rise to a short fine seta, setae ~0.5 length of those on elytra.
Venter with less dense pubescence than dorsum. Mentum with scattered punctures
similar in size to the small punctures on the vertex, interspaces smooth and shining, punctures
0.5 diameter apart. Submentum and gula with several large punctures equal in size to the large
punctures on the vertex, interspaces smooth and shining. Prosternum and epimeron irregularly
punctate, punctures 3 times larger than those on mentum, interspaces alutaceous, prosternal
punctures separated by 0.75 diameter, those on the epimeron by 0.25 to 0.5 diameter. Prosternal
punctures very faint. Meso- and metasternum irregularly punctate, with large faint punctures
similar in size to those on prosternum and epimeron, interspaces alutaceous becoming smooth on
metasternal disc, punctures separated by ~1 diameter. Mesosternal punctures aggregated toward
posterior margin. Abdominal sternite 1 with faint, almost obsolete, punctures occurring in all but
the anterior 0.33 of the structure which is completely alutaceous. Interspaces alutaceous,
punctures separated by 1 diameter. Abdominal segments 2-4 with punctures aligned in irregular
lateral rows. Punctures equal in size to those on abdominal sternite 1. Interspaces alutaceous,
punctures separated by 0.5 diameter. Hypopygidium densely shallowly punctate, punctures
equal to those on other abdominal sternites, interspaces alutaceous with faint microreticulation,
punctures separated by 0.25 diameter of puncture.
Head somewhat wider than long, broadly triangular, with fronotclypeal region projecting
slightly anteriorly. Vertex with deep broad concavity. Antennal club compact, broadly oval,
asymmetrical with the last antennomere longer than previous two combined. Antennomeres 6-8
79
strongly flattened into disc-like structures, their combined length equal to the length of
antennomere 9. Antennal scape asymmetrical, broadly hemispherical, 2 times as long as pedicel.
Pedicel very similar in shape to scape only smaller. Antennal segment 3 subequal in length to
pedicel. Segments 4 and 5 subquadrate, 0.5 the length of segment 3. Segments 6-8 flattened,
disc-like in shape. Antennal club large, equal in length to 2X’s the pedicel, slightly
asymmetrical, terminal segment equal to preceding two segments. Prosternal process narrowed
slightly between coxae, evenly rounded at apex. Mesosternum extending to midway between
mesocoxae with a concave apex. Metasternum much wider than long (W:L = 3:1).
Metepisternum only slightly concave medially, anterior third strongly produced anteriolaterally
and separated from rest of metepisternum by a slightly raised carina. First abdominal sternite
with broad almost truncate process ending in a small point between metacoxae, ~2X’s longer
than second sternite. Sternites 2-4 subequal in length. Hypopygidium subequal in length to first
abdominal sternite.
Protibia with apical tooth as long as second tarsomere. Outer apical notch with almost
90° angle, notch depth shallow, equal to length of first tarsomere. Inner apical spine subequal in
length to first tarsomere. Apical border of tibia smooth, no armature present. Mesotibia heavily
armored with two rows of slender spines, one row along entire lateral edge and another row
laterally oriented on ventral surface. Apical border armored with short spines, 0.33-0.25 length
of lateral spines. Outer apical process elongate and robust, 0.75 length of inner apical spine.
Inner apical spine equal in length to tarsomeres 1-2. Metatibia heavily armored with four rows
of slender elongate spines, one dorsal row, one ventral row, and two lateral rows. Spines of
varying lengths, but most longer than those on the mesotibia. Apical border armored with short
spines, as in mesotibia, but more numerous. Outer apical process elongate and robust, 0.75
80
length of inner apical spine. Inner apical spine equal in length to tarsomeres 1-2 and half of
tarsomere 3.
Male genitalia moderately sclerotized. Anal sclerite with large broadly curved
membranous region anteriodorsally (Fig. 7), and a more heavily sclerotized well-demarcated
posteriodorsal region, apex densely fimbriate, ventrally with a deep medial concavity
approaching apex. Spiculum gastrale more lightly sclerotized than other genitalic structures
(Fig. 47), lateral region broadly rounded and widely explanate, medial region elevated and
extended beyond basal margin posteriorly, apical border finely crenulate with ≤10 setae, apico-
lateral margins deeply concave, spiculum attached posteriorly to sclerites. Tegmen evenly
rounded apically (Fig. 86), much longer than wide (w:l ~ 1.5:6), lateral row of setae visible from
the median fossa around the apex, inner row of hair incomplete apically extending from apex of
median fossa to just prior to apex, basal notch projecting forward, basal margin slightly concave,
entire apex of median fossa border densely ciliate. Median lobe large and robust, slightly greater
than 0.5 the length of the tegmen, apical opening not well-developed extending posteriorly to ~
0.25 the length of the median lobe (Fig. 127). Ejaculatory rods small, less than half the length of
median lobe. Basal piece not fused to apical rod. Apical rods fused into single stout rod with
ventrally curved apex (Fig. 168).
Female genitalia lightly sclerotized. Gonocoxites with sclerotized basal border with two
lateral prominences, basal border giving rise to two medial oblique sclerotized ridges extending
apicolaterally, ridges long ~0.33 length of basal border. Gonocoxal apices widely separated with
intragonocoxal invagination evenly rounded at base, gonocoxal tips evenly rounded at apex, each
apex with a small lateral depression that gives rise to 5-6 short seta (Fig. 206). Valvifers
membranous with some sclerotization along medial border, evenly tapering to lateral apex.
81
Variation. Females are slightly larger than males, can be up to 3.2mm long and
somewhat more robust. Some individuals examined had deeper punctation on the ventral surface
than others, however, the patterns and interspaces are alike. Males typically have a slightly more
produced outer apical process on the protibia. Gonocoxites of ovipositor with between 5-7
terminal setae.
Seasonality/Habitat. The Ghana material was collected from flight intercept traps and
fungi from mid-March through July. These specimens were all within the Bobiri Forest Reserve,
which is composed primarily of upper Guinean wet forest. The specimens reported from
Angola, Zaire and Sudan are from areas dominated by lower Guinean rain forest.
Distribution. Specimens were studied from western Africa, however Kirejtshuk reported
its distribution east and southward to Angola, Zaire, and Sudan (pers. comm.)
Notes. Label data from the Ghana specimens indicated individuals collected from
“earthstar fungi”. This record indicates the occurrence of the species in the fungal family
Geasteraceae. This is the first and only known host record for this species.
Pocadius antennuliferus Cline new species
(Figs. 8, 48, 87, 128, 169)
Type Material Examined. HOLOTYPE ♂ (CMN): BRAZIL: Pará; Carajas, Serra;
Norte, III-1985; 6 15’S 50 75’W; N. Degallier; FIT, carrion, dung.
Diagnosis. This species is easily distinguished from all other Neotropical Pocadius by
the following suite of characters: head surface deeply punctate and shining with interspaces
completely smooth, clypeus distinctly indentate medially, protibial outer apical notch deep and
distinct; prosternal process when viewed laterally with drastic posterior declivity and short
posterior face, terminal antennomere greatly enlarged and asymmetrical with two lateral
82
depressed regions, metasternal disc with faint minute punctation; tegmen with incomplete rows
of inner and lateral setae, median lobe with apex narrowly rounded, and ejaculatory rods
comprised of three distinct pieces.
Description. Length 3.1mm, Width 2.4mm, Depth 1.3mm. Body moderately convex,
surface shining, dark reddish-brown in color, with legs and antennae lighter. Pronotum and
elytra margins with moderate fimbriae, setae subequal to width of antennal scape. Dorsal and
ventral pubescence moderate, fine and inconspicuous.
Head surface deeply, irregularly punctate, punctures larger on vertex, becoming
somewhat smaller towards orbits and fronotclypeal region. Larger punctures 3-4 X diameter of
eye facet, smaller punctures 2 X diameter. Interspaces smooth and shining. Each smaller
puncture gives rise to short fine golden seta. Pronotal surface with large punctures equal in size
to large punctures on vertex of head, interspersed with relatively few smaller punctures, equal to
smaller ones on vertex. Interspaces alutaceous to finely microreticulate, larger punctures ~0.5
diameters apart. Each puncture gives rise to a moderate golden seta. Scutellar surface with very
few vague shallowly impressed small punctures equal in size to smaller ones on vertex,
interspaces are alutaceous to granular. Elytral surface with serial rows of alternating large and
small deep punctures. Smaller punctures are 1.5 X diameter of smaller ones on pronotum, larger
punctures are ~1.5-2 X diameter of smaller ones. Smaller punctures giving rise to an straight
erect fine seta, larger punctures giving rise to a decumbent fine seta. Interspaces narrow between
punctures of a given row and between different rows. Within a row, small punctures are
separated by ~0.75 puncture width, and large punctures by 0.5 puncture width. Large rows are
separated by 1.0-1.5 large puncture diameters. Interspaces moderately shining, alutaceous to
finely microreticulate in sculpture. Pygidium densely punctate, punctures equal in size to those
83
on pronotum, each puncture giving rise to a short fine golden seta. Interspaces narrow, 0.5-0.75
diameter, with alutaceous to granular sculpture.
Venter with similar short fine dispersed pubescence as dorsum. Mentum with small very
shallow scattered punctures, equal in size to smaller ones vertex. Interspaces smooth to finely
alutaceous. Submentum and gula similar in punctation to mentum but with punctures more
widely dispersed. Prosternum and epimeron deeply irregularly punctate, punctures 2X larger
than larger ones on vertex, interspaces granular with microreticulate areas with apex of
prosternal process becoming smooth, prosternal punctures separated by ~0.5 diameter, those on
the epimeron by 0.5-0.75 diameter. Mesosternum with shallow punctures, about 0.5 diameter of
those on prosternum, interspaces alutaceous to granular, separated by about 0.5 to 1 diameter and
mostly aggregated near metasternum. Metasternum irregularly punctate, with faint small
punctures on disc similar in size to smaller ones on vertex and larger punctures equal to those on
prosternum, interspaces faintly alutaceous on disc becoming granular with microreticulate areas
laterally, disc punctures separated by ~2-4 diameters and lateral punctures by ~1 diameter.
Abdominal sternite 1 with small faint, almost obsolete punctures, punctures equal to smaller ones
on vertex, interspaces alutaceous, separated by ~1-2 diameters. Abdominal sternites 2-4 with
two irregular indistinct rows of punctures, one row near anterior margin and the other near
posterior margin, punctures similar in size to those on abdominal sternite 1, rows separated by ~2
puncture diameters, punctures within rows separated by ~1 puncture width. Hypopygidium with
more deep punctures than other sternites, diameter similar in size to those on sternites 2-4,
interspaces mostly alutaceous to granular, punctures separated by ~1-2 puncture diameters.
Head slightly wider than long (W:L = 1.5:1), fronotclypeal region projecting anteriorly
with clypeus distinctly indentate. Vertex with concavity between orbits near fronotclypeal
region. Labrum with deep incision at anterior margin. Antennal club compact, swollen apically,
84
distinctly asymmetrical with the last antennomere larger than the previous two combined.
Antennomeres 4-8 more or less compact, with 6-8 characteristically disc-like. Antennal scape
asymmetrical, somewhat hemispherical, subequal in length to pedicel. Pedicel cylindrical in
shape. Antennal segment 3 subequal in length to pedicel. Antennal club large, ~0.85 length of
segments 1-8 combined. Antennal grooves very deep and excavate, converging posteriorly.
Lateral mental-submental sulcus prominent, at lower level than submentum, medially the sulcus
is punctate with microreticulations and laterally with oblique to longitudinal microreticulations.
Mentum with anterior angles distinct, anterior margin broadly with medial acuminate apex,
pentagonal, entire structure flattened.
Pronotum widest in posterior third (L:W = 1:2.1), anterior margin broadly concave,
posterior margin slightly convex almost truncate, lateral margins somewhat arcuate anteriorly,
posterior angles nearly indistinct. Scutellum large, triangular, apex acuminate. Prosternal
process somewhat narrowed between procoxae, apex acuminate, in lateral aspect the posterior
third has a well-defined declivity, medially convex, and anteriorly with well-developed carina,
posterior apical wall reduced. Mesosternum small, extending to midway between mesocoxae,
evenly concave for reception of metasternum. Metasternum much wider than long (W:L ~ 3:1).
Metepisternum with slight medial constriction, oblique line dividing anterior 0.125 of structure.
First abdominal sternite with narrowed acuminate process between metacoxae. First sternite
~2X’s longer than second sternite. Sternites 2-4 subequal in length. Hypopygidium subequal in
length to first abdominal sternite.
Protibia with apical tooth prominent, slightly longer than tarsomeres 1. Outer apical
notch with ~110° angle, notch depth moderate, equal to length of tarsomere 1. Inner apical spine
short, 0.75 length to tarsomeres 1. Protibia not heavily armored but with characteristic dense
patch of stiff setae along the inner apical region. Mesotibia more heavily armored than protibia
85
with more dense stiff setae and a row of numerous slender spines along entire lateral edge. Outer
apical process not well-developed, subequal to protibial process. Inner apical spine equal in
length to tarsomeres 1-2 combined. Metatibia with armature similar to that of mesotibia, but
outer apical process well-developed, subequal to tarsomeres 1-2 combined.
Male genitalia well-sclerotized. Anal sclerite with large broadly curved region
anteriodorsally (Fig. 8); apex somewhat fimbriate; ventrally with a broad medial concavity
approaching apex in a broadly convex manner. Spiculum gastrale with wide narrowly curved
lateral flanges, medial margins slightly concave, long stiff setae originating from apex (Fig. 48).
Tegmen evenly rounded apically (Fig. 87), longer than wide (w:l = 1.0:2.31), lateral row of
setae visible from the median fossa to prior to the apex, large shallow concavity in apical third,
basal notch perpendicular, basal margin nearly straight. Median lobe robust, ~0.4 the length of
the tegmen, apex narrowly rounded, apical opening well-developed with inner structure complex
(Fig. 128). Ejaculatory rods not fused to basal piece or each other, overall elongate and
comprised of three distinct segments. Basal piece of internal sac sclerites with rounded with
apico-medial projection (Fig. 169).
Female genitalia not observed.
Variation. Known only from the holotype.
Seasonality/Habitat. Holotype collected in March.
Distribution. Known only from the type locality.
Notes. No fungal host is known for this species.
Etymology. Specific epithet denotes the greatly enlarged terminal antennomere with
distinct sensillar regions.
Pocadius ashei Cline new species
86
(Figs. 9, 88, 129, 170, 207)
Type Material Examined. HOLOTYPE ♀ (SNEC): BOLIVIA: LaPaz; 6km W
Yanacachi, 2150m; 16°25.95’S 67°47.07’W, 23-24-I-2001; J.S. Ashe, R.S. Hanley, ex. flight;
intercept trap, BOL1AH01 048 / SM0553606; KUNHM-ENT [barcode label]; HOLOTYPE;
Pocadius; ashei; A.R. Cline des. 2004. 3 PARATYPE ♀ (SNEC): same data label as holotype
but with paratype labels and the following barcode numbers: SM0553601, SM0553600,
SM0553614. 1 PARATYPE ♂ (SNEC): BOLIVIA: LaPaz; 9.4km E Chulumani, Apa Apa
Ecol.; Reserve, 2110m, 16°20.98’S; 67°30.28’W, 17-19-I-2001, J.S. Ashe; R.S. Hanley, ex.
flight intercept trap; BOL1AH01 022 / SM0553649; KUNHM-ENT / PARATYPE; Pocadius;
ashei; A.R. Cline des. 2004.
Diagnosis. This species is readily distinguished from other Neotropical Pocadius by the
reddish brown elongate pubescence covering the habitus, elongate pronotal and elytral fimbriae,
dorsal surface densely punctate with shining interspaces throughout, inner protibial patch of
short setae extending to more than midway proximad on the tibia, lateral setae on all tibia
elongate, erect and semi-erect rows of elytral setae, antennomeres 4-6 trapezoidal and subequal
in length, densely punctate scutellum, the extremely high W:L ratio of the pronotum, aedeagus
with tegmen having complete rows of inner and lateral elongate setae; ejaculatory rods fused to
each other in the apical 025; basal piece of internal sac sclerites consisting of paired sharply
curved pieces; ovipositor with deep intragonocoxal invagination, numerous terminal primary
setae, short baculi at gonocoxal base, and gonocoxae with lateral convexity.
Description. Length 4.2mm, Width 2.55mm, Depth 1.5mm. Body robust and convex,
surface shining, reddish brown to dark reddish brown in color, venter somewhat lighter in color
than dorsum. Pronotum and elytra margins with long fimbriae, setae longer than length of
87
antennal scape. Dorsal and ventral pubescence quite long and conspicuous on the appendages,
all pubescence reddish brown.
Head surface deeply, densely, irregularly punctate, large and small punctures interspersed
on vertex, interspaces for all punctures <0.25 large puncture diameter. Larger punctures 4 X
diameter of eye facet, smaller punctures ~2 X diameters, interspaces smooth and shining. Most
punctures give rise to a long seta, with those near the orbits longer. Pronotal surface with large
and small interspersed punctures, equal in size to large and small punctures on vertex;
interspaces smooth and shining, about 0.5-1.0 diameter apart. Each puncture gives rise to a long
seta, most seta are slightly curved. Scutellar surface with numerous moderately impressed small
punctures, punctures equal to smaller ones on pronotum, some punctures giving rise to setae,
interspaces smooth and shining. Elytral surface with serial rows of alternating large and small
deep punctures. Smaller punctures are equal in size to smaller ones on pronotum, larger
punctures are ~2 times diameter of smaller ones. Smaller punctures giving rise to an erect, long
seta, larger punctures giving rise to a semi-erect long seta. Interspaces narrow between
punctures of a given row and wide between different rows. Within a row, small punctures are
separated by ~0.5-1 puncture width, and large punctures by ~0.25 puncture width. Larger rows
are separated by ~2.0 large puncture diameters. Interspaces mostly smooth shining. Pygidium
densely punctate, punctures subequal in size to larger ones on pronotum, each puncture giving
rise to a short seta. Interspaces narrow, ~0.25 diameter, with smooth to somewhat alutaceous
sculpture and shining.
Venter with similar long pubescence as dorsum. Mentum with several shallowly
impressed punctures, equal in size to smaller ones on vertex, each giving rise to a seta;
interspaces smooth to alutaceous. Submentum and gula with less developed punctation as
mentum but with interspaces similar in sculpture. Prosternum and epimeron with moderately
88
impressed irregular punctures, punctures slightly larger in size to those on mentum, interspaces
alutaceous with microreticulate areas, prosternal punctures separated by 0.5 diameter, and those
on the epimeron by 0.5-1 diameter. Mesosternum with shallow punctures, subequal to those on
prosternum, interspaces smooth to alutaceous, separated by ~0.5 diameter, with most punctures
aggregated near metasternal border. Metasternum irregularly punctate with moderately
impressed punctures on disc similar in size to those on mesosternum, interspaces smooth and
shining on metasternal disc becoming more alutaceous laterally, punctures separated by ~1-2
diameters, lateral punctures much larger than those on disc ~2-3X’s diameter of those on
metasternal disc and more closely spaced. Abdominal sternite 1 with moderately impressed
punctures, punctures equal to those on metasternal disc, interspaces smooth to alutaceous and
shining, separated by ~1-2 diameters. Abdominal sternites 2-4 with two irregular rows of
punctures, one row near anterior margin and the other near posterior margin, punctures similar in
size to those on metasternal disc, rows separated by ~2 puncture diameters, punctures within
rows separated by ~0.5 punctures width, interspaces smooth to granular and shining. Punctures
between rows are irregularly organized and do not form definite rows. Hypopygidium with
moderately deep punctures, similar in size to those on sternites 2-4, interspaces mostly smooth to
alutaceous and shining, punctures separated by ~0.25 puncture diameters.
Head transverse, wider than long (W:L = 1.5:1), fronotclypeal region moderately
projecting anteriorly. Vertex with broad distinct concavity between orbits near fronotclypeal
region. Antennal club compact, somewhat circular to oblong, asymmetrical, with the last
antennomere twice as long as the previous two combined. Antennomeres 4-6 trapezoidal and
subequal in length, and 7-8 characteristically disc-like. Antennal scape asymmetrical,
hemispherical, 1.5 times as long as pedicel. Pedicel cylindrical in shape. Antennal segment 3
subequal in length to pedicel, and tapering proximally. Antennal club large, ~0.85 length of
89
segments 1-8 combined. Antennal grooves very deep and narrowly excavate, strongly
converging posteriorly. Lateral mental/submental sulcus and ridge prominent, at level of
submentum, ridge is sculptured with lateral areas of longitudinal microreticulation and the sulcus
is finely granulate. Mentum with anterior angles obsolete, anterior margin broadly hemispherical
with short central acute point, entire structure somewhat convex when viewed laterally.
Pronotum widest near middle (L:W = 1:2.07), anterior margin broadly deeply concave,
posterior margin moderately broadly convex, lateral margins less arcuate posteriorly. Scutellum
large, broadly triangular, apex broadly rounded. Prosternal process somewhat narrowed between
procoxae, apex acuminate, in lateral aspect there is only a moderate convexity over the procoxae.
Posterior apical wall prominent and slightly oblique. Mesosternum extending to midway
between mesocoxae, broadly shallowly concave for reception of the metasternum. Metasternum
width to length ratio is ~2.47:1.0. Metepisternum with medial constriction, oblique line dividing
anterior 0.16 of structure into metepisternal axillary space. Elytral humeri moderately produced,
lateral margin very narrow. First abdominal sternite with narrow process between metacoxae.
First sternite ~2X’s longer than second sternite. Sternites 2-4 subequal in length. Hypopygidium
subequal in length to first abdominal sternite.
Protibia with apical tooth moderately prominent, subequal in length to tarsomere 1.
Outer apical notch absent. Inner apical spine subequal in length to tarsomeres 1 and part of 2
combined. Protibia moderately armored with characteristic dense patch of stiff setae along the
inner apical region reaching midway to femoral articulation. Mesotibia more heavily armored
than protibia with more dense stiff setae and two rows of numerous slender spines along entire
lateral edge. Outer apical process moderately prominent, slightly longer and more robust than
protibial process. Inner apical spine equal in length to tarsomeres 1-2 combined. Metatibia with
armature similar to that of mesotibia, but apical spine more robust.
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Male genitalia well-sclerotized. Anal sclerite with elongate apical fimbriae; ventrally
with a narrow medial concavity approaching apex in a truncate manner (Fig. 9). Spiculum
gastrale not observed. Tegmen broadly rounded apically (Fig. 88), longer than wide (w:l =
1.0:2.4), lateral row of setae visible from the median fossa to around the apex, elongate
somewhat rectangular concavity in apical third, basal notch perpendicular, basal margin
obsoletely concave, inner row of setae complete. Median lobe elongate, 0.7 length of tegmen,
apex narrowly rounded, apical opening well-developed with double bilobed internal structure
(Fig. 129). Ejaculatory rods not fused to basal piece but fused to each other in apical 0.33, basal
portion perpendicular and apical portion curved outward. Basal piece distinctive with paired
crescent shaped portions laterally and medially with a bilobed globular portion that is less
sclerotized than the lateral crescent shaped portions (Fig. 170).
Female genitalia moderately sclerotized. Paraprocts large and broadly flanged with
sclerotization only along median line and not extending to baso-lateral angles. Gonocoxite with
one basal lateral prominence, basal ridge well-sclerotized with two short and sub-parallel baculi,
and lateral margin curved. Intragonocoxal invagination 0.5 length of total gonocoxite length.
Gonocoxal apices with recurved “tooth” absent. Three to four primary setae originate from
small depressions at the gonocoxal apices (Fig. 207).
Variation. None observed.
Seasonality/Habitat. All specimens were collected from mid to late January from
montane forests in Andean Bolivia.
Distribution. Known from the type locality near LaPaz, Bolivia.
Notes. No host data is available for this species.
Etymology. Specific epithet honors Steve Ashe, director of the Snow Entomological
Museum, for his generosity and understanding during the course of this study.
91
Pocadius barclayi Cline new species
(Figs. 10, 49, 89, 130, 171, 208)
Type Material Examined. HOLOTYPE ♂ (BMNH): INDONESIA:; SULAWESI
UTARA,; Dumoga-Bone N.P.; 25 February 1985. / Flight; interception; trap 1 / Plot A, ca 200m;
Lowland forest / [upside down] R.Ent.Soc.Lond.; PROJECT WALLACE; B.M. 1985-10 /
HOLOTYPE; Pocadius; barclayi; A.R. Cline des. 2004. 1 PARATYPE (BMNH): same data
labels as holotype but 24 February 1985, and also having a pink colored label with number 80.5.
1 PARATYPE (BMNH): same data labels as holotype but 26 February 1985. 1 PARATYPE
(BMNH): same data labels as holotype but April 1985. 1 PARATYPE (BMNH): INDONESIA:;
SULAWESI UTARA,; Dumoga-Bone N.P.; 9-16 May 1985 / Lowland forest; edge ca 200m /
malaise; trap / [upside down] R.Ent.Soc.Lond.; PROJECT WALLACE; B.M. 1985-10. 2
PARATYPES (BMNH): INDONESIA; SULAWESI UTARA; Gng. Ambang F.R.; nr.
Kotamobagu; Jan. 1985 / Lower montane; forest; 1200-1400m / in mature puffballs / [upside
down] R.Ent.Soc.Lond.; PROJECT WALLACE; B.M. 1985-10.
Diagnosis. This species is most similar to P. martini from the Philippines but differs
from it and the other Old World species by the following suite of characters: surface with mostly
granular sculpture; metasternum with small densely distributed punctures on disc becoming
greatly enlarged and more widely spaced laterally; terminal antennomere more than twice as
long as previous two segments combined; mesosternum distinctly carinate along midline;
mentum hemispherical; pronotum with anterior margin broadly slightly concave; scutellum
comparatively large; tarsomeres greatly thickened with relatively few setae; tegmen elongate
with inner median row of setae angulate to apex; median lobe elongate with complex apical
opening; internal sac with ejaculatory rods adjacent and large basal piece with small lunate
92
lateral LSAMessory pieces; and ovipositor with sharp medial gonocoxal incision and numerous
apical setae on gonocoxae.
Description. Length 3.4mm, Width 2.0mm, Depth 1.4mm. Body moderately convex,
surface somewhat shining, light brown to brown in color, with elytral apices darker or body
uniformly dark brown with legs lighter. Pronotum and elytra margins with short fimbriae, setae
subequal to length of antennal scape. Dorsal and ventral pubescence quite short and
inconspicuous.
Head surface shallowly, irregularly punctate, large and small punctures interspersed on
vertex. Larger punctures 3-4 X diameter of eye facet, smaller punctures ~2 X diameters,
interspaces alutaceous to granular. Each smaller puncture gives rise to a short decumbent golden
seta. Pronotal surface with large and small interspersed punctures, equal in size to large and
small punctures on vertex. Interspaces alutaceous to granular, about 0.5-1.0 diameter apart.
Each puncture gives rise to a short decumbent golden seta, most seta are slightly curved.
Scutellar surface with very few vague shallowly impressed punctures, some punctures giving rise
to setae, and the interspaces are granular. Elytral surface with serial rows of alternating large and
small deep punctures. Smaller punctures are equal in size to larger ones on pronotum, larger
punctures are ~1.5 times diameter of smaller ones. Smaller punctures giving rise to a semi-erect,
oblique short golden seta, larger punctures giving rise to a decumbent short golden seta.
Interspaces narrow between punctures of a given row and between different rows. Within a row,
small punctures are separated by ~0.5 puncture width, and large punctures by ~0.25 puncture
width. Larger rows are separated by ~1.0 large puncture diameters. Interspaces moderately
shining being mostly granular in sculpture. Pygidium densely punctate, punctures equal in size
to larger ones on pronotum, each puncture giving rise to a short stiff golden seta. Interspaces
narrow, ~0.5 diameter, with alutaceous to mostly granular sculpture.
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Venter with similar short sparse golden pubescence as dorsum. Mentum with several
moderately impressed punctures, equal in size to larger ones on vertex, each giving rise to a short
seta. Interspaces alutaceous to granular with some microreticulate areas. Submentum and gula
with less developed punctation as mentum but with interspaces similar in sculpture. Prosternum
and epimeron with moderately impressed irregular punctures, punctures slightly equal in size to
those on mentum, interspaces granular with microreticulate areas, prosternal punctures separated
by 0.25-0.5 diameter, and those on the epimeron by 0.25 to 0.5 diameter. Mesosternum with
shallow punctures, subequal to those on prosternum, interspaces alutaceous to granular,
separated by about 1 diameter, with most punctures aggregated near metasternal border.
Metasternum irregularly punctate, with moderately impressed punctures on disc similar in size to
those on mesosternum, interspaces granular on metasternal disc and laterally, punctures
separated by ~1 diameters, lateral punctures 3-3.5 X’s diameter of those on metasternal disc and
more widely separated. Abdominal sternite 1 with faint, almost obsolete punctures, punctures
equal to those on metasternal disc, interspaces alutaceous to granular, separated by ~1-2
diameters. Abdominal sternites 2-4 with two irregular rows of punctures, one row near anterior
margin and the other near posterior margin, punctures similar in size to those on metasternal
disc, rows separated by ~2-3 puncture diameters, punctures within rows separated by ~0.25-0.5
punctures width. Punctures between the rows are irregularly organized and not forming definite
rows. Hypopygidium with moderately deep punctures, similar in size to those on sternites 2-4,
interspaces mostly granular, punctures separated by ~1 puncture diameters.
Head transverse, wider than long (W:L = 1.5:1), fronotclypeal region moderately
projecting anteriorly. Vertex with broad distinct concavity between orbits near fronotclypeal
region. Eyes large and protruding. Antennal club compact, somewhat circular to oblong,
asymmetrical, with the last antennomere twice as long as the previous two combined.
94
Antennomeres 4-8 more or less compact, with 4-5 somewhat trapezoidal, and 6-8
characteristically disc-like. Antennal scape asymmetrical, greatly hemispherical, 1.8 times as
long as pedicel. Pedicel cylindrical in shape. Antennal segment 3 equal in length to pedicel.
Antennal club large, ~0.65 length of segments 1-8 combined. Each club segment with dense
short setae, and only relatively few protruding setae. Antennal grooves very deep and narrowly
excavate, strongly converging posteriorly. Lateral mental ridge prominent, at level of
submentum, ridge is sculptured with small closely packed minute punctures and some lateral
areas of microreticulation. Mentum with anterior angles obsolete, anterior margin broadly
hemispherical, entire structure somewhat convex when viewed laterally.
Pronotum widest near middle (L:W = 1:1.83), anterior margin broadly shallowly
concave, posterior margin moderately broadly convex, lateral margins less arcuate posteriorly.
Scutellum large, oblong and broadly hemispherical, apex broadly rounded. Prosternal process
somewhat narrowed between procoxae, apex acuminate, in lateral aspect the anterior half slopes
steeply dorsad, and the posterior half flattens into a short shelf, moderate convexity over
procoxae. Posterior apical wall prominent and straight to slightly oblique. Mesosternum
extending to midway between mesocoxae, deeply evenly concave for reception of the
metasternum, a distinct longitudinal median carina is present along entire length. Metasternum
width to length ratio is ~2.25:1.0. Metepisternum with medial constriction, oblique line dividing
anterior 0.20 of structure. Elytral humeri moderately produced, lateral margin very narrow. First
abdominal sternite with broad process between metacoxae. First sternite ~2X’s longer than
second sternite. Sternites 2-4 subequal in length. Hypopygidium subequal in length to first
abdominal sternite.
Protibia with apical tooth prominent, slightly longer than tarsomere 1. Outer apical notch
with ~100° angle, notch depth shallow, equal to length of tarsomere 1. Inner apical spine
95
subequal in length to tarsomeres 1-2 combined. Protibia moderately armored with characteristic
dense patch of stiff setae along the inner apical region. Mesotibia more heavily armored than
protibia with more dense stiff setae and two rows of numerous slender spines along entire lateral
edge. Outer apical process not prominent, subequal to protibial process. Inner apical spine equal
in length to tarsomeres 1-2 and part of 3 combined. Metatibia with armature similar to that of
mesotibia, but apical spine more elongate and robust.
Male genitalia well-sclerotized. Anal sclerite with apex forming a distinct pointed lobe;
apex fimbriate; ventrally with a narrow medial concavity approaching apex (Fig. 10). Spiculum
gastrale with broad lateral flanges, medial margins parallel, elongate setae originating from apex
(Fig. 49). Tegmen narrowly rounded apically (Fig. 89), much longer than wide (w:l = 1.0:3.85),
lateral row of setae visible from the median fossa to prior to the apex, elongate oval concavity in
apical third, basal notch perpendicular, basal margin concave, median fossa elongate. Median
lobe elongate, equal in length to the tegmen, apex acuminate, apical opening well-developed
with distinct proximal concavity (Fig. 130). Ejaculatory rods not fused to basal piece, adjacent
with distal region slightly curved outward. Basal piece distinctive with small lateral projections
and basal lateral LSAMessory pieces that are lunate in shape (Fig. 171).
Female genitalia moderately sclerotized. Paraprocts large with sclerotization only along
median line and not extending to baso-lateral angles. Gonocoxite with one basal lateral
prominence, basal ridge well-sclerotized with two elongate oblique baculi. Gonocoxal apices
with recurved “tooth” absent. Four to five setae originate from small depressions on the
gonocoxal apices (Fig. 208).
Variation. The specimen collected in March is much smaller than the other specimens.
The two specimens collected from Ambang F.R. are completely dark brown and do not exhibit
the darker elytral apices as in the other specimens.
96
Seasonality/Habitat. Specimens were collected in lowland wet forests from January to
mid-May.
Distribution. Known from the Utara (northern) region of Sulawesi (a.k.a. Celebes).
Notes. The genitalia of the holotype are contained in a glycerin vial beneath the
specimen, which is card-mounted. Host data is listed as “mature puffballs” for the Ambang F.R.
specimens.
Etymology. Specific epithet honors Maxwell Barclay, curator at the Natural History
Museum London, for his efforts and generosity during the course of this study.
Pocadius basalis Schaeffer
(Figs. 11, 50, 90, 131, 172, 209)
Type Material Examined. HOLOTYPE (USNM) ♂: Huach Mts.; VIII-29, Ariz. /
Brooklyn; Museum; Coll. 1929 / Type No.; 42564; U.S.N.M. / Pocadius; basalis; Type, Schffr.
Specimen in average condition but lacking both antennal clubs, and right metatarsi.
PARATYPES (USNM) ♀, 3 specimens with same label data as Holotype. PARATYPES
(USNM), 2 specimens with the following labels: Douglas, Ariz.; 20-VIII-1960; J. H. Russell /
ex. puffball; 61-24953 / Pocadius; basalis; Schaeffer.
Non-Type Material Examined. >200 specimens total: (5) from LSAM: Hck. Hwy.
Mi.24, Sta.; Catalina Mts., ARIZ.; VIII-8-1959, F.G. Werner; in hole in puffball / ♀. (5) from
LSAM: ARIZONA Pajarito; Mts. Pena Blanca; Cyn. 15 Aug 1970; K. Stephan coll. (1) from
LSAM: ARIZONA: Pajarito; Mts. Pena Blanca Cyn.; Aug. 15 1971; K. Stephan leg. (4) from
LSAM: U.S.A. ARIZ; Santa Cruz Co.; Pena Blanca; Pajarito Mts. / Lot No. 521; July. 28, 1961;
R.H. Arnett Jr.; E. VanTassell. (4) Tex. Jeff Davis Co.; Davis Mts. Resort; 25 June 2000; R.
Turnbow. (2) AZ: Cochise Co., Rustler Park Camp; 20-21 Aug 1981; J. Liebherr; El. 8000’. (8)
97
from UAC: AZ: Fajarito Mts.; Sycamore Cn. At Tank Springs; 6 Aug. 1978; F.G. Nerner; in
puffball. (8) from UAC: Peña Blanca, 10mi. W. Nogales, ARIZ.; Aug.1, 1961; Werner, Nutting.
(3) from UAC: Ruby, Ariz.; Aug.17, 1959; M.W. Larson. (1) from UAC: 4mi. N of Sonita,
Ariz.; VIII-12-1947; in puffball; L.R. Gillogly collr. (28) from UAC: Casa Blanca Cn.; 7mi. NE
Patagonia, Ariz.; 5 Aug. 1978; F. Werner. (128) from CAS: Huachuca Mts., Carr Cn., Ariz.;
VIII (22-24) 1961; fungi; L.R. Gillogly collr. (5) from CAS: Sta. Catalina Mts., Bear Wallow,
Ariz.; VIII-20-1961. (7) from CAS: Skeleton Cyn., Hidalgo Co., New Mexico; VIII-13-1965; G.
Forister.
Diagnosis. This species is most similar to P. niger known from New Mexico and the
Pinal and Sierra Ancha Mts. in Arizona. These species are putative sister taxa and the variation
in color in P. basalis approaches that of P. niger, but the head and pronotum are never black.
Pocadius basalis has a well-developed apical notch on the inner margin of the profemora that P.
niger does not. Clear differences can be seen in both the male and female genitalia, specifically
the median lobe of P. basalis has an acuminate tip and shallow apical opening, the tegmen
contains setae that extend around the apex and also on the lateral margins of the medial fossa,
whereas P. niger does not. Females possess 3 terminal gonocoxal setae in a small shallow
depression, whereas in P. niger there are 2 terminal setae in an oblique groove.
Redescription. Length 4.1mm, Width 2.7mm, Depth 2.1mm. Body moderately convex,
surface shining, reddish-brown to dark brown in color, sometimes with elytral humeri lighter.
Pronotum and elytra margins with elongate fimbriae, setae longer than width of antennal scape.
Dorsal and ventral pubescence quite long.
Head surface deeply, irregularly punctate, punctures larger on vertex, becoming
somewhat smaller towards orbits and fronotclypeal region. Larger punctures 6-7 X diameter of
eye facet, smaller punctures 3-4 X diameter. Interspaces smooth to finely alutaceous, shining.
98
Each puncture gives rise to an elongate curved golden seta. Labrum distinctly granular on entire
dorsal surface. Pronotal surface with large punctures equal in size to large punctures on vertex
of head, interspersed with relatively few smaller punctures, ~0.75 size of larger ones.
Interspaces alutaceous to finely microreticulate, about 1-1.5 diameters apart. Each puncture
gives rise to a long golden seta, most seta are curved but some are rather straight. Scutellar
surface with very few vague shallowly impressed punctures, some punctures giving rise to setae,
and the interspaces are granular. Elytral surface with serial rows of alternating large and small
deep punctures. Smaller punctures are equal in size to those on pronotum, larger punctures are
~1.5 times diameter of smaller ones. Smaller punctures giving rise to a semi-erect long golden
seta, larger punctures giving rise to a semi-erect long golden seta. Interspaces narrow between
punctures of a given row and between different rows. Within a row, small punctures are
separated by ~1 puncture width, and large punctures by 0.3-0.8 puncture width. Rows are
separated by 0.5-1.0 puncture diameters. Interspaces always shining but variable from smooth to
finely microreticulate in sculpture. Pygidium densely punctate, punctures equal in size to those
on pronotum, each puncture giving rise to a moderately long golden seta. Interspaces narrow,
0.7-1.2 diameters, with granular sculpture.
Venter with similar long golden pubescence as dorsum. Mentum with large very shallow
punctures, equal in size to those on vertex, each giving rise to an elongate seta. Interspaces
granular to finely microreticulate. Submentum and gula similar in punctation to mentum but
with interspaces completely granular. Prosternum and epimeron deeply irregularly punctate,
punctures slightly larger than those on mentum, interspaces alutaceous with microreticulate
areas, prosternal punctures separated by 0.5 diameter, those on the epimeron by 0.25 to 0.5
diameter. Mesosternum with shallow punctures, about 0.75 diameter of those on prosternum,
interspaces alutaceous to smooth, separated by about 0.5 to 1 diameter. Metasternum irregularly
99
punctate, with moderate faint punctures on disc similar in size to those on mesosternum,
interspaces smooth on metasternal disc becoming microreticulate to granular laterally, punctures
separated by ~1-2 diameters. Abdominal sternite 1 with large faint, almost obsolete punctures,
punctures equal to large punctures on elytra, interspaces smooth to alutaceous, separated by ~1
diameter. Abdominal sternites 2-4 with two irregular rows of punctures, one row near anterior
margin and the other near posterior margin, punctures similar in size to those on metasternum,
rows separated by 1.5-2 puncture diameters, punctures within rows separated by ~0.5 punctures
width. Rows on abdominal sternite 4 becoming less organized. Hypopygidium with moderately
deep punctures, similar in size to those on sternites 2-4, interspaces alutaceous to granular,
punctures separated by 1-2 puncture diameters.
Head wider than long (W:L = 1.55:1), fronotclypeal region moderately projecting
anteriorly. Vertex with shallow concavity between orbits near fronotclypeal region. Labrum
with shallow concavity at anterior margin. Antennal club compact, oval, only slightly
asymmetrical with the last antennomere subequal to the previous two combined. Antennomeres
5-8 more or less compact, with 6-8 characteristically disc-like. Antennal scape asymmetrical,
somewhat hemispherical, 1.7 times as long as pedicel. Pedicel subcylindrical in shape. Antennal
segment 3 subequal in length to pedicel. Antennal club large, ~0.75 length of segments 1-8
combined. Antennal grooves very deep and widely excavate, slightly converging posteriorly.
Lateral mental/submental ridge prominent, at level of submentum, ridge is divided longitudinally
by rounded crest, medially the ridge is transversely sculptured with microreticulations and
laterally with oblique to longitudinal microreticulations. Mentum with anterior angles obsolete,
anterior margin angulate, triangular, entire structure somewhat convex.
Pronotum widest near middle (L:W = 1:1.9), anterior margin broadly trapezoidal,
posterior margin moderately convex, lateral margins less arcuate posteriorly. Scutellum large,
100
obtusely triangular, apex rounded. Prosternal process somewhat narrowed between procoxae,
apex somewhat acuminate, in lateral aspect the anterior and posterior ends are prominent and
convex medially. Posterior apical wall prominent and only slightly oblique. Mesosternum
extending to midway between mesocoxae, evenly concave for reception of the metasternum.
Metasternum width to length ratio is ~2.6:1.0. Metepisternum with slight medial constriction,
oblique line dividing anterior 0.2 of structure. Elytral humeri moderately produced, lateral
margin very narrow. First abdominal sternite with acuminate process between metacoxae. First
sternite ~2X’s longer than second sternite. Sternites 2-3 subequal in length, the fourth slightly
larger than the preceding two. Hypopygidium subequal in length to first abdominal sternite.
Protibia with apical tooth very prominent, slightly longer than tarsomeres 1-2 combined.
Outer apical notch with ~105° angle, notch depth moderate, equal to length of tarsomere 1.
Inner apical spine subequal in length to tarsomeres 1-2 combined. Protibia heavily armored with
characteristic dense patch of stiff setae along the inner apical region. Mesotibia more heavily
armored than protibia with more dense stiff setae and a row of numerous slender spines along
entire lateral edge. Outer apical process elongate and robust, larger than protibia process, and
bifid apically. Inner apical spine equal in length to tarsomeres 1-2 combined. Metatibia with
armature similar to that of mesotibia.
Male genitalia well-sclerotized. Anal sclerite with large broadly curved region
anteriodorsally (Fig. 11); apex somewhat fimbriate; ventrally with a broad medial concavity
approaching apex. Spiculum gastrale with wide lateral flanges, medial margins concave
proximally, short stiff setae originating from apex (Fig. 50). Tegmen evenly rounded apically
(Fig. 90), much longer than wide (w:l = 1.0:2.55), lateral row of setae visible from the median
fossa to prior to the apex, small shallow concavity in apical third, basal notch perpendicular,
basal margin nearly straight, apex of median fossa border with small patch of fine setae. Median
101
lobe large and robust, ~0.66 the length of the tegmen, apex acuminate, apical opening well-
developed with proximal concavity (Fig. 131). Ejaculatory rods not fused to basal piece, curved
inward and expanded outward at basal piece. Basal piece of rods with deep medial concavity
extending almost half length of structure, proximally with four sharp projections (Fig. 172).
Female genitalia moderately sclerotized. Paraprocts large with sclerotization only along
median line to baso-lateral angles. Gonocoxite with two basal lateral prominences, basal ridge
well-sclerotized. Gonocoxal apices with recurved “tooth” present. Three setae originate from
small depressions on the gonocoxal apices (Fig. 209).
Variation. Color varies considerably among members of this species. Most individuals
are brown to dark reddish brown, some individuals have slightly lighter humeri. Some variation
exists in the armature of the mesotibial, some individuals exhibit 1-2 fewer lateral spines than
others, but overall armature is still quite impressive. Some variation in the length and degree of
setation on the male anal sclerite was observed, with a few males having more longer setae than
illustrated.
Seasonality/Habitat. Adults are known to occur from June to August, with the only
known host record as “puffballs”. This species can be considered an alpine taxon with
individuals collected up to 8000’ in elevation.
Distribution. Known from southern Arizona, New Mexico, and western Texas (see
Cline 2003a).
Notes. This species likely occurs in northern Mexico and future collecting will
undoubtedly reveal records for its occurrence there. Pocadius basalis exhibits a peculiar
sympatry with both P. helvolus and P. niger both of which occur within its known range.
Pocadius bicolor Cline n. sp.
102
(Fig. 210)
Type Material Examined. HOLOTYPE ♀ (CMN): BRAZIL: Para; Carajas, Serra;
Norte, III-1985; 6 15’S 50 25’W; N. Degallier; FIT, carrion, dung.
Diagnosis. The bicolored appearance of this species is unlike any other in the genus,
with the pronotum distinctly lighter than the rest of the body.
Redescription. Length 2.7mm, Width 2.1mm, Depth 1.35mm. Body moderately
convex, surface shining, pronotum and legs lighter than rest of body, pubescence light colored
mostly golden, pronotal and elytral fimbriae not exceptionally elongate (~0.5 as long as
scutellum) and sparsely distributed along margins. Pygidium and hypopygidium densely deeply
punctate.
Head surface densely irregularly punctate with small and large punctures, some larger
punctures with their surface rugose, interspaces smooth to granular and ~0.5 diameter apart on
disc. Larger punctures equal to ~4 eye facet diameters, smaller punctures scattered on vertexal
disc and ~0.5 diameter of large puncture. Pronotal surface with large and small punctures
interspersed throughout. Large and small punctures similar in size to corresponding large and
small punctures on head. Interspaces 1-2 puncture diameters apart, alutaceous to granular. Each
smaller puncture bearing a semierect seta. Scutellar surface faintly punctate, most punctures
similar in size to the smaller punctures on the pronotum, interspaces granular to finely
microreticulate and separated by ~1-1.5 diameter of puncture width. Eytral surface with serial
rows of alternating large and small shallow punctures. Large punctures ~1.5X diameter of larger
punctures on head, each giving rise to a decumbent seta. Small punctures ~0.5 diameter of large
punctures, each giving rise to an erect seta. Interspaces alutaceous to finely microreticulate
between all punctures, larger punctures separated from each other by 1 puncture diameter,
smaller punctures separated from each other by 1-2 puncture diameters. Large and small
103
puncture rows separated from each other by ~1 small puncture diameters. Pygidium densely
irregularly punctate, subequal in size to small punctures on elytra, interspaces rugose with some
microreticulation, punctures separated by ~0.25-0.5 puncture diameters. Each puncture giving
rise to a short straight seta.
Venter more scarcely pubescent than dorsum with the exception of that on the abdominal
segments, which is similar to that of the dorsum. Mentum with only a few faintly impressed
minute punctures, punctures subequal in size to the small punctures on the vertex, interspaces
smooth to faintly alutaceous, punctures ~1 diameter apart. Submentum and gula with similar
sized punctures as mentum, each giving rise to a short seta, interspaces 1-2 diameters apart with
surface alutaceous and having transverse microreticulations. Mental/submental sulcus with
oblique microreticulations present. Prosternum and epimeron deeply impressed with large
punctures, punctures similar in size to larger ones on pronotum, interspaces alutaceous with
slight microreticulation present, ~0.25-0.5 diameter apart. Mesosternum with punctures
dispersed throughout, ~0.5-.75 those of prosternum, interspaces alutaceous to rugose.
Metasternal disc faintly punctate with smaller punctures 0.5 diameter to those on prosternum,
interspaces smooth to alutaceous, becoming more rugose laterally. Abdominal sternite 1 with
moderately impressed punctures, 1.5-2 diameters of those on metasternal disc, interspaces
alutaceous to rugose, separated by ~1 puncture diameter, abdominal process with more
reticulation present. Abdominal segments 2-4 with punctures scattered in an irregular transverse
row near the anterior border, similar in size to those on abdominal sternite 1, interspaces mostly
alutaceous to rugose, 0.5 - 1 diameter apart. Hypopygidium more densely deeply punctate than
rest of abdominal sternites, punctures equal in size to those on metasternal disc, interspaces
rugose with faint microreticulation, punctures separated by ~0.5 diameter.
104
Head wider than long (W:L = 1.5:1), fronotclypeal region moderately pronounced.
Vertex with deep broad concavity extending between each orbit from the posterior region of the
frons to the clypeal region. Eyes very large, bulbous and convex, finely faceted. Labrum
transverse with sharp medial incision on anterior margin. Mentum pentagonal in shape and
somewhat convex. Antennal club compact, large, ovate, asymmetrical with the last antennomere
longer than previous two segments combined, entire club subequal in length to antennal stem.
Terminal antennomere with distinct sensillar region extending across entire apex on ventral
surface in a U-shaped manner. Antennomeres 10 more narrow than 9. Antennomeres 6-8
flattened, segment 6 less flattened than 7 and 8, their combined length slightly longer than length
of antennomere 9. Antennal scape asymmetrical, broadly hemispherical, 1.2X’s longer than
pedicel, >12 setae originating from it. Pedicel more cylindrical in shape than scape and thinner.
Antennal segment 3 subequal in length and similar in shape to pedicel, narrowed proximally.
Segment 4 cuboidal and 5 more or less trapezoidal. Antennal grooves deep and somewhat curved
posteriomedially. Prosternal process narrowed slightly between coxae, evenly rounded over
procoxae with flattened declivity posterior to procoxae, apical border oblique when viewed
laterally. Mesosternum extending to anterior 0.33 of mesocoxae with a broadly concave apex,
with no visible carina present. Mesepisternum slightly wider than mesosternum, ~2 times wider
than mesepimeron. Metasternum width to length ratio 3.33:1. Metepisternum rather broad and
well-developed, moderately concave medially, axillary space impunctate ~0.05 length of entire
structure. First abdominal sternite with narrowly rounded process ending in a small point
between metacoxae, ~2X’s longer than second sternite. Sternites 2-4 subequal in length.
Hypopygidium subequal in length to first abdominal sternite.
Protibia finely crenulate along entire lateral edge. Apical tooth as long as tarsomere 1-2
combined. Outer apical notch with 85° angle. Longer inner apical spine subequal in length to
105
tarsomeres 1-2 combined. Apical border of tibia smooth, no armature present. Mesotibia
armored with a row slender spines along entire lateral edge, spines ~0.75 length of lateral setae
on mesotibia; apical border armored with 2-3 short spines, ~0.5 length of lateral spine; outer
apical process robust, subequal length of inner apical spine; inner apical spine equal in length to
tarsomeres 1-2.5 combined. Metatibia more heavily armored than mesotibia, lateral slender
spines more elongate and numerous. Spines of varying lengths, but most longer than those on
the mesotibia. Apical border armored with a few short spines, as in mesotibia, but more
apparent. Outer apical process similar in length and robustness as that on mesotibia, but shorter
in length to inner apical spine. Inner apical spine equal in length to tarsomeres 1-2 combined.
All tarsomeres thin and elongate.
Male genitalia not observed.
Female genitalia overall moderately sclerotized. Gonocoxite with well sclerotized basal
border with two lateral prominences. Gonocoxal apices narrowly separated with intragonocoxal
invagination narrowly acute basally, gonocoxal tips broadly rounded at apex, each apex without
a small lateral recurved “tooth”, apical pits giving rise to 2 primary seta (Fig. 210). Valvifers
membranous, relatively long compared to gonocoxite, with sclerotization along medial border,
evenly tapering to lateral apex.
Variation. Known only from the holotype.
Seasonality/Habitat. Holotype collected in March in equatorial Brazil.
Distribution. Known only from the type locality in north central Brazil.
Notes. No fungal host data is available for this species.
Etymology. Specific epithet denotes the distinctive bicolored habitus.
Pocadius brevis Reitter
106
(Figs. 12, 51, 91, 132, 173, 211)
Non-Type Material Examined. 27 specimens with the following label data: Baños de
Ciego, Montero, Santa Clara, Cuba, 14-IV-1918 / beetles collected from inside puffball (20
specimens USNM, 7 specimens LSAM). Cayamas, Cuba (USNM). CUBA: Pinar del Rio,
Sierra del Rosario, ca. 15km S. Cinco Pesos Range 1, 29 June 1990, 420m, M.A. Ivie colr.
(MTEC).
Diagnosis. This species appears related to the Eastern North American species P.
helvolus. Pocadius brevis, unlike P. helvolus, is always light brown to light brown-red with
antennal segments 3-11 darker. The mentum is much wider than long, with no prominent
anterio-lateral angles. Clear differences are also visible in the male genitalia, i.e. the median
lobe is markedly narrowed apically, the tegmen has an incomplete inner line of setae, the
ejaculatory rods are more slender and nearly approximate basally, the spiculum gastrale is
greatly expanded laterally with numerous projecting setae apically.
Redescription. Length 3.9mm, Width 2.2mm, Depth 1.5mm. Body modestly convex,
surface shining, light brownish red with antennal segments 3-9 darker, pubescence light colored
mostly golden, pronotal and elytral fimbriae not exceptionally elongate and sparsely distributed
along margins.
Head surface densely irregularly punctate with small and large punctures, some larger
punctures with their surface microreticulate, interspaces smooth to alutaceous and ~0.25-0.5
diameter apart. Larger punctures equal to ~4-5 eye facet diameters, smaller punctures scattered
on vertex and ~0.5 diameter of large puncture. Pronotal surface with large and small punctures
interspersed throughout. Large and small punctures similar in size to corresponding large and
small punctures on head. Interspaces 0.25-0.5 puncture diameter apart, smooth to alutaceous.
Each puncture bearing an erect or semierect seta. Scutellar surface faintly punctate, punctures
107
similar in size to the smaller punctures on the pronotum, interspaces rugose and separated by
~0.5 diameter of puncture width. Eytral surface with serial rows of alternating large and small
shallow punctures. Large punctures ~1.5X diameter of larger punctures on head, each giving rise
to a decumbent seta. Small punctures ~0.25-0.33 diameter of large punctures, each giving rise to
a semierect seta. Interspaces smooth to alutaceous between all punctures, larger punctures
separated from each other by 0.5 puncture diameter, smaller punctures separated from each other
by 1-2 puncture diameters. Large rows separated from each other by ~2-2.5 large puncture
diameters. Pygidium densely irregularly punctate, punctures equal in size to small punctures on
elytra, interspaces rugose with some microreticulation, punctures separated by ~0.5 puncture
diameters. Each puncture giving rise to a short straight seta, setae similar in length of those on
elytra.
Venter with more scarce somewhat shorter pubescence (~0.5 - 0.75 length of dorsal
vestiture) than dorsum with the exception of that on the abdominal segment which is subequal to
that of the dorsum. Mentum with only a few punctures near anterior margin, punctures similar in
size to the small punctures on the pronotum, interspaces smooth to alutaceous, punctures 0.5 - 1
diameter apart. Submentum and gula with similar sized punctures as mentum, each giving rise to
a short seta, interspaces 1-2 diameters apart and surface alutaceous. Prosternum and epimeron
faintly impressed with moderate sized punctures, punctures ~2 times larger than those on
mentum, interspaces alutaceous with slight microreticulation present, ~1 diameter apart.
Mesosternum with punctures mostly dispersed but with distinct aggregation near posterior
margin with metasternum, interspaces and anterior portion of mesosternum alutaceous to
rugulose. Metasternal disc moderately punctate with moderate sized punctures equal to those on
prosternum, interspaces alutaceous to rugose. Abdominal sternite 1 with faint punctures, equal
in size to those on metasternum, interspaces alutaceous to rugose, separated by ~1-2 puncture
108
diameters, most punctures located in anterior 0.5 of sternite. Abdominal segments 2-4 with
punctures scattered in an irregular lateral row near the anterior border, similar in size to those on
abdominal sternite 1, interspaces mostly rugose, 0.5 - 1 diameter apart. Hypopygidium more
densely deeply punctate than rest of abdominal sternites, punctures equal in size to those on
metasternum, interspaces rugose with faint microreticulation, punctures separated by 0.25-0.5
diameter.
Head wider than long (W:L = 1.5:1), fronotclypeal region well pronounced. Vertex with
deep broad concavity extending between each orbit from the posterior region of the frons to the
clypeal region. Labrum transverse with medial incision on anterior margin. Mentum
hemispherical in shape and slightly convex. Antennal club compact, large, ovate, slightly
asymmetrical with the last antennomere subequal to previous two segments combined, entire
club equal in length to 0.85 length of antennal stem. Terminal antennomere with distinct
hemispherical sensillar region midway to apex on ventral surface. Antennomeres 9-10 similar in
shape. Antennomeres 6-8 flattened, segment 6 more trapezoidal than 7 and 8, their combined
length slightly longer than length of antennomere 9. Antennal scape asymmetrical, broadly
hemispherical, only slightly longer than pedicel, >12 setae originating from it. Pedicel more or
less similar in shape as scape but thinner. Antennal segment 3 ~0.75 length of pedicel, narrowed
proximally. Segments 4 and 5 more or less globular, together ~0.75 the length of segment 3.
Antennal grooves deep and somewhat curved posteriomedially.
Pronotum widest near middle (L:W = 1:1.9). Anterior margin deeply trapezoidal,
posterior margin moderately convex, lateral margin more arcuate anteriorly than posteriorly.
Prosternal process narrowed slightly between coxae, evenly rounded with a moderately acute
apex. Posterior margin well-developed and perpendicular. Mesosternum extending to anterior
0.33 of mesocoxae with an indentate apex, with no visible carina present. Mesepisternum equal
109
slightly wider than mesosternum, ~2.4 times wider than mesepimeron. Metasternum width to
length ratio 2.5:1. Metepisternum rather broad and well-developed, width to length 1:3,
moderately concave medially, anterior third strongly produced anteriomedially with axillary
space (0.2 of metepisternum) impunctate with surface granular with microreticulation.
Metasternal axillary space minute, extending medially closely to the mesocoxal border.
Metacoxal lines not deviating much from metacoxal border. Median metasternal line
incomplete, extending only 0.75 length of structure, rounded abdominal sternite line encircling
junction between metasternum and first abdominal sternite. First abdominal sternite with
broadly rounded process ending in a small point between metacoxae, ~2.5X’s longer than second
sternite. Sternites 2-4 subequal in length. Hypopygidium subequal in length to first abdominal
sternite. Mentum with
Protibia finely crenulate along lateral edge. Apical tooth as long as tarsomere 1 and half
of tarsomere two. Outer apical notch with no distinct angle or notch but oriented in a lateral
manner. Longer inner apical spine subequal in length to first tarsomere. Apical border of tibia
smooth, no armature present. Mesotibia armored with a row slender spines along entire lateral
edge, spines ~0.5 length of lateral setae on mesotibia. Apical border armored with 2-3 short
spines, ~0.5 length of lateral spines. Outer apical process not robust, ~0.5 length of inner apical
spine. Inner apical spine equal in length to tarsomeres 1-1.5 combined. Metatibia more heavily
armored than mesotibia, lateral slender spines more elongate and numerous. Spines of varying
lengths, but most longer than those on the mesotibia. Apical border armored with a few short
spines, as in mesotibia, but more apparent. Outer apical process more elongate and robust than
on mesotibia, subequal in length to inner apical spine. Inner apical spine equal in length to
tarsomeres 1-2 combined. All tarsomeres elongate, protarsomeres combined greater than 0.5
length of protibia.
110
Male genitalia well sclerotized. Anal sclerite with fossa anteriodorsally for reception of
the tegmen with apical border nearly truncate (Fig. 12), apex of sclerite moderately fimbriate
with elongate setae. Spiculum gastrale with well-pronounced medial regions (Fig. 51), lateral
region hemispherical and moderately explanate, apical border rounded with >10 setae, spiculum
attached posterio-medially to sclerites. Tegmen evenly broadly rounded apically (Fig. 91),
longer than wide (w:l ~ 1:2.2), lateral row of setae visible anterior to the median fossa but not
attaining apex, inner row of hair present but not connecting apically, basal notch of phallobase
absent, basal margin angulate. Median lobe moderately large, ~0.5 the length of the tegmen,
entire structure becoming narrowed apically, apical opening not well-developed (Fig. 132).
Ejaculatory rods not fused, proximal pair more or less straight, apical pair with distinct medial
concavity (Fig. 173).
Female genitalia overall moderately sclerotized. Gonocoxites with sclerotized basal
border with two lateral prominences. Gonocoxal apices moderately separated with
intragonocoxal invagination narrowly rounded at base, gonocoxal tips rounded at apex, each
apex without a small lateral recurved “tooth”, apical pits giving rise to 2short seta (Fig. 211).
Valvifers membranous, relatively small compared to gonocoxites, with some sclerotization along
medial border, evenly tapering to lateral apex.
Variation. No demonstrable differences between geographic regions or between males
and females.
Seasonality/Habitat. Specimens studied were collected between April and June.
Distribution. Found throughout most of Cuba.
Notes. The only known host for this species comes from the generic data label “inside
puffball”.
111
Pocadius carltoni Cline new species
(Figs. 13, 52, 92, 133, 174, 212)
Type Material Examined. HOLOTYPE ♂ (SNEC): NICARAGUA : Rio San Juan
Dept.; 60km SE San Carlos, Refugio; Bartola 100m, 10°58.40’N 84°20.30’W; 25-V-2002, R.
Brooks, Z. Falin; S. Chatzimanolis, ex. pyrethrum; fogging fungusy logs, NIC1BFC02 065 /
SM0531370; KUNHM-ENT [barcode label]. 5 PARATYPES (3 SNEC, 2 LSAM): same data
labels as holotype except barcode numbers are SM0531374, SM0531376, SM0531375, and
SM0531368 . 4 PARATYPES (SNEC) NICARAGUA : Rio San Juan Dept.; 60km SE San
Carlos, Refugio; Bartola 100m, 10°58.40’N 84°20.30’W; 29-V-2002, R. Brooks, Z. Falin; S.
Chatzimanolis, ex. puffball fungus; NIC1BFC02 126 / SM0557674 [other barcode numbers
include SM0557682, SM0557675, SM0557684, and SM0557688; KUNHM-ENT.
Diagnosis. This species is similar to P. jelineki that is known from Costa Rica and parts
of Nicaragua, however, it differs from this and other Neotropical members of the genus by the
following attributes: large densely spaced punctures on the metasternal disc, more globular
shaped terminal antennomere and corresponding sensillar region, erect and semi-erect rows of
alternating setae on the elytra, strongly oblique apical wall of prosternal process when viewed
laterally, and the basal piece of the aedeagal ejaculatory rods are not elbowed, but have sharply
produced lateral flanges.
Description. Length 2.85mm, Width 2.1mm, Depth 1.3mm. Body moderately convex,
surface shining, unicolorous light brown, legs and venter only slightly lighter. Pronotum and
elytra margins with moderate fimbriae, setae subequal to length of antennal scape. Dorsal and
ventral pubescence fine, moderately short and sparsely distributed.
Head surface deeply, irregularly punctate, interspersed large and small punctures on
vertex, becoming somewhat more dense towards orbits and fronotclypeal region. Larger
112
punctures 4-5 X diameter of eye facet, smaller punctures ~3 X diameter. Interspaces smooth to
finely alutaceous and shining, becoming slightly granular near orbits. Most punctures give rise
to a moderate length straight golden seta. Pronotal surface with large punctures equal in size to
large punctures on vertex of head, interspersed with relatively few smaller punctures, similar in
size to those on vertex. Interspaces smooth to finely alutaceous and shining, about 1-1.5
diameters apart. Each puncture gives rise to a moderate length golden seta, most seta are
straight. Scutellar surface with deeply to moderately impressed smaller punctures similar to
smaller ones on pronotum and vertex, some punctures giving rise to short seta, interspaces
smooth to alutaceous. Elytral surface with serial rows of alternating large and small deep
punctures. Smaller punctures are equal in size to those on pronotum, larger punctures are ~2-2.5
times diameter of smaller ones. Smaller punctures giving rise to an erect golden seta, larger
punctures giving rise to a semi-erect golden seta that is slightly longer than the erect seta.
Interspaces broad between punctures of a given row and between different rows. Within a row,
small punctures are separated by ~1.5 puncture diameters, and large punctures by ~1 puncture
width. Rows are separated by 1.0-1.5 small puncture diameters. Interspaces always shining but
variable from smooth to finely alutaceous in sculpture. Pygidium deeply densely punctate,
punctures equal in size to larger ones on pronotum, each puncture giving rise to a short stiff
golden seta. Interspaces narrow, 0.25-0.5 diameter, with granular sculpture.
Venter with similar sparse golden pubescence as dorsum. Mentum with large shallow
punctures, equal in size to large ones on vertex, each giving rise to a short seta, interspaces
alutaceous to finely granular. Submentum and gula similar in punctation to mentum but with
interspaces completely granular. Prosternum and epimeron moderately impressed with irregular
punctures, punctures slightly larger than those on mentum, interspaces alutaceous with granular
and microreticulate areas, prosternal punctures separated by 0.5-1 diameter, those on the
113
epimeron by 0.25 to 0.5 diameter. Mesosternum with shallow punctures, subequal in diameter of
those on prosternum, interspaces alutaceous to granular, separated by ~0.5-1 diameter.
Metasternum irregularly punctate, with deeply impressed large punctures on disc similar in size
to large ones on pronotum, interspaces smooth to alutaceous on metasternal disc becoming
mostly alutaceous to granular laterally, punctures separated by ~1 diameter on disc. Abdominal
sternite 1 with faint, almost obsolete punctures, punctures 0.75 diameter of those on
metasternum, interspaces alutaceous to granular, separated by ~0.5-1 diameter. Abdominal
sternites 2-4 with two or three irregular rows of punctures, one row near anterior margin and the
other two near posterior margin, punctures similar in size to those on sternite 1, rows separated
by 0.5 puncture diameter, punctures within rows separated by ~0.25-0.5 diameter. Rows on
abdominal sternite 4 becoming less organized. Hypopygidium with moderately deep punctures,
similar in size to those on sternites 2-4, interspaces alutaceous to granular, punctures separated
by 0.5-1 diameter.
Head slightly wider than long (W:L = 1.3:1), fronotclypeal region moderately projecting
anteriorly. Vertex with moderate concavity between orbits near fronotclypeal region. Labrum
with shallow incision at anterior margin. Antennal club compact, obovate, asymmetrical with
the last antennomere longer than the previous two combined. Antennomeres 6-8 disc-like, 4-5
cuboidal to slightly trapezoidal. Antennal scape asymmetrical, hemispherical, 1.2 times as long
as pedicel. Pedicel subcylindrical in shape. Antennal segment 3 subequal in length to pedicel,
and similarly shaped. Antennal club large, ~0.90 length of segments 1-8 combined. Antennal
grooves very deep and widely excavate, slightly converging posteriorly. Mental submental ridge
slightly produced, at level of submentum, the ridge is laterally sculptured with oblique to
longitudinal microreticulations. Mentum with single medial anterior angle, entire structure
triangular and somewhat convex.
114
Pronotum widest in posterior third (L:W = 1:1.8), anterior margin broadly trapezoidal,
posterior margin broadly convex, lateral margins less arcuate posteriorly. Scutellum large,
obtusely triangular, apex rounded. Prosternal process somewhat narrowed between procoxae,
apex somewhat acuminate, in lateral aspect the anterior and posterior ends are prominent and
slightly convex medially. Posterior apical wall prominent and sharply oblique. Mesosternum
extending to midway between mesocoxae, evenly concave for reception of the metasternum.
Metasternum wider than long (W:L = 2.6:1.0). Metepisternum with slight medial constriction,
oblique line dividing anterior 0.20 of structure. First abdominal sternite with moderately broad
process between metacoxae. First sternite ~2X’s longer than second sternite. Sternites 2-3
subequal in length, the fourth slightly larger than the preceding two. Hypopygidium subequal in
length to first abdominal sternite.
Protibia distinctly crenulate along lateral edge. Apical tooth very prominent, slightly
longer than tarsomeres 1 and part of 2 combined. Outer apical notch with ~95° angle, notch
depth shallow, subequal to length of tarsomere 1. Inner apical spine short, equal in length to
tarsomere 1. Protibia not heavily armored, but with characteristic dense patch of stiff setae along
the inner apical region. Mesotibia more heavily armored than protibia with more dense stiff
setae and a row of numerous slender spines along entire lateral edge. Outer apical process
elongate and robust, larger than protibia process. Inner apical spine equal in length to tarsomeres
1-2 combined. Metatibia with armature similar to that of mesotibia, but outer apical spine much
longer.
Male genitalia well-sclerotized. Anal sclerite with large broadly curved region
anteriodorsally; apex moderately fimbriate; ventrally with a broad medial concavity approaching
apex (Fig. 13). Spiculum gastrale with wide rounded lateral flanges, medial margins concave
proximally, sclerotized region more or less hourglass-shaped, short stiff setae originating from
115
apex (Fig. 52). Tegmen evenly rounded apically (Fig. 92), much longer than wide (w:l =
1.0:2.83), lateral row of setae visible from the median fossa to prior to around the apex, large
shallow concavity in apical third with inner row of setae attaining apex of concavity, basal notch
perpendicular, basal margin nearly straight. Median lobe large and robust, ~0.85 the length of
the tegmen, apex rounded, apical opening well-developed with proximal concavity, base tapering
to acute point (Fig. 133). Ejaculatory rods not fused to basal piece, straight and slightly outward
at basal piece (Fig. 174). Basal piece of rods with sharply produced lateral flanges and a slightly
curved central region.
Female genitalia moderately sclerotized. Paraprocts moderately large with sclerotization
only along median line to apico-lateral angles. Gonocoxite with two basal lateral prominences,
basal ridge well-sclerotized. Gonocoxal apices with recurved “tooth” absent. Three primary
setae originate from small depressions on the gonocoxal apices. Intragonocoxal invagination
deep, ~0.66 the length of the gonocoxite (Fig. 212).
Variation. No demonstrable variation within the type series.
