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ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2016 Magnolia Press
Zootaxa 4072 (3): 343
–
357
http://www.mapress.com/j/zt/
Article
343
http://doi.org/10.11646/zootaxa.4072.3.4
http://zoobank.org/urn:lsid:zoobank.org:pub:A358F7A0-12B8-401D-B6FE-ADBF1469B786
A revised phylogenetic classification of the ant subfamily Formicinae
(Hymenoptera: Formicidae), with resurrection of the genera
Colobopsis and Dinomyrmex
PHILIP S. WARD
1,4
, BONNIE B. BLAIMER
2
& BRIAN L. FISHER
3
1
Department of Entomology and Nematology, University of California at Davis, Davis, CA 95616, USA. E-mail: psward@ucdavis.edu
2
Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560, USA.
E-mail: bonnieblaimer@gmail.com
3
Department of Entomology, California Academy of Sciences, San Francisco, CA 94118, USA. E-mail: bfisher@calacademy.org
4
Corresponding author. E-mail: psward@ucdavis.edu
Abstract
The classification of the ant subfamily Formicinae is revised to reflect findings from a recent molecular phylogenetic study
and complementary morphological investigations. The existing classification is maintained as far as possible, but some
tribes and genera are redefined to ensure monophyly. Eleven tribes are recognized, all of which are strongly supported as
monophyletic groups: Camponotini, Formicini, Gesomyrmecini, Gigantiopini, Lasiini (= Prenolepidii syn. n.), Melo-
phorini (= Myrmecorhynchini syn. n.; = Notostigmatini syn. n.), Myrmelachistini stat. rev. (= Brachymyrmicini syn. n.),
Myrmoteratini, Oecophyllini, Plagiolepidini, and Santschiellini stat. rev. Most of the tribes remain similar in content, but
the generic composition of Lasiini, Melophorini, and Plagiolepidini is changed substantially. Species that have been
placed in the genus Camponotus belong to three separate lineages. To ensure monophyly of this large, cosmopolitan genus
we institute the following changes: Colobopsis and Dinomyrmex, both former subgenera of Camponotus, are elevated to
genus level (stat. rev.), and two former genera, Forelophilus and Phasmomyrmex, are demoted to subgenus status (stat.
n. and stat. rev., respectively) under Camponotus; two erstwhile subgenera of Phasmomyrmex, Myrmorhachis and Myr-
macantha, become junior synonyms (syn. n.) of Camponotus (Phasmomyrmex); and the Camponotus subgenus Myrmog-
onia becomes a junior synonym (syn. n.) of Colobopsis. Dinomyrmex, represented by a single species from southeast Asia,
D. gigas, is quite distinctive, but Camponotus and Colobopsis exhibit more subtle differences, despite being well separated
phylogenetically. We identify morphological features of the worker caste that are broadly useful for distinguishing these
two genera. Colobopsis species on the islands of New Caledonia and Fiji—regions with few native Camponotus species—
tend to exceed these diagnostic bounds, but in this case regionally applicable character differences can be used to distin-
guish the two clades. Despite confusing similarities in the worker caste Colobopsis and Camponotus retain diagnostic dif-
ferences in their larvae and pupae.
Key words: ant taxonomy, phylogenomics, morphology, convergence, divergence, Camponotus
Introduction
The ant subfamily Formicinae is a large and successful group, comprising about 3030 described species, distributed
globally across a wide range of terrestrial environments (Brown 2000; Bolton 2003; AntCat 2015). The subfamily
includes such well-known taxa as wood ants and their relatives (Formica), carpenter ants (Camponotus), weaver
ants (Oecophylla), and honeypot ants (Myrmecocystus), and a diverse array of about fifty other genera. The females
(workers and gynes) of this subfamily are readily distinguished from all other ants by the presence of an acidopore, a
nozzle-shaped structure at the apex of the seventh abdominal sternum used to spray formic acid (Bolton 1994).
Formicine workers have a flexible promesonotal suture (secondarily immobile in a few taxa), closed metacoxal
cavities, single petiolar node, complete tergosternal fusion of the petiole (second abdominal segment), and no
functional sting; abdominal segments 4–6 are very large relative to the sternites, which they overlap laterally and
usually also ventrally (Bolton 2003). A diagnosis of Formicinae males is provided by Boudinot (2015).
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Early attempts at a higher classification of Formicinae (e.g., Forel 1912; Emery 1925) were based largely on
the features of the proventriculus, an internal organ regulating movement of liquids from the crop to the midgut
(Eisner 1957). A more synthetic approach, involving a larger suite of morphological characters, was initiated by
Agosti (1991) and further developed and expanded by Bolton (2003), who recognized nine tribes and two informal
tribe groups: the lasiine tribe group (with Lasiini, Myrmoteratini and Plagiolepidini) and the formicine tribe group
(with Camponotini, Formicini, Gigantiopini, Melophorini, Notostigmatini and Oecophyllini). Two tribes,
Gesomyrmecini and Myrmecorhynchini, were unplaced to tribe group. This classification has been accepted to the
present day, with 51 extant genera being distributed among these eleven tribes (AntCat 2015).
