Revision of Philoscaptus brèthes, 1919 and description of a new genus (Coleoptera, Melolonthidae, Dynastinae)

Abstract

The genus Philoscaptus Brèthes, 1919, is revised and found to be monotypic, including P. bonariensis (Burmeister) a species found mainly in open areas along the Paraná River basin, in Brazil, Uruguay, Argentina and Paraguay (new country record). The new genus Saccharoscaptus is described for P. laminifer Dechambre, differentiated by mandibles with one weak rounded apical tooth, apex of clypeus with no denticles, frons excavate, male with blade like process on clypeus and small pronotal tubercles. A key for males of the Neotropical Pentodontini genera is given and the current status of Ligyrus Burmeister and Tomarus Erichson is discussed and a checklist with the known species included in each genus is also provided.

Full text

ISSN 2443-437X (electrónico)
ISSN 1317-5262 (impreso)
© 2015, Sociedad Venezolana de Entomología
ENTOMOTROPICA
Vol. 30(1): 1-11
on line 20-Abril-2015
Revision of Philoscaptus Brèthes, 1919 and description of a new genus
(Coleoptera, Melolonthidae, Dynastinae)
urn:lsid:zoobank.org:pub:4094B56C-A1A3-4350-BE91-534DC4BC6976
Miguel Angel Morón Rios1, Paschoal Coelho Grossi2
1Instituto de Ecologia, A.C. Xalapa, México. E-mail: moronrios.miguelangel@gmail.com
2Universidade Estadual de Maranhao, Brasil.
Abstract
M R MA, G PC. 2015. Revision of Philoscaptus Brèthes, 1919 and description of a new genus
(Coleoptera, Melolonthidae, Dynastinae). E 30(1): 1-11.
e genus Philoscaptus Brèthes, 1919, is revised and found to be monotypic, including P. bonariensis (Burmeister)
a species found mainly in open areas along the Paraná River basin, in Brazil, Uruguay, Argentina and Paraguay
(new country record). e new genus Saccharoscaptus is described for P. laminifer Dechambre, dierentiated by
mandibles with one weak rounded apical tooth, apex of clypeus with no denticles, frons excavate, male with blade
like process on clypeus and small pronotal tubercles. A key for males of the Neotropical Pentodontini genera is
given and the current status of Ligyrus Burmeister and Tomarus Erichson is discussed and a checklist with the
known species included in each genus is also provided.
Additional key words: Ligyrus, Pentodontini, Saccharoscaptus, South America, taxonomy, Tomarus.
Resumen
M R MA, G PC. 2015. Revisión de Philoscaptus Brèthes, 1919 y descripción de un nuevo género
(Coleoptera, Melolonthidae, Dynastinae). E 30(1): 1-11.
Se revisa el género Philoscaptus Brèthes, 1919, que de acuerdo con el presente estudio es monotípico, que incluye
a P. bonariensis (Burmeister), especie distribuída en áreas abiertas de la cuenca del río Paraná, en Brasil, Uruguay,
Argentina y Paraguay (nuevo registro). El nuevo género Saccharoscaptus se describe para P. laminifer Dechambre,
el cual se distingue por tener mandíbulas con un diente apical poco redondeado, ápice del clípeo sin dentículos,
frente excavada, y los machos con un proceso laminar en el clípeo y pequeños tubérculos pronotales. Se incluye
una clave actualizada para los machos de los géneros de Pentodontini Neotropicales, y se discute el estado actual
de Ligyrus Burmeister, 1847 y Tomarus Erichson, 1847 con una lista actualizada de las especies asignadas a cada
uno de estos géneros.
Palabras clave adicionales: Ligyrus, Pentodontini, Saccharoscaptus, Sudamérica, taxonomía, Tomarus.