Seasonality/Habitat. All specimens were collected in late May in southern Nicaragua
near the Costa Rican border in a lowland tropical forest.
Distribution. Known only from the type locality in southern Nicaragua.
Notes. Specimens were collected from both puffballs and fungusy logs. The latter
suggests that this species may be residing in members of bracket and/or shelf fungi that dominate
most old decaying wood.
Etymology. Specific epithet honors Christopher Carlton, director of the Louisiana State
Arthropod Museum, for his guidance and support during the course of this study.
Pocadius centralis Cline new species
116
(Figs. 14, 53, 93, 134, 175, 213)
Type Material Examined. HOLOTYPE ♂ (CMN): HONDURAS: Cortés; 25km N
Cofradia, P.N.; Cosuco, 15.IX.-7.X.1994;S.&J. Peck, cloud forest; flight inter. Trap, 94-62 /
HOLOTYPE; Pocadius; centralis; A.R. Cline des. 2004. 1 ♂ PARATYPE (CMN): MEX.: SLP,
1700m; 40km W. Xilitla; 12.VI-6.VIII.83; S&J Peck, FIT; pine-oak forest / PARATYPE;
Pocadius; centralis; A.R. Cline des. 2004. 1 ♀ PARATYPE (CMN): GUAT.: Zacapa; 3.5km SE
LaUnion; 1500m, 25-27.VI.1993; J. Ashe & R. Brooks; cloud forest, FIT / PARATYPE;
Pocadius; centralis; A.R. Cline des. 2004. 1 ♀ PARATYPE (SNEC): HONDURAS:
Ocotepeque ; 24km E Ocotepeque ; El Güisayote, 14-16.VI.1994 ; 2170m, 14°25’N, 89°04’W ;
J. Ashe, R. Brooks #117; ex. flight intercept traps/ PARATYPE; Pocadius; centralis; A.R. Cline
des. 2004.
Diagnosis. This species is similar in coloration to P. maquipucunensis in its dark
brown/black surface, however, it differs from it and the other Neotropical species by the
following suite of characters: antennal club symmetrical with terminal antennomeres not greatly
enlarged; terminal antennomeres with circular depressed region medially near base; metasternum
deeply densely punctate throughout with greatly enlarged punctures; pronotum densely deeply
punctate; protibia with distinct outer apical tooth but rather shallow apical notch; pubescence
very fine and light colored; abdominal process broad; males with pygidial apex distinctly
concave medially; tegmen with inner row of setae complete and laterally with elongate setae;
median lobe elongate oval with large bilobed internal structure; internal sac sclerites with one
basal piece having long curved lateral arms; and ovipositor with extremely deep intragonocoxal
invagination, ~0.75 length of gonocoxite.
Description. Length 3.45mm, Width 2.25mm, Depth 1.2mm. Body moderately convex,
surface shining, dark brown/black in color, legs and antennal club somewhat lighter. Pronotum
117
and elytra margins with moderately long fimbriae with that of the elytra being distinctly longer
than that of the pronotum. Dorsal and ventral pubescence fine and light colored.
Head surface deeply, irregularly punctate, punctures mostly larger on vertex, becoming
somewhat smaller towards orbits, occiput and fronotclypeal region. Larger punctures 4X
diameter of eye facet, smaller punctures 2X diameter; interspaces smooth and shining. Most
vertexal punctures give rise to a moderately long curved seta. Pronotal surface with large
punctures equal in size to large punctures on vertex of head, interspersed with relatively few
smaller punctures, similar in size to smaller ones on vertex; interspaces smooth and shining,
large punctures ~0.75-1 diameter apart. Most punctures give rise to an elongate seta, most seta
are decumbent. Scutellar surface with very few vague shallowly impressed punctures, ~0.5
diameter of small vertexal puncture, some punctures giving rise to minute setae, interspaces
smooth to finely alutaceous. Elytral surface with serial rows of alternating large and small deep
punctures. Smaller punctures are equal in size to smaller ones on pronotum, larger punctures are
~1.25-1.5X diameter of smaller ones. Both smaller and larger punctures giving rise to an
elongate decumbent golden seta. Interspaces moderately broad between punctures of a given
row and between different rows. Within a row, small punctures are separated by ~1.5 puncture
diameter, and large punctures by 1 puncture diameter. Rows are separated by ~1.0 puncture
diameter; interspaces always shining but variable from smooth to finely alutaceous in sculpture.
Pygidium densely punctate, punctures equal in size to larger ones on pronotum, each puncture
giving rise to a moderately long golden seta; interspaces narrow, 0.33-0.5 diameters, with
smooth to finely alutaceous sculpture.
Venter with similar pubescence as dorsum. Mentum with few shallow punctures, equal
in size to smaller ones on vertex; interspaces smooth to finely microreticulate. Submentum and
gula similar in punctation to mentum but somewhat larger with interspaces more alutaceous.
118
Prosternum and epimeron deeply irregularly punctate, punctures subequal to larger ones on
pronotum; interspaces alutaceous with microreticulate areas, prosternal punctures separated by
0.5 diameter, those on the epimeron 0.33-0.5 diameter. Mesosternum with moderately impressed
punctures, subequal to those on prosternum, interspaces alutaceous to finely microreticulate,
separated by about 0.5 diameter. Metasternum deeply densely irregularly punctate, with
punctures on disc similar in size to larger ones on pronotum, interspaces smooth to finely
alutaceous on metasternal disc becoming more alutaceous and microreticulate laterally, disc
punctures separated by ~0.5-1 diameter. Abdominal sternite 1 with punctures equal to large
punctures on elytra, interspaces mostly alutaceous, separated by ~1 diameter with only few
punctures present on abdominal process. Abdominal sternites 2-4 with two irregular rows of
punctures, one row near anterior margin and the other near posterior margin, punctures subequal
to those on metasternum, rows separated by 1.5-2 puncture diameters, punctures within rows
separated by ~0.5 puncture diameter. Rows on abdominal sternite 4 becoming less organized.
Hypopygidium with deep punctures, similar in size to those on sternites 2-4, interspaces
alutaceous to granular, punctures separated by ~0.5 puncture diameter.
Head slightly wider than long (W:L = 1.3:1), fronotclypeal region moderately projecting
anteriorly. Vertex with shallow extensive concavity between orbits near fronotclypeal region.
Antennal club compact, oval, symmetrical with the last antennomere subequal in length to the
previous two combined. Antennomeres 4-8 more or less compact, with 7-8 characteristically
disc-like and 4-6 trapezoidal. Antennal scape asymmetrical, somewhat hemispherical, ~2X as
long as pedicel. Pedicel subcylindrical in shape. Antennal segment 3 equal in length to pedicel.
Antennal club large, ~0.70 length of segments 1-8 combined. Antennal grooves very deep and
widely excavate, slightly converging posteriorly. Lateral mental/submental ridge prominent, at
level of submentum, ridge with longitudinal microreticulations laterally and densely punctate
119
medially. Mentum with anterior angles faint but present, anterior margin obsoletely angulate,
entire structure somewhat pentagonal, convex when viewed laterally.
Pronotum transverse, widest in posterior third (L:W = 1:2.0), anterior margin broadly
deeply trapezoidal, posterior margin moderately convex, lateral margins less arcuate posteriorly.
Scutellum large, obtusely triangular, apex rounded. Prosternal process somewhat narrowed
between procoxae, apex acuminate, in lateral aspect the anterior and posterior ends are
prominent with a modest convexity medially; the posterior end being somewhat prolonged.
Posterior apical wall short and slightly oblique. Mesosternum extending to midway between
mesocoxae, evenly concave for reception of the metasternum. Metasternum width to length ratio
is ~2.8:1.0. Metepisternum with slight medial constriction, oblique line dividing anterior 0.15 of
structure. Elytral humeri moderately produced, lateral margin very narrow. First abdominal
sternite with broad process between metacoxae. First sternite ~2X’s longer than second sternite.
Sternites 2-3 subequal in length, the fourth slightly larger than the preceding two.
Hypopygidium subequal in length to first abdominal sternite.
Protibia distinctly crenulate along lateral edge. Apical tooth moderately prominent,
slightly longer than tarsomeres 1. Outer apical notch with ~100° angle, notch depth shallow,
subequal to length of tarsomere 1. Inner apical spine subequal in length to tarsomeres 1.
Protibia not heavily armored but with characteristic dense patch of stiff setae along the inner
apical region. Mesotibia more heavily armored than protibia with more dense stiff setae and a
row of numerous slender spines along entire lateral edge. Outer apical process moderately
robust, only slightly larger than protibia process. Inner apical spine equal in length to tarsomere
1 and part of 2 combined. Metatibia with armature similar to that of mesotibia.
Male genitalia well-sclerotized. Anal sclerite with large broadly curved region
anteriodorsally (Fig. 14); apex with elongate moderately dense fimbria; ventrally with a broad
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medial concavity approaching apex in a narrowly concave manner. Spiculum gastrale with wide
lateral flanges apically oriented, medial margins concave , elongate stiff setae originating from
apex (Fig. 53). Tegmen narrowly rounded apically (Fig. 93), much longer than wide (w:l = 1:3),
lateral row of setae visible from the median fossa to prior to the apex, inner row of setae
complete, elongate oval concavity in apical third, basal notch perpendicular, basal margin
concave. Median lobe large and oval, ~0.75 the length of the tegmen, apex acuminate, apical
opening well-developed with bilobed internal structure (Fig. 134). Ejaculatory rods not fused to
basal piece or each other, distinctly curved outward apically. Basal piece of rods with as one
fused piece with long basally projecting lateral arms and centrally with a short medio-basal
bilobed projection (Fig. 175).
Female genitalia moderately sclerotized. Paraprocts large with sclerotization only along
median line to apico-lateral angles. Gonocoxite with one broad lateral prominence, basal ridge
well-sclerotized. Gonocoxal apices with recurved “tooth” present. Two primary setae originate
from small depressions on the gonocoxal apices. Intragonocoxal invagination extremely deep,
~0.75 length of gonocoxite (Fig. 213).
Variation. None observed.
Seasonality/Habitat. Specimens are known to occur from June to October in mid to
high elevation forests.
Distribution. Known range is from southern Mexico to Honduras.
Notes. No fungal records exist for this species. It appears to favor montane
environments.
Etymology. Specific epithet is a derivative of the region from which the species is
known, i.e. Central America.
121
Pocadius cochabambus Cline new species
(Figs. 15, 54, 135, 176)
Type Material Examined. HOLOTYPE ♂(CMN): BOLIVIA: Cochabamba;
Cochabamba, 67.5km NE Est. Biol.; Valle del Sajita, Univ. De San Simon; 300m, 17°6’33”S
64°47’52”W; 7-9-II-1999, R. Hanley, ex. flight; intercept trap, BOL1H99 057 / HOLOTYPE;
Pocadius; cochabambus; A.R. Cline des. 2004.
Diagnosis. This species is most similar to P. ashei, but differs from it and the other
Neotropical fauna by the following suite of characters: antennal club greatly widened with large
oblong sensillar region; habitus not shining with surface almost entirely alutaceous to granular in
texture; all tibial spines elongate; elytral fimbria elongate; densely punctured pronotum; serial
row of small punctures on elytra often with multiple punctures not just a single row; the extreme
W:L ratio of the metasternum of 3.2:1.0; aedeagus with tegmen having incomplete row of inner
setae; basal piece of internal sac sclerites U-shaped curved structure and a median section with
an elongate blunt process.
Description. Length 3.45mm, Width 2.25mm, Depth 1.05mm. Body moderately
convex, surface dull, uniformly light brown in color. Pronotum and elytra margins with elongate
fimbriae, setae longer than width of antennal scape. Dorsal and ventral pubescence quite long.
Head surface deeply, densely, irregularly punctate with large and small punctures,
punctures mostly larger on vertex, becoming somewhat smaller towards orbits, occiput and
fronotclypeal region. Larger punctures 4-5X diameter of eye facet, smaller punctures 1-2X
diameter; interspaces alutaceous to granulate and dull. Each puncture gives rise to an elongate
somewhat curved golden seta. Pronotal surface with large punctures equal in size to large
punctures on vertex of head, interspersed with numerous smaller punctures, ~0.5 size of larger
ones; interspaces alutaceous and dull, all punctures ~0.25-0.5 large diameter apart. Each
122
puncture gives rise to a long golden seta, most seta are decumbent. Scutellar surface with several
shallowly impressed punctures equal to smaller ones on pronotum, some punctures giving rise to
setae; interspaces are granular. Elytral surface with serial rows of alternating large and small
deep punctures. Smaller punctures are equal in size to smaller ones on pronotum, larger
punctures are ~2-2.5X diameter of smaller ones. Smaller punctures giving rise to an elongate
erect long golden seta, larger punctures giving rise to a decumbent to adpressed moderately long
golden seta that is much shorter than those derived from the smaller punctures. Interspaces
broad between punctures of a given row and between different rows. Within a row, small
punctures are separated by ~2 puncture diameters, and large punctures by 1 puncture diameter.
Larger rows are separated by 2 large puncture diameters. Interspaces always dull with
alutaceous to granular sculpture. Pygidium densely punctate, punctures equal in size to smaller
ones on pronotum, each puncture giving rise to a moderately long golden seta’ interspaces
narrow, ~0.5 diameter, with mostly granular sculpture.
Venter with similar long golden pubescence as dorsum. Mentum with obsolete shallow
punctures, equal in size to smaller ones on vertex, each giving rise to a moderately elongate seta;
interspaces granular. Submentum and gula similar in punctation and surface sculpture as
mentum. Prosternum and epimeron moderately impressed with irregular punctures, punctures
2X larger than those on mentum with epimeron punctures slightly larger than those on
prosternum, interspaces granular with microreticulate areas, prosternal punctures separated by
0.5 diameter, those on the epimeron by 0.25 to 0.5 diameter. Mesosternum with shallow
punctures, subequal in diameter to those on prosternum, interspaces alutaceous to granular,
separated by 0.5 diameter. Metasternum irregularly punctate, with moderately impressed
punctures on disc similar in size to those on mesosternum, interspaces granular on metasternal
disc becoming granular and microreticulate laterally, punctures separated by ~1-2 diameters.
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Metepisternum deeply punctate with large irregular punctures, 2X larger than those on
metasternal disc. Abdominal sternite 1 with faint punctures, punctures equal to those on
metasternal disc, interspaces alutaceous to granular, separated by ~1 diameter. Abdominal
sternites 2-4 with two irregular rows of punctures, one row near anterior margin and the other
near posterior margin, punctures similar in size to those on metasternal disc, rows separated by
1.5-2 puncture diameters, punctures within rows separated by ~0.5 punctures width. Rows on
abdominal sternites 3-4 becoming less organized. Hypopygidium with moderately deep
punctures, similar in size to those on sternites 2-4, interspaces granular with microreticulations,
punctures separated by 0.5 puncture diameter.
Head slightly wider than long (W:L = 1.26:1), fronotclypeal region moderately projecting
anteriorly. Vertex with shallow concavity between orbits near fronotclypeal region. Antennal
club compact, greatly widened apically, greatly asymmetrical with the last antennomere longer
than the previous two combined. Antennomeres 4-8 more or less compact, with 6-8
characteristically disc-like, 4-5 cuboidal to slightly trapezoidal. Antennal scape asymmetrical,
somewhat hemispherical, 1.8 times as long as pedicel. Pedicel subcylindrical in shape. Antennal
segment 3 ~.85 length to pedicel. Antennal club large, ~0.75 length of segments 1-8 combined.
Antennal grooves very deep and widely excavate, slightly converging posteriorly. Lateral
mental/submental ridge prominent and elongate, at level of submentum, ridge sculptured
laterally with longitudinal microreticulations and the mental/submental sulcus if granulate in
sculpture. Mentum with anterior angles visible, anterior margin angulate, entire structure
pentagonal, when viewed laterally slightly convex.
Pronotum widest near posterior angles (L:W = 1:2.0), anterior margin broadly concave,
posterior margin moderately convex, lateral margins not distinctly arcuate. Scutellum large,
obtusely triangular, apex rounded. Prosternal process narrowed between procoxae, apex
124
acuminate, in lateral aspect there is an even convexity over the procoxae. Posterior apical wall
prominent and slightly oblique. Mesosternum extending to midway between mesocoxae, evenly
concave for reception of the metasternum. Metasternum much wider than long W:L = 3.2:1.0.
Metepisternum with slight medial constriction, oblique line dividing anterior 0.18 of structure.
First abdominal sternite with broad process between metacoxae. First sternite ~2X’s longer than
second sternite. Sternites 2-3 subequal in length, the fourth slightly larger than the preceding
two. Hypopygidium subequal in length to first abdominal sternite.
Protibia distinctly crenulate along lateral edge. Apical tooth prominent, slightly longer
than tarsomeres 1. Outer apical notch with ~135° angle, notch depth shallow, equal to length of
one-third of tarsomere 1. Inner apical spine subequal in length to tarsomeres 1-2 combined.
Protibia heavily armored with characteristic dense patch of stiff setae along the inner apical
region. Mesotibia more heavily armored than protibia with more dense stiff setae and a row of
numerous slender spines along entire lateral edge. Outer apical process robust, equal in length to
that of the protibia. Inner apical spine equal in length to tarsomeres 1-2 and part of 3 combined.
Metatibia with armature similar to that of mesotibia, but apical spine not as long, only equal in
length to tarsomeres 1-2 combined.
Male genitalia well-sclerotized. Anal sclerite with large broadly curved region
anteriodorsally (Fig. 15); apex with elongate dense fimbriae; ventrally with a broad medial
concavity approaching apex in a convex manner. Spiculum gastrale with wide lateral flanges,
medial margins perpendicular, moderately long stiff setae originating from apex (Fig. 54).
Tegmen broadly rounded apically (Fig. 94), much longer than wide (w:l = 1:3), lateral row of
setae visible from the median fossa to prior to the apex, inner row of setae incomplete, small
shallow concavity barely visible in apical third, basal notch perpendicular, basal margin concave.
Median lobe large and robust, ~0.66 the length of the tegmen, rounded, apical opening well-
125
developed with bilobed internal structure (Fig. 135). Ejaculatory rods not fused to basal piece or
each other, curved inward and expanded outward at proximally and distally. Basal piece of rods
with U-shaped curved structure and a central elongate blunt section (Fig. 176).
Female genitalia not observed.
Variation. Known only from the holotype.
Seasonality/Habitat. Collected in early February from a lowland forest.
Distribution. Known only from the type locality in western Bolivia.
Notes. No host data is available for this species.
Etymology. Specific epithet is a derivative of the type locality, i.e. Cochabamba,
Bolivia.
Pocadius coxus Cline new species
(Figs. 16, 55, 95, 136, 177)
Type Material Examined. HOLOTYPE ♂ (SNEC): BRAZIL: Rondonia; 9km NE
Cacaulandia; XII-1996 to I-1997; K. Vulinec & D. Mellow / HOLOTYPE; Pocadius; coxus;
A.R. Cline des. 2004.
Diagnosis. This species is most similar to P. maquipucunensis in overall shape and
dorsal coloration, but differs from it and other Neotropical species by the following combination
of characters: procoxae light tan in color, terminal antennomere with medial and lateral elliptical
sensillar regions, antennomere 3 ~0.5 the length of pedicel, dorsal habitus with primarily
granular surface sculpture but shining, sharply oblique apical wall of prosternal process when
viewed laterally, ejaculatory rods distinctly elongate with medial swellings that almost abut, and
basal piece of internal sac sclerites with two lateral arms and a medial “t-shaped” arm.
126
Description. Length 3.6mm, Width 2.1mm, Depth 1.35mm. Body moderately convex,
surface shining, dark brown to black in color, with venter and legs somewhat lighter, and
procoxae light tan in color. Pronotum and elytra margins with moderately long fimbriae, setae
slightly longer than width of antennal scape. Dorsal and ventral pubescence golden and
moderately long.
Head surface deeply, densely, irregularly punctate, punctures larger on vertex with
interspersed smaller punctures, becoming predominantly smaller towards orbits and
fronotclypeal region. Larger punctures 4-5 X diameter of eye facet, smaller punctures ~2X
diameter. Interspaces alutaceous to granular and moderately shining. Each puncture gives rise
to a moderately long curved golden seta. Pronotal surface with large punctures equal in size to
large punctures on vertex of head, interspersed with relatively few smaller punctures, ~0.5-0.75
size of larger ones. Interspaces alutaceous to granular, about 0.5-1 diameter apart. Each
puncture gives rise to a moderately long golden seta, most seta are curved. Scutellar surface with
very shallowly impressed punctures that are mostly aggregated in anterior one-third of structure,
some punctures giving rise to short setae, and the interspaces are mostly granular. Elytral
surface with serial rows of alternating large and small deep punctures. Smaller punctures are
equal in size to those on pronotum, larger punctures are ~2 times diameter of smaller ones.
Smaller punctures giving rise to an erect moderately long golden seta, larger punctures giving
rise to a semi-erect shorter golden seta. Interspaces wide between punctures of a given row and
between different rows. Within a row, small punctures are separated by ~2 puncture width, and
large punctures by 0.5-0.75 puncture diameter. Large rows are separated by 1 large puncture
diameter. Interspaces shining but variable from alutaceous to finely microreticulate in sculpture.
Pygidium densely punctate, punctures equal in size to smaller ones on pronotum, each puncture
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giving rise to a moderately long golden seta. Interspaces narrow, 0.25-0.5 diameter, with mostly
granular sculpture.
Venter with similar golden pubescence as dorsum. Mentum with large very shallow
punctures, equal in size to larger ones on vertex, each giving rise to a short seta; interspaces
granular to finely microreticulate. Submentum and gula similar in punctation to mentum but
with interspaces more microreticulate. Prosternum and epimeron deeply irregularly punctate,
punctures 1.5X larger than those on mentum, interspaces alutaceous to granular with
microreticulate areas, prosternal punctures separated by 0.5-1 diameter, those on the epimeron by
0.25 to 0.5 diameter. Mesosternum with shallow punctures, subequal to those on prosternum,
interspaces alutaceous to granular, separated by about 0.5 to 1 diameter and aggregated near
metasternal border. Metasternum irregularly punctate, with moderate faint punctures on disc
similar in size to those on mesosternum, interspaces granular with microreticulations on
metasternal disc, appearing rugulose, and becoming more microreticulate laterally, punctures
separated by ~1-2 diameters. Abdominal sternite 1 with faint, almost obsolete punctures,
punctures equal to those on mesosternum, interspaces alutaceous to granular, separated by ~1-2
diameters. Abdominal sternites 2-4 with two irregular, disorganized rows of punctures, one row
near anterior margin and the other near posterior margin, punctures similar in size to those on
metasternum, rows separated by ~2 puncture diameter, punctures within rows separated by ~0.5-
1 punctures diameter. Rows on abdominal sternite 4 becoming less organized. Hypopygidium
with deeper punctures than the other abdominal sternites, similar in size to those on sternites 2-4,
interspaces mostly granular with microreticulate regions, punctures separated by 0.25-0.5
puncture diameter.
Head slightly wider than long (W:L = 1.35:1.0), fronotclypeal region moderately
projecting anteriorly. Vertex with shallow concavity between orbits near fronotclypeal region.
128
Labrum with medial incision at anterior margin. Antennal club compact, oval, only slightly
asymmetrical with the last antennomere slightly longer than the previous two combined.
Antennomeres 4-8 more or less compact, with 6-8 characteristically disc-like and 4-5 more or
less trapezoidal. Antennal scape asymmetrical, somewhat hemispherical, 1.2 times as long as
pedicel. Pedicel subcylindrical in shape. Antennal segment 3 ~0.5 length of pedicel. Antennal
club large, subequal in length to segments 1-8 combined. Each club segment with dense short
setae, and only relatively few protruding setae (2-4 in number). Antennal grooves very deep and
widely excavate, slightly converging posteriorly. Lateral mental/submental ridge prominent, at
level of submentum, the ridge is heavily sculptured with numerous longitudinal
microreticulations and the sulcus is almost entirely granular in sculpture. Mentum with anterior
angles faint, anterior margin broadly angulate, appearing pentagonal ventrally and slightly
convex laterally.
Pronotum widest near posterior angles (L:W = 1:1.8), anterior margin broadly
trapezoidal, posterior margin moderately convex, lateral margins arcuate anteriorly. Scutellum
large, triangular, apex narrowly rounded. Prosternal process narrowed between procoxae, apex
acuminate, in lateral aspect there is a steep convexity over the procoxae with a sharp declivity
and then flattened region posterior to the coxae. Posterior apical wall prominent and greatly
oblique. Mesosternum extending to midway between mesocoxae, evenly concave for reception
of the metasternum. Metasternum much wider than long (W:L = 3:1). Metepisternum with
distinct medial constriction, oblique line dividing anterior 0.125 of structure. First abdominal
sternite with broad process between metacoxae. First sternite ~2X’s longer than second sternite.
Sternites 2-3 subequal in length, the fourth slightly larger than the preceding two.
Hypopygidium subequal in length to first abdominal sternite.
129
Protibia distinctly crenulate along lateral edge. Apical tooth prominent, subequal in
length to tarsomere 1 and part of 2 combined. Outer apical notch absent. Inner apical spine
subequal in length to tarsomeres 1 and part of 2 combined. Protibia not heavily armored but with
characteristic dense patch of stiff setae along the inner apical region. Mesotibia more heavily
armored than protibia with more dense stiff setae and a row of numerous slender spines along
entire lateral edge. Outer apical process elongate and robust, larger than protibia process. Inner
apical spine equal in length to tarsomeres 1-2 combined. Metatibia with armature similar to that
of mesotibia, with inner apical spine slightly longer.
Male genitalia well-sclerotized. Anal sclerite with apex coming to a more or less acute
point and somewhat fimbriate, ventrally with a broad medial concavity approaching apex (Fig.
16). Spiculum gastrale with wide lateral flanges, medial margins concave straight, long stiff
setae originating from apex (Fig. 55). Tegmen evenly rounded apically (Fig. 95), much longer
than wide (w:l = 1.0:2.9), lateral row of setae visible from the median fossa to around the apex,
oblong shallow concavity in apical third with short setae around lateral border, basal notch
perpendicular, basal margin concave. Median lobe large and robust, ~0.75 the length of the
tegmen, apex narrowly rounded, apical opening well-developed with simple internal structure
(Fig. 136). Ejaculatory rods not fused to basal piece, elongate and straight with medial swelling.
Basal piece of internal sac sclerites with two lateral arms and a medial arm that is “t-shaped”
(Fig. 177).
Female genitalia not observed.
Variation. Known only from the holotype.
Seasonality/Habitat. Specimen collected during December/January in lowland tropical
forest.
Distribution. Known from the type locality in western Brazil.
130
Notes. No host data is available for this species.
Etymology. Specific epithet denotes the distinctive light colored procoxae.
Pocadius crypsis Cline new species
(Figs. 17, 56, 96, 137, 178, 214)
Type Material Examined. HOLOTYPE ♂(SNEC): GUYANA: Region 8; Iwokrama
Forest, Turtle Mt. Base; camp, 4°43’5”N 58°43’5”W, 50m; 1 JUN 2001, R. Brooks, Z. Falin;
GUY1BF01 104, ex. in puffballs / SM0226858; KUNHM-ENT [barcode label] / HOLOTYPE;
Pocadius; crypsis; A.R. Cline des. 2004. 51 PARATYPES (SNEC) with the same data label as
the holotype but with PARATYPE designation labels and the following barcode numbers:
SM0226877, SM0226898, SM0226847, SM0386268, SM0386267, SM0226864, SM0226855,
SM0226856, SM0386265, SM0226857, SM0226853, SM0226852, SM0226851, SM0226850,
SM0226849, SM0226848, SM0226861, SM0226862, SM0226863, SM0226864, SM0226865,
SM0226867, SM0226866, SM0226869, SM0226897, SM0226878, SM0226876, SM0226875,
SM0226896, SM0226895, SM022688, SM0226884, SM0226886, SM0226882, SM0226881,
SM0226879, SM0226891, SM0226892, SM0226893, SM0226860, SM0226870, SM0226871,
SM0226872, SM0226873, SM0226874, SM0226868, SM0226889, SM0226894, SM0226890,
SM0226887, SM0226859. 1 PARATYPE (SNEC): GUYANA: Region 8; Iwokrama Forest,
1km W Kurupukari; Iwokrama Field Stn., 60m; 4°40’19”N 58°41’4”W, 28-29 MAY 2001; R.
Brooks, Z. Falin, GUY1BF01 064; ex. flight intercept trap / SM0565301; KUNHM-ENT
[barcode label] / PARATYPE; Pocadius; crypsis; A.R. Cline des. 2004. 1 PARATYPE (SNEC):
GUYANA: Region 8; Iwokrama Forest, 1km W Kurupukari; Iwokrama Field Stn., 60m;
4°40’19”N 58°41’4”W, 21 MAY 2001; R. Brooks, Z. Falin, GUY1BF01 005; ex. Acromyrmex
hystrix refuse pile / SM0568734; KUNHM-ENT [barcode label] / PARATYPE; Pocadius;
131
crypsis; A.R. Cline des. 2004. 1 PARATYPE (SNEC): GUYANA: Region 8; Iwokrama Forest,
Pakatau Hills; 70m, 4°44’54”N 59°1’36”W; 25-29 MAY 2001, R. Brooks; Z. Falin, GUY1BF01
061; ex. flight intercept trap / SM0569499; KUNHM-ENT [barcode label] / PARATYPE;
Pocadius; crypsis; A.R. Cline des. 2004.
Diagnosis. This species is most similar to P. coxus, but can be distinguished from them
by the following suite of characters: deep distinct protibial notch and heavily armored tibiae,
mentum transversely hemispherical, alternating rows of erect and semi-erect setae on elytra,
metasternal disc with minute faintly impressed punctures appearing nearly glabrous, body
uniformly light tan in color, terminal antennomere enlarged with small circular sensillar region,
elytral fimbriae distinctly elongate, epimeron with two distinct rows of large punctures, mentum
with numerous large punctures, habitus shining with mostly smooth surface sculpture, each
ejaculatory rod split into two adjacent rods, basal piece of internal sac sclerites with two short
lateral arms and a simple medial arm, anal sclerite with elongate fimbria, median lobe short and
robust with widely diverging internal structure, and tegmen with inner row of setae minute and
extending around apex of apical tegminal fossa.
Description. Length 3.75mm, Width 2.25mm, Depth 1.35mm. Body moderately
convex, surface shining, uniformly light brown in color. Pronotum and elytra margins with
elongate fimbriae that of the elytra being longer than the pronotum, all setae longer than width of
antennal scape. Dorsal and ventral pubescence long.
Head surface deeply, densely, irregularly punctate, punctures larger on vertex, becoming
somewhat smaller towards orbits and fronotclypeal region. Larger punctures 4-5 X diameter of
eye facet, smaller punctures ~2X diameter; interspaces smooth with some microreticulation and
shining. Each puncture gives rise to an elongate straight red-brown seta. Pronotal surface with
large punctures equal in size to large punctures on vertex of head, interspersed with relatively
132
few smaller punctures, ~0.5 size of larger ones; interspaces smooth with some fine
microreticulations, about 0.5-1 diameter apart. Each puncture gives rise to a long brown-red
seta, most seta are rather straight. Scutellar surface with shallowly impressed small punctures,
equal to small ones on pronotum some punctures giving rise to short straight setae, interspaces
smooth to finely alutaceous. Elytral surface with serial rows of alternating large and small deep
punctures. Smaller punctures are equal in size to those on pronotum, larger punctures are ~2-
2.5X diameter of smaller ones. Smaller punctures giving rise to an erect long red-brown seta,
larger punctures giving rise to a semi-erect long red-brown seta. Interspaces broad between
punctures of a given row and between different rows. Within a row, small punctures are
separated by ~2-2.5 puncture diameters, and large punctures by 0.75-1 puncture diameter. Large
rows are separated by 1-1.5 large puncture diameters. Interspaces always shining but variable
from smooth to finely alutaceous in sculpture. Pygidium densely punctate, punctures equal in
size to larger ones on pronotum, each puncture giving rise to a moderately long red-brown seta;
interspaces narrow, 0.25-0.5 diameter, with smooth to finely alutaceous sculpture and some faint
areas of microreticulation.
Venter with similar long red-brown pubescence as dorsum. Mentum with numerous
large deeply impressed punctures, equal in size to larger ones on vertex, each giving rise to a
moderately long seta, interspaces smooth with finely microreticulate areas. Submentum and gula
with little to no punctation punctation; interspaces alutaceous to faintly granular. Prosternum
and epimeron deeply irregularly punctate, punctures on epimeron slightly larger than those on
mentum and those on prosternum slightly smaller than mental punctures, interspaces alutaceous
with microreticulate areas, prosternal punctures separated by 0.5-1 diameter, those on the
epimeron by 0.25 to 0.5 diameter and organized into two distinct rows. Mesosternum with
shallow punctures, equal in size to those on mentum, interspaces smooth to alutaceous, separated
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by about 0.5 to 1 diameter and aggregated near metasternum. Metasternum irregularly punctate,
faint minute punctures on disc similar in size to small ones on vertex those laterally on
metasternum 2-3X diameter of those on disc, interspaces smooth to alutaceous on metasternal
disc becoming more granular laterally, punctures separated by ~2-3 diameters on disc.
Abdominal sternite 1 with large moderately impressed punctures, punctures equal to large
punctures on vertex, interspaces smooth to alutaceous, separated by ~1-2 diameter. Abdominal
sternites 2-4 with two irregular rows of punctures, one row near anterior margin and the other
near posterior margin, punctures similar in size to those on abdominal sternite 1, rows separated
by 0.5-1 puncture diameter, punctures within rows separated by ~0.25-0.5 punctures width.
Rows on abdominal sternites 3-4 becoming less organized. Hypopygidium with moderately
deep punctures, similar in size to those on sternites 2-4, interspaces smooth to alutaceous,
punctures separated by 1-2 puncture diameters.
Head wider than long (W:L = 1.44:1), fronotclypeal region moderately projecting
anteriorly. Labrum with broad concavity at anterior margin. Antennal club compact, ovoid,
asymmetrical with the last antennomere longer than the previous two combined. Antennomeres
4-8 more or less compact, with 6-8 characteristically disc-like and 4-5 mostly cuboidal.
Antennal scape asymmetrical, somewhat hemispherical, 1.6 times as long as pedicel. Pedicel
subcylindrical in shape. Antennal segment 3 subequal in length to pedicel. Antennal club large,
~0.85 length of segments 1-8 combined. Lateral mental/submental ridge moderately prominent,
at level of submentum, ridge is longitudinally sculptured with microreticulations and the sulcus
with alutaceous to finely granular surface sculpturing. Mentum with anterior angles obsolete,
anterior margin hemispherical, entire structure transversely hemispherical when viewed ventrally
and somewhat convex laterally.
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Pronotum widest in posterior third (L:W = 1:1.9), anterior margin shallowly
trapezoidal, posterior margin moderately convex, lateral margins less arcuate posteriorly.
Scutellum large, triangular, apex acute. Prosternal process narrowed between procoxae, apex
acuminate, in lateral aspect there is a slight even convexity over the procoxae. Posterior apical
wall prominent and oblique. Mesosternum extending to midway between mesocoxae, evenly
concave for reception of the metasternum. Metasternum much wider than long, W:L = 3.3:1.0.
Metepisternum with slight medial constriction, oblique line dividing anterior 0.18 of structure.
First abdominal sternite with acuminate process between metacoxae. First sternite ~2X’s longer
than second sternite. Sternites 2-3 subequal in length, the fourth slightly larger than the
preceding two. Hypopygidium subequal in length to first abdominal sternite.
Protibia distinctly crenulate along lateral edge. Apical tooth very prominent, slightly
longer than tarsomeres 1 and part of 2 combined. Outer apical notch with ~95° angle, notch
depth deep, equal to length of tarsomere 1 and part of 2 combined. Inner apical spine subequal
in length to tarsomere 1. Protibia heavily armored with characteristic dense patch of stiff setae
along the inner apical region and other elongate stiff setae. Mesotibia more heavily armored than
protibia with more dense stiff setae and a row of numerous slender spines along entire lateral
edge. Outer apical process elongate and robust, larger than protibia process. Inner apical spine
equal in length to tarsomeres 1-2 combined. Metatibia with heavier armature than that of
mesotibia with more setae and spines and the inner apical spine equal to tarsomeres 1-2 and part
of 3 combined.
Male genitalia well-sclerotized. Anal sclerite with large narrowly curved region
ventrally and apical fimbria elongate (Fig. 17). Spiculum gastrale with wide lateral flanges,
medial margins concave, inner raised area extending past posterior margin, and numerous short
stiff setae originating from apex (Fig. 56). Tegmen evenly rounded apically (Fig. 96), longer
135
than wide (w:l = 1.0:2.4), lateral row of setae visible from the median fossa to around the apex,
shallow concavity in apical third with minute setae laterally and around apex, basal notch
perpendicular, basal margin broadly concave. Median lobe short and robust, ~0.5 the length of
the tegmen, apex coming to an acuminate point, apical opening well-developed with internal
structure broadly separated (Fig. 137). Ejaculatory rods not fused to basal piece and split into
two adjacent pieces, the distal pieces abutting apically. Basal piece of rods with two lateral and
one medial arm (Fig. 178).
Female genitalia moderately sclerotized. Paraprocts large with sclerotization only along
median line to apico-lateral angles. Gonocoxite with two basal lateral prominences, basal ridge
well-sclerotized. Gonocoxal apices with recurved “tooth” absent. Two primary setae originate
from small depressions on the gonocoxal apices. Intragonocoxal invagination moderately deep,
~0.5 length of gonocoxite (Fig. 214).
Variation. Some male specimens were less than 3mm in length.
Seasonality/Habitat. Specimens were collected from May and June in lowland tropical
forests.
Distribution. Known from localities in central Guyana.
Notes. Host data is indicated by the generic term “puffball” on some data labels. One
specimen was collected from an Acromyrmex hystrix refuse pile, which could be LSAMidental
or indicate that the refuse was sufficiently decayed to potentially have fungal mycelia intermixed
within it, thereby providing the individual a fungus meal.
Etymology. Specific epithet denotes the cryptic, uniform coloration of the habitus,
which is undoubtedly excellent camouflage in the field.
Pocadius decoratus Kirejtshuk
136
(Figs. 97, 138)
Type Material Examined. (1) HOLOTYPE ♂ (ZISP): [N] Vietnam, mountains, 50km;
NE Thai-Nguen, 300m; 9/1/1964, O. Kabokav / Holotypus Pocadius; decoratus; det. Kirejtshuk
1983.
Non-Type Material Examined. None.
Diagnosis. Differs from the other Pocadius fauna by its distinctive color pattern of
orange and black on the pronotum and elytra, and also in genitalic features.
Redescription. Length 4.1mm, Width 2.1mm, Depth 2.0mm. Body moderately convex,
surface very shining, orange in color with distinct black markings on the pronotum and elytra,
setae short and fine. Pronotum and elytra with margins scarcely fimbriate. Pygidium and
hypopygidium scarcely setate, but posterior margins fimbriate.
Head surface deeply punctate, punctures becoming dense near orbits, punctures becoming
smaller near fronto-clypeal region. Only relatively few punctures giving rise to small fine pale
setae. Interspaces smooth to slightly alutaceous. Smaller punctures about 0.75 the size of larger
ones, large punctures about 7-8 times the size of facets. Eyes with small fine facets. Pronotal
surface with large and small punctures interspersed throughout. Large and small punctures
similar in size to respective large and smaller punctures on head. Interspaces 1-2 puncture
diameters apart, smooth to alutaceous. Each puncture bearing a short fine pale seta. Scutellar
surface scarcely punctate with small punctures similar in size to the smaller punctures on the
head, granular near apex, interspaces alutaceous. Eytral surface with serial rows of alternating
large and small shallow punctures. Large punctures 2-3 times diameter of the large punctures on
head, each giving rise to a single short fine seta. Small punctures 0.20 diameter of large
punctures, each giving rise to a short fine seta. Interspaces mostly smooth between all punctures,
larger punctures separated from each other by < 0.5 puncture diameter, smaller punctures
137
separated from each other by 2 puncture diameters. Large and small puncture rows separated
from each other by ~1 large puncture diameter. Pygidium moderately punctate, punctures equal
in size to small punctures on elytra, interspaces alutaceous with some minute microreticulation,
punctures separated by ~1 puncture diameter. Each puncture giving rise to a short stiff seta,
setae ~0.5 length of those on elytra.
Venter similar in pubescence to dorsum. Mentum alutaceous with no distinct punctation.
Submentum and gula with scattered small punctures equal in size to the smaller punctures on the
vertex, interspaces alutaceous with some microreticulation. Prosternum and epimeron irregularly
punctate, punctures approximately equal in size to larger punctures on vertex, interspaces mostly
smooth with faint alutaceous areas, prosternal punctures separated by 0.5 diameter, those on the
epimeron by 0.25 to 0.5 diameter. Prosternal punctures quite deep. Mesosternum with very faint
small punctures. Metasternum irregularly punctate, with small faint punctures on disc similar in
size to those on vertex, interspaces smooth on metasternal disc, punctures separated by ~3
diameters. Large punctures, equal large punctures on elytra, on lateral region of metasternum,
interspaces smooth and ~1-2 diameters apart. Abdominal sternite 1 with faint large punctures
occurring in all but the anterior 0.33 of the structure which is completely alutaceous. Interspaces
alutaceous, punctures separated by 1 diameter. Abdominal segments 2-4 with punctures aligned
in irregular lateral rows. Punctures equal in size to those on abdominal sternite 1. Interspaces
alutaceous, punctures separated by 0.25 – 0.5 diameter. Hypopygidium densely deeply punctate,
punctures equal to those on other abdominal sternites, interspaces smooth to alutaceous with
faint microreticulation, punctures separated by 0.25- 0.5 diameter of puncture.
Head much wider than long, broadly triangular, with fronotclypeal region projecting
slightly anteriorly. Vertex with shallow concavity in the fronto-clypeal region. Eyes large with
small fine facets. Labrum transverse with minute medial incision on anterior margin. Antennal
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club compact, oblong oval, asymmetrical with the last antennomere longer than previous two
combined. Antennomeres 6-8 strongly flattened into disc-like structures, their combined length
equal to the length of antennomere 9. Antennal scape asymmetrical, globular, somewhat
hemispherical, 3 times as long as pedicel. Pedicel subcylindrical in shape, ~0.75 length of scape.
Antennal segment 3 subequal in length to pedicel. Segment 4 subquadrate, 0.5 the length of
segment 3. Antennal club large, ~0.75 length of segments 1-8 combined, asymmetrical, terminal
segment equal to preceding two segments. Antennal grooves moderately deep.
Pronotum widest near posterior angles (L:W = 1:2.97 ), anterior margin truncate,
posterior margin broadly convex with distinct “lobe” over scutellum. Scutellum large and
broadly hemispherical. Prosternal process somewhat narrowed between coxae, evenly rounded
at apex, in lateral aspect the apex is slightly depressed behind the procoxae. Mesosternum
extending to midway between mesocoxae. Mesepisternum larger than mesosternum.
Metasternum width to length ratio is ~3:1. Metepisternum narrow, only slightly concave
medially, anterior third moderately produced anteriolaterally. Elytral humeri moderately
produced, lateral margin very narrow. First abdominal sternite with broadly truncate process
between metacoxae, first sternite ~2X’s longer than second sternite. Sternites 2-4 subequal in
length. Hypopygidium subequal in length to first abdominal sternite.
Protibia with apical tooth prominent, slightly longer than tarsomeres 1-2 combined.
Outer apical notch with 90° angle, notch depth equal to tarsomere 1-2 combined. Inner apical
spine subequal in length to first tarsomere. Protibia with very little armature overall. Mesotibia
more heavily armored than protibia with a row of slender spines along entire lateral edge and
other spines scattered on ventral surface. Outer apical process elongate and robust, larger than
protibia process. Inner apical spine equal in length to tarsomeres 1 and 0.5 of 2. Metatibia
heavily armored with four rows of slender elongate spines, one dorsal row, one ventral row, and
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two lateral rows. Spines of varying lengths, but most longer than those on the mesotibia. Outer
apical process elongate and robust, equal in length to inner apical spine, projecting more
posteriorly than pro- or mesotibial processes. Inner apical spine subequal in length to tarsomeres
1-2 combined.
Male genitalia moderately well-sclerotized. Anal sclerite large broadly curved apically
(not figured); apex moderately fimbriate; ventrally with a broad medial concavity approaching
apex. Spiculum gastrale not observed. Tegmen evenly broadly rounded apically (Fig. 97),
much longer than wide (w:l ~ 1:3), lateral row of setae visible from the median fossa around the
apex, inner row of hair absent, basal notch of phallobase only slightly concave. Median lobe
large and robust, ~0.5 the length of the tegmen, apical opening well-developed extending
posteriorly (Fig. 138). Ejaculatory rods not observed.
Variation. None documented, only known from male Holotype.
Seasonality/Habitat. The holotype, and only existing specimen of the species, was
collected in September in a wet tropical forest.
Distribution. Known only from the type locality.
Notes. No host records are available for this species. Due to the attachment of the
genitalia directly to the specimen card mount with a water insoluble adhesive, some characters of
the “aedeagus” were not readily visible. No attempts were made to remove the genitalia from
the card mount due to the fragility of the organ and the extensive amount of adhesive present.
Pocadius dimidiatus Jelínek
(Figs. 18, 57, 98, 139, 179, 215)
Type Material Examined. PARATYPE ♂ (NMP): ARGENTINA: prov. Entre Rios;
Conocordia; Hayward lgt.
140
Non-Type Material Examined. >20 specimens with the following data labels:
ARGENTINA: Salta Prov.; 45km W Salta, 1950m; 1-29-XII-1987, El Alisal; S&J Peck, moist
ravine thicket, malaise FIT (CMN). ARGENTINA: Salta Prov.; El Rey Nat. Park, 900m; 11-15-
XII-1987, S&J Peck; Aguas Negras Trail, Prosopis forest, malaise-FIT (CMN). ARGENTINA:
Salta Prov.; 17km N La Caldera, Alto de la Sierra; 1550m, 2-30-XII-1987; S&J Peck, malaise-
FIT; subtropical humid forest (CMN). ARGENTINA: Salta Prov.; El Rey Nat. Park, 900m; Rio
La Sala, S&J Peck; 5-15-XII-1987, malaise-FIT; humid mossy Chaco forest.
Diagnosis. Jelínek (1977) noted that specimens of this species were in the museum at
LaPlata under the name P. helvolus, the eastern North American species, which was likely the
basis of Blackwelder’s (1945) LSAMount of P. helvolus in Argentina. The species is easily
distinguished from any North American species by the greatly enlarged terminal antennomere as
noted by Jelínek . However, due to the presence of the enlarged terminal antennomere in other
Neotropical species, there is a necessity for a more detailed diagnosis using both external and
genitalic characters. Pocadius dimidiatus has the following characters delimiting the species:
proportionately elongate pronotal and elytral fimbriae and overall long dorsal pubescence,
pronotum with trapezoidal anterior margin and simple almost truncate posterior margin,
prosternal process with prolonged apex, proportionately elongate tegmen to body size of adult,
large robust median lobe, ejaculatory rods fused medially, and both the lateral and inner rows of
setae on the tegmen are incomplete.
Redescription. Length 3.2 mm, Width 1.8mm, Depth 1.4mm. Body moderately
convex, surface shining, reddish-brown to dark brown in color, most specimens with elytral
apices and lateral regions dark brown. Pronotum and elytra margins with elongate fimbriae,
setae much longer than width of antennal scape. Dorsal and ventral pubescence golden and quite
long.
141
Head surface deeply, irregularly, densely punctate, punctures larger on vertex, becoming
somewhat smaller towards fronotclypeal region. Larger punctures 4-5 X’s diameter of eye facet,
smaller punctures 3-4 X’s diameter of facet. Interspaces smooth to finely alutaceous, shining.
Each smaller puncture gives rise to an elongate curved golden seta. Eyes finely faceted.
Pronotal surface with large punctures equal to slightly larger in size to large punctures on vertex
of head, interspersed with relatively few smaller punctures, ~0.75 size of larger ones.
Interspaces smooth to alutaceous, about 0.5-1 diameters apart. Each puncture gives rise to an
elongate golden seta, most seta are slightly curved. Pronotal fimbria elongate, longer than length
of scape. Scutellar surface with shallowly impressed smaller punctures, somewhat smaller than
small punctures on pronotum, interspaces smooth, no punctures in apical 0.25. Elytral surface
with serial rows of alternating large and small deep punctures. Smaller punctures are equal in
size to smaller punctures on pronotum, larger punctures are ~2 times diameter of smaller ones.
Smaller punctures giving rise to an erect elongate curved golden seta, larger punctures giving
rise to a shorter decumbent golden seta. Interspaces more narrow between punctures of a given
row than those of different rows. Within a row, small punctures are separated by ~1 puncture
width, and large punctures by ~1 puncture width. Rows are separated by 1.0 large puncture
width. Interspaces always shining but variable from alutaceous to finely microreticulate in
sculpture. Elytral fimbria elongate, setae about 1.25X length of pronotal fimbria. Pygidium
densely punctate, punctures equal in size to smaller punctures on pronotum, each puncture giving
rise to a short stiff golden seta. Interspaces narrow, 0.7-1.2 diameters, with alutaceous to
granular sculpture.
Venter less densely setose surface as dorsum, however, setae still elongate like those on
dorsum. Mentum with large very faint punctures, equal in size to large punctures on vertex,
some but not all of punctures giving rise to moderately long golden seta, interspaces smooth to
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alutaceous. Submentum and gula much less densely punctate than mentum with interspaces
smooth to granular and more than 3 puncture diameters apart. Prosternum and epimeron faintly
irregularly punctate, punctures slightly larger than those on mentum, interspaces alutaceous to
rugulose, prosternal punctures separated by 0.5 - 1 diameter, those on the epimeron by >1
diameter. Punctures on prosternal process giving rise to elongate golden setae. Mesosternum
with shallow punctures, about 1.25 diameter of those on prosternum, interspaces alutaceous to
granular, separated by about 0.25-0.5diameter. Metasternum irregularly punctate, with large
punctures laterally, becoming slightly smaller and more faintly impressed on disc, lateral
punctures similar in size to those on mesosternum, interspaces smooth on metasternal disc
becoming microreticulate to granular laterally, punctures separated by ~1 diameter. Abdominal
sternite 1 with large faint, almost obsolete punctures, punctures equal to smaller punctures on
metasternal disc, interspaces alutaceous, separated by ~1-1.5 diameter. Abdominal sternites 2-4
with two irregular rows of punctures, one row near anterior margin and the other near posterior
margin, punctures similar in size to larger punctures on metasternum, rows separated by 0.5
puncture diameter, punctures within rows separated by ~0.5 punctures width. Hypopygidium
with moderately deep punctures, similar in size to those on sternites 2-4, interspaces alutaceous
to granular, punctures separated by 0.5 puncture diameter.
Head much wider than long (L:W = 1:1.5), fronotclypeal region moderately projecting
anteriorly. Vertex with shallow concavity between orbits near fronotclypeal region. Labrum
with deep medial incision at anterior margin. Antennal club compact, obovate, asymmetrical
with the last antennomere greater in length to the previous two combined. Antennomeres 6-8
characteristically disc-like. Antennal scape asymmetrical, slightly hemispherical, 1.9 times as
long as pedicel. Pedicel subcylindrical in shape. Antennal segment 3 slightly longer in length
than pedicel. Antennal club large, ~0.75 length of segments 1-8 combined. Each club segment
143
with dense short setae, and a small dense sensillar region near the anterior border of the ventral
surface of antennomeres 11. Antennal grooves very deep and widely excavate, slightly
converging posteriorly. Lateral mental channel prominent and widely excavated, at level of
submentum, densely punctate with small punctures each giving rise to curved golden setae.
Mentum with anterior angles almost obsolete, anterior margin angulate, entire structure
somewhat pentagonal and convex ventrally.
Pronotum widest near middle (L:W = 1:2), anterior margin trapezoidal, posterior
margin simple, lateral margins less arcuate posteriorly, anterior and posterior angles present.
Scutellum large, elongate triangular, apex narrowly rounded. Prosternal process somewhat
narrowed between procoxae, apex acuminate, in lateral aspect the anterior and posterior ends are
prominent and slightly convex medially. Posterior apical wall prominent and only slightly
oblique. Mesosternum extending to midway between mesocoxae, sharply concave for reception
of the metasternum. Metasternum width to length ratio is ~1.6:1. Metepisternum with slight
medial concavity, oblique line dividing anterior 1/8 of structure. Elytral humeri moderately
produced, lateral margin very narrow. First abdominal sternite with acuminate process between
metacoxae. First sternite ~2X’s longer than second sternite. Sternites 2-3 subequal in length, the
fourth slightly larger than the preceding two. Hypopygidium subequal in length to first
abdominal sternite.
Protibia with apical tooth not prominent, slightly longer than tarsomere 1. Inner apical
spine subequal in length to tarsomeres 1 and half of 2 combined. Outer apical notch absent.
Protibia heavily armored with characteristic dense patch of stiff setae along the inner apical
region. Mesotibia more heavily armored than protibia with more dense stiff setae and a row of
numerous slender spines along entire lateral edge. Outer apical process somewhat developed,
larger than protibia process. Inner apical spine equal in length to tarsomeres 1-2 combined.
144
Metatibia with more developed armature than that of mesotibia, outer apical process well-
developed.
Male genitalia well-sclerotized. Anal sclerite with large broadly curved region
anteriodorsally (Fig. 18); apex sparsely fimbriate; ventrally with a broad medial concavity
approaching apex. Spiculum gastrale with wide lateral flanges, medial margins concave
proximally, short stiff setae originating from angulate apical margin (Fig. 57). Tegmen evenly
rounded apically (Fig. 98), much longer than wide (w:l = 1.0:2.1), lateral row of setae visible
from the median fossa to around the apex, small shallow concavities in apical third, basal notch
perpendicular, basal margin concave. Median lobe elongate, ~0.75 the length of the tegmen,
apex acuminate, apical opening well-developed (Fig. 139). Ejaculatory rods not fused to basal
piece, but fused to each other in apical third, concave medially. Basal piece of rods in three
pieces, two lateral asymmetrical tear-drop pieces, and a medial bilobed globular section (Fig.
179).
Female genitalia well sclerotized. Paraprocts large with sclerotization only along median
line to baso-lateral angles. Gonocoxite with two basal lateral prominences, basal ridge well-
sclerotized with two short medial baculi. Gonocoxal apices with recurved “tooth” absent. Three
setae originate from small depressions on the lateral region of the gonocoxal apices (Fig. 215).
Variation. Some variability exists in the amount of dark coloration on the lateral and
apical regions of the elytra.
Seasonality/Habitat. The type and non-type specimens were all collected during
December. The species is found in both lowland and mid- to high elevation wet forests.
Distribution. The type series is from eastern Argentina and the non-type material from
northwestern Argentina. Leschen and Carlton (1994) mistakenly identified specimens of the
new species P. peruensis as members of this species, indicating these individuals denoted a range
145
extension for P. dimidiatus. This is not surprising as like other New World Pocadius this species
has the elytral apices darker in color than the rest of the body. However, following dissection of
the Peruvian material, the specimens were determined as undescribed. Thus, this species does
appear to be restricted to north and central Argentina.
Notes. No host records exist for this species.
Pocadius dominicus Cline new species
(Figs. 19, 58, 99, 140, 180, 216)
Type Material. HOLOTYPE ♂ (MTEC): DOM. REP.: PR. Hato Mayor; Par. Nac. Los
Haitises; 02 JULY 1992 – 16 JULY 1993; D. Sikes & R. Rosenfield; flight intercept trap /
HOLOTYPE; Pocadius; dominicus; A.R. Cline des. 2004. Holotype with genitalia dissected and
contained under specimen in genitalia vial. PARATYPE (MTEC): DOMINICAN REPUBLIC:
Prov. Hato Mayor; Par. Nac. Los Hatises; 22-31 JULY 1993, FIT; D. Sikes & R. Rosenfield.
PARATYPE (MTEC): DOMINIC. REP.: Prov Hato; Mayor, P.N. Los Hatises; JULY 1992 - 16
JULY 1993; flight intercept trap; D.S. Sikes colr. PARATYPE (LSAM): DOM. REP.: Prov
Hato Mayor; Par. Nac. Los Hatises; 16-APR-01-JULY-1992; FIT #1, bosque humido; M. Ivie,
D. Sikes, Lanier. PARATYPE (CMN): DOM. REP.: LaVega Prov.; 10km NE Jaraboca; Raquet
Club, 550m, FIT; 20-VII-4-VIII-1995, mixed; for., S&J Peck, 95-37. PARATYPE (CMN) 3
specimens: DOM. REP.: LaVega Prov.; PN. A. Bermudez Cienaga; 19-VII-2-VIII-1995, 1100m;
trop. evgrn. for., FIT; S&J Peck, 95-36. PARATYPE (LSAM) 2 specimens: DOM. REP.:
LaVega Prov.; PN. A. Bermudez Cienaga; 19-VII-2-VIII-1995, 1100m; trop.evgrn.for., FIT;
S&J Peck, 95-36. PARATYPE (ARC) 1 specimen: DOMINICAN REPUBLIC: La; Vega,
Cordillera Central; 4.1km SW El Convento; 18-50-37N 70-42-48W; 1730m, 31 May 2003 / J.
Rawlins, R. Davidson; C. Young, C. Nunez, P.; Acevedo, dense secondary; evergreen forest
146
with; pine, hand collected; Sample 22242 / Carnegie Museum; Specimen Number; CMNH-
347,548. PARATYPE (LSAM) 1 specimen: label data same as above but specimen number is
CMNH-347,284. 12 PARATYPES (CMNH): same as above but specimen numbers as follows:
CMNH-347,410, CMNH-347,617, CMNH-347,448, CMNH-347,520, CMNH-347,842, CMNH-
347,354, CMNH-347,247, CMNH-347,280, CMNH-347,843, CMNH-347,881, CMNH-347,891,
and CMNH-347,238.
Diagnosis. This species is easily distinguished from any other Neotropical Pocadius by
the following characters: body coloration dull orange-yellow with well-demarcated apical 0.66 of
elytra black, nearly triangular mentum, male hypopygidium with central asetose region, terminal
antennomere with distinct circular sensillar region, densely setose tegmen with inner setal row
attaining apex in convergent manner, and lateral pores on gonocoxites.
Description. Length 4.0 mm, Width 1.9mm, Depth 0.9mm. Body slightly convex,
surface shining, orange-yellow in color with antennal club somewhat darker and apical 0.66
well-demarcated black. Pronotal and elytral margins with dense moderately elongate fimbriae.
Dorsal and ventral pubescence moderately elongate. Head, pronotum and elytra densely
punctate with surface rugulose.
Head surface densely irregularly punctate, with both large and small punctures evenly
dispersed across vertex. Larger punctures 3-4 X diameter of eye facet, smaller punctures ~1.5 X
diameter. Interspaces granular, moderately shining. Each smaller puncture gives rise to an
elongate curved golden seta. Pronotal surface with large punctures equal in size to large
punctures on vertex of head, interspersed with relatively few smaller punctures, approximately
2X diameter small punctures on vertex. Interspaces finely granular, about 0.25-0.5 diameter
apart. Each puncture gives rise to a moderately long golden seta, seta curved and decumbent.
Scutellar surface with more shallowly impressed small punctures, similar in size to small
147
punctures on pronotum, some punctures giving rise to setae, interspaces granular. Elytral surface
with serial rows of alternating large and small moderately impressed punctures. Smaller
punctures equal in size to smaller punctures on pronotum, larger punctures are ~4 times diameter
of smaller ones. Smaller punctures giving rise to a semi-erect moderately long golden seta,
larger punctures giving rise to a short decumbent golden seta. Interspaces narrow between
punctures of a given row and between different rows. Within a row, small punctures are
separated by ~1 puncture diameter, and large punctures by 0.25-0.5 puncture diameter. Larger
rows are separated by 0.5-1 large puncture diameter. Interspaces somewhat shining but granular
in surface sculpture. Pygidium somewhat densely punctate, punctures equal in size to larger
punctures on pronotum, each puncture giving rise to a rather short stiff golden seta. Interspaces
narrow, 0.75 diameter, with granular surface sculpture.
Venter with less dense shorter golden pubescence than dorsum. Mentum with faintly
impressed punctures, equal in size to larger punctures on vertex, each giving rise to a short
straight seta, interspaces mostly alutaceous. Submentum and gula with similar punctation as
mentum but with interspaces more granular. Prosternum and epimeron more regularly punctate,
punctures equal to large punctures on vertex, interspaces granular with microreticulate areas,
prosternal punctures separated by 0.5-1.0 diameter, those on the epimeron by 0.5 diameter.
Mesosternum with moderately impressed punctures, congregated along posterior margin only,
interspaces granular, separated by approximately 1 diameter. Metasternum irregularly punctate,
with a few smaller more faint punctures on disc similar in size to those on mesosternum, lateral
punctures larger, interspaces granular, punctures separated by ~1 diameter laterally. Abdominal
sternite 1 with somewhat faintly impressed punctures, punctures equal to those on lateral region
of metasternum, interspaces granular, separated by ~1 diameter. Abdominal sternites 2-4 with
irregular rows of punctures, one distinct row near posterior margin, punctures similar in size to
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those on metasternum, punctures separated by ~0.75-1 puncture diameter. Hypopygidium with
deep punctures, similar in size to those on sternites 2-4, interspaces granular, punctures separated
by 0.25-0.5 puncture diameter, central region of hypopygidium with only a few setal bearing
punctures present.
Head wider than long (W:L = 1.33:1), fronotclypeal region moderately projecting
anteriorly. Vertex with deep indistinct concavity between orbits near fronotclypeal region.
Labrum with deep medial incision at anterior margin. Antennal club compact, obovate,
asymmetrical with the last antennomere greater in length than the previous two combined.
Antennomeres 6-8 flattened. Antennal scape asymmetrical, distinctly hemispherical, 1.66 times
as long as pedicel. Pedicel subcylindrical in shape, somewhat swollen anteriorly. Antennal
segment 3 shorter in length than pedicel. Antennal club moderately large, ~0.6 length of
segments 1-8 combined. Each club segment with dense short setae, and several protruding
elongate setae. Antennal grooves deeply excavate, slightly converging posteriorly. Lateral
mental ridge prominent and elongate, at level of submentum, ridge with longitudinal
microreticulations laterally and medially granular surface. Mentum with anterior angles absent,
anterior margins angulate and coming together in acuminate apex, overall triangular in
appearance, entire structure with slight convexity.
Pronotum widest at middle (L:W = 1:2), anterior margin distinctly trapezoidal,
posterior margin highly convex, lateral margins less arcuate posteriorly, anterior and posterior
angles distinct. Scutellum large, hemispherical, apex broadly rounded. Prosternal process
greatly narrowed between procoxae, apex acuminate, in lateral aspect the anterior and posterior
ends are not prominent with moderate convexity medially over procoxae. Posterior apical wall
prominent and perpendicular. Mesosternum extending to approximately midway between
mesocoxae, broadly concave for reception of the metasternum. Metasternum much wider than
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long (W:L = 3:1). Metepisternum with broad medial concavity, oblique line dividing anterior
0.16 of structure into axillary space. First abdominal sternite with acuminate process between
metacoxae. First sternite ~1.8 X’s longer than second sternite. Sternites 2-3 subequal in length,
the fourth slightly larger than the preceding two. Hypopygidium subequal in length to first
abdominal sternite.
Protibia with apical tooth moderately prominent, approximately equal to length of first
tarsomere Outer apical notch with ~100° angle, notch depth moderately deep, equal to length of
tarsomere 1 and part of 2. Inner apical spine subequal in length to tarsomere 1 and part of 2.
Protibia moderately armored with elongate stiff setae on the lateral margin and with
characteristic dense patch of stiff setae along the inner apical region. Mesotibia more heavily
armored than protibia with more dense stiff setae and a row of numerous slender spines along
entire lateral edge. Outer apical process distinct, larger than protibia process. Inner apical spine
equal in length to tarsomeres 1. Metatibia more heavily armored than mesotibia with longer
more densely distributed stiff setae, and more distinct outer apical tooth and inner apical spine,
the latter of these is subequal in length to tarsomeres 1 and 2 combined.
Male genitalia well-sclerotized. Anal sclerite with broadly curved region
anteriodorsally (Fig. 19); apex with elongate fimbria; ventrally with a more broad medial
concavity approaching apex. Spiculum gastrale with broad rounded lateral flanges, medial
margin approximate, numerous short stiff setae originating from apex (Fig. 58). Tegmen evenly
rounded apically (Fig. 99), much longer than wide (w:l = 1.0:2.2), lateral row of setae visible
from the median fossa around the apex, no concavity in apical third, basal notch oriented
perpendicular, inner row of setae attaining apex in triangular fashion. Median lobe becoming
narrow apically, ~0.4 the length of the tegmen, apex acuminate, apical opening moderately well-
developed (Fig. 140). Ejaculatory rods not fused to basal piece or to each other, more or less
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straight without medial concavity. Basal piece of rods fused basally with elongate paired apical
projections (Fig. 180).
Female genitalia well sclerotized. Paraprocts with sclerotization along median border
only. Gonocoxite with two basal lateral prominences, basal ridge well-sclerotized. Gonocoxal
apices with recurved “tooth” not well-developed. Several elongate setae originate from small
depressions on lateral margin of the gonocoxal apices. Numerous pores present laterally along
gonocoxites (Fig. 216).
Variation. No demonstrable variation exists except for the denuded asetose central
region visible only on the male hypopygidium.
Seasonality/Habitat. Though some FIT’s were serviced year-round, specimens appear
most likely to be collected from mid-April through August as evidenced by other less long-term
FIT sampling protocols.
Distribution. Known only from the Dominican Republic on the island of Hispaniola.
Notes. No host data available for this species.
Etymology. Specific epithet is a derivative of the type locality, i.e. the Dominican
Republic.
Pocadius endroedyi Cline new species
(Figs. 20, 59, 100, 141, 181, 217)
Type Material. HOLOTYPE ♂ (TMSA): S. Afr., Transvaal; Pretoria, Waterkloof;
25.43S – 28.11E / 11-3-1993, E-Y: 2879; ex puffballs; leg. Endrödy-Younga. 27 PARATYPE
(TMSA): same data label as holotype. 1 PARATYPE (BMNH): same data label as holotype. 5
PARATYPE (LSAM): same data label as holotype. ! PARATYPE (LSAM): TANGANYIKA
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Terr.; Ukerewe Island; leg. Father Conrad. 1 PARATYPE (LSAM): Sartruggens; Marico; Dr.
Brauns / Transvaal; 15-I-1921.
Diagnosis. This species is well distinguished from P. africanus by the following
attributes: more robust overall body size, more elongate antennal club with different sensillar
region on terminal antennomere, much smaller axillary space on metepisternum, anterior
pronotal margin broadly evenly concave, head and pronotum much more densely punctate with
smooth interspaces, prosternal process with posterior apical wall in lateral aspect with distinct
concavity, ejaculatory rods not fused basally, more narrowed elongate median lobe of aedeagus,
inner row of setae on tegmen not complete apically, lateral row of setae on tegmen with elongate
central tuft, eighth abdominal sternite with apical margin with crenulations, and numerous setae
originating from each gonocoxal apex.
Redescription. Length 3.9 mm, Width 1.8mm, Depth 1.3mm. Body moderately
convex, surface shining, reddish-brown to brown in color. Pronotal and elytral margins with
sparse moderately elongate fimbriae. Dorsal and ventral pubescence not elongate. Head,
pronotum and elytra densely deeply punctate.
Head surface deeply, irregularly punctate, with both large and small punctures evenly
dispersed across vertex. Larger punctures 3-4 X diameter of eye facet, smaller punctures 1-2 X
diameter. Interspaces smooth to finely alutaceous, shining. Each smaller puncture gives rise to
an elongate curved golden seta. Pronotal surface with large punctures equal in size to large
punctures on vertex of head, interspersed with relatively few smaller punctures, similar in size to
small punctures on vertex. Interspaces finely microreticulate to granular, about 0.5-1 diameter
apart. Each puncture gives rise to a moderately long golden seta, most seta curved and
decumbent. Scutellar surface with few vague shallowly impressed punctures, some punctures
giving rise to setae, interspaces alutaceous to granular. Elytral surface with serial rows of
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alternating large and small deep punctures. Smaller punctures are equal in size to smaller
punctures on pronotum, larger punctures are ~3-4 times diameter of smaller ones. Smaller
punctures giving rise to a semi-decumbent moderately long golden seta, larger punctures giving
rise to a short almost adpressed golden seta. Interspaces narrow between punctures of a given
row and between different rows. Within a row, small punctures are separated by ~1-1.5 puncture
diameters, and large punctures by 0.33-0.5 puncture diameter. Rows are separated by 0.5 large
puncture diameter. Interspaces somewhat shining but granular in surface sculpture. Pygidium
densely punctate, punctures equal in size to larger punctures on pronotum, each puncture giving
rise to a rather short golden seta. Interspaces narrow, 0.5-1 diameter, with alutaceous to granular
surface sculpture.
Venter with similar moderately long golden pubescence as dorsum. Mentum with small
faintly impressed punctures, equal in size to smaller punctures on vertex, each giving rise to a
short straight seta, interspaces alutaceous to somewhat granular. Submentum and gula with
similar punctation as mentum but with interspaces more granular. Prosternum and epimeron
deeply regularly punctate, punctures equal to large punctures on vertex, interspaces granular with
microreticulate areas, prosternal punctures separated by 0.25-0.5 diameter, those on the epimeron
by 0.25 diameter. Mesosternum with moderately impressed punctures, congregated along
posterior margin, interspaces alutaceous to granular, separated by about 0.5-1 diameter.
Metasternum deeply irregularly punctate, with smaller more faint punctures on disc similar in
size to those on mesosternum with lateral punctures slightly larger, interspaces alutaceous on
metasternal disc becoming granular laterally, punctures separated by ~1-2 diameters. Abdominal
sternite 1 with large somewhat faintly impressed punctures, punctures equal to those on
metasternum, interspaces alutaceous to granular, separated by ~1-2 diameters. Abdominal
sternites 2-4 with irregular rows of punctures, one distinct row near posterior margin, punctures
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similar in size to those on metasternum, punctures separated by ~0.5-0.75 puncture diameter.
Hypopygidium with deep punctures, similar in size to those on sternites 2-4, interspaces
granular, punctures separated by 0.25-0.5 puncture diameter.
Head wider than long (W:L = 1.4:1), fronotclypeal region moderately projecting
anteriorly. Vertex with small but indistinct concavity between orbits near fronotclypeal region.