The first molecular phylogenetic analysis of Formicinae, based on two mitochondrial genes (Johnson et al.
2003), yielded results that were mostly concordant with the Bolton (2003) classification, but taxon and gene
sampling was quite limited. Later studies based on multiple nuclear genes and more extensive taxon sampling
(Brady et al. 2006; Moreau et al. 2006; Moreau & Bell 2013) indicated that some of the recognized formicine taxa,
such as the genus Camponotus and tribes Lasiini and Plagiolepidini, are not monophyletic. A recent study of
Formicinae, employing a phylogenomic data set of almost 1000 genes and a broad sampling of taxa, has given a
much more comprehensive and robust picture of the evolution of this group (Blaimer et al. 2015). We propose a
revised classification of the subfamily based on the results of this study and on complementary morphological
investigations of the tribe Camponotini.
Material and methods
Taxon sampling, gene sampling, and methods of phylogenetic analysis are detailed in Blaimer et al. (2015). For the
phylogenomic study we sequenced 959 UCE (ultraconserved element) loci in 82 formicines, representing 48 of 51
currently recognized genera, and in eight outgroup taxa. For the same set of taxa we also generated a more
“traditional” dataset of 10 nuclear genes, by Sanger sequencing. Bayesian and maximum likelihood (ML) analyses
were carried out on both datasets, yielding similar results but with notably better resolution and stronger node
support in the phylogenomic tree, which we use as the reference phylogeny.
Morphological analysis focused on the tribe Camponotini, in which we introduce several genus-level changes.
We examined specimens of approximately 85 species of Camponotus (Colobopsis) and about 200 species of
Camponotus sensu stricto, in an effort to discern diagnostic differences between the two lineages. Most specimens
were examined directly but we also took advantage of images, especially on AntWeb (www.antweb.org), to
scrutinize taxa for which direct examination was not possible.
The following metric measurements and indices were employed:
HW Head width: maximum width of head, excluding the eyes.
HL Head length: midline length of head from the posterior margin to a line across the anterior clypeal
margin (medial indentations on either margin do not decrease length).
EL Eye length: length of eye measured in a full-face view of the head.
ASM Minimum distance between the antennal sclerites (inter-torular distance).
CLW Clypeus width: width of clypeus, taken at the anterior tentorial pits.
CLL Clypeus length: maximum measurable length of clypeus, taken along the midline, from a line drawn
across posterior margin to a line across the anterior margin (medial indentations on either margin do not
decrease length).
CI Cephalic index: HW/HL
REL Relative eye length: EL/HL
The first three measurements were taken in a full-face (dorsal) view of the head, with the posterior margin of
the head and the anterior clypeal margin in the same focal plane. The last three measurements (ASM, CLW, CLL)
were taken in an anterodorsal view of the head, such that the measurement of clypeus length was maximized.
Voucher specimens from molecular and morphological studies are deposited in UCDC (Bohart Museum of
Entomology, University of California, Davis), CASC (California Academy of Sciences, San Francisco) and USNM
(National Museum of Natural History, Washington, DC).
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REVISED CLASSIFICATION OF FORMICINAE
Results
Phylogeny. The new phylogeny of the Formicinae (Figure 1) reveals six strongly supported, species-rich clades,
and five other species-poor, long-branched lineages whose positions in the tree are less certain. This pattern is
obtained with both the 10-gene and 959-gene data sets, but the latter provides greater resolution across the tree and
stronger branch support (Blaimer et al. 2015). Of the six well-supported clades, Myrmelachistini is sister to all
other Formicinae, and Lasiini is consistently recovered as sister to the remaining taxa. The phylogenomic data
support Melophorini as the next branch in this series, but the relationships among the three remaining groups
(Plagiolepidini, Formicini, and Camponotini) are less clear. Phylogenetic relationships within each of these six
clades are quite well resolved, however, with most nodes having 100% bootstrap support.
The five taxonomically isolated lineages correspond to the genera Gesomyrmex, Gigantiops, Myrmoteras,
Oecophylla, and Santschiella. The phylogenomic data do suggest partial resolution of their positions in the
formicine tree, including placement of Myrmoteras as sister to Camponotini, and a sister-group relationship of
Oecophylla and Gesomyrmex as well as Gigantiops and Santschiella (Figure 1). All of these taxa are situated on
long branches, however, and the nodes where they join the tree are subtended by very short branches. Some of the
putative sister group relationships might therefore be artifacts of long-branch attraction (Bergsten 2005), base
frequency heterogeneity (Jermiin et al. 2004), or other confounding factors.