ENTOMOTROPICA 30(1): 1-11
2
Introduction
Pentodontini is a worldwide tribe of Dynastinae,
the American Pentodontini comprises 30
genera, being more diverse in South America
with 21 genera (Krajcik 2012), and almost all
lacking studies or comprehensive taxonomic
studies after the revisions of Endrodi (1969,
1985). Philoscaptus Bréthes, 1919 has been
known by two species, P. bonariensis (Burmeister,
1847) from open areas in Southern Brazil,
Uruguay, and Northern Argentina, while P.
laminifer Dechambre, 1979 was known from
unique male specimen collected during XIX
century with vague location “Para, Brazil“.
Philoscaptus as other genera in Pentodontini is
a small to medium sized brownish beetle with
body convex and simply striated elytra, and with
frontal small tubercles on head. The concise
morphological diagnosis makes it difficult to
identify the genus.
The status of Philoscaptus is reviewed for the
two species included in the genus. Based on a
series of morphological characters we transfer
P. bonariensis to a new genus here described,
add new distributional data and discus its
distribution and agricultural relevance. A key
to the American Pentodontini is incorporated
to and includes the genera Ligyrus Burmeister,
1847 and Tomarus Erichson, 1847, with
comments on the polemic synonymic status of
these genera proposed by Ratcliffe (2002) and a
preliminary checklist of the species assigned to
each one.
Material and methods
We use the classification of families of
Scarabaeoidea proposed by Endrödi (1966),
Morón (1997) and Cherman and Morón
(2014). Diagnostic characters and terminology
used in the descriptions follows in part Endrödi
(1969), Dechambre (1979) and Ratcliffe (1981).
A total of 24 specimens were studied by us from
the following collections:
CEMT: Coleção Zoológica do Mato Grosso,
Universidade Federal do Mato Grosso, Cuiabá,
Brazil (Fernando Zagury Vaz-de-Mello).
DZUP: Coleção Entomológica Padre Jesús
Santiago Moure, Universidade Federal do
Paraná, Curitiba, Paraná, Brazil (Lúcia Massutti
de Almeida).
EPGC: Everardo and Paschoal Grossi
Collection, Nova Friburgo, Rio de Janeiro,
Brazil.
FCUR: Facultad de Ciencias, Universidad de
la República. Montevideo, Uruguay (Enrique
Morelli).
IOC: Instituto Oswaldo Cruz, Rio de Janeiro,
Rio de Janeiro (Jane Margareth Costa).
MNRJ: Museu Nacional do Rio de Janeiro, Rio
de Janeiro, Brazil (Marcela Laura Monné).
MZUSP: Museu de Zoologia da Universidade
de São Paulo, São Paulo, Brazil (Sônia Aparecida
Casari).
MXAL: Miguel A. Morón Collection, Xalapa,
Veracruz, Mexico.
Philoscaptus Brèthes, 1919
Type species: Podalgus bonariensis Burmeister
1847: 118 (by monotypy).
Diagnosis. Mandibles exposed, with three
rounded teeth on outer border; apex of clypeus
narrowed, with two short denticles; frons convex
with small tubercle; pronotum widely convex;
sexual dimorphism only in pygidium and last
sternite.
Body: Stubby, somewhat oval, wider posteriorly;
dorsal surface convex. Head: Small with frontal
tubercle present, narrow, high, and weakly
incised; clypeus convergent, with apex bidentate
and elevate (Figure 1). Mandibles external
border clearly tridentate with upturned apex
(Figure 2); teeth rounded, inner tooth narrower
and longer. Maxilla slender, elongated, apex
narrowly rounded and with one or two hooklets

Moron rios, Coelho Grossi. Neotropical genera Philoscaptus and Saccharoscaptus
3
on internal margin of lacinia (Figure 3). Mentum
strongly tapering anteriorly, emarginated or not.
Antennae 10 joined, club short. Pronotum:
Small sized, less than half of elytral length,
narrower than both elytron together; surface
simply convex with no tubercle. Elytra: Shape
subparallel, distinctly diverging posteriorly and
wider than pronotum. Scutellum triangular,
with subequal sides, smooth or with scattered
punctures. Venter: Surface densely setose;
prosternal process elongate, blade-like, apex
rounded. Legs: Anterior tibiae tridentate,
teeth equidistant; protarsi not swollen in either
sex; middle tibiae slender with two external
carinae, proximal weakly marked, distal strongly
crenulated, apical border with four teeth, being
one longer ventrally; posterior tibiae with apical
border expanded, irregularly notched, with
3–4 truncate distal teeth. Sexual dimorphism:
Pygidium of male more convex than in female;
last sternite of female twice longer than in male.