Eyes large, moderately protruding. No prominence over antennal insertion. Labrum with small
shallow concavity at anterior margin. Antennal club compact, elongate oval, more or less
symmetrical with the last antennomere subequal to the previous two combined. Antennomeres
6-8 more or less compact, with 7-8 characteristically disc-like. Antennal scape asymmetrical,
distinctly hemispherical, 1.5 times as long as pedicel. Pedicel subcylindrical in shape. Antennal
segment 3 shorter in length to pedicel. Antennal club large, ~0.75 length of segments 1-8
combined. Each club segment with dense short setae, and only relatively few protruding
elongate setae (2-4 in number). Antennal grooves very deep and moderately excavate, slightly
converging posteriorly. Lateral mental ridge prominent, at level of submentum, ridge with
longitudinal microreticulations laterally and medially is granular. Mentum with anterior angles
obsolete, anterior margin broadly hemispherical, entire structure flattened with no convexity.
Pronotum widest in posterior 0.33 (L:W = 1:1.8), anterior margin broadly evenly
concave, posterior margin convex, lateral margins less arcuate posteriorly, anterior and posterior
angles distinct. Scutellum moderately large, triangular, apex acutely rounded. Prosternal
process narrowed between procoxae, apex somewhat acuminate, in lateral aspect the anterior and
posterior ends are prominent with little convexity medially over procoxae. Posterior apical wall
prominent with distinct concavity. Mesosternum extending to approximately midway between
mesocoxae, deeply incised for reception of the metasternum. Metasternum much wider than
long (W:L = 2.9:1.0). Metepisternum with slight medial concavity, curved oblique line dividing
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anterior 1/7 of structure. Elytral humeri moderately produced, lateral margin very narrow. First
abdominal sternite with broad truncate process between metacoxae. First sternite ~2X’s longer
than second sternite. Sternites 2-3 subequal in length, the fourth slightly larger than the
preceding two. Hypopygidium subequal in length to first abdominal sternite.
Protibia with apical tooth not prominent, approximately 0.5 length of first tarsomere
Outer apical notch with ~90° angle, notch depth moderate, equal to 0.75 length of tarsomere 1.
Inner apical spine subequal in length to tarsomeres 1. Protibia not heavily armored but with
characteristic dense patch of stiff setae along the inner apical region. Mesotibia more heavily
armored than protibia with more dense stiff setae and a row of numerous slender spines along
entire lateral edge. Outer apical process only faintly distinct, only slightly larger than protibia
process. Inner apical spine equal in length to tarsomeres 1. Metatibia with armature similar to
that of mesotibia, but with longer more distinct outer apical tooth and inner apical spine.
Male genitalia well-sclerotized. Anal sclerite with narrowly curved region
anteriodorsally (Fig. 20); apex densely fimbriate; ventrally with a more narrow medial concavity
approaching apex; posterior apex with well-defined acuminate apex. Spiculum gastrale with
angulate lateral flanges, medial margin biconcave, numerous short stiff setae originating from
apex, apical margin with crenulations near edge (Fig. 59). Tegmen evenly rounded apically
(Fig. 100), much longer than wide (w:l = 1.0:3.0), lateral row of setae visible from the median
fossa around the apex, narrow elongate shallow concavity in apical third, basal notch oriented
somewhat posteriorly, inner row of setae not attaining apex and expanded basally. Median lobe
narrow, ~0.5 the length of the tegmen, apex acuminate, apical opening not well-developed (Fig.
141). Ejaculatory rods not fused to basal piece or to each other. Basal piece of rods with deep
medial projection extending almost half length of ejaculatory rods, proximally with swollen base
(Fig. 181).
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Female genitalia moderately sclerotized. Paraprocts relatively small compared to
gonocoxites, with sclerotization along median line to midway to baso-lateral angles. Gonocoxite
with two basal lateral prominences, basal ridge well-sclerotized. Gonocoxal apices with
recurved “tooth” absent. Several setae originate from small depressions on lateral margin of the
gonocoxal apices (Fig. 217).
Variation. The single specimen from Tanzania is slightly lighter in color than those
from Zimbabwe and South Africa.
Seasonality/Habitat. Specimens were collected from December through March
Distribution. The species range extends south from the Tanzanian island of Ukerewe
which lies in the southern reaches of Lake Victoria through Salisbury, Zimbabwe to the
northeastern region of South Africa.
Notes. Specimens from South Africa and Zimbabwe were collected from puffballs.
Etymology. Specific epithet honors Sebastian Endrödy-Younga, former nitidulid
specialist, for his work on the fauna of Africa.
Pocadius ephite Leschen and Carlton
(Figs. 21, 60, 101, 142, 182)
Type Material Examined. HOLOTYPE ♂ (SNEC): COSTA RICA: Alajuela; Peñas
Blancas 875m; 19 May 1989, J. Ashe,; R. Brooks, R. Leschen; ex., flight intercept / Snow
Entomol. Mus.; Costa Rica Exped. #267 / HOLOTYPE; Pocadius ephite; R. Leschen & C.
Carlton.
Non-Type Material Examined. None.
Diagnosis. The original description of this species (Leschen and Carlton 1994)
suggested similarities with P. fumatus from South America, with deviations in elytral vestiture
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and protibial armature. The authors also briefly diagnosed P. ephite from the two other species
described in the paper, i.e. P. jelineki and P. maquipucunensis, by “the structure of the rods in
the ejaculatory duct of the aedeagus, protibial morphology, and vestiture without curved ends.”
This diagnosis must be modified as other Pocadius are known to have vestiture without curved
ends and details of the structure/morphology of the protibia and ejaculatory rods were not fully
discussed. Pocadius ephite is one of the more oval shaped members, with an extremely low L:W
ratio, making initial identification moderately successful. Also, the large robust very
asymmetrical antennal club, in particular the terminal antennomere, the evenly rounded anterior
margin of the pronotum, the rounded apex of the abdominal process, as well as the large
umbilicate punctures with smooth shining interspaces on the metasternal disc help to externally
define this species. These characters, in combination with the apical fossa of the tegmen
attaining the tegminal apex, readily identify P. ephite from the other Neotropical members of the
genus.
Redescription. Length 2.9mm, Width 2.1mm, Depth 1.2mm. Body moderately convex,
surface shining, light brown to reddish brown. Pronotum and elytra with moderately long
fimbriae, setae ~1.5 width of antennal scape. Dorsal and ventral pubescence moderate in length,
fine and relatively sparsely distributed.
Head surface irregularly punctate, punctures larger on vertex, becoming somewhat
smaller laterally and posteriorly, and minute punctures on clypeal-labral region. Large punctures
equal to ~4 eye facet diameters, smaller punctures ~3 diameters, and minute punctures ~1
diameter. Interspaces smooth to alutaceous, shining except on clypeal-labral region where the
interspaces are alutaceous to granular. Each vertexal puncture gives rise to a single fine short
seta. Eyes finely faceted. Pronotal surface with large punctures about 1.3X’s diameter of large
punctures on vertex of head, interspersed with smaller punctures, equal to size of large vertex
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punctures. Interspaces smooth to finely alutaceous, shining, about 0.25-0.5 diameter apart. Each
puncture gives rise to a fine golden seta that is either semi-erect or erect. Scutellar surface with
very few vague shallowly impressed large punctures, some punctures giving rise to setae, and the
interspaces are alutaceous, impunctate in apical third. Elytral surface with serial rows of
alternating large and small deep punctures. Punctures equal in size to corresponding large and
small punctures on pronotum. Smaller punctures giving rise to an erect golden seta, larger
punctures giving rise to a semi-erect golden seta. Interspaces widely spaced between punctures
of a given row and between different rows. Within a row, small punctures are separated by ~1-
1.5 puncture width, and large punctures by 0.75-1 puncture width. Rows are separated by ~1
large puncture width. Interspaces shining but variable from smooth to finely alutaceous.
Pygidium densely punctate, punctures equal in size to larger ones on pronotum, each puncture
giving rise to a short stiff golden seta. Interspaces narrow, 0.25-0.5 diameters apart.
Venter with shorter more sparsely distributed pubescence than dorsum. Mentum with
shallow small punctures faintly visible, equal in size to small punctures on vertex. Interspaces
granular to alutaceous. Submentum similar in punctation to mentum but with interspaces
completely granular. Gula impunctate with alutaceous surface. Prosternum and epimeron
deeply irregularly punctate, punctures slightly larger than those on mentum, interspaces
alutaceous prosternal punctures separated by 0.5 diameter, those on the epimeron by 0.5-1
diameter. Mesosternum with large shallow punctures, about 2-3 diameter of those on
prosternum, interspaces alutaceous, separated by ~1 diameter. Metasternum irregularly punctate,
with large punctures on disc similar in size to those on mesosternum, interspaces alutaceous to
granular. Abdominal sternite 1 with faint punctures, punctures 0.5-0.75 diameter of metasternal
punctures, interspaces alutaceous, separated by ~1 diameter. Abdominal sternites 2-4 with 2-3
irregular rows of punctures, two rows near anterior margin and the other near posterior margin,
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rows separated by 1 puncture width, punctures within rows separated by ~0.25 punctures width.
Hypopygidium with moderately deep punctures, similar in size to those on sternites 2-4,
interspaces alutaceous, punctures separated by 1-2 puncture diameters, each puncture giving rise
to a moderately long golden seta.
Head much wider than long (W:L = 2:1), fronotclypeal region only slightly projecting
anteriorly. Vertex with concavity between orbits broad and deep. Labrum with moderately
broad concavity medially at anterior margin. Antennal club compact, globular, greatly
asymmetrical with the last antennomere much larger than the previous two segments combined.
Antennomeres 4-6 more or less compact and somewhat cuboidal, with 7-8 characteristically
flattened and disc-like. Antennal scape moderately asymmetrical, somewhat hemispherical, 2
times as long as pedicel. Pedicel subcylindrical in shape. Antennal segment 3 equal in length to
pedicel. Antennal club large, ~0.95 length of segments 1-8 combined. Antennal grooves
moderately deep and widely excavate, slightly converging posteriorly. Lateral mental ridge
prominent, at level of submentum, medially the ridge has granular sculpture with some faint
microreticulations. Mentum with anterior angles obsolete, anterior margin broadly
hemispherical with well defined medial apical tip, entire structure slightly convex.
Pronotum widest near middle (L:W = 1:1.8), anterior margin broadly shallowly
trapezoidal, posterior margin moderately convex, lateral margins less arcuate posteriorly.
Scutellum large, obtusely triangular, apex broadly rounded. Prosternal process somewhat
narrowed between procoxae, apex acuminate, in lateral aspect the anterior and posterior ends are
moderately prominent and only slightly convex medially. Posterior apical wall prominent and
slightly oblique. Mesosternum extending to midway between mesocoxae, evenly and somewhat
deeply concave for reception of the metasternum. Metasternum width to length ratio is ~2.5:1.0.
Metepisternum with medial margin angulate, oblique line dividing anterior 0.125 of structure.
159
Elytral humeri moderately produced, lateral margin very narrow. First abdominal sternite with
moderately narrow acuminate process between metacoxae. First sternite ~2X’s longer than
second sternite. Sternites 2-3 subequal in length, the fourth slightly larger than the preceding
two. Hypopygidium subequal in length to first abdominal sternite.
Protibia with apical tooth very prominent, equal in length to first tarsomere and half of
second combined. Outer apical notch with ~90° angle, notch depth deep, equal to length of
tarsomere 1 and part of two combined. Inner apical spine subequal in length to first tarsomere
and half of second combined. Protibia only somewhat armored but with characteristic patch of
stiff setae along the inner apical region. Mesotibia somewhat more heavily armored than
protibia with more dense stiff setae and a row of numerous slender spines along entire lateral
edge. Outer apical process elongate and robust, slightly larger than protibia process. Inner
apical spine equal in length to first tarsomere and half of second combined. Metatibia with
armature similar to that of mesotibia, but outer apical process and spines slightly longer.
Male genitalia well-sclerotized. Anal sclerite with broadly curved region
anteriodorsally (Fig. 21); apex somewhat fimbriate; ventrally with a broad elongate medial
concavity approaching apex. Spiculum gastrale with wide elongate lateral flanges, medial
margins concave, numerous short stiff setae originating from apex (Fig.60). Tegmen evenly
rounded apically (Fig. 101), longer than wide (w:l = 1.0:2.75), lateral row of setae visible from
the apex of the median fossa around the tegminal apex, large broad shallow concavity in apical
half, basal notch perpendicular, basal margin angulate with basal concavity. Median lobe
elongate oval, ~0.45 the length of the tegmen, apex broadly rounded, apical opening well-
developed with trilobed concavity (Fig. 142). Ejaculatory rods not fused to basal piece, nearly
straight with medially projecting apical prominence. Basal piece of rods fused with shallow
medial concavity extending one fourth length of structure, evenly rounded proximally (Fig. 182).
160
Female genitalia not observed.
Variation. Known only from holotype male.
Seasonality/Habitat. Type collected from mid-May in a wet forest in the Alajuela
Province of Costa Rica.
Distribution. Known only from type locality.
Notes. Holotype has abdomen completely dissected from body, also protarsi and
mesotarsi on left side missing. Two vials are under specimen, first contains abdomen, second
contains male genitalia except for anal sclerite and spiculum gastrale which are contained in the
first vial. No specimens outside of the type locality around Monteverde have been collected,
suggesting this species is a cloud forest endemic. This is common in Central and South
American Nitidulidae, as other genera (i.e. Stelidota, Epuraea, Psilotus (Cline 2004b)) also have
endemic species in mid- to high elevation cloud forests.
Pocadius falini Cline new species
(Figs. 22, 61, 102, 143, 183, 218)
Type Material Examined. HOLOTYPE ♂ (SNEC): PARAGUAY: Itapua; Yatai, prop.
Hostettler family ; San Rafael Reserve, 100m ; 26°38’17”S 55°39’50”W ;21-25 NOV 2000, Z.H.
Falin; PAR1F00 040; ex. flight intercept trap / SM0258682; KUNHM-ENT [barcode label] /
HOLOTYPE; Pocadius; falini; A.R. Cline des. 2004. 3 PARATYPES (SNEC): same data label
as holotype but with barcode # SM0258640, SM0564420, SM0564350and PARATYPE
designation labels.
Diagnosis. This species is similar to P. crypsis and P. coxus but can be differentiated
from them and the other Neotropical fauna by the following characters: male pygidium with
distinctly concave apical margin, uniformly dark reddish brown coloration with antennal club
161
darker, posterior prosternal wall perpendicular, anterior pronotal margin nearly truncate, elongate
elytral fimbriae, alternating erect and semi-erect rows of setae on elytra, mentum distinctly
pentagonal, pronotum with trapezoidal anterior margin, protibia with deep outer apical notch,
terminal antennomere with two elliptical sensillar regions, abdominal sternites 2-4 with three
organized rows of punctures, ejaculatory rods elongate and not split, basal piece of internal sac
sclerites with lateral arms wrapping around middle ovoid section, tegmen with two apical
depressions, and median lobe large and robust with acute apex and complex internal structure.
Description. Length 4.35mm, Width 2.7mm, Depth 2.0mm. Body large and robust,
convex, surface moderately shining, dark chestnut brown in color with venter somewhat lighter.
Pronotum and elytra margins with elongate fimbriae, setae longer than width of antennal scape.
Dorsal and ventral pubescence quite long.
Head surface densely, deeply, irregularly punctate, punctures larger on vertex, becoming
somewhat smaller towards orbits and fronotclypeal region. Larger punctures ~5X diameter of
eye facet, smaller punctures ~2X diameter; interspaces smooth to finely alutaceous, shining.
Each puncture gives rise to an elongate curved golden seta. Pronotal surface with large
punctures ~1.5 diameter of large punctures on vertex of head, interspersed with relatively few
smaller punctures, ~0.5 size of larger ones; interspaces alutaceous to finely microreticulate, about
0.75-1 diameter apart. Each puncture gives rise to a long, curved, golden seta. Scutellar surface
with very few vague shallowly impressed punctures, some punctures giving rise to setae, and the
interspaces are alutaceous and 3-4 diameters apart. Elytral surface with serial rows of alternating
large and small deep punctures, rows not clearly organized, rows of smaller punctures sometimes
with multiple smaller punctures instead of the typical single row. Smaller punctures minute, 0.5-
0.75 diameter of smaller ones on pronotum, larger punctures are ~2X diameter of smaller ones.
Smaller punctures giving rise to an erect long golden seta, larger punctures giving rise to a semi-
162
erect long golden seta; interspaces narrow between punctures of a given row and broad between
different rows. Within a row, small punctures are separated by ~1 puncture width, and large
punctures by 0.5-0.75 puncture diameter. Larger rows are separated by1-2 large puncture
diameters. Interspaces moderately shining but variable from alutaceous to finely granulate with
some microreticulations in sculpture. Pygidium densely punctate, punctures equal in size to
larger ones on pronotum, each puncture giving rise to a moderately long golden seta; interspaces
narrow, 0.25-0.5 diameter, with alutaceous to granular sculpture.
Venter with similar long golden pubescence as dorsum. Mentum with minute very
shallow punctures, equal in size to smaller ones on vertex, each giving rise to a golden seta.
Interspaces granular to finely microreticulate. Submentum and gula similar in punctation to
mentum but with interspaces more granular. Prosternum and epimeron deeply irregularly
punctate, punctures on epimeron equal to large ones on vertex those on prosternum slightly
smaller, interspaces alutaceous to granular with microreticulate areas, prosternal punctures
separated by 0.5-1 diameter, those on the epimeron by 0.25 to 0.5 diameter. Mesosternum with
shallow punctures, equal in size to large ones on vertex, interspaces alutaceous to granular,
separated by about 0.5 to 1 diameter, and aggregated near the metasternum. Metasternum
irregularly punctate, with moderately faint large punctures on disc 1.5 diameter of those on
mesosternum, interspaces alutaceous to granular on metasternal disc and similarly sculptured
laterally, punctures separated by ~0.5-1 diameter on disc. Abdominal sternite 1 with large
moderately faint punctures, punctures equal to those on metasternum, interspaces alutaceous to
granular, separated by ~1 diameter. Abdominal sternites 2-4 with three irregular rows of
punctures, one row near anterior margin another near posterior margin and midway between the
anterior and posterior margins, punctures similar in size to those on metasternum, rows separated
by 1 puncture diameter, punctures within rows separated by ~0.5 punctures width.
163
Hypopygidium with moderately deep punctures, similar in size to those on sternites 2-4,
interspaces granular, punctures separated by 0.5-1 puncture diameter.
Head slightly wider than long (W:L = 1.66:1), fronotclypeal region moderately projecting
anteriorly. Labrum with broad concavity at anterior margin. Antennal club compact, ovoid,
asymmetrical with the last antennomere longer than the previous two combined. Antennomeres
7-8 characteristically disc-like, 6 is trapezoidal, and 4-5 are somewhat cuboidal. Antennal scape
asymmetrical, somewhat hemispherical, 1.8 times as long as pedicel. Pedicel subcylindrical in
shape. Antennal segment 3 subequal in length to pedicel. Antennal club large, ~0.75 length of
segments 1-8 combined. Antennal grooves very deep and widely excavate, slightly converging
posteriorly. Lateral mental/submental ridge prominent, at level of submentum, ridge with
oblique and longitudinal microreticulations and sulcus with minute punctures and granulate
sculpturing. Mentum with anterior angles narrowly rounded, anterior margin angulate, entire
structure pentagonal when viewed ventrally and somewhat convex laterally.
Pronotum widest near middle (L:W = 1:1.77), anterior margin distinctly trapezoidal,
posterior margin moderately convex. Scutellum large, triangular, apex acute. Prosternal process
narrowed between procoxae, apex acuminate, in lateral aspect the posterior portion is 0.5 length
of anterior portion and there is a moderate convexity over the procoxae. Posterior apical wall
prominent and perpendicular. Mesosternum extending to midway between mesocoxae, evenly
concave for reception of the metasternum. Metasternum much wider than long (W:L = 3:1).
Metepisternum with slight medial constriction, oblique line dividing anterior 0.20 of structure.
First abdominal sternite with broad process between metacoxae. First sternite ~2X’s longer than
second sternite. Sternites 2-3 subequal in length, the fourth slightly larger than the preceding
two. Hypopygidium subequal in length to first abdominal sternite.
164
Protibia with apical tooth very prominent, slightly longer than tarsomere 1and part of 2
combined. Outer apical notch with ~90° angle, notch depth deep, equal to length of tarsomere 1
and part of 2 combined. Inner apical spine subequal in length to tarsomeres 1. Protibia not
heavily armored but with characteristic dense patch of stiff setae along the inner apical region.
Mesotibia more heavily armored than protibia with more dense stiff setae and a row of numerous
slender spines along entire lateral edge. Outer apical process moderately robust, somewhat
larger than protibia process. Inner apical spine equal in length to tarsomeres 1-2 and part of 3
combined. Metatibia more heavily armored than either protibia or mesotibia with longer and
more numerous setae and spines, inner apical spine slightly shorter than mesotibial spine.
Male genitalia well sclerotized. Anal sclerite with broadly curved region ventrally (Fig.
22); apex moderately fimbriate; apex with an acute apex. Spiculum gastrale with wide lateral
flanges, medial margins concave, numerous long stiff setae originating from apex (Fig. 61).
Tegmen evenly narrowly rounded apically (Fig. 102), much longer than wide (w:l = 1.0:3.1),
lateral row of setae visible from the apex of the median fossa around the tegminal apex, two
shallow concavities in apical half with numerous short stiff setae laterally but not extending
around apex, basal notch perpendicular, basal margin with slight concavity. Median lobe
elongate with slight medial constriction, ~0.5 the length of the tegmen, apex with acute point,
apical opening well-developed with inverted bilobed structure (Fig. 143). Ejaculatory rods
greatly elongate and not fused to basal piece, nearly straight with somewhat enlarged apex.
Basal piece of internal sac sclerites globular with lateral arms partially wrapping around central
region (Fig. 183).
Female genitalia moderately sclerotized. Paraprocts large with sclerotization only along
median line to apico-lateral angles. Gonocoxite with two basal lateral prominences, basal ridge
well-sclerotized. Gonocoxal apices with recurved “tooth” absent. Two primary setae originate
165
from small depressions on the gonocoxal apices. Intragonocoxal invagination extremely deep,
~0.75 length of gonocoxite (Fig. 218).
Variation. Some specimens are slightly lighter in color than the holotype.
Seasonality/Habitat. Specimens were collected in late November in a lowland tropical
forest.
Distribution. Known from the type locality in southern Paraguay.
Notes. No host information is available for this species.
Etymology. Specific epithet honors Zack Falin, collection manager of the Snow
Entomological Collection, for the collection of the specimens and his generosity during the
course of this study.
Pocadius femoralis Cline new species
(Figs. 219)
Type Material Examined. HOLOTYPE ♀ (SNEC): VIETNAM: Dong Nai; Cat Tien
National Park, near Park Hqtrs.; 11°25’23”N, 107°25’41”E, 120m; 27-31 MAY 1999, B.
Hubley, N. Tatmic; VIET1H95-99 039, ex. malaise trap / SM0189129; KUNHM-ENT [this is a
bar code label] / HOLOTYPE; Pocadius; femoralis; A.R. Cline des. 2004.
Diagnosis. This species is somewhat similar to P. decoratus and P. martini, which are
also known from SE Asia. This species can be easily delimited from all other Old World species
by the following suite of characters: small body size, unique color pattern including all femora
and tibia darkly colored, integument sparsely shining, scutellum not triangular, broad abdominal
process, metasternal disc with few minute punctures, outer protibial notch extremely deep,
prosternal process enlarged laterally posterior to procoxae, and ovipositor with greatly reduced
gonocoxal extensions that are present only as short extensions.
166
Description. Length 3.0mm, Width 2.0mm, Depth 1.2mm. Body moderately convex;
surface sparsely shining; head, prosternum, sides of pronotum, elytral apices and legs dark brown
with rest of body light red-brown. Pronotum and elytra margins with rather short fimbriae, setae
subequal in length to that of antennal scape. Dorsal and ventral pubescence moderately long and
fine.
Head surface densely, deeply, irregularly punctate, punctures larger on vertex, becoming
somewhat smaller towards orbits and fronotclypeal region, very few smaller punctures present
and not at all on vertex. Larger punctures 4-5X diameter of eye facet, smaller punctures 2X
diameter. Interspaces smooth to finely alutaceous, shining. Each puncture gives rise to a
moderately long curved golden seta. Pronotal surface with large punctures equal in size to large
punctures on vertex of head, interspersed with relatively few smaller punctures, ~0.33 size of
larger ones, smaller punctures closely adjacent to the anterior margin of the larger ones;
interspaces alutaceous, ~1-1.5 diameters apart. Most punctures give rise to a moderately long
curved golden seta. Scutellar surface with very few vague shallowly impressed small punctures,
equal to small ones on pronotum, some punctures giving rise to setae, interspaces alutaceous to
granular. Elytral surface with serial rows of alternating large and small deep punctures. Smaller
punctures are equal in size to smaller ones on pronotum, larger punctures are ~2.5 times diameter
of smaller ones. Smaller punctures giving rise to an erect long golden seta, larger punctures
giving rise to a shorter decumbent golden seta. Interspaces narrow between punctures of a given
row and between different rows. Within a row, small punctures are separated by ~1 puncture
diameter, and large punctures by 0.25 puncture diameter. Large rows are separated by 0.75-1.0
puncture diameter. Interspaces sparsely shining but variable from alutaceous to granular with
finely microreticulate regions. Pygidium densely punctate, punctures equal in size to large ones
167
on pronotum, each puncture giving rise to a moderately long golden seta; interspaces narrow,
0.25 diameters, with granular sculpture.
Venter with similar golden pubescence as dorsum. Mentum with minute shallow
punctures, equal in size to smaller ones on vertex, each giving rise to a seta; interspaces granular.
Submentum and gula with punctures 2X diameter of those on mentum and with interspaces
completely granular. Prosternum and epimeron deeply irregularly punctate, punctures equal to
larger ones on vertex on epimeron and slightly smaller on prosternum, interspaces alutaceous to
granular with some microreticulate areas, prosternal punctures separated by 0.5-1 diameter, those
on the epimeron by 0.25 diameter. Mesosternum with shallow punctures, equal to larger ones on
vertex, interspaces alutaceous to granular separated by about 0.5diameter and aggregated near
metasternum. Metasternum irregularly punctate, with moderate faint punctures on disc similar in
size to smaller ones on lateral region of head, interspaces smooth on metasternal disc becoming
microreticulate and granular laterally, punctures separated by ~1-2 diameters on disc and more
densely aggregated laterally. Abdominal sternite 1 with large faint, almost obsolete punctures,
punctures equal to large ones on vertex, interspaces alutaceous with numerous microreticulate
regions, separated by ~1 diameter. Abdominal sternites 2-4 with two irregular rows of large
punctures, one row near anterior margin and the other near posterior margin, punctures similar in
size to those on mesosternum, rows separated by 0.5 puncture diameter, punctures within rows
separated by ~0.25-0.5 punctures diameter. Rows on abdominal sternite 4 becoming less
organized. Hypopygidium with moderately deep punctures, similar in size to those on sternites
2-4, interspaces mostly granular, punctures separated by 0.25-0.5 puncture diameter.
Head wider than long (W:L = 1.5:1), fronotclypeal region moderately projecting
anteriorly. Labrum with narrow medial incision at anterior margin. Antennal club compact,
oval, slightly asymmetrical with the last antennomere slightly longer than the previous two
168
combined. Antennomeres 4-8 more or less compact, with 7-8 characteristically disc-like, and 4-
6 appearing trapezoidal. Antennal scape asymmetrical, distinctly hemispherical, 1.5 times as
long as pedicel. Pedicel barrel-like in shape. Antennal segment 3 equal in length to pedicel.
Antennal club large, ~0.90 length of segments 1-8 combined. Antennal grooves very deep and
widely excavate, slightly converging posteriorly. Lateral mental/submental ridge prominent, at
level of submentum, ridge with all oblique microreticulations and corresponding sulcus with
minute punctures and microreticulations. Mentum with anterior angles obsolete, anterior margin
broadly hemispherical, entire structure transversely hemispherical ventrally and laterally
somewhat convex.
Pronotum widest near posterior angles (L:W = 1:2), anterior margin broadly shallowly
concave, posterior margin moderately convex, lateral margins slightly arcuate to anterior angles.
Scutellum large, obtusely hemispherical, apical angle obsolete. Prosternal process somewhat
narrowed between procoxae, apex widely expanded posterior to procoxal cavities and somewhat
acuminate, in lateral aspect the structure is mostly flattened with a slight convexity over the
coxae. Posterior apical wall moderately prominent and perpendicular. Mesosternum extending
to midway between mesocoxae, evenly convexly shaped for reception of the metasternum.
Metasternum much wider than long (W:L = 3.2:1.0). Metepisternum with slight medial
constriction, oblique line dividing anterior 0.10 of structure. First abdominal sternite with broad
process between metacoxae. First sternite ~2X’s longer than second sternite. Sternites 2-3
subequal in length, the fourth slightly larger than the preceding two. Hypopygidium subequal in
length to first abdominal sternite.
Protibia with apical tooth prominent, slightly longer than tarsomeres 1 and part of 2
combined. Outer apical notch with ~85° angle, notch depth deep, equal to length of tarsomere 1-
2 combined. Inner apical spine short, subequal in length to tarsomere 1. Protibia not heavily
169
armored but with characteristic dense patch of stiff setae along the inner apical region.
Mesotibia more heavily armored than protibia with more dense stiff setae and a row of numerous
slender spines along entire lateral edge. Outer apical process moderately elongate and robust,
slightly larger than protibia process. Inner apical spine equal in length to tarsomeres 1-2
combined. Metatibia with armature similar to that of mesotibia, but outer apical process and
inner apical spine longer and more robust.
Male genitalia not observed.
Female genitalia moderately sclerotized (Fig. 218). Paraprocts elongate with
sclerotization only along median line and not extending to baso-lateral angles. Gonocoxite with
one basal lateral prominences, basal ridge well-sclerotized with two short oblique baculi
extending apically. Gonocoxal apices extremely reduced, present only as short extensions,
narrowly separated by short concavity. Two primary setae originate from small depressions near
the gonocoxal apices.
Variation. Known only from the single holotype female.
Seasonality/Habitat. Specimen collected from late May in the forested region of Dong
Nai.
Distribution. Known only from the type locality in southern Vietnam.
Notes. No fungal hosts are known for this species.
Etymology. Specific epithet denotes the distinctive darkened femora.
Pocadius ferrugineus (Fabricius)
(Figs. 23, 62, 103, 144, 184, 220)
170
Non-Type Material Examined. >250 specimens from throughout the Palearctic
including material from Austria, Belgium, Czech Republic, Denmark, England, France,
Germany, Hungary, Italy, Lithuania, Poland, Russia, Spain, Sweden, Switzerland, and Ukraine.
Diagnosis. This species can be distinguished from the sympatric Palearctic P. adustus by
the following characters: distinctly shorter pronotal and elytral fimbria, protibial notch deeper,
antennal pedicel shorter, terminal antennomere with different sensillar region, anal sclerite more
broadly convex, apico-lateral angles of eighth abdominal sternite less developed, tegmen with
more diffused/scattered inner rows of setae, median lobe more rectangular with almost
perpendicular sides, basal piece of ejaculatory rods more elongate, gonocoxites of ovipositor
with fewer elongate setae.
Redescription. Length 3.8 mm, Width 2.1mm, Depth 1.8mm. Body moderately
convex, surface somewhat shining, reddish-brown to dark reddish brown in color, sometimes
with head, pronotum and antennal club slightly darker. Pronotum and elytra margins with
moderately long fimbria, setae only slightly longer than width of antennal scape. Dorsal and
ventral pubescence moderate in length.
Head surface deeply, irregularly punctate, punctures larger on vertex with a few
interspersed smaller ones, becoming predominantly smaller towards orbits and fronotclypeal
region. Larger punctures 5 X diameter of eye facet, smaller punctures ~3 X diameter.
Interspaces smooth to alutaceous becoming somewhat granular in fronotclypeal depression,
moderately shining. Each smaller puncture gives rise to an elongate curved golden seta. Eyes
finely faceted. Pronotal surface with large punctures equal in size to large punctures on vertex of
head, interspersed with relatively few smaller punctures, equal in size to smaller punctures on
head. Interspaces smooth to alutaceous, large punctures about 0.25-0.5 diameters apart. Each
smaller puncture gives rise to a moderately long somewhat curved golden seta. Scutellar surface
171
with few moderately impressed punctures ~0.75 diameter of large punctures on pronotum, some
but not all punctures giving rise to setae, interspaces smooth with some microreticulation
present. Elytral surface with serial rows of alternating large and small deep punctures. Smaller
punctures are equal in size to smaller ones on pronotum, larger punctures are ~2.5-3 times
diameter of smaller ones. Smaller punctures giving rise to a semi-decumbent moderately long
golden seta, larger punctures giving rise to a shorter completely decumbent golden seta.
Interspaces not narrow between punctures of a given row and between different rows. Within a
row, small punctures are separated by ~2 puncture diameters, and large punctures by 0.5-0.75
puncture width. Rows are separated by 1.0-1.5 large puncture diameters. Interspaces
moderately shining but variable from smooth to somewhat granular in sculpture. Pygidium
densely punctate, punctures equal in size to smaller ones on pronotum, each puncture giving rise
to a short stiff golden seta. Interspaces narrow, 0.33-0.5 diameter, with alutaceous to granular
sculpture.
Venter with similar moderate golden pubescence as dorsum. Mentum with large very
shallowly impressed punctures, equal in size to larger ones on vertex, each giving rise to a short
golden seta. Interspaces alutaceous to finely granular. Submentum and gula similar in
punctation to mentum but with interspaces becoming more granular. Prosternum and epimeron
irregularly punctate, punctures equal to those on mentum, interspaces alutaceous to granular,
prosternal punctures separated by 0.5 diameter, those on the epimeron by 0.5-0.75 diameter.
Mesosternum with smaller somewhat deeper punctures than prosternum, about 0.75 diameter of
those on prosternum, interspaces alutaceous to granular, separated by about ~1 diameter.
Metasternum irregularly punctate, with moderate faint punctures on disc similar in size to those
on mesosternum, interspaces smooth to slightly alutaceous on metasternal disc becoming
microreticulate to granular laterally, punctures separated by ~1-3 diameters on disc. Abdominal
172
sternite 1 with moderately impressed punctures, punctures equal to those on metasternum,
interspaces alutaceous to granular, separated by ~0.5-1 diameter. Abdominal sternites 2-4 with a
faintly defined irregular row of punctures along posterior margin, the rest of the sternite has
moderately impressed punctures, interspaces 0.25-0.66 diameters apart and mostly granular
surface sculpture. Hypopygidium with moderately deep punctures, similar in size to those on
sternites 2-4, interspaces granular, punctures separated by 0.25-0.33 puncture diameter.
Head wider than long (L:W = 1:1.8), fronotclypeal region moderately projecting
anteriorly. Vertex with distinct deep concavity between orbits near fronotclypeal region. Eyes
large, moderately protruding. No prominence over antennal insertion. Labrum with small
shallow concavity at anterior margin. Antennal club compact, oval, mostly symmetrical with the
last antennomere subequal to the previous two combined. Antennomeres 6-8 more or less
compact, with 7-8 characteristically disc-like. Antennal scape broadly asymmetrical, somewhat
hemispherical, 2.1 times as long as pedicel. Pedicel subcylindrical, somewhat barrel-shaped.
Antennal segment 3 subequal in length to pedicel, becoming narrowed proximally. Antennal club
somewhat large, ~0.55 length of segments 1-8 combined. Each club segment with dense short
setae, and only relatively few protruding setae. Antennal grooves deep and widely excavate,
slightly converging posteriorly. Lateral mental ridge prominent, at level of submentum, medially
the ridge is longitudinally sculptured with microreticulations and laterally with oblique to
longitudinal microreticulations. Mentum with anterior angles obsolete, anterior margin broadly
hemispherical, lateral margins nearly perpendicular, entire structure flattened.
Pronotum widest near posterior angles (L:W = 1:2.0), anterior margin broadly
trapezoidal to moderately concave, posterior margin moderately convex with concavity near
posterior corners, lateral margins more or less evenly arcuate to posterior angles. Scutellum
large, obtusely triangular, apex rounded. Prosternal process somewhat narrowed between
173
procoxae, apex somewhat acuminate, in lateral aspect the structure is convex posterior-medially.
Posterior apical wall prominent and only slightly oblique. Mesosternum extending to 0.75 length
of mesocoxae, evenly and broadly concave for reception of the metasternum. Metasternum
width to length ratio is ~2.25:1.0. Metepisternum with only slight medial constriction, oblique
line dividing anterior 0.18 of structure. Elytral humeri moderately produced, lateral margin very
narrow. First abdominal sternite with broadly rounded process between metacoxae. First
sternite ~1.8X’s longer than second sternite. Sternites 2-3 subequal in length, the fourth slightly
larger than the preceding two. Hypopygidium 0.88 length to first abdominal sternite.
Protibia with apical tooth slightly prominent, subequal in length to 0.5 tarsomere 1.
Outer apical notch with ~130° angle, notch depth moderately deep, equal to length of tarsomere
1 and part of 2 combined. Inner apical spine subequal in length to tarsomeres 1 and small part of
2 combined. Protibia not heavily armored but with characteristic dense patch of stiff setae along
the inner apical region. Mesotibia more heavily armored than protibia with more dense stiff
setae and a row of numerous slender spines along entire lateral edge. Outer apical process
moderately elongate, larger than protibia process, and apical margin with row of flat teeth. Inner
apical spine equal in length to tarsomeres 1-2 combined. Metatibia with armature similar to that
of mesotibia, including row of flat teeth on apical margin.
Male genitalia well-sclerotized. Anal sclerite with large broadly curved region
anteriodorsally (Fig. 23); apex fimbriate; ventrally with a broad medial concavity approaching
apex. Spiculum gastrale with wide broadly convex lateral flanges, medial margins narrowly
concave, short stiff setae originating from apex and also anteapically (Fig. 62). Tegmen evenly
rounded apically (Fig. 103), much longer than wide (w:l = 1.0:2.25), lateral row of setae visible
from the median fossa around the apex, small shallow concavity in apical third, basal notch
perpendicular, basal margin angulate. Median lobe large and robust, ~0.5 the length of the
174
tegmen, apex acuminate, lateral margins nearly perpendicular, apical opening well-developed
(Fig. 144). Ejaculatory rods not fused to basal piece or each other, straight with slight outward
expansion apically. Basal piece of rods separated, with two lunate shaped pieces coming to
sharp apical point (Fig. 184).
Female genitalia moderately sclerotized. Paraprocts large with sclerotization only along
median line. Gonocoxite with two basal lateral prominences, basal ridge well-sclerotized.
Baculi extending medially from gonocoxites base, length equal to ~0.5 gonocoxal invagination.
Gonocoxal apices with recurved “tooth” present. No primary elongate setae originate from small
grooves along the side of the gonocoxal tooth apices (Fig. 220).
Variation. Overall length of specimens can vary greatly, but typically females are
somewhat larger than males.
Seasonality/Habitat. Specimens that were available for this study indicate that most
collections of this species occurred from May through October in forested regions of the
Palearctic.
Distribution. This species has the most extenisve distribution of any known Pocadius.
Pocadius ferrugineus is known from Scandinavia south through the Iberian peninsula, west
through Turkey parts of the Middle East and Central Asia, and as far west and north as the
Kamchatka peninsula in northwestern Russia. Kirejtshuk (1992) also proposes the species is
known from Malaysia, however, this record seems suspect and should be confirmed.
Notes. This species has the most catholic diet of Lycoperdaceae genera known amongst
the Pocadius having been reported from species of Lycoperdon, Calvatia, Bovista,
Langermannia, and Scleroderma, as well as non-Gasteromycetes taxa (Audisio 1993).
Pocadius fulvipennis Erichson
175
(Figs. 24, 63, 104, 145, 185, 221)
Type Material Examined. HOLOTYPE ♀ (Humboldt, Berlin): 8641 / HOLOTYPUS;
Pocadius; fulvipennis; Erichson, 1843 / fulvipennis; Er.; Mexico, Ehrenb.
Non-Type Material Examined. >400 specimens from western and southwestern North
America. Most specimens were collected from Mendocino, Marin, Santa Cruz, Trinity, Sonoma,
San Matco, and Alameda Counties in California. Other specimens were identified from Arizona,
Oregon and Washington in the U.S. and British Columbia, Canada. The following are new
distribution records for the species: IDAHO: Valley Co.; Round Valley, David M. Ward Jr.
collr.; ex. puffball; 25-JUN-1995. IDAHO: Bonner Co.; Sandpoint, N.M. Downie; VII-8-1971
(FMNH).
Diagnosis. Easily distinguished from the other Nearctic species by the following
characters: pronotum and head always darker in color than elytra, though elytral apices may
sometimes be as dark as the head and pronotum; terminal antennomeres with distinct sensillar
region, prominent tibial armature, granular mental-submental sulcus, relatively broad abdominal
process, very slender concavity for reception of the tegmen on the anal sclerite, eighth abdominal
sternite with dense elongate brush of setae apically, apex of tegmen characteristically indentate,
median lobe with well-developed acuminate apex and concave lateral margins, ejaculatory rods
thick and robust and abutting each other and basal piece, and gonocoxites with two elongate
primary setae.
Redescription. Length 3.8 mm, Width 2.0mm, Depth 1.1mm. Body moderately
convex, surface shining, dark reddish-brown to dark brown in color, elytra always lighter in
color, varying from light brown to light red-brown. Pronotum and elytra margins with extremely
elongate fimbriae, setae more than 2X’s longer than width of antennal scape. Dorsal and ventral
pubescence overall quite elongate and light golden in color.
176
Head surface deeply, irregularly punctate, punctures larger on vertex, becoming
somewhat smaller towards orbits and with only small punctures on fronotclypeal region. Larger
punctures 4-5 X’s diameter of eye facet, smaller punctures ~3 X’s diameter. Interspaces smooth
to alutaceous, shining, 0.5-0.75 diameter of large puncture apart. Each smaller puncture gives
rise to an elongate moderately curved golden seta. Eyes finely faceted. Pronotal surface densely
punctate with large punctures equal in size to large punctures on vertex of head, interspersed
with relatively few smaller punctures, ~0.75 size of larger ones. Interspaces smooth to
alutaceous, about 0.25-0.75 large puncture diameters apart. Each puncture gives rise to a long
curved golden seta. Scutellar surface with large impressed punctures mostly in basal 0.5 of
structure, some punctures giving rise to setae, with interspaces alutaceous to granular. Elytral
surface with serial rows of alternating large and small deep punctures. Smaller punctures are
equal in size to large ones on pronotum, larger punctures are ~2.5 times diameter of smaller
ones. Smaller punctures giving rise to a semi-erect long golden seta, larger punctures giving rise
to a decumbent shorter golden seta. Interspaces moderately well separated between punctures of
a given row and between different rows. Within a row, small punctures are separated by 1-2
puncture diameters, and large punctures by 0.75-1.0 puncture width. Large puncture rows are
separated by 1.0-1.5 puncture diameters. Interspaces shining but variable from finely alutaceous
to granular in sculpture. Pygidium densely punctate, punctures equal in size to larger ones on
pronotum, each puncture giving rise to a moderately long straight golden seta. Interspaces
narrow, 0.5-0.75 diameters, with micro-reticulate to granular surface sculpture.
Venter with somewhat shorter golden pubescence as dorsum. Mentum with large
moderately impressed punctures, equal in size to larger ones on vertex, each giving rise to a
moderately long seta. Interspaces smooth to finely alutaceous. Submentum and gula similar in
punctation to mentum but with interspaces alutaceous to microreticulate. Prosternum and
177
epimeron deeply irregularly punctate, punctures slightly larger than those on mentum, ~1.5X
diameter, interspaces microreticulate with some granular areas, prosternal punctures separated by
0.25-0.5 diameter, those on the epimeron by ~0.5 diameter. Mesosternum with deeply impressed
punctures, about 1.25-1.5 diameter of those on prosternum, interspaces alutaceous to granular,
separated by about 0.5 diameter. Metasternum irregularly punctate, with moderately deep
punctures on disc similar in size to those on mentum, interspaces smooth to alutaceous on
metasternal disc becoming alutaceous to granular laterally, punctures separated by ~1-2
diameters on disc. Abdominal sternite 1 with large moderately impressed punctures, punctures
equal to those on metasternum, interspaces alutaceous with some microreticulation present,
separated by ~1-1.5 diameter. Abdominal sternites 2-4 with one irregular disorganized rows of
punctures near the posterior margin, punctures similar in size to those on metasternum, punctures
separated by 0.5-1 diameters. Hypopygidium with moderately deep punctures, similar in size to
those on sternites 2-4, interspaces alutaceous to granular with some microreticulation present,
punctures separated by 0.25-0.5 puncture diameters.
Head slightly wider than long (W:L = 1:1.3), fronotclypeal region moderately projecting
anteriorly. Vertex with shallow and broad concavity between orbits near fronotclypeal region.
Labrum with narrow incision at anterior margin. Antennal club compact, oval, symmetrical with
the last antennomere subequal in length to the previous two combined. Antennomeres 6-8 more
or less compact, with 7-8 characteristically disc-like. Antennal scape asymmetrical, distinctly
hemispherical, ~2 times as long as pedicel. Pedicel subcylindrical, approximately barrel-shaped.
Antennal segment three ~0.8 length of pedicel. Antennal segments four and five cuboid and
equal in length. Antennal club large, ~0.8 length of segments 1-8 combined. Each club segment
with several dense short setae protruding. Antennal grooves very deep and widely excavate,
converging posteriorly. Lateral mental ridge prominent and elongate, at level of submentum,
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medially the ridge is granular with few transversely sculptured microreticulations. Mentum
broadly pentagonal with anterior angles broad but distinct, lateral sides short but perpendicular,
anterior margin coming to acute medial apex, entire structure flattened.
Pronotum widest near posterior angles (L:W = 1:2.2), anterior deeply distinctly
trapezoidal, posterior margin broadly convex, lateral margins arcuate from posterior to anterior
angles. Scutellum large, broadly triangular, apex broadly rounded. Prosternal process broadly
narrowed between procoxae, apex broadly rounded, in lateral aspect the anterior and posterior
ends are not prominent and convex in posterior 0.33 and angulate in anterior 0.66. Posterior
apical wall not prominent, short, and perpendicular. Mesosternum extending to midway between
mesocoxae, evenly broadly concave for reception of the metasternum. Metasternum width to
length ratio is ~2.38:1.0. Metepisternum with slight medial constriction, oblique line dividing
anterior 0.125 of structure. Elytral humeri moderately produced, lateral margin very narrow.
First abdominal sternite with broadly rounded process between metacoxae. First sternite ~2X’s
longer than second sternite. Sternites 2-4 subequal in length. Hypopygidium subequal in length
to first abdominal sternite.
Protibia with apical tooth prominent, slightly longer than tarsomeres one and part of two
combined. Outer apical notch with ~105° angle, notch depth moderate, subequal to length of
tarsomere 1. Inner apical spine subequal in length to tarsomere one and part of two combined.
Protibia heavily armored with characteristic dense patch of stiff setae along the inner apical
region. Mesotibia more heavily armored than protibia with more dense stiff setae and an
additional row of numerous slender spines along entire lateral edge. Outer apical process
elongate and robust, larger than protibia process, equal to tarsomere one and two combined.
Inner apical spine equal in length to tarsomeres 1-2 combined. Metatibia with armature similar
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to that of mesotibia, but setae longer and apical spine and process slightly more robust and
longer.
Male genitalia well-sclerotized. Anal sclerite with large broadly curved region
anteriodorsally (Fig. 24); apex somewhat fimbriate; ventrally with a narrow medial concavity
approaching apex. Spiculum gastrale with broad wide lateral flanges, medial margins concave
proximally, elongate stiff setae originating from apex (Fig. 63. Tegmen broadly emarginate
apically (Fig. 104), much longer than wide (w:l = 1.0:3.2), lateral row of setae visible from the
median fossa to around the apex, basal notch perpendicular, basal margin of phallobase broadly
concave, three inner rows of setae present. Median lobe large and robust with lateral margins
concave and apex sharply acuminate, ~0.66 the length of the tegmen, apical opening not well-
developed (Fig. 145). Ejaculatory rods not fused to basal piece but directly adjacent to it, and
adjacent to each other apically and basally, slightly curved outwardly. Basal piece of rods with
deep medial concavity extending almost 0.75 length of structure, proximally with three sharp
projections, fused basally but not apically (Fig. 185).
Female genitalia moderately sclerotized. Paraprocts large with sclerotization along
median line to baso-lateral angles. Gonocoxite with two basal lateral prominences, basal ridge
well-sclerotized. Two elongate baculi extending down median region of gonocoxites, ~0.5
length of gonocoxal extension. Gonocoxal apices with recurved “tooth” present. Two primary
setae originate from small depressions on the sides of the gonocoxal teeth (Fig. 221).
Variation. Some specimens have elytral lateral and apical margins darker than rest of
elytra, some specimens may also have a lighter pronotal lateral margin. Some specimens from
Idaho are more slender in overall body shape, but have all other features similar to holotype.
Females are slightly larger than males, and the male pygidial apex broadly shallowly concave.
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Seasonality/Habitat. Most specimens have been collected from lowland
deciduous/mixed deciduous forests from March through October.
Distribution. This species occurs from British Columbia, Canada in the north southward
through the western U.S. and into parts of Mexico. Other than the holotype, the author has
encountered relatively few specimens from Mexico (e.g. six specimens). Clearly more collection
efforts in NW Mexico and Baja California are needed to verify the southern limits of the range of
this species.
Notes. Larval specimens of this species were successfully reared from Lycoperdon
puffball spores mixed 0.5:0.5 with rust spores by Lorin Gillogly. Gillogly noted on labels that
the process of maturation from larvae to adult took approximately 1-2 months on this medium.
However, no definitive information was provided regarding the stage of larval development from
which this rearing process was initiated. Specimens reared on this diet were smaller than others
of the species. Other host records include Calvatia giganteum.
Pocadius fumatus Jelínek
(Figs. 25, 64, 105, 146, 186, 222)
Type Material Examined. 1 PARATYPE (Czech Republic): Brazil, Sao Paulo; J. Mráz
lgt.
Non-Type Material Examined. 12 specimens (CMN): ARG: Salta Prov.; 45km W
Salta, 1950m; 1-29-XII-87, El Alisal; S&J Peck, moist ravine; thicket, malaise-FIT.
Diagnosis. As suggested by Jelínek (1977) P. fumatus is closely related to P.
dimidiatus, but can be readily distinguished from it by the following suite of characters: surface
of head smooth and shining with no rugose or microreticulate regions, overall large body size,
terminal antennomere less broadly hemispherical and with distinct depressed area, protibial outer
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apical tooth only somewhat anteapical with very short small outer apical notch, serial row of
large punctures on elytra widely separated, tegmen robust with complete inner row of setae and
elongate apical furrow, median lobe with shallow bilobed opening, spiculum gastrale broadly
flanged with elongate apical fimbria, ejaculatory rods fused basally, basal piece separated into
two large sickle-shaped pieces, ovipositor with short baculi on gonocoxae and two primary and
two secondary apical gonocoxal setae.
Redescription. Length 4.7 mm, Width 2.8mm, Depth 1.2mm. Body somewhat convex,
surface shining, tan-brown to dark brown in color with posterior 0.66-0.75 of elytral dark brown,
and sometimes with pronotal disc somewhat darker. Pronotum and elytra margins with
moderately long fimbriae, setae subequal to slightly longer than width of antennal scape. Dorsal
and ventral pubescence moderately long.
Head surface deeply, densely, irregularly punctate, punctures larger on vertex, becoming
somewhat smaller towards orbits and fronotclypeal region, some smaller punctures interspersed
with large ones on vertex. Larger punctures 5-6 X diameter of eye facet, smaller punctures 2-3
X diameter. Interspaces smooth and shining, ~0.5 diameter apart. Each smaller puncture gives
rise to a moderately elongate golden seta. Eyes finely faceted. Pronotal surface with large
punctures ~1.5 diameter to large punctures on vertex of head, interspersed with relatively few
smaller punctures, which are subequal to size of larger ones on vertex. Interspaces smooth to
finely alutaceous, about 0.25-0.5 diameters apart. Each smaller puncture gives rise to a thin
golden seta, most seta are curved and not elongate. Scutellar surface evenly punctate with very
moderately impressed punctures equal to large punctures on pronotum, some punctures giving
rise to short thin golden setae, interspaces smooth to alutaceous. Elytral surface with serial rows
of alternating large and small moderately impressed punctures. Smaller punctures are equal in
size to smaller ones on pronotum, larger punctures are slightly larger than larger punctures on
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pronotum. Smaller punctures giving rise to a semi-erect to erect moderately long golden seta,
larger punctures giving rise to a shorter decumbent golden seta. Interspaces narrow between
punctures of a given row and wide between different rows. Within a row, small punctures are
separated by ~1 puncture width, and large punctures by 0.33 puncture width. Larger puncture
rows are separated by ~2 large puncture diameters. Interspaces always shining but variable from
mostly smooth to finely alutaceous in sculpture. Pygidium densely punctate, punctures equal in
size smaller ones on elytra, each puncture giving rise to a short semi-erect golden seta.
Interspaces narrow, 0.33-0.5 diameters, with smooth to finely microreticulate sculpture.
Venter with similar moderately long golden pubescence as dorsum. Mentum with few
very shallow punctures, equal in size to smaller ones on vertex, each giving rise to a short seta.
Interspaces smooth to finely microreticulate. Submentum and gula similar in punctation to
mentum but with interspaces completely having more distinct microreticulation present.
Prosternum and epimeron shallowly irregularly punctate, punctures larger than those on mentum,
interspaces smooth to alutaceous with some microreticulate areas, prosternal punctures separated
by 0.25-0.5 diameter, those on the epimeron by 0.5-0.75 diameter. Mesosternum with shallow
punctures aggregated along meso- metasternal border, equal in diameter to those on prosternum,
interspaces mostly alutaceous, separated by about 0.5 diameter. Metasternum irregularly
punctate, with moderately deep punctures on disc similar in size to those on mesosternum,
interspaces smooth on metasternal disc becoming microreticulate to granular laterally, punctures
separated by ~1 diameter. Abdominal sternite 1 with large faint, almost obsolete punctures,
punctures subequal to those on metasternum, interspaces mostly alutaceous with abdominal
process rugose, separated by ~0.5-1 diameter. Abdominal sternites 2-4 with an irregular row of
punctures near posterior margin, punctures ~0.66 in size to those on metasternum, punctures
within row separated by ~0.5 punctures width. Hypopygidium with moderately deep punctures,
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similar in size to those on sternites 2-4, interspaces smooth to alutaceous with some
microreticulate areas, punctures separated by 0.5-1 puncture diameter.
Head wider than long (W:L = 1.43:1), fronotclypeal region moderately projecting
anteriorly. Vertex with broad shallow concavity between orbits near fronotclypeal region.
Labrum with deep distinct incision at anterior margin. Antennal club compact, oval, broadly
asymmetrical with the last antennomere longer than the previous two combined. Antennomeres
6-8 more or less compact, with 7-8 characteristically disc-like. Antennal scape somewhat
asymmetrical, broadly hemispherical, 1.33 times as long as pedicel. Pedicel subcylindrical in
shape and elongate. Antennal segment 3 equal in length to pedicel. Antennal club large, ~0.68
length of segments 1-8 combined. Each club segment with dense short setae, and several longer
protruding setae. Antennal grooves very deep and widely excavate, slightly converging
posteriorly. Lateral mental ridge prominent, at level of submentum, medially the ridge is
granularly sculptured with few microreticulations and laterally with oblique to longitudinal
microreticulations. Mentum with anterior angles somewhat distinct and obtuse, anterior margin
more or less angled to a central point, having an overall broadly pentagonal appearance, entire
structure with little to no central convexity.
Pronotum widest in posterior third (L:W = 1:2), anterior margin broadly shallowly
concave, posterior margin moderately convex, lateral margins less arcuate posteriorly.
Scutellum large, triangular, apex narrowly rounded. Prosternal process somewhat narrowed
between procoxae, apex somewhat acuminate, in lateral aspect the anterior and posterior ends are
prominent with a distinct medial convexity. Posterior apical wall prominent and oblique (though
in some Argentinean specimens not as dramatic as illustrated in Jelínek 1977). Mesosternum
extending to midway between mesocoxae, evenly broadly concave for reception of the
metasternum. Metasternum width to length ratio is ~2.6:1.0. Metepisternum with slight medial
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constriction, oblique line dividing anterior 0.15 of structure. Elytral humeri moderately
produced, lateral margin very narrow. First abdominal sternite with acuminate process between
metacoxae. First sternite ~2X’s longer than second sternite. Sternites 2-4 subequal in length.
Hypopygidium subequal in length to first abdominal sternite.
Protibia with apical tooth prominent, subequal in length to tarsomeres 1-2 combined.
Outer apical notch absent. Inner apical spine subequal in length to tarsomeres 1 and most of 2
combined. Protibia not heavily armored but with characteristic dense patch of stiff setae along
the inner apical region. Mesotibia more heavily armored than protibia with more dense stiff
setae and a row of numerous slender spines along entire lateral edge. Outer apical process
elongate and robust, larger than protibial process. Inner apical spine equal in length to
tarsomeres 1-2 combined. Metatibia with armature similar to that of mesotibia, but with setae
and spines slightly longer.
Male genitalia well-sclerotized. Anal sclerite transverse (Fig. 25); apex somewhat
fimbriate; ventrally with a broad shallow medial concavity approaching apex. Spiculum gastrale
with wide lateral flanges, medial margins straight, moderately long stiff setae originating from
apex (Fig. 64). Tegmen evenly rounded apically (Fig. 105), much longer than wide (w:l =
1.0:2.56), lateral row of setae visible from the median fossa to prior to around the apex, elongate
shallow concavity in apical third, basal notch obsolete, basal margin nearly straight, inner row of
tegminal setae distinct and reaching apex. Median lobe large and robust, ~0.66 the length of the
tegmen, apex narrowly rounded, apical opening well-developed with proximal and lateral
concavity (Fig. 146). Ejaculatory rods not fused to basal piece but fused to each other in distal
third, enlarged and expanded outward distally. Basal piece of rods with two separate sclerites,
each more or less sickle shaped with distal swollen region (Fig. 186).
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Female genitalia moderately sclerotized. Paraprocts large with sclerotization only along
median line to baso-lateral angles. Gonocoxite with two basal lateral prominences, basal ridge
well-sclerotized. Gonocoxal apices without recurved “tooth” present. Two primary and three
secondary setae originate from small depressions on the gonocoxal apices (Fig. 222).
Variation. Some specimens with dark area on elytra more clearly delimited than others,
two specimens with lighter area extending posteriorly on elytra near suture.
Seasonality/Habitat. This species is known from moist-wet forested areas. No
seasonality information was recorded from the type series (Jelínek 1977), however, the
Argentinean specimens suggest the species is active throughout December.
Distribution. Previously this species was known only from Brazil, the Argentina locality
above denotes a new country record and range extension for the species.
Notes. No host fungal information is available for this species. The Argentinean
specimens are evidence that this species is sympatric with the closely related P. dimidiatus,
which is known from Argentina. Specimens of both species were collected at the same locality
45km W of Salta.
Pocadius fusiformis Cline new species
(Figs. 26, 65, 106, 147, 187, 223)
Type Material Examined. HOLOTYPE ♂ (BMNH): INDONESIA; SULAWESI
UTARA; Danau Mooat 1200m.; nr. Kotamobagu; 16 Feb. 1985 / Lower montane; forest; 1200-
1400m / puffballs / HOLOTYPE; Pocadius; fusiformis; A. Cline des. 2004. 1 PARATYPE ♂
(BMNH): same data as holotype. 3 PARATYPES ♀ (BMNH): INDONESIA; SULAWESI
UTARA; Gng. Ambang F.R.; nr. Kotamobagu; Jan. 1985 / Lower montane; forest; 1200-1400m
/ in mature puffballs. All specimens have a label turned upside down under the rest of the data
186
labels, but prior to a type label with the following information: R. Ent. Soc. Lond.; PROJECT
WALLACE; B.M. 1985-10. One of the female paratypes collected in January also has an
additional label with the following information: 2 examples in; Mus. Bogoriense.
Diagnosis. This species is easily distinguished from the other Old World species by the
following suite of characters: the characteristic color pattern consisting of a dark brown-black
scutellum, pronotal disc, and elytra except for the anterior-medial fourth with the rest of the body
light tan-brown; very short elytral pubescence that is decumbent in both the large and small
serially punctate rows; greatly enlarged terminal antennomere; swollen/thickened tarsomeres;
anal sclerite ending in acute apical point; elongate narrow tegmen and median lobe; ejaculatory
rods with convex distal region; and ovipositor gonocoxites without recurved apical tooth and
broadly rounded baso-lateral angles.
Description. Length 4.1mm, Width 2.3mm, Depth 1.5mm. Body moderately convex,
surface shining, dark brown/black in color with elytral humeri, lateral pronotal margins, anterior
half of head, venter, and appendages lighter. Pronotal and elytral margins with short fimbriae,
setae shorter than length of antennal scape. Dorsal and ventral pubescence quite short and fine,
light yellow in color.
Head surface moderately deeply, irregularly punctate, punctures larger on vertex with
some interspersed smaller punctures, becoming somewhat smaller towards orbits and
fronotclypeal region. Larger punctures 3-4X diameter of eye facet, smaller punctures ~2X
diameter; interspaces smooth with some microreticulation present and distinctly shining. Each
smaller puncture gives rise to a short seta. Pronotal surface with large punctures equal in size to
large punctures on vertex of head, interspersed with relatively few smaller punctures, equal in
size to smaller ones on head; interspaces smooth with areas of fine microreticulation, about 1-2
diameters apart. Each puncture gives rise to a short decumbent seta. Scutellar surface with few
187
vague, shallowly impressed small punctures, equal to smaller ones on vertex, some punctures
giving rise to short setae; interspaces smooth with fine microreticulate areas. Elytral surface
with serial rows of alternating large and small deep punctures. Smaller punctures are equal in
size to smaller ones on pronotum, larger punctures are ~3 times diameter of smaller ones.
Smaller punctures giving rise to a decumbent short seta, larger punctures giving rise to a short
decumbent seta. Interspaces broad between punctures of a given row and between different
rows. Within a row, small punctures are separated by ~2-3 puncture diameters, and large
punctures by ~1 puncture diameter. Larger rows are separated by ~2 large puncture diameters.
Interspaces always shining but variable from smooth to finely microreticulate in sculpture.
Pygidium densely punctate, punctures equal in size to larger ones on pronotum, each puncture
giving rise to a short straight seta. Interspaces narrow, 0.25-0.5 diameter, shining with heavily
microreticulate sculpture.
Venter with similar short pubescence as dorsum. Mentum with few moderately large
shallow punctures, equal in size to larger ones on vertex, each giving rise to a short seta;
interspaces smooth to alutaceous and finely microreticulate. Submentum and gula with smaller
punctation, punctures ~0.5-0.75 diameter of those on mentum, and with interspaces more
granular. Prosternum and epimeron irregularly punctate, punctures moderately impressed, those
on epimeron equal to large punctures on elytra and those of prosternum to smaller elytral
punctures, interspaces alutaceous with microreticulate areas, prosternal punctures separated by
0.5 diameter, those on the epimeron by 0.25 diameter. Mesosternum with shallow punctures,
about 0.75 diameter of those on epimeron, interspaces alutaceous to somewhat granular,
separated by about 0.5 to 1 diameter, and aggregated near metasternum. Metasternum densely
irregularly punctate, with faint punctures on disc similar in size to those on mesosternum,
interspaces granular on metasternal disc becoming microreticulate laterally, punctures separated
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by ~0.5-1 diameter. Abdominal sternite 1 with large faint, almost obsolete punctures, punctures
equal to those on metasternal disc, interspaces granular with microreticulations, separated by
~0.5-1 diameter. Abdominal sternites 2-4 with irregular rows of punctures, punctures similar in
size to those on metasternum, rows separated by 1-2 puncture diameters, punctures within rows
separated by ~0.5 puncture diameter. Rows on abdominal sternite 4 becoming less organized.
Hypopygidium with moderately deep punctures, similar in size to those on sternites 2-4,
interspaces granular, punctures separated by 0.25-0.5 puncture diameter.
Head wider than long (L:W = 1:1.5), fronotclypeal region moderately projecting
anteriorly. Vertex with deep transverse concavity between orbits near fronotclypeal region.
Labrum with shallow concavity at anterior margin. Antennal club compact, somewhat globular,
asymmetrical with the last antennomere greater in length than the previous two combined.
Antennomeres 4-8 more or less compact, with 7-8 characteristically disc-like and 4-6
trapezoidal. Antennal scape asymmetrical, somewhat hemispherical, 1.4 times as long as
pedicel. Pedicel subcylindrical in shape. Antennal segment 3 subequal in length to pedicel.
Antennal club large, ~0.65 length of segments 1-8 combined. Antennal grooves very deep and
widely excavate, slightly converging posteriorly. Lateral mental/submental ridge moderately
prominent, at level of submentum, ridge with longitudinal microreticulations and the
corresponding sulcus with minute punctures and granular sculpture. Mentum with anterior
angles present, anterior margin angulate coming to an acute apex, entire structure transversely
pentagonal ventrally and somewhat convex laterally.
Pronotum widest near middle (L:W = 1:2), anterior margin broadly trapezoidal to
evenly concave, posterior margin moderately convex, lateral margins slightly arcuate to anterior
and posterior angles. Scutellum large, obtusely triangular, apex broadly rounded. Prosternal
process somewhat narrowed between procoxae, apex acuminate, in lateral aspect the posterior
189
end is prominent and slightly upturned and steeply convex medially over the procoxae. Posterior
apical wall prominent and perpendicular. Mesosternum extending to midway between
mesocoxae, evenly concave for reception of the metasternum. Metasternum wider than long
(L:W = 1:2.9). Metepisternum with slight medial constriction, oblique line dividing anterior
0.13 of structure. First abdominal sternite with broad process between metacoxae. First sternite
~2X’s longer than second sternite. Sternites 2-3 subequal in length, the fourth slightly larger
than the preceding two. Hypopygidium subequal in length to first abdominal sternite.
Protibia with apical tooth prominent, slightly longer than tarsomeres 1 and part of 2
combined. Outer apical notch with ~115° angle, notch depth shallow, equal to length of half of
tarsomere 1. Inner apical spine subequal in length to tarsomeres 1-2 combined. Protibia not
heavily armored but with characteristic dense patch of stiff setae along the inner apical region.
Mesotibia more heavily armored than protibia with more dense stiff setae and a row of numerous
slender spines along entire lateral edge. Outer apical process moderate, subequal in length to
that of protibia process. Inner apical spine equal in length to tarsomeres 1-2 combined.
Metatibia with armature similar to that of mesotibia, but with inner apical spine equal to
tarsomeres 1-2 and part of 3 combined.
Male genitalia well-sclerotized. Anal sclerite with elongate narrowly curved region
anteriodorsally (Fig. 26); apex sparsely fimbriate; ventrally with a medial concavity approaching
apex in a broadly concave manner. Spiculum gastrale with narrow lateral flanges curved
apically, medial margins biconcave, moderately long stiff setae originating from apex (Fig. 65).
Tegmen evenly rounded apically (Fig. 106), much longer than wide (w:l = 1:3.7), lateral row of
setae visible from the median fossa to prior to the apex, small shallow concavity in apical third,
inner row of setae incomplete not attaining apex, basal notch perpendicular, basal margin deeply
concave. Median lobe elongate, ~0.85 the length of the tegmen, apex rounded, apical opening
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well-developed with short bilobed internal structure (Fig. 147). Ejaculatory rods not fused to
basal piece or each other but adjacent to one another almost the entire length, both rods swollen
apically. Basal piece of rods with two elongate apico-lateral extensions (Fig. 187).
Female genitalia moderately sclerotized. Paraprocts large with sclerotization only along
median line to apico-lateral angles. Gonocoxite with no basal lateral prominences, basal ridge
well-sclerotized. Gonocoxal apices with recurved “tooth” absent. Four primary setae originate
from small depressions on the gonocoxal apices. Intragonocoxal invagination shallow, <0.25
length of gonocoxite (Fig. 223).
Variation. Some specimens have more of the elytra and pronotal margin lighter in color.
Females tend to be slightly larger.
Seasonality/Habitat. Specimens were collected during January and February from
montane forests.
Distribution. Known from the type locality on the northern peninsula of Sulawesi,
Indonesia.
Notes. Host information exists only as collected in “puffballs.”
Etymology. Specific epithet denotes the overall fusiform body shape.
Pocadius globularis Cline n. sp.
(Figs. 27, 66, 107, 148, 188, 224)
Type Material Examined. HOLOTYPE ♂ (CDFA): HONDURAS: Fca. Morazan; San
Anotnio de Oriente; El Zamorano; 5 MAY 1993; rcol. R. Cave / Bovistos / HOLOTYPE;
Pocadius; globularis; A.R. Cline des. 2004. 9 PARATYPES (3 FSCA, 2 CAS, 2 LSAM, 2
USNM): same data labels as holotype but with PARATYPE designation labels.
191
Diagnosis. Similar to P. helvolus, but can be distinguished by the following characters:
body overall globular in shape with low L:W ratio, uniformly dark reddish brown color,
pronotum with trapezoidal anterior margin and widest point near posterior angles, terminal
antennomere with central circular sensillar region, metasternal disc smooth and shining but
heavily punctate with large punctures, scutellum heavily punctate, elytra with both large and
small rows of serial punctures bearing semi-erect setae, apical wall of prosternal process oblique
when viewed laterally, median lobe large and robust with acute apex, tegmen with elliptical
apical fossa and elongate curved inner row of setae not attaining apex, basal piece of internal sac
sclerites with two distinct pieces the more distal of which has two medial sharp projections.
Redescription. Length 3.45 mm, Width 2.25mm, Depth 1.2mm. Body moderately
convex, surface shining, reddish-brown to dark reddish brown in color. Pronotum and elytra
margins with moderately long fimbriae, setae equal in length to antennal scape. Dorsal and
ventral pubescence moderately long.
Head surface deeply, densely, irregularly punctate, punctures larger on vertex,
interspersed with somewhat smaller punctures that become more abundant towards orbits and
fronotclypeal region. Larger punctures 3-4 X diameter of eye facet, smaller punctures 1-2 X
diameter. Interspaces alutaceous to granular, ~0.5-1 diameter apart, and somewhat shining.