Revised tribal classification. The tribal classification of Formicinae is modified in accordance with these
molecular phylogenetic results. The new classification was briefly outlined in Blaimer et al. (2015) and is here
given a more formal treatment. We strive to maintain the existing classification as far as possible, while also
ensuring that all recognized tribes are monophyletic. The major clades are treated as six tribes (Camponotini,
Formicini, Lasiini, Melophorini, Myrmelachistini, Plagiolepidini) whose composition is discussed below. The five
taxonomically isolated genera are each assigned to their own tribe; this is a cautionary approach, justified by the
uncertainty of their relationships to one another, and to other Formicinae. The resulting schema comprises 11
tribes, with high confidence in the monophyly of each one. This provides a framework for future work on the
morphological and social characteristics of each clade.
We cite the author and year of publication of each tribe name; equivalent information for genus names is
available in AntCat (http://antcat.org/). Genera known only from fossils are signified with a dagger (†); most of
these are unplaced to tribe and treated as incertae sedis within the subfamily.
Tribe Camponotini Forel 1878
= Polyrhachidini Ashmead 1905
Genera: Calomyrmex, Camponotus, †Chimaeromyrma, Colobopsis, Dinomyrmex, Echinopla, Opisthopsis,
Overbeckia, Polyrhachis, †Pseudocamponotus.
Comments. The composition of this tribe remains unchanged, although some generic boundaries have been
modified (see below). Overbeckia, not sequenced in this study, is likely a junior synonym of Camponotus (Bolton
2003). All members of this tribe have a unique, vertically inherited bacterial symbiont, Blochmannia, whose
evolutionary history mirrors that of the ants (Wernegreen et al. 2009). Morphologically the workers of
Camponotini can be recognized by the combination of distinctive mandibular dentition (5–8 teeth, with the third
tooth from apex not reduced in size), antennal insertions well separated from the posterior clypeal margin, and
twelve antennal segments (Bolton 1994, 2003).
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FIGURE 1. Phylogeny of the ant subfamily Formicinae based on 959 UCE (ultraconserved element) loci (Blaimer et al. 2015).
Support values are maximum likelihood bootstrap percentages. The tree depicts six major clades, here treated as tribes, and five
isolated genera subtended by long branches (Gesomyrmex, Gigantiops, Myrmoteras, Oecophylla, and Santschiella, in bold
font). Note that species assigned to Camponotus occur as three separate lineages within the tribe Camponotini, and that
Phasmomyrmex and Forelophilus are embedded within Camponotus (sensu stricto).
Tribe Formicini Latreille 1809
Genera: Alloformica, Bajcaridris, Cataglyphis, †Cataglyphoides, †Conoformica, Formica, Iberoformica,
Polyergus, Proformica, †Protoformica, Rossomyrmex.
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Comments. This is another distinctive group whose composition is unchanged. These ants are characterized
by closely approximated metacoxae, elliptical to slit-shaped propodeal spiracle, presence of ocelli in workers, and
a double row of stout setae on the metatibia (Bolton (2003). This clade is mostly confined to the Palearctic and
Nearctic regions.
Tribe Gesomyrmecini Ashmead 1905
= Dimorphomyrmii Emery 1895
= Gesomyrmini Forel 1912
= †Sicelomyrmicini Wheeler 1929
Genera: Gesomyrmex, †Prodimorphomyrmex, †Sicilomyrmex.
Comments. The tribe Gesomyrmecini is here restricted to Gesomyrmex and two similar fossil taxa (Wheeler
1915). Bolton (2003) also placed Santschiella in Gesomyrmecini, but the molecular results do not support a close
relationship between Gesomyrmex and Santschiella (Blaimer et al. 2015). The similarities between the two—very
large eyes, widely separated antennal insertions, and scapes that pass below the eyes (Bolton 2003)—must be
interpreted as due to convergence.
Tribe Gigantiopini Ashmead 1905
Genus: Gigantiops.
Comments. This remains a monotypic tribe, represented by a single taxonomically isolated species,
Gigantiops destructor, restricted to northern South America. The molecular phylogeny recovers Gigantiops as
sister to Santschiella (Figure 1), a monotypic African genus, but support for this relationship is not strong, so we
retain a separate tribe for Santschiella. Although both genera share distinctively large eyes, they differ markedly in
configuration of the frontoclypeal complex, with the antennal insertions of Gigantiops being located close to one
another and between the eyes (Bolton 2003).
Tribe Lasiini Ashmead 1905
= Acanthomyopsini Donisthorpe 1943
= Prenolepidii Forel 1912 syn. n.
Genera: Cladomyrma, Euprenolepis (tribal transfer), †Glaphyromyrmex, Lasius, Myrmecocystus, Nylanderia
(tribal transfer), Paraparatrechina (tribal transfer), Paratrechina (tribal transfer), Prenolepis (tribal transfer),
Pseudolasius (tribal transfer), Zatania (tribal transfer).