Male genitalia: Parameres symmetric, stout,
elongate, curved ventrally (Figure 5).
Remarks. Philoscaptus includes only P.
bonariensis (Burmeister) and looks similar to
some species of Hylobothynus Ohaus, 1910, but
it can be distinguished from these by the last
segment of maxillary palps, not triangularly
expanded. It also can be confused with Neoryctes
Arrow, 1908, but it differs in the mandibles
broadly curved and basal margin of pronotum
absent or incomplete.
Philoscaptus bonariensis (Burmeister, 1847)
(Figures 1–5, 15-18)
Podalgus bonariensis Burmeister, 1847: 118
Material examined: BRAZIL: Rio Grande
do Sul, Pelotas, 30-x-1962 (3 females)
(DZUP and EPGC); São Leopoldo, no more
data (1 specimen) (MZSP). PARAGUAY:
Guairá, Villarrica, xii-1924 (2 males) (FIOC).
URUGUAY: Canelones, Las Piedras, 07-x-
1967 (1 male, 5 females) (CEMT, MXAL).
Diagnosis. Apex of clypeus with two small
denticles; frons convex with bifid or transverse
tubercle; pronotum widely convex in both sexes.
Redescription. Length: 16.8–20.3 mm. Width:
9.8–12.0 mm. Body: Wider posteriorly, 1.6–
1.7 longer than wide; sides weakly convergent
posteriorly (Figures 15-16). Color: Light to dark
brown, ventrally usually lighter. Head: 2.4–2.7
times narrower than pronotal width; frons with
dorsal surface flat to slightly concave. Apex of
distal maxillary palps subconical. Antennal club
short, as long as antennomeres 2–7 together.
Pronotum: Surface completely punctate,
densely at pronotal declivity; punctures rounded,
c-shaped and sometimes converging; anterior
border with wide margin and translucent blade;
posterior margin absent. Elytra: Surface convex
and moderately punctate by rows of shallow and
“o” shape punctures; elytral epipleural margin
ascending and reaching humeri. Abdomen:
Prepygidium posterior border with scarce
rugosities (stridulatory surface). Pygidium 2.2
times wider than long, apical border slightly
prominent in female, widely rounded in male.
Male genitalia: Parameres with rounded apex,
briefly toothed on lateral borders (Figures 4–5,
17-18).
Distribution. Argentina: Buenos Aires
(Buenos Aires); Santa Fé (Santa Fé). Paraguay:
Guairá (Villarrica) (New Country Record).
Brazil: Rio Grande do Sul (Pelotas and São
Leopoldo). Uruguay: Canelones (Colonia,
Tacuarembo) (Figure 13).
Biology. According to Morelli (1997) the
larvae of P. bonariensis live under soil of native
grasslands and gramineous crops in Uruguay,
feeding on roots and organic debris, and may
be located in vertical galleries at 25 cm depth.
Second and third instar were collected at the
same time with young adults in the soil during
the same season that suggests overlapping of 2
year life cycle generations.

ENTOMOTROPICA 30(1): 1-11
4
Saccharoscaptus Morón & Grossi n. gen.
(Figures 6–12)
Type species: Philoscaptus laminifer Dechambre,
1979.
Diagnosis. Mandibles exposed, with one weak
rounded apical tooth; apex of clypeus narrowly
rounded, with no denticles; frons excavate; male
with blade like process on clypeus and one pair
of small pronotal tubercles.
Description. Body: Stubby, suboval, wider
posteriorly; dorsal surface convex. Head: Small;
frontal carina absent, frons excavate; clypeus
convergent, rounded and weakly reflexed
(Figure 6). Mandibles almost flat, with external
edge notched and with one apical rounded
tooth (Figure 7). Maxillae slender, elongated,
apex digitiform, without hooklets on internal
margin of lacinia (Figure 8). Mentum weakly
tapering anteriorly. Antennae 10 joined.