Each smaller puncture gives rise to a short golden seta. Pronotal surface with large punctures 1.5
diameter to large punctures on vertex of head, interspersed with numerous smaller punctures,
~0.5-0.75 size of larger ones. Interspaces alutaceous to finely granular, about 0.5-1.5 diameters
apart. Each puncture gives rise to a short straight golden seta. Scutellar surface with numerous
shallowly impressed punctures equal to smaller punctures on pronotum, some punctures giving
rise to setae, with the interspaces smooth to alutaceous. Elytral surface with serial rows of
alternating large and small deep punctures. Smaller punctures are equal in size to smaller ones
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on pronotum, larger punctures are ~2 times diameter of smaller ones. Smaller punctures giving
rise to a semi-erect somewhat decumbent moderately long golden seta, larger punctures giving
rise to an adpressed moderately long golden seta. Interspaces broad between punctures of a
given row and between different rows. Within a row, small punctures are separated by ~1-2
puncture diameters, and large punctures by ~1 puncture diameter. Large puncture rows are
separated by ~2 large puncture diameters. Interspaces only shining and mostly alutaceous with
some microreticulation in sculpture. Smaller puncture rows may contain more than a single row
of punctures, in particular near suture. Pygidium densely punctate, punctures equal in size to
larger ones on pronotum, each puncture giving rise to a short straight golden seta, interspaces
narrow, 0.25-0.5 diameter, with smooth sculpture.
Venter with similar moderately long golden pubescence as dorsum. Mentum with small
shallow punctures, equal in size to smaller ones on vertex, each giving rise to a moderately long
seta, interspaces smooth to alutaceous. Submentum and gula similar in sparse punctation to
mentum but with interspaces becoming granular. Prosternum and epimeron with moderately
deep irregular punctation, punctures on epimeron equal to large ones on pronotum and those on
prosternum equal to smaller ones on pronotum, interspaces almost completely granular,
prosternal punctures separated by 0.5-2 diameters, those on the epimeron by ~0.5-1 diameter.
Mesosternum with shallow punctures, similar in size as those on prosternum, interspaces
alutaceous to finely granular, separated by about 0.5 to 1 diameter, and aggregated along
posterior border next to metasternum. Metasternum irregularly punctate, with moderately
impressed punctures on disc similar in size to larger ones on pronotum, interspaces smooth on
metasternal disc becoming more finely alutaceous laterally, punctures separated by ~1 diameter.
Abdominal sternite 1 with small faint, almost obsolete punctures, punctures equal to those on
mentum, interspaces smooth with microreticulate areas, separated by ~1-2 diameters.
193
Abdominal sternites 2-4 with indistinct irregular rows of punctures, punctures separated by ~0.5-
1 puncture diameter. Hypopygidium with moderately deep punctures, similar in size to those on
sternites 2-4, interspaces smooth with microreticulate regions, punctures separated by 0.25-0.5
puncture diameters.
Head wider than long (L:W = 1:1.41), fronotclypeal region moderately projecting
anteriorly. Vertex with broad deep concavity between orbits near fronotclypeal region. Labrum
with moderately deep medial incision at anterior margin. Antennal club compact, oval,
symmetrical with the last antennomere subequal to the previous two combined. Antennomeres
4-5 more or less cuboid, 6 trapezoidal and 7-8 characteristically disc-like. Antennal scape
moderately asymmetrical, slightly hemispherical, 0.9 times as long as pedicel. Pedicel
subcylindrical in shape. Antennal segment 3 equal in length to pedicel and tapering proximally.
Antennal club large, ~0.80 length of segments 1-8 combined. Antennal grooves very deep and
widely excavate, slightly converging posteriorly. Lateral mental/submental ridge prominent, at
level of submentum, with both oblique and longitudinal microreticulations, corresponding sulcus
widely excavate. Mentum with anterior angles distinct, anterior margin angulate with well
defined apical angle, entire structure broadly pentagonal when viewed ventrally and flattened
when viewed laterally.
Pronotum widest near posterior angles (L:W = 1:1.9), anterior margin broadly deeply
trapezoidal, posterior margin moderately convex, lateral margins evenly arcuate to anterior
angles. Scutellum large, obtusely triangular, apex rounded. Prosternal process narrowed
between procoxae, apex acuminate, in lateral aspect the posterior end is somewhat recurved and
there is a modest convexity medially over the coxae. Posterior apical wall prominent and
oblique. Mesosternum extending to midway between mesocoxae, evenly broadly concave for
reception of the metasternum. Metasternum wider than long (W:L = 2.77:1.0). Metepisternum
194
with slight medial constriction, oblique line dividing anterior 0.2 of structure. First abdominal
sternite with acuminate process between metacoxae. First sternite ~2X’s longer than second
sternite. Sternites 2-3 subequal in length, the fourth slightly larger than the preceding two.
Hypopygidium subequal in length to first abdominal sternite.
Protibia with apical tooth prominent, slightly longer than tarsomere 1. Outer apical notch
with ~100° angle, notch depth moderate, equal to length of tarsomere 1. Inner apical spine equal
in length to tarsomeres 1. Protibia not heavily armored but with characteristic dense patch of
stiff setae along the inner apical region. Mesotibia more heavily armored than protibia with
more dense stiff setae and a row of numerous slender spines along entire lateral edge. Outer
apical process similar in length to protibia, and bifid apically. Inner apical spine equal in length
to tarsomeres 1 and part of 2 combined. Metatibia with armature similar to that of mesotibia, but
outer apical process slightly longer, and inner apical spine equal to tarsomeres 1-2 combined.
Male genitalia well-sclerotized. Anal sclerite with large broadly curved region
anteriodorsally that is truncate apically (Fig. 27); apex densely fimbriate and broadly rounded.
Spiculum gastrale with wide posteriorly directed lateral flanges, medial margins only deeply
concave, sparse long setae originating from apex (Fig. 66). Tegmen evenly rounded apically
(Fig. 107), longer than wide (w:l = 1.0:2.6), lateral row of setae visible from the median fossa to
around the apex, elongate shallow concavity in apical third with sinuate arranged row of small
setae not attaining the apex of the fossa, basal margin concave. Median lobe large and robust,
~0.75 the length of the tegmen, apex broadly rounded with acuminate tip, apical opening well-
developed (Fig. 148), with broadly diverging internal structure. Ejaculatory rods not fused to
basal piece, and nearly straight with slight outward orientation apically. Basal piece in two parts,
apical piece with deep medial concavity and multiple sharp projections, proximal piece separated
into two inwardly curved structures each having a medial sharp projection (Fig. 188).
195
Female genitalia moderately sclerotized. Paraprocts large with sclerotization only along
median line to apico-lateral angles. Gonocoxite with two basal lateral prominences, basal ridge
well-sclerotized. Gonocoxal apices with recurved “tooth” present. Three primary setae originate
from small depressions on the sides of the gonocoxal teeth. Introgonocoxal invagination
extremely deep, ~0.75 length of gonocoxite (Fig. 224).
Variation. None observed.
Seasonality/Habitat. Specimens collected in early may in mid-elevation forest.
Distribution. Known only from the type locality in central Honduras.
Notes. Host data include all specimens collected from a “bovista” type of
Gasteromycetes.
Etymology. Specific epithet denotes the overall globular body shape.
Pocadius helvolus Erichson
(Figs. 28, 67, 108, 149, 189, 225)
Type Material Examined. LECTOTYPE ♂ (Humboldt, Berlin): 8642 / LECTOYPUS;
Pocadius; helvolus; Erichson 1843; Jelínek det. 1971. / helvolus; Er.; Am. Sept. Auf. Kn. [The
bottom (3rd) label is hand-written and only partially legible]. The translation of the last line
“Am. Sept. Mus. Kn.” likely is “America Septrionalis Museum Knoch’schen” as revealed in part
by Erichson’s (1843: 320) original description and the typed label data of the paralectotype
below.] . PARALECTOTYPE ♀ (Humboldt, Berlin): Hist. Coll. (Coleoptera); Nr. 8642;
Pocadius helvolus Erichs.; Americ. Sept. Mus. Knoch; Zool. Mus. Berlin /
PARALECTOTYPUS; Pocadius; helvolus; Erichson 1843; Jelínek det. 1971.
Non-Type Material Examined. ~1500 specimens from throughout North America.
196
Diagnosis. Though somewhat variable in color pattern and overall body pubescence, the
following characters clearly define P. helvolus from all other species: the almost complete
alutaceous to granular surface sculpturing of the pronotum and elytra, distinct antennal sensillar
region on ventral surface of terminal antennomere, terminal antennomeres broadly evenly
rounded or truncate apically, eyes comparatively small (width ~.10 width of head), posterior face
of prosternal process when viewed laterally with slight concavity, tegmen with 1 inner row of
setae meeting near apex, median lobe with simple deeply incised apical opening, ejaculatory rods
straight with basal piece having two separate distinct curved parts adjacent to the baso-lateral
margin, and ovipositor with single baso-lateral prominence on gonocoxites and three elongate
primary setae.
Redescription. Length 3.9 mm, Width 2.1mm, Depth 1.5mm. Body moderately
convex, surface only slightly shining, reddish-brown to dark reddish brown in color, sometimes
with elytral apices and lateral area much darker brown to black. Pronotum and elytra margins
with moderately elongate fimbriae, setae longer than width of antennal scape. Dorsal and ventral
pubescence quite long.
Head surface moderately, irregularly punctate, punctures larger on vertex, becoming
somewhat smaller towards orbits and fronotclypeal region. Larger punctures 4 X diameter of
eye facet, smaller punctures 2 X diameter. Interspaces smooth to finely alutaceous, ~0.5-0.75
diameters apart, and shining becoming completely granular near occiput. Each smaller puncture
gives rise to an elongate curved golden seta, very few smaller punctures on vertex. Pronotal
surface with large punctures equal in size to large punctures on vertex of head, interspersed with
numerous smaller punctures, ~0.5-0.75 size of larger ones. Interspaces alutaceous to granular,
about 0.5-1 diameters apart. Each puncture gives rise to a long straight golden seta. Scutellar
surface with numerous shallowly impressed punctures equal to smaller punctures on pronotum,
197
some punctures giving rise to setae, with the interspaces are granular. Elytral surface with serial
rows of alternating large and small deep punctures. Smaller punctures are equal in size to
smaller ones on pronotum, larger punctures are ~2 times diameter of smaller ones. Smaller
punctures giving rise to an erect long golden seta, larger punctures giving rise to a semi-erect
long golden seta. Interspaces narrow between punctures of a given row and between different
rows. Within a row, small punctures are separated by ~1-1.5 puncture width, and large
punctures by 0.5-0.75 puncture width. Large puncture rows are separated by 1.50-2 large
puncture diameters. Interspaces only slightly shining and mostly granular in sculpture.
Pygidium densely punctate, punctures equal in size to larger ones on pronotum, each puncture
giving rise to a moderately long straight golden seta, interspaces narrow, 0.25-0.5 diameters,
with granular sculpture.
Venter with similar long golden pubescence as dorsum. Mentum with few smaller
shallow punctures, equal in size to smaller ones on vertex, each giving rise to a moderately long
seta, interspaces smooth to alutaceous. Submentum and gula similar in sparse punctation to
mentum but with interspaces becoming granular. Prosternum and epimeron with moderately
deep irregular punctation, punctures equal to large ones on vertex, interspaces almost completely
granular, prosternal punctures separated by 0.25-0.5 diameter, those on the epimeron by ~0.5-1
diameter. Mesosternum with shallow punctures, similar in size as those on prosternum,
interspaces alutaceous to granular, separated by about 0.5 to 1 diameter, and aggregated along
posterior border next to metasternum. Metasternum irregularly punctate, with moderately
impressed punctures on disc similar in size to those on mesosternum, interspaces alutaceous to
granular on metasternal disc becoming completely granular laterally, punctures separated by ~1
diameter. Abdominal sternite 1 with large faint, almost obsolete punctures, punctures equal to
those on metasternum, interspaces granular, separated by ~0.5 diameter. Abdominal sternites 2-
198
4 with two distinct irregular rows of punctures, punctures similar in size to those on
metasternum, punctures in row separated by 1 puncture diameter, other punctures on sternite
diffusedly dispersed and separated by ~0.5 punctures width. Hypopygidium with moderately
deep punctures, similar in size to those on sternites 2-4, interspaces granular, punctures separated
by 0.25-0.5 puncture diameters.
Head slightly wider than long (L:W = 1:1.34), fronotclypeal region moderately projecting
anteriorly. Vertex with broad moderately deep concavity between orbits near fronotclypeal
region. Eyes comparatively small, moderately protruding. Labrum with relatively deep medial
incision at anterior margin. Antennal club compact, oval, symmetrical with the last antennomere
subequal to the previous two combined. Antennomeres 4-5 more or less cuboid, 6 only
moderately flattened with 7-8 characteristically disc-like. Antennal scape moderately
asymmetrical, slightly hemispherical, 2 times as long as pedicel. Pedicel subcylindrical in shape.
Antennal segment 3 subequal in length to pedicel. Antennal club large, ~0.75 length of segments
1-8 combined. Each club segment with dense short setae, and several protruding setae.
Antennal grooves very deep and widely excavate, slightly converging posteriorly. Lateral
mental ridge prominent, at level of submentum, medially the ridge is granular in sculpture with
microreticulations laterally. Mentum with anterior angles broadly rounded, anterior margin
broadly hemispherical with somewhat defined apex, lateral sides short and somewhat
convergent, entire structure pentagonal and flattened.
Pronotum widest near posterior angles (L:W = 1:2), anterior margin broadly
trapezoidal, posterior margin moderately convex, lateral margins evenly arcuate to anterior
angles. Scutellum large, obtusely triangular, apex rounded. Prosternal process narrowed
between procoxae, apex acuminate, in lateral aspect the anterior and posterior ends are
prominent and convex medially. Posterior apical wall prominent and only slightly concave.
199
Mesosternum extending to midway between mesocoxae, evenly broadly concave for reception of
the metasternum. Metasternum width to length ratio is ~3:1.0. Metepisternum with slight
medial constriction, oblique line dividing anterior 0.2 of structure. First abdominal sternite with
acuminate process between metacoxae. First sternite ~2X’s longer than second sternite.
Sternites 2-3 subequal in length, the fourth slightly larger than the preceding two.
Hypopygidium subequal in length to first abdominal sternite.
Protibia with apical tooth very prominent, slightly longer than tarsomeres 1 and part of 2
combined. Outer apical notch with ~100° angle, notch depth moderate, subequal to length of
tarsomere 1. Inner apical spine subequal in length to tarsomeres 1 and part of 2 combined.
Protibia not heavily armored but with characteristic dense patch of stiff setae along the inner
apical region. Mesotibia more heavily armored than protibia with more dense stiff setae and a
row of numerous slender spines along entire lateral edge. Outer apical process elongate and
robust, larger than protibia process, and bifid apically. Inner apical spine equal in length to
tarsomeres 1-2 combined. Metatibia with armature similar to that of mesotibia, but outer apical
process slightly longer.
Male genitalia well-sclerotized. Anal sclerite with large broadly curved region
anteriodorsally (Fig. 28); apex fimbriate and acuminate; ventrally with a broad medial concavity
approaching apex in truncate fashion. Spiculum gastrale with wide lateral flanges, medial
margins only slightly concave proximally, sparse long setae originating from apex (Fig. 67).
Tegmen evenly rounded apically (Fig. 108), somewhat longer than wide (w:l = 1.0:1.6), lateral
row of setae visible from the median fossa to around the apex, small shallow concavity in apical
third, basal margin angulate, inner row of setae complete and meeting near apex. Median lobe
large and robust, ~0.50 the length of the tegmen, apex broadly rounded, apical opening with
well-developed medial opening (Fig. 149). Ejaculatory rods not fused to basal piece, and nearly
200
straight. Basal piece with deep medial concavity extending almost two-thirds length of structure,
proximally with broad basal convexity and two separate lateral pieces that are inwardly curved
(Fig. 189).
Female genitalia moderately sclerotized. Paraprocts large with sclerotization only along
median line. Gonocoxite with one basal lateral prominence, basal ridge well-sclerotized.
Gonocoxal apices with recurved “tooth” present. Three primary setae originate from small
depressions on the sides of the gonocoxal teeth (Fig. 225).
Variation. Specimens from Colorado, Manitoba, and the Northwest Territory have the
ejaculatory rods more slender and slightly inwardly curved, however, other genitalic features are
similar.
Seasonality/Habitat. Specimens can be collected nearly year-round depending on the
collecting locality. Specimens from the southern U.S. and Mexico have been collected in all
months except January and February, with most specimens collected from April through
October.
Distribution. Known range extends from the Northwest Territories, Saskatchewan,
Manitoba and Quebec, Canada south through the eastern U.S. to central Florida in the east and
central Mexico in the west. This range includes all states on or east of the 100th parallel in the
U.S.A, as well as the following western states that may lie west of that line: Nebraska, Montana,
Wyoming, and Arizona. Mexico records were indicated by Parsons (1943) as Puente de Ixtla,
Durango and Guanajuata. New records from Mexico include the following: (78) from CAS:
Cocula; Jalisco, Mex.; VII-2-60; in puffball; L.R. Gillogly collr. (41) from CAS: Cocula;
Jalisco, Mex.; VII-22-60; in puffball; L.R. Gillogly collr. (29) from CAS: Guadalajara; Jalisco,
Mex.; 9 July 1960; in puffball; Gilloglys collrs. (3) from CAS: Delgollado, Jalisco; Mexico,
VII-2-1963; in puffball; L.R. Gillogly collr. (8) from CAS: LaVenotsa, Oaxaca; Mexico, VII-
201
19-1963; ex. shaggy-mane; L.R. Gillogly collr. (3) from CAS: Tapalpa; Jalisco; Mexico; VII-4-
1960; ex. fungi; L.R. Gillogly collr. (1) from University of Guadalajara: MEXICO: Jalisco,
Zapopan; Nextipac, Las Agujas, CUCBA; 1600m, 14-VII-1994; ex. Lycoperdon, D. Pérez col.
(17) from Montana St. University: MEX.: Jalisco, 20mi.; SW Encarnacion de Diaz; 09 JUL
1982, 2000m; M.A. Ivie colr.
Notes. Specimens have been collected from numerous Lycoperdaceae as well as from
“gilled fungi” and fungus covered logs.
Pocadius insularis Cline new species
(Figs. 29, 68, 109, 150, 190)
Type Material Examined. HOLOTYPE ♂ (CMN): Simla, 5 mi. N.; Arima, Trinidad;
W.I. Aug.20, 1969; H. & A. Howden / HOLOTYPE; Pocadius; insularis; A.R. cline des. 2004.
Diagnosis. This species is most similar to P. dimidiatus but can be differentiated from it
and the rest of the Neotropical fauna by the following suite of characters: nearly circular antennal
club without enlarged terminal antennomere; pronotum widest near posterior angles with anterior
margin broadly shallowly concave; mentum transversely hemispherical; prosternal process in
lateral view highly convex over procoxal cavities and with strongly oblique apical wall; overall
minute body size; and aedeagus with tegmen having a complete inner row of setae, nearly
perpendicular basal margin; basal piece of internal sac sclerites with three distinct parts.
Description. Length 3.3mm, Width 1.9mm, Depth 1.2mm. Body moderately convex,
surface slightly shining, uniformly light reddish-brown in color. Pronotum and elytra margins
with moderately long sparse fimbriae, setae subequal to length of antennal scape. Dorsal and
ventral pubescence short to moderately long, dorsal surface with heavily granulate and
microreticulate sculpturing.
202
Head surface deeply, irregularly punctate, punctures larger on vertex, becoming
somewhat smaller towards orbits and fronotclypeal region. Larger punctures 3-4X diameter of
eye facet, smaller punctures 1-2X diameter. Interspaces granular with microreticulations,
sparsely shining. Each puncture gives rise to a moderately long curved golden seta. Pronotal
surface with large punctures equal in size to large punctures on vertex of head, interspersed with
relatively few smaller punctures, ~0.5 size of larger ones. Interspaces granular with
microreticulations, about 0.5-1 diameter apart. Each puncture gives rise to a moderately long
curved golden seta. Scutellar surface with few shallowly impressed punctures, some punctures
giving rise to setae, and the interspaces are granular. Elytra with serial rows of alternating large
and small deep punctures. Smaller punctures are equal in size to smaller ones on pronotum,
larger punctures are ~2.5 times diameter of smaller ones. Smaller punctures giving rise to a
semi-erect long golden seta, larger punctures giving rise to a short to minute semi-erect golden
seta. Interspaces narrow between punctures of a given row and between different rows. Within
a row, small punctures are separated by ~1 puncture width, and large punctures by 0.5 puncture
width. Larger rows are separated by 0.5-1.0 large puncture diameters. Interspaces sparsely
shining with granular to microreticulate sculpture. Pygidium densely punctate, punctures equal
in size to larger ones on pronotum, each puncture giving rise to a moderately long golden seta.
Interspaces narrow, 0.25-0.5 diameter, with granular sculpture.
Venter with similar moderately long pubescence as dorsum. Mentum with large shallow
punctures, equal in size to larger ones on vertex, each giving rise to a moderately long seta.
Interspaces granular to finely microreticulate. Submentum and gula punctures somewhat smaller
than mentum but with interspaces similar. Prosternum and epimeron faintly irregularly punctate,
punctures subequal to those on mentum, interspaces granular with microreticulate areas,
prosternal punctures separated by 0.5-2 diameters, those on the epimeron by ~0.5-1 diameter.
203
Mesosternum with shallow punctures, subequal to those on prosternum, interspaces alutaceous to
granular, separated by about 0.5 to 1 diameter, and aggregated near metasternal border.
Metasternum irregularly punctate, with moderate faint punctures on disc similar in size to
smaller ones on vertex, interspaces dull on metasternal disc becoming more microreticulate to
granular laterally, punctures separated by ~1-3 diameters on disc. Abdominal sternite 1 with
small faint, almost obsolete punctures, punctures equal to those on metasternum, interspaces
mostly granular, separated by ~1-2 diameters. Abdominal sternites 2-4 with irregular rows of
punctures, punctures similar in size to those on metasternum, rows separated by 1.5-2 puncture
diameters, punctures within rows separated by ~0.5 punctures width. Hypopygidium with
moderately deep punctures, similar in size to those on sternites 2-4, interspaces granular with
microreticulations, punctures separated by 0.5-1 diameter.
Head slightly wider than long (W:L = 1.27:1), fronotclypeal region moderately projecting
anteriorly. Vertex with deep concavity between orbits near fronotclypeal region. Labrum with
shallow concavity at anterior margin. Antennal club compact, circular, symmetrical with the last
antennomere subequal to the previous two combined. Antennomeres 4-8 more or less compact,
with 7-8 characteristically disc-like, 4-5 cuboidal, and 6 trapezoidal. Antennal scape
asymmetrical, hemispherical, 1.75 times as long as pedicel. Pedicel similar in shape to scape but
not as hemispherical. Antennal segment 3 longer in length to pedicel. Antennal club relatively
small, ~0.65 length of segments 1-8 combined. Antennal grooves very deep and widely
excavate, slightly converging posteriorly. Lateral mental/submental ridge prominent, at level of
submentum, ridge and corresponding sulcus with longitudinal microreticulations. Mentum with
anterior angles obsolete, entire structure broadly hemispherical when viewed ventrally and
flattened in lateral aspect.
204
Pronotum widest near posterior angles (L:W = 1:1.7), anterior margin broadly
shallowly concave, posterior margin moderately convex, lateral margins slightly arcuate
anteriorly. Scutellum moderately large, obtusely triangular, apex rounded. Prosternal process
narrowed between procoxae, apex acuminate, in lateral aspect the anterior and posterior ends are
prominent and there is a steep convexity medially. Posterior apical wall prominent and slightly
oblique. Mesosternum extending to midway between mesocoxae, evenly concave for reception
of the metasternum. Metasternum wider than long (W:L = 2.6:1.0). Metepisternum with slight
medial constriction, oblique line dividing anterior 0.17 of structure. First abdominal sternite with
broad process between metacoxae. First sternite ~2X’s longer than second sternite. Sternites 2-3
subequal in length, the fourth slightly larger than the preceding two. Hypopygidium subequal in
length to first abdominal sternite.
Protibia with apical tooth not prominent, subequal to tarsomere 1. Outer apical notch
with ~95° angle, notch depth moderate, subequal to length of tarsomere 1. Inner apical spine
subequal in length to tarsomeres 1 and part of 2 combined. Protibia not heavily armored but with
characteristic dense patch of stiff setae along the inner apical region. Mesotibia more heavily
armored than protibia with more dense stiff setae and a row of numerous slender spines along
entire lateral edge. Outer apical process robust, larger than protibia process, subequal to
tarsomeres 1-2 combined. Inner apical spine equal in length to tarsomeres 1-2 combined.
Metatibia with armature similar to that of mesotibia.
Male genitalia well-sclerotized. Anal sclerite with large broadly curved region
anteriodorsally (Fig. 29); apex densely fimbriate; ventrally with a broad medial concavity
approaching apex in a truncate manner. Spiculum gastrale with wide lateral flanges, medial
margins deeply concave, moderately long stiff setae originating from apex (Fig. 68). Tegmen
evenly rounded apically (Fig. 109), much longer than wide (w:l = 1:2.6), lateral row of setae
205
visible from the median fossa to around the apex, inner row of setae complete, small shallow
concavity in apical third, basal notch perpendicular, basal margin nearly perpendicular. Median
lobe large and robust, ~0.66 the length of the tegmen, apex narrowly rounded, apical opening
well-developed with deeply cleft bilobed internal structure (Fig. 150). Ejaculatory rods not fused
to basal piece or each other, curved inward and expanded outward basally and apically. Basal
piece of internal sac sclerites with two short paired basal lateral sections each having a medial
projection, and a central section that is concave apically and has a basally projecting extension
(Fig. 190).
Female genitalia not observed.
Variation. Known only from the holotype male.
Seasonality/Habitat. Known from late August in a lowland tropical forest.
Distribution. Known from the type locality on the island of Trinidad.
Notes. No host information is available for this species.
Etymology. Specific epithet is a derivative of the Latin “insula” meaning “living on an
island”, alluding to the type locality of the island of Trinidad.
Pocadius jelineki Leschen and Carlton
(Figs. 30, 69, 110, 151, 191, 226)
Type Material Examined. HOLOTYPE ♂ (SNEC): COSTA RICA: Puntarenas; Monte
Verde, 1580m; 13 May 1989, J. Ashe; R. Brooks, R. Leschen; ex. Lycoperdon sp. / Snow
Entomol. Mus.; Costa Rica Exped. #159 / HOLOTYPE; Pocadius jelineki; R. Leschen & C.
Carlton. 6 PARATYPES (SNEC): same data labels as holotype. 2 PARATYPES (SNEC):
COSTA RICA:; Puntarenas, Monte Verde; 1400m, 14-16-July-1989; ex. flight intercept trap;
Robert E. Beer. 1 PARATYPE (SNEC): COSTA RICA: Puntarenas; Monte Verde, 1550m; 22
206
May 1989, J. Ashe; R. Brooks, R. Leschen; ex. Lycoperdon / Snow Entomol. Mus.; Costa Rica
Exped. #375. 1 PARATYPE (SNEC): COSTA RICA: Puntarenas; Reserva Biologica de;
Monteverde, 1550m; nr. Quelorada cuecha / on Sendero rio; 13 May 1989, J. Ashe; R. Brooks,
R. Leschen; ex. Lycoperdon / Snow Entomol. Mus.; Costa Rica Exped. #160.
Non-Type Material Examined. >100 specimens: 16 specimens (CMN) Costa Rica:
Punt., Monteverde, 1400m, 21-24-VIII, H. & A. Howden, FIT. Costa Rica: Monteverde, flight
intercept, 1520m, 18-25-VI-1983, D.H. Lindeman. Costa Rica: Punt., Monteverde, 1400m, 18-
20-VIII-1987, H. & A. Howden, FIT. Costa Rica: Monteverde, unbaited pit traps, 3-VII-1983,
D.H. Lindeman. Costa Rica: 1520m, Monteverde, FIT, 9-13-VII-1983, D.H. Lindeman. Costa
Rica: 1520m, Monteverde, FIT, 2-9-VII-1983, D.H. Lindeman. Costa Rica: Punt., Monteverde,
1400m, 15-17-VIII-1987, H. & A. Howden. Costa Rica: Monteverde, flight intercept, 1520m,
11-18-VI-1983, D.H. Lindeman. 8 specimens (SNEC): COSTA RICA: Guanacaste; Cacao
Biological Station, 1050m; 10°55’38”N, 85°27’7”W; 10 JUL 2000; J. Ashe, R. Brooks, Z. Falin;
CR1ABF00 089; ex: in giant puffball mushroom. 5 specimens (SNEC): NICARAGUA:
Granada Dept.; Res. Nat. Volcan Mombacho, 1150m, 11°50.05’N 85°58.83’W; 3-VI-2002, R.
Brooks, Z. Falin, S. Chatzimanolis, ex. puffball fungus; NIC1BFC02 171.
Diagnosis. In the original diagnosis two characters were given to distinguish this
species: the curved apices of the setae and the elbowed posterior ejaculatory rods. The first of
these characters is seen in numerous other Pocadius and therefore is of little diagnostic value and
the latter is only useful for discerning males of the species. Other characters that more
thoroughly define the species include: metepisternal axillary space very narrow (~.10 length of
structure), antennal segment 2-3 elongate, terminal antennomere with characteristic sensillar
region, eyes comparatively large and protruding, pronotum widest near middle, metatibial spurs
elongate, eighth abdominal sternite with elongate narrow lateral flanges projecting anteriorly,
207
median lobe with complex apical opening, and ovipositor with narrowed paraprocts, gonocoxal
base with one lateral prominence, inner gonocoxal margin angulate, gonocoxal apices lacking
“tooth”, and 4-5 primary elongate setae projecting from the gonocoxites apices.
Redescription. Length 3.1 mm, Width 1.7mm, Depth 1.1mm. Body moderately
convex, surface shining, brown to dark brown in color, venter distinctly lighter. Pronotum and
elytra margins with elongate fimbriae, setae longer than width of antennal scape. Dorsal and
ventral pubescence quite long.
Head surface deeply, irregularly punctate, both small and large punctures on vertex,
becoming proportionately smaller towards orbits and fronotclypeal region. Larger punctures 4 X
diameter of eye facet, smaller punctures 3 X diameter. Interspaces smooth to finely alutaceous,
shining. Each smaller puncture gives rise to an moderately long curved golden seta. Eyes finely
faceted. Pronotal surface with large punctures equal in size to large punctures on vertex of head,
interspersed with relatively few smaller punctures, ~0.75 size of larger ones. Interspaces smooth
to alutaceous, about 0.5-0.75 diameters apart. Each puncture gives rise to a moderately long
golden seta, most seta are curved but some are rather straight. Scutellar surface with very few
moderately impressed punctures equal to smaller ones on pronotum, some punctures giving rise
to short golden decumbent setae, interspaces smooth to alutaceous. Elytral surface with serial
rows of alternating large and small deep punctures. Smaller punctures are equal in size to
smaller ones on pronotum, larger punctures are ~1.75 times diameter of smaller ones. Smaller
punctures giving rise to semi-erect longer curved golden seta, larger punctures giving rise to a
decumbent long curved golden seta. Interspaces narrow between punctures of a given row and
between different rows. Within a row, small punctures are separated by ~1 puncture width, and
large punctures by 0.25-0.5 puncture diameters. Larger puncture rows are separated by 1 larger
puncture diameter. Interspaces always shining but variable from smooth to finely alutaceous in
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sculpture. Pygidium densely punctate, punctures equal in size to smaller ones on vertex, each
puncture giving rise to a short straight golden seta. Interspaces narrow, 0.75-1.0 diameters, with
smooth to alutaceous sculpture.
Venter with short more sparsely distributed pubescence as dorsum. Mentum with small
shallowly impressed punctures, equal in size to smaller ones on vertex, each giving rise to a short
straight seta, interspaces alutaceous. Submentum and gula similar in punctation to mentum but
with interspaces alutaceous to microreticulate. Prosternum and epimeron deeply irregularly
punctate, punctures 1.5-1.75X larger than those on mentum, interspaces alutaceous with
microreticulate areas, prosternal punctures separated by 0.5 diameter, those on the epimeron by
0.33 to 0.66 diameter. Mesosternum with shallow punctures, about 0.75 diameter of those on
prosternum, interspaces alutaceous to microreticulate, separated by about 0.5 to 1 diameter,
aggregated near posterior margin. Metasternum irregularly punctate, with moderate faint
punctures on disc similar in size to those on prosternum, interspaces smooth to alutaceous on
metasternal disc becoming more microreticulate laterally, punctures separated by ~1-2 diameters.
Abdominal sternite 1 with large faint, almost obsolete punctures, punctures equal to those on
mesosternum, interspaces smooth to alutaceous, separated by ~1 diameter. Abdominal sternites
2-4 with two irregular rows of punctures, one row near posterior margin, punctures similar in
size to those on abdominal sternite 1, rows separated by ~1-1.5 puncture diameters, punctures
within rows separated by ~0.33 punctures diameters. Hypopygidium with moderately deep
punctures, similar in size to those on sternites 2-4, interspaces alutaceous, punctures separated by
0.25 puncture diameter.
Head slightly wider than long (L:W = 1:1.6), fronotclypeal region moderately projecting
anteriorly. Vertex with broad shallow concavity between orbits near fronotclypeal region.
Labrum with deep median incision at anterior margin. Antennal club compact, obovate,
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distinctly asymmetrical with the last antennomere much greater in length than the previous two
antennomeres combined. Antennomeres 6-8 more or less compact, with 7-8 characteristically
disc-like. Antennomeres 4-5 more or less cuboid, slightly narrowed proximally Antennal scape
asymmetrical, somewhat hemispherical, 1.4 times as long as pedicel. Pedicel elongate
cylindrical in shape. Antennal segment 3 subequal in length to pedicel. Antennal club large,
~0.95 length of segments 1-8 combined. Each club segment with dense short setae, terminal
antennomeres with deep circular sensillar region. Antennal grooves very deep and widely
excavate, slightly converging posteriorly. Lateral mental ridge prominent, at level of
submentum, medially ridge has setal bearing small punctures and alutaceous to granular
interspaces and laterally with longitudinal to oblique microreticulations. Mentum with anterior
angles apparent, anterior margin broadly hemispherical with acuminate apex, lateral margins
short and perpendicular, entire structure pentagonal and somewhat convex.
Pronotum widest near middle (L:W = 1:2.1), anterior margin broadly shallowly
trapezoidal, posterior margin moderately convex, lateral margins less arcuate posteriorly.
Scutellum large, triangular, apex narrowly rounded. Prosternal process somewhat narrowed
between procoxae, apex somewhat acuminate, in lateral aspect the anterior and posterior ends are
prominent and convex medially. Posterior apical wall short but prominent and only slightly
oblique. Mesosternum extending to midway between mesocoxae, evenly broadly concave for
reception of the metasternum. Metasternum width to length ratio is ~2.67:1.0. Metepisternum
rather broad with two medial constrictions, oblique line dividing anterior 0.10 of structure.
Elytral humeri moderately produced, lateral margin very narrow. First abdominal sternite with
broadly rounded process between metacoxae. First sternite ~2X’s longer than second sternite.
Sternites 2-4 subequal in length. Hypopygidium subequal in length to first abdominal sternite.
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Protibia with apical tooth very prominent, slightly longer than tarsomeres 1 and part of 2
combined. Outer apical notch with ~100° angle, notch depth shallow, equal to length of
tarsomere 1. Inner apical spine subequal in length to tarsomeres 1-2 combined. Protibia heavily
armored with characteristic dense patch of stiff setae along the inner apical region. Mesotibia
more heavily armored than protibia with more dense stiff setae and a row of numerous slender
spines along entire lateral edge. Outer apical process elongate and robust, larger than protibia
process. Inner apical spine equal in length to tarsomeres 1-2 combined. Metatibia with more
armature than that of mesotibia, including more and longer spines, and apical spur equal to
tarsomeres equal in length to one two and part third tarsomere combined.
Male genitalia well-sclerotized. Anal sclerite with large broadly curved region
anteriodorsally (Fig. 30); apex fimbriate; ventrally with a broad medial concavity approaching
apex. Spiculum gastrale with narrow anteriorly projecting lateral flanges, medial margins
concave proximally, numerous short stiff setae originating from apex (Fig. 69). Tegmen evenly
rounded apically (Fig. 110), much longer than wide (w:l = 1.0:2.55), lateral row of setae visible
from the median fossa around the apex, small shallow concavity in apical third, basal notch
perpendicular, basal margin slightly concave, inner row of setae complete coming together near
apex. Median lobe moderately large, ~0.50 the length of the tegmen, apex acuminate, apical
opening well-developed (Fig. 151). Ejaculatory rods not fused to basal piece or each other,
mostly straight and perpendicular. Basal piece of rods visible as two distinct elbowed parts (Fig.
191).
Female genitalia moderately sclerotized. Paraprocts narrow with concave lateral
margins, sclerotization only along median line to gonocoxal base. Gonocoxite with one basal
lateral prominence, basal ridge well-sclerotized. Gonocoxal apices without recurved “tooth”.
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Four or five primary setae originate from small depressions laterally on the gonocoxal apices
(Fig. 226).
Variation. Some variability in size of specimens, though not directly correlated to
gender.
Seasonality/Habitat. The species is known from the mid-elevation tropical wet forests
from mid-May through late July.
Distribution. Known from Costa Rica and southern Nicaragua.
Notes. Specimens are known from Lycoperdon puffballs.
Pocadius kirejtshuki Cline new species
(Figs. 31, 70, 111, 152, 227)
Type Material Examined. HOLOTYPE ♀ (BMNH): AUSTRALIA; C.E. Clarke;
Collection; B.M. 1957-24 / Katanda; 24-6-38; C.E.C. / Katanda / HOLOTYPE; Pocadius;
kirejtshuki; A.R. Cline des. 2004. 2 PARATYPES (BMNH): AUSTRALIA; C.E. Clarke;
Collection; B.M. 1957-24 / Katanda / PARATYPE; Pocadius; kirejtshuki; A.R. Cline des. 2004.
1 PARATYPE (BMNH): Moreton Bay / Pocadius; sp.; det. Jelínek , 1970 / PARATYPE;
Pocadius; kirejtshuki; A.R. Cline des. 2004.
Diagnosis. Distinct from the other Old World fauna by the following suite of characters:
thickened tarsomeres; sides and apex of elytra darker than rest of body; terminal antennomere
enlarged, greater than previous two combined, and with sharply pointed apex; apical wall of
prosternal process oblique; aedeagus with tegmen having an elliptical apical fossa with few
elongate thick associated setae; median lobe with prolonged acuminate apex and complex
internal structure; spiculum with numerous ridges along the apical border; and ovipositor with
gonocoxal apices abutting distally with numerous primary setae originating apico-laterally..
212
Description. Length 4.2mm, Width 2.25mm, Depth 1.35mm. Body slightly convex,
surface moderately shining to dull, reddish-brown to dark brown in color, with elytral sides and
apex darker. Pronotum and elytra margins with moderately long fimbriae, setae subequal to
length of antennal scape. Dorsal and ventral pubescence moderately long.
Head surface with moderately impressed irregular punctures, punctures larger on vertex,
becoming somewhat smaller towards orbits and fronotclypeal region. Larger punctures 3-4X
diameter of eye facet, smaller punctures 1-2X diameter. Interspaces granular with some
microreticulation, slightly shining. Each puncture gives rise to a moderately long curved golden
seta. Pronotal surface with large punctures equal in size to large punctures on vertex of head,
interspersed with relatively few smaller punctures, ~0.5 size of larger ones. Interspaces granular
to finely microreticulate, about 0.5-1 diameter apart. Each puncture gives rise to a moderately
long golden seta, most seta curved. Scutellar surface with few vague shallowly impressed
punctures, some punctures giving rise to setae, and the interspaces nearly completely
microreticulate. Elytral surface with serial rows of alternating large and small deep punctures.
Smaller punctures are equal in size to those on pronotum, larger punctures are ~2-3 times
diameter of smaller ones. Smaller punctures giving rise to a semi-erect to decumbent
moderately long golden seta, larger punctures giving rise to a semi-erect to decumbent long
golden seta. Interspaces narrow between punctures of a given row and between different rows.
Within a row, small punctures are separated by ~1-2 puncture diameter, and large punctures by
0.5-1 puncture diameter. Larger rows are separated by ~1.0 large puncture diameter. Interspaces
slightly shining but variable from granular to finely microreticulate in sculpture. Pygidium
densely punctate, punctures equal in size to larger ones on pronotum, each puncture giving rise
to a short golden seta. Interspaces narrow, 0.5-0.75 diameter, with granular to microreticulate
sculpture.
213
Venter with similar pubescence as dorsum. Mentum with minute shallow punctures,
equal in size to smaller ones on vertex, each giving rise to a short seta. Interspaces granular to
finely microreticulate in areas. Submentum and gula similar in punctation to mentum but with
interspaces more microreticulate. Prosternum and epimeron shallowly irregularly punctate,
punctures slightly larger than those on mentum, interspaces granular with microreticulate areas,
prosternal punctures separated by 1-3 diameters, those on the epimeron by 2-4 diameters.
Mesosternum with faint shallow punctures, equal in diameter to those on prosternum, interspaces
granular, separated by about ~1 diameter, and aggregated near metasternum. Metasternum
faintly irregularly punctate with minute punctures on disc similar in size to those on mentum,
interspaces granular on metasternal disc becoming granular with microreticulations laterally,
punctures separated by ~2-3 diameters. Abdominal sternite 1 with large faint punctures, equal to
large punctures on elytra, interspaces granular to microreticulate, separated by ~1-2 diameters.
Abdominal sternites 2-4 with irregular rows of punctures, one defined row near anterior margin,
punctures similar in size to those on abdominal sternite 1. Hypopygidium with moderately deep
punctures, similar in size to those on sternites 2-4, interspaces granular to microreticulate,
punctures separated by 0.5-1 diameter.
Head wider than long (W:L = 1.65:1), fronotclypeal region moderately projecting
anteriorly. Vertex with shallow concavity between orbits near fronotclypeal region. Labrum
with shallow concavity at anterior margin. Antennal club compact, obovate with acute apical
point, somewhat asymmetrical with the last antennomere longer than the previous two combined.
Antennomeres 5-8 more or less compact, with 7-8 characteristically disc-like, 5-6 trapezoidal,
and 4 cuboidal. Antennal scape asymmetrical, somewhat hemispherical, 1.9 times as long as
pedicel. Pedicel barrel-shaped. Antennal segment 3 longer in length than pedicel. Antennal club
large, ~0.80 length of segments 1-8 combined. Antennal grooves very deep and widely
214
excavate, distinctly converging posteriorly. Lateral mental/submental ridge prominent, at level
of submentum, ridge with few longitudinal microreticulations and the corresponding sulcus with
none. Mentum with anterior angles present, anterior margin with acute apex, entire structure
somewhat compactly pentagonal when viewed ventrally and flattened when viewed laterally.
Pronotum widest near middle (L:W = 1:1.9), anterior margin broadly concave with the
central region slightly convex, posterior margin moderately convex, lateral margins somewhat
more arcuate anteriorly. Scutellum large, hemispherical, apex broadly rounded. Prosternal
process somewhat narrowed between procoxae, apex somewhat acuminate, in lateral aspect the
posterior end is prominent with a slight convexity medially. Posterior apical wall prominent and
only slightly oblique. Mesosternum extending to midway between mesocoxae, evenly concave
for reception of the metasternum. Metasternum width to length ratio is ~2.6:1.0. Metepisternum
with slight medial constriction, oblique line dividing anterior 0.10 of structure. First abdominal
sternite with moderately narrow process between metacoxae. First sternite ~2X’s longer than
second sternite. Sternites 2-3 subequal in length, the fourth slightly larger than the preceding
two. Hypopygidium subequal in length to first abdominal sternite.
Protibia with apical tooth moderately prominent, slightly longer than tarsomeres 1. Outer
apical notch absent. Inner apical spine subequal in length to tarsomeres 1 and part of 2
combined. Protibia not heavily armored but with characteristic dense patch of stiff setae along
the inner apical region. Mesotibia more heavily armored than protibia with more dense stiff
setae and a row of numerous slender spines along entire lateral edge. Outer apical process
robust, larger than protibia process. Inner apical spine equal in length to tarsomeres 1-2 and part
of 3 combined. Metatibia with armature similar to that of mesotibia.
Male genitalia well-sclerotized. Anal sclerite with large broadly curved region
anteriodorsally (Fig. 31) and recurved convexity apically; apex densely fimbriate. Spiculum
215
gastrale with wide posterior projecting lateral flanges, medial margins convex, short stiff setae
originating from finely ridged apex (Fig. 70). Tegmen narrowly rounded apically (Fig. 111),
much longer than wide (w:l = 1.0:3.1), sparse lateral row of setae visible from the median fossa
to prior to the apex, elliptical shallow concavity in apical third with few thick associated setae,
basal notch perpendicular, basal margin deeply concave. Median lobe large and robust but not
elongate, ~0.45 the length of the tegmen, apex acuminate and slightly prolonged, apical opening
well-developed with large complex internal structure (Fig. 152). Ejaculatory rods not observed.
Female genitalia moderately sclerotized. Paraprocts large with sclerotization only along
median line. Gonocoxite with two basal lateral prominences, basal ridge well-sclerotized.
Gonocoxal apices with recurved “tooth” absent. Several primary setae originate from small
depressions on the gonocoxal apices. Gonocoxal apices abutting in distal third of structure (Fig.
227).
Variation. Some specimens are uniformly brown without darkened elytral apices.
Seasonality/Habitat. Collected in June in wet forests.
Distribution. Known from the type localities in Australia.
Notes. No host information is available for this species.
Etymology. Specific epithet honors Alexander Kirejtshuk, research scientist at the
Zoological Institute in St. Petersburg, a world authority on Nitidulidae.
Pocadius luisalfredoi Cline new species
(Figs. 32, 71, 112, 153, 192, 228)
Type Material Examined. HOLOTYPE ♂ (Univ. of Guadelajara): MEXICO: México,
Atlanta; Sn. J. Tepecoculco; 29-IX-1991; Col. J.L. Navarrete #1108; ex. L. aff. compactum /
HOLOTYPE; Pocadius; luisalfredoi; A. Cline des. 2004. 4 PARATYPES (2 Univ. of
216
Guadelajara; 2 LSAM): MEXICO: México; Atlanta, San Juan; Tepecoculco; 5-X-1991 / ex.
Lycoperdon; aff. compactum; #1110 / PARATYPE; Pocadius; luisalfredoi; A. Cline des. 2004.
1 PARATYPE (Univ. of Guadelajara): MEXICO: Guerrera; km. 10 carr. Teti; pac, “El Peral”;
23-VIII-1986 / Bosque Mesopilo; 2180 msnm; ex. Lycoperdon; #72 / PARATYPE; Pocadius;
luisalfredoi; A. Cline des. 2004. 2 PARATYPES (CAS): Cerro de Garnica; Mich., Mex., VII-
25-1963 / puffball / G.M. Gillogly collector / L.R. Gillogly Collection / Lorin R. Gillogly;
Collection; Donated To The; Calif. Academy Of Sciences; May 1990 / PARATYPE; Pocadius;
luisalfredoi; A. Cline des. 2004. 2 PARATYPES (CAS): same as previous but J.J. Gillogly
collector. 5 PARATYPES (CAS): Summit bet.; Mexico and Puebla / VIII-14-1965; in puffball /
L.R. Gillogly Collector / L.R. Gillogly Collection / Lorin R. Gillogly; Collection; Donated To
The; Calif. Academy Of Sciences; May 1990 / PARATYPE; Pocadius; luisalfredoi; A. Cline des.
2004. 15 PARATYPES (CAS): 15mi. E. of; Zitauaro, Mex.; VIII-12-1965 / gill; fungus / L.R.
Gillogly Collector / L.R. Gillogly Collection / Lorin R. Gillogly; Collection; Donated To The;
Calif. Academy Of Sciences; May 1990 / PARATYPE; Pocadius; luisalfredoi; A. Cline des.
2004. 6 PARATYPES (CAS): Paracutin / July 15, 1960 / puffball / A.R. Gillogly Collector /
L.R. Gillogly Collection / Lorin R. Gillogly; Collection; Donated To The; Calif. Academy Of
Sciences; May 1990 / PARATYPE; Pocadius; luisalfredoi; A. Cline des. 2004. 2 PARATYPES
(CAS): Paracho; Mich., Mex.; July 15, 1960 / Boletus / A.R. Gillogly Collector / L.R. Gillogly
Collection / Lorin R. Gillogly; Collection; Donated To The; Calif. Academy Of Sciences; May
1990 / PARATYPE; Pocadius; luisalfredoi; A. Cline des. 2004. 31 PARATYPES (CAS): 27mi.
N.; Guadalajara; Mex. VIII-65 / puffball / L.R. Gillogly Collector / L.R. Gillogly Collection /
Lorin R. Gillogly; Collection; Donated To The; Calif. Academy Of Sciences; May 1990 /
PARATYPE; Pocadius; luisalfredoi; A. Cline des. 2004. 8 PARATYPES (Univ. Guadalajara):
MEXICO: Jalapa; Zapopan, Bosque la; Primavera; Rancho Tres Rios; 20°41’1.4”N,
217
103°36’32.8”W; 28 August 1999 / collected by: R. Leschen & J.L. Navarette-Heredia; on
Lycoperdon; pyriforme / PARATYPE; Pocadius; luisalfredoi; A. Cline des. 2004.
Diagnosis. This species is more similar to others from the Nearctic fauna, including P.
niger, P. basalis, and P. fulvipennis than it is to those of the Neotropical fauna. However, it
differs from the other Nearctic members by the following suite of characters: characteristic color
pattern with dark pronotal stripe and large light colored region on elytra; densely deeply punctate
head and pronotum with smooth surface sculpturing; elongate pronotal and elytral fimbriae;
disproportionately long and slender tibiae; hemispherical mentum; elytral pubescence with
alternating rows of elongate decumbent apically curved golden setae; male pygidium indentate
along posterior margin; tegmen with complete row of inner setae attaining apex; median lobe
robust and globular with correspondingly large apical opening and simple internal structure;
ejaculatory rods oriented in a converging manner and with internal sac basal piece “U-shaped”;
and ovipositor with deep widely divergent gonocoxal appendages bearing two primary setae in
depressions next to apical tooth.
Description. Length 3.6mm, Width 1.8mm, Depth 1.2mm. Body moderately convex,
surface shining, dark brown-black with the following areas much lighter in color: legs, lateral
third of pronotal margins, middle area of elytra, and transverse portions of each abdominal
sternite. Pronotum and elytra margins with extremely elongate fimbriae, setae longer than 2X
length of antennal scape. Dorsal and ventral pubescence sparse but quite long.
Head surface deeply, irregularly punctate, punctures mostly larger on vertex, becoming
somewhat smaller towards orbits and fronotclypeal region. Larger punctures 5-6 X diameter of
eye facet, smaller punctures ~3X diameter. Interspaces smooth to finely alutaceous, shining.
Each puncture gives rise to an elongate curved golden seta. Pronotal surface with large
punctures equal in size to large punctures on vertex of head, interspersed with numerous smaller
218
punctures, equal to smaller ones on head. Interspaces smooth to finely alutaceous, about 1-2
diameters apart. Each puncture gives rise to an elongate curved golden seta. Scutellar surface
with numerous moderately impressed punctures, most punctures giving rise to elongate setae,
interspaces smooth. Elytral surface with serial rows of alternating large and small deep
punctures. Smaller punctures are equal in size to smaller ones on pronotum, larger punctures are
~2-3X diameter of smaller ones. Smaller punctures giving rise to a semi-erect elongate curved
golden seta, larger punctures giving rise to a semi-erect elongate curved golden seta. Interspaces
broad between punctures of a given row and between different rows. Within a row, small
punctures are separated by 3-4 puncture diameters, and large punctures by ~1 puncture diameter.
Larger rows are separated by 2-3 puncture diameters. Interspaces always shining but variable
from smooth to finely alutaceous in sculpture. Pygidium densely punctate, punctures equal in
size to larger ones on pronotum, each puncture giving rise to a moderately long golden seta;
interspaces narrow, 0.25-0.5 diameter, with smooth to microreticulate sculpture.
Venter with much shorter more sparsely distributed golden pubescence than dorsum.
Mentum with few small very shallow punctures, equal in size to smaller ones on vertex;
interspaces alutaceous with finely microreticulate areas. Submentum and gula similar in
punctation to mentum but with interspaces more granular than alutaceous. Prosternum and
epimeron shallowly irregularly punctate, punctures slightly larger than those on mentum,
interspaces alutaceous to granular with microreticulate areas, prosternal punctures separated by
2-4 diameters, those on the epimeron by 1-2 diameters. Mesosternum with shallow punctures,
about 1.5 diameter of those on prosternum, interspaces alutaceous to granular, separated by ~1
diameter. Metasternum irregularly punctate, with faint small punctures on disc similar in size to
those on mentum, interspaces alutaceous to finely granular on metasternal disc becoming
somewhat more granular laterally, punctures separated by ~2-4 diameters. Abdominal sternite 1
219
with large faint punctures, punctures equal to large punctures on pronotum, interspaces
alutaceous to finely microreticulate, separated by 1-2 diameter. Abdominal sternites 2-4 with
two irregular rows of punctures, one row near anterior margin and the other midway to the
posterior margin, punctures similar in size to those on abdominal sternite 1, rows separated by
0.5-1 diameter, punctures within rows separated by ~0.5 punctures width. Rows on abdominal
sternite 4 becoming less organized. Hypopygidium with deeply impressed punctures, similar in
size to those on sternites 2-4, interspaces smooth to granular, punctures separated by 0.25-0.5
diameter.
Head wider than long (W:L = 1.6:1), fronotclypeal region moderately projecting
anteriorly. Vertex with deep concavity between orbits near fronotclypeal region. Labrum with
deep incision at anterior margin. Antennal club compact, oval, symmetrical with the last
antennomere subequal to the previous two combined. Antennomeres 4-8 more or less compact,
with 7-8 characteristically disc-like, 5-6 trapezoidal, and 4 cuboidal. Antennal scape
asymmetrical, somewhat hemispherical, 1.5 times as long as pedicel. Pedicel barrel-shaped.
Antennal segment 3 subequal in length to pedicel. Antennal club large, ~0.70 length of segments
1-8 combined. Antennal grooves very deep and widely excavate, slightly converging
posteriorly. Lateral mental/submental ridge moderately prominent, at level of submentum, ridge
with few longitudinal microreticulations and the corresponding sulcus with granular sculpturing.
Mentum with anterior angles obsolete, anterior margin broadly somewhat angulate, entire
structure obtusely triangular when viewed ventrally and somewhat convex laterally.
Pronotum widest near middle (L:W = 1:1.9), anterior margin broadly trapezoidal,
posterior margin moderately convex, lateral margins less arcuate posteriorly. Scutellum large,
obtusely triangular, apex rounded. Prosternal process somewhat narrowed between procoxae,
apex somewhat acuminate, in lateral aspect the anterior and posterior ends are prominent with a
220
slight convexity medially. Posterior apical wall not prominent and concave. Mesosternum
extending to midway between mesocoxae, evenly concave for reception of the metasternum.
Metasternum wider than long (W:L = 2.6:1.0). Metepisternum with slight medial constriction,
oblique line dividing anterior 0.20 of structure. First abdominal sternite with acuminate process
between metacoxae. First sternite ~2X’s longer than second sternite. Sternites 2-3 subequal in
length, the fourth slightly larger than the preceding two. Hypopygidium subequal in length to
first abdominal sternite.
Protibia with apical tooth prominent, equal in length to tarsomeres 1 and part of 2
combined. Outer apical notch with ~100° angle, notch depth deep, equal to length of tarsomere
1 and part of 2 combined. Inner apical spine subequal in length to tarsomeres 1 and part of 2
combined. Protibia not heavily armored but with characteristic dense patch of stiff setae along
the inner apical region. Mesotibia more heavily armored than protibia with more dense stiff
setae and a row of numerous slender spines along entire lateral edge. Outer apical process
robust, larger than protibia process. Inner apical spine equal in length to tarsomeres 1-2
combined. Metatibia with armature similar to that of mesotibia.
Male genitalia well-sclerotized. Anal sclerite with large broadly curved region
anteriodorsally (Fig. 32); apex densely fimbriate. Spiculum gastrale with wide curved lateral
flanges, medial margins concave, numerous short stiff setae originating from apex (Fig. 71).
Tegmen evenly rounded apically (Fig. 112), longer than wide (w:l = 1.0:2.1), lateral row of setae
visible from the median fossa to prior to the apex, large shallow concavity in apical third with
inner row of setae attaining apex, basal notch perpendicular, basal margin deeply concave.
Median lobe large and robust, ~0.66 the length of the tegmen, apex rounded, apical opening
well-developed with simple internal structure (Fig. 153). Ejaculatory rods not fused to basal
221
piece or each other, oriented at angles to each other. Basal piece U-shaped with inner margin of
arms sinuate (Fig. 192).
Female genitalia moderately sclerotized. Paraprocts moderately large with sclerotization
along median line and approximating baso-lateral angles. Gonocoxite with one basal lateral
prominence, basal ridge well-sclerotized. Gonocoxal apices with recurved “tooth” present. Two
primary setae originate from small depressions on the gonocoxal apices (Fig. 228).
Variation. This species is quite variable in size, with males typically smaller than
females. Color varies with some individuals being darker than others, and some without a
darkened pronotal disc.
Seasonality/Habitat. Specimens are known to occur from mid-July through October,
mostly in low to mid-elevation forests.
Distribution. Known from the type localities in southern Mexico from
Notes. This species is known from the fungal hosts, Lycoperdon compactum and L.
pyriforme.
Etymology. Specific epithet honors Luis Alfredo, the son of Jose Luis Navarette Heredia.
Pocadius majusculus Kirejtshuk
(Figs. 33, 72, 113, 154, 193, 229)
Type Material Examined. (1) Holotype ♂: North Thailand; Doi Sutep; 19-6-1958,
1100m; B. Degerbøl leg.; Pr. 5/0 (1-7-59) / Holotypus Pocadius; majusculus; det. Kirejtshuk
1982. (1) Paratype ♀: Zoologisk Museum, Kobenhaven; Loc. Doi Sutep, N. Thailand; 1100m;
Dat.: 19-6-1968; Leg.: B. Degerbøl; Pr. 510, Journ. 1/759 / Pocadites; sp.; ♀ / Paratypus
Pocadius; majusculus; det. Kirejtshuk 1982.
222
Diagnosis. The large size, sparse pubescence, and relatively small head outwardly
delineate this species from other members of the genus in the Old World. Other characters
delimiting the species include: large umbilicate elytral punctures widely spaced, pronotum
markedly narrowing from base to apex with a shallow anterior emargination, tegmen truncate
apically with setae of similar size extending from margins, pygidial punctation complex with
large irregular punctures bearing no setae and smaller circular punctures bearing short stiff setae
anteriorly to each large puncture, and the gonocoxites with large blunt apices and no recurrent
lateral “teeth”.
Redescription. Length 5.4mm, Width 3.7mm, Depth 1.7mm. Body uniformly
brownish-orange, appendages only slightly lighter, surface somewhat shining, pubescence short
and scattered. Pronotum and elytra with margins fimbriate. Pygidium and hypopygidium
sparingly setate, but posterior margins distinctly fimbriate.
Head surface deeply densely punctate, punctures large becoming smaller and more
densely packed towards orbits, and minute and dense in fronotclypeal region. Only relatively
few punctures giving rise to small fine pale setae. Interspaces alutaceous with some
microreticulation. Smaller punctures about 0.75 the size of larger ones, large punctures about 7-
8 times the size of facets, minute punctures about 0.20 the size of larger punctures. Pronotal
surface with large and small punctures interspersed throughout, all punctures deep. Larger
punctures ~1.5 times diameter of large punctures on vertex, smaller punctures ~0.33 the size of
larger punctures. Interspaces 0.25 - 0.5 diameter apart, smooth to alutaceous with some
microreticulation. Each puncture bearing a short fine pale seta. Scutellar surface scarcely
punctate with small punctures similar in size to the smaller punctures on the pronotum,
microreticulate near apex, interspaces alutaceous. Eytral surface with serial rows of alternating
large and small shallow punctures. Large punctures similar in size to large pronotal punctures,
223
each giving rise to a single short fine semierect seta. Small punctures 0.20 diameter of large
punctures, each giving rise to a short fine seta. Small punctures in either single or irregular
double rows. Interspaces mostly alutaceous between all punctures, larger punctures separated
from each other by 0.25 puncture diameter, smaller punctures separated from each other by 1-2
puncture diameters. Large and small puncture rows separated from each other by ~0.5 large
puncture diameter. Pygidium densely punctate, punctures equal in size to small punctures on
elytra, interspaces alutaceous with some minute microreticulation, punctures separated by ~0.5
puncture diameter. Each puncture giving rise to a short stiff seta, setae equal to length of those
on elytra.
Venter with less pubescence than dorsum. Mentum with sparse minute punctation,
punctures similar in size to small punctures on elytra. Submentum and gula with scattered small
punctures equal in size to those on mentum, interspaces alutaceous with some microreticulation.
Prosternum and epimeron irregularly faintly punctate, punctures approximately equal in size to
larger punctures on vertex, interspaces alutaceous, prosternal punctures separated by 0.25 - 0.5
diameter, those on the epimeron similarly separated. Mesosternum with very faint large
punctures laterally, interspaces alutaceous. Metasternum irregularly punctate, with small faint
punctures on disc similar in size to large punctures on vertex, interspaces smooth to alutaceous
on metasternal disc, punctures separated by ~2-3 diameters. Abdominal sternite 1 with faint
large punctures, only tip of abdominal process lacking punctation. Interspaces alutaceous,
punctures separated by 0.5 - 1 diameter. Abdominal segments 2-4 with punctures aligned in
irregular lateral rows. Punctures equal in size to those on abdominal sternite 1. Interspaces
alutaceous, punctures separated by 0.25 diameter. Hypopygidium densely deeply punctate,
punctures equal to those on other abdominal sternites, interspaces alutaceous with
microreticulation, punctures separated by 0.25 diameter of puncture.
224
Head only slightly wider than long, somewhat triangular, with fronotclypeal region
projecting anteriorly. Vertex with moderate concavity in the frontoclypeal region. Labrum
transverse with indented medial region on anterior margin. Antennal club compact, oblong oval,
densely setose, asymmetrical with the last antennomere longer than previous two combined.
Antennomeres 6-8 strongly flattened into disc-like structures, their combined length less than the
length of antennomere 9. Antennal scape asymmetrical, broadly hemispherical, 2 times as long
as pedicel. Pedicel cylindrical in shape, ~0.5 length of scape. Antennal segment 3
subcylindrical, subequal in length to pedicel. Segments 4-5 subquadrate, narrowing basally, each
0.5 the length of segment 3. Antennal club large, ~0.75 length of segments 1-8 combined,
asymmetrical, terminal segment larger than preceding two segments. Antennal grooves
moderately deep, antennal ledge extending from gula laterally along mentum and transversely
microreticulate. Mentum with apex acutely pointed anteriorly.
Pronotum widest near base (W:L = 1.875:1), anterior margin broadly shallowly concave,
posterior margin convex and somewhat undulate with faint “lobe” over scutellum. Prosternal
process somewhat narrowed between coxae, evenly rounded at apex, in lateral aspect the apex is
somewhat depressed behind coxae. Scutellum large, elongate, with rounded apex. Mesosternum
extending to midway between mesocoxae, posterior border truncate. Mesepisternum larger than
mesosternum. Metasternum width to length ratio is ~2.8:1. Metepisternum somewhat narrow,
only slightly concave medially, anterior third produced anteriolaterally. First abdominal sternite
with broadly rounded almost truncate process between metacoxae with medial acute tip, first
sternite ~2X’s longer than second sternite. Sternites 2-4 subequal in length. Hypopygidium
subequal in length to first abdominal sternite. Humeri moderately produced. Elytral margin
narrow.
225
Protibia distinctly curved laterally, and faintly crenulate along posterior 0.75 of lateral
edge. Apical tooth prominent, subequal to tarsomere 2. Outer apical notch with >90° angle,
notch depth equal to tarsomere 1 and 0.5 of 2 combined. Inner apical spine subequal in length to
first tarsomere. Protibia with very little armature overall. Mesotibia more heavily armored than
protibia with a row of slender spines along entire lateral edge and another row medially to lateral
spines on ventral surface. Outer apical process elongate and robust, larger than protibia process,
projecting more posteriorly than protibial process. Inner apical spine equal in length to
tarsomeres 1 and 0.5 of 2. Metatibia heavily armored with numerous rows of slender elongate
spines. Spines of varying lengths, but mostly longer than those on the mesotibia. Outer apical
process elongate and robust, equal in length to inner apical spine, projecting more posteriorly
than mesotibial processes. Inner apical spine subequal in length to tarsomeres 1-2 combined.
Male genitalia well sclerotized. Anal sclerite with large broadly curved medial concavity
approaching apex (Fig. 33). Spiculum gastrale with moderately wide lateral flanges, medial
margins deeply concave, short stiff setae originating from apex (Fig. 72. Tegmen truncate
apically (Fig. 113), much longer than wide (w:l = 1.0:2.5), lateral row of setae visible from the
median fossa to prior to around apex, large concavity in apical third, basal notch perpendicular,
basal margin nearly straight, apex of median fossa border with extended patch of fine setae.
Median lobe large and robust, ~0.6 the length of the tegmen, apex acuminate, apical opening
well-developed with proximal invagination well-developed (Fig. 154). Ejaculatory rods fused
basally, but not fused to basal piece, straight and elongate, basal piece not observed (Fig. 193).
Female genitalia well-sclerotized. Gonocoxites with large blunt apices and no recurrent
lateral “teeth”. Inner gonocoxal cavity parallel-sided, not acute basally. Gonocoxite base with
ridge of sclerotization, and two medial sclerotized ridges. Paraprocts moderately sclerotized
with inner margin heavily sclerotized to apex (Fig. 229).
226
Variation. None documented, only one Paratype studied.
Seasonality/Habitat. Known from mid-June.
Distribution. Known from the type locality in northern Thailand.
Notes. Pocadius majusculus is the largest species in the genus. No host records have
been recorded for this species.
Pocadius maquipucunensis Leschen and Carlton
(Figs. 34, 73, 114, 155, 194, 230)
Type Material Examined. HOLOTYPE ♂ (SNEC): ECUADOR: Pichiincha;
Maquipucuna For. Res.; 50km NW Quito, 1600m; 21 Dec. 1991, C. Carlton; R. Leschen #34; ex.
Lycoperdon / HOLOTYPE; Pocadius; maquipucunensis; R. Leschen & C. Carlton. 13
PARATYPES: same data label as holotype. 1 PARATYPE: ECUADOR: Pichincha;
Maquipucuna For. Res.; 50km NW Quito, 1720m; 23 Dec.1991, C. Carlton; R. Leschen #64, ex.
FIT.
Non-Type Material Examined. 2 specimens with the following label data. (CMN):
ECU: Pich., 16km E Sanot Domingo, Tinalandia, 4-V-25-VII-1985, S&J Peck, 680m, malaise-
FIT, rainforest.
Diagnosis. The original description of this species suggested that the black coloration
was sufficient to distinguish it from other species of the genus. The author’s stated that the
somewhat similarly colored P. niger is mostly black with red maculae on the elytra. The present
study demonstrates that P. niger has individuals completely black and lack red elytral maculae.
Also, new species described here from the Neotropics have individuals that are completely black
in color. The following characters provide a more reliable diagnosis of the species: metacoxae
broadly separated by rounded almost truncate abdominal process, terminal antennomere much
227
longer than previous two segments combined and with characteristic sensillar region, prosternal
process in lateral view with very slight convexity over procoxal cavity and with short slightly
oblique posterior face, scutellum broader than long with large punctures aggregated in basal half
and smaller punctures in apical half, elytra with extremely narrowly spaced rows of large and
small punctures both within and between rows, metasternal disc with central glabrous apunctate
region, tegmen much longer than wide and with two anterior shallow fossae, median lobe evenly
rounded at apex with complex apical opening, ejaculatory rods with basal piece thickened and
adjacent basally, and ovipositor with broad gonocoxites having more or less converging apical
processes, and three primary setae extending from apical pit (not asetose as remarked upon in the
original description).
Redescription. Length 3.8 mm, Width 2.6mm, Depth 1.4mm. Body moderately
convex, surface shining, dark pitchy brown-black to black in color, venter and appendages
lighter. Pronotum and elytra margins with elongate fimbriae, setae longer than width of antennal
scape. Dorsal and ventral pubescence quite long, but ventral setae somewhat shorter and more
sparsely distributed than dorsal ones.
Head surface deeply, irregularly punctate, punctures larger on vertex, becoming
somewhat smaller and more densely aggregated towards orbits and fronotclypeal region. Larger
punctures 5-6 X diameter of eye facet, smaller punctures 3-4 X diameter. Interspaces smooth to
finely alutaceous, shining. Each puncture gives rise to an elongate curved golden seta. Pronotal
surface with large punctures equal in size to large punctures on vertex of head, interspersed with
relatively few smaller punctures, ~0.85 size of larger ones. Interspaces alutaceous to granular,
about 0.5-1 diameters apart. Each puncture gives rise to a long golden seta, most seta are curved
but some are rather straight. Scutellar surface with shallowly impressed punctures, larger
punctures aggregated in basal half becoming smaller posteriorly, some punctures giving rise to
228
setae, interspaces alutaceous to granular. Elytral surface with serial rows of alternating large and
small deep punctures. Smaller punctures are equal in size to smaller ones on pronotum, larger
punctures are ~1.5 times diameter of smaller ones. Smaller punctures giving rise to a semi-erect
long golden seta, larger punctures giving rise to a semi-erect long golden seta. Interspaces
extremely narrow between punctures of a given row and between different rows. Within a row,
small punctures are separated by ~0.75 puncture width, and large punctures by 0.25-0.5 puncture
diameter. Large puncture rows are separated by 1.0 large puncture diameter. Interspaces shining
but variable from alutaceous to granular in sculpture. Pygidium densely punctate, punctures
equal in size to larger ones on pronotum, each puncture giving rise to a long straight golden seta.
Interspaces narrow, 0.25-0.5 diameter, with smooth to alutaceous sculpture.