Comments. Apart from retention of Cladomyrma, Lasius and Myrmecocystus, the composition of this tribe is
greatly changed. It is now comprised of those three genera and a cluster of taxa known as the Prenolepis genus
group. Lasius and Myrmecocystus are each other’s closest relatives, and in turn they are the sister group of the
Prenolepis genus group. The latter has been the subject of recent phylogenetic and taxonomic studies by LaPolla
and colleagues (e.g., LaPolla et al. 2010, 2012).
Tribe Melophorini Forel 1912
= Myrmecorhy nchini Wheeler 1917 syn. n.
= Notostigmatini Bolton 2003 syn. n.
Genera: Lasiophanes (tribal transfer), Melophorus, Myrmecorhynchus (tribal transfer), Notoncus (tribal transfer),
Notostigma (tribal transfer), Prolasius (tribal transfer), Pseudonotoncus (tribal transfer), Stigmacros (tribal
transfer), Teratomyrmex (tribal transfer).
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Comments. This tribe formerly contained Melophorus only, but that genus is now known to be embedded
within a larger, well-supported clade of ants (Figure 1), which is confined to southern South America, Australia,
New Zealand, and New Guinea, and for which the oldest available tribal name is Melophorini. A satisfactory
morphological circumscription of this group awaits further study.
Tribe Myrmelachistini Forel 1912 stat. rev.
= Brachymyrmicini Emery 1925 syn. n.
Genera: Brachymyrmex (tribal transfer), Myrmelachista (tribal transfer).
Comments. These two genera form a robustly supported clade that is sister to all other Formicinae (Figure 1).
Shared worker characters include a reduced antennal count (9–10 segments), five mandibular teeth, petiole inclined
anteriorly and/or with long posterior peduncle, and anterior tergosternal fusion of the third abdominal segment
(Bolton 2003). Both genera have the plesiomorphic palp formula 6,4 but this has been reduced to five maxillary
palp segments in some Myrmelachista species (Longino 2006).
Tribe Myrmoteratini Emery 1895
= Myrmoteratini Forel 1912
Genus: Myrmoteras.
Comments. This is another isolated genus, with numerous distinctive features including elongate, trap-jaw
mandibles and very large eyes (Bolton 2003). It appears to be sister to the tribe Camponotini (Figure 1), but this
requires confirmation.
Tribe Oecophyllini Emery 1895
= Oecophyllini Forel 1912
Genus: Oecophylla.
Comments. The placement of the genus Oecophylla (weaver ants) in the formicine phylogeny remains
uncertain, and we continue to maintain tribal status for this taxonomically distinctive group. A sister group
relationship with Gesomyrmex (Figure 1) is plausible and deserves further investigation.
Tribe Plagiolepidini Forel 1886
= Bregmatomyrminii Wheeler 1929
Genera: Acropyga (tribal transfer), Agraulomyrmex, Anoplolepis (tribal transfer), Aphomomyrmex,
Bregmatomyrma, Lepisiota, Petalomyrmex, Plagiolepis, Tapinolepis.
Comments. A number of genera previously placed in Plagiolepidini, such as Brachymyrmex, Myrmelachista,
Nylanderia, and Prenolepis, are here transferred elsewhere (to either Lasiini or Myrmelachistini), and the tribe is
now comprised exclusively of Old World taxa, except for the cosmopolitan Acropyga. The placement of
Bregmatomyrma has not been evaluated with sequence data, and its retention in Plagiolepidini remains provisional.
A taxonomic revision of the plagiolepidine genera is overdue.
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REVISED CLASSIFICATION OF FORMICINAE
Tribe Santschiellini Forel 1917 stat. rev.
Genus: Santschiella (tribal transfer)
Comments. We resurrect the tribe Santschiellini for this monotypic West African genus due to uncertainty
about its position within the formicine tree (Blaimer et al. 2015). It appears to be the sister group of the Neotropical
genus Gigantiops but there is only moderate support for this hypothesis. See previous discussion under
Gesomyrmecini and Gigantiopini.
Unplaced to tribe
The following fossil genera cannot be placed with confidence in any of the 11 extant tribes and are here considered
incertae sedis in Formicinae: †Camponotites, †Curtipalpulus, †Drymomyrmex, †Eoleptocerites, †Eurytarsites,
†Fushuniformica, †Heeridris, †Huaxiaformica, †Imhoffia, †Kyromyrma, †Leptogasteritus, †Leucotaphus,
†Liaoformica, †Longiformica, †Magnogasterites, †Orbicapitia, †Ovalicapito, †Ovaligastrula, †Protrechina,
†Sinoformica, †Sinotenuicapito, †Wilsonia.