Pronotum: Small sized, less than half of elytral
length, narrower than both elytron together.
Elytra: Shape subparallel, distinctly diverging
posteriorly and wider than pronotum. Scutellum
triangular, with subequal sides and v-shaped
deep impression. Venter: Surface densely setose;
Figures 1-10. Philoscaptus bonariensis: head dorsal view (1), right mandible dorsal view (2), right maxilla dorsal view (3),
parameres distal view (4), parameres lateral view (5). Saccharoscaptus laminifer:head dorsal view (6), right mandible dorsal
view (7), right maxilla dorsal view (8), parameres distal view (9), parameres lateral view (10). Scale lines 1 mm.

Moron rios, Coelho Grossi. Neotropical genera Philoscaptus and Saccharoscaptus
5
prosternal process strong, elongated, nearly
semi-columnar, apex rounded. Legs: Anterior
tibiae tridentate, teeth equidistant, protarsi not
swelling in both sexes; middle tibiae slender with
two external carinae, proximal weakly marked,
distal moderately crenulated, apex with 4–5
teeth, being one longer laterally; posterior tibiae
with apex nearly truncate, briefly expanded, with
7–8 small teeth. Sexual dimorphism: Strongly
evident on head and pronotum. Male genitalia:
Parameres symmetric, flattened, elongate, and
curved ventrally (Figure 10).
Remarks. Saccharoscaptus can be confused in a
first view with some species of Bothynus Hope,
1837, but is readily distinguish from it by its
mandibles only weakly notched externally, and
strongly tridentate in Bothynus. This character
also differentiates Saccharoscaptus from
Philoscaptus, as well as the clypeus not dentate
and a bituberculated pronotum. This genus
includes only S. laminifer (Dechambre).
Etymology. Derived from the Greeks words
saccharon, sugar and scapto to dig ( Jaeger
1955) in reference to the larval habits recently
observed in Bolivia (Copa-Bazan and Morón
2014), where they feed on roots of sugar cane.
Saccharoscaptus laminifer (Dechambre, 1979)
new combination
(Figures 6-12, 19-22)
Philoscaptus laminifer Dechambre 1979: 103
Examined material. BRAZIL: Pará,
Jacareacanga, ii-1970 (2 males) (DZUP and
EPGC); BOLIVIA: Santa Cruz, Warnes, El
Naranjal, La Jupia, 308 m, viii-2011 (5 males, 5
females) (MXAL, EPGC).
Diagnosis. Clypeus with rounded apex and
blade-like preapical projection in male; frons
widely concave in male, and scarcely concave in
female; pronotum widely convex in female, with
small pair of tubercles in male.
Redescription. Length: 19.1–22.4 mm. Width:
9.5–10.4 mm. Body: Wider posteriorly, 2.0–
Figures 11-12. Dorso lateral view of head and pronotum
of Saccharoscaptus laminifer: male (11), female (12). Scale
lines 1 mm.
Figure 13. Distribution of Philoscaptus bonariensis.
Figure 14. Distribution of Saccharoscaptus laminifer.

ENTOMOTROPICA 30(1): 1-11
6
Figures 15-18. Philoscaptus bonariensis, dorsal habitus of male from Villarica, Paraguay (15), distal view of parameres (16),
lateral view of parameres (17), dorsal habitus of female from Pelotas, Brazil (18). Scale lines 5 mm, except gs. 15-16 =
1 mm.

Moron rios, Coelho Grossi. Neotropical genera Philoscaptus and Saccharoscaptus
7
Figures 19-22. Saccharoscaptus laminifer male from Jacareacanga, Brazil, dorsal habitus (19), lateral habitus (20), distal
view of parameres (21), lateral view of parameres (22). Scale lines 5 mm, except gs. 21-22 = 1 mm.