Venter with shorter more sparsely distributed golden pubescence as dorsum. Mentum
with small very shallow punctures, equal in size to smaller ones on vertex, each giving rise to a
moderately long seta. Interspaces alutaceous to granular. Submentum and gula with similar
vaguely impressed small punctation as mentum but with interspaces completely granular.
Prosternum and epimeron deeply irregularly punctate, punctures 1.5-1.75X larger than those on
mentum, interspaces granular, prosternal and epimeron punctures separated by 0.5 diameter.
Mesosternum with shallow punctures, similar in diameter to those on prosternum, interspaces
granular, separated by about 0.5 to 1 diameter and mostly aggregated along posterior border.
Metasternum irregularly punctate, with central glabrous apunctate region on disc, those on rest of
metasternum similar in size to those on mesosternum, interspaces alutaceous to granular
becoming more granular laterally, punctures separated by ~1-1.5 diameters. Abdominal sternite
1 with large faint punctures, punctures equal to those on metasternum, interspaces alutaceous to
granular with some microreticulation anteriorly, separated by ~1-2 diameter. Abdominal
sternites 2-4 with an irregular row of punctures near posterior margin as well as other dispersed
229
punctures, punctures somewhat smaller in size to those on metasternum, dispersed punctures
separated by 0.75-1 puncture diameter, punctures within rows separated by ~0.5 punctures
diameter. Hypopygidium with moderately deep punctures, similar in size to those on abdominal
sternite 1, interspaces alutaceous to granular, punctures separated by 0.75-1 puncture diameter.
Head slightly wider than long (W:L = 1.56:1), fronotclypeal region moderately projecting
anteriorly. Vertex with shallow broad wide concavity between orbits near fronotclypeal region.
Labrum with shallow medial incision at anterior margin. Antennal club compact, broadly
obovate, strongly asymmetrical with the last antennomere much greater in length than the
previous two combined. Antennomeres 6-8 more or less compact, and characteristically disc-
like. Antennal scape asymmetrical, somewhat hemispherical, 1.75 times as long as pedicel.
Pedicel subcylindrical in shape. Antennal segment 3 shorter in length than pedicel. Antennal
club large, ~0.73 length of segments 1-8 combined. Each club segment with dense short setae,
and only relatively few protruding setae. Antennal grooves very deep and widely excavate,
slightly converging posteriorly. Lateral and medial mental ridges prominent with very broadly
excavate furrow, at level of submentum, medially the ridge is granularly sculptured with several
setal bearing punctures, and laterally with oblique to longitudinal microreticulations. Mentum
with anterior angles reduced, anterior margin angulate and coming to a distinct medial acuminate
apex, lateral margins very short almost obsolete , basal margin straight, entire structure
pentagonal and moderately convex.
Pronotum widest near middle (L:W = 1:1.83), anterior margin broadly concave and only
slightly trapezoidal, posterior margin moderately convex, lateral margins less arcuate posteriorly.
Scutellum large, obtusely broadly triangular, much wider than long, apex broadly rounded.
Prosternal process somewhat narrowed between procoxae, apex acuminate, in lateral aspect the
anterior and posterior ends are not prominent and there is only a slight medial convexity over the
230
procoxae. Posterior apical wall prominent, but short, and only slightly oblique. Mesosternum
extending to just before midway between mesocoxae, evenly deeply concave for reception of the
metasternum. Metasternum width to length ratio is ~3.2:1.0. Metepisternum broad with slight
medial concavity, oblique line dividing anterior 0.15 of structure. Elytral humeri moderately
produced, lateral margin very narrow. First abdominal sternite with broadly convex to truncate
process between metacoxae. First sternite ~2X’s longer than second sternite. Sternites 2-3
subequal in length, the fourth slightly larger than the preceding two. Hypopygidium subequal in
length to first abdominal sternite.
Protibia with apical tooth very prominent, subequal to tarsomeres 1 and half of 2
combined. Outer apical notch with ~95-100° angle, notch depth deep, equal to length of
tarsomere 1 and part of 2. Inner apical spine subequal in length to tarsomeres 1-2 combined.
Protibia not heavily armored but with characteristic dense patch of stiff setae along the inner
apical region. Mesotibia more heavily armored than protibia with more dense stiff setae and a
row of numerous slender spines along entire lateral edge. Outer apical process elongate but not
robust, larger than protibia process. Inner apical spine equal in length to tarsomeres 1-2
combined. Metatibia with armature similar to that of mesotibia, but with apical tooth and spines
longer.
Male genitalia well-sclerotized. Anal sclerite with large broadly curved region
anteriodorsally (Fig. 34); apex with elongate fimbria; ventrally with a broad medial concavity
approaching apex. Spiculum gastrale with moderately wide lateral flanges, medial margins
broadly concave proximally, numerous moderately long stiff setae originating from apex (fig.
73). Tegmen more acutely rounded apically (Fig. 114), much longer than wide (w:l = 1.0:2.9),
lateral row of setae visible from the median fossa to prior to around the apex, two small shallow
concavities in apical third, inner row of setae incomplete, not attaining apex, basal margin of
231
phallobase angulate. Median lobe large and robust, ~0.66 the length of tegmen, apex broadly
rounded, apical opening well-developed (Fig. 155). Ejaculatory rods not fused to basal piece or
each other, apical rods curved inward medially and expanded outward at basal piece. Basal
piece of rods as two distinct elongate parts, thicker than apical ejaculatory rods and abutting
basally (Fig. 194).
Female genitalia moderately sclerotized. Paraprocts large with sclerotization only along
median line to gonocoxite. Gonocoxite with one basal lateral prominence, basal ridge well-
sclerotized. Gonocoxal apices with recurved “tooth” absent. Usually three but in a few
specimens four primary setae originate from small depressions at the gonocoxal apices (Fig.
230).
Variation. Some specimens are more dark brown than black. Male pygidium with
apical margin truncate.
Seasonality/Habitat. The species is known from mid-elevation tropical rainforests from
May through July and again in December.
Distribution. Known only from Amazonian Ecuador in the Pichincha Province.
Notes. Specimens have been collected from Lycoperdon sp.
Pocadius martini Kirejtshuk
(Fig. 231)
Type Material Examined. (1) Holotype ♀ (ZMUC): PHILIPPINES, Palawan;
Mantalingajan; Pinigisan, 630 meter; 4 Sept. 1961; Noona Dan Exp., 61-62 / caught in malaise
trap / Holotypus Pocadius martini det. Kirejtshuk 1982.
Non-Type Material Examined. None.
232
Diagnosis. Easily distinguished from any other Pocadius due to the extremely short
antennal stem (i.e. antennomeres 1-8), which is subequal in length to the antennal club (i.e.
antennomeres 9-11); and the antennal club is distinctly asymmetrical. Pocadius martini is also
one of the smaller members of the genus at ~3.1mm in length.
Redescription. Length 3.1mm, Width 1.6mm, Depth 0.7mm. Body slightly convex,
surface shining, light tan to reddish brown in color, setae pale, short and fine. Pronotum and
elytra with margins scarcely fimbriate. Pygidium and hypopygidium scarcely setate, but
posterior margins finely fimbriate.
Head surface shallowly, diffusely punctate with only relatively few punctures giving rise
to small fine pale setae. Interspaces granular to alutaceous. Smaller punctures about 0.75-0.5
the size of larger ones, large punctures about 6-7 times the size of facets. Eyes with small fine
facets. Pronotal surface with large and small punctures interspersed throughout. Large
punctures similar in size to respective large punctures on head, smaller punctures about 0.75 size
of large ones. Interspaces ~1 puncture diameter apart, granular with some microreticulation
present on disc. Each puncture bearing a short fine pale seta. Scutellar surface scarcely punctate
with small punctures similar in size to the smaller punctures on the pronotum, interspaces
alutaceous. Eytral surface with serial rows of alternating large and small shallow punctures.
Large punctures ~3 times diameter of the large punctures on head, each giving rise to a single
short fine decumbent seta. Small punctures 0.25 diameter of large punctures, each giving rise to
a short fine decumbent seta. Interspaces faintly granular to smooth between all punctures, larger
punctures separated from each other by <0.5 puncture diameter, smaller punctures separated
from each other by 1.5-2 puncture diameters. Large and small puncture rows separated from
each other by ~0.33 large puncture diameter. Pygidium densely punctate, punctures equal in size
to small punctures on elytra, interspaces granular with some minute microreticulation, punctures
233
separated by ~0.5-0.25 puncture diameter. Each puncture giving rise to a short stiff semi-
decumbent seta, setae ~0.33 length of those on elytra.
Venter similar in pubescence to dorsum, with femora having longest setae. Mentum with
small fine punctures, equal in size to small punctures on head, interspaces distant with granular
to alutaceous texture. Submentum and gula also with scattered small punctures equal in size to
the small punctures on mentum, interspaces granular to alutaceous. Prosternum and epimeron
more deeply and irregularly punctate, punctures approximately 0.75 size larger punctures on
vertex, interspaces mostly alutaceous with some microreticulation, prosternal punctures
separated by 0.25 diameter, those on the epimeron by 0.25 to 0.5 diameter. Mesosternum with
very faint small punctures in a linear series, punctures equal to large punctures on head,
interspaces smooth to alutaceous. Metasternum deeply irregularly punctate, with large punctures
similar in size to those on elytra, disc less densely punctate than lateral regions, interspaces
smooth to alutaceous, punctures separated by ~1-2 diameters. Abdominal sternite 1 with faint
large punctures occurring in all but the anterior 0.33 of the structure which has faint transverse
microreticulate sculpturing. Interspaces alutaceous, punctures separated by 1-2 diameters.
Abdominal segments 2-4 with punctures aligned in irregular transverse rows, punctures equal in
size to those on abdominal sternite 1. Interspaces alutaceous to granular, punctures separated by
~1 diameter. Hypopygidium densely deeply punctate, punctures slightly larger than those on
other abdominal sternites, interspaces smooth to alutaceous with faint microreticulation,
punctures separated by ~0.5 diameter of puncture.
Head much wider than long (W:L = 1.4:1), broadly triangular, with fronotclypeal region
projecting slightly anteriorly. Vertex with broad deep concavity in the fronto-clypeal region.
Eyes large with small fine facets. Labrum transverse with minute medial incision on anterior
margin. Mentum with ventral convexity, giving somewhat convex appearance, structure broadly
234
pentagonal with acute apex. Lateral mental ridge not prominent, at level of submentum, ridge
with longitudinal microreticulation on surface. Antennal club compact, oblong oval,
asymmetrical with the last antennomere much longer than previous two combined, subequal in
length to segments 1-8 combined. Antennal scape asymmetrical, somewhat hemispherical, 2
times as long as pedicel. Pedicel subcylindrical in shape. Antennomere three elongate, longer
than either the pedicel or segment 4. Antennomeres 4-5 subquadrate, subequal to one another.
Antennomeres 6-8 flattened into disc-like structures, their combined length equal to the length of
antennomere 9. Antennal grooves moderately shallow.
Pronotum transverse, widest near posterior angles (L:W = 1:2.8 ), anterior margin
broadly shallowly emarginate, posterior margin mostly truncate with faint “lobe” over scutellum,
lateral margins slightly tapering anteriorly. Scutellum large and broadly triangular. Prosternal
process somewhat narrowed between coxae, acute apex, in lateral aspect the apical 0.33 is
straight and more dorsad behind the procoxae (Fig. ?). Mesosternum extending to 0.66 between
mesocoxae. Mesepisternum wider than mesosternum. Metasternum transverse, W:L ~3.5:1.
Metepisternum moderate, only slightly concave medially, anterior third moderately produced
anteriolaterally. Anterior 0.20 cut-off to form axillary space which lacks punctures. Elytral
humeri slightly produced, lateral margin narrow. First abdominal sternite with broadly rounded
process between metacoxae, first sternite ~2X’s longer than second sternite. Sternites 2-4
subequal in length. Hypopygidium subequal in length to first abdominal sternite. All femora
elongate, widest at middle to distal 0.66.
Protibia with apical tooth prominent, slightly longer than tarsomere 1. Outer apical notch
with ~90° angle, notch depth equal to tarsomere 1 and part of 2 combined. Inner apical spine
subequal in length to tarsomeres 1 and part of 2. Protibia with very little armature overall.
Mesotibia more heavily armored than protibia with a row of slender spines along entire lateral
235
edge and other spines scattered on ventral surface. Outer apical process elongate and robust,
larger than protibia process. Inner apical spine equal in length to tarsomeres 1 and 0.5 of 2.
Metatibia heavily armored with several rows of slender elongate spines. Spines of varying
lengths, but mostly longer than those on the mesotibia. Outer apical process elongate and robust,
equal in length to inner apical spine, projecting more posteriorly than pro- or mesotibial
processes. Inner apical spine subequal in length to tarsomeres 1-2 combined.
Female genitalia overall moderately sclerotized. Gonocoxites with sclerotized basal
border with two lateral prominences, basal border giving rise to two medial oblique sclerotized
ridges extending apicolaterally, ridges short ~1/8 length of basal border. Gonocoxal apices
moderately separated with intragonocoxal invagination evenly rounded at base, gonocoxal tips
evenly rounded basally, each apex with a small lateral depression that gives rise to 4 short seta.
Valvifers membranous with some sclerotization along medial border, evenly tapering to lateral
apex (Fig. 231).
Male genitalia not observed
Variation. None documented, only Holotype studied.
Seasonality/Habitat. Holotype collected in early September.
Distribution. Known from the type locality on the southwest island of Palawan.
Notes. This species does not superficially resemble any of the new species from the
Malaysian archipelago and likely represents an island endemic. More collecting from the other
Philippine islands is needed to further define this fauna, which will likely include other island
endemics as observed in the Greater Antille island chain. No host records are known for this
species.
236
Pocadius monticolis Lechanteur
(Fig. 232)
Type Material Examined. (1) Paratype ♀ (Prague, Czech Rep.): Blukwa; Mont Wago;
7900m; 7-5-56 / Ituri, Zaire / Paratype / F. Lechanteur det., 1958; Pocadius; monticola; n. sp. /
Mus. Nat. Prague; 65627; Inv.
Non-Type Material Examined. None.
Diagnosis. Differs from other Pocadius by unique dark medial line on the pronotum in
combination with orange-brown humeri, a dark antennal club, the protibial outer apical spur
without a prominent notch, heavy armature on all tibia, the elongate sub-cylindrical shape of the
antennal pedicel, and genitalic features such as .
Redescription. Length 4.9mm, Width 2.4mm, Depth 1.6mm. Body dark brown/black
with the following areas orange/brown: lateral 0.33 of pronotum, anterior 0.33 of elytra from
humeri to near suture, venter and appendages except antennal club which is dark. Pronotum and
elytra with margins fimbriate. Pygidium and hypopygidium moderately setate, with posterior
margins densely fimbriate.
Head surface deeply densely punctate, large and small punctures interspersed throughout
vertex. Large punctures ~4-5 times the diameter of an eye facet, smaller punctures about 0.5 size
of larger punctures, interspaces alutaceous and granular, ~ 1 diameter apart. Smaller punctures
giving rise to elongate pale brown setae. Fronotclypeal region with broad shallow concavity on
vertex. Pronotal surface with large and small punctures interspersed throughout, with the
pronotal disc having more abundance of smaller punctures than lateral regions. Elongate pale
brown setae derived from each small puncture. Larger punctures ~1.5 times diameter of large
punctures on vertex, smaller punctures ~0.33 the size of larger punctures. Interspaces 0.5-1
diameter apart, mostly granular texture. Scutellar surface faintly punctate with punctures similar
237
in size to the larger punctures on the vertex, most punctures aggregated in anterior 0.5,
interspaces granular, a few long setae derived from some of the punctures. Eytral surface with
serial rows of alternating large and small shallow punctures. Large punctures somewhat
transverse, ~1.5 times the size of the large pronotal punctures, each giving rise to a single short
stiff semidecumbent seta, ~0.5 the length of the setae derived from the smaller punctures. Small
punctures 0.25 diameter of large punctures, each giving rise to an elongate pale brown seta,
similar in length to those on head and pronotum. Large and small punctures in single rows.
Interspaces mostly alutaceous to granular between all punctures, larger punctures separated from
each other by 0.25 puncture diameter, smaller punctures separated from each other by 1-2
puncture diameters. Large and small puncture rows separated from each other by ~0.75-1 large
puncture diameter. Pygidium densely irregularly punctate, punctures equal in size to small
punctures on elytra, interspaces alutaceous, punctures separated by ~0.25 puncture diameter.
Some but not all punctures giving rise to a short stiff seta, setae equal to length of those from
large punctures on elytra.
Venter with shorter finer pubescence than dorsum. Mentum with faint large punctation,
punctures similar in size to ~0.75 the size of the large vertex punctures, some of the punctures
giving rise to moderately long setae, interspaces granular. Submentum and gula with scattered
small punctures equal to 0.5 size of those on mentum, interspaces alutaceous to granular, with
moderately long setae derived from them. Prosternum and epimeron irregularly faintly punctate,
punctures approximately equal in size to mentum punctures, interspaces alutaceous, prosternal
punctures separated by 0.25 - 0.5 diameter, those on the epimeron similarly separated. Prosternal
punctures giving rise to elongate setae the are slightly curved apically. Mesosternum with very
faint large punctures along posterior margin, interspaces alutaceous. Metasternum irregularly
punctate, with small faint punctures on disc similar in size to small elytral punctures, interspaces
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smooth to alutaceous on metasternal disc, punctures separated by ~2-3 diameters, each punctures
giving rise to a straight moderately long seta. Abdominal sternite 1 with faint medium sized
punctures on the abdominal process that are about 1.5 times larger than those on metasternum,
interspaces wide, ~3 diameters apart, mostly alutaceous with transverse microreticulation visible;
large pictures posteriorly and laterally on sternite 1, equal in size to those on elytra. Abdominal
segments 2-4 with large punctures aligned in irregular lateral rows. Punctures similar in size to
larger punctures on abdominal sternite 1. Interspaces alutaceous, punctures separated by 0.25
diameter. Hypopygidium densely deeply punctate, punctures equal to those on abdominal
sternites 2-4, interspaces alutaceous with some microreticulation, punctures separated by 0.25-
0.5 diameter of puncture. Head only slightly wider than long, somewhat triangular, with
fronotclypeal region projecting anteriorly. No prominence over antennal insertion. Labrum
transverse with broad indented medial region on anterior margin. Antennal club compact,
elongate oval, densely setose, asymmetrical, with the last antennomere longer than previous two
combined. Antennomeres 6-8 strongly flattened into disc-like structures, their combined length
less than the length of antennomere 9. Antennal scape somewhat asymmetrical, subcylindrical, 2
times as long as pedicel, numerous elongate setae arising from anterior margin of scape. Pedicel
cylindrical in shape, ~0.5 length and width of scape, widest near middle. Antennal segment 3
subcylindrical, about 0.75 length of pedicel. Segments 4-5 subquadrate, narrowing basally, each
~0.5 the length of segment 3. Antennal club large, subequal in length of segments 1-8 combined,
asymmetrical, terminal segment larger than preceding two segments. Antennal grooves
moderately deep and somewhat medially curved posteriorly, antennal ledge extending from
submentum laterally along mentum with alutaceous and microreticulate surface. Mentum with
apex evenly rounded anteriorly, hemispherical, somewhat convex.
239
Pronotum transverse, widest near posterior angles (W:L = ~2:1), anterior margin broadly
deeply concave, posterior margin broadly convex. Prosternal process somewhat narrowed
between coxae, acute apex, in lateral aspect the anterior portion is depressed before coxal
cavities, with a perpendicular apical wall. Scutellum large, broadly triangular with rounded
apex. Mesosternum not extending to midway between mesocoxae, posterior border deeply
concave. Mesepisternum equal in width to mesosternum. Mesepimeron ~0.5 the width of
mesepisternum. Metasternum width to length ratio is ~2.3:1. Metepisternum somewhat broad,
only slightly concave medially, anterior third produced anteriomedially, anterior 1/6 cut-off from
rest of metepisternum by faint oblique incomplete carina. First abdominal sternite with broadly
triangular process between metacoxae with medial acute tip, first sternite ~1.75’s longer than
second sternite. Sternites 2-4 subequal in length. Hypopygidium ~1.2 times the length of first
abdominal sternite. Humeri slightly produced with faint to obsolete narrow elytral margin.
Protibia with slight lateral curvature, faintly crenulate along lateral edge. Apical tooth
somewhat prominent, subequal to tarsomere 2. Outer apical notch indistinct, notch depth equal
to 0.5 length of tarsomere 1. Inner apical spine subequal in length to first tarsomere and 0.5 of
second. Protibia with moderate armature overall, a row of short stiff spines along inner edge.
Mesotibia more heavily armored than protibia with several rows of slender spines along lateral
edge and medial and ventral surfaces. Outer apical process somewhat robust, subequal to
protibial process, projecting slightly more posteriorly than protibial process. Inner apical spine
equal in length to tarsomere 1. Metatibia heavily armored with numerous rows of slender
elongate spines. Spines of varying lengths, but mostly longer than those on the mesotibia. Outer
apical process robust, projecting more posteriorly than mesotibial processes. Inner apical spine
subequal in length to tarsomeres 1 and 0.5 of the second combined.
Male genitalia not observed.
240
Female genitalia moderately sclerotized (Fig. 232). Paraprocts large with sclerotization
along median margin. Gonocoxite with one basal lateral prominence, basal ridge moderately
well-sclerotized. Gonocoxal apices with recurved “tooth” present. Two primary setae originate
from small depressions on the gonocoxal apices (Fig. ?). Intragonocoxal invagination deep and
narrow.
Variation. None documented, only 1 Paratype studied.
Seasonality/Habitat. Known to the author only from the single paratype female that was
collected in May.
Distribution. Known only from the type locality.
Notes. No host information is known for this species.
Pocadius niger Parsons
(Figs. 35, 74, 115, 156, 195, 233)
Type Material Examined. HOLOTYPE (USNM): Las Vegas HS; 5-8, N.M. / Barber
&; Schwarz Coll. / Holotype No; 54656. 4 PARATYPES (USNM): same data labels as
holotype, except last label reads “Paratype No; 54656”.
Non-Type Material Examined. 22 specimens (CAS, LSAM, ACC) with the following
label data: Peterson Ranch; [illegible handwriting]; Sierra Ancha Mts. / Gila Co., AZ; VIII-20-
1947 / puffball / L.R. Gillogly collector. ARIZ. Rustler Park; Chiricahua Mts., Cochise Co.,
Aug. 11, 1970, 8500ft.
Diagnosis. This species can be distinguished from the other New World fauna by the
following characters: body coloration black often with orange brown humeri; prosternal process
evenly rounded over procoxae with posterior face straight to slightly concave; terminal
antennomere shorter than segments 9-10 combined and evenly rounded apically, apico-lateral
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protibial notch deep and distinct; eyes smaller and not greatly protruding; elytral fimbriae
markedly elongate and sparsely distributed; median lobe of aedeagus oval in shape with evenly
rounded apex; basal structure of ejaculatory rods with medial convexity along basal margin;
ovipositor gonocoxites with one lateral prominence and with a W:L ~1:1.
Redescription. Length 3.6 mm, Width 1.9mm, Depth 1.3mm. Body moderately
convex, surface somewhat shining, dark brown/black in color often with elytral humeri and basal
portion of elytra lighter. Pronotum and elytra margins with elongate sparsely distributed
fimbriae, setae much longer than width of antennal scape. Dorsal and ventral pubescence quite
long.
Head surface deeply, irregularly punctate, large and smaller punctures evenly distributed
across vertex. Larger punctures ~5 X diameter of eye facet, smaller punctures 4 X diameter,
interspaces finely alutaceous and somewhat shining. Each smaller puncture gives rise to an
elongate curved golden seta. Pronotal surface with large punctures equal in size to large
punctures on vertex of head, interspersed with numerous smaller punctures, ~0.75 size of larger
ones, interspaces alutaceous with some microreticulation, about 0.5 diameters apart. Each
puncture gives rise to a long golden seta. Scutellar surface with few vague shallowly impressed
punctures similar in size to smaller ones on pronotum, some punctures giving rise to setae, and
the interspaces are alutaceous to somewhat granular. Elytral surface with serial rows of
alternating large and small moderately deep punctures. Smaller punctures are equal in size to
smaller ones on pronotum, larger punctures are ~1.5-2 times diameter of smaller ones. Smaller
punctures giving rise to an erect long golden seta, larger punctures giving rise to a semi-erect
long golden seta. Interspaces narrow between punctures of a given row and between different
rows. Within a row, small punctures are separated by ~1 puncture width, and large punctures by
0.3-0.5 puncture width. Large puncture rows are separated by 2 large puncture diameters.
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Interspaces moderately shining but variable from smooth to somewhat rugose in sculpture.
Pygidium densely punctate, punctures equal in size to smaller ones on pronotum, each puncture
giving rise to a moderately long golden seta. Interspaces narrow, 0.25-0.5 diameters, with
alutaceous to granular sculpture.
Venter with similarly long golden pubescence as dorsum. Mentum with few small very
shallow punctures, equal in size to smaller ones on vertex, each giving rise to an short stiff seta.
Interspaces alutaceous to finely microreticulate. Submentum and gula similar in punctation to
mentum but with interspaces having more microreticulation present. Prosternum and epimeron
deeply irregularly punctate, punctures similar in size to larger ones on vertex, interspaces
granular with conspicuous microreticulate areas, prosternal punctures separated by 0.25-0.5
diameter, those on the epimeron similarly spaced. Mesosternum with shallow punctures, similar
in size to those on prosternum, interspaces alutaceous to rugose, separated by about 0.5 to 1
diameter, and mostly aggregated near metasternal border. Metasternum deeply irregularly
punctate, with punctures on disc slightly larger in size to those on mesosternum, interspaces
alutaceous to rugose on metasternal disc becoming microreticulate to granular laterally,
punctures separated by ~0.75-1 diameter. Abdominal sternite 1 with some large and smaller
punctures intermixed, larger punctures equal to those on metasternum and smaller punctures
equal to those on mentum, interspaces alutaceous to granular, separated by ~1 diameter.
Abdominal sternites 2-4 with two irregular rows of punctures near the anterior and posterior
margin and other dispersed punctures, punctures similar in size to smaller ones on abdominal
sternite 1, punctures within a row separated by ~1 puncture width, other punctures more widely
spaced. Hypopygidium with moderately deep punctures, similar in size to those on sternites 2-4,
interspaces alutaceous to granular, punctures separated by 0.5-1 puncture width.
243
Head slightly wider than long (W:L = 1.5:1), fronotclypeal region moderately projecting
anteriorly. Vertex with broad deep concavity between orbits near fronotclypeal region. Labrum
with shallow but broad incision at anterior margin. Antennal club compact, elongate to oval,
symmetrical with the last antennomere shorter than the previous two segments combined.
Antennomeres 6-8 more or less compact, with 7-8 characteristically disc-like. Antennal scape
asymmetrical, faintly hemispherical, 2 times as long as pedicel. Pedicel short, subcylindrical in
shape. Antennal segment 3 equal in length to pedicel. Antennal club moderately large, ~0.68
length of segments 1-8 combined. Antennal grooves very deep and widely excavate, slightly
converging posteriorly. Lateral mental/submental ridge prominent, at level of submentum, ridge
sculptured with small minute punctures medially and laterally with oblique to longitudinal
microreticulations. Mentum with anterior angles distinct, anterior margin angular coming to
more or less distinct apex, entire structure pentagonal and flattened when viewed laterally.
Pronotum widest near middle (L:W = 1:1.73), anterior margin broadly trapezoidal,
posterior margin moderately convex, lateral margins less arcuate posteriorly, anterior and
posterior angles distinct. Scutellum large, elongate triangular, apex narrowly rounded.
Prosternal process somewhat narrowed between procoxae, apex somewhat acuminate, in lateral
aspect the anterior and posterior ends are prominent and evenly convex medially. Posterior
apical wall prominent and perpendicular with a slight concavity. Mesosternum extending to
midway between mesocoxae, evenly concave for reception of the metasternum. Metasternum
width to length ratio is ~2.51:1.0. Metepisternum with slight medial constriction, oblique line
dividing anterior 0.20 of structure. Elytral humeri moderately produced, lateral margin very
narrow. First abdominal sternite with narrow acuminate process between metacoxae. First
sternite ~2X’s longer than second sternite. Sternites 2-3 subequal in length. Hypopygidium
subequal in length to first abdominal sternite.
244
Protibia with apical tooth very prominent, slightly longer than tarsomeres 1-2 combined.
Outer apical notch with ~95° angle, notch depth deep, equal to length to tarsomere 1 and part of
2 combined. Inner apical spine subequal in length to tarsomere 1 and part of 2 combined.
Protibia heavily armored with characteristic dense patch of stiff setae along the inner apical
region and numerous elongate setae along lateral margin. Mesotibia more heavily armored than
protibia with more rows of dense stiff setae and a row of numerous slender spines along entire
lateral edge. Outer apical process elongate and robust, larger than protibia process. Inner apical
spine equal in length to tarsomeres 1-2 combined. Metatibia with more than that of mesotibia,
numerous spines adorning the lateral and medial borders, but with outer apical process similar to
that of mesotibia.
Male genitalia well-sclerotized. Anal sclerite with large broadly curved region
anteriodorsally (Fig. 35); apex somewhat fimbriate; ventrally with a broad medial concavity
approaching apex in a truncate manner. Spiculum gastrale with wide lateral flanges having small
concavity near apical-lateral region, medial margins concave, extremely short stiff setae
originating from apex (Fig. 74). Tegmen evenly rounded apically (Fig. 115), longer than wide
(w:l = 1.0:1.77) becoming somewhat narrowed basally, lateral row of setae visible from apex of
the median fossa to prior to the tegmen apex, circular/oval shallow concavity in apical third,
basal notch perpendicular, basal margin slightly concave, inner row of setae not attaining apex.
Median lobe oval and robust, ~0.50 the length of the tegmen, apex evenly rounded, apical
opening well-developed with deep proximal concavity (Fig. 156). Ejaculatory rods not fused to
basal piece, slightly curved inward medially and expanded outward at basal piece. Basal piece
of rods with biconcavity along apical margin, proximally with two sharp projections and two
rounded projections (Fig. 195).
245
Female genitalia moderately sclerotized. Paraprocts large and widely flared with
sclerotization along median and basal margins. Gonocoxite with one basal lateral prominence,
basal ridge moderately well-sclerotized. Gonocoxal apices with recurved “tooth” present. Two
primary setae originate from small depressions on the gonocoxal apices (Fig. 233).
Variation. Some specimens have the reddish areas on the elytral humeri lacking, giving
rise to an all dark brown/black color pattern.
Seasonality/Habitat. Specimens have only been collected in the month of August in all
localities in Arizona and New Mexico. This species occurs at some of the highest altitudes in the
genus, with specimens collected from above 8000’.
Distribution. This species is known from southern New Mexico through parts of
southern and south-central Arizona.
Notes. Host records for this species only include the generic term “puffball” in the label
data.
Pocadius nigerrimus Cline new species
(Figs. 36, 75, 116, 157, 196)
Type Material Examined. HOLOTYPE ♂ (SNEC): PARAGUAY: Itapúa; Yataí, prop.
Hostettler family,; San Rafael Reserve, 100m; 26°38’17”S, 55°38’50”W; 21-25 NOV 2000, Z.H.
Falin; PAR1F00 040; ex. flight intercept trap / SM0257887; KUNHM-ENT [“bar-code” label] /
HOLOTYPE; Pocadius; nigerrimus; A.R. Cline des. 2004.
Diagnosis. This species can be delimited from the other Neotropical fauna by the
following suite of characters: uniformly dark reddish brown/black coloration; thickened
tarsomeres; elongate pronotal and elytral fimbriae; elytral punctures with little dissimilarity in
the diameter of the large and small serial rows of punctures; elytral setae erect and semi-erect;
246
widely globular antennal club with transversely expanded terminal antennomere; male pygidium
deeply indentate along apical border; tegmen with incomplete sinuately organized inner row of
setae; median lobe globular but with narrowed apex; ejaculatory rods elongate with expanded
proximal and distal regions; basal piece of internal sac sclerites with two separate lateral pieces
and a central elongate piece that overlaps the two lateral arms.
Description. Length 4.5mm, Width 2.7mm, Depth 1.8mm. Body moderately convex,
surface shining, uniformly dark reddish-brown/black. Pronotum and elytra margins with
elongate fimbriae, setae longer than width of antennal scape. Dorsal and ventral pubescence
somewhat sparsely distributed but quite long.
Head surface deeply, irregularly punctate, punctures larger on vertex, becoming
somewhat smaller towards orbits and fronotclypeal region. Larger punctures 5-6X diameter of
eye facet, smaller punctures 1-2X diameter. Interspaces smooth to finely granular, shining.
Each puncture gives rise to an elongate curved golden seta. Pronotal surface with large
punctures subequal in size to large punctures on vertex of head, interspersed with relatively few
smaller punctures, ~0.5 size of larger ones. Interspaces alutaceous to finely microreticulate,
about 0.5-1 diameter apart. Each puncture gives rise to a long curved golden seta. Scutellar
surface with many shallowly impressed punctures, some punctures giving rise to setae, and the
interspaces are alutaceous to finely microreticulate. Elytral surface with serial rows of
alternating large and small deep punctures. Smaller punctures are equal in size to those on
pronotum, larger punctures are ~2-3 times diameter of smaller ones. Smaller punctures giving
rise to an erect long golden seta, larger punctures giving rise to a semi-erect somewhat shorter
golden seta. Interspaces broad between punctures of a given row and between different rows.
Within a row, small punctures are separated by ~2-3 puncture diameters, and large punctures by
1 puncture diameter. Larger rows are separated by ~2 large puncture diameters. Interspaces
247
always shining but variable from smooth to finely microreticulate in sculpture. Pygidium
densely punctate, punctures equal in size to larger ones on pronotum, each puncture giving rise
to a moderately long golden seta. Interspaces narrow, 0.25-0.75 diameter, with granular to
microreticulate sculpture.
Venter with somewhat shorter and sparser golden pubescence as dorsum. Mentum with
large small shallow punctures, equal in size to smaller ones on vertex, each giving rise to a short
seta. Interspaces smooth with areas of fine microreticulations. Submentum and gula similar in
punctation to mentum. Prosternum and epimeron shallowly irregularly punctate, epimeron
punctures equal to large punctures on vertex and those on prosternum equal to smaller punctures
on vertex, interspaces alutaceous with microreticulate areas, prosternal punctures separated by
~1-2 diameter, those on the epimeron by 0.25 to 0.5 diameter. Mesosternum with shallow
punctures, equal in diameter of those on epimeron, interspaces alutaceous to granular, separated
by about 0.5 to 1 diameter, and aggregated near metasternum. Metasternum irregularly punctate,
with minute faint punctures on disc similar in size to smaller ones on vertex, interspaces finely
alutaceous to granular on metasternal disc becoming somewhat smoother laterally, disc
punctures separated by ~1-2 diameters. Abdominal sternite 1 with large faint punctures,
punctures equal to large punctures on pronotum, interspaces alutaceous with microreticulations
present, separated by ~1-2 diameters. Abdominal sternites 2-4 with two irregular rows of
punctures, one row near anterior margin and the other near posterior margin, punctures similar in
size to those on abdominal sternite 1, rows separated by ~2-3 puncture diameters, punctures
within rows separated by ~1-2 puncture diameter. Rows on abdominal sternite 4 becoming less
organized. Hypopygidium with moderately deep punctures, similar in size to those on sternites
2-4, interspaces alutaceous to granular, punctures separated by 0.5-1 diameter.
248
Head wider than long (W:L = 1.5:1), fronotclypeal region moderately projecting
anteriorly. Vertex with shallow concavity between orbits near fronotclypeal region. Labrum
with deep broad medial incision at anterior margin. Antennal club compact, globular,
asymmetrical with the last antennomere longer than the previous two combined. Antennomeres
4-8 more or less compact, with 6-8 characteristically disc-like, and 4-5 cuboidal. Antennal scape
asymmetrical, somewhat hemispherical, 1.6 times as long as pedicel. Pedicel subcylindrical in
shape. Antennal segment 3 longer in length to pedicel. Antennal club moderately large, ~0.65
length of segments 1-8 combined. Antennal grooves very deep and widely excavate, slightly
converging posteriorly. Lateral mental/submental ridge prominent, at level of submentum, ridge
with well-developed longitudinal microreticulations throughout and the corresponding sulcus
completely granular. Mentum with anterior angles obsolete, anterior margin broadly
hemispherical, entire structure widely hemispherical when viewed ventrally and somewhat
convex when viewed laterally.
Pronotum widest near middle (L:W = 1:1.7), anterior margin deeply broadly
trapezoidal, posterior margin moderately convex, lateral margins less arcuate posteriorly.
Scutellum large, obtusely triangular, apex rounded. Prosternal process somewhat narrowed
between procoxae, apex somewhat acuminate, in lateral aspect the anterior and posterior ends are
prominent with a slight convexity medially. Posterior apical wall not prominent and with a
medial concavity. Mesosternum extending to midway between mesocoxae, evenly concave for
reception of the metasternum. Metasternum width to length ratio is ~3.0:1.0. Metepisternum
with slight medial constriction, oblique line dividing anterior 0.15 of structure. First abdominal
sternite with broad truncate process between metacoxae. First sternite ~2X’s longer than second
sternite. Sternites 2-3 subequal in length, the fourth slightly larger than the preceding two.
Hypopygidium subequal in length to first abdominal sternite.
249
Protibia with apical tooth not prominent, subequal to tarsomere 1. Outer apical notch
with ~115° angle, notch depth shallow, equal to length of tarsomere 1. Inner apical spine
subequal in length to tarsomeres 1 and part of 2 combined. Protibia moderately heavily armored
with characteristic dense patch of stiff setae along the inner apical region and numerous elongate
setae along the lateral margin. Mesotibia more heavily armored than protibia with more dense
stiff setae and a row of numerous slender spines along entire lateral edge. Outer apical process
elongate and robust, larger than protibia process. Inner apical spine equal in length to tarsomeres
1-2 combined. Metatibia with heavier armature than that of mesotibia, and inner apical spine
equal to tarsomeres 1-2 and part of 3 combined.
Male genitalia well-sclerotized. Anal sclerite with large broadly curved region
anteriodorsally (Fig. 36); apex somewhat fimbriate; ventrally with a broad medial concavity
approaching apex in a truncate manner. Spiculum gastrale with wide lateral flanges, medial
margins convex proximally, numerous short stiff setae originating from apex (Fig. 75). Tegmen
evenly rounded apically (Fig. 116), much longer than wide (w:l = 1.0:2.55), lateral row of setae
visible from the median fossa to prior to the apex, large elliptical concavity in apical third, basal
notch perpendicular, basal margin nearly straight. Median lobe large and robust, ~0.75 the
length of the tegmen, apex narrowing, apical opening well-developed with large internal
structure (Fig. 157). Ejaculatory rods not fused to basal piece or each other, straight, and
expanded outward basally and apically. Basal piece of internal sac sclerites with two separate
lateral pieces and a large elongate central piece that overlaps the two lateral ones (Fig. 196).
Female genitalia not observed.
Variation. Known from the holotype male.
Seasonality/Habitat. The holotype was collected in late November in a lowland tropical
forest.
250
Distribution. Known from the type locality in southeastern Paraguay.
Notes. No host information is available for this species.
Etymology. Specific epithet is derived from the Latin “niger” meaning “black” or “dark”,
and “imus” meaning “ a likeness”, denoting the dark almost black habitus.
Pocadius nobilis Reitter
(Figs. 37, 76, 117, 158, 197, 234)
Type Material Examined. HOLOTYPE (BMNH), two female specimens on same
paper card with the following data labels: Type; H.T. [circular label with red trim] / Japan.; G.
Lewis.; 1910-320 / Pocadius; nobilis m. HOLOTYPE (RNH) ♂: Japan; leg. Lewis; Coll. Reitter
/ nobilis; Japan, Rtts. [upside down yellow label] / Holotypus 1873; Pocadius; nobilis; Reitter
[rectangular label with red trim] /P. nobilis; m.; Japan [upside down label].
Non-Type Material Examined. 10 specimens (BMNH): 3 with same data labels as
holotype but without type labels, 1 with same data label as holotype but with additional label of
“Higo.”, 1 with same data label as holotype but with additional label of “Hitoyoshi.; 3.V. –
8.V.81.”, 1 with the following labels: Japan, (BMNH) 1 specimen: in puffball / JAPAN:; J.E.A.
Lewis.; B.M. 1933-490, 1 with the following labels: JAPAN; Kobe; Shinohara; 30-IX-’30;
J.E.A. Lewis / JAPAN:; J.E.A. Lewis.; B.M. 1933-490. 1 specimen (RMNH ) with the
following labels: KOREA, No. 693; Kangwon Prov.; Mts. Kumgang-san; singled, 16-IX-1980;
leg. Topál and Forró. 4 specimens (ZMHB): no specimens with specific label data, only ID
labels.
Diagnosis. The peculiar color pattern, particularly the dark medial longitudinal “stripe”
on the pronotum serves to distinguish P. nobilis from the other Oriental and Old World
Pocadius. The deep punctation of the pygidium and non-trapezoidal anterior pronotal margin
251
serves to differentiate this species from the Palearctic species P. adustus and P. ferrugineus.
These features in conjunction with densely setose tegmen, sub-parallel sided median lobe with
angular apical margins, and basally fused ejaculatory rods and deeply cleft and separated basal
piece of ejaculatory rods clearly delimit this species.
Redescription. Length 3.6mm, Width 2.0mm, Depth 0.9 mm. Body convex, shining,
light reddish brown with the following regions being much darker brown to almost black:
antennal club, temples of head, pronotal disc from apex to base, scutellum, lateral region of
elytra from humeri to apices as well as apical 0.5 of elytra from suture to lateral explanation
leaving only the medial basal region lighter in color. Pubescence of modest length, golden,
sparsely covering dorsum. Pronotum more broadly explanate than elytra, sides fimbriate.
Head approximately equal in length and width, broadly triangular, with clypeolabral
region projecting anteriorly. Head surface irregularly punctate, both large and small punctures
interspersed throughout vertex, becoming somewhat more densely packed near orbits, becoming
more obsolete near clypeus, interspaces smooth to alutaceous. Smaller punctures ~3 times larger
than eye facets, larger punctures between 3-5 times diameter of smaller punctures. Large
punctures microreticulate. Each puncture giving rise to a single apically curved seta. Labrum
impunctate, smooth to alutaceous in texture. Labrum transverse with small medial incision on
anterior margin. Vertex with deep somewhat narrow concavity extending between antennal
insertions from the medial region of the frons to the clypeal region. Clypeal region convex,
distinctly bulging dorsally. Antennal club compact, broadly oval, asymmetrical with the last
antennomere subequal to previous two combined. Antennomere 8 strongly flattened into disc-
like structure, 6 and 7 also flattened and disc-like but not to the extent of 8, their combined
length subequal to the length of antennomere 9. Antennal scape asymmetrical, broadly convex
dorsally, 2 times as long as pedicel. Pedicel subcylindrical in shape. Antennal segment 3
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subequal in length to pedicel, gradually expanded apically. Segments 4 and 5 subquadrate, 0.5
the length of segment 3. Antennal grooves deep and somewhat curved posteriomedially.
Mentum with anterior margin broadly evenly convex, convex medially, surface with minute faint
punctures sparsely distributes, interspaces smooth with some microreticulation laterally.
Submentum and gula, elongate, parallel-sided, surface with faint punctation, punctures equal in
size to the large punctures on the vertex, interspaces alutaceous with microreticulation. Lateral
mental-gular ledge well-developed, evenly rounded along outer edge,0.5 as wide as gula, widest
anterior to middle, surface with distinct oblique microreticulation.
Pronotum widest near middle, evenly rounded laterally, anterior and posterior apices
somewhat rounded, anterior margin broadly shallowly concave, posterior margin with deep
concavity near posterior apices becoming broadly convex near middle. Pronotal surface with
large and small punctures interspersed throughout. Large and small punctures similar in size to
respective large and small punctures on head. Interspaces 1 to 2 puncture diameters apart,
smooth to alutaceous. Prosternum and epimeron irregularly punctate, punctures equal in size to
large punctures on head, interspaces alutaceous. Prosternal process somewhat narrowed between
coxae, acutely rounded at apex, basal 0.33 elevated but not carinate. Prosternal process surface
with indistinct punctation, surface completely alutaceous and granular, with apical margin finely
granular. Scutellum large, broadly rounded at apex. Scutellar surface sparsely punctate with
minute punctures ~0.5 the diameter of the small punctures on the head, interspaces mostly
smooth. Elytra L:W = 1.1:1, narrowly margined, humeri moderately produced, apices separately
rounded with a complete subapical line present. Eytral surface with serial rows of alternating
large and small shallow punctures. Large punctures ~2 times diameter of the large punctures on
head, each giving rise to a decumbent seta. Small punctures 0.20 diameter of large punctures,
each giving rise to a decumbent seta. Interspaces mostly alutaceous between all punctures, larger
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punctures separated from each other within a row by 0.5 puncture diameter, smaller punctures
separated from each other by 2-3 puncture diameters. Large and small puncture rows separated
from each other by ~2 small puncture diameters. Mesosternum extending to midway between
mesocoxae with a faint medial carina, deeply concave apical margin, lateral margin broadly
shallowly concave. Mesosternal punctures aggregated toward posterior margin. Anterior
surface completely granular with faint microreticulation. Mesepisternum only slightly larger
than mesepimeron. Metasternum width to length ratio is 2.5:1. Metasternal punctures faint,
equal in size to large punctures on head, interspaces on disc smooth to alutaceous, becoming
completely alutaceous laterally. Metepisternum rather broad, width to length 1:3.8, surface with
faint large punctation similar in size to those on metasternum, interspaces alutaceous with some
microreticulation. First abdominal sternite with broad almost truncate process ending in a small
point between metacoxae, ~2.5X’s longer than second sternite. Sternites 2-4 subequal in length.
Abdominal sternite 1 with faint umbilicate punctures occurring in all but the anterior 0.33 of the
structure which is alutaceous. Abdominal segments 2-4 with punctures aligned in irregular
lateral rows. Punctures equal in size to those on abdominal sternite 1, but not umbilicate.
Interspaces alutaceous, punctures separated by 0.5 diameter. Pygidium broadly triangular with
sparsely fimbriate margins. Pygidial surface densely punctate, punctures equal in size to large
punctures on elytra, interspaces microreticulate, punctures separated by 0.25-0.20 puncture
diameter. Each puncture giving rise to a short seta, ~0.25 length of those on elytra.
Hypopygidium smaller in length than first abdominal sternite. Hypopygidium densely punctate,
punctures equal to those on other abdominal sternites, interspaces almost rugulose, punctures
separated by 0.25 diameter of puncture.
Protibia with apical tooth approximately 1.5 times length of second tarsomere. Outer
apical notch indistinct. Inner apical spine subequal in length to 1.5 length of first tarsomere.
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Apical border of tibia with one large tooth. Mesotibia modestly armored with one row of slender
spines along entire lateral edge and other rows of stiff hairs on the ventral and dorsal surfaces.
Apical border armored with short spines, 0.33 length of lateral spines. Outer apical tooth short
and robust, 0.5 length of inner apical spine. Inner apical spine equal in length to tarsomeres 1-2.
Metatibia similarly armored as mesotibia. Apical border armored with short spines, as in
mesotibia, but more numerous. Outer apical process robust, 0.5 length of inner apical spine.
Inner apical spine equal in length to tarsomeres 1-2.
Male genitalia moderately sclerotized. Anal sclerite with large broadly curved region
anteriodorsally (Fig. 37), apex moderately fimbriate; ventrally with a deep medial concavity
approaching apex in a truncate manner. Spiculum gastrale with (Fig. 76) lateral region broadly
rounded and widely explanate, medial border evenly concave, apical border with numerous
elongate setae. Tegmen broadly rounded to truncate apically (Fig. 117), longer than wide (w:l ~
1.3.1), lateral row of setae visible from the median fossa to prior to the apex, inner row of setae
incomplete apically extending from apex of median fossa to just prior to apex, basal notch
perpendicular when viewed laterally, basal margin slightly concave. Median lobe large and
robust, sides subparallel laterally becoming angulate apically, slightly greater than 0.6 the length
of the tegmen, apical opening well-developed (Fig. 158). Ejaculatory rods fused to each other
basally but not fused to basal piece, basal piece deeply cleft apically and becoming narrowed
proximally (Fig. 197).
Female genitalia overall moderately sclerotized. Gonocoxites with well sclerotized basal
border and two lateral prominences, basal border giving rise to two medial oblique sclerotized
baculi extending apicolaterally, ridges long ~0.33 length of basal border. Gonocoxal apices
narrowly separated with intragonocoxal invagination evenly rounded at base, gonocoxal tips
evenly rounded at apex, each apex with a small lateral depression that gives rise to 4-5 primary
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seta (Fig. 234). Paraprocts membranous with some sclerotization along medial border, evenly
tapering to lateral apex.
Variation. Male specimens tend to have a broader apico-lateral notch on the protibia.
Seasonality/Habitat. Specimens are known to occur from May through September.
Distribution. This species may be found on the major islands of Japan. The Korean
specimen is a new distribution record for the species and extends the range into mainland Asia.
Notes. The only host information given is that of “puffball.”
Pocadius okinawaensis Cline new species
(Figs. 38, 77, 118, 198, 235)
Type Material Examined. HOLOTYPE ♂ (FMNH): Ryukyu Is.: OKINAWA;
Katsudake, IX-28-1945; leg. E. Ray / eating hole; in Lycoperdon; fungus / Pocadius; nobilis Rtt;
Kirejtshuk det. 1994 / HOLOTYPE; Pocadius; okinawaensis; A.R. Cline des. 2004. 12
PARATYPES (FMNH): same data labels as holotype, but with PARATYPE designation labels.
Diagnosis. This species is most similar to P. nobilis known from Japan, but can be
differentiated from it and the other Palearctic fauna by the following suite of characters: terminal
antennomere slightly longer than preceding two combined, irregularly shaped, and with distinct
centrally located sensillar region; elytra with dark markings laterally and apically; head and
pronotum with surface smooth and shining; metasternal disc with minute dispersed punctures
and smooth shining surface; elytra with alternating rows of semi-erect and decumbent setae;
outer apical notch of protibia indistinct; tegmen with a double row of inner setae not attaining the
apex; median lobe with elongate deeply cleft internal structure; ejaculatory rods diverging
apically and parallel basally; basal piece of internal sac sclerite complex with two deeply cleft
lateral arms; ovipositor with short intragonocoxal invagination and 3-4 primary setae.
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Description. Length 3.75mm, Width 2.55mm, Depth 1.5mm. Body moderately convex,
surface shining, reddish-brown to dark reddish brown in color, with lateral and apical portion of
elytra darker. Pronotum and elytra margins with moderately long fimbriae, setae longer equal to
length of antennal scape. Dorsal and ventral pubescence moderately long.
Head surface irregularly punctate with moderately impressed small and large punctures,
most larger ones on vertex, becoming somewhat smaller towards orbits and fronotclypeal region.
Larger punctures 3-4X diameter of eye facet, smaller punctures 1-2X diameter. Interspaces
smooth, shining. Each puncture gives rise to a curved golden seta. Pronotal surface with large
punctures equal in size to large punctures on vertex of head, interspersed with relatively few
smaller punctures, ~0.5 size of larger ones. Interspaces smooth and shining, about 1-2 diameters
apart. Each puncture gives rise to a curved golden seta. Scutellar surface with very few vague
shallowly impressed small punctures, some punctures giving rise to setae, interspaces smooth to
finely alutaceous. Elytral surface with serial rows of alternating large and small deep punctures.
Smaller punctures are equal in size to smaller ones on pronotum, larger punctures are ~2-3 times
diameter of smaller ones. Smaller punctures giving rise to a semi-erect long golden seta, larger
punctures giving rise to a decumbent long golden seta. Interspaces broad between punctures of a
given row and between different rows. Within a row, small punctures are separated by ~2-3
puncture diameter, and large punctures by 0.5-1 puncture diameter. Larger rows are separated
by 2-3 puncture diameters. Interspaces always shining but variable from smooth to finely
alutaceous in sculpture. Pygidium densely punctate, punctures equal in size to larger ones on
pronotum, each puncture giving rise to a short golden seta; interspaces narrow, 0.25-0.5
diameter, with smooth to finely alutaceous sculpture.
Venter with shorter sparser pubescence as dorsum. Mentum with large very shallow
punctures, equal in size to larger ones on vertex, each giving rise to a short seta; interspaces
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alutaceous with finely microreticulate areas. Submentum and gula similar in punctation to
mentum but with interspaces more granular. Prosternum and epimeron shallowly irregularly
punctate, punctures on epimeron slightly larger than those on mentum and those on prosternum
equal to smaller punctures on vertex, interspaces alutaceous to granular with microreticulate
areas, prosternal punctures separated by 1-2 diameter, those on the epimeron by 0.5-1 diameter.
Mesosternum with moderately impressed punctures, equal to those on epimeron, interspaces
alutaceous to granular, separated by about ~1 diameter and aggregated near metasternal border.
Metasternum irregularly punctate, mostly impunctate on disc but with some moderately faint
large and small punctures on disc similar in size to those on vertex, interspaces smooth on
metasternal disc becoming alutaceous to finely granular laterally, punctures separated by ~1-3
diameters. Abdominal sternite 1 with large faint punctures, punctures equal to large punctures
on elytra, interspaces smooth to alutaceous, separated by ~0.5-1 diameter. Abdominal sternites
2-4 with irregular rows of punctures, punctures similar in size to those on sternite 1.
Hypopygidium with moderately deep punctures, similar in size to those on sternites 2-4,
interspaces alutaceous to granular, punctures separated by ~0.5 diameter.
Head wider than long (W:L = 1.5:1), fronotclypeal region moderately projecting
anteriorly. Vertex with shallow concavity between orbits near fronotclypeal region. Labrum
with shallow concavity at anterior margin. Antennal club compact, obovate, slightly
asymmetrical with the last antennomere slightly longer than the previous two combined.
Antennomeres 4-8 more or less compact, with 6-8 characteristically disc-like, and 4-5
trapezoidal. Antennal scape asymmetrical, somewhat hemispherical, 2X as long as pedicel.
Pedicel subcylindrical in shape. Antennal segment 3 subequal in length to pedicel. Antennal
club moderately large, ~0.65 length of segments 1-8 combined. Antennal grooves very deep and
widely excavate, slightly converging posteriorly. Lateral mental/submental ridge prominent with
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a corresponding widely excavate sulcus, ridge at level of submentum, ridge with faint
longitudinal microreticulations and sulcus with alutaceous to granular sculpture. Mentum with
anterior angles obsolete, anterior margin broadly convex, entire structure hemispherical in
ventral view and somewhat convex laterally.
Pronotum widest near posterior angles (L:W = 1:1.9), anterior margin shallowly
concave, posterior margin moderately convex, lateral margins slightly arcuate anteriorly.
Scutellum large, obtusely triangular, apex rounded. Prosternal process somewhat narrowed
between procoxae, apex somewhat acuminate, in lateral aspect the anterior and posterior ends are
not prominent and there is a slight convexity medially, appearing almost flat. Posterior apical
wall moderately prominent and oblique. Mesosternum extending to midway between
mesocoxae, evenly concave for reception of the metasternum. Metasternum wider than long
(W:L = 2.9:1.0). Metepisternum with slight medial constriction, oblique line dividing anterior
0.20 of structure. First abdominal sternite with acuminate process between metacoxae. First
sternite ~2X’s longer than second sternite. Sternites 2-3 subequal in length, the fourth slightly
larger than the preceding two. Hypopygidium subequal in length to first abdominal sternite.
Protibia with apical tooth not prominent, equal to 0.5 length of tarsomere 1. Outer apical
notch absent. Inner apical spine subequal in length to tarsomeres 1 and part of 2 combined.
Protibia not heavily armored but with characteristic dense patch of stiff setae along the inner
apical region. Mesotibia somewhat more heavily armored than protibia with some dense stiff
setae and a row of numerous slender spines along entire lateral edge. Outer apical process not
robust, subequal to protibia process. Inner apical spine equal in length to tarsomere 1 and part of
2 combined. Metatibia with armature similar to that of mesotibia.
Male genitalia well-sclerotized. Anal sclerite with large broadly curved region
anteriodorsally (Fig. 38); apex fimbriate; ventrally with a broad medial concavity approaching
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apex in a truncate manner. Spiculum gastrale with wide lateral flanges, medial margins deeply
concave proximally, few short stiff setae originating from apex (Fig. 77). Tegmen evenly
rounded apically (Fig. 118), much longer than wide (w:l = 1.0:2.67), lateral row of setae visible
from the median fossa to prior to the apex, large elliptical shallow concavity in apical third with
double row of inner setae not attaining apex, basal notch perpendicular, basal margin slightly
concave. Median lobe large and robust, ~0.66 the length of the tegmen, apex narrowed, apical
opening well-developed with deeply cleft bilobed internal structure (Fig. 159). Ejaculatory rods
not fused to basal piece or each other, parallel basally and divergent from one another apically.
Basal piece of internal sac sclerite with two lateral deeply cleft extensions (Fig. 198).
Female genitalia moderately sclerotized. Paraprocts large with sclerotization only along
median line. Gonocoxite with two basal lateral prominences, basal ridge well-sclerotized.
Gonocoxal apices with reduced recurved “tooth” present. Three to four primary setae originate
from terminal pits on the gonocoxal apices. Intragonocoxal invagination shallow (Fig. 235).
Variation. None observed.
Seasonality/Habitat. All specimens were collected in late September.
Distribution. Known only from the type locality.
Notes. Specimens were collected from a Lycoperdon puffball.
Etymology. Specific epithet is a derivative of the type locality, i.e. Okinawa.
Pocadius pecki Cline new species
Figs. (39, 78, 119, 160, 199)
Type Material Examined. HOLOTYPE ♂ (CMN): VEN: Miranda: 400m; 35km N
Altagracia; Guatopo NP, Aqua Blanca; 31-V-7-VI87-2; S&J Peck, ravine FIT / HOLOTYPE:
Pocadius; pecki; A.R. Cline des. 2004. PARATYPE (CMN): VENEZUELA: Tachira;
260
Pregonera, Presa Las; Cuevas, 650m, 9-31-VII-; 1989, S&J Peck, rain-; forest, ex: FIT, 89-255 /
PARATYPE: Pocadius; pecki; A.R. Cline des. 2004.
Diagnosis. This species can be delimited from the rest of the Neotropical fauna by the
following suite of characters: mentum obtusely triangular with curved lateral margins; pronotum
widest near middle with anterior margin feebly concave; prosternal process with apical wall
concave in lateral aspect; elytra with alternating rows of erect and semi-erect setae; metasternum
faintly punctate with minute punctures, nearly glabrous; terminal antennomere only slightly large
than preceding two segments combined, not drastically asymmetrical, and with large sensillar
region; male pygidium with indentate posterior margin; aedeagus with tegmen having a complete
outer row and incomplete inner row of setae, ejaculatory rods with sharply excised distal regions,
basal piece of internal sac sclerites with paired lateral arms each having two sharp projections.
Description. Length 3.50mm, Width 2.27mm, Depth 1.5mm. Body moderately convex,
surface shining, uniformly light golden brown in color. Pronotum and elytra margins with
elongate fimbriae, setae longer than width of antennal scape. Dorsal and ventral pubescence
moderately long.
Head surface irregularly punctate with moderately impressed small and large punctures,
most larger ones on vertex, becoming somewhat smaller towards orbits and fronotclypeal region.
Larger punctures 3-4X diameter of eye facet, smaller punctures 1-2X diameter; interspaces
smooth to finely alutaceous, shining. Each puncture gives rise to a curved golden seta. Pronotal
surface with large punctures equal in size to large punctures on vertex of head, interspersed with
numerous smaller punctures, equal to smaller punctures on vertex; interspaces smooth to finely
alutaceous and shining, ~0.5-1 diameter apart. Each puncture gives rise to a curved golden seta.
Scutellar surface with numerous shallowly impressed small punctures, some punctures giving
rise to setae, interspaces smooth to finely microreticulate apically. Elytral surface with serial
261
rows of alternating large and small deep punctures, the first few rows of smaller punctures are
confusedly arranged and may appear to have more than a single row of punctures. Smaller
punctures are equal in size to smaller ones on pronotum, larger punctures are ~3 times diameter
of smaller ones. Smaller punctures giving rise to an erect long golden seta, larger punctures
giving rise to a semi-erect moderately long golden seta. Interspaces broad between punctures of
a given row and between different rows. Within a row, small punctures are separated by ~2-3
puncture diameters, and large punctures by 1 puncture diameter. Larger rows are separated by 2-
3 puncture diameters. Interspaces always shining but variable from smooth to finely alutaceous
in sculpture. Pygidium densely punctate, punctures equal in size to larger ones on pronotum,
each puncture giving rise to a moderately long golden seta; interspaces narrow, 0.25-0.5
diameter, with alutaceous to finely microreticulate sculpture.
Venter with somewhat shorter sparser pubescence as dorsum. Mentum with minute
shallowly impressed punctures, equal in size to smaller ones on vertex, each giving rise to a short
seta; interspaces alutaceous with finely microreticulate areas. Submentum and gula similar in
punctation to mentum but with interspaces more granular. Prosternum and epimeron irregularly
punctate, punctures on epimeron equal to larger ones on pronotum and those on prosternum
equal to smaller ones on pronotum, interspaces alutaceous to microreticulate, prosternal
punctures separated by 1-2 diameter, those on the epimeron by 0.5 diameter. Mesosternum with
shallowly impressed punctures, equal to those on epimeron, interspaces alutaceous to granular,
separated by about ~1 diameter and aggregated near metasternal border. Metasternum
irregularly punctate, mostly impunctate on disc but with some moderately faint minute broadly
dispersed punctures on disc similar in size to smaller ones on vertex, interspaces alutaceous on
metasternal disc becoming more granular laterally, punctures separated by ~3-5 diameters on
disc. Abdominal sternite 1 with moderate sized faint punctures, punctures equal to large ones on
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vertex, interspaces alutaceous, separated by ~1-2 diameters. Abdominal sternites 2-4 with 2
irregular rows of punctures, one near anterior margin and the other near the posterior margin
with some intervening punctures, punctures similar in size to those on sternite 1. Hypopygidium
with moderately deep punctures, 1.5 diameter of those on sternites 2-4, interspaces alutaceous to
granular, punctures separated by ~0.5-1 diameter.
Head slightly wider than long (W:L = 1.4:1), fronotclypeal region moderately projecting
anteriorly. Vertex with shallow concavity between orbits near fronotclypeal region. Labrum
with minute medial incision at anterior margin. Antennal club compact, obovate, slightly
asymmetrical with the last antennomere slightly longer than the previous two combined.
Antennomeres 4-8 more or less compact, with 6-8 characteristically disc-like, 5 trapezoidal, and
4 cuboidal. Antennal scape asymmetrical, somewhat hemispherical, 2X as long as pedicel.
Pedicel subcylindrical in shape. Antennal segment 3 equal in length to pedicel. Antennal club
not large, ~0.5 length of segments 1-8 combined. Antennal grooves very deep and widely
excavate, slightly converging posteriorly. Lateral mental/submental ridge prominent with a
corresponding widely and deeply excavate sulcus, ridge at level of submentum, ridge with
numerous distinct longitudinal microreticulations and sulcus with alutaceous to granular
sculpture and some longitudinal microreticulations. Mentum with anterior angles obsolete,
anterior margin broadly curved with acute apex, entire structure obtusely triangular in ventral
view and somewhat convex laterally.
Pronotum widest in posterior third (L:W = 1:1.9), anterior margin concave, posterior
margin moderately convex, lateral margins more arcuate anteriorly. Scutellum large, obtusely
triangular, apex rounded. Prosternal process somewhat narrowed between procoxae, apex
somewhat acuminate, in lateral aspect the anterior and posterior ends are prominent and there is a
distinct convexity medially. Posterior apical wall moderately prominent and concave.
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Mesosternum extending to midway between mesocoxae, evenly concave for reception of the
metasternum. Metasternum much wider than long (W:L = 3.1:1.0). Metepisternum with slight
medial constriction, oblique line dividing anterior 0.18 of structure. First abdominal sternite with
acuminate process between metacoxae. First sternite ~2X’s longer than second sternite.
Sternites 2-3 subequal in length, the fourth slightly larger than the preceding two.
Hypopygidium subequal in length to first abdominal sternite.
Protibia with apical tooth prominent, equal to length of tarsomere 1. Outer apical notch
reduced, equal to 0.5 length of tarsomere 1. Inner apical spine subequal in length to tarsomere 1.
Protibia not heavily armored but with characteristic dense patch of stiff setae along the inner
apical region. Mesotibia much more heavily armored than protibia with numerous dense stiff
setae and a row of numerous slender spines along entire lateral edge. Outer apical process
robust, larger than protibial process. Inner apical spine equal in length to tarsomere 1-2 and part
of 3 combined. Metatibia with armature similar to that of mesotibia.
Male genitalia well-sclerotized. Anal sclerite with large broadly curved region
anteriodorsally (Fig. 39); apex fimbriate; ventrally with a broad medial concavity approaching
apex in a convex manner. Spiculum gastrale with wide lateral flanges, medial margins concave,
relatively few long stiff setae originating from apex (Fig. 78). Tegmen narrowly rounded
apically (Fig. 119), longer than wide (w:l = 1:2.6), lateral row of setae visible from the median
fossa to prior to the apex, large elliptical shallow concavity in apical third with single row of
inner setae not attaining apex, basal notch perpendicular, basal margin convex. Median lobe
large and robust, ~0.66 the length of the tegmen, apex narrowed, apical opening well-developed
with shallowly cleft bilobed internal structure (Fig. 160). Ejaculatory rods not fused to basal
piece or each other, slightly curved outward apically and basally with the apical ends sharply
264
excised. Basal piece of internal sac sclerite with paired curved structures having two lateral
sharp projections (Fig. 199).
Female genitalia not observed.
Variation. No demonstrable variation.
Seasonality/Habitat. Specimens known from May to July in lowland rainforest.
Distribution. Known from the type localities in Venezuela.
Notes. No host information available for this species.
Etymology. Specific epithet honors Stewart Peck, Professor of Biology at Carleton
University, whose devotion to beetle collecting is unsurpassed.
Pocadius peruensis Cline new species
(Figs. 40, 79, 120, 161, 200, 236)
Type Material Examined. HOLOTYPE ♂ (SNEC) PERU: Tambopata Prov.; Madre de
Dios Dpto.; 15km NE Puerto Maldonaldo Reserva; Cuzco Amazonica / 12° 33’S 69° 03’W;
220m, Plot# Z1U16; 22 June 1989, J.S. Ashe, R.A. Leschen #207; ex. flight intercept trap. 1
PARATYPE (SNEC): PERU: Tambopata Prov.; 15km NE Pto. Maldonaldo; 22 June 1989,
200m; J. Ashe, R. Leschen, #210; ex: Geastrum. 2 PARATYPES (SNEC): PERU: Tambopata
Prov.; 15km NE Pto. Maldonaldo; 17 June 1989, 200m; J. Ashe, R. Leschen, #210; ex:
Lycoperdonales. 1 PARATYPE (SNEC): PERU: Tambopata Prov.; Madre de Dios Dpto.; 15km
NE Puerto / Maldonaldo Reserva; Cuzco Amazonica; 12°33’S 69°03’W; 200m, Plot# Z2E15 /
15 June 1989; R.A. Leschen #061; ex. Flight intercept trap. 1 PARATYPE (FSCA): PERU:
Loreto Pr., nr.; jct. Rio Maranon &; Ocayali, 73.5°W 4.8°S; 6-20-VIII-1994; P. Skelley, day
catch.
265
Diagnosis. This species can be delimited from the rest of the Neotropical fauna by the
following suite of characters: ;anal sclerite with broad flanges pointed apically; tegmen with
small depression apically, inner row of setae not attaining apex, and a distinctly concave
posterior margin when viewed laterally; median lobe with medial constriction; basal piece of
internal sac sclerites U-shaped with small piece separating the two arms of the U; ovipositor with
paraprocts acute apically, and gonocoxal extensions abutting midway down the intragonocoxal
invagination.
Description. Length 4.1mm, Width 2.7mm, Depth 2.1mm. Body moderately convex,
surface shining, light golden to reddish-brown, sometimes with elytral apices and lateral margin
darker. Pronotum and elytra margins with elongate fimbriae, setae longer than length of antennal
scape. Dorsal and ventral pubescence quite long.
Head surface irregularly punctate with deeply impressed small and large punctures, most
larger ones on vertex, becoming somewhat smaller towards orbits and fronotclypeal region.
Larger punctures ~4X diameter of eye facet, smaller punctures 1-2X diameter. Interspaces
alutaceous to finely granular, moderately shining. Each puncture gives rise to a curved golden
seta. Pronotal surface with large punctures equal in size to large punctures on vertex of head,
interspersed with numerous smaller punctures, ~0.5 size of larger ones; interspaces alutaceous to
finely granular and moderately shining, ~0.25-0.5 diameter apart. Each puncture gives rise to a
curved golden seta. Scutellar surface with numerous vague shallowly impressed minute
punctures equal to smaller ones on vertex, some punctures giving rise to setae, interspaces
alutaceous with some microreticulation apically. Elytral surface with serial rows of alternating
large and small deep punctures. Smaller punctures are equal in size to smaller ones on
pronotum, larger punctures are ~2 times diameter of smaller ones. Smaller punctures giving rise
to an erect long golden seta, larger punctures giving rise to a semi-erect long golden seta.
266
Interspaces narrow between punctures of a given row and between different rows. Within a row,
small punctures are separated by ~1-2 puncture diameter, and large punctures by 0.25-0.5
puncture diameter. Larger rows are separated by 1-1.5 larger puncture diameters. Interspaces
somewhat shining but variable from alutaceous to finely granular in sculpture. Pygidium densely
punctate, punctures equal in size to larger ones on pronotum, each puncture giving rise to a short
golden seta; interspaces narrow, ~0.5-1 diameter, with finely granular sculpture.
Venter with shorter sparser pubescence as dorsum. Mentum with minute shallow
punctures, equal in size smaller ones on vertex, each giving rise to a short seta; interspaces finely
granular. Submentum and gula similar in punctation to mentum but with interspaces more
granular with some microreticulation present. Prosternum and epimeron irregularly punctate
with moderately impressed punctures, punctures on epimeron equal to larger ones on pronotum
and those on prosternum equal 0.75 diameter of those on epimeron, interspaces granular,
prosternal punctures separated by 1 diameter, those on the epimeron by ~0.5 diameter.