Genus-level changes in the tribe Camponotini
The new phylogeny has important implications for the classification of Camponotini. Camponotus itself is non-
monophyletic, appearing as three separate lineages: Camponotus sensu stricto, Camponotus gigas, and
Camponotus (Colobopsis). C. (Colobopsis) is sister to all other members of the tribe, and is well separated from
true Camponotus. The latter remains technically non-monophyletic, however, because two taxa, Forelophilus and
Phasmomyrmex, are found to be nested within it (Figure 1).
The following changes are made to ensure monophyly of each recognized genus in Camponotini. New and
revived combinations implicitly include the junior synonyms of the species names transferred below. Author and
year of publication for species names can be found in AntCat (http://antcat.org/).
Camponotus Mayr 1861
Type species: Formica ligniperda, designated by Bingham (1903: 347).
For generic synonymy see Bolton (2003) and AntCat (2015).
As a result of new phylogenetic findings we institute the following genus-level changes to Camponotus.
1Forelophilus Kutter (type species Forelophilus overbecki, by monotypy) is demoted to subgenus (stat. n.)
under Camponotus, creating the following new combinations in Camponotus: overbecki, philippinensis,
stefanschoedli. One of these new combinations, C. overbecki (Kutter), becomes a secondary junior homonym,
here replaced with C. javaensis (nom. n.).
2Phasmomyrmex Stitz (type species Phasmomyrmex sericeus (=P. buchneri), by monotypy) is demoted to
subgenus (stat. rev.) under Camponotus, creating the following revived combinations (unless noted as new)
in Camponotus: aberrans, buchneri, buchneri griseus (comb. n.), paradoxus (comb. n.), paradoxus cupreus
(comb. n.), wolfi. One of these new combinations, C. buchneri griseus (Santschi), becomes a secondary junior
homonym, here replaced with C. buchneri camerounensis (nom. n.). The two erstwhile subgenera of
Phasmomyrmex, Myrmorhachis Forel and Myrmacantha Emery, are here treated as junior synonyms (syn. n.)
of Camponotus (Phasmomyrmex).
3 The subgenera Colobopsis Mayr and Dinomyrmex Ashmead are removed from Camponotus, and treated as
separate genera (see below). The Camponotus subgenus Myrmogonia Forel becomes a junior synonym of
Colobopsis.
With these changes Camponotus now comprises 45 valid subgenera (including the nominate subgenus), with
three losses (Colobopsis, Dinomyrmex, Myrmogonia; see below) and two gains (Forelophilus, Phasmomyrmex).
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As others have commented (Bolton 1995), many of the Camponotus subgenera are poorly defined and undoubtedly
represent artificial groupings (Brady et al. 2000; Clouse et al. 2015). Nevertheless, until a global revision of the
genus is carried out, we prefer to retain the subgeneric classification—at least some of the groups are distinctive
and evidently monophyletic, and thus serve to keep clusters of closely related species together.
Colobopsis Mayr 1861 stat. rev.
Type species: Formica truncata, designated by Bingham (1903: 342).
Myrmogonia Forel (as subgenus of Camponotus). Type species Camponotus laminatus, designated by Wheeler (1913: 81).
Syn. n.
Dolophra Wu & Wang. Type species Dolophra politae, by original designation. Junior synonym of Camponotus: Bolton (1995:
27); of Camponotus (Colobopsis): Bolton (2003: 113).
Diagnosis, minor worker. Generally small, HW 0.65–1.10 (exceptions: cylindrica group and the Fijian radiation,
where HW 0.90–1.70), with rounded head and relatively small eyes, REL 0.20–0.32; head width three-quarters of
more of head length (CI 0.75–0.98; except one Fijian species, C. polynesica, where CI ~0.72); antennal
insertions—and hence also the frontal carinae—relatively relatively well separated, ASM/HW 0.36–0.47 (except
cylindrica group, New Caledonia radiation and the Fijian radiation, where ASM/HW 0.31–0.39), ASM/CLW
usually 0.66–0.98 (except some New Caledonian and most Fijian species where ASM/CLW is in the range of 0.60–
0.66); frontal carinae relatively short, usually not strongly sinuate, the antennal insertions occurring at about
midlength of the frontal carinae; clypeus more or less subquadrate, as long as wide or slightly wider than long
(CLW/CLL 0.96–1.32), with sides parallel or diverging moderately towards the anterior margin (clypeus broader in
Fijian species of the bryani and dentata groups where CLW/CLL ~ 1.46, and in the conica and vitrea groups, sensu
Emery (1925), where CLW/CLL 1.40–1.50 and clypeus more trapezoidal in form); anterolateral extremities of
clypeus differentiated from rest of clypeus by a sulcus or impression running from the anterior tentorial pit to the
clypeal margin, the suture between clypeus and malar region of head often weak here, so that the clypeus appears
to lack the anterolateral extensions often conspicuous in Camponotus minors (compare Figures 2–5 with Figure
15).