ENTOMOTROPICA 30(1): 1-11
8
2.2 longer than wide; sides weakly convergent
posteriorly (Figures 19-20). Color: Dark brown
to black, ventrally dark reddish brown. Head:
2.2–2.3 times narrower than pronotal width;
dorsal surface widely concave in male to slightly
concave in female (Figures 11–12). Apex of
distal maxillary palpus rounded. Antennal
club slightly longer than antennomeres 2–7
together in male; slightly shorter than same
antennomeres in female. Pronotum: Shape
convex, with scattered deep, round punctures,
denser in females, males with two weak tubercles
near the middle of anterior half; anterior
border with wide margin and translucent
blade; posterior margin absent. Elytra: Surface
convex and moderately punctate by rows of
deep and ocellate punctures; elytral epipleural
margin ascending and weakly reaching humeri.
Abdomen: Prepygidium densely covered with
small, shallow punctures, without rugosities on
the middle of posterior border. Pygidium 2.7
times wider than long, apical border slightly
prominent in female, widely rounded in male.
Parameres with flattened, acute apex, strongly
toothed on lateral borders (Figures 9–10, 21-
22).
Remarks. This species was originally described
in Philoscaptus in which it clearly does not
belong based on the character commented above
(Table 1). Female is examined by the first time,
and distinctive characters are commented in the
diagnosis and redescription. Sexual dimorphism
is much more accentuated than in Philoscaptus.
Distribution. Brazil (Pará); Bolivia (Santa
Cruz) (New Coutry Record). (Figure 14).
Biology. According to Copa-Bazan and Morón
(2014) during 2011 an important increase in the
larval populations of S. laminifer living in the soil
of sugar cane fields in northern of department
of Santa Cruz, Bolivia was recorded. Important
damage to the roots of sugar cane in a wide
surface of such fields was observed, with 150–
200 larvae/m2 in combination with an unusual
dry season. Regions of Jacareacanga, Pará and
Santa Cruz are separated by more than 1 300
km in straight line, so that, it is much probable
that this species has a wide distribution along
the southern part of the Amazonian biome.
Key to genera of american Pentodontini based on
males
1. Propygidium with stridulatory surface ............. 2
1’ Propygidium without stridulatory surface .........
...................................................................... 10
2 Pygidium much shorter than prepygidium ........
........................................................................ 3
2’ Pygidium as long as prepygidium or longer than
it ...................................................................... 4
3 Foretibia tridentate. Pronotum with apical
tubercle and pit. Apex of clypeus narrowed, with
2-4 teeth ........................... Bothynus Hope, 1837
Characters S. laminifer P. bonariensis
External border of mandibles Rounded with weak notch With 3 rounded teeth
Apex of clypeus Rounded, briey projected Narrowed with 2 short teeth
Male pronotum With 2 small tubercles Convex
Male clypeus With blade like projection Without projections
Male frons Broadly excavate Convex, with transverse tubercle
Female frons Shallowly excavate Convex, with transverse tubercle
External border of paramere With 2 well dened teeth; distal part
expanded
Sinuate, with weak tooth; distal part
narrowed
Table 1. Dierential characters of adults of Philoscaptus and Saccharoscaptus.

Moron rios, Coelho Grossi. Neotropical genera Philoscaptus and Saccharoscaptus
9
3’ Foretibia bidentate. Pronotum simply convex.
Apex of clypeus narrowly rounded ....................
..............................................Heteroconus Kolbe,
1900 (= Parabothynus Endrödi, 1968).
4 Body length less than 15 mm ............................
...........................Heteronychus Burmeister, 1847
4’ Body length greater than 15 mm. ................. ..5
5 Pronotum with wide hollow and rounded
tubercles .................... Gillaspytes Howden, 1980
5’ Pronotum simply convex ................................. 6
6 Antenna with 9 segments. Frons with short horn
in males; females with frons convex ...................
.............................. Eremobothynus Ohaus, 1910
6’ Antenna with 10 segments. Frons with transverse
carina or tubercles ............................................ 7
7 External border of mandibles with three teeth or
rounded lobes .................................................. 8
7’ External border of mandibles simply curved ... 9
8 Apical segment of maxillary palp expanded,
triangular .................. Hylobothynus Ohaus, 1910
8’ Apical segment of maxillary palp cylindrical .......