Mesosternum with few moderately impressed punctures, equal to those on epimeron, interspaces
granular, separated by about ~1 diameter and aggregated near metasternal border. Metasternum
irregularly punctate, disc punctures similar in size to smaller ones on vertex, interspaces
alutaceous to finely granular on metasternal disc becoming more granular laterally, punctures
separated by ~1-2 diameters. Abdominal sternite 1 with large faint punctures, punctures equal to
large punctures on pronotum, interspaces granular, separated by ~1-2 diameters. Abdominal
sternites 2-4 with two irregular but distinct rows of punctures, one near anterior margin and the
other near the posterior margin, punctures similar in size to those on sternite 1. Hypopygidium
with moderately deep punctures, similar in size to those on sternites 2-4, interspaces granular,
punctures separated by ~0.25 diameter.
267
Head wider than long (W:L = 1.75:1), fronotclypeal region moderately projecting
anteriorly. Vertex with shallow concavity between orbits near fronotclypeal region. Labrum
with deep medial incision at anterior margin. Antennal club compact, obovate, slightly
asymmetrical with the last antennomere longer than the previous two combined. Antennomeres
4-8 more or less compact, with 6-8 characteristically disc-like, and 4-5 trapezoidal. Antennal
scape asymmetrical, somewhat hemispherical, 1.8X as long as pedicel. Pedicel subcylindrical in
shape. Antennal segment 3 subequal in length to pedicel. Antennal club moderately large, ~0.65
length of segments 1-8 combined. Antennal grooves very deep and widely excavate, slightly
converging posteriorly. Lateral mental/submental ridge prominent with a corresponding widely
excavate sulcus, ridge at level of submentum, ridge with distinct oblique and longitudinal
microreticulations and sulcus with transverse to oblique microreticulations. Mentum with
anterior angles distinct, anterior margin angulate, entire structure pentagonal in ventral view and
somewhat convex laterally.
Pronotum widest near indistinct posterior angles (L:W = 1:1.9), anterior margin
shallowly concave, posterior margin moderately convex, lateral margins slightly arcuate
anteriorly. Scutellum large, obtusely triangular, apex rounded. Prosternal process somewhat
narrowed between procoxae, apex somewhat acuminate, in lateral aspect the anterior end is more
prominent than the posterior end and there is a moderate convexity medially. Posterior apical
wall prominent and slightly oblique. Mesosternum extending to midway between mesocoxae,
deeply concave for reception of the metasternum. Metasternum much wider than long (W:L =
3.2:1.0). Metepisternum with slight medial constriction, oblique line dividing anterior 0.13 of
structure. First abdominal sternite with acuminate process between metacoxae. First sternite
~2X’s longer than second sternite. Sternites 2-3 subequal in length, the fourth slightly larger
than the preceding two. Hypopygidium subequal in length to first abdominal sternite.
268
Protibia with apical tooth prominent, equal to length of tarsomere 1. Outer apical notch
present, depth equal to length of tarsomere 1, notch angle ~75°. Inner apical spine subequal in
length to tarsomeres 1 and part of 2 combined. Protibia not heavily armored but with
characteristic dense patch of stiff setae along the inner apical region. Mesotibia somewhat more
heavily armored than protibia with dense stiff setae and a row of numerous slender spines along
entire lateral edge. Outer apical process robust, larger than protibial process. Inner apical spine
equal in length to tarsomere 1-2 and part of 3 combined. Metatibia with armature similar to that
of mesotibia, but with more numerous slender spines.
Male genitalia well-sclerotized. Anal sclerite with large broadly curved region
anteriodorsally (Fig. 40); apex densely fimbriate; ventrally with a narrow medial concavity
approaching apex in a curved manner. Spiculum gastrale with wide lateral flanges that are
pointed apically, medial margins perpendicular, 5-7 stiff setae originating from apex (Fig. 79).
Tegmen narrowly rounded apically (Fig. 120), much longer than wide (w:l = 1.0:3.1), lateral
row of setae visible from the median fossa to around the apex, small circular shallow concavity
in apical third with single row of inner setae not attaining apex, basal notch perpendicular, basal
margin distinctly concave. Median lobe large and robust with medial constriction, ~0.66 the
length of the tegmen, apex narrowly rounded, apical opening well-developed with simple
internal structure (Fig. 161). Ejaculatory rods not fused to basal piece or each other, slightly
converging basally and divergent from one another apically. Basal piece of internal sac sclerite
U-shaped with small piece present between arms of U (Fig. 200).
Female genitalia moderately sclerotized. Paraprocts somewhat narrowed, acute apically,
with sclerotization only along median line to base. Gonocoxite with one basal lateral
prominence, basal ridge well-sclerotized. Gonocoxal apices without recurved “tooth”. Two or
269
three primary setae originate from small depressions on the gonocoxal apices. Gonocoxal
extensions approximate midway down the intragonocoxal invagination (Fig. 236).
Variation. Three of the paratypes have the elytral sides more markedly darker than the
holotype specimen.
Seasonality/Habitat. Known from lowland tropical forests in June and August.
Distribution. Known from the type localities in Peru.
Notes. This species is known from the fungi Lycoperdonales and Geastrum.
Etymology. Specific epithet is a derivative of the type locality, i.e. Peru.
Pocadius rubidus Erichson
(Figs. 41, 80, 121, 162, 201, 237)
Type Material Examined. HOLOTYPE ♂ (Humboldt): 8643 / Holotype; Pocadius;
rubidus; ERICHSON, 1843 / rubidus; Er.; [illegible handwriting on last line].
Non-Type Material Examined. 12 specimens (USNM): 2 with the following label data:
ROSAS - F.C. SUD; Provincia de Buenos Aires; Juan B. Daguerre / ARGNETINA; 1968 colln.;
J. Daguerre. 3 with the following label data: BsAsTanolil; P. Köelerls / ARGNETINA; 1968
colln.; J. Daguerre. 5 with the following label data: BsAs; Zelolya [sp?]; X-45, J. Daguerre /
ARGNETINA; 1968 colln.; J. Daguerre. 3 with the following label data: Cordoba; Alta Gracia;
III-45, J. Daguerre / ARGNETINA; 1968 colln.; J. Daguerre.
Diagnosis. One of the larger South American species, P. rubidus can be distinguished
from the other New World Pocadius by the following suite of characters: elytral surface
distinctly rugose to microreticulate and densely punctate; pronotum with posterior angles
indistinct and broadly rounded; antennal club with terminal antennomeres not asymmetrical or
longer than previous two segments combined; body large and robust; dorsum densely pubescent
270
with elongate golden setae; metasternal disc glabrous with few widely distributed minute
punctures; outer protibial tooth large and robust; anal sclerite with characteristically curved setae
along apical border;
Redescription. Length 4.0 mm, Width 2.4mm, Depth 1.9mm. Body convex, surface
shining, reddish-brown to dark brown in color, sometimes with apical third of elytra dark brown
to piceous. Pronotum and elytra margins with elongate fimbriae, setae much longer than length
of antennal scape. Dorsal and ventral pubescence quite long and conspicuous.
Head surface deeply, irregularly punctate, large and smaller punctures densely aggregated
throughout head. Larger punctures 4-5 X diameter of eye facet, smaller punctures 3 X diameter.
Interspaces smooth to finely alutaceous, shining. Most smaller punctures gives rise to an
elongate curved golden seta. Eyes finely faceted. Pronotal surface with large punctures equal in
size to large punctures on vertex of head, interspersed with numerous smaller punctures, ~0.33-
0.50 size of larger ones. Interspaces smooth to alutaceous, about 0.33-0.5 diameters apart. Each
smaller puncture gives rise to a long golden seta, most seta are moderately curved. Scutellar
surface with very numerous shallowly impressed punctures, somewhat larger than smaller
punctures on pronotum, most punctures giving rise to setae, with the interspaces are alutaceous
to granular. Elytral surface with serial rows of alternating large and small deep punctures, rows
of smaller punctures confusedly dispersed such that in some places there appears to be 2 rows of
smaller punctures between the larger ones. Smaller punctures are equal in size to smaller ones
on pronotum, larger punctures are ~2 times diameter of smaller ones. Smaller punctures giving
rise to an erect long golden seta, larger punctures giving rise to a semi-erect long golden seta.
Interspaces narrow between punctures of a given row and between different rows. Within a row,
small punctures are separated by ~0.75-1 puncture width, and large punctures by 0.1-0.25
puncture diameters. Larger rows are separated by 1.0 large puncture width. Interspaces
271
moderately shining but variable from rugose/granular to microreticulate in sculpture, the
sculpturing is characteristically deep and distinct. Pygidium densely punctate, punctures equal in
size to smaller ones on elytra, each puncture giving rise to a shorter golden seta. Interspaces
narrow, 0.25-0.33 diameter, with granular sculpture.
Venter with similar long golden pubescence as dorsum. Mentum with moderately large
shallow punctures, equal in size to larger ones on vertex, each giving rise to an elongate seta.
Interspaces alutaceous to granular. Submentum and gula similar in punctation to mentum but
with interspaces completely granular with some microreticulation present. Prosternum and
epimeron more deeply irregularly punctate than mentum, punctures slightly larger than those on
mentum, interspaces granular with microreticulate areas, prosternal punctures separated by ~0.5
diameter, those on the epimeron by 0.25 to 0.5 diameter. Mesosternum with more deeply
impressed punctures, slightly larger than those on prosternum, interspaces completely granular,
separated by about 0.5 diameter and mostly aggregated along metasternal border. Metasternum
irregularly punctate, with minute faint punctures on disc similar in size to smaller ones on venter,
interspaces alutaceous to granular on metasternal disc becoming granular to microreticulate
laterally, punctures separated by >2 diameters around disc and becoming more dense laterally.
Abdominal sternite 1 with large faint, almost obsolete punctures, punctures equal to those on
prosternum, interspaces alutaceous to granular, separated by ~0.75 - 1.5 diameter. Abdominal
sternites 2-43 with three irregular rows of punctures, one row near anterior margin, one midway
between anterior and posterior border, and the other near posterior margin, punctures similar in
size to those on abdominal segment 1, rows separated by 1-1.5 puncture diameters, punctures
within rows separated by ~0.25-0.5 punctures width. Rows on abdominal sternite 4 becoming
much less organized, in particular the middle row. Hypopygidium with moderately deep
272
punctures, similar in size to those on sternites 2-4, interspaces mostly granular, punctures
separated by 0.5 puncture diameters.
Head wider than long (L:W = 1:1.45), fronotclypeal region moderately projecting
anteriorly. Vertex with shallow broad indistinct concavity between orbits near fronotclypeal
region. Eyes large, moderately protruding. Antennal club compact, elongate oval, mostly
symmetrical with the last antennomere subequal in length to the previous two combined.
Antennomeres 4-5 more or less compact, with 6-8 characteristically disc-like. Antennal scape
asymmetrical, shortened and somewhat hemispherical, 1.3 times as long as pedicel. Pedicel
subcylindrical in shape. Antennal segment 3 shorter in length to pedicel. Antennal club
moderately large, ~0.55 length of segments 1-8 combined. Each club segment with dense short
setae, and only relatively few protruding setae. Antennal grooves very deep and widely
excavate, slightly converging posteriorly. Lateral mental ridge prominent, at level of
submentum, medially the ridge is rugose sculptured with a few small punctures and laterally with
oblique to longitudinal microreticulations. Mentum with anterior angles obsolete, anterior
margin angulate, overall shape is transversely triangular, entire structure when viewed laterally is
flattened.
Pronotum widest near in posterior 0.33, transverse (L:W = 1:2), anterior margin broadly
concave, posterior margin moderately convex, lateral margins somewhat arcuate anteriorly,
anterior angles distinct, posterior angles indistinct and broadly rounded. Scutellum large,
obtusely triangular, apex rounded. Prosternal process somewhat narrowed between procoxae,
apex somewhat acuminate, in lateral aspect the anterior and posterior ends are not prominent,
convex medially with a steeper declivity in anterior 0.66. Posterior apical wall not prominent,
oblique with little concavity. Mesosternum extending to midway between mesocoxae, evenly
concave for reception of the metasternum. Metasternum width to length ratio is ~3.1:1.0.
273
Metepisternum with slight medial constriction, oblique line dividing anterior 0.20 of structure.
Elytral humeri moderately produced, lateral margin very narrow. First abdominal sternite with
acuminate process between metacoxae. First sternite ~2X’s longer than second sternite.
Sternites 2-4 subequal in length. Hypopygidium subequal in length to first abdominal sternite.
Protibia with apical tooth somewhat prominent, slightly longer than tarsomeres 1. Outer
apical notch indistinct. Inner apical spine subequal in length to tarsomeres 1 and part of 2
combined. Protibia not heavily armored but with characteristic dense patch of stiff setae along
the inner apical region. Mesotibia somewhat more heavily armored than protibia with a few
more dense stiff setae and a row of numerous short slender spines along entire lateral edge.
Outer apical process elongate and robust, larger than protibia process. Inner apical spine equal in
length to tarsomeres 1 and part of 2 combined. Metatibia with heavy armature, lateral row of
spines 2-2.5 X longer than those on mesotibia.
Male genitalia well-sclerotized. Anal sclerite with large broadly curved region
anteriodorsally (Fig. 41); apex fimbriate with characteristically curved setae; ventrally with a
broad medial concavity approaching apex in a convex manner. Spiculum gastrale with wide
tapering lateral flanges, medial margin evenly concave, long stiff setae originating from apex
(Fig. 80). Tegmen evenly rounded apically (Fig. 121), much longer than wide (w:l = 1.0:2.44),
lateral row of setae visible from the median fossa to prior to the apex, small shallow elliptical
concavity in apical third, basal notch perpendicular, basal margin concave, inner row of setae
almost attaining apex. Median lobe large and robust, ~0.66 the length of the tegmen, apex
broadly rounded, lateral sides with slight constriction basally, apical opening well-developed
with proximal concavity (Fig. 162). Ejaculatory rods not fused to each other or to basal piece,
curved inward and expanded outward basally and apically. Basal piece of rods with lateral
inward projecting arms (Fig. 201).
274
Female genitalia moderately sclerotized. Paraprocts large with sclerotization only along
median line to baso-lateral angles. Gonocoxite with two basal lateral prominences, basal ridge
well-sclerotized with two short widely diverging baculi. Gonocoxal apices with distinct
recurved “tooth” present. Two primary and at least two secondary setae originate from small
depressions on the gonocoxal apices (Fig. 237).
Variation. The specimens from Cordoba have a different color pattern than the other
Argentinean material. In these individuals the basal half of the elytra are lighter brown than the
rest of the body. A similar pattern is seen in P. helvolus, a Nearctic member of the genus, where
individuals in the same region have some specimens with darker bodies and lighter elytral bases.
Seasonality/Habitat. Specimens have been collected in March and October.
Distribution. Known from northern and central Argentina.
Notes. No host records are available for this species.
Pocadius tepicensis Cline new species
(Figs. 42, 81, 122, 163, 202, 238)
Type Material Examined. HOLOTYPE ♂ (CAS): 20mi. E. Tepic, Nay.; Mex. VIII-3-
65 / L.R. Gillogly Collector / gill; fungus / L.R. Gillogly Collection / Lorin R. Gillogly;
Collection; Donated To The; Calif. Academy Of Sciences; May 1990 / HOLOTYPE; Pocadius;
tepicensis; A. Cline des. 2004. 12 PARATYPES (CAS): same data labels as holotype.
Diagnosis. This species can be delimited from the rest of the Neotropical fauna by the
following suite of characters: terminal antennomere longer than preceding two and with V-
shaped sensillar region; mentum hemispherical with an obsoletely acute apex; metasternal disc
deeply densely punctate with large punctures; prosternal process in lateral aspect with sharp
declivity posterior to coxal cavities to a flattened posterior third, posterior wall strongly oblique;
275
protibia lacking an outer apical notch; indentate male pygidium; and basal piece of ejaculatory
rods L-shaped lateral arms and scoop-shaped central region.
Description. Length 3.75mm, Width 2.15mm, Depth 1.35mm. Body slightly convex,
surface shining, uniformly reddish-brown to dark brown in color. Pronotum and elytra margins
with moderately elongate fimbriae, setae slightly longer than length of antennal scape. Dorsal
and ventral pubescence moderately long.
Head surface irregularly punctate with deeply impressed small and large punctures, most
larger ones on vertex, becoming somewhat smaller towards orbits and fronotclypeal region.
Larger punctures 4-5X diameter of eye facet, smaller punctures ~2X diameter; interspaces
smooth to finely alutaceous, shining. Each puncture gives rise to a curved golden seta. Pronotal
surface with large punctures equal in size to large punctures on vertex of head, interspersed with
relatively few smaller punctures, ~0.5 size of larger ones; interspaces smooth to finely alutaceous
and shining, ~0.5 diameter apart. Each puncture gives rise to a curved golden seta. Scutellar
surface with vague shallowly impressed small punctures, some punctures giving rise to setae,
interspaces smooth to finely alutaceous. Elytral surface with serial rows of alternating large and
small deep punctures. Smaller punctures are 0.75 diameter to smaller ones on pronotum, larger
punctures are ~3-4X diameter of smaller ones. Smaller punctures giving rise to an erect curved
golden seta, larger punctures giving rise to an erect curved golden seta. Interspaces narrow
between punctures of a given row and broad between different rows. Within a row, small
punctures are separated by ~2 puncture diameters, and large punctures by 0.25-0.5 puncture
diameter. Larger rows are separated by ~3 larger puncture diameters. Interspaces always
shining but variable from smooth to finely alutaceous in sculpture. Pygidium densely punctate,
punctures equal in size to larger ones on pronotum, each puncture giving rise to a short golden
seta; interspaces narrow, ~0.5 diameter, shining, with alutaceous to finely granular sculpture.
276
Venter with shorter sparser pubescence as dorsum. Mentum with minute very shallow
punctures, equal in size to smaller ones on vertex, each giving rise to a seta; interspaces
alutaceous to granular. Submentum and gula similar in punctation to mentum but with
interspaces more granular. Prosternum with moderately impressed punctures and epimeron
deeply irregularly punctate, punctures on epimeron equal to larger ones on pronotum and those
on prosternum 0.75 diameter of those on epimeron, interspaces alutaceous to granular, prosternal
punctures separated by ~1 diameter, those on the epimeron by 0.25-0.5 diameter. Mesosternum
with moderately impressed punctures, equal to those on epimeron, interspaces alutaceous to
granular, separated by ~0.5-1 diameter and aggregated near metasternal border. Metasternum
irregularly punctate, heavily punctate on disc with moderately to deeply impressed large
punctures similar in size to those on mesosternum, interspaces alutaceous to granular on
metasternal disc becoming more granular laterally, punctures separated by ~0.5-1 diameter.
Abdominal sternite 1 with few large faint punctures, punctures equal to those on metasternal
disc, interspaces alutaceous, separated by ~1-2 diameters. Abdominal sternites 2-4 with 3
irregular rows of punctures, one row near anterior margin, one near the posterior margin, and a
confusedly organized row medially, punctures similar in size to those on sternite 1.
Hypopygidium with moderately deep punctures, similar in size to those on sternites 2-4,
interspaces alutaceous to granular, punctures separated by ~0.5 diameter.
Head much wider than long (W:L = 1.9:1), fronotclypeal region somewhat projecting
anteriorly. Vertex with deep concavity between orbits near fronotclypeal region. Labrum with
deep medial incision at anterior margin. Antennal club compact, ovate, slightly asymmetrical
with the last antennomere slightly longer than the previous two combined. Antennomeres 4-8
more or less compact, with 6-8 characteristically disc-like, and 4-5 cuboidal. Antennal scape
asymmetrical, somewhat hemispherical, 1.75X as long as pedicel. Pedicel subcylindrical in
277
shape. Antennal segment 3 ~0.75 length of pedicel. Antennal club moderately large, ~0.65
length of segments 1-8 combined. Antennal grooves very deep and widely excavate, slightly
converging posteriorly. Lateral mental/submental ridge prominent with a corresponding widely
excavate sulcus, ridge at level of submentum, ridge with distinct longitudinal and oblique
microreticulations and sulcus with transverse and oblique microreticulations and granular
sculpture. Mentum with anterior angles obsolete, anterior margin broadly convex with an acute
apex, entire structure obtusely triangular in ventral view and somewhat convex laterally.
Pronotum widest near middle (L:W = 1:1.75), anterior margin shallowly concave,
posterior margin moderately convex, lateral margins slightly arcuate anteriorly and less so
posteriorly. Scutellum large, obtusely triangular, apex rounded. Prosternal process somewhat
narrowed between procoxae, apex somewhat acuminate, in lateral aspect the anterior and
posterior ends are prominent and there is a distinct convexity medially that abruptly declines to a
flattened posterior third. Posterior apical wall moderately prominent and sharply oblique.
Mesosternum extending to midway between mesocoxae, broadly concave to almost truncate for
reception of the metasternum. Metasternum wider than long (W:L = 2.6:1.0). Metepisternum
with slight medial constriction, oblique line dividing anterior 0.20 of structure. First abdominal
sternite with broad process between metacoxae. First sternite ~2X’s longer than second sternite.
Sternites 2-3 subequal in length, the fourth slightly larger than the preceding two.
Hypopygidium subequal in length to first abdominal sternite.
Protibia with apical tooth not prominent, equal to 0.5 length of tarsomere 1. Outer apical
notch shallow, equal to 0.25 length of tarsomere 1 with an ~100°. Inner apical spine subequal in
length to tarsomeres 1. Protibia not heavily armored but with characteristic dense patch of stiff
setae along the inner apical region. Mesotibia somewhat more heavily armored than protibia
with some dense stiff setae and a row of numerous slender spines along entire lateral edge.
278
Outer apical process not robust, subequal to protibia process. Inner apical spine equal in length
to tarsomere 1 and part of 2 combined. Metatibia with armature similar to that of mesotibia, but
with inner apical spine equal to tarsomeres 1-2 combined.
Male genitalia well-sclerotized. Anal sclerite with large broadly curved region
anteriodorsally (Fig. 42); apex moderately fimbriate; ventrally with a broad medial concavity
approaching apex in a truncate manner. Spiculum gastrale with wide lateral indentate flanges,
medial margins deeply concave apically, few short stiff setae originating from apex (Fig. 81).
Tegmen evenly broadly rounded apically (Fig. 122), longer than wide (w:l = 1.0:2.7), lateral row
of setae visible from the median fossa to prior to the apex, large elliptical shallow concavity in
apical third with single row of inner setae not attaining apex, basal notch perpendicular, basal
margin perpendicular to slightly oblique. Median lobe large and robust, ~0.7 the length of the
tegmen, apex evenly broadly narrowed, apical opening well-developed with deeply cleft bilobed
internal structure (Fig. 163). Ejaculatory rods not fused to basal piece or each other, oriented
parallel to each other. Basal piece of internal sac sclerite with lateral arms L-shaped and central
region evenly depressed or scoop-like (Fig. 202).
Female genitalia moderately sclerotized. Paraprocts large with sclerotization only along
median line to apico-lateral angles. Gonocoxite with two basal lateral prominences, basal ridge
well-sclerotized. Gonocoxal apices with recurved “tooth” absent. Four primary setae originate
from small depressions on the gonocoxal apices. Intragonocoxal invagination extremely deep,
~0.8 length of gonocoxite (Fig. 238).
Variation. None observed.
Seasonality/Habitat. All specimens were collected in early August.
Distribution. Known from the type locality only.
279
Notes. The fungal host data available suggests that this species was not collected in a
Gasteromycetes, but rather a mushroom or some other Agaricales. This is the only species
known to occur solely from a non-Gasteromycetes host.
Etymology. Specific epithet is a derivative of the type locality, i.e. Tepic, Mexico.
Pocadius testaceous Grouvelle
(Figs. 43, 82, 123, 164, 203, 239)
Type Material Examined. HOLOTYPE ♀ (MNHN) : Mandar; Bengale / Type /
Museum Paris; 1917; Coll. Grouvelle / 76 / Pocadius; testaceus; ty. Grouv / Holotypus Pocadius;
testaceus Grouvelle; det. Kirejtshuk 1994.
Non-Type Material Examined. 1 ♂ and 1 ♀ (BMNH): CEYLON, C. Prov.; Knucles,
1600m; 28-VI-1983; Ole Mehl Leg. 1 ♂ (RNH, Leiden): India or. Biró, 1902 / Matheran; 800m.
Diagnosis. This species can be distinguished from all other Pocadius by the following
suite of characters: body uniformly testaceous in color; terminal antennomere enlarged,
asymmetrical and longer than previous two segments combined; prosternal process in lateral
aspect with posterior face long and with slight concavity ventrally, posterior region of prosternal
process with marked declivity behind procoxae; metasternal disc glabrous with few widely
spaced minute punctures; mesosternum convex medially; abdominal process broad between
metacoxae; anal sclerite with an acute apical point; inner margin of spiculum gastrale convex;
apex of tegmen truncate to broadly rounded; basal piece of ejaculatory rods with deeply cleft
apices; ovipositor with narrow not broadly flanged paraprocts, gonocoxites with elongate baculi
and prolongations narrowly separated and abutting, intragonocoxal invagination shallow.
280
Redescription. Length 3.3 mm, Width 1.9mm, Depth 1.0mm. Body moderately
convex, surface shining, uniformly testaceous in color. Pronotum and elytra margins with short
fimbriae, setae subequal to length of antennal scape. Dorsal and ventral pubescence short.
Head surface moderately deeply, irregularly punctate, punctures more widely dispersed
on vertex, becoming somewhat more congregated towards orbits and fronotclypeal region.
Larger punctures 4-5 X diameter of eye facet, smaller punctures 3 X diameter. Interspaces
smooth to finely alutaceous, shining. Some punctures give rise to a short somewhat curved
golden seta. Eyes finely faceted. Pronotal surface with large punctures equal in size to large
punctures on vertex of head, interspersed with relatively few smaller punctures, ~0.50 - 0.75 size
of larger ones. Interspaces alutaceous to finely microreticulate, about 1-1.5 diameters apart on
disc. Some punctures gives rise to a short somewhat curved golden seta. Scutellar surface with
very few vague shallowly impressed punctures, punctures similar in size to smaller ones on
pronotum, some punctures giving rise to setae, and the interspaces are alutaceous. Elytral
surface with serial rows of alternating large and small deep punctures. Smaller punctures are
equal in size to larger ones on pronotum, larger punctures are ~1.5 times diameter of smaller
ones. Smaller punctures giving rise to a semi-erect to erect golden seta, larger punctures giving
rise to a decumbent golden seta. Interspaces narrow between punctures of a given row and
between different rows. Within a row, small punctures are separated by ~1 puncture width, and
large punctures by 0.5-0.75 puncture width. Larger rows are separated by 1.5 large puncture
diameters. Interspaces always shining but mostly alutaceous in sculpture. Pygidium densely
punctate more so in the apical 0.33, punctures equal in size to smaller ones on pronotum, each
puncture giving rise to a short stiff golden seta. Interspaces narrow, 0.25-0.5 diameters, with
granular sculpture.
281
Venter with similar short golden pubescence as dorsum. Mentum with shallow minute
punctures, equal in size to smaller ones on vertex, interspaces alutaceous to finely
microreticulate. Submentum and gula similar in punctation to mentum but with interspaces more
granular with some microreticulation present. Prosternum and epimeron shallowly irregularly
punctate, punctures 1.5X larger than those on mentum, interspaces alutaceous to granular,
prosternal punctures separated by 0.5 diameter, those on the epimeron by 0.5 diameter.
Mesosternum with shallow punctures, about 0.75 diameter of those on prosternum, interspaces
alutaceous to granular, separated by about 0.5 to 1 diameter, with most punctures congregated
near the metasternal border. Metasternum irregularly punctate, with faint minute punctures on
disc similar in size to smaller ones on venter, interspaces alutaceous on metasternal disc
becoming more microreticulate to granular laterally, punctures separated by >2-3 diameters
around disc. Abdominal sternite 1 with faint small, almost obsolete punctures, punctures equal
to larger ones on vertex, interspaces alutaceous, separated by ~1-2 diameters. Abdominal
sternites 2-4 with one distinct irregular rows of punctures near anterior margin, punctures in the
row separated by 1 puncture diameters, other non-row punctures more diffusely distributed.
Hypopygidium with more deeply impressed punctures, similar in size to those on sternites 2-4,
interspaces alutaceous to granular, punctures separated by 0.5-1 puncture width.
Head slightly wider than long (W:L = 1.28:1), fronotclypeal region distinctly projecting
anteriorly. Vertex with shallow broad indistinct concavity between orbits near fronotclypeal
region. Eyes large and protruding. Antennal club compact, somewhat oval, slightly
asymmetrical with the last antennomere longer than the previous two combined. Antennomeres
6-8 more or less compact, with 7-8 characteristically disc-like. Antennal scape asymmetrical,
distinctly hemispherical, 1.3 times as long as pedicel. Pedicel subcylindrical in shape. Antennal
segment 3 slightly shorter in length to pedicel. Antennal club large, ~0.95 length of segments 1-8
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combined. Each club segment with dense short setae, and only relatively few protruding setae.
Antennal grooves very deep and widely excavate, slightly converging posteriorly. Lateral
mental ridge prominent, at level of submentum, medially the ridge is granular in sculpture with
some microreticulation and laterally with longitudinal microreticulations. Mentum with anterior
angles distinct, anterior margin angular coming to a distinct medial point, lateral sides
subparallel, overall pentagonal in shape, when viewed laterally the entire structure is flattened.
Pronotum widest near posterior angles (L:W = 1:2.1), anterior margin broadly
shallowly concave, posterior margin moderately convex, lateral margins slightly arcuate
anteriorly, anterior and posterior angle distinct. Scutellum large, obtusely triangular, apex
broadly rounded. Prosternal process somewhat narrowed between procoxae, apex somewhat
acuminate, in lateral aspect the anterior and posterior ends are prominent and convex medially
over procoxae with distinct declivity behind coxal cavities. Posterior apical wall prominent,
mostly straight with a slight concavity ventrally. Mesosternum extending to midway between
mesocoxae, evenly concave for reception of the metasternum, somewhat convex ventrally but
not carinate. Metasternum width to length ratio is ~2.57:1.0. Metepisternum with slight medial
constriction, oblique line dividing anterior 0.10 of structure. Elytral humeri moderately
produced, lateral margin very narrow. First abdominal sternite with broadly convex process
between metacoxae. First sternite ~2X’s longer than second sternite. Sternites 2-4 subequal in
length. Hypopygidium subequal in length to first abdominal sternite.
Protibia with apical tooth distinct, slightly longer than tarsomere 1. Outer apical notch
with ~100° angle, notch depth shallow, equal to length of tarsomere 1. Inner apical spine
subequal in length to tarsomere 1 and part of 2 combined. Protibia not heavily armored but with
characteristic dense patch of stiff setae along the inner apical region. Mesotibia more heavily
armored than protibia with more dense stiff setae and a row of numerous slender spines along
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entire lateral edge. Outer apical process elongate and robust, larger than protibia process. Inner
apical spine equal in length to tarsomeres 1-2 combined. Metatibia with armature similar to that
of mesotibia, but lateral setae somewhat longer.
Male genitalia well-sclerotized. Anal sclerite with large broadly curved region
anteriodorsally (Fig. 43) ending in a distinct apical point; apex somewhat fimbriate; ventrally
with a broad medial concavity approaching apex in more or less truncate manner. Spiculum
gastrale with wide narrowly rounded lateral flanges, medial margins convex, short stiff setae
originating from apex (Fig. 82). Tegmen broadly rounded to truncate apically (Fig. 123), much
longer than wide (w:l = 1.0:2.56), lateral row of setae visible from the median fossa to prior to
the apex, broad shallow concavity in apical third, basal notch perpendicular, basal margin
concave, inner row of setae not attaining apex. Median lobe large and elongate, ~0.75 the length
of the tegmen, apex acuminate, apical opening well-developed with proximal concavity (Fig.
164), lateral margins slightly angulate becoming wider posteriorly. Ejaculatory rods not fused to
each other or to basal piece, straight and subparallel. Basal piece of rods not fused to each other,
deeply cleft in apical 0.66 (Fig. 203).
Female genitalia moderately sclerotized. Paraprocts not broadly flanged with
sclerotization only along median line to baso-lateral angles. Gonocoxite with two basal lateral
prominences, basal ridge well-sclerotized. Gonocoxal apices without recurved “tooth”, and
intragonocoxal invagination shallow and narrow. Four primary setae originate from small
depressions on the gonocoxal apices (Fig. 239).
Variation. Specimens from Ceylon have slightly longer dorsal pubescence than the
specimen from India/Biró, however all other characters are consistent.
Seasonality/Habitat. Specimens from Ceylon were collected in June, and the species is
likely known from the more tropical regions of India.
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Distribution. Known from India and Ceylon.
Notes. No fungal hosts are available for this species.
Pocadius torresi Jelínek
(Figs. 44, 84, 124, 165, 240)
Type Material Examined. (5) PARATYPES 3 ♀, 2 ♂: two females and one male with
the following data label: Argent. prov.; Buenos Aires; J. Basq coll.; one female and one male
with the following data label: Argentina; LA PLATA; ex. coll. Mus. La Plata. All specimens
with the following labels: Paratypus; Pocadius; torresi; sp. n.; Jelínek det. 1976. / Mus. Nat.
Prague; Inv. #’s 65614 – 65618 respectively.
Non-Type Material Examined. One with the following label data (CMN): ARG: Salta
Prov.; El Rey Nat. Park, 900m; 7-XII-1987, S&J Peck; Hasteria, Chaco Forest; leaf and wood
litter.
Diagnosis. Differs from other Pocadius by the absence of setae in the apico-medial
regions of the inner row and median lobe fossa, the almost symmetrical antennal club with rather
short terminal antennomere, and lack of a well-developed protibial apical-lateral notch.
Redescription. Length 4.2mm, Width 2.8mm, 1.8Depth mm. Body uniformly brown-
orange, head and pronotum somewhat darker and less shining than elytra. Pronotum and elytra
with margins with dense and elongate fimbria. Pygidium and hypopygidium densely setate, with
posterior margins moderately fimbriate.
Head surface moderately punctate, large and small punctures interspersed throughout
vertex, becoming more densely packed near orbits and posterior margin. Large punctures ~5
times the diameter of an eye facet, smaller punctures about 0.33 size of larger punctures,
interspaces mostly alutaceous, ~ 1-2 diameters apart. Smaller punctures giving rise to very
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elongate pale golden setae, setae longer than length of eye, overall head very densely pubescent.
Fronotclypeal region with broad shallow concavity between the orbits. Pronotal surface with
large and small punctures interspersed throughout, with the punctures similar in size to
respective large and small punctures on vertex. Elongate golden setae derived from each small
puncture. Interspaces 2-3 diameters apart becoming somewhat more dense laterally, mostly
granular to alutaceous texture. Scutellar surface with faint large punctures similar in size to the
larger punctures on the vertex aggregated along anterior margin, other punctures aggregated in
anterior 0.75 are smaller, similar in size to smaller punctures on vertex , interspaces granular, a
few elongate setae derived from some of the smaller punctures. Eytral surface with serial rows
of alternating large and small shallow punctures. Large punctures irregularly rounded, ~1.5
times the size of the large pronotal punctures, each giving rise to a single erect elongate curved
seta. Small punctures 0.25 diameter of large punctures, each giving rise to a smaller semi-erect
to decumbent seta, setal length about 0.75 length of setae derived from larger punctures. Large
and small punctures in single rows. Interspaces mostly alutaceous between all punctures, larger
punctures separated from each other by 0.5 puncture diameter, smaller punctures separated from
each other by 1-2 puncture diameters. Large and small puncture rows separated from each other
by ~1-1.5 large puncture diameter. Pygidium densely irregularly punctate, punctures equal in
size to large punctures on pronotum, interspaces alutaceous with clear transverse
microreticulation present, punctures separated by ~0.25 puncture diameter. Some but not all
punctures giving rise to a moderately long setae, setae equal to length of those from small
punctures on elytra, becoming longer laterally.
Venter with shorter pubescence than dorsum, setae about 0.5 to 0.75 length of pronotal
setae. Mentum with faint large punctation, punctures similar in size to ~0.75 the size of the large
vertex punctures, some of the punctures giving rise to moderately long setae, interspaces
286
granular with some microreticulation present. Submentum and gula with very faint scattered
small punctures equal to 0.5 size of those on mentum, interspaces very granular, with no setae
derived from them. Prosternum and epimeron with very faintly impressed punctures
approximately 1.25 size of mentum punctures, interspaces alutaceous, prosternal punctures
separated by 0.5 diameter, those on the epimeron similarly separated. Prosternal punctures
giving rise to moderately long setae the are curved apically. Mesosternum with very faint large
punctures along posterior margin, interspaces alutaceous. Metasternum irregularly punctate,
with punctures on disc similar in size to small mentum punctures, interspaces smooth to
alutaceous on metasternal disc becoming more granular laterally, punctures separated by ~3
diameters becoming somewhat more dense laterally, no disc punctures giving rise to setae, but
lateral punctures giving rise to long setae. Metepisternum with dense elongate setae, setae longer
than anywhere else on venter. Abdominal sternite 1 with very faint medium sized punctures that
are slightly larger than those on the metasternum, interspaces ~2 diameters apart, mostly
alutaceous. Abdominal segments 2-4 with similar faint punctures as on sternite 1, however they
are aligned in irregular lateral rows, interspaces alutaceous to granular, punctures separated by
0.25-0.5 diameter, each puncture giving rise to a long seta. Hypopygidium more densely deeply
punctate than other sternites, punctures ~0.75 the size of those on abdominal sternites 2-4,
interspaces alutaceous to granular, punctures separated by ~0.5-1 diameter of puncture.
Head wider than long (W:L = 1.25:1), somewhat triangular, with fronotclypeal region
projecting anteriorly. Labrum transverse with sharp triangular medial incision on anterior
margin. Antennal club compact, oval, densely setose, almost symmetrical, with the last
antennomere subequal to the previous two combined. Antennomeres 6-8 flattened into disc-like
structures, their combined length equal to the length of antennomere 9. Antennal scape
asymmetrical, hemispherical, 1.75 times as long as pedicel, numerous elongate setae arising from
287
anterior margin of scape. Pedicel subcylindrical in shape tapering proximally, widest near
middle. Antennal segment 3 subcylindrical, about 0.75 length of pedicel, tapering proximally as
in the pedicel. Segments 4-5 similar in shape to segment 3, each ~0.5 the length of segment 3.
Antennal club large, subequal in length to ~0.60 the length of segments 1-8 combined. Antennal
grooves quite deep and medially curved posteriorly, antennal ledge extending from submentum
laterally along mentum with a longitudinal and oblique microreticulate surface. Mentum with
apex evenly narrowly rounded anteriorly with short medial angle, lateral angles faint but present,
overall pentagonal and somewhat convex.
Pronotum broadly convex, widest near middle (W:L = ~1.9:1), anterior margin broadly
deeply concave to trapezoidal, posterior margin broadly convex with slight undulation near
humeri. Prosternal process narrowed between coxae, somewhat acute apex, in lateral aspect the
process is almost evenly convex, with greatest height achieved at midline of procoxae. Posterior
wall well-developed and slightly oblique. Scutellum large, elongate triangular with rounded
apex. Mesosternum extending to midway between mesocoxae, posterior border truncate.
Mesepisternum wider than mesosternum. Mesepimeron ~0.33 the width of mesepisternum.
Metasternum width to length ratio is ~2.1:1. Metepisternum broad, not concave medially,
anterior third produced anteriomedially, anterior 0.13 cut-off from rest of metepisternum by faint
incomplete carina. First abdominal sternite with triangular process between metacoxae with
produced medial acute tip, first sternite ~2 X’s longer than second sternite. Sternites 2-4
subequal in length. Hypopygidium equal to the length of the first abdominal sternite.
Protibia with slight lateral curvature, crenulate along lateral edge. Apical tooth
prominent, subequal to tarsomeres 1 and 2 combined. Outer apical notch indistinct, notch depth
equal to 0.25 length of tarsomere 1. Inner apical spine subequal in length to first tarsomere and
0.5 of second. Protibia with well-developed armature overall, several rows of short stiff spines
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visible. Mesotibia more heavily armored than protibia with several rows of slender spines along
lateral edge and medial and ventral surfaces. Outer apical process more robust, somewhat larger
than protibial process, projecting more posteriorly than protibial process. Inner apical spine
equal in length to tarsomere 1 and 2 combined.
Metatibia heavily armored with numerous rows of slender more elongate spines. Spines of
varying lengths, but most longer than those on the mesotibia. Outer apical process robust,
projecting more posteriorly than mesotibial process. Inner apical spine subequal in length to
tarsomeres 1, 2 and 0.5 of the third combined.
Male genitalia moderately sclerotized. Anal sclerite with very broad tegminal fossa, apex
and lateral borders well-sclerotized, ~35 elongate setae on apical border (Fig. 44). Spiculum
gastrale with ~20 setae extending from the apical margin, lateral flange widely rounded and not
well-developed, lateral flange on same plane as rest of sclerite, spiculum attached posterio-
medially (Fig. 83). Tegmen not elongate (W:L = ), large median lobe fossa that is longer than
0.5 the length of the tegmen, cilia along apico-lateral regions of medina lobe fossa but not apico-
medially, vaguely defined apical fossa with no cilia around it, tegmen apex truncate, lateral edge
of tegmen with long setae derived from small pits, inner row of setae originating apico-laterally
but also absent apico-medially, posterior prominence projecting forward, posterior margin
shallowly concave (Fig. 124). Median lobe quite large, greater than half the length of the
tegmen, with bilobed apical opening (Fig. 165). Internal sac sclerites not observed.
Female genitalia moderately to well sclerotized. Gonocoxites elongate, well-developed,
intragonocoxal invagination wide, greater than width of gonocoxite process, invagination with
acute excavate base, gonocoxite apices with faintly developed recurrent “tooth” present, three
primary setae extending from apical pit of gonocoxal apices, basal gonocoxal ridge well-
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sclerotized, oblique sclerotized baculi short, basal sclerotized ridge with only one basal
prominence (Fig. 240).
Variation. No noticeable variation between the five specimens studied.
Seasonality/Habitat. Known from early December from non-type material.
Distribution. Known from the type series in the Buenos Aries province of Argentina
and La Salta Province.
Notes. No host information is available for this species.
Pocadius wappesi Cline n. sp.
(Figs. 45, 84, 125, 166, 204, 241)
Type Material Examined. HOLOTYPE ♂ (UASC): BOLIVIA: Santa Cruz; 4-6k SSE
Buena Vista; F&F Hotel, 2-12 Feb.; 2000, JE Wappes / transition tropical; forest, 420-450m /
HOLOTYPE; Pocadius; wappesi; A.R. Cline des. 2004. 1 PARATYPE (CDFA): same data
label as holotype but with PARATYPE designation label. 2 PARATYPES (SNEC): BOLIVIA:
Santa Cruz Dept.; 3.7km SSE Buena Vista; Hotel Flora-Fauna, 400-440m; 17°29.949’S
63°33.152’W; R. Leschen, 9-Nov. 2002; Ex. Lycoperdon fungus, #054 / PARATYPE; Pocadius;
wappesi; A.R. Cline des. 2004. 5 PARATYPES (CMN): BOLIVIA: Sta. Cruz, 5km; SSE Buena
Vista, Hotel; Flora y Fauna, 17°29.925’S; 63°39.128’W, 440m forest,; FIT, 15-24 December
2003; S&J Peck 03-131 / PARATYPE; Pocadius; wappesi; A.R. Cline des. 2004.
Diagnosis. This species can be delimited from the rest of the Neotropical fauna by the
following suite of characters: mentum robustly pentagonal; terminal antennomere without well-
defined sensillar region; prosternal process in lateral view with pronounced declivity posteriorly
past coxal cavities to flattened posterior third, posterior wall prominent and oblique; all tibia
well-armored; elytra with alternating rows of semi-erect setae; dorsal surface almost completely
290
granular in surface sculpture and only moderately shining; abdominal process acute; metasternal
disc distinctly punctate; aedeagus with tegmen having a complete inner row of setae; ejaculatory
rods partially fused to each other in apical 0.33, and curving inward basally; basal piece of
internal sac sclerites with paired short curved structures and a globular central piece that is
concave basally and distally; ovipositor with short intragonocoxal invagination, ~0.33 length of
gonocoxite, three elongate primary setae, and two short oblique baculi.
Redescription. Length 3.8 mm, Width 2.25mm, Depth 1.55mm. Body moderately
convex, surface not to only moderately shining, brown to dark brown in color, with the elytral
apices darker in some specimens. Pronotum and elytra margins fimbriate, setae equal to slightly
longer than length of antennal scape. Dorsal and ventral pubescence moderately long.
Head surface irregularly punctate with moderately impressed small and large punctures,
most larger ones on vertex, becoming somewhat smaller towards orbits and fronotclypeal region.
Larger punctures ~4X diameter of eye facet, smaller punctures 1-2X diameter; interspaces
alutaceous to granular, somewhat shining. Each puncture gives rise to a curved golden seta.
Pronotal surface with large punctures equal in size to large punctures on vertex of head,
interspersed with relatively few smaller punctures, ~0.5 size of larger ones; interspaces granular
and somewhat shining, ~0.25-0.75 diameter apart. Each puncture gives rise to a curved golden
seta. Scutellar surface with few vague shallowly impressed small punctures, equal to smaller
ones on pronotum, some punctures giving rise to setae, interspaces granular with some
microreticulation present. Elytral surface with serial rows of alternating large and small deep
punctures. Smaller punctures are equal in size to smaller ones on pronotum, larger punctures are
~3 times diameter of smaller ones. Smaller punctures giving rise to a semi-erect long golden
seta, larger punctures giving rise to a semi-erect long golden seta. Interspaces narrow between
punctures of a given row and between different rows. Within a row, small punctures are
291
separated by ~1-2 puncture diameters, and large punctures by ~0.5 puncture diameter. Larger
rows are separated by 1-2 larger puncture diameters. Interspaces scarcely shining and granular
in sculpture. Pygidium densely punctate, punctures equal in size to smaller ones on pronotum,
each puncture giving rise to a short golden seta; interspaces narrow, 0.25-0.5 diameter, with
granular sculpture.
Venter with shorter pubescence than dorsum. Mentum with minute very shallow almost
obsolete punctures, equal in size to smaller ones on vertex, each giving rise to a short seta;
interspaces alutaceous to finely granular. Submentum and gula similar in punctation to mentum
but with interspaces more granular. Prosternum and epimeron shallowly irregularly punctate,
punctures on epimeron equal to larger ones on pronotum and those on prosternum 0.50 diameter
of those on epimeron, interspaces granular with microreticulate areas, prosternal punctures
separated by 0.5-1 diameter, those on the epimeron by 0.25-0.5 diameter. Mesosternum with
faintly impressed punctures, equal to those on epimeron, interspaces granular, separated by about
~1 diameter and most but not all aggregated near metasternal border. Metasternum irregularly
punctate, large punctures on disc similar in size to those on mesosternum; interspaces alutaceous
to granular on metasternal disc becoming more granular laterally, punctures separated by ~1-2
diameters on disc. Abdominal sternite 1 with large faint punctures, punctures equal to those on
metasternum, interspaces granular, separated by ~1-2 diameter. Abdominal sternites 2-4 with
irregular rows of punctures, punctures similar in size to those on sternite 1. Hypopygidium with
moderately deep punctures, similar in size to those on sternites 2-4, interspaces granular,
punctures separated by ~0.5 diameter.
Head much wider than long (W:L = 1.7:1), fronotclypeal region moderately projecting
anteriorly. Vertex with shallow concavity between orbits near fronotclypeal region. Labrum
with deep medial incision at anterior margin. Antennal club compact, obovate, slightly
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asymmetrical with the last antennomere slightly longer than the previous two combined.
Antennomeres 4-8 more or less compact, with 6-8 characteristically disc-like, and 4-5
trapezoidal. Antennal scape asymmetrical, somewhat hemispherical, 1.8X as long as pedicel.
Pedicel subcylindrical in shape. Antennal segment 3 subequal in length to pedicel. Antennal
club moderately large, ~0.60 length of segments 1-8 combined. Antennal grooves moderately
deep and excavate, slightly converging posteriorly. Lateral mental/submental ridge prominent
with a corresponding excavate sulcus, ridge at level of submentum, ridge with longitudinal
microreticulations and sulcus with alutaceous to granular sculpture and faint oblique
microreticulations. Mentum with anterior angles present, anterior margin angulate, entire
structure pentagonal in ventral view and somewhat convex laterally.
Pronotum widest near middle (L:W = 1:2.2), anterior margin shallowly concave almost
truncate, posterior margin moderately convex, lateral margins more arcuate anteriorly.
Scutellum large, obtusely triangular, apex rounded. Prosternal process somewhat narrowed
between procoxae, apex somewhat acuminate, in lateral aspect the posterior end is prominent
and there is a slight convexity medially, posterior to coxal cavities there is a sharp declivity to a
flattened posterior thirds of the structure. Posterior apical wall prominent and strongly oblique.
Mesosternum extending to midway between mesocoxae, evenly concave for reception of the
metasternum. Metasternum wider than long (W:L = 3.2:1.0). Metepisternum with slight medial
constriction, oblique line dividing anterior 0.14 of structure. First abdominal sternite with
acuminate process between metacoxae. First sternite ~2X’s longer than second sternite.
Sternites 2-3 subequal in length, the fourth slightly larger than the preceding two.
Hypopygidium subequal in length to first abdominal sternite.
Protibia with apical tooth moderately prominent, subequal to length of tarsomere 1.
Outer apical notch absent. Inner apical spine subequal in length to tarsomere 1. Protibia heavily
293
armored with numerous setae and with characteristic dense patch of stiff setae along the inner
apical region. Mesotibia more heavily armored than protibia with numerous dense stiff setae and
a row of numerous slender spines along entire lateral edge. Outer apical process robust, larger
than protibial process. Inner apical spine equal in length to tarsomere 1 and part of 2 combined.
Metatibia with heavier armature than that of mesotibia, and inner apical spine equal to length of
tarsomeres 1-2 combined.
Male genitalia well-sclerotized. Anal sclerite with large broadly curved region
anteriodorsally (Fig. 45); apex fimbriate; ventrally with a broad medial concavity approaching
apex in a broadly convex to truncate manner. Spiculum gastrale with wide lateral flanges,
medial margins concave, many elongate stiff setae originating from apex (Fig. 84). Tegmen
evenly rounded apically (Fig. 125), longer than wide (w:l = 1:2.3), lateral row of setae visible
from the median fossa to prior to the apex, large elliptical shallow concavity in apical third with
single row of inner setae completely attaining apex, basal notch perpendicular, basal margin
slightly concave. Median lobe large and robust, ~0.66 the length of the tegmen, apex moderately
narrowed, apical opening well-developed with widely bilobed internal structure (Fig. 166).
Ejaculatory rods not fused to basal piece but to each other in apical 0.33, slightly divergent from
one another basally where they are inwardly curved. Basal piece of internal sac sclerite with
paired short curved structures laterally and a medial globular structure with concavities apically
and basally (Fig. 204).
Female genitalia moderately sclerotized (Fig. ?). Paraprocts moderately large with
sclerotization only along median line to apico-lateral angles. Gonocoxite with two basal lateral
prominences, basal ridge well-sclerotized. Gonocoxal apices without recurved “tooth”. Three
primary setae originate from small depressions on the gonocoxal apices. Intragonocoxal
invagination shallow, not greater than 0.33 entire length of gonocoxite (Fig. 241).
294
Variation. No demonstrable variation.
Seasonality/Habitat. All specimens were collected in a lowland tropical transition
forest from November through February.
Distribution. Known only from the type locality.
Notes. Specimens were collected from Lycoperdon fungi.
Etymology. Specific epithet honors Jim Wappes for introducing me to fieldwork in
Bolivia and his generosity during the course of this research.
Pocadius yunnanensis Grouvelle ‘nomen reinstated’
(Fig. 242)
Type Material Examined. HOLOTYPE (MNHN) 2 specimens on same card mount:
Yunnan / MUSEUM PARIS; Yunnan; H. Donckier 1907 / Pocadius; yunnanis; Gr. ex. type
[specific epithet published as yunnanensis in 1910 and again in 1913 catalog by author]. CO-
TYPE (BMNH) ♀: Yunnan; Mission / Yunnan; 1908-55 / [circular label] Co-type / Pocadius;
yunnanensis; ty. Grouv.
Diagnosis. The following suite of characters clearly delimit P. yunnanensis from all
other Old World species: circular antennal club, obsoletely carinate mesosternum, broad
abdominal process extending between metacoxae, smaller less protuberant eyes, pronotum
densely punctate, protibia with outer apical notch indistinct, and gonocoxae with shallow
incision between extensions, gonocoxal apices with 3 primary setae, and sclerotized baculi at
base of gonocoxite reduced.
Redescription. Length 3.2 mm, Width 1.3mm, Depth 0.9mm. Body moderately
convex, surface shining, brown to dark brown in color, with the extreme lateral pronotal margins
and basal 0.25 of elytra excluding the humeri orange-brown. Pronotum and elytra margins
295
fimbriate, setae equal to length of antennal scape. Dorsal and ventral pubescence moderately
long.
Head surface deeply, irregularly punctate, large and small punctures evenly dispersed on
vertex, becoming more densely aggregated towards orbits and fronotclypeal region. Larger
punctures 4 X diameter of eye facet, smaller punctures 3 X diameter. Interspaces alutaceous and
shining. Each smaller puncture gives rise to a golden seta. Pronotal surface with large punctures
slightly larger in size to large punctures on vertex of head, interspersed with smaller punctures,
roughly equal in size to larger ones on vertex. Interspaces smooth to alutaceous, about 0.25-0.5
diameters apart. Each puncture gives rise to a golden seta, most seta are slightly curved but
some are rather straight. Scutellar surface with vague shallowly impressed punctures similar in
size to smaller ones on pronotum, some punctures giving rise to setae, and the interspaces are
alutaceous to granular. Elytral surface with serial rows of alternating large and small deep
punctures. Smaller punctures are equal in size to smaller ones on pronotum, larger punctures are
~2 times diameter of smaller ones. Smaller punctures giving rise to a semi-erect moderately long
golden seta, larger punctures giving rise to a semi-erect shorter golden seta, all setae have curved
apices. Interspaces narrow between punctures of a given row and between different rows.
Within a row, small punctures are separated by ~1 puncture width, and large punctures by 0.5
puncture width. Rows are separated by 0.5-0.75 large puncture diameters. Interspaces always
shining but variable from smooth to alutaceous in sculpture. Pygidium densely punctate,
punctures equal in size to smaller ones on pronotum, each puncture giving rise to a short stiff
golden seta. Interspaces narrow, 0.33 diameters, with granular sculpture.
Venter with similar moderately long golden pubescence as dorsum. Mentum with several
somewhat deeply impressed punctures, equal in size to larger ones on vertex, each giving rise to
a golden seta. Interspaces granular to finely microreticulate. Submentum and gula with more
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shallowly impressed punctation to mentum but with interspaces completely granular.
Prosternum and epimeron deeply irregularly punctate, punctures slightly larger than those on
mentum, interspaces alutaceous to granular, prosternal punctures separated by 0.25 diameter,
those on the epimeron by 0.25 diameter. Mesosternum with shallow punctures, similar in
diameter to those on prosternum, interspaces alutaceous to granular, separated by about 0.5
diameter, all punctures aggregated along metasternal border. Metasternum irregularly punctate,
with moderately faint punctures on disc similar in size to those on mesosternum, interspaces
granular throughout, punctures separated by ~0.5-1 diameters. Abdominal sternite 1 with large
faint, almost obsolete punctures, punctures equal to those on metasternum, interspaces granular,
separated by ~1 diameter. Abdominal sternites 2-4 with two irregular rows of punctures, one
row near anterior margin and the other near posterior margin, punctures similar in size to those
on metasternum, rows separated by 2-3 puncture diameters, punctures within rows separated by
~0.33 punctures width, punctures not in rows diffusedly dispersed. Hypopygidium with
moderately deep punctures, similar in size to those on sternites 2-4, interspaces granular,
punctures separated by 0.5 puncture diameters.
Head slightly wider than long (W:L = 1.65:1), fronotclypeal region moderately projecting
anteriorly. Vertex with broad shallow obsoletely defined concavity between orbits near
fronotclypeal region. Eyes large, moderately protruding. Labrum with shallow obsolete
indentation at anterior margin. Antennal club compact, nearly circular, not asymmetrical, with
the last antennomere subequal to the previous two combined. Antennomeres 6-8 more or less
compact and characteristically disc-like. Antennal scape asymmetrical, somewhat
hemispherical, 1.9 times as long as pedicel. Pedicel subcylindrical in shape. Antennal segment
3 subequal in length to pedicel. Antennal club moderately large, ~0.55 length of segments 1-8
combined. Each club segment with dense short setae, and only relatively few protruding setae.
297
Antennal grooves very deep and widely excavate, slightly converging posteriorly. Lateral
mental ridge prominent and elongate, at level of submentum, ridge extends completely to apex of
gula, all microsculpturing of ridge surface is longitudinal. Mentum with anterior angles visible,
anterior margin angular to narrowly rounded apex, overall pentagonal in shape, entire structure
flattened when viewed laterally.
Pronotum widest near posterior angles (L:W = 1:1.9), anterior margin broadly concave
somewhat truncate, posterior margin moderately convex, lateral slightly arcuate anteriorly.
Scutellum large, obtusely triangular, apex broadly rounded. Prosternal process somewhat
narrowed between procoxae, apex somewhat acuminate, in lateral aspect the anterior and
posterior ends are prominent and convex medially. Posterior apical wall short and slightly
oblique. Mesosternum extending to midway between mesocoxae, evenly concave for reception
of the metasternum, with faintly produced medial carina. Metasternum width to length ratio is
~2.66:1.0. Metepisternum with slight medial constriction, oblique line dividing anterior 0.125 of
structure. Elytral humeri moderately produced, lateral margin very narrow. First abdominal
sternite with broadly rounded process between metacoxae. First sternite ~2X’s longer than
second sternite. Sternites 2-4 subequal in length. Hypopygidium subequal in length to first
abdominal sternite.
Protibia with apical tooth prominent, slightly longer than tarsomeres 1. Outer apical
notch indistinct. Inner apical spine subequal in length to tarsomeres 1 and part of 2 combined.
Protibia not heavily armored but with characteristic dense patch of stiff setae along the inner
apical region. Mesotibia more heavily armored than protibia with more dense stiff setae and a
row of numerous slender spines along entire lateral edge. Outer apical process elongate and
robust, larger than protibia process. Inner apical spine equal in length to tarsomeres 1-2
combined. Metatibia with more armature than that of mesotibia, spines longer and more robust.
298
Male genitalia not observed.
Female genitalia moderately sclerotized. Paraprocts moderately large with sclerotization
only along median line to lateral angles. Gonocoxite with two basal lateral prominences and one
medial prominence, basal ridge well-sclerotized. Gonocoxal apices without recurved “tooth”.
Three primary setae originate from small depressions on the gonocoxal apices. Intragonocoxal
invagination shallow, not greater than 0.33 entire length of gonocoxite (Fig. 242).
Variation. No demonstrable variation between the three specimens observed.
Seasonality/Habitat. No seasonality information or specific habitat information is
known for this species.
Distribution. Known only from the type locality of Yunnan, China.
Notes. Kirejtshuk (1984) synonomized this species with P. nobilis Reitter without giving
detailed morphological explanations for this change, however distinct differences are available
that preclude the inclusion of P. yunnanensis with P. nobilis. Kirejtshuk stated that his
taxonomic change was “based on a study of many specimens of P. nobilis, and the type series of
P. yunnanensis from (RNH)…”. Specimens of P. yunnanensis from RNH possessing label data
Kirejtshuk mentioned were not located during the course of this study. However, one Co-type
specimen with the exact label data provided by Kirejtshuk was borrowed from the BMNH. This
specimen and the two specimens from Paris are the basis for the reinstatement of P. yunnanensis
as a valid species. No host data are available for this species.
POCADIUS PHYLOGENY
Following the phylogenetic analyses a total of 1131 equally parsimonious trees were
constructed, which included all 46 in-group species and 9 out-group taxa. Figure 243 represents
the strict consensus tree from this analysis.
299
Fig. 6. Anal sclerite,
P. adustus.
Fig. 8. Anal sclerite,
P. antennuliferus.
Fig. 7. Anal sclerite,
P. africanus.
Fig. 9. Anal sclerite,
P. ashei.
Fig. 10. Anal sclerite,
P. barclayi.
Fig. 11. Anal sclerite,
P. basalis.
Fig. 12. Anal sclerite,
P. brevis.
Fig. 13. Anal sclerite,
P. carltoni.
Fig. 14. Anal sclerite,
P. centralis.
Fig. 17. Anal sclerite,
P. crypsis.
Fig. 15. Anal sclerite,
P. cochabambus.
Fig. 16. Anal sclerite,
P. coxus.
300
Fig. 18. Anal sclerite,
P. dimidiatus
.
Fig. 19. Anal sclerite,
P
.
dominicus
.
Fig. 20. Anal sclerite,
P
.
end
r
oedyi
.
Fig. 22. Anal sclerite,
P. falini.
Fig. 21. Anal sclerite,
P. ephite.
Fig. 23. Anal sclerite,
P. ferrugineus.
Fig. 24. Anal sclerite,
P. fulvipennis.
Fig. 25. Anal sclerite,
P. fumatus.
Fig. 26. Anal sclerite,
P. fusiformis.
Fig. 27. Anal sclerite,
P. globularis.
Fig. 28. Anal sclerite,
P. helvolus.
Fig. 29. Anal sclerite,
P. insularis.
301
Fig. 30. Anal sclerite,
P.
j
elineki.
Fig. 31. Anal sclerite,
P. kire
j
tshuki.
Fig. 32. Anal sclerite,
P. luisalfredoi.
Fig. 33. Anal sclerite,
P. majusculus.
Fig. 35. Anal sclerite,
P. niger.
Fig. 34. Anal sclerite,
P. ma
q
ui
p
ucunensis.
Fig. 37. Anal sclerite,
P. nobilis.
Fig. 38. Anal sclerite,
P. okinawaensis.
Fig. 36. Anal sclerite,
P. nigerrimus.
Fig. 39. Anal sclerite,
P. pecki. Fig. 40. Anal sclerite,
P. peruensis. Fig. 41. Anal sclerite,
P. rubidus.
302
Fig. 42. Anal sclerite,
P. tepicensis.
Fig. 43. Anal sclerite,
P. testaceous.
Fig. 44. Anal sclerite,
P. torresi.
Fig. 45. Anal sclerite,
P. wappesi.
Fig. 46. Spiculum gastrale,
P. adustus.
Fig. 47. Spiculum gastrale,
P. africanus.
Fig. 50. Spiculum gastrale,
P. basalis.
Fig. 49. Spiculum gastrale,
P. barclayi.
Fig. 48. Spiculum gastrale,
P. antennuliferus.
Fig. 51. Spiculum gastrale,
P. brevis.
Fig. 52. Spiculum gastrale,
P. carltoni.
Fig. 53. Spiculum gastrale,
P. centralis.
303
Fig. 54. Spiculum gastrale, Fig. 55. Spiculum gastrale,
P. coxus. Fig. 56. Spiculum gastrale,
P. crypsis.
P. cochabambus.
Fig. 57. Spiculum gastrale,
P. dimidiatus.
Fig. 58. Spiculum gastrale,
P. dominicus.
Fig. 59. Spiculum gastrale,
P. endroedyi.
Fig. 60. Spiculum gastrale,
P. ephite.
Fig. 61. Spiculum gastrale,
P. falini.
Fig. 62. Spiculum gastrale,
P. ferrugineus.
Fig. 64. Spiculum gastrale,
P. fumatus.
Fig. 65. Spiculum gastrale,
P. fusiformis.
Fig. 63. Spiculum gastrale,
P. fulvipennis.
304
Fig. 66. Spiculum gastrale,
P. globularis.
Fig. 67. Spiculum gastrale,
P. helvolus.
Fig. 68. Spiculum gastrale,
P. insularis.
Fig. 69. Spiculum gastrale,
P. jelineki.
Fig. 70. Spiculum gastrale,
P. kirejtshuki.
Fig. 71. Spiculum gastrale,
P. luisalfredoi.
Fig. 72. Spiculum gastrale,
P. majusculus.
Fig. 73. Spiculum gastrale,
P. maquipucunensis.
Fig. 74. Spiculum gastrale,
P. niger.
Fig. 75. Spiculum gastrale,
P. nigerrimus.
Fig. 76. Spiculum gastrale,
P. nobilis.
Fig. 77. Spiculum gastrale,
P. okinawaensis.
305
Fig. 78. Spiculum gastrale,
P. pecki.
Fig. 79. Spiculum gastrale,
P. peruensis.
Fig. 80. Spiculum gastrale,
P. rubidus.
Fig. 81. Spiculum gastrale,
P. tepicensis.
Fig. 82. Spiculum gastrale,
P. testaceous.
Fig. 83. Spiculum gastrale,
P. torresi.
Fig. 86. Tegmen,
P. africanus.
Fig. 84. Spiculum gastrale,
P. wappesi.
Fig. 85. Tegmen,
P. adjustus.
Fig. 87. Tegmen,
P. antennuliferus.
Fig. 88. Tegmen,
P. ashei.
Fig. 89. Tegmen,
P. barclayi.
.
306
Fig. 90. Tegmen,
P. basalis.
Fig. 91. Tegmen,
P. brevis.
Fig. 92. Tegmen,
P. carltoni.
Fig. 93. Tegmen,
P. centralis.
Fig. 94. Tegmen,
P. cochabambus.
Fig. 95. Tegmen,
P. coxus.
Fig. 96. Tegmen,
P. crypsis.
Fig. 97. Tegmen,
P. decoratus.
Fig. 98. Tegmen,
P. dimidiatus.
Fig. 99. Tegmen,
P. dominicus.
Fig. 100. Tegmen,
P. endroedyi.
Fig. 101. Tegmen,
P. ephite.
307
Fig. 102. Tegmen,
P. falini.
Fig. 103. Tegmen,
P. ferrugineus.
Fig. 104. Tegmen,
P. fulvipennis.
Fig. 105. Tegmen,
P. fumatus.
Fig. 106. Tegmen,
P. fusiformis.
Fig. 107. Tegmen,
P. globularis.
Fig. 110. Tegmen,
P. jelineki.
Fig. 108. Tegmen,
P. helvolus.
Fig. 109. Tegmen,
P. insularis.
Fig. 113. Tegmen,
P. majusculus.
Fig. 112. Tegmen,
P. luisalfredoi.
Fig. 111. Tegmen,
P. kirejtshuki.
308
Fig. 114. Tegmen,
P. maquipucunensis.
Fig. 115. Tegmen,
P. niger.
Fig. 116. Tegmen,
P. nigerrimus.
Fig. 117. Tegmen,
P. nobilis.
Fig. 118. Tegmen,
P. okinawaensis.
Fig. 119. Tegmen,
P. pecki.
Fig. 120. Tegmen,
P. peruensis.
Fig. 121. Tegmen,
P. rubidus.
Fig. 122. Tegmen,
P. tepicensis.
Fig. 125. Tegmen,
P. wappesi.
Fig. 123. Tegmen,
P. testaceous.
Fig. 124. Tegmen,
P. torresi.
309
Fig. 126. Median Lobe,
P. ad
j
ustus.
Fig. 127. Median Lobe,
P. africanus.
Fig. 128. Median Lobe,
P. antennuliferus.
Fig. 129. Median Lobe,
P. ashei.
Fig. 130. Median Lobe,
P. barclayi.
Fig. 131. Median Lobe,
P. basalis.
Fig. 134. Median Lobe,
P. centralis.
Fig. 132. Median Lobe,
P. brevis.
Fig. 133. Median Lobe,
P. carltoni.
Fig. 135. Median Lobe,
P. cochabambus.
Fig. 136. Median Lobe,
P. coxus.
Fig. 137. Median Lobe,
P. crypsis.
310
Fig. 138. Median Lobe,
P. decoratus.
Fig. 139. Median Lobe,
P. dimidiatus.
Fig. 140. Median Lobe,
P. dominicus.
Fig. 143. Median Lobe,
P. falini.
Fig. 141. Median Lobe,
P. endroedyi.
Fig. 142. Median Lobe,
P. ephite.
Fig. 144. Median Lobe,
P. ferrugineus.
Fig. 145. Median Lobe,
P. fulvipennis.
Fig. 146. Median Lobe,
P. fumatus.
Fig. 147. Median Lobe,
P. fusiformis.
Fig. 148. Median Lobe,
P. globularis.
Fig. 149. Median Lobe,
P. helvolus.
311
Fig. 150. Median Lobe,
P. insularis.
Fig. 151. Median Lobe,
P. jelineki.
Fig. 152. Median Lobe,
P. kirejtshuki.
Fig. 153. Median Lobe,
P. luisalfredoi.
Fig. 154. Median Lobe,
P. majusculus.
Fig. 155. Median Lobe,
P. maquipucunensis.
Fig. 156. Median Lobe,
P. niger.
Fig. 157. Median Lobe,
P. nigerrimus.
Fig. 158. Median Lobe,
P. nobilis.
Fig. 159. Median Lobe,
P. okinawaensis. Fig. 160. Median Lobe,
P. pecki.
Fig. 161. Median Lobe,
P. peruensis.
312
Fig. 162. Median Lobe,
P. rubidus.
Fig. 164. Median Lobe,
P. testaceous.
Fig. 163. Median Lobe,
P. tepicensis.
Fig. 165. Median Lobe,
P. torresi.
Fig. 166. Median Lobe,
P. wappesi.
Fig. 167. Internal Sac
Sclerites, P. adjustus.
Fig. 170. Internal Sac
Sclerites, P. ashei.
Fig. 168. Internal Sac
Sclerites, P. africanus.
Fig. 169. Internal Sac
Sclerites, P. antennuliferus.
Fig. 171. Internal Sac
Sclerites, P. barclayi.
Fig. 172. Internal Sac
Sclerites, P. basalis.
Fig. 173. Internal Sac
Sclerites, P. brevis.
313
Fig. 174. Internal Sac
Sclerites, P. carltoni.
Fig. 175. Internal Sac
Sclerites, P. centralis.
Fig. 176. Internal Sac
Sclerites, P. cochabambus.
Fig. 179. Internal Sac
Sclerites, P. dimidiatus.
Fig. 177. Internal Sac
Sclerites, P. coxus.
Fig. 178. Internal Sac
Sclerites, P. crypsis.
Fig. 181. Internal Sac
Sclerites, P. endrodi.
Fig. 182. Internal Sac
Sclerites, P. ephite.