Diagnosis, major worker. Head generally phragmotic, varying from strongly truncate and marginate (Figure 6)
to weakly truncate (Figure 7), the truncated portion incorporating part of the clypeus, the malar region of the head
capsule and the upper surface of the mandibles. Clypeus elongate-rectangular, the anterolateral extremities
separated from the clypeus by a well-marked sulcus and appearing to form an independent triangular sclerite.
Additional diagnostic features. Dimorphic worker caste, with few or no intermediates between major and
minor workers, except in the cylindrica group (Emery 1925); larva with distinctive ventral trough (praesaepium),
overhung posteriorly by a protruding welt of the second abdominal segment (Wheeler & Wheeler 1953, 1982);
pupa naked (Wheeler 1904). Brendon Boudinot has recently found that male Colobopsis have distinctive genitalia,
with the shape of the digitus distinguishing them from Camponotus males (Boudinot, in prep.).
The elevation of Colobopsis to the rank of genus generates the following new combinations (unless noted as
revived) in Colobopsis: abdita, anderseni, annetteae, aruensis, aurata, aureliana, badia (unresolved junior
primary homonym), badia saginata, brachycephala, †brodiei, bryani, calva, camela, cerberula, ceylonica,
clerodendri, conica (comb. rev.), conithorax, corallina (comb. rev.), cotesii, cristata, culmicola, culmicola
haweisi, custodula, cylindrica (comb. rev.), dentata (comb. rev.), desecta (comb. rev.), elysii, equa, etiolata,
excavata, fijiana, flavolimbata, gasseri (comb. rev.), gundlachi, guppyi, horrens, horripila, hosei, hosei mima,
howensis, hunteri, impressa (comb. rev.), kadi, karawaiewi, laminata, laotsei, lauensis, leonardi, leonardi
gracilenta, leonardi grisea, levuana, loa, loa belli, longi, maafui, macarangae, macrocephala, manni, markli,
mathildeae, mississippiensis, mutilata, mutilata stitzi, newzealandica, nigrifrons (comb. rev.), nipponica, obliqua,
oceanica, papago, perneser, phragmaticola, politae, polynesica, pylartes, pylartes fraxinicola, pylora, quadriceps,
quadriceps convexior, quadriceps curvata, quadriceps nanula, reepeni, riehlii (comb. rev.), rothneyi, rothneyi
krafti, rothneyi makilingi, rothneyi taivanae, rotunda, rufifrons (comb. rev.), rufifrons leucopa, sadina,
sanguinifrons, saundersi, saundersi krama, schmeltzi, schmitzi, severini, shohki, smithiana, solenobia, sommeri
(comb. rev.), stricta (comb. rev.), trajana, tricolor (comb. rev.), triton, truncata (comb. rev.), umbratilis,
vitiensis, vitrea, vitrea angustula, vitrea carinata, vitrea latinota (unresolved junior primary homonym), vitrea
oebalis, vitrea praelutea, vitrea praerufa, vitrea vittatula, and wildae.
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Comments. As now conceived the genus Colobopsis comprises 94 valid species (93 extant, 1 fossil) and 23
subspecies. Based on original descriptions and images, Camponotus bifossus and Camponotus tritschleri,
nominally Colobopsis, are retained in Camponotus, subgenus indeterminate; and Camponotus cordincola is
retained in Camponotus, and transferred to subgenus Pseudocolobopsis. The eight Fijian species in the subgenus
Myrmogonia (Sarnat & Economo 2012) are transferred to Colobopsis, based on both genetic (e.g., Clouse et al.
2015) and morphological evidence. Three Australian species, armstrongi, cameratus, and macareaveyi, previously
assigned to subgenus Myrmogonia, are retained in Camponotus, subgenus indeterminate. McArthur’s (2012)
treatment of Colobopsis, which involved numerous ad hoc and poorly justified transfers of species from other
Camponotus subgenera into Camponotus (Colobopsis), is here ignored.
Biology. Most species of Colobopsis are strictly arboreal, nesting in cavities in dead branches or twigs and
employing phragmotic major workers to block the nest entrance (Forel 1892; Wheeler 1904; Creighton 1967). In
some Fijian species, with reduced phragmosis, nests can also be found in rotten wood and in epiphytic ant-plants
(Sarnat & Economo 2012). Phragmosis is also reduced in some Southeast Asian species nesting in live stems; at
least one species, Colobopsis macarangae, apparently lacks a major worker subcaste (Dumpert 1996). In the field,
collections of Colobopsis can be readily distinguished from those of Camponotus if pupae are available: these are
always naked in Colobopsis (Wheeler 1904; Ward, pers. obs.), while those of Camponotus are enclosed in cocoons.