..................................Philoscaptus Brèthes, 1919
9 Pronotum with basal margin complete ..............
........................................ Neoryctes Arrow, 1908
9’ Pronotum without basal margin ........................
............................Indieraligus Dechambre, 1979
10 Mandibles exposed dorsally ........................... 11
10’ Mandibles mostly hidden under clypeus. ....... 26
11 External border of mandibles without
projections ..................................................... 12
11’ External border of mandibles with 2-3 teeth, or
wide lobes ...................................................... 21
12 Clypeus with transverse carina immediately
behind apex of clypeus ................................... 13
12’ Clypeus without transverse carina behind apex of
clypeus ........................................................... 14
13 Clypeal carina tridentate. Body length 15-17
mm .............................. Aphonus LeConte, 1856
13’ Clypeal carina entire or sinuate. Body length 18-
30 mm ........................Orizabus Fairmaire, 1878
14 Pronotum simply convex, without tubercle or
fovea .............................................................. 15
14’ Pronotum convex, but with fovea or protuberances
on anterior half .............................................. 20
15 Frons simply convex ...................................... 16
15’ Frons convex with protuberances .................. 18
16 Fronto-clypeal keel absent. Body length 13-15
mm. ..................................Euetheola Bates, 1888
16’ Fronto-clypeal keel transverse or swelling
present. Body length 20-23 mm .................... 17
17 Elytra convex with striate surface. Abdominal
spiracles small ................. Aphonides Rivers, 1889
17’ Elytra swollen with surface nearly smooth.
Abdominal spiracles large ..................................
..................................... Gorditus Ratclie, 2010
18 Frons with prominence on disc ...................... 19
18’ Frons with two tubercles on fronto-clypeal keel
...............................Denhezia Dechambre, 2006
19 Frons with a small truncate tubercle ..................
........................Endroedianibe Chalumeau, 1981
19’ Frons with a conical acuminate tubercle ............
................................. Hiekeianus Endrödi, 1978
20 Head with dimorphic structures. Pronotum with
small or large tubercles, but without subapical
fovea .............................................................. 21
20’ Head similar in both sexes. Pronotum with small
apical tubercle and subapical fovea ....................
..............Tomarus Erichson, 1847, revised status
21 Male with conical horn on head and bifurcate
tubercle on pronotum. Female when known with
small tubercle on clypeus ............................... 22
21’ Male with truncate, short horn on clypeus, and
concave frons. Female with transverse keel on
clypeus ...................... Saccharoscaptus, new genus
22 Clypeus with a triangle shaped apex, truncate.
Elytra without distinct striae and punctures ......
......... Heteroglobus Dupuis & Dechambre, 2006
22’ Clypeus with a triangle shaped apex, emarginated
or bidentated. Elytra with distinct striae and
punctures .................ronistes Burmeister, 1847
23 Clypeal apex acute, unidentate ..........................
...................................... Oxygrylius Casey, 1915
23’ Clypeal apex bidentate or rounded ................ 24
24 External border of mandibles with rounded
lobes .............................. Barutus Ratclie, 1981
24’ External borders of mandibles with 3 teeth ... 25
25 Pronotum convex, but with small apical tubercle
and subapical fovea. Apex of clypeus narrowed .
............ Ligyrus Burmeister 1847, revised status.
25’ Pronotum simply convex. Apex of clypeus
bidentate .............................. Aceratus Prell, 1936
26 Clypeal apex truncate or rounded .................. 27
26’ Clypeal apex acute .........Oxyligyrus Arrow, 1908
27 Posterior femur greatly thickened ......................
................................... Pentodina Endrödi, 1968
27’ Posterior femur not thickened ....................... 28
28 Apex of clypeus rounded ............................... 29

ENTOMOTROPICA 30(1): 1-11
10
28’ Apex of clypeus broadly truncate................... 30
29 Center of head with strong, transverse keel .......
................... Collagenus Ratclie & Hardy, 2005
29’ Center of head slightly tumid............................