Fig. 180. Internal Sac
Sclerites, P. dominicus.
Fig. 183. Internal Sac
Sclerites, P. falini.
Fig. 184. Internal Sac
Sclerites, P. ferrugineus.
Fig. 185. Internal Sac
Sclerites, P. fulvipennis.
314
Fig. 186. Internal Sac
Sclerites, P. fumatus.
Fig. 187. Internal Sac
Sclerites, P. fusiformis.
Fig. 188. Internal Sac
Sclerites, P. globularis.
Fig. 189. Internal Sac
Sclerites, P. helvolus.
Fig. 190. Internal Sac
Sclerites, P. insularis.
Fig. 191. Internal Sac
Sclerites, P. jelineki.
Fig. 192. Internal Sac
Sclerites, P. luisalfredoi.
Fig. 193. Internal Sac
Sclerites (basal piece not
observed), P. majusculus.
Fig. 194. Internal Sac
Sclerites, P. maquipucunensis.
Fig. 195. Internal Sac
Sclerites, P. niger.
Fig. 196. Internal Sac
Sclerites, P. nigerrimus.
Fig. 197. Internal Sac
Sclerites, P. nobilis.
315
Fig. 198. Internal Sac
Sclerites, P. okinawaensis. Fig. 199. Internal Sac
Sclerites, P. pecki.
Fig. 200. Internal Sac
Sclerites, P. peruensis.
Fig. 203. Internal Sac
Sclerites, P. testaceous.
Fig. 201. Internal Sac
Sclerites, P. rubidus.
Fig. 202. Internal Sac
Sclerites, P. tepicensis.
Fig. 204. Internal Sac
Sclerites, P. wappesi. Fig. 205. Ovipositor,
P. adustus. Fig. 206. Ovipositor,
P. africanus.
Fig. 207. Ovipositor,
P. ashei.
Fig. 209. Ovipositor,
P. basalis.
Fig. 208. Ovipositor,
P. barclayi.
316
Fig. 211. Ovipositor,
P. brevis.
Fig. 210. Ovipositor,
P. bicolor. Fig. 212. Ovipositor,
P. carltoni.
Fig. 215. Ovipositor,
P. dimidiatus.
Fig. 213. Ovipositor,
P. centralis.
Fig. 214. Ovipositor,
P. crypsis.
Fig. 218. Ovipositor,
P. falini.
Fig. 216. Ovipositor,
P. dominicus.
Fig. 217. Ovipositor,
P. endroedyi.
Fig. 219. Ovipositor,
P. femoralis.
Fig. 221. Ovipositor,
P. fulvipennis.
Fig. 220. Ovipositor,
P. ferrugineus.
317
Fig. 222. Ovipositor,
P. fumatus.
Fig. 223. Ovipositor,
P. fusiformis.
Fig. 224. Ovipositor,
P. globularis.
Fig. 225. Ovipositor,
P. helvolus.
Fig. 226. Ovipositor,
P. jelineki.
Fig. 227. Ovipositor,
P. kirejtshuki.
Fig. 228. Ovipositor,
P. luisalfredoi.
Fig. 229. Ovipositor,
P. majusculus.
Fig. 230. Ovipositor,
P. maquipucunensis.
Fig. 233. Ovipositor,
P. niger.
Fig. 231. Ovipositor,
P. martini.
Fig. 232. Ovipositor,
P. monticolis.
318
Fig. 234. Ovipositor,
P. nobilis.
Fig. 235. Ovipositor,
P. okinawaensis. Fig. 236. Ovipositor,
P. peruensis.
Fig. 238. Ovipositor,
P. tepicensis. Fig. 239. Ovipositor,
P. testaceous.
Fig. 237. Ovipositor,
P. rubidus.
Fig. 240. Ovipositor,
P. torresi.
Fig. 242. Ovipositor,
Fig. 241. Ovipositor,
P. wappesi.
P. yunnanensis.
319
The tree length was reported to be 674, CI = 0.24, RI = 0.57, RC = 0.14, and HI = 0.76. There is
an unresolved polytomy basally that contains P. adustus, P. africanus, P. endroedyi, P.
ferrugineus, P. monticolis, P. nobilis, P. okinawaensis, P. rubidus, P. yunnanensis, and P.
rubidus. This unresolved grouping contains two Palearctic members from Europe and Asia (P.
adustus and P. ferrugineus), two Palearctic members from the Orient (P. nobilis, and P.
okinawaensis), three members from Africa (P. africanus, P. endroedyi, and P. monticolis), and
one member from the Neotropics (P. rubidus). Two clades are supported excluding the basal
polytomy: one clade including the remainder of the Old World species (i.e. those from the
Orient, SE Asia, and Australia), and the other containing all of the New World species (except P.
rubidus). Within the Old World clade, the oriental species P. yunnanensis is most basal with the
Australian P. kirejtshuki in a more derived but basal position. The clade P. testaceous + P.
barclayi + P. fusiformis are known from India and Sulawesi respectively. The clade P.
decoratus + P. femoralis + P. majusculus + P. martini are from mainland SE Asia and the
Philippines. The New World clade has a basal rooting with P. fulvipennis from western North
America, and P. globularis from Honduras. Following these two basal taxa is a clade containing
P. niger, P. luisalfredoi, P. basalis, P. brevis, and P. dominicus, which occur in Central America
and the Caribbean. The final large clade of taxa from the New World is rooted with P. helvolus,
P. centralis, and P. carltoni from the Nearctic and Central America, and a large polytomy of
exclusively South American species. The most derived grouping includes P. ashei, P. fumatus,
and P. wappesi, which are from Bolivia and Brazil.
The sup-optimal resolution of the tree shown in Fig. 243 prompted a more in-depth look
at the out-group taxa chosen. The genera Teichostethus, Hebascus, and Hyleopocadius are all
Neotropical taxa that are highly derived and most likely derivatives of Pocadius. Thus, the
original matrix was reconfigured with those genera excluded from a subsequent analysis. The
320
Epuraea
10 Carpophilus
1 Thalycra
Pocadionta
Axyra
2 Lasiodactylus
Hyleopocadius
1 Hebascus
Teichostethus
P. adustus
P. africanus
P. endroedyi
P. ferrugineus
P. monticolis
P. nobilis
P. okinawaensis
P. rubidus
P. yunnanensis
1 P. kirejtshuki
P. testaceous
1 1 P. barclayi
1
4 1 P. fusiformis
P. decoratus
1 P. femoralis
1 P. majusculus
P. martini
P. fulvipennis
P. globularis
1
P. niger
1 P. luisalfredoi
1 P. basalis
1
1 P. brevis
2 P. dominicus
P. helvolus
P. centralis
1 P. carltoni
P. antennuliferus
1 P. bicolor
1 P. cochabambus
P. coxus
P. crypsis
P. dimidiatus
P. ephite
P. falini
P. insularis
1 P. jelineki
P. maquipucunensis
P. nigerrimus
P. pecki
P. peruensis
P. tepicensis
P. torresi
P. ashei
1 P. fumatus
P. wappesi
Figure 243. Strict consensus tree of 46 Pocadius species and nine outgroup taxa (TL = 674, CI =
0.24, RI = 0.57, RC = 0.14, HI = 0.76). Bremer support is indicated above branches with values
≥1.
321
strict consensus tree of the resulting analysis is shown in Fig. 244. The tree length was reported
to be 623, CI = 0.25, RI = 0.57, RC = 0.14, and HI = 0.75. The resulting topology was
significantly different at several nodes. First, the basal polytomy was more clearly defined,
leaving only P. ferrugineus and two African species (P. endroedyi and P. africanus) as basally
unresolved. However, P. endroedyi and P. africanus did group together in a single clade. Also,
there was more basal clarity within the Palearctic and Oriental fauna conferring a more logical
evolutionary progression as discussed below. Most importantly, P. rubidus, moved from a basal
position with the Palearctic fauna to the basal member of the New World clade. The Oriental,
SE Asian, and Australian clade of P. yunnanensis + P. kirejtshuki + P. testaceous + P. barclayi +
P. fusiformis + P. decoratus + P. femoralis + P. majusculus + P. martini remained consistent in
both analyses with no change in the overall topology. Likewise, the two imbedded New World
clades of P. niger + P. luisalfredoi + P. basalis + P. brevis + P. dominicus and the derived P.
helvolus clade with the South American polytomy did not change. Thus the removal of these
three derived out-group genera most notably changed the basal nodes within the overall
Pocadius lineage and two important tree statistics changed as well, namely the dramatic decrease
in tree length (623 in this analysis instead of 674 in the previous analysis) as well as the total
number of most parsimonious trees produced (534 total trees in this analysis compared to 1131
total trees in the previous analysis). Though these two tree values were found to be different, the
other tree statistics remained constant.
As Fig. 244 illustrates, there are still some unresolved nodes in both basal and derived
regions of the tree. To help further resolve these polytomies, all outgroups except Epuraea and
Carpophilus were excluded. These two genera are known to be basal within the Nitidulidae as
evidenced by both morphological (Kirejtshuk 1986c) evidence and molecular data (Cline,
unpublished data). The morphological evidence is primarily concerned with features of the male
322
Epuraea
Carpophilus
Thalycra
Pocadionta
Axyra
Lasiodactylus
P. africanus
P. ferrugineus
P. adustus
P. rubidus
P. fulvipennis
P. endroedyi
P. monticolis
P. nobilis
P. okinawaensis
P. yunnanensis
P. kirejtshuki
P. testaceous
P. barclayi
P. fusiformis
P. decoratus
P. femoralis
P. majusculus
P. martini
P. globularis
P. niger
P. luisalfredoi
P. basalis
P. brevis
P. dominicus
P. helvolus
P. centralis
P. carltoni
P. antennuliferus
P. bicolor
P. cochabambus
P. coxus
P. crypsis
P. dimidiatus
P. ephite
P. falini
P. insularis
P. jelineki
P. maquipucunensis
P. nigerrimus
P. pecki
P. peruensis
P. tepicensis
P. torresi
P. ashei
P. fumatus
P. wappesi
11
2
2
6
2
12
1
1
1
1
1 1 1
1
1 1
1
2
1
1
1
1 1 2
1
1
1
2
1
Figure 244. Strict consensus tree of 46 Pocadius species and six out-group taxa (TL = 623, CI =
0.25, RI = 0.57, RC = 0.14, HI = 0.75). Bremer support is indicated above branches with values
≥1.
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genitalia and the molecular evidence is based on 6 genes representing 7kb. Axyra,
Lasiodactylus, Pocadionta, and Thalycra are all within the same subfamily as Pocadius, e.g.
Nitidulinae, whereas the other two genera are in the Epuraeinae and Carpophilinae. The strict
consensus tree of the resulting analysis is shown in Fig. 245. The tree length was reported to be
568, CI = 0.25, RI = 0.54, RC = 0.14, and HI = 0.75. The resulting topology was significantly
different at some of the basal nodes. First, and foremost, the basal polytomy was resolved with
the two Palearctic species, i.e. P. adustus and P. ferrugineus, being most basal with the three
African species, i.e. P. africanus, P. endroedyi, and P. monticolis, subsequently more derived.
The Oriental taxa, i.e. P. nobilis, P. okinawaensis, and P. yunnanensis, follow with the latter of
these species giving rise to the SE Asian and Australian fauna, i.e. P. kirejtshuki + P. testaceous
+ P. barclayi + P. fusiformis + P. decoratus + P. femoralis + P. majusculus + P. martini. This
SE Asian and Australian fauana remained consistent in both analyses with no change in the
overall topology.. The rest of the tree topology remained constant as well with a derived
polytomy of the South American fauna as the most derived grouping. The trees represented in
Figs. 244 and 245 not only differ in tree topology, but also in tree lenth (Fig. 244 = 623 and Fig.
245 = 568) and the number of trees generated in the analaysis (Fig. 244 = 534 and Fig. 245 =
238). Though these two tree values were found to be different, the other tree statistics remained
constant.
As with the previous analyses, the reduction in outgroups did not resolve the polytomy in
the derived South American clade of P. antennuliferus + P. bicolor + P. cochabambus + P.
coxus + P. crypsis + P. dimidiatus + P. ephite + P. falini + P. insularis + P. jelineki + P.
maquipucunensis + P. nigerrimus + P. pecki + P. peruensis + P. tepicensis + P. torresi + P.
ashei + P. fumatus + P. wappesi. To help resolve this issue the dataset containing the 46
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P. africanus
P. ferrugineus
Epuraea
Carpophilus
P. adustus
P. endroedyi
P. monticolis
P. nobilis
P. okinawaensis
P. carltoni
P. rubidus
P. femoralis
P. yunnanensis
P. fulvipennis
P. kirejtshuki
P. testaceous
P. barclayi
P. fusiformis
P. decoratus
P. majusculus
P. martini
P. globularis
P. niger
P. luisalfredoi
P. basalis
P. brevis
P. dominicus
P. helvolus
P. centralis
P. bicolor
P. antennuliferus
P. cochabambus
P. coxus
P. crypsis
P. dimidiatus
P. ephite
P. falini
P. insularis
P. jelineki
P. maquipucunensis
P. nigerrimus
P. pecki
P. peruensis
P. tepicensis
P. torresi
P. ashei
P. fumatus
P. wappesi
33
1
1
1
1
1
1
1
1 1 1
1
1
1
1
1
1
1
1
1 1 2
1
1
1
1
1
Figure 245. Strict consensus tree of 46 Pocadius species and two out-group taxa (TL = 568, CI =
0.25, RI = 0.54, RC = 0.14, HI = 0.75). Bremer support is indicated above branches with values
≥1.
325
Pocadius and two outgroups (Epuraea and Carpophilus) was successively weighted through four
iterations (Fig. 246). Following these iterations, the South American polytomy was spilt into
several more basal taxa and two derived clades. From the previous polytomy, P. pecki grouped
with P. carltoni to form the basal lineage (P. carltoni originating from Central American and P.
pecki from Venezuela in northern South America). Pocadius ephite follows this basal lineage as
well as the clade P. maquipucunensis + P. jelineki + P. tepicensis. Pocadius falini and P. bicolor
follow the maquipucunensis clade, leaving two clades: 1) the clade of P. crypsis + P.
antennulifus + P. torresi + P. peruensis + P. cochabambus + P. dimidiatus and 2) the clade of
P. insularis + P. nigerrimus + P. coxus + P. ashei + P. fumatus + P. wappesi. The unresolved
highly derived polytomy of P. ashei + P. fumatus + P. wappesi from the previous analyses also
became resolved in the successive weighting protocol, indicating a basal P. ashei and more
derived sister grouping of P. wappesi and P. fumatus.
A final phylogenetic analysis was performed using only external characters to help
decipher the relative value of both external and internal (i.e. genitalia) features. Figure 247
illustrates the strict consensus tree of this analysis. For this analysis all outgroups except
Hebascus, Hyleopocadius and Teichostethus were used. This was deemed appropriate and a
compromise from the above three analyses. From a total of 65 trees, the tree length was reported
to be 407, CI = 0.25, RI = 0.62, RC = 0.15, and HI = 0.75. The resulting topology was most
similar to the prior analyses regarding the Old World taxa, and demonstrated numerous changes
in the New World taxa. Most notable in the Old World analysis was the placement of P.
yunnanensis near P. adustus at the base of the Pocadius group and the placement of P.
ferrugineus near where P. yunnanensis was near the more derived SE Asian/Australian clade.
The SE Asia/Australia clade remained intact with some rearrangements within the clade. The
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Epuraea
Carpophilus
Figure 246. Succesive weighting (4 iterations) of the strict consensus tree of 46 Pocadius species
and two out-group taxa (TL = 568, CI = 0.25, RI = 0.54, RC = 0.14, HI = 0.75).
P. africanus
P. ferrugineus
P. adustus
P. endroedyi
P. monticolis
P. nobilis
P. okinawaensis
P. yunnanensis
P. kirejtshuki
P. testaceous
P. barclayi
P. fusiformis
P. femoralis
P. decoratus
P. majusculus
P. rubidus
P. martini
P. fulvipennis
P. globularis
P. niger
P. luisalfredoi
P. basalis
P. brevis
P. dominicus
P. helvolus
P. carltoni
P. centralis
P. ephite
P. pecki
P. jelineki
P. maquipucunensis
P. bicolor
P. falini
P. tepicensis
P. antennuliferus
P. crypsis
P. cochabambus
P. peruensis
P. torresi
P. dimidiatus
P. insularis
P. coxus
P. nigerrimus
P. ashei
P. fumatus
P. wappesi
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Figure 247. Strict consensus tree of external charcters only for the 46 Pocadius species and six
out-group taxa (TL = 407, CI = 0.25, RI = 0.62, RC = 0.15, HI = 0.75). Bremer support is
indicated above branches with values ≥1.
Epuraea
Carpophilus
Pocadionta
Thalycra
11
6
Axyra
Lasiodactylus
2
P. adustus
2
P. africanus
P. ferrugineus
P. rubidus
P. fulvipennis
P. endroedyi
P. yunnanensis
12
P. monticolis
P. nobilis
P. okinawaensis
P. femoralis
P. majusculus
1
P. kirejtshuki
P. testaceous
P. decoratus
P. martini
P. barclayi
1 P. fusiformis
P. fumatus
P. basalis
P. wappesi
2 P. dimidiatus
P. ephite
P. falini
P. globularis
P. helvolus
P. niger
P. luisalfredoi
P. jelineki
P. pecki
1
P. bicolor
P. brevis
P. maquipucunensis
2 P. dominicus
P. crypsis
P. centralis
P. carltoni
P. peruensis
P. cochabambus
P. nigerrimus
P. antennuliferus
P. coxus
P. insularis
P. tepicensis
P. torresi
P. ashei
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New World taxa, however, notably changed based on external characters alone. Pocadius
fumatus and P. wappesi rooted the New World clade with a large polytomy following. However,
interesting sister groupings became evident that were not clear with the combined analysis. The
following sister relationships were deduced, P. maquipucunensis and P. bicolor, and P. brevis
and P. dominicus, P. crypsis and P. peruensis, P. centralis and P. cochabambus, P. insularis and
P. nigerrimus, and P. coxus and P. tepicensis. A terminal clade of P. torresi + P. antennuliferus
+ P. ashei + P. coxus + P. tepicensis also is well resolved. The influence of external versus
internal characters on the phylogenetic reconstruction of Pocadius and the ramifications of the
resulting external character only tree will be further discussed in the following section.
CHECKLIST OF POCADIUS ERICHSON
Pocadius adustus Reitter. (Distribution: Europe, northern Africa)
Pocadius africanus Kraatz (Distribution: western Africa)
Pocadius antennuliferus Cline n. sp. (Distribution: Brazil)
Pocadius ashei Cline n. sp. (Distribution: Andean Bolivia)
Pocadius barclayi Cline n. sp. (Distribution: Sulawesi region of Indonesia)
Pocadius basalis Schaeffer (Distribution: SW United States of America)
Pocadius bicolor Cline n. sp. (Distribution: Brazil)
Pocadius brevis Grouvelle (Distribution: Cuba)
Pocadius carltoni Cline n. sp. (Distribution: Nicaragua)
Pocadius centralis Cline n. sp. (Distribution: southern Mexico, Honduras, Guatemala)
Pocadius cochabambus Cline n. sp. (Distribution: western Bolivia)
Pocadius coxus Cline n. sp. (Distribution: Brazil)
Pocadius crypsis Cline n. sp. (Distribution: Guyana)
Pocadius decoratus Kirejtshuk (Distribution: Vietnam)
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Pocadius dimidiatus Jelínek (Distribution: Argentina)
Pocadius dominicus Cline n. sp. (Distribution: Dominican Republic)
Pocadius endroedyi Cline n. sp. (Distribution: Tanzania, Zimbabwe, South Africa)
Pocadius ephite Leschen and Carlton (Distribution: Costa Rica)
Pocadius falini Cline n. sp. (Distribution: Paraguay)
Pocadius femoralis Cline n. sp. (Distribution: Vietnam)
Pocadius ferrugineus (Fabricius) (Distribution: Europe to the Russian Far East, northern Africa)
Pocadius fulvipennis Erichson (Distribution: western North America, northern Mexico)
Pocadius fumatus Jelínek (Distribution: Brazil, Argentina)
Pocadius fusiformis Cline n. sp. (Distribution: Sulawesi region of Indonesia)
Pocadius globularis Cline n. sp. (Distribution: Honduras)
Pocadius helvolus Erichson (Distribution: eastern North America, central and northern Mexico)
Pocadius insularis Cline n. sp. (Distribution: Trinidad)
Pocadius jelineki Leschen and Carlton (Distribution: Costa Rica, Nicaragua)
Pocadius kirejtshuki Cline n. sp. (Distribution: Australia)
Pocadius luisalfredoi Cline n. sp. (Distribution: southern Mexico)
Pocadius majusculus Kirejtshuk (Distribution: Thailand)
Pocadius maquipucunensis Leschen and Carlton (Distribution: Ecuador)
Pocadius martini Kirejtshuk (Distribution: Philippines)
Pocadius monticolis Lechanteur (Distribution: Central Africa)
Pocadius niger Parsons (Distribution: SW United States of America)
Pocadius nigerrimus Cline n. sp. (Distribution: Paraguay)
Pocadius nobilis Reitter (Distribution: Japan)
Pocadius okinawaensis Cline n. sp. (Distribution: Okinawa)
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Pocadius pecki Cline n. sp. (Distribution: Venezuela)
Pocadius peruensis Cline n. sp. (Distribution: Peru)
Pocadius rubidus Erichson (Distribution: Argentina)
Pocadius tepicensis Cline n. sp. (Distribution: southern Mexico)
Pocadius testaceous Grouvelle (Distribution: India, Ceylon)
Pocadius torresi Jelínek (Distribution: Argentina)
Pocadius wappesi Cline n. sp. (Distribution: central Bolivia)
Pocadius yunnanensis Grouvelle (Distribution: China)
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CHAPTER 4. DISCUSSION
“The number of minute and obscurely-colored beetles is exceedingly great. It is sufficient
to disturb the composure of an entomologist's mind, to look forward to the future
dimensions of a complete catalogue.”
Charles Darwin, The Voyage of the Beagle
The treatment herein of Pocadius represents the first-ever complete revision of a
globally distributed nitidulid genus, including taxonomic descriptions, keys, biological
notes, geographic ranges, host fungal associations, cladistic phylogenies, phenological
data, and remarks on the evolution of both the genus and tribe. Pocadius species
diversity more than doubled following this revision (46 total species, 25 of which are
new, and one reinstated species). Most new species were discovered from Neotropical
and SE Asian material. The availability of large amounts of small beetle material to
study is due in part to new techniques involved with collecting beetles from the upper
canopy (e.g. chemical fogging, cranes, and inflatable rafts), and other methods for mass
collecting beetles passively (i.e. flight intercept traps, malaise traps, Townes traps, etc.).
These new collecting methodologies in combination with biodiversity initiatives have
generated interest in obscure but diverse groups of beetles such as Nitidulidae.
MONOPHYLY OF POCADIUS AND ITS PLACEMENT IN NITIDULINAE
In all cladistic reconstructions Pocadius was found to be monophyletic.
Synapomorphies uniting all Pocadius species include: a row of punctures along the
anterior metacoxal line, flattened shape of antennomeres 6-8 (though 6 is usually less
flattened than 7 and 8, the latter two being disc-like), an elevated mentum in posterior
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0.66 of structure that forms various geometric shapes, abdominal segments 2-4 with a
row of closely spaced punctures near the posterior margin, a short epicranial stem visible
dorsally on the larval head, and recurved body of mature larvae (2nd instars and further in
development). Thus, the morphological characters chosen for this revision appear to
provide substantial evidence supporting a unified Pocadius. More characters are likely to
come to light once males and females of all species are collected and studied. The
present phylogeny relies heavily on genitalic characters (37% of characters are genitalic,
60% external features, and 3% larval), and with 9 species (~20% of the total number of
known Pocadius) represented by only males or females there is little doubt that the lack
of one sex or the other has affected the tree topology due to the coding of these characters
as “?'” in the matrix.
The delimitation of appropriate outgroups for the phylogenetic analyses proved
more difficult than originally thought, and is likely the cause of the basal polytomy in
Pocadius seen in the reconstruction in Figure 243. These problems are due to the basal
position of Pocadius within the Nitidulinae and Pocadiini (it is not uncommon for a
widely distributed genus to be ancestral in a particular clade). The Pocadiini outgroups
chosen were Hebascus, Hyleopocadius, and Teichostethus. These Pocadiini members are
likely derived from a Pocadius ancestor as they exhibit many derived characters not seen
in Pocadius, including: a more highly modified ovipositor (Hebascus and
Teichostethus), a more loosely joined antennal club (all three genera), pronotal pits and
setal tufts (Hyleopocadius), widely separated metacoxae (Hyleopocadius), elongate
antennal funicle (all three genera), tegmen with apical split (Teichostethus), variously
enlarged maxillary palpi (all three genera), more convex body form (all three genera), an
association with members of the Agaricales, and are restricted to the New World tropics.
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Other Pocadiini taxa also exhibit some of the above characters but were unavailable for
study due to their rarity in collections and the lack of both males and females in museum
holdings. Further work with other Pocadiini members may resolve some of these issues
as more specimens become available.
The Nitidulinae outgroups (Axyra, Lasiodactylus, Pocadionta, and Thalycra) are
also somewhat problematic (see Fig. 244), but less so than the Pocadiini outgroups as
more resolution is obtained basally within Pocadius with the exclusion of the non-
Pocadius Pocadiini taxa, and little further resolution is obtained with the exclusion of the
Nitidulinae outgroups (compare Figs 244 and 245). However, the exclusion of the
Nitidulinae taxa (Fig. 245) does place the two Palearctic species in closer relation (i.e. P.
ferrugineus and P. adustus), as well as placing two of the African species together (i.e. P.
africanus and P. endroedyi) with the other African taxa derived from them (i.e. P.
monticolis); clearly, a more logical configuration. The relationship of the African taxa
would likely be more highly supported, however no males of P. monticolis were available
for study.
The placement of Pocadius within the Nitidulinae was not resolved by this
analysis; however, it was not the major objective of the dataset that was constructed.
Some interesting finds within the Nitidulinae, however, were made based on the dataset
at hand. The Thalycra-complex of genera, which was thought to be sister to Pocadiini
(Kirejtshuk and Leschen 1998), is represented by Thalycra and Pocadionta and is
supported by high Bremer values. However, there is no clear sister relationship to
Pocadius or the other Pocadiini taxa as demonstrated in any of the present
reconstructions. A molecular analysis (in prep.), which is focused more on higher-level
relationships within the Nitidulidae, likewise does not support a close relationship
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between members of Pocadiini and the Thalycra complex. Thus, both morphological and
preliminary molecular evidence does not support a close relationship between these
purported sister tribes. A logical presumption may be an early split of the Thalycra
complex from a common Pocadiini ancestor with subsequent isolation and evolutionary
divergence. This hypothesis, though not tested here, is somewhat supported by
Kirejtshuk’s (1995) conjecture on the mode of life evolution within Nitidulidae.
ORIGIN AND DISTRIBUTION OF POCADIUS
This revision suggests several interesting species distribution patterns and the
subsequent adaptive radiation of the genus. Pocadius, according to the phylogenetic
reconstruction (excluding the reconstruction in Fig. 243 which includes outgroups
Hebascus, Hyleopocadius, and Teichostethus), appears to have originated in the
Palearctic somewhere in Europe and/or northern Africa. The consistent placement of P.
adustus (European/North Africa), P. ferrugineus (European/Asian/North Africa), P.
africanus (western Africa), P. endroedyi (southern Africa), and P. monticolis (central
Africa) clearly denote the western Palearctic or African origin of the genus. From this
region, Pocadius radiated into the rest of the Old World, including all of Asia and
extending into the Malaysian archipelago and Australia. No relationships between the
African and/or Palearctic fauna with the South American fauna, suggests that Pocadius
originated after the break-up of Laurasia and Gondwanaland and the separation of the
current continents. The species from SE Asia, including P. testaceous (India), P.
barclayi (Sulawesi), P. fusiformis (Sulawesi), P. decoratus (Vietnam), P. femoralis
(Vietnam), P. majusculus (Thailand), and P. martini (Philippines) all form a
monophyletic grouping that is most derived within the Old World taxa, thus suggesting
that these species represent the most recent adaptive radiation in the Old World, that of
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penetration into the Old World tropics via radiation from the Palearctic. No sister
relationship between P. endroedyi (South Africa) and P. kirejtshuki (Australia) again
suggests that Pocadius evolved sometime after the current continents separated in the
Mesozoic.
A significant find was a new species from Australia, i.e. P. kirejtshuki, which
extended the known range of the genus into a continent previously known to contain an
undescribed Pocadius (Lawrence 1991). This species is known from the more tropical
eastern and northern areas of Australia and is absent from the more arid western region.
Pocadius kirejtshuki is most closely allied to the SE Asian Pocadius fauna suggesting a
migration from mainland Asia into Australia with subsequent reproductive isolation,
rather than a ancient vicariant event associated with the break-up of Gondwanaland. This
does not necessarily suggest that a Pocadius species could not have evolved in such a
manner and subsequently become extinct, but the present species does not appear to have
arisen in this fashion. Also, a lack of Pocadius in the New World south temperate (Chile
and parts of Argentina) regions suggests that this genus was not present during the
breakup of Gondwanaland. However, there is one shared character between P. endroedyi
and P. kirejtshuki, that of the male anal sclerite with an apical produced tip. The SE
Asian and Australian clade are derived from the Oriental species P. nobilis, P.
okinawaensis, and P. yunnanensis in all phylogenetic reconstructions. This evolutionary
track, or pathway, is not surprising due to the high mountains and arid environments in
Central Asia that do not typically support the preferred fungal hosts for this genus. Thus,
the Old World taxa appear to have evolved from western Palearctic or African ancestors
with expansion into the Orient and subsequent radiation into tropical SE Asia and
Australia.
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The evolution of Pocadius in the New World is a bit more perplexing, and,
according to the phylogeny, likely occurred relatively recently in a geological context.
This recent origin and radiation of species is further substantiated by a lack of Pocadius
(and Pocadiini) in Dominican amber deposits (Cline pers. observation), as well as the
derived polytomy of the Neotropical fauna that suggests not enough time has occurred for
characters to evolve which delineate natural groups. Pocadius rubidus is the only
Neotropical taxon without clear affinities to the rest of the New World fauna based on the
complete dataset with all outgroups included (see Fig. 243.). However, with subsequent
exclusion of the Pocadiini and Nitidulinae outgroup taxa this species becomes established
as the basal taxon of the New World clade (see. Figs. 244, 245, and 246). The position of
P. rubidus is not readily indicative of a Palearctic to Nearctic to Neotropical evolution as
indicated by the rest of the taxa. Removing genitalic characters from the dataset (Fig.
247) subsequently removes P. rubidus from this basal position and places it within a
more derived polytomy. Thus, I suggest that P. rubidus may have reverted to more
plesiomorphic genitalic forms, which in turn placed this species basally in the New
World clade. Character reversion is evident in P. rubidus in the following genitalic
features: development of apico-lateral region of ovipositor gonostylus; degree of fusion
of gonocoxites; number of lateral protuberances on ovipositor gonocoxite base;
development of intragonocoxal invagination; development of oblique baculi on
gonocoxite base; development of tegminal lateral setae; development of subapical furrow
on tegmen; and the number of components of the basal portion of the ejaculatory rods.
Further analysis of males and females of all New World taxa are needed to fully resolve
this issue. Also, one cannot completely abandon the hypothesis that P. rubidus may
indeed be of more ancient Gondwanaland origin with long-term extreme isolation from
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the rest of the Old World fauna and thus retention of some characters more associated
with the Old World fauna.
Other than P. rubidus, the New World fauna follows a pattern of North American
to Central American to South American (as shown in Fig. 246) speciation. There are
exceptions to this general pattern such as the fauna of the Greater Antilles (i.e. P. brevis
and P. dominicus), which is discussed further below. Pocadius fulvipennis, like P.
rubidus, has a basal position in the New World clade; however this species has a more
logical connection to the Old world fauna via a Beringian land bridge link. Following the
arrival of P. fulvipennis, two separate groups originated, one from the Southwestern U.S.,
Mexico, and the Greater Antilles, and another from Eastern North America and the rest of
the Neotropics. The first of these groupings is interesting as it is corroborated by a
peculiar geologic event, i.e. the passing of the Caribbean plate between North and South
America prior to the formation of the Isthmus of Panama. Pocadius luisalfredoi is
known from southern Mexico and, in combination with the two southwestern U.S.
species (P. niger and P. basalis) is sister to the Greater Antilles group (P. brevis from
Cuba and P. dominicus from Hispaniola). The migration of the Caribbean plate between
North and South America was nearing completion approximately 30-40 million years ago
(see Rosen 1985 and Donnelly 1988 for discussion of this geological event and its
pertinence to biogeography). I suggest this event is a good geological marker for
understanding not only the origin of the Greater Antilles Pocadius group, but also for
establishing a conservative estimate for the invasion of South America and thus the time
needed for specialist fungivores to spread and colonize tropical environments. The
placement of P. globularis (Honduras) between P. fulvipennis and the SW U.S. / Greater
Antilles group suggests that there was early colonization of the southernmost reaches of
338
the Tertiary Mexican peninsula sometime prior to the completion of the Panamanian
Isthmus. Other insect groups, including various beetle lineages, have exhibited this
Central American and Greater Antillean connection (Liebherr 1988, Browne, Peck and
Ivie 1993, Gaimari and Erwin 2000, Morrone and Marquez 2001, and Davis, Scholtz and
Philips 2002).
The multiplicity of Pocadius species within South America shows only a vaguely
clear biogeographic pattern of radiation. The successively weighted phylogeny (Fig.
246) does illustrate some trends, but these cannot be quantitatively substantiated. First,
this phylogeny suggests an origin of the South American Neotropical fauna from the
Eastern North American species P. helvolus, and subsequently the widespread Central
American species P. centralis. The southward radiation is continued through southern
Nicaragua (P. carltoni) and eventually into northern South America (P. pecki).
Interestingly, no specimens of Pocadius are known from Panama, despite observations
and identifications of thousands of Panamanian Nitidulidae via loans from many regions
of Panama and personal collecting on Barro Colorado Island.
Upon entering South America via the Panamanian Isthmus, Pocadius underwent a
significant series of speciation events (16 of 46 species, e.g. ~35% of known Pocadius,
occur in South America). According to the phylogeny (Fig. 246), South America was
invaded from species most closely related to P. helvolus (Eastern North America and
Northern Mexico), P. centralis (widespread in Central America), and P. carltoni
(Nicaragua). Three Central American species are embedded within the South American
grouping (i.e. P. ephite, Costa Rica; P. jelineki, Costa Rica and Nicaragua; and P.
tepicensis, Southern Mexico), and may represent speciation in Central America
concurrent or following the invasion of South America. Due to a lack of
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specimens/species known from Panama, Colombia, and Chile, no clear patterns are
readily apparent in the South American fauna and may in part be due to the relatively
recent radiation of the genus into the continent.
HOST FUNGAL EVOLUTION
Prior to this work, most coleopterists (Lawrence 1991), including notable nitidulid
researchers (Parsons 1943, Audisio 1980, Kirejtshuk 1992), suggested that Pocadius was
a specialist solely on members of gasteromycetes fungi in the Lycoperdaceae, in
particular on members of Lycoperdon and Calvatia. This suggestion seemed well-
founded as most collection records at the time consisted of no host data or host data that
only included gasteromycetes genera (Parsons 1936, Leschen and Carlton 1994, and
Kubisz 1995). Only Audisio (1993) suggested that this may not be a clear trend as the
two European species have also been collected from Basidiomycetes, in particular on the
fruiting bodies of Agaricales. However, Kirejtshuk and Leschen (1998) did not consider
Audisio’s recent finding and concurred that members of the Pocadius complex specialize
on fungi with a concealed hymenium. This statement can only be concluded to mean that
members of Pocadius of Pocadiini feed exclusively on gasteromycetes. Members of
Pocadius, for which host information is available, demonstrate that host fungal
preferences do not exclusively include gasteromycetes fungi; these taxa include P.
carltoni, P. crypsis (found in ant refuse pile), P. ferrugineus, P. helvolus, P. tepicensis,
and P. torresi. Ashe and his colleagues utilize a field collecting technique in which
“fungusy logs” are sprayed with a pyrethroid insecticide and the resulting beetles
collected. This technique has produced abundant Pocadius specimens and indicates that
some species may be utilizing polypores (i.e. bracket fungi) and other Basidiomycetes as
a secondary food source and refugia. Thus, the data presented herein suggests that
340
Pocadius adults are, at best, facultative specialists on gasteromycetes and that only the
larval stage appears to be obligately associated with fungi that have a concealed
hymenium. Larvae have only been collected from Lycoperdales. Pocadius adults may
utilize other fungal substrates for food and refuge and then switch to the more ephemeral
Lycoperdales for oviposition and larval development. Field observations suggest that
only puffballs near or at full maturity contain larval Pocadius.
Members of gasteromycetes are globally distributed, in particular Lycoperdon,
Calvatia, Scleroderma, and Geastrum, which have been recorded in a separate database
as the favored adult and larval fungal host (i.e. ~90% of all Pocadius recorded with host
fungal data). These gasteromycetes appear to be able to resist desiccation and elevational
atmospheric conditions, and are found in most environments except xeric conditions.
This ubiquity and resistance to desiccation is likely the cause for the success of Pocadius
worldwide. Schubert (1988) suggested that northern South America and the Antilles
were much more arid during the Tertiary period than present day conditions suggest. The
spread of Pocadius into these regions coincides with this dry time and their association
with desiccation resistant fungi, such as those in the gasteromycetes, suggests that
Pocadius was able to survive this period through the utilization of these fungal taxa.
Pocadius africanus and P. peruensis, as reported here, are the first records of the genus
on members of Geasteraceae.
Pocadius larvae are known to survive in lab culture for approximately 6-8 weeks
(Gillogly unpub. data), but typically takes less than 3 weeks to finish their development.
The larvae leave the host fungus through an exit hole where they pupate in the
surrounding leaf litter or soil. Pupation time is unknown; however, I predict this to be a
plastic time interval that is influenced strongly by external environmental factors,
341
especially rainfall because this will trigger the development of fungal hosts. Larvae of P.
jelineki were studied in the field and an interesting developmental pattern was exhibited
by the different larval stages in a species of Lycoperdon. As the larvae matured, the older
more advanced stages migrated away from the hymenium towards the inside of the
fungus near the developing spores while the younger less developed stages were more
embedded within the hymenium and absent from the internal cavity where spore
development was taking place. This may represent a division of the fungal resource such
that competition between early larval instars is lessened. Larvae of P. helvolus from
Louisiana also exhibited this peculiar life history trait in Lycoperdon pyriforme.
Hibbett et al. (1997) performed a phylogenetic analysis to understand the
evolutionary relationships of gilled mushrooms and puffballs. Their analysis suggested a
polyphyletic gasteromycetes with puffballs and other forms with an enclosed spore-
bearing structure independently evolving at least four times. The genera Lycoperdon and
Calvatia group together and are sister to Scleorderma, a boletoid member according to
Binder and Bresinsky (2002). Geastrum, the newly recorded host fungus from this study,
was shown not to be closely related to any of the three genera listed above (see Hibbett et
al. 1997). Thus, fungal host use in Pocadius does not appear to have any phylogenetic
significance and the use of varied fungal genera may be do in part to their convergence
on body forms with an enclosed spore-bearing structure that is likely a safe refugia for
larval development.
PHENOLOGICAL CONSIDERATIONS
Species in sympatry may be temporally distributed such that there is no seasonal
overlap of adults and thus reproductive isolation can be maintained. Most species of
Pocadius that are sympatric do not exhibit temporal displacement. However, some
342
exceptions to this can be found, though caution must be considered as few specimens
from some regions are known and temporal data may be due to collecting bias (supported
by the fact that widespread common species have broad temporal occurrences, whereas
more endemic and/or isolated species have a shorter temporal occurrence). The
European species, P. adustus and P. ferrugineus, overlap in all months except early in the
season when only P. adustus is known from March and April with P. ferrugineus not
occurring until May. In Vietnam, P. decoratus occurs in September whereas P. femoralis
occurs four months earlier in May. In the Southwestern United States, P. basalis and P.
niger overlap in all months except P. basalis occurs in the months of June and July prior
to the occurrence of P. basalis in August. The Brazilian fauna appears to have two pairs
of species that coexist at two different time periods; P. antennuliferus and P. bicolor both
occur in March, whereas P. coxus and P. fumatus occur earlier in December and January.
As more specimens are collected the temporal reproductive boundaries will undoubtedly
become more defined.
FUTURE/CONTINUING RESEARCH
The research described herein, as well as published works (Cline 2003b, 2004b,
Cline and Carlton 2004a, 2004b, Ewing and Cline 2004 and Ewing and Cline 2005),
unpublished works (Nitidulidae and Kateretidae of Nova Scotia and Prince Edward
Island, Majka and Cline 2005 submitted; Phalacridae and Corylophidae of Nova Scotia
and Prince Edward Island, Majka and Cline in prep.; review of North American
Kateretidae Cline in prep.; and revision of Smicripidae, Cline in prep.), and study at
numerous museums and institutes will help make advancements in Cucujoidea taxonomy
and systematics possible. A catalogue of Nearctic Nitidulidae and Kateretidae is
currently underway, in collaboration with C. Ewing. Several new genera and numerous
343
new species of Nitidulinae have been identified from the New and Old World tropics and
will be described in the near future. Also, faunistic and ecological studies in the
Neotropics through the IBISCA (Investigations in the BIodiversity of Soil and Canopy
Arthropods) working group will help study large scale phenomena via nitidulid beetle
diversity patterns. A future revision of the genus Stelidota will help shed light on its
biology and classification, as well as the efficacy for which nitidulid species can be used
in biodiversity and conservation studies (see Anderson and Ashe 2000 for a commentary
on using soil beetles in conservation practices and Didham et al. 1998 on the use of soil-
inhabiting beetles in addressing biodiversity responses with respect to forest
fragmentation).
An ongoing project with Michael Whiting is underway to test the status and
monophyly of nitidulid subfamilies, which has never been rigorously tested and will
likely indicate that the Nitidulinae is paraphyletic, the Cybocephalinae will be separated
as a distinct family (especially in light of the numerous larval and adult apomorphies now
evident, see Lawrence et al. 1999a and 1999b for a more or less comprehensive list), and
the Maynipeplinae will likely be a basal member of the Cillaeinae.
Larval studies in Cucujoidea were also initiated during this research and will be
continued for Pocadius and expanded into other Nitidulinae. A previously unknown
larval form of Macrostola was identified in collaboration with a Neotropical pollination
study (see Garcia-Robledo 2004). This is the first-ever known larva of the genus
Macrostola, and one of only a very few larvae known for Neotropical Cillaeinae taxa
(Cline and Carlton unpublished data). The larval nitidulid holdings at the California State
Collection of Arthropods, where Cline currently is employed, are among the best in the
nation due to the efforts of Iris Savage whose illustrations and specimens have been given
344
to Cline to further develop and publish. Larval characters remain enigmatic for many
beetle taxa, but which provide a wealth of new character systems that are valuable for
understanding phylogenetic relationships.
Undoubtedly a solid foundation of Coleoptera taxonomy and systematics was
established during the course of this dissertation and will provide a lifetime of
opportunities to pursue research on little known groups of beetles. With fewer and fewer
traditional beetle taxonomists being educated and trained, the role of a morphological
taxonomist and systematist will be continually sought for biodiversity, ecological,
systematic, and conservation efforts. Thus, the research undertaken herein is not
considered a single self-contained study, but rather an ongoing program that will sustain
innumerable future endeavors.
345
REFERENCES CITED
Anderson, R.S. and J.S. Ashe. 2000. Leaf litter inhabiting beetles as surrogates for
establishing priorities for conservation of selected montane cloud forests in Honduras,
Central America (Coleoptera: Staphylinidae, Curculionidae). Biodiversity and
Conservation 9: 617-653.
Andrews, F.G. 2002. Latridiidae Erichson 1842 [pp. 395-398]. In: American Beetles. Volume
2. Polyphaga: Scarabaeoidea through Curculionoidea (R.H. Arnett, M.C. Thomas, P.E.
Skelley, and J. Howard Frank, editors.). CRC Press, Boca Raton. 861pp.
Arnettt, R.H. Jr. 1963. The beetles of the United States: A manual for identification. The
American Entomological Institute. Ann Arbor, Michigan. xii + 1112pp.
Audisio. P. 1979. The Nearctic species of the genus Cateretes (Coleoptera: Nitidulidae). The
Coleopterists Bulletin 33: 48.
Audisio, P. 1980. Magyarország Allatvilága (Hungariae Fauna), VIII. Kötet, Coleoptera III, 9
Füzet: Nitidulidae. Hungary Fauna. 140, pp. 171 + 6, Akadémiai Kiadò, Budapest.
Audisio, P. 1981. The Nitidulidae (Coeloptera) of the Hortobagy National Park. The Fauna of
the Hortobagy National Park, 1981, Akadamie Kiadò 135-138.
Audisio, P. 1984a. Necessità di ridefinizione delle sottofamiglie nei Nitidulidae a nuove
prospettive per la ricostruzione filogenetica del gruppo (Coleoptera). Bolletino di
Zoologia 51, Supplement 5.
Audisio, P. 1987. The Nitidulidae (Coeloptera) fauna of the Kiskunsag National Park. The
Fauna of the Kiskunsag National Park, 1987, Akadamie Kiadò, Budapest. 189-192.
Audisio, P. 1989. Notes on the genus Brachyleptus Motschulsky (Coeloptera: Kateretidae).
Folia Entomologica Hungarica. 50: 9-14.
Audisio, P. 1993. Coleoptera, Nitidulidae – Kateridae. Fauna d’Italia. Vol. 32. Edizioni
Calderini: Bologna. 971pp.
Audisio, P. 1994. Brachypterinae Erichson, 1845 (Insecta, Coleoptera) and Brachypterinae
Zwick, 1973 (Insecta, Plecoptera): proposed removal of homonymy. Bulletin of
Zoological Nomenclature 51: 309-311.
Audisio, P. 1995. Comments on the proposal to remove homonymy between
Brachypterinae Erichson, [1845] (Insecta, Coleoptera) and Brachypterinae Zwick,
1973 (Insecta, Plecoptera), and proposed precedence of Kateretidae Ganglbauer, 1899
over Brachypterinae Erichson, [1845]. Bulletin of Zoological Nomenclature 52: 179-
181.
Audisio, P. 1996. Kateretidae and Nitidulidae (Coleoptera) from the Bükk National Park. The
Fauna of the Bükk National Park, 1996, Akadamie Kiadò, Budapest 293-298.
346
Audisio, P., M.C. Angelici, and V. Sbordoni. 1984. Studio sistematico su Meligethes exilis
Sturm, in base a dati elettroforetici, morfo-ecologici e biogeografici (Coleoptera:
Nitidulidae). Fragmenta Enomologica 17: 359-372.
Audisio, P., A. DeBiase, G. Antonini, C. Belfiore, and M. Oliverio. 2001. Morphological,
molecular and ecological evidence of a new Euro-Anatolian species of the Meligethes
coracinus complex (Coleoptera: Nitidulidae). Insect Systematics and Evolution 31:
361-385.
Audisio, P., A. DeBiase, G. Antonini, M. Oliverio, M. Ketmaier, and E. DeMatthews.
2002. Specific distinction by allozymic data of sympatric sibling specie of the pollen-
beetle genus Meligethes (Coleoptera: Nitidulidae). Italian Journal of Zoology 69: 65-
69.
Audisio P., A. DeBiase, P. Romanelli, M.C. Angelici, V. Ketmaier, & D. De Matthaeis.
2000. Molecular re-examination of the taxonomy of the Meligethes viridescens species
complex (Coleoptera : Nitidulidae). Biochemical Systematics & Ecology 28:1-13.
Audisio, P. and J. Jelínek. 1993. Two new genera of Nitidulidae from the Oriental region,
with notes on phylogeny of the "Axyroid-group" (Coleoptera, Nitidulidae, Nitidulinae).
Revue Suisse de Zoologie 100: 405-423.
Audisio, P., J. Jelínek, A. Mariotti, and A. DeBiase. 2000. The Coleoptera and Kateretidae
from Anatolian, Cuacasian, and Middle East Regions. Biogeographia 21: 241-354.
Audisio, P. and A.G. Kirejtshuk. 1983. Revision of the genera Ithyra Reitter and Neothalycra
Grouvelle (Coleoptera: Nitidulidae). Revue de Zoologie africaine 97: 365-378.
Beutel, R.G. and S.A. Ślipiński. 2001. Comparative study of head structures of larvae of
Sphindiddae and Protocucujidae (Coloeptera: Cucujoidea). European Journal of
Entomology 98: 219-232.
Binder, M. and A. Bresinsky. 2002. Derivation of a polymorphic lineage of Gasteromycetes
from boletoid ancestors. Mycologia 94: 85-98.
Blackwelder, R.E. 1945. Checklist of the coleopterous insects of Mexico, Central America, the
West Indies, and South America. Bulletin of the United States National Museum, part 3,
no. 185, iv + 207 pp.
Blatchley, W.S. 1910. The Coleoptera or Beetles Known to Occur in Indiana. The Nature
Publishing Co., Indianapolis, IN. 1386pp.
Blumberg, D. 1973. Survey and distribution of Cybocephalidae (Coleoptera) in Israel.
Entomophaga 18: 125-131.
347
Blumberg, D. and E. Swirski. 1982. Comparative biological studies on two species of
predatory beetles of the genus Cybocephalus (Coleoptera: Cybocephalidae).
Entomophaga 27: 67-76.
Böving, A.G. and F.C. Craighead. 1931. An illustrated synopsis of the principal larval forms
of the order Coleoptera. Entomologica Americana 11: 1-351.
Böving, A.G. and J.G. Rozen. 1962. Anatomical and systematic study of the mature larvae of
the Nitidulidae (Coleoptera). Entomologiske Meddelelser 31: 265-299.
Bousquet, Y. 2002. Monotomidae Laporte 1840 [pp. 319-321]. In: American Beetles. Volume
2. Polyphaga: Scarabaeoidea through Curculionoidea (R.H. Arnett, M.C. Thomas, P.E.
Skelley, and J. Howard Frank, editors.). CRC Press, Boca Raton. 861pp.
Bowestead, S. and R.A.B. Leschen. 2002. Corylophidae LeConte 1852 [pp. 390-394]. In:
American Beetles. Volume 2. Polyphaga: Scarabaeoidea through Curculionoidea (R.H.
Arnett, M.C. Thomas, P.E. Skelley, and J. Howard Frank, editors.). CRC Press, Boca
Raton. 861pp.
Bronstein, J.L. and Y. Ziv. 1997. Costs of two non-mutualistic species in a yucca/yucca moth
mutualism. Oecologia 112: 379-385.
Browne, D.J., Peck, S.B., and Ivie, M.A. 1993. The longhorn beetles (Coleoptera,
Cerambycidae) of the Bahama Islands with an analysis of species-area relationship,
distribution patterns, origin of the fauna and an annotated species list. Tropical Zoology
6:.27-53.
Bruck, D.J. and L.C. Lewis. 2002. Carpophilus freemani (Coleoptera: Nitidulidae) as a vector
of Beauveria bassiana. Journal of Invertebrate Pathology 80: 188-190.
Burkhardt, D. and S.A. Slipinski. 1991. A review of the Passandridae of the world
(Coleoptera: Cucujoidea). III. Genera Anisocerus, Aulonosoma, Passandrella,
Passandrina, Scalidiopsis, and Taphroscelidia. Revue Suisse Zoologie 98: 453-497.
Burkhardt, D. and S.A. Slipinski. 1995. A review of the Passandridae of the world
(Coleoptera: Cucujoidea). IV. Revue Suisse Zoologie 102: 995-1044.
Casey, T. L. 1916. Some random studies among Clavicornia. Memoirs on the Coleoptera 7:
35-292.
Chiao, E. and J.V. McHugh. 2000. Larval Sphindidae (Coleoptera: Cucujoidea): Phylogenetic
implications and new descriptions. Invertebrate Taxonomy 14: 807-824.
Chujo, M. 1992. Nitidulidae from Chejudo Island. Esakia 32: 19-24.
Chujo, M. 1994. Nitidulidae from Korea (Coleoptera, Insecta). Esakia 34: 195-202.
348
Cline, A.R. 2003a. New distribution records for Pocadius basalis Schaeffer (Coleoptera:
Nitidulidae: Nitidulinae) from Southwestern United States. The Coleopterists Bulletin
57: 390.
Cline, A.R. 2003b. A new sap beetle (Coleoptera: Nitidulidae) to the United States with
a revised key to the Camptodes Erichson occurring in America north of Mexico. Insecta
Mundi 17: 101-102.
Cline, A.R. 2004a. New state records for two species of Thalycra Erichson (Coeloptera:
Nitidulidae) with notes on species sympatry. The Coleopterists Bulletin 58: 137-138.
Cline, A.R. 2004b. A New Species of Psilotus Fischer vonWaldheim (Coleoptera: Nitidulidae:
Nitidulinae) from Peru, with new distribution records for other Psilotus species.
Proceedings of the Entomological Society of Washington 106(4): 890-898.
Cline, A.R. and C.E. Carlton. 2004a. Two new species of Epuraea (Orthopeplus)
(Coleoptera: Nitidulidae) from Mexico. The Coleopterists Bulletin 58: 261-270.
Cline, A.R. and C.E. Carlton. 2004b. Review of Lasiodactylus Perty with Descriptions
of Three New Species (Coleoptera: Nitidulidae: Nitidulinae) The Coleopterists Bulletin
58(3): 355-368.
Cline, A.R. and R.A.B. Leschen. 2005. Coleoptera associated with the oyster mushroom,
Pleurotus ostreatus Fries, in America north of Mexico. Southeastern Naturalist 4(3):
409-420.
Connell, W.A. 1956. Nitidulidae of Delaware. Delaware Agricultural Experiment Station
Bulletin 318: 1-67.
Connell, W.A. 1981. Bibliography of Carpophilus humeralis (Fab.) in support of a revision of
the genus Carpophilus Stephens. Bulletin of the Entomological Society of America 27:
263-266.
Cooper, M.C. 1982. The species of the genus Pria Stephens (Coeloptera: Nitidulidae).
Zoological Journal of the Linnaean Society 75: 327-390.
Crowson, R.A. 1954. The classification of the families of British Coleoptera. The
Entomologist’s Monthly Magazine 90: 57-63.
Crowson, R.A. 1955. The Natural Classification of the Families of Coleoptera. Nathaniel
Lloyd and Co., London, 187 pp.
Crowson, R.A. 1960. The phylogeny of Coleoptera. Annual Review of Entomology 5:111-94.
Crowson, R.A. 1964a. A review of the classification of Cleroidea (Coeloptera), with
description of two new genera of Peltidae and of several new larval types. Transactions
of the Royal Entomological Society of London (B) 116: 275-327.
349
Crowson, R.A. 1964b. A new genus of Australian clavicorn Coleoptera, probably a new
family. Proceedings of the Linnean Society of New South Wales 89: 241-245.
Crowson, R.A. 1966. Further observations on Peltidae (Coleoptera: Cleroidea), with
definitions of a new subfamily and of four new genera. Proceedings of the royal
Entomological Society of London (B) 35: 119-127.
Crowson, R.A. 1970. Further observation on Cleroidea (Coleoptera). Proceedings of the Royal
Entomological Society of London (B) 39: 1-20.
Crowson, R.A. 1973. Further observations on Phloeostichidae and Cavognathidae, with
definitions of new genera from Australia and New Zealand. Coleopterists Bulletin 27:
54-62.
Crowson, R.A. 1981. The Biology of the Coleoptera. Academic Press, London. 802pp.
Crowson, R.A. 1990. A new genus of Boganiidae (Coleoptera) from Australia, with
observations on glandular openings, cycad associations and geographical distribution in
the family. Journal of the Australian Entomological Society 29: 91-99.
Crowson, R.A. and T. SenGupta. 1969. The systematic position of Propalticidae and of
Carinophloeus Lefkovitch (Coleoptera: Clavicornia) with description of a new species of
Propalticus and of its supposed larva. Proceedings of the Royal Entomological Society
of London (B) 38: 132-140.
Currie, C.R., J.R. Spence, and W.J.A. Volney. 1996. Biology and life history of Epuraea
obliquus Hatch (Coleoptera: Nitidulidae) on western gall rust. The Canadian
Entomologist 128: 177-186.
Davis, A.L.V.; Scholtz, C.H.; Philips, T.K. 2002. Historical biogeography of scarabaeine
dung beetles. Journal of Biogeography 29: 1217-1256.
DeBiase, A., G. Antonini, E. Mancini, and P. Audisio. 2003. Molecular taxonomy of two
sympatric sibling species of the pollen-beetle genus Meligethes (Coeloptera: Nitidulidae).
Zootaxa 190: 1-16.
Didham, R.K., P.M. Hammond, J.H. Lawton, P. Eggleton, and N.E. Stork. 1998. Beetle
species responses to tropical forest fragmentation. Ecological Monographs 68: 295-323.
Donisthorpe, H. 1935. The British fungicolous Coleoptera. Entomologists Monthly Magazine
71: 21-31.
Donnelly. T.W. 1988. Geological Constraints on Caribbean Biogeography, In: Zoogeography
of Caribbean Insects (J.K. Liebherr Editor). Cornell University Press, Ithaca, NY. 285pp.
350
Dorsey, C.K. and J.G. Leach. 1956. The bionomics of certain insects associated with Oak
Wilt with particular reference to the Nitidulidae. Journal of Economic Entomology 49:
219-230.
Downie, N.M. and R.H. Arnettt. 1996. The Beetles of Northeastern North America. Volume
II. The Sandhill Crane Press, Gainesville, FL. 1721pp.
Drea, J.J. and R.W. Carlson. 1988. Establishment of Cybocephalus sp. (Coleoptera:
Nitidulidae) from Korea on Unaspis euonymi (Homoptera: Diaspididae) in the eastern
United States. Proceedings of the Entomological Society of Washington 90: 307-309.
Endrödy-Younga, S. 1968. Monographie der Paläarktischen arten der familie Cybocephalidae
(Coleoptera: Clavicornia). Acta Zoologica Academiae Scientarium Hungaricae 14: 27-
115.
Endrödy-Younga, S. 1978. Systematic revision and phylogeny of some Meligethinae genera
from the Ethiopian Region (Coleoptera: Nitidulidae). Entomologica Germanica 4: 295-
316.
Endrödy-Younga, S. 1982. Cybocephalids of Réunion and Mauritius Islands (Coleoptera:
Cybocephalidae). Annals of the Transvaal Museum 33: 259-264.
Endrödy-Younga, S. 1984. A new species of Cybocephalus (Coeloptera: Cucujoidea:
Cybocephalidae) from Israel. Israel Journal of Entomology 18: 1-2.
Endrödy-Younga, S. 1991. Boganiidae (Coeloptera: Cucujoidea) associated with cycads in
South Africa: two new species and a new synonym. Annals of the Transvaal Museum
35: 285-292.
Erichson, W.F. 1843. Versuch einer systematischen Eintheilung der Nitidularian. Zeitschrift
für Entomologie 4: 225-361.
Ewing, C.P. and A.R. Cline. 2004. New records and taxonomic updates for adventive sap
beetles (Coleoptera: Nitidulidae) in Hawaii. Bishop Museum Occasional Papers
(Records for the Hawaii Biological Survey for 2003) 79: 40-45.
Ewing, C.P. and A.R. Cline. 2005. Key to adventive sap beetles (Coleoptera: Nitidulidae) in
Hawaii, with notes on records and habits. The Coleopterists Bulletin 59: 167-183.
Fabricius, J. 1775. Systema entomologiae sistens insectorum classes, ordines, genera, species,
adjectis synonymis, locis, descriptionibus, observationibus. Korte, Flensburgi et Lipsiae.
832pp.
Farris, J.S. 1982. Outgroups and parsimony. Systematic Zoology 31(3): 328-334.
351
Ferrer, J., H. Wendt, and P. Audisio. 2000. Provisional checklist of Scarabaeidae,
Nitidulidae, Tenebrionidae, and Bruchidae (Coloeptera) from the Brandberg Massif,
Namibia. Cimbebasia Memoir 9: 385.
Garcia-Robledo, C. 2004. Beetle pollination and fruit predation of Xanthosoma daguense
(Araceae) in an Andean cloud forest in Colombia. Journal of Tropical Ecology 20: 459-
469.
Gillogly, L.R. 1955. A review of the genus Mystrops Erichson (Coleoptera, Nitidulidae).
Revista Brasileira de Entomologia 3: 191-204.
Gillogly, L.R. 1965. A key to the genera of the subfamily Nitidulinae (Nitidulidae, Coleoptera)
and description of a new genus and a new species. Occasional Papers of the Bureau of
Entomology California Department of Agriculture 8: 1-24.
Gillogly, L.R. 1969. Nitidulidae (Coleoptera) collected by the Noona Dan expedition in the
Philippine and Bismarck Islands. Entomologiske Meddelesler 37: 207-224.
Gillogly, L.R. 1972. A new species of Mystrops from Costa Rica (Coleoptera: Nitidulidae).
The Pan-Pacific Entomologist 48: 116-120.
Goodrich, M.A. 2002a. Byturidae Jacquelin duVal 1858 [pp. 354-355]. In: American Beetles.
Volume 2. Polyphaga: Scarabaeoidea through Curculionoidea (R.H. Arnett, M.C.
Thomas, P.E. Skelley, and J. Howard Frank, editors.). CRC Press, Boca Raton. 861pp.
Goodrich, M.A. 2002a. Biphyllidae LeConte 1861 [pp. 356-357]. In: American Beetles.
Volume 2. Polyphaga: Scarabaeoidea through Curculionoidea (R.H. Arnett, M.C.
Thomas, P.E. Skelley, and J. Howard Frank, editors.). CRC Press, Boca Raton. 861pp.
Grouvelle, A. 1894. Insectes du Bengale: Clavicornes. Annales de la Société Entomologique
de Belgique 20:578-587.
Grouvelle, A. 1896. Descriptions de Clavicornes D’Afrique et de Madagascar. Annales de la
Societe Entomologique de France 65: 71-94.
Grouvelle, A. 1897. Clavicornes Nouveaux des Indes Orientales et Pays Voisins. Annali del
Museo Civico di Storia Naturale di Genova (Nitidulidae part) 18: 342-374.
Grouvelle, A. 1898. Clavicornes Nouveaux d’Amerique. 2nd memoire. Annales de la Societe
Entomologique de France 67: 344-381.
Grouvelle, A. 1899a. Nitidulides de L’Afrique Occidentale (Cameroun). Annales de la Société
Entomologique de France 68:125-135.
Grouvelle, A. 1899b. Descriptions de clavicornes D’Afrique et de la region Malgache.
Annales de la Société Entomologique de France 68:136-185.
352
Grouvelle, A. 1899c. Clavicornes Nouveaux. Annales de la Société Entomologique de
Belgique 89: 299-301.
Grouvelle, A. 1901. Supplément a la liste des Coléoptéres de la Guadeloupe. Extrait des
Annales de la Société Entomologique de France 71:756-758.
Grouvelle, A. 1905a. Clavicornes Nouveaux du Musée Civique de Génes. Annali del Museo
Civico di Storia Naturale di Genova 2: 308-333.
Grouvelle, A. 1905b. Quelques clavicorns nouveaux de la République Argentine recueilles par
M. Charles Bruch. De la Revista del Museo de La Plata 12: 121-133.
Grouvelle, A. 1906. Nitidulides nouveaux du British Museum (Coleoptera). Bulletin de la
Societe Entomologique de France 201-215.
Grouvelle, A. 1908a. Coleopteres de la Region Indienne: Rhysodidae, Trogositidae,
Nitidulidae, Colydiidae, Cucujidae. Extrait des Annales de la Societe Entomologique de
France (Nitidulidae part) 6: 323-397.
Grouvelle, A. 1908b. Supplément aux Coléoptères de la Gaudeloupe. Extrait des Annales de la
Société Entomologique de France 77: 41-42.
Grouvelle, A. 1910. Nitidulides et Cryptophagides de L’Asie et des Indes Orientales. Notes
from the Leyden Museum 32: 241-256.
Grouvelle, A. 1913. Byturidae, Nitidulidae pars 56 [pp. 1-223]. In: Coleopterorum Catalogus
(W. Junk and S. Schenkling, Editors). Berlin.
Grouvelle, A. 1914a. Nitidulidae des Philippines Recoltés Par C.F. Baker. The Philippine
Journal of Science 9: 535-542.
Grouvelle, A. 1914b. Descriptions de Coléoptères Africains. Annales de la Société
Entomologique de France 83: 141-144.
Grouvelle, A. 1915. Clavicornes Africans du Musée D’Historie Naturelle de Luxembourg
recoltés par M. Ed. Luja de Luxembourg. Gesselschaft Luxembourger Naturfreunde 25:
103-123.
Grouvelle, A. 1916. Description des Clavicornes Nouveaux de la République Argentine. De la
Revista del Museo de La Plata 23: 234-247.
Grouvelle, A. 1919. Descriptions de Coléoptères de L’Afrique australe. Memoirs de
Entomologique de Paris 47-61.
Habeck, D. 2002a. Nitidulidae Latrielle 1802 [pp.311-315]. In: American Beetles. Volume 2.
Polyphaga: Scarabaeoidea through Curculionoidea (R.H. Arnett, M.C. Thomas, P.E.
Skelley, and J. Howard Frank, editors.). CRC Press, Boca Raton. 861pp.
353
Habeck, D. 2002b. Brachypteridae Erichson 1845 [pp.309-310]. In: American Beetles. Volume
2. Polyphaga: Scarabaeoidea through Curculionoidea (R.H. Arnett, M.C. Thomas, P.E.
Skelley, and J. Howard Frank, editors.). CRC Press, Boca Raton. 861pp.
Hayashi, N. 1978. A contribution to the knowledge of the larvae of Nitidulidae occurring in
Japan (Coleoptera, Cucujidea). Insecta Matsumurana 14: 1-98.
Heer, O. 1841. Fauna Coleopterum Helvetica, pars 1, fascicle 3, pp. 361-652. Turici (Zurigo).
Hennig, W. 1965. Phylogenetic Systematics. Annual Review of Entomology 10: 97-116.
Hennig, W. 1966. Phylogenetic Systematics. University of Illinois Press, Urbana. 263pp.
Hetschko, A. 1930. Pars 109. Cucujidae, Thorictidae (suppl.), Cossyphodidae (suppl.) [pp. 1-
122]. In: Coleopterum Catalogus (S. Schenkling, editors). Berlin.
Hibbett, D.S., E.M. Pine, E. Langer, and M.J. Donoghue. 1997. Evolution of gilled
mushrooms and puffballs inferred from ribosomal DNA sequences. Proceedings of the
National Academy of Sciences USA 94: 12002-12006.
Hisamatsu, S. 1958. A revision of the Japanese Cychramus (Coleoptera: Nitidulidae). New
Entomologist 7: 7-11.
Hisamatsu, S. 1976. A new genus of Cateretinae from Taiwan (Coleoptera: Nitidulidae).
Transactions of the Shikoku Entomological Society 13: 19-23.
Hood, W.M. 2000. Overview of the small hive beetle, Aethina tumida, in North America. Bee
World 81: 129-137.
Hölldobler, B. and E.O. Wilson. 1990. The Ants. Belknap Press of Harvard University Press,
Cambridge, MA. 732pp.
Horn, G.H. 1879. Revision of the Nitidulidae of the United States. Transactions of the
American Entomological Society 7: 267-336.
Howden, H.F. 1961. A revision of the New World species of Thalycra Erichson, with a
description of a new genus and notes on generic synonymy (Coleoptera: Nitidulidae).
The Canadian Entomologist, Supplement 25. 61pp.
Huth, C.J. and O. Pellmyr. 1997. Non-random fruit retention in Yucca filamentosa:
consequences for an obligate mutualism. Oikos 78: 576-584.
Jacquelin DuVal, C. 1858. Manuel Entomologique. Genera des Coléoptères d’Europe. Paris,
Vol. 2. 1-285.
354
Jelínek, J. 1964. Nitidulidae of the Klapperich’s Expedition in Afghanistan. Annotationes
Zoologicae et Botanicae 8: 1-5.
Jelínek, J. 1965a. Ergebnisse der Albanien-Expedition 1961 des Deutschen Entomologischen
Institutes. Beiträge zur Entomologie 15: 673-688.
Jelínek, J. 1965b. Ergebnisse der zoologischen Forschungen von Dr. Z. Kaszab in der
Mongolei, 33, Coleoptera: Nitidulidae. Reichenbachia 7(16): 135-145.
Jelínek, J. 1966. Ergebnisse der zoologischen Forschungen von Dr. Z. Kaszab in der Mongolei,
68, Coleoptera: Nitidulidae II. Reichenbachia 7(33): 291-294.
Jelínek, J. 1967a. Beiträge zur Kenntnis der Fauna Afghanistans, Nitidulidae: Coleoptera.
Acta Musei Moraviae, Supplement 52 147-150.
Jelínek, J. 1967b. 30th Result of the zoological expedition of the National Museum in Prague to
Turkey. Acta Entomologica Musei Nationalis Pragae 37: 23-30.
Jelínek, J. 1969. Drei neue arten der gattung Mystrops Erichson (Coleoptera: Nitidulidae).
Acta Entomologica Bohemoslovaca 66: 366-372.
Jelínek, J. 1974. Generic reclassification of oriental Cryptarchinae (Coleoptera, Nitidulidae).
Acta Entomologica Bohemoslovoca 71: 187-196.
Jelínek, J. 1975. Redescriptions of genera Hebascus Erichson and Teichostethus Sharp with
designations of their type-species (Coleoptera: Nitidulidae). Annotationes Zoologicae et
Botanicae 101:1-12.
Jelínek, J. 1976. Description and revision of the genus Anamartus gen. n. (Coeloptera:
Nitidulidae). Acta Entomologica Bohemoslovaca 73: 17-31.
Jelínek, J. 1977a. Revision of South American species of the genus Pocadius Er. with
description of new genus (Coleoptera, Nitidulidae). Acta Entomologica Musei Nationalis
Pragae 39: 29-44.
Jelínek, J. 1977b. Revision of the genus Epuraea Erichson from Africa with remarks to related
genera (Coloeoptera, Nitidulidae). Acta Entomologica Musei Nationalis Pragae 39:
345-397.
Jelínek, J. 1978. Ergebnisse der Bhutan-Expedition 1972 des Naturhistorischen Museums in
Basel. Entomologica Basiliensia 3: 171-218.