Distribution. Colobopsis occurs in the New World from southern United States to Costa Rica; across the
southern and central Palearctic from the western Mediterranean to Japan; throughout the Oriental and Australian
biogeographic regions as far south as Tasmania; and into the Pacific as far east as New Caledonia, Vanuatu, and
Fiji. The genus is notably absent from the Afrotropics and most of the Neotropics.
Distinguishing Colobopsis from Camponotus. Despite their phylogenetic distance, morphological
distinctions between Colobopsis and Camponotus have been obscured by extensive evolution within each group,
including convergent evolution of phragmotic heads in the major workers of some Camponotus, and variable
development of phragmosis in Colobopsis (Figures 5–6). Some recent taxonomic treatments have confused the two
lineages. For example, of the 11 species assigned by McArthur & Shattuck (2001) to the “Camponotus
macrocephalus group” eight are Colobopsis (anderseni, annetteae, conithorax, gasseri, howensis, macrocephala,
sanguinifrons, vitrea) and three are Camponotus (janeti, janforrestae, mackayensis). Similarly, of the four newly
described species of phragmotic “Camponotus” from New Guinea in Klimes & McArthur (2014), one is
Colobopsis (rotunda), while the other three are Camponotus.
To reduce future confusion we provide a key for separating minor workers of Colobopsis from those of
Camponotus. It is helpful to segregate the New Caledonian and Fijian species, since Colobopsis tends to “break the
rules” in these island situations (Figures 8–13).
1 Not occurring in Fiji or New Caledonia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- Occurring in Fiji . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- Occurring New Caledonia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2 Generally small species, HW 0.65–1.10 (except cylindrica-group of Southeast Asia with HW 1.20–1.70, and facies as in Fig-
ures 4 and 5); either antennal insertions relatively well separated, such that ASM/HW 0.36–0.47 and ASM/CLW 0.66–0.98,
and/or clypeus relatively narrow, such that CLW/CLL 0.96–1.32; antennal insertions occurring at about midlength of frontal
carinae; anterolateral extremities of clypeus set off from rest of clypeus by a sulcus or impression, so clypeus appears to lack
prominent anterolateral extensions (Figures 2–5) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Colobopsis
- Small to large species, HW 0.70–3.00; antennal insertions less well separated, such that ASM/HW 0.22–0.35 and ASM/CLW
0.35–0.68; clypeus variable in shape but in smaller species with HW 0.70–1.35 (e.g., Camponotus (Myrmamblys), C. (Myr-
mentoma) and C. (Pseudocolobopsis)) clypeus tending to be relatively broad, such that CLW/CLL 1.25–1.62, although excep-
tions occur (e.g., in some C. (Pseudocolobopsis) species) (Figures 14–15); antennal insertions usually occurring in front of
midlength of frontal carinae; clypeus typically with prominent anterolateral extensions (Figure 15) . . . . . . . . . . . Camponotus
3 With conspicuous long setae, gracile legs, and a shield-shaped clypeus with prominent anterolateral extensions (Figure 16). . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Camponotus chloroticus
- Without the combination of conspicuous long setae and gracile legs; clypeus lacking prominent anterolateral extensions (Fig-
ures 8–11) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Colobopsis
4 Small species, HW 0.68–1.04; antennal insertions more widely separated (ASM/HW 0.34–0.39 and ASM/CLW 0.64–0.77)
(Figures 18–19); clypeus tending to be less broad (CLW/CLL 1.15–1.40) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Colobopsis
- Small to medium-sized species, HW 0.75–2.10; antennal insertions less well separated (ASM/HW 0.25–0.29 and ASM/CLW
0.46–0.55); clypeus varying in shape, but if HW < 1.05 (e.g., Camponotus pulchellus complex) (Figure 17) then clypeus tend-
ing to be broader (CLW/CLL 1.25–1.60) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Camponotus
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FIGURES 2–7. Selected species of Colobopsis, full-face view of head of minor worker (Figs. 2–5) and lateral view of soldier
(Figs. 6–7). 2, Colobopsis etiolata, Texas, USA (CASENT0104949); 3, C. truncata, Bulgaria (CASENT0179881); 4. C.
cylindrica group, Brunei (CASENT0280269); 5, C. quadriceps, Papua New Guinea (CASENT0280264); 6, C. truncata, Italy
(CASENT0249998); 7, C. quadriceps, Papua New Guinea (CASENT0280263). Images from AntWeb (www.antweb.org);
photographers Aprile Nobile (2), Erin Prado (3), Estella Ortega (4, 5, 7) and Ryan Perry (6).
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FIGURES 8–13. Species of Colobopsis from Fiji. Full-face view of head of minor worker (Figs. 8–11) and lateral view of
minor worker (Figs. 12–13). 8, Colobopsis vitiensis (CASENT0280250); 9, C. polynesica (CASENT0280252); 10. C. schmelzi
(CASENT0280259); 11, C. bryani (CASENT0280258); 12, C. schmelzi (CASENT0280259); 13, C. bryani
(CASENT0280258). Images from AntWeb (www.antweb.org); photographers Shannon Hartman (8–9) and Estella Ortega (10–
13).