................................. Coscinocephalus Prell, 1936
30 Head lacking horns of tubercles ........................
....................................... Parapucaya Prell, 1934
30’ Head with small horns or tubercles next to eyes.
Pronotum declivous anteriorly at middle ...........
...........................................Pucaya Ohaus, 1910
e genera Ligyrus and Tomarus
Following Escalona and Joly (2006) we disagree
with the synonymy of Ligyrus Burmeister, 1847
under Tomarus Erichson, 1847 according with
the month of publication (Ratcliffe 2002),
because such emendation is unnecessary,
is not justified with names in wide usage
during decades, and cause confusion instead
to promote stability (International Code of
Zoological Nomenclature 1999: article 23).
But the study of many species allowed us to
confirm that both names really correspond to
related but distinct genera. The morphological
characters traditionally used in keys and generic
descriptions, as well as ambiguous taxonomic
criteria, offer dichotomical options to reach the
genus Ligyrus by two different ways (Ratcliffe
1981, Ratcliffe and Hardy 2005), results that in
part support the existence of two valid genera.
Detailed systematic studies based on adults
and larvae are necessary to support an accurate
revision of both genera, but in the meantime a
preliminary checklist is proposed as follows.
Tomarus Erichson, 1847
Type species Scarabaeus ebenus DeGeer, 1774.
Outer side of mandibles simply curved;
pronotum with a pit; larva undescribed.
Group A: Protarsi enlarged.
Tomarus ebenus (DeGeer 1774:317).
T. similis (Endrödi 1968: 166).
Group B: Protarsi normal.
Tomarus bituberculatus (Palisot de Beauvois
1805: 103).
T. maimon Erichson 1847:96 revised status.
T. gyas Erichson 1848:561 revised status.
T. laevicollis (Bates 1888:316) new
combination
T. subtropicus Blatchley 1922:30 new
combination.
T. maternus Prell 1937:89 new combination.
T. pumilus Prell 1937:90 new combination.
T. selanderi Cartwright 1959:537 new
combination.
T. rosettae Endrödi 1968:168 new
combination.
T. discrepans Escalona & Joly 2006:125 new
combination.
Ligyrus Burmeister 1847
Type species: Scarabaeus gibbosus DeGeer, 1774
Outer side of mandibles with 2 teeth
Group A (or subgenus Ligyrus s.str.). Pronotum
with or without pit; protarsi normal; larva
without septula on raster.
Ligyrus gibbosus (DeGeer 1774: 322).
L. cuniculus (Fabricius 1801: 1801).
L. fossor (Latreille 1833: 11).
L. villosus (Burmeister 1847: 120).
L. nasutus (Burmeister 1847:120).
L. neglectus (LeConte1847: 87).
L. rubripes (Boheman 1858: 57).
L. burmeisteri Steinheil 1872: 560.
L. bidentulus Fairmaire 1892: 244.
L. scitulus Casey 1915: 20.
L. cicatricosus Prell 1937: 90.
L. pullus Prell 1937: 89 new combination
L. gianucai Dechambre & Lumaret 1985:
107.
L. adoceteus Ratcliffe & Cave 2010: 7 new
combination.

Moron rios, Coelho Grossi. Neotropical genera Philoscaptus and Saccharoscaptus
11
Group B (or subgenus Ligyrodes Casey, 1915).
Pronotum without pit; protarsi enlarged; Larva
with septula on raster.
L. relictus (Say 1825: 194).
L. sallaei Bates 1888: 318.
L. peruvianus Endrödi 1970: 106.
Acknowledgements
Fernando Copa (IIAV, Santa Cruz, Bolivia) and
Jorge Frana (INTA, Rafaela, Argentina) send
to MAM samples for study. Sara L. Rivera-
Gasperín (Xalapa, Veracruz) provided the
illustrations in figures 1-12. PCG is supported
by a grant from UEMA of the Profix program.
Caio Antunes and Hingrid Yara (MZUSP)
are thanked for images of the specimens of
Philoscaptus from that institution.