Jelínek, J. 1979a. A new genus of Neotropical Cateretinae (Coleoptera, Nitidulidae). Acta
Entomologica Bohemoslovaca 76: 188-202.
Jelínek, J. 1979b. Insects of Saudi Arabia, Coleoptera: Fam. Nitidulidae. Fauna of Saudi
Arabia 1: 223-227.
355
Jelínek, J. 1981a. Review of the genus Anister (Coleoptera, Nitidulidae). Acta Entomologica
Bohemoslovaca 78: 183-188.
Jelínek, J. 1981b. Results of the Czechoslovak-Iranian entomological expeditions to Iran 1970
and 1973, Coleoptera: Nitidulidae. Acta Entomologica Musei Nationalis Pragae 40:
105-119.
Jelínek, J. 1982. New and little known taxa of Nitidulidae (Coleoptera). Acta Musei nationalis
Pragae 38: 171-200.
Jelinek, J. 1984. Revision of the genus Stelidota from Asia, Australia, and Pacific area
(Coleoptera: Nitidulidae). Acta Entomologica Bohemoslovaca 81: 132-156.
Jelínek, J. 1988. Coleoptera: Nitidulidae of Saudi Arabia (Part 2). Fauna of Saudi Arabia 9:
42-51.
Jelínek, J. 1996. Coleoptera: Cucujoidea 1 (Kateretidae, Nitidulidae, Rhizophagidae, and
Sphindidae) [pp. 485-493]. In: Terrestrial Invertebrates of the Pálava Biosphere Reserve
of UNESCO, III (R. Rozkošný and J. Vaňhara, Editors). Folia Fac. Nat. Univ.
Masarykianae Brunensis, Bilogia. Volume 94.
Jelínek, J. 1999a. Contribution to taxonomy of the beetle subfamily Nitidulinae (Coleoptera:
Nitidulidae). Folia Heyrovskayana 7: 251-281.
Jelínek, J. 1999b. Brachypteridae and Nitidulidae (Coeloptera) from the Aggtelek National
Park. The Fauna of the Aggtelek National Park, 1999. 233-238.
Jelínek, J. and P. Audisio. 2003. Type fixations and nomenclatural corrections in some taxa of
Palearctic Nitidulidae and Keteretidae (Coleoptera). Folia Heyrovskyana 11: 159-171.
Juzwik, J. 1986. Relationship between nitidulids and Ceratocystis fagacearum during late
summer and autumn in Minnesota. Plant Disease 70: 424-426.
Kirejtshuk, A.G. 1979. New species of the Cryptarchopria Jelinek genus (Coleoptera:
Nitidulidae: Meligethinae) from Vietnam and its variability. Proceedings of the
Academy of Sciences of the Ukranian S.S.R. Series B. Geological, Chemical, and
Biological Sciences (5): 383-387.
Kirejtshuk, A.G. 1980. New species of beetles of the subfamily Meligethinae from the
Ethiopian region (Coleoptera: Nitidulidae). Extrait de la Revue de Zoologie Africaine
94: 249-294.
Kirejtshuk, A.G. 1981. Preliminary revision of the Cryptarchinae genera of the afrotropical
region, with descriptions of a new genus, a new subgenus, and some new species. Revue
de Zoologie Africaine 95: 765-805.
356
Kirejtshuk, A.G. 1982. Systematic position of the genus Calonecrus J. Thomson and notes on
the phylogeny of the family Nitidulidae (Coleoptera). Entomologicheskoye Obozrenie
59: 833-851.
Kirejtshuk, A.G. 1984a. New taxa of Nitidulidae (Coleoptera) from the Indo-Malayan fauna.
Annales Historico-Naturales Musei Nationales Hungarici 76: 169-195.
Kirejtshuk, A.G. 1984b. New species of beetles of the families Nitidulidae and
Cybocephalidae (Coleoptera) in the East Palearctic fauna. Zoologicheskij Zhurnal
(Moscow) 63: 517-533.
Kirejtshuk, A.G. 1985. New species of Cyllodes Erichson and Viettherchnus gen. n.
(Coleoptera: Nitidulidae) of the fauna of Vietnam and adjacent territories. Nasekomye
Vietname 157-164. (In Russian)
Kirejtshuk, A.G. 1986a. On the polyphyly of the Carpophilinae with description of a new
subfamily, Cillaeinae (Coleoptera: Nitidulidae). The Coleopterists Bulletin 40: 217-221.
Kirejtshuk, A.G. 1986b. New genera and species of (Nitidulidae, Coleoptera) from the
Australian region. I. Entomologicheskoye Obozreniye 3: 559-573. (In Russian)
Kirejtshuk, A.G. 1986c. Analysis of genitalia structure for the phylogeny reconstruction and
supporting the system of the family Nitidulidae (Coleoptera). Trudy VEO [Proceedings
of the All-Union Entomological Scoiety] 68: 22-28. (in Russian)
Kirejtshuk, A.G. 1986d. Revision of the genus Aethina Erichson (Coleoptera, Nitidulidae)
from the Oriental and Palearctic region. USSR Academy of Sciences, Proceedings of the
Zoological Institute in Leningrad 140: 44-82.
Kirejtshuk, A.G. 1987. New species in the Cyllodes generic complex (Coeloptera: Nitidulidae)
from Indochina and neighboring areas. Entomofauna Vietnama Nauka Moscow 137-
172.
Kirejtshuk, A.G. 1988a. New genera and species of (Nitidulidae, Coleoptera) from the
Australian region. II. Entomological Review 67:129-156. (Translated from
Entomologicheskoye Obozreniye. 1987. 4:773-799. In Russian.)
Kirejtshuk, A.G. 1988b. New Palearctic genus and species of the family Kateretidae
(Coleoptera) and notes on the synonomy. Zoologigicheskii Zhurnal 68: 145-149.
Kirejtshuk, A.G. 1989. New taxa of the Nitidulidae (Coleoptera) of the East Hemisphaera
(Part III). USSR Academy of Sciences. Proceedings of the Zoological Institute,
Leningrad 208: 64-89.
Kirejtshuk, A.G. 1990a. Revision of the Australian genus Idaethina Reitter (Coleoptera:
Nitidulidae). Journal of the Australian Entomological Society 29:1-9.
357
Kirejtshuk, A.G. 1990b. New genera and species of the Nitidulidae beetles (Coleoptera:
Nitidulidae) from Australian region. III. Revue d’Entomologie de l’URSS 69: 857-878.
(In Russian)
Kirejtshuk, A.G. 1990c. New species and taxonomic notes on Nitidulidae of Indochina and
adjacent territories. Part 1. USSR Academy of Sciences, Proceedings of the Zoological
Institute in Leningrad 209: 61-98.
Kirejtshuk, A.G. 1990d. New taxa of the Nitidulidae (Coleoptera) of the Eastern Hemisphere.
Part 4. USSR Academy of Sciences, Proceedings of the Zoological Institute in
Leningrad 211: 84-103.
Kirejtshuk, A.G. 1992. Nitidulidae [pp.114-209]. In: Identification key to the insects of the
Far East of the USSR. Volume 3. Part 2. (P.A. Lehr, editor). Nauka Pren, St. Petersburg,
Russia.
Kirejtshuk, A.G. 1993. On old and new South African Meligethes species (Coeloptera:
Nitidulidae). Mitteilungen Müenchener Entomologischen Gesellschaft 83: 47-75.
Kirejtshuk, A.G. 1994a. Revision of the genus Neopallodes Reitter 1884 (Coleoptera:
Nitidulidae) from the Palearctic and Indo-Malayan regions. Tropical Zoology 7: 225-
253.
Kirejtshuk, A.G. 1994b. New taxa of the Nitidulidae (Coleoptera) of the Eastern Hemisphere.
Part 5. USSR Academy of Sciences, Proceedings of the Zoological Institute in
Leningrad 258: 3-49.
Kirejtshuk, A.G. 1995. System, evolution of the way of life, and phylogeny of the order
Coleoptera. I. Entomological Review 74: 12-31.
Kirejtshuk, A.G. 1996. Some results of study on the Nitidulidae from Namibia and adjacent
territories. Part 1. (Coloeptera: Cucujoidea, Nitidulidae). Mitteilungen aus dem
Zoologischen Museum in Berlin 72: 21-52.
Kirejtshuk, A.G. 1997a. New Palearctic nitidulid beetles, with notes on synonomy and
systematic position of some species (Coleoptera: Nitidulidae). Zoosystematica Rossica
6: 255-268.
Kirejtshuk, A. G. 1997b. On the evolution of anthophilous Nitidulidae (Coleoptera) in tropical
and subtropical regions. Bonner Zoologische Beiträge 47: 111-134.
Kirejtshuk, A.G. 1997c. Notes on nitidulid beetles (Coleoptera: Nitidulidae) collected by O.N.
Kabakov in Vietnam and Laos. Archives of the Kharkov Entomological Society 5(2):
13-23. (In Russian)
358
Kirejtshuk, A.G. 1998a. Nitidulidae (Coleoptera) of the Himalayas and northern Indochina,
Part 1: Subfamily Epuraeinae, Theses Zoologicae, Vol. 28. Koeltz Scientific Books,
Koenigstein. 489 pp.
Kirejtshuk, A.G. 1998b. Position of the subfamily Maynipeplinae subfamily n. from equatorial
Africa in the classification of sap-beetles (Coleoptera, Nitidulidae) with notes on the
evolution and structural modifications. Entomologicheskoe Obozrenie 77: 540-554. [In
Russian; translation in Entomological Review 78: 793-807.]
Kirejtshuk, A.G. 2000. On origin and early evolution of the superfamily Cucujoidea
(Coleoptera: Polyphaga): Comments on the family Helotidae. The Kharkov
Entomological Society Gazette 8(1): 8-38.
Kirejtshuk, A.G. 2001. Notes on the systematics of the African Nitidulidae (Coleoptera).
Annales Historico-Naturales Musei Nationalis Hungarici 93: 17-89.
Kirejtshuk, A.G. and P. Audisio. 1995. Preliminary revision of South African Meligethes
Subg. Lariopsis Kirejtshuk (Coleoptera: Nitidulidae: Meligethinae). Fragmenta
Entomologica, Rome 27: 191-254.
Kirejtshuk, A.G., D.G. James, and R. Heffer. 1997. Description and biology of a new species
of Cybocephalus Erichson (Coloeptera: Nitidulidae), a predator of Australian citrus
whitefly. Australian Journal of Entomology 36: 81-86.
Kirejtshuk, A.G. and J. Jelinek. 2000. Preliminary review of genera of the tribe Mystropini
with redescriptions and new descriptions of some genera, subgenera, and species
(Coleoptera: Nitidulidae: Nitidulinae). Folia Heyrovskyana 8: 171-192.
Kirejtshuk, A.G. and A.H. Kirk-Spriggs. 1996. Viettherchnus Kirejtshuk, 1985 and
Ceramphosia gen. n. from the Indo-Malayan region (Coleoptera: Nitidulidae).
Zoosystematica Rossica 4: 131-138.
Kirejtshuk, A.G. and T. Kvamme. 2002. Revision of the subgenus Lasiodites Jelinek, 1999,
stat. nov. of the genus Phenolia Erichson, 1843 from Africa and Madagascar (Coleoptera,
Nitidulidae). Mitteilungen aus dem Museum für Naturkunde in Berlin, Zoologische
Reihe 78: 3-70.
Kirejtshuk, A.G., & J.F. Lawrence. 1992a. Cychramptodini, a new tribe of Nitidulidae
(Coleoptera) from Australia. Journal of the Australian Entomological Society 31: 29-46.
Kirejtshuk, A.G., & J.F. Lawrence. 1992b. Review of the Thalycrodes-complex of genera
(Coleoptera, Nitidulidae), endemic to the Australian region. Journal of the Australian
Entomological Society 31: 119-142.
Kirejtshuk, A.G., & J.F. Lawrence. 1999. Notes on the Aethina complex (Coeloptera:
Nitidulidae: Nitidulinae), with a review of Aehtina (Cleidorura) subgen. nov. and Aethina
(Idaethina) Gemminger et Harold. Annales Zoologici 49: 233-254.
359
Kirejtshuk, A.G. & R.A.B. Leschen. 1998. Review of the Thalycra Complex (Coleoptera:
Nitidulidae: Nitidulinae) with three new genera and notes on mycophagy. Annales
Zoologici 48: 253-273.
Klimasweski, J. and J.C. Watt. 1997. Coleoptera: Family-group review and keys to
identification. Fauna of New Zealand. Vol. 37. Manaaki Press, New Zealand. 199pp.
Kirk-Spriggs, A.H. 1996. Pollen Beetles, Coleoptera: Kateretidae and Nitidulidae:
Meligethinae; In W.R. and R.R. Askew eds. Handbooks for the Identification of British
Insects. Royal Entomological Society, London. 157pp.
Kitching, I.J., P.L. Forey, C.J. Humphries, and D.M. Williams. 2000. Cladistics: The theory
and practice of parsimony analysis. The Systematics Association, Special Issue.
Volume 20. Oxford University Press, Oxford.
Kubisz, D. 1995. Remarks on the Occurrence of Pocadius adustus Reitter in Poland
(Coleotpera: Nitidulidae). Acta Entomologica Silesiana 3: 13-15.
Lachance, M.A., W.T. Starmer, C.A. Rosa, J.M. Bowles, J.S.F. Barker, and D.H. Janzen.
2001. Biogeography of the yeasts of ephemeral flowers and their insects. FEMS Yeast
Research 1: 1-8.
Lameere, A. 1938. Évolution des Coléoptères. Bulletin et Annales de la Société
Entomologique Belgique 78: 355-362.
Latrielle, P.A. 1802. Historie Naturelle, Générale et particuliére des Crustacés et des Insectes.
Families nautrelles des genres. F. Dufart, Paris. Vol.3. 387pp. + 21 plates.
Latrielle, P.A. 1807. Genera Crustaceorum et Insectorum secundum ordinem naturalem in
familias disposita, iconibus exemplisque plurimis explicate. Parisiis, Köng ed. 2. 289pp.
Lawrence, J.F. 1982. Coleoptera [pp. 482-553]. In: Synopsis and Classification of Living
Organisms, Vol. 2. (S. P. Parker, editor). McGraw-Hill, New York.
Lawrence, J.F. 1991. Nitidulidae (Cucujoidea) (including Brachypteridae, Cateretidae,
Cybocephalidae, Smicripidae): sap beetles, dried fruit beetles [pp. 456-460]. In:
Immature Insects, Volume 2 (F. W. Stehr, editor). Kendall Hunt Pub. Co. Dubuque,
Iowa.
Lawrence, J.F. 1995. Two new species of Rhopalobrachium Boheman (Coleoptera:
Phloeostichidae: Hymaeinae) from Australia and Chile [pp. 433-447]. In. Biology,
Phylogeny, and Classification of Coleoptera: Papers celebrating the 80th birthday of Roy
A. Crowson. Volume 1 (I.J. Pakaluk and S.A. Slipinski, editors). 558pp. Muzeum i
Instytut Zoologii PAN: Warszawa. 1092pp.
360
Lawrence, J.F., A.M. Hastings, M.J. Dallwitz, T. Paine, & E.J. Zurcher. 1999a. ‘Beetle
Larvae of the World: Descriptions, Illustrations and Information Retrieval for Families
and Subfamilies. CD-ROM, Version 1.1 for MS-Windows’. CSIRO Publishing,
Melbourne.
Lawrence, J.F., A.M. Hastings, M.J. Dallwitz, T. Paine, and E.J. Zurcher. 1999b. Beetles
of the World: A key and information system for families and subfamilies. CD-ROM,
Version 1.1 for MS-Windows. CSIRO Publishing, Melbourne.
Lawrence, J.F., and A.F. Newton, Jr. 1982. Evolution and Classification of Beetles. Annual
Review of Ecology and Systematics 13: 261-290.
Lawrence, J.F., and A.F. Newton, Jr. 1995. Families and subfamilies of Coleoptera (with
selected genera, notes, references and data on family-group names) [pp. 779–1006]. In.
Biology, Phylogeny, and Classification of Coleoptera: Papers celebrating the 80th
birthday of Roy A. Crowson. Volume 1 (I.J. Pakaluk and S.A. Slipinski, editors). 558pp.
Muzeum i Instytut Zoologii PAN: Warszawa. 1092pp.
LeConte, J.L. 1878. Additional descriptions of new species. Proceedings of the American
Philosophical Society 17: 373-434.
Leng, C.W. 1920. Catalogue of the Coleoptera of America North of Mexico. Cosmos Press,
Cambridge, MA. x + 470pp.
Leschen, R.A.B. 1988. Pallodes austrinus, a new species of Nitidulidae (Nitidulinae) with
discussions of Pallodes mycophagy. Journal of the New York Entomological Society
96: 452-458.
Leschen, R.A.B. 1996. Phylogeny and revision of the genera of Cryptophagidae (Coleoptera:
Cucujoidea). Kansas Science Bulletin 55: 549-634.
Leschen, R.A.B. 1999. Systematics of Convex Nitidulinae (Coleoptera: Nitidulidae):
Phylogenetic Relationships, Convexity, and the Origin of Phallalophagy. Invertebrate
Taxonomy 13: 845-882.
Leschen, R.A.B. 2003. Erotylidae (Insecta: Coleoptera: Cucujoidea): Phylogeny and Review.
In: Fauna of New Zealand. Number 47. (T. K. Crosby, editor). Manaaki Whenua Press,
Lincoln, Canterbury, New Zealand. 107pp.
Leschen, R.A.B. and C.E. Carlton. 1994. Three new species and a new record of neotropical
Pocadius Erichson 1843 (Coleoptera Nitidulidae). Tropical Zoology 7:209-216.
Leschen, R.A.B. and C.E. Carlton. 1996. Slime-production in mycophagous Nitidulidae
(Coleoptera) including a new species of Eusphaerius. Journal of Natural History 30:
1861-1873.
361
Leschen, R.A.B. and C.E. Carlton. 2004. A new tribe, genus and species of nitidulid beetle
(Coleoptera: Nitidulidae: Nitidulinae) from Bolivia. Coleopterists Bulletin 58(3): 443-
451.
Leschen, R.A.B. J.F. Lawrence and S. A. Ślipiński. 2005. Basal classification of Cucujoidea
(Coleotpera: Polyphaga): cladistic analysis, keys and review of new families.
Invertebrate Systematics 19: 17-73.
Leschen, R.A.B. and P.E. Skelley. 2002. Cryptophagidae Kirby 1837 [pp. 338-342]. In:
American Beetles. Volume 2. Polyphaga: Scarabaeoidea through Curculionoidea (R.H.
Arnett, M.C. Thomas, P.E. Skelley, and J. Howard Frank, editors.). CRC Press, Boca
Raton. 861pp.
Liebherr, J.K. 1988. Zoogeography of Caribbean Insects. Cornell University Press, Ithaca,
NY, 285pp.
Lima, I.M.M. 2002. Record of Cybocephalus sp. (Coleoptera: Nitidulidae) preying on pest
species of Diaspididae (Hemiptera), in the state of Alagoas, Brazil. Neotropical
Entomology 31: 157-159.
Linnaeus, C. 1758. Systema Naturae per regna tria naturae secundum classes, ordines, genera,
species, cum characteribus, differentilis, synonymis, locis. Holmiae, Lauretii Salvii, 8 (X
Ed.). 824pp.
Listabarth, C. 1996. Pollination of Bactris by Phyllotrox and Epurea. Implications of the
palm breeding beetles on pollination at the community level. Biotropica 28: 69-81.
Maddison, W.P., M.J. Donoghue, and D.R. Maddison. 1984. Outgroup analysis and
parsimony. Systematic Zoology 33(1): 83-103.
Matadha, D., G.C. Hamilton, M.G. Hughes, and J.H. Lashomb. 2003. Distribution of
natural enemies of Euonymus Scale, Unaspis euonymi (Comstock) (Homoptera:
Diaspididae), in New Jersey. Environmental Entomology 32: 602-607.
McHugh, J.V. 1993. A revision of Eurysphindus LeConte (Coleoptera: Cucujoidea:
Sphindidae) and a review of sphindid classification and phylogeny. Systematic
Entomology 18: 57-92.
McHugh, J.V. 2002. Sphindidae Jacquelin duVal 1861. pp. 305-308. In R.H. Arnett, M.C.
Thomas, P.E. Skelley, and J. Howard Frank eds. American Beetles. Volume 2.
Polyphaga: Scarabaeoidea through Curculionoidea. CRC Press, Boca Raton. 861pp.
Morrone, J.J. and Marquez, J. 2001. Halffter's Mexican Transition Zone, beetle generalized
tracks, and geographical homology. Journal of Biogeography 28: 635-650.
Murray, A. 1864. Monograph of the family Nitidulariae. Transactions of the Linnaean Society
of London 24: 211-439.
362
Murray, A. 1867. List of Coeloptera received from Old Calabar, on the west coast of Africa.
Annals and Magazine of Natural History 19: 167-179.
Murray, A. 1868. Description of a new genus of Nitidulidae. Coleopterologische Hefte 4: 78.
Nakane, T. 1959. Entomological results from the scientific survey of the Tokara Islands. VIII.
Coleoptera: Clavicornia - Nitidulidae, Languriidae, Erotylidae, and Endomychidae.
Scientific Report of Kyoto Prefecture University 3: 53-61.
Naskrecki, P. and R.K. Colwell. 1995. New genus and two new species of Melicharini from
Venezuela (Acari: Mesostigmata: Ascidae). Annals of the Entomological Society of
America 88: 284-293.
Navarette-Heredia, J.L. 2003. Notes on Mexican Psilopyga and Oxycnemus (Coleoptera:
Nitidulidae). Entomological News 114: 81-85.
Nei, M. 1978. Estimation of average heterozygosity and genetic distance from a small
number of individuals. Genetics 89: 583-590.
Nixon, K.C. and J.M. Carpenter. 1996. On outgroups. Cladistics 9: 413-426.
Pakaluk, J., S.A. Slipinski, and J.F. Lawrence. 1995. Current classification and family-group
names in Cucujoidea (Coleoptera). Genus 5: 223-268.
Parsons, C.T. 1936. Notes on North American Nitidulidae: Pocadius. Psyche 2:114-118.
Parsons, C.T. 1943. A revision of the Nearctic Nitidulidae (Coleoptera). Bulletin of the
Museum of Comparative Zoology 92: 121-278.
Parsons, C.T. 1972. On the mesosternum in some Nitidulidae (Coleoptera), with a key to the
New World Amphicrossus. The Coleopterists Bulletin 26: 103-114.
Payne, J.A. 1965. A summer carrion study of the baby pig Sus scrofa Linnaeus. Ecology 46:
592-602.
Payne, J.A. and E.W. King. 1970. Coleoptera associated with carrion. Entomologists
Monthly Magazine 105: 224-232.
Phillips, T.K. and M.A. Ivie. 2002. Bothrideridae Erichson 1845 [pp. 358-362]. In: American
Beetles. Volume 2. Polyphaga: Scarabaeoidea through Curculionoidea (R.H. Arnett,
M.C. Thomas, P.E. Skelley, and J. Howard Frank, editors.). CRC Press, Boca Raton.
861pp.
Poole, R.W. and P. Gentili Eds. 1996. Nomina Insecta Nearctica. Volume 1: Coleoptera,
Strepsiptera. Entomological Information Services. Rockville, MD. 827pp.
363
Price, M.B. 2002. Smicripidae Horn 1879 [pp. 316-318]. In: American Beetles. Volume 2.
Polyphaga: Scarabaeoidea through Curculionoidea (R.H. Arnett, M.C. Thomas, P.E.
Skelley, and J. Howard Frank, editors.). CRC Press, Boca Raton. 861pp.
Reitter, E. 1873. Systematische Eintheilung der Nitidularian. Verhandlungen naturgischichte
Vereines in Brünn 12: 5-194.
Reitter, E. 1874a. Beschreibungen neuer Käfer-Arten nebst synonymischen Notizen.
Verhandlungen der Kaiserlich-Koniglichen Zoologisch-Botanischen Gesellschaft 24:
509-528.
Reitter, E. 1874b. Diagnosen der bekannten Cybocephalus-Arten. Verhandlungen des
Naturforschenden Vereines in Brünn 12: 1-10.
Reitter, E. 1875. Beschereibungen neuer Nitidulidae. Verhandlungen des Naturforschenden
Vereines in Brünn 13: 111.
Reitter, E. 1876a. Neue Nitidularier. Deutsche Entomologische Zeitschrift 20: 305-311.
Reitter, E. 1876b. Neue exotische Nitidulidae. Stettiner Entomologische Zeitung 37: 317-320.
Reitter, E. 1876c. Neue Clavicornien. Stettiner Entomologische Zeitung 37: 363-368.
Reitter, E. 1877. Beiträge zur Käferfauna von Japan. Deutsche Entomologische Zeitschrift
21: 369-384.
Reitter, E. 1880. Neun neue Clavicornier (Coleoptera). Verhandlungen des Naturforschenden
Vereines in Brünn 18: 1-6.
Reitter, E. 1883. Die Nitiduliden Japans. Wiener Entomologische Zeitung 3(9): 257-303.
Reitter, E. 1884. Platychorodes, nov. gen. Nitidulidarum. Deutsche Entomologische
Zeitschrift 28: 261-262.
Rosen, D.E. 1976. A Vicariance Model of Caribbean Biogeography. Systematic Zoology 24:
431-464.
Rosen, D.E. 1985. Geological Hierarchies and Biogeographic Congruence in the Caribbean.
Annals of the Missouri Botanical Garden 72: 636-659.
Scariot, A.O. and E. Lieras. 1991. Reproductive biology of the palm Acrocomia aculeata in
Central Brazil. Biotropica 23: 12-22.
Schaeffer, C.F. 1911. Pocadius basalis Schaeffer. Journal of the New York Entomological
Society 19:117.
364
Schubert, C. 1988. Climatic changes during the last glacial maximum in northern South
America and the Caribbean: A review. Interciencia. Caracas. 13 (3): 128-137.
SenGupta, T. 1967. A new subfamily of Languriidae (Coleoptera) based on four genera, with a
key to species of Toramus. Proceedings of the Royal Entoological Society of London,
Series B 36: 167-176.
SenGupta, T. 1968a. Review of the genera of the tribe Loberini (Coleoptera: Languriidae).
Breviora 303: 1-27.
SenGupta, T. 1968b. Revision of the genera of Cladoxenini (= Cladoxeninae Arrow) and
Thallisellini trib. nov. of the family Languriidae (Coleoptera: Clavicornia). Journal of
Natural History 2: 463-475.
SenGupta, T. 1969. On the taxonomy of Erotylidae (Insecta: Coleoptera: Clavicornia), with
descriptions of two new larvae. Proceedings of the Zoological Society of Calcutta 22:
97-107.
SenGupta, T. 1988. Review of the genera of the family Rhizophagidae (Clavicornia:
Coleoptera) of the world. Memoirs of the Zoological Survey of India 17: 1-58.
SenGupta, T. 1979. A new subfamily of Merophysiidae (Clavicornia: Coleoptera) and
description of two new species of Gomya Dajoz and its larva. Revue Suisse de
Zoologie 86: 691-698.
SenGupta, T. and R.A. Crowson 1966. A new family of cucujoid beetles, based on six
Australian and one New Zealand genera. Annals and Magazine of Natural History
9: 61-85.
SenGupta, T. and R.A. Crowson 1967. The systematic position of Eicolyctus Sahlberg
(Coleoptera: Languriidae). Proceedings of the Royal Entomological Society of London
(B) 36: 87-93.
SenGupta, T. and R.A. Crowson. 1969a. Further observations on the family Boganiidae with
definition of two new families Cavognathidae and Phloeostichidae. Journal of Natural
History 3: 371-590.
SenGupta, T. and R.A. Crowson. 1969b. On a new family of Clavicornia (Coeloptera) and a
new genus of Languriidae. Proceedings of the Royal Entomological Society of London
(B) 38: 125-131.
SenGupta, T. and R.A. Crowson 1971. A review of the classification of the family
Languriidae (Coleoptera: Clavicornia) and the place of Languriidae in the natural system
of Clavicornia. Memoirs of the Zoological Survey of India 15: 1-42.
365
SenGupta, T. and R.A. Crowson 1973. A review of the classification of Cerylonidae
(Coleoptera: Clavicornia). Transactions of the Royal Entomological Society of London
124: 365-446.
SenGupta, T. and R.A. Crowson 1979. The coleopteran family Sphindidae. Entomologists
Monthly Magazine 113: 177-191.
Sharp, D. 1890. Nitidulidae [pp. 265-388]. In: Biologia Centrali-Americana. Insecta,
Coleoptera II. Part 1 (F. D. Godman and O. Salvin, editors). Dulau and Co., London.
717pp.
Sharp, D. 1900. Monotomidae [pp. 563-579, pl. XVIII]. In. Biologia Centrali-Americana.
Insecta, Coleoptera. II. Part 1 (F.D. Godman and O. Salvin, editors.). Dulau and Co.,
London. 717pp.
Sharp, D. 1908. Nitidulidae [pp 435-509 + 3 pls]. In: Fauna Hawaiiensis, vol. 3 (D.A. Sharp,
editor). Cambridge University Press, London .
Shubeck, P.P., N.M. Downie, R.L. Wentzel, and S.B. Peck. 1981. Species composition and
seasonal abundance of carrion beetles in an oak-beech forest in the Great Swamp
National Wildlife Refuge (N.J.). Entomological News 92: 7-16.
Shockley, F.W. and A.R. Cline. 2004. A contribution to the inventory of Coleoptera of
Missouri: New Records from Benton County. Journal of the Kansas Entomological
Society 77: 280-284.
Slipinski, S.A. 1987. A review of the Passandridae of the world (Coleoptera: Cucujoidea). I.
Genus Passandra Dalman. Annali del Museo di Civico Storia Naturelle di Genova 86:
553-603.
Slipinski, S.A. 1989. A review of the Passandridae of the world (Coleoptera: Cucujoidea). II.
Genus Catogenus. Polskie Pismo Entomologiczne 59: 85-129.
Slipinski, S.A. 1990. A monograph of the world Cerylonidae (Coleoptera: Cucujoidea) Part I –
Introduction and higher classification. Annali del Museo Civico di Storia Naturale
“Giacomo Doria” 33: 1-273.
Slipinski, S.A. 1998. Revision and phylogeny of Protocucujidae (Coleoptera: Cucujoidea).
Annales Zoologici 48: 275-298.
Slipinski, S.A. and J. Pakaluk. 1991. Problems in the classification of the Cerylonid series of
Cucujoidea (Coleoptera) Pp. 79-88. In: M. Zunino, X. Belles and M. Blas, eds.
Advances in Coleopterology. European Association of Coleopterology. Silvestrelli and
Cappelletto. Torino.
366
Spornraft, V.K. and A.G. Kirejtshuk. 1993. Über alte und neue südafrikanische Meligethes-
Arten (Coleoptera, Nitidulidae). Mitteilungen Müenchener Entomologischen
Gesellschaft. 83: 47-75.
Steiner, W. 2002. Phalacridae Leach 1815 [pp. 335-337]. In: American Beetles. Volume 2.
Polyphaga: Scarabaeoidea through Curculionoidea (R.H. Arnett, M.C. Thomas, P.E.
Skelley, and J. Howard Frank, editors.). CRC Press, Boca Raton. 861pp.
Stephens, J.F. 1839. A Manual of British Coeloptera, or beetles; containing a brief description
of all the species of beetles hitherto ascertained to inhabit Great Britain and Ireland;
together with a notice of their chief localities, times and places of appearances, etc.
London. 443pp.
Thomas, M.C. 1984a. Two new genera of Neotropical Laemophloeinae (Coleoptera:
Cucujidae). The Florida Entomologist 67: 437-453.
Thomas, M.C. 1984b. A new Neotropical genus and species of rostrate Laemophloeinae
(Coleoptera: Cucujidae) with discussion of the systematic position of the subfamily. The
Coleopterists Bulletin 38:67-83.
Thomas, M.C. 1984c. A new species of apterous Telephanus (Coleoptera: Silvanidae) with a
discussion of phylogenetic relationships of the Silvanidae. The Coleopterists Bulletin
38: 43-55.
Thomas, M.C. 2002a. Cucujidae Latrielle 1802 [pp. 329-330]. In: American Beetles. Volume
2. Polyphaga: Scarabaeoidea through Curculionoidea (R.H. Arnett, M.C. Thomas, P.E.
Skelley, and J. Howard Frank, editors.). CRC Press, Boca Raton. 861pp.
Thomas, M.C. 2002b. Silvanidae Kirby 1837 [pp. 322-326]. In: American Beetles. Volume 2.
Polyphaga: Scarabaeoidea through Curculionoidea (R.H. Arnett, M.C. Thomas, P.E.
Skelley, and J. Howard Frank, editors.). CRC Press, Boca Raton. 861pp.
Thomas, M.C. 2002c. Laemophloeidae Ganglbauer 1899 [pp. 331-334]. In: American Beetles.
Volume 2. Polyphaga: Scarabaeoidea through Curculionoidea (R.H. Arnett, M.C.
Thomas, P.E. Skelley, and J. Howard Frank, editors.). CRC Press, Boca Raton. 861pp.
Thomson, C.G. 1859. Skandinaviens Coleoptera, snoptiskt bearbetade. 1: 290pp.
Tian, M. and H. Pang. 1994. A new species of Cybocephalus (Coleoptera: Cybocephalidae)
from Taiwan. Chinese Journal of Entomology 14: 401-403.
Tomaszweska, W. 2000. Morphology, phylogeny, and classification of adult Endomychidae
(Coleoptera: Cucujoidea). Annales Zoologici 50: 449-558.
Tomaszweska, W. and S.A. Slipinski. 1995. A review of the family Hobartiidae (Coeloptera:
Cucujoidea). Genus 6: 303-325.
367
Watrous, L.E. and Q.D. Wheeler. 1981. The outgroup comparison method of character
analysis. Systematic Zoology 30: 1-11.
Vandenberg, N.J. 2002. Coccinellidae Latrielle 1807 [pp. 371-389]. In: American Beetles.
Volume 2. Polyphaga: Scarabaeoidea through Curculionoidea (R.H. Arnett, M.C.
Thomas, P.E. Skelley, and J. Howard Frank, editors.). CRC Press, Boca Raton. 861pp.
Verhoeff, K.W. 1923. Beiträge zur Kenntnis der Coleopteren-Larven mit besonderer Berück-
sichtigung der Clavicornia. 89(A): 1-109.
Yu, G. and M. Tian. 1995. Notes on the genus Cybocephalus Erichson from China
(Coleoptera: Cybocephalidae). Entomologia Sinica 2: 35-38.
368
APPENDIX. DATA MATRIX
1 2
1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0
adustus 0 0 0 1 0 0 0 1 1 1 0 1 0 1 1 0 1 0 0 0
africanus 0 1 0 1 1 0 0 1 1 1 0 1 0 1 1 0 2 0 0 0
antennuliferus 0 1 0 1 1 1 1 1 1 1 2 1 0 1 1 0 2 0 0 0
ashei 0 1 0 1 1 1 1 1 1 1 1 1 0 1 1 0 2 0 0 0
barclayi 0 0 0 1 0 0 0 1 1 1 0 1 0 1 1 0 1 0 0 0
basalis 0 1 0 1 1 0 1 1 1 1 1 1 0 1 1 1 2 1 1 1
bicolor 0 1 1 1 1 1 0 1 1 1 2 1 0 1 1 1 2 0 1 1
brevis 0 1 0 1 1 0 1 1 1 1 2 1 1 1 1 1 2 0 1 1
carltoni 1 0 0 1 1 0 1 1 1 1 0 1 0 1 1 0 2 0 1 0
centralis 1 0 0 1 1 0 1 1 1 1 1 1 0 1 1 1 2 1 1 0
cochabambus 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 0
coxus 0 1 0 1 1 1 1 1 1 1 1 1 1 1 1 0 2 0 1 0
crypsis 0 0 0 1 1 1 1 1 1 1 1 1 0 1 1 1 2 0 1 1
decoratus 0 0 0 1 0 0 0 1 1 1 0 1 0 1 1 0 1 0 0 0
dimidiatus 1 0 0 1 1 1 0 1 1 1 1 1 0 1 1 1 2 1 1 1
dominicus 0 1 0 1 1 0 1 1 1 1 0 1 1 1 1 1 2 0 0 1
endrodi 0 1 0 1 1 0 0 1 1 1 0 1 1 1 1 0 2 0 0 0
ephite 1 1 0 1 1 1 0 1 1 1 1 1 0 1 1 1 2 0 1 1
falini 0 0 0 1 1 1 1 1 1 1 0 1 1 1 1 1 2 0 1 1
femoralis 0 0 0 1 1 0 1 1 1 1 0 1 0 1 1 0 2 0 0 0
ferrugineus 0 1 0 1 1 0 1 1 1 1 0 1 0 1 1 0 2 0 0 0
fulvipennis 0 1 0 1 1 0 1 1 1 1 1 1 0 1 1 1 2 1 1 1
fumatus 1 0 0 1 1 1 0 1 1 1 2 1 0 1 1 0 2 0 1 1
fusifromis 0 0 0 1 0 0 0 1 1 1 0 1 0 1 1 0 1 0 0 0
globularis 0 0 0 1 1 1 1 1 1 1 0 1 0 1 1 0 2 0 1 0
helvolus 1 0 0 1 1 0 0 1 1 1 1 1 0 1 1 0 2 0 1 1
insularis 0 1 0 1 1 1 0 1 1 1 1 1 1 1 1 0 2 0 1 1
jelineki 0 0 0 1 1 1 0 1 1 1 1 1 1 1 1 1 2 0 0 1
kirejtshuki 0 0 0 1 0 0 0 1 1 1 0 1 0 1 1 0 1 0 0 0
luisalfredoi 0 1 0 1 1 0 0 1 1 1 1 1 1 1 1 1 2 0 0 1
majusculus 0 0 0 1 1 0 0 1 1 1 0 1 0 1 1 0 2 0 0 1
maquipucunensis 0 1 0 1 1 1 1 1 1 1 2 1 0 1 1 1 2 0 1 1
martini 0 0 0 1 1 0 0 1 1 1 0 1 0 1 1 0 2 0 0 0
monticolis 0 0 0 1 1 0 0 1 1 1 0 1 0 1 1 0 2 0 0 0
niger 0 1 0 1 1 0 1 1 1 1 1 1 0 1 1 0 2 1 1 1
nigerrimus 0 1 0 1 1 1 1 1 1 1 1 1 0 1 1 0 2 0 1 1
nobilis 0 0 0 1 1 0 0 1 1 1 0 1 0 1 1 1 2 0 0 0
okinawaensis 0 0 0 1 1 0 0 1 1 1 0 1 0 1 1 0 2 0 0 0
pecki 0 0 0 1 1 0 0 1 1 1 1 1 0 1 1 1 2 0 1 1
peruensis 0 0 1 1 1 1 0 1 1 1 1 1 0 1 1 1 2 1 1 1
rubidus 0 0 0 1 0 1 0 1 1 1 1 1 1 1 1 0 2 0 1 1
tepicensis 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 0 2 0 0 1
testaceous 2 0 0 1 0 0 0 1 1 1 1 1 0 1 1 0 1 2 0 0
torresi 0 1 0 1 1 1 0 1 1 1 1 1 0 1 1 0 2 0 0 1
wappesi 1 0 0 1 1 0 0 1 1 1 0 1 0 1 1 0 2 0 1 1
yunnanensis 0 0 0 1 0 0 0 1 1 1 1 1 0 1 1 0 1 0 0 0
Thalycra 1 0 0 0 0 0 0 0 0 1 0 1 1 0 0 1 - 1 1 1
Pocadionta 1 0 0 0 1 1 0 0 0 1 0 1 0 0 0 0 2 0 1 1
Lasiodactylus 0 1 0 0 0 0 0 1 0 0 0 1 0 1 1 0 - 0 1 1
Hebascus 0 0 0 0 1 0 1 1 0 0 0 0 1 2 1 0 2 0 1 1
Teichostethus 0 0 0 0 1 1 1 1 0 0 0 1 0 1 1 0 2 0 1 0
Hyleopocadius 0 1 0 0 0 1 1 1 0 0 2 1 0 1 1 0 0 0 0 0
Axyra 0 1 0 0 0 0 0 1 0 0 0 1 0 1 1 0 0 0 0 0
Epuraea 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 - 0 0 0
Carpophilus 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 - 0 0 0
369
3 4
1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0
adustus 0 0 1 1 1 1 1 0 1 1 0 1 0 1 0 0 0 1 1 1
africanus 2 0 1 1 1 1 1 0 1 1 0 1 0 1 0 0 0 2 1 1
antennuliferus 2 1 1 1 1 2 1 0 2 1 1 1 1 2 0 1 2 2 1 1
ashei 2 0 1 1 1 2 1 1 2 1 1 1 0 2 0 1 2 2 1 1
barclayi 2 0 1 1 1 1 1 1 1 1 0 1 0 1 0 0 1 2 1 1
basalis 1 1 1 1 1 1 1 1 2 1 2 1 1 1 0 1 2 2 1 1
bicolor 2 0 1 1 1 2 1 1 2 1 1 1 1 2 0 1 2 3 1 1
brevis 2 0 1 1 1 1 1 1 1 1 0 1 1 1 0 1 1 1 1 1
carltoni 2 1 1 1 1 2 1 1 2 1 2 1 1 2 0 1 2 2 1 1
centralis 0 0 1 1 1 2 1 1 2 1 1 1 1 2 0 1 2 2 1 1
cochabambus 2 1 1 1 1 2 1 1 2 1 1 1 1 2 0 1 1 2 1 1
coxus 2 1 1 1 1 2 1 1 2 1 1 1 1 2 0 1 1 2 1 1
crypsis 2 1 1 1 1 2 1 1 2 1 0 1 1 2 0 0 1 2 1 1
decoratus 2 0 1 1 1 2 1 0 1 1 0 1 1 1 0 0 0 2 1 1
dimidiatus 2 1 1 1 1 2 1 1 2 1 1 1 1 2 0 1 2 2 1 1
dominicus 2 0 1 1 1 1 1 1 2 1 2 1 1 2 0 1 1 1 1 1
endrodi 0 0 1 1 1 2 1 0 1 1 0 1 0 1 0 0 2 3 1 1
ephite 2 0 1 1 1 2 1 1 2 1 2 1 1 1 0 1 2 2 1 1
falini 2 0 1 1 1 2 1 1 2 1 1 1 1 2 0 1 2 1 1 1
femoralis 2 0 1 1 1 2 1 1 2 1 1 1 0 2 0 0 1 1 1 1
ferrugineus 1 0 1 1 1 2 1 0 1 1 0 1 0 2 0 0 0 1 1 1
fulvipennis 2 1 1 1 1 2 1 0 2 1 1 1 1 2 0 1 1 2 1 1
fumatus 2 0 1 1 1 2 1 0 2 1 1 1 1 2 0 1 2 2 1 1
fusifromis 0 0 1 1 1 2 1 1 2 1 1 1 0 2 0 0 1 1 1 1
globularis 2 0 1 1 1 2 1 0 2 1 1 1 1 2 0 1 1 2 1 1
helvolus 2 0 1 1 1 2 1 0 2 1 1 1 1 2 0 1 1 3 1 1
insularis 1 0 1 1 1 2 1 0 1 1 0 1 1 2 0 1 2 2 1 1
jelineki 2 0 1 1 1 2 1 1 2 1 1 1 1 2 0 1 2 2 1 1
kirejtshuki 0 0 1 1 1 1 1 0 2 1 1 1 0 1 0 0 1 2 1 1
luisalfredoi 2 1 1 1 1 2 1 1 2 1 1 1 0 1 0 1 2 3 1 1
majusculus 2 1 1 1 1 1 1 0 2 1 1 1 1 1 0 0 1 1 1 1
maquipucunensis 1 1 1 1 1 2 1 1 2 1 1 1 1 2 0 0 0 2 1 1
martini 0 0 1 1 1 1 1 0 2 1 1 1 1 1 0 0 1 2 1 1
monticolis 2 1 1 1 1 1 1 0 1 1 0 1 0 1 0 0 0 1 1 1
niger 2 1 1 1 1 2 1 0 2 1 1 1 1 2 0 1 2 3 1 1
nigerrimus 2 1 1 1 1 2 1 1 1 1 0 1 1 2 0 1 2 3 1 1
nobilis 0 0 1 1 1 2 1 0 1 1 0 1 0 2 0 0 1 2 1 1
okinawaensis 2 0 1 1 1 2 1 1 1 1 0 1 1 2 0 0 0 2 1 1
pecki 2 0 1 1 1 2 1 1 2 1 2 1 1 1 0 1 2 3 1 1
peruensis 2 1 1 1 1 2 1 1 2 1 1 1 1 2 0 1 1 2 1 1
rubidus 2 0 1 1 1 2 1 0 2 1 2 1 1 2 0 0 2 3 1 1
tepicensis 2 1 1 1 1 2 1 1 2 1 1 1 1 2 0 0 2 2 1 1
testaceous 2 0 1 1 1 1 1 0 2 1 1 1 1 1 0 0 1 1 1 1
torresi 2 1 1 1 1 2 1 1 2 1 2 1 1 2 0 1 1 2 1 1
wappesi 1 0 1 1 1 2 1 1 2 1 1 1 1 2 0 1 1 2 1 1
yunnanensis 1 0 1 1 1 1 1 0 1 1 0 1 1 1 0 0 2 2 1 1
Thalycra 0 0 1 0 1 0 1 0 0 0 1 1 0 0 1 0 0 1 1 0
Pocadionta 0 0 1 0 1 2 1 0 0 1 1 1 0 2 1 0 2 2 1 0
Lasiodactylus 0 0 1 1 1 0 1 0 0 0 1 1 1 0 0 0 2 1 2 0
Hebascus 2 1 1 1 1 2 1 0 0 0 3 1 1 2 0 0 2 3 1 0
Teichostethus 2 0 1 1 1 2 1 0 0 2 3 1 0 2 1 0 2 3 1 0
Hyleopocadius 2 1 1 1 1 2 1 1 0 1 3 2 1 1 1 0 2 3 1 1
Axyra 2 0 1 0 1 1 1 0 0 0 4 1 0 1 0 0 0 0 0 0
Epuraea 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0
Carpophilus 0 0 1 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0
370
5 6
1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0
adustus 0 0 1 0 1 0 1 1 1 1 2 0 0 0 1 0 0 0 0 1
africanus 0 0 2 0 1 0 1 1 1 1 1 0 1 0 1 1 0 0 0 1
antennuliferus 1 1 1 0 1 1 1 1 1 0 2 0 1 1 1 1 1 1 1 1
ashei 1 0 1 0 1 0 1 1 1 2 2 0 1 0 1 1 1 1 1 1
barclayi 0 1 1 0 1 1 1 1 1 1 2 1 0 0 1 1 1 1 0 1
basalis 1 0 1 0 1 0 1 1 1 2 2 0 1 0 1 0 0 0 1 1
bicolor 1 1 1 0 1 1 1 1 1 1 2 0 ? 1 1 1 1 1 1 1
brevis 1 0 1 0 1 0 1 1 1 2 2 0 1 0 1 0 0 0 1 1
carltoni 1 0 1 0 1 1 1 1 1 2 2 0 1 0 1 1 1 1 1 1
centralis 1 1 1 0 1 1 1 1 1 2 2 0 0 0 1 0 0 0 0 1
cochabambus 1 0 1 0 1 1 1 1 1 1 2 0 1 0 1 1 1 1 0 1
coxus 1 1 1 0 1 1 1 1 1 2 2 0 1 0 1 1 1 1 1 1
crypsis 1 1 1 0 1 1 1 1 1 2 2 0 1 1 1 1 1 1 1 1
decoratus 1 0 1 0 1 0 1 1 1 1 2 1 1 0 1 0 1 1 0 1
dimidiatus 1 1 1 0 1 1 1 1 1 2 2 0 1 0 1 1 1 1 1 1
dominicus 1 0 1 0 1 0 1 1 1 2 2 0 1 0 1 0 0 0 1 1
endrodi 1 0 1 0 1 0 1 1 1 1 2 0 1 0 1 0 0 0 0 1
ephite 1 0 1 0 1 1 1 1 1 2 2 0 1 0 1 1 1 1 1 1
falini 1 1 1 0 1 1 1 1 1 2 2 0 0 0 1 1 1 1 1 1
femoralis 1 0 1 0 1 0 1 1 1 2 2 1 ? 0 1 1 1 1 0 1
ferrugineus 0 0 1 0 1 0 1 1 1 1 2 0 1 0 1 0 0 0 0 1
fulvipennis 1 0 1 0 1 0 1 1 1 2 2 0 1 0 1 0 0 0 1 1
fumatus 1 0 1 0 1 0 1 1 1 1 2 0 1 0 1 1 1 1 1 1
fusifromis 0 1 1 0 1 1 1 1 1 1 2 1 1 0 1 1 1 1 0 1
globularis 1 0 1 0 1 1 1 1 1 2 2 0 2 0 1 0 0 0 1 1
helvolus 1 0 1 0 1 0 1 1 1 2 2 0 1 0 1 0 0 0 1 1
insularis 1 1 1 0 1 1 1 1 1 1 2 0 1 0 1 0 1 1 1 1
jelineki 1 1 1 0 1 1 1 1 1 1 2 0 1 0 1 1 1 1 1 1
kirejtshuki 1 0 1 0 1 0 1 1 1 1 2 1 1 0 1 1 1 1 1 1
luisalfredoi 1 0 1 0 1 1 1 1 1 1 2 0 0 0 1 0 0 0 1 1
majusculus 1 1 1 0 1 1 1 1 1 1 2 1 1 0 1 1 1 1 1 1
maquipucunensis 1 1 1 0 1 1 1 1 1 1 2 0 1 0 1 1 1 1 1 1
martini 1 1 1 0 1 1 1 1 1 1 2 1 ? 0 1 1 1 1 0 1
monticolis 0 0 1 0 1 0 1 1 1 1 2 0 ? 0 1 1 0 1 0 1
niger 1 0 1 0 1 0 1 1 1 2 2 0 1 0 1 0 0 0 1 1
nigerrimus 1 1 1 0 1 1 1 1 1 2 2 0 1 0 1 1 1 1 1 1
nobilis 1 0 1 0 1 0 1 1 1 1 2 0 1 0 1 0 0 0 0 1
okinawaensis 1 1 1 0 1 1 1 1 1 1 2 0 1 0 1 1 1 1 0 1
pecki 1 0 1 0 1 0 1 1 1 1 2 0 1 0 1 1 1 1 1 1
peruensis 1 1 1 0 1 1 1 1 1 2 2 0 1 0 1 1 1 1 1 1
rubidus 1 0 1 0 1 0 1 1 1 2 2 0 1 1 1 0 0 0 1 1
tepicensis 1 1 1 0 1 1 1 1 1 1 2 0 0 0 1 1 1 1 1 1
testaceous 1 0 1 0 1 0 1 1 1 1 2 0 1 0 1 1 1 1 0 1
torresi 1 1 1 0 1 1 1 1 1 2 2 0 1 0 1 0 0 1 1 1
wappesi 1 0 1 0 1 0 1 1 1 1 2 0 1 0 1 1 1 1 1 1
yunnanensis 1 0 1 0 1 0 1 1 1 0 2 0 ? 0 1 0 0 0 1 1
Thalycra 0 0 1 0 1 0 1 1 1 2 2 0 2 0 1 0 2 0 1 0
Pocadionta 0 0 0 0 1 0 1 1 1 2 0 0 2 0 1 0 2 0 1 0
Lasiodactylus 0 0 1 0 1 0 1 0 0 2 1 1 2 0 1 0 0 0 0 0
Hebascus 1 2 1 0 1 2 1 1 1 2 2 0 2 0 2 0 0 0 0 0
Teichostethus 1 2 1 0 1 2 1 1 1 2 2 0 2 0 1 0 0 0 0 0
Hyleopocadius 1 1 0 1 1 1 1 0 1 2 2 0 2 0 1 0 0 0 0 0
Axyra 0 0 1 0 1 0 1 1 0 2 2 1 2 0 0 0 0 0 0 1
Epuraea 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0
Carpophilus 0 0 0 0 1 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0
371
7 8
1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0
adustus 1 1 0 0 2 0 0 0 0 0 1 1 0 0 0 2 2 1 1 1
africanus 1 1 1 1 2 0 0 0 0 1 1 0 0 0 0 2 2 0 1 1
antennuliferus 1 1 1 1 2 0 0 0 0 1 1 0 0 0 0 2 2 1 1 1
ashei 1 1 0 0 ? 0 0 0 0 1 1 2 ? 0 1 2 2 1 1 1
barclayi 1 1 1 1 2 0 0 0 0 1 1 2 0 0 1 2 2 1 1 1
basalis 1 1 2 2 4 0 0 1 0 1 1 0 1 1 2 0 2 1 1 1
bicolor 1 1 2 2 ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ?
brevis 1 1 0 1 3 0 0 0 0 1 1 1 0 0 0 2 2 1 1 1
carltoni 1 1 1 1 2 0 0 0 0 1 2 1 1 0 1 2 2 1 1 1
centralis 1 1 1 1 2 1 0 0 0 1 1 0 0 0 1 2 2 1 1 1
cochabambus 1 1 1 1 2 1 0 1 0 1 1 1 0 0 0 2 2 1 1 1
coxus 1 1 0 0 2 0 0 0 0 1 1 0 1 0 0 2 2 1 1 1
crypsis 1 1 1 2 2 0 0 0 0 1 1 1 0 0 1 2 2 1 1 1
decoratus 1 1 2 2 ? ? 0 1 0 1 ? ? ? 0 2 0 2 ? 1 1
dimidiatus 1 1 0 0 2 1 0 0 0 1 1 2 0 0 0 1 2 1 1 1
dominicus 1 1 2 1 2 0 0 0 0 1 1 0 0 0 1 2 2 1 1 1
endrodi 1 1 1 1 1 0 0 0 0 1 1 1 0 0 0 0 2 0 1 1
ephite 1 1 2 2 2 0 0 0 0 1 1 0 0 0 0 2 2 1 1 1
falini 1 1 2 2 2 0 0 0 1 1 1 2 1 0 0 1 2 1 1 1
femoralis 1 1 2 2 2 ? 0 ? ? ? ? ? 0 ? ? ? ? ? ? ?
ferrugineus 1 1 1 1 2 0 0 0 0 0 1 1 0 0 0 0 2 1 1 1
fulvipennis 1 1 0 0 2 0 0 2 0 1 1 0 0 0 0 2 2 1 1 1
fumatus 1 1 0 0 2 0 0 0 0 1 1 1 0 0 1 2 2 1 1 1
fusifromis 1 1 1 1 2 0 0 0 0 1 1 0 0 1 1 2 2 1 1 1
globularis 1 1 1 2 2 0 0 0 0 1 1 2 1 0 0 2 2 1 1 1
helvolus 1 1 1 1 2 0 0 0 0 1 1 0 0 0 1 2 2 1 1 1
insularis 1 1 1 1 2 0 0 0 0 1 1 2 0 0 1 2 2 1 1 1
jelineki 1 1 0 1 2 0 0 0 0 1 2 1 0 0 1 2 2 1 1 1
kirejtshuki 1 1 0 0 1 0 0 0 0 1 ? ? 0 0 0 2 2 ? 1 1
luisalfredoi 1 1 2 2 2 0 0 0 0 1 1 0 0 0 1 2 2 1 1 1
majusculus 1 1 2 2 2 0 0 1 0 1 ? ? 0 0 2 2 2 1 1 1
maquipucunensis 1 1 2 2 2 0 0 0 0 1 1 0 0 0 0 1 2 1 1 1
martini 1 1 2 2 ? ? 0 ? ? ? ? 0 ? ? ? ? ? 1 1 1
monticolis 1 1 1 1 ? ? 0 ? ? ? ? ? ? ? ? ? ? 1 1 1
niger 1 1 2 2 4 0 0 0 0 1 1 0 1 0 0 2 2 1 1 1
nigerrimus 1 1 1 1 2 0 0 0 0 1 1 2 0 0 0 2 2 1 1 1
nobilis 1 1 1 1 2 0 0 1 0 1 1 1 0 0 0 2 2 1 1 1
okinawaensis 1 1 0 0 2 0 0 0 0 1 1 0 1 0 0 2 2 1 1 1
pecki 1 1 1 1 2 0 0 0 0 1 1 1 1 0 0 1 2 1 1 1
peruensis 1 1 1 1 3 1 0 0 1 1 1 1 1 0 0 2 2 1 1 1
rubidus 1 1 0 0 2 0 0 0 0 1 1 0 0 0 0 2 2 1 1 1
tepicensis 1 1 0 0 2 0 0 0 0 1 2 2 1 0 0 2 2 1 1 1
testaceous 1 1 1 1 2 0 0 1 0 1 1 1 0 0 0 2 2 1 1 1
torresi 1 1 1 1 2 0 0 0 0 1 ? ? 0 0 0 2 2 ? 1 1
wappesi 1 1 0 0 2 0 0 0 0 1 1 2 0 1 0 2 2 1 1 1
yunnanensis 1 1 0 0 ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ?
Thalycra 2 1 0 0 1 - 2 0 0 1 - - 0 0 - 0 2 - 1 1
Pocadionta 2 1 0 0 1 - 1 0 0 1 - - 0 0 - 0 2 - 1 1
Lasiodactylus 1 1 0 0 1 - 0 0 0 1 - - 0 0 - 0 2 - 1 2
Hebascus 1 1 0 0 1 - 0 0 0 1 - - 0 0 - 0 3 - 1 1
Teichostethus 1 1 0 1 1 - 0 0 0 1 - - 0 0 - 0 2 - 1 1
Hyleopocadius 1 1 2 2 1 - 0 0 0 1 - - 0 0 - 0 2 - 1 1
Axyra 1 1 0 1 1 - 0 0 0 1 - - 0 0 - 0 2 - 1 1
Epuraea 0 0 0 0 0 0 0 0 0 0 - - 0 0 - 0 1 - 1 1
Carpophilus 1 0 0 0 0 0 0 0 0 0 - - 0 0 - 0 0 - 0 0
372
9 10
1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0
adustus 0 0 0 0 0 0 0 1 1 1 1 0 0 2 1 2 0 2 1 1
africanus 0 0 0 0 0 0 0 0 0 0 1 0 1 2 0 2 1 2 1 1
antennuliferus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? 1 1
ashei 1 1 1 2 1 0 1 0 1 2 1 1 1 2 1 2 1 2 1 1
barclayi 1 1 0 0 2 0 1 1 0 1 1 0 1 2 1 2 0 2 1 1
basalis 1 0 1 2 0 1 1 0 1 2 0 2 0 2 1 2 1 2 1 1
bicolor 1 0 0 1 1 0 1 1 1 2 1 1 1 2 0 2 1 2 1 1
brevis 0 1 0 0 1 0 0 0 1 1 1 2 1 2 0 2 1 2 1 1
carltoni 1 0 0 1 1 1 0 1 1 1 1 2 1 2 0 2 0 2 1 1
centralis 1 0 0 1 1 0 0 1 1 1 1 2 0 2 0 2 0 2 1 1
cochabambus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? 1 1
coxus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? 1 1
crypsis 1 0 0 1 1 0 1 0 1 2 1 1 1 2 0 2 1 2 1 1
decoratus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? 1 1
dimidiatus 1 0 1 1 1 1 1 1 1 2 1 1 1 2 1 2 1 2 1 1
dominicus 1 0 0 2 1 1 0 0 1 2 1 2 1 2 1 2 1 2 1 1
endrodi 1 0 0 0 1 0 0 0 1 0 1 0 1 2 1 2 1 2 1 1
ephite ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? 1 1
falini 1 ? 1 1 1 0 1 1 1 1 1 2 ? ? 0 2 1 2 1 1
femoralis 0 2 0 0 1 0 0 1 0 0 1 1 1 2 0 2 0 2 1 1
ferrugineus 0 2 0 0 0 1 0 0 0 0 0 0 0 2 1 2 0 2 1 1
fulvipennis 1 0 0 1 1 0 0 0 1 1 1 0 0 2 0 2 1 2 1 1
fumatus 1 0 0 1 1 0 1 0 1 1 1 1 1 2 0 2 1 2 1 1
fusifromis 1 1 0 1 2 0 0 1 1 1 1 1 1 2 1 2 0 2 1 1
globularis 1 0 0 1 1 0 0 0 1 2 1 2 0 2 0 2 1 2 1 1
helvolus 1 0 0 1 1 0 0 0 0 2 1 2 0 2 1 2 0 2 1 1
insularis ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? 1 1
jelineki 0 0 0 2 1 0 0 1 1 2 1 2 1 2 1 2 0 2 1 1
kirejtshuki 1 1 0 1 1 0 0 0 1 2 1 1 ? 2 1 2 1 2 1 1
luisalfredoi 1 0 0 1 1 1 1 0 1 2 1 2 0 2 0 2 0 2 1 1
majusculus 1 2 0 1 1 0 0 1 0 0 0 1 1 2 0 2 0 2 1 1
maquipucunensis 0 0 0 1 1 0 0 1 1 2 1 2 1 2 1 2 0 2 1 1
martini 1 0 0 1 1 0 0 1 0 0 1 2 1 2 1 2 1 2 1 1
monticolis 1 0 0 1 0 0 1 1 0 1 1 1 1 2 0 2 0 2 1 1
niger 1 0 0 1 1 0 0 1 1 2 1 2 0 2 0 2 0 2 1 1
nigerrimus ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? 1 1
nobilis 1 0 0 1 1 0 0 1 1 1 1 0 1 2 1 2 1 2 1 1
okinawaensis 1 0 0 1 1 0 0 1 1 1 1 0 1 2 0 2 1 2 1 1
pecki ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? 1 1
peruensis 1 1 1 1 2 0 1 1 1 2 1 1 1 2 0 2 0 2 1 1
rubidus 0 0 1 1 1 0 1 1 1 1 1 1 0 2 0 2 1 2 1 1
tepicensis 1 0 0 1 1 0 0 0 1 2 1 2 1 2 1 2 1 2 1 1
testaceous 1 1 0 0 2 0 1 1 1 1 1 0 1 2 1 2 0 2 1 1
torresi 1 0 0 1 1 0 0 1 1 1 1 1 0 2 0 2 0 2 1 1
wappesi 1 0 0 1 1 0 1 0 1 2 1 1 1 2 0 2 1 2 1 1
yunnanensis 1 1 0 1 1 0 0 0 1 1 1 1 1 2 0 2 1 2 1 1
Thalycra 0 - 0 0 0 0 0 1 0 0 - - 0 0 0 1 0 1 0 0
Pocadionta 0 - 0 0 0 0 0 1 0 0 - - 0 0 0 1 0 1 ? ?
Lasiodactylus 0 - 1 0 - 0 0 0 1 0 - - 0 0 0 0 0 1 ? ?
Hebascus 0 - 0 0 0 0 0 1 0 0 - - 2 2 0 2 0 2 ? ?
Teichostethus 0 - 0 0 0 0 0 1 0 0 - - 1 2 0 2 0 2 ? ?
Hyleopocadius 1 - 0 0 - 1 0 1 0 0 - - 0 1 0 1 0 2 ? ?
Axyra 0 - 0 0 - 0 0 0 0 0 - - 0 0 0 0 0 1 ? ?
Epuraea 0 - 0 0 - 0 0 0 0 0 - - 0 0 0 1 0 0 0 0
Carpophilus 0 - 0 0 - 0 0 0 0 0 - - 0 0 0 0 0 0 0 0
373
11
1 2 3 4 5 6 7 8 9 0 1 2 3 4
adustus 1 1 0 0 1 0 0 0 1 0 1 0 1 1
africanus 1 1 0 0 0 0 0 1 0 0 1 0 1 1
antennuliferus 1 1 1 0 2 1 0 1 1 ? 1 0 1 1
ashei 1 1 ? 1 1 0 0 1 1 1 1 1 1 1
barclayi 1 1 1 0 1 0 1 1 0 1 1 0 1 1
basalis 1 1 1 0 1 1 0 2 1 0 1 0 1 1
bicolor 1 ? ? ? ? ? 0 ? ? 1 1 ? 1 1
brevis 1 1 1 0 1 1 0 1 1 0 1 0 1 1
carltoni 1 1 1 0 1 0 0 1 1 0 1 0 1 1
centralis 1 1 1 0 1 0 0 1 1 0 1 0 1 1
cochabambus 1 1 1 0 1 1 0 1 1 ? 1 1 1 1
coxus 1 1 1 0 1 0 0 1 1 ? 1 1 1 1
crypsis 1 1 1 0 2 0 0 1 1 1 1 0 1 1
decoratus 1 1 ? ? ? 0 0 2 1 ? 1 1 1 1
dimidiatus 1 1 1 1 1 1 0 1 1 1 1 1 1 1
dominicus 1 1 1 0 1 0 0 1 1 0 1 1 1 1
endrodi 1 1 0 0 0 0 0 1 0 0 1 1 1 1
ephite 1 1 1 0 1 0 0 1 1 ? 1 0 1 1
falini 1 1 1 0 1 0 0 1 1 0 1 1 1 1
femoralis 1 ? 1 0 1 ? 0 ? ? 1 1 ? 1 1
ferrugineus 1 0 0 0 1 0 0 0 1 0 1 0 1 1
fulvipennis 1 1 1 0 1 0 0 0 1 0 1 0 1 1
fumatus 1 1 1 1 1 0 0 1 1 1 1 1 1 1
fusifromis 1 1 1 0 1 1 1 1 0 1 1 0 1 1
globularis 1 1 1 0 1 0 0 1 1 0 1 1 1 1
helvolus 1 1 1 0 1 0 0 1 1 0 1 0 1 1
insularis 1 1 1 0 1 0 0 1 1 ? 1 1 1 1
jelineki 1 1 1 0 1 0 0 1 1 0 1 1 1 1
kirejtshuki 1 1 0 ? ? 0 1 1 1 0 1 0 1 1
luisalfredoi 1 1 1 0 1 0 0 1 1 0 1 1 1 1
majusculus 1 1 1 1 1 1 1 2 1 1 1 1 1 1
maquipucunensis 1 1 1 0 1 0 0 1 1 0 1 1 1 1
martini ? ? ? ? ? ? 0 ? ? 0 1 ? 1 1
monticolis ? ? ? ? ? ? 0 ? ? 0 1 ? 1 1
niger 1 1 1 0 1 0 0 1 1 0 1 1 1 1
nigerrimus 1 1 1 0 1 0 0 1 1 ? 1 1 1 1
nobilis 1 1 1 1 1 1 0 0 1 1 1 0 1 1
okinawaensis 1 1 1 0 1 0 0 0 1 1 1 0 1 1
pecki 1 1 1 0 1 0 0 1 1 ? 1 0 1 1
peruensis 1 1 1 0 1 0 0 1 1 1 1 0 1 1
rubidus 1 1 1 0 1 1 0 0 1 0 1 1 1 1
tepicensis 1 1 1 0 1 1 0 1 1 0 1 0 1 1
testaceous 1 1 1 0 1 1 0 1 0 1 1 0 1 1
torresi 1 1 1 ? ? 0 0 1 1 1 1 1 1 1
wappesi 1 1 1 1 1 1 0 0 1 1 1 1 1 1
yunnanensis ? ? ? ? ? ? 0 ? ? 1 1 ? 1 1
Thalycra 0 1 0 - - - 0 - 1 - 0 - 0 1
Pocadionta ? 1 0 - - - 0 - 1 - ? - 1 2
Lasiodactylus ? 1 0 - - - 0 - 1 - ? - 0 1
Hebascus ? 1 1 - - - 0 - 1 - ? - 1 1
Teichostethus ? 1 1 - - - 0 - 1 - ? - 1 1
Hyleopocadius ? 1 1 - - - 0 - 1 - ? - 1 2
Axyra ? 1 0 - - - 0 - 1 - 0 - 1 1
Epuraea 0 0 0 - - - 0 - 0 - 0 - 0 0
Carpophilus 0 0 0 - - - 0 - 0 - 0 - 0 0
374
VITA
Andrew R. Cline was born in Springfield, Illinois, in March of 1974, and spent
the first ten years of his life in the “Land of Lincoln.” The Cline family then moved to
northern Alabama, following an employment opportunity for Roger Cline, Andrew’s
father. In the Decatur area, Andrew and his two brothers spent much time enjoying
outdoor activities and gaining an appreciation for the natural beauty of the southern
states. In May of 1995 Andrew married JoAnna Caudle. Andrew and JoAnna were
blessed with their first child, Joshua David, in July of 2002.
Andrew received his Bachelor of Science degree in biology/ecology at the
University of Alabama – Huntsville in 1996, and following graduation joined the
department as a part-time instructor. In 1998, he began graduate studies at the University
of Missouri, and in 2000 was awarded a Master of Science degree in entomology with a
specialization in insect behavior, and also was awarded the Stone Award for Outstanding
Master of Science student in the Department of Entomology.
Andrew joined the Entomology Department at LSU in the fall of 2000, pursuing
doctoral work under the direction of Dr. Chris Carlton. During doctoral research he
engaged in biodiversity, ecological, taxonomic and systematic research on nitidulids and
other beetles. Andy took part in three international biodiversity studies including: the
INBio initiative in Costa Rica, the IBISCA working group in Panama, and a project to
document the fauna of Nova Scotia and Prince Edward Island. In a pedagogical context,
Andy taught numerous biology and entomology courses at his undergraduate and
graduate institutions, and attended a National Science Foundation teaching workshop in
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June of 2004, which was devoted to the advancement of undergraduate instruction in
science.
Funding for Andrew’s doctoral research came from a number of sources
including: a National Science Foundation Doctoral Dissertation Improvement Grant, an
Ernst Mayr Grant from the Museum of Comparative Zoology at Harvard University,
visiting systematist grants from the Canadian Museum of Nature and Field Museum, an
Organization for Tropical Studies and Smithsonian Tropical Research Institute A.W.
Mellon Grant, a Sigma Xi Grant in Aid of Research, Florida Entomological Society
Grants, LSU Entomology Department Funds, and the Louisiana State University
Agricultural Center and Graduate School.
The dissertation presented herein is not the culmination of an academic career but
rather the beginning of another phase of scientific investigation. Much has changed since
Andrew began his journey in higher education and this work is dedicated to those who
have provided inspiration and encouragement throughout those many years.
Andrew is currently an Associate Insect Biosystematist at the Plant Pest
Diagnostics Branch of the California Department of Food and Agriculture. At his new
post he is able to not only aid in the detection of potential beetle pests of California, but
also has an active research program involving the taxonomy and systematics of little
known groups of beetles.
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