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FIGURES 14–19. Selected species of Camponotus (Figs. 14–17) and New Caledonia Colobopsis (Figs. 18–19), full-face view
of head of minor worker. 14, Camponotus (Pseudocolobopsis) claviscapus, Costa Rica (CASENT0249388); 15, C.
(Myrmentoma) decipiens, Texas, USA (CASENT0249367); 16. C. (Tanaemyrmex) chloroticus, Fiji (CASENT0171139); 17, C.
(Myrmamblys) pulchellus_cf, New Caledonia (CASENT0280237); 18, Colobopsis indet. (CASENT0280248); 19, Colobopsis
camela (CASENT0280242). Images from AntWeb (www.antweb.org); photographers Will Ericson (14–15), Eli Sarnat (16) and
Shannon Hartman (17–19).
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Dinomyrmex Ashmead 1905 stat. rev.
Type species: Formica gigas, by original designation.
Based on its phylogenetic position, as a lineage separate from both Camponotus and Colobopsis, and sister to all
other camponotines except Opisthopsis and Colobopsis, Dinomyrmex is here resurrected as a genus. This generates
the following revived combinations: gigas, gigas borneensis.
Dinomyrmex is a distinctive camponotine, confined to southeast Asia, and recognized by the combination of
very large size (HW 3.25–5.35), elongate antennae and legs, and the presence of a metapleural gland. The species
also has characteristic mandibular dentition, with teeth occurring on both the masticatory and basal margins of the
mandible (Emery 1925).
Concluding remarks
This study continues a series of attempts to revise the higher classification of ants in accordance with new
molecular phylogenetic information, with the goal of promoting a ranked phylogenetic classification (Ward et al.
2010, 2015; Brady et al. 2014; Schmidt & Shattuck 2014), just as systematists are doing for other groups such as
flowering plants (Stephens 2015). Establishing higher taxa that are monophyletic, diagnosable, and—for any given
rank—mutually exclusive can be challenging in the face of variable evolutionary rates and extensive convergence
(Ward 2011), but such classifications are likely to be more stable and informative over the long term than non-
phylogenetic alternatives.
It is important, however, to avoid the trap of essentialist thinking (Hillis 2006). The features that evolved along
a particular branch of the tree of life—the synapomorphies of the clade that succeeds that branch—are not always
easily discovered, nor immutable. Such features may undergo further evolution to the point where they are no
longer recognizable. Thus the higher taxa that we decide to name (as genera, tribes, etc.) will not always be readily
diagnosable by “gold standard” synapomorphies. Morphological circumscription may require conditional
statements and subclauses that account for exceptions. We encountered this situation when attempting to define
Colobopsis on the basis of worker morphology. Diagnostic attributes that hold up under most circumstances for this
genus do not apply to the island radiations that have occurred in New Caledonia and Fiji. Here there are few or no
species of Camponotus (sensu stricto), and Colobopsis appears to have expanded into regions of morphospace not
occupied elsewhere. Any diagnosis of Colobopsis needs to take these exceptions into account.
Our solution was to insert qualifying clauses for the island species, and to treat them separately in the key.
While this might be considered rather arbitrary, it is an effective way to deal with the vagaries of adult worker
morphology. For practical reasons worker morphology is the basis of most ant taxonomy, even though worker
features can be prone to deceptive convergence and divergence. In this instance the larval, pupal, and genetic
(DNA sequence) characteristics of Colobopsis continue to distinguish it globally from Camponotus, so we can be
confident that these island radiations are correctly attributed to Colobopsis.
There are still some issues remaining to be addressed in the higher classification of the Formicinae. The genus
Prolasius may be paraphyletic relative to Ter at om y r m e x , based on cox1 (cytochrome c oxidase I) sequence data
(Alan Andersen, pers. comm.). The new molecular phylogeny of the subfamily reveals three other genera that are
likely non-monophyletic (Nylanderia, Prenolepis and Lepisiota) as well as an apparently undescribed genus related
to Agraulomyrmex (Blaimer et al. 2015). More detailed scrutiny of these taxa, including more comprehensive
taxon sampling, is needed to resolve the taxonomic problems presented. We can therefore anticipate some
additional changes at the genus level, but it is hoped that the basic framework adopted here for the Formicinae
remains robust and useful.
Acknowledgments
For contributing ant specimens for morphological and molecular work we are grateful to Alan Andersen, Rumsaïs
Blatrix, Marek Borowiec, Chris Burwell, Alfred Buschinger, Dinah Davidson, Stéphane DeGreef, Flavia Esteves,
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Linda