Collection managers cited are thanked for the
loaned material which forms the basis of this
study. This paper is a contribution to the research
line “Coleopteros Lamelicornios de América
Latina” supported by Instituto de Ecología, A.
C. (account 902-08-011).
References
C, MA, M MA. 2014. Validación de la
familia Melolonthidae Leach, 1819 (Coleoptera
Scarabaeoidea). Acta Zoologica Mexicana 30(1): 201–
220.
C-B F, M M A. 2014. Importancia
del gusano blanco (Coleoptera: Melolonthidae;
Dynastinae) en cultivos de caña de azúcar en Santa
Cruz, Bolivia. pp. 31–44. En: Pérez-Domínguez, J. F.
y A. Aragón (eds.).Memoria IX Mesa Redonda sobre
Plagas Subterráneas. Guadalajara, Jalisco, México.
Publicación especial INIFAP y BUAP, México.
D RP. 1979. Nouveaux Dynastidae
Pentodontini américains (Coleoptera Scarabaeoidea).
Revue Francaise d’Entomologie 1: 101–105.
E S. 1966. Monographie der Dynastinae (Col.
Lam.) I Teil. Entomologische Abhandlungen 33: 1–457.
E S. 1969. Monographie der Dynastinae. 4.
Tribus: Pentodontini (Coleoptera, Lamellicornia).
Entomologische Abhandlungen 37: 1–145.
E S. 1985. e Dynastinae of the World. Dr. W.
Junk Publ. Dordrecht. 800 pp.
E H E, J L J. 2006. El género Ligyrus
Burmeister, 1847 en Venezuela (Coleoptera:
Scarabaeidae: Dynastinae: Pentodontini). Boletín de
la Sociedad Entomológica Aragonesa 39: 111–137.
ICZN. 1999. International Code of Zoological
Nomenclature, 4th edition. International Trust for
Zoological Nomenclature, London. 306 pp.
J EC. 1955. A source-book of biological names and
terms. ird Edition. Charles C. omas Publisher,
Springeld, Illinois. 323 pp.
K M. 2012. Checklist of the world Scarabaeoidea.
Animma. X supplement 5. Plzen, Czech Republic.
278 pp.
K FT. 2002. Case 3202 Podalgus Burmeister, 1847
and Philoscaptus Bréthes, 1919 (Insecta, Coleoptera):
proposed conservation by the designation of Podalgus
cuniculus Burmeister, 1847 as the type species of
Podalgus. Bulletin of Zoological Nomenclature 59(2):
93–95.
M E. 1997. Descripción de los estados inmaduros
y notas sobre la biología de Diloboderus abderus Sturm
y Philoscaptus bonariensis Burmeister (Coleoptera:
Melolonthidae: Dynastinae). Acta Zoológica Mexicana
71: 57–70.
M MA. 1997. Capítulo 1. Antecedentes. [pp.1-8].
En: Atlas de los Escarabajos de México. Coleoptera
Lamellicornia. Vol. I. Familia Melolonthidae. (M.
A. Morón, B. C. Ratclie y C. Deloya, editores)
CONABIO y Sociedad Mexicana de Entomología,
México. 280 pp.
R BC. 1981. Barutus hartmanni, a new genus and
species from Panama with a key to the New World
genera of Pentodontini (Coleoptera: Scarabaeidae:
Dynastinae). e Coleopterists Bulletin 35: 463–472.
R BC. 2002. Dynastinae Mac Leay 1819 pp.64–
67. In: American beetles, Volume 2 (R. H. Arnett, M.
omas, P. E. Skelley and J. H. Frank, editors) CRC
Press, Boca Raton, FL. 861 pp.
R B C, H M. 2005. Collagenus dasysternus,
a new genus and species of Dynastinae from
eastern Venezuela with a key to the New World
genera of Pentodontini (Coleoptera: Scarabaeidae:
Dynastinae). e Coleopterists Bulletin 549(1): 143–
150.
Recibido: 31-01-2014.
Aceptado: 09-07-2014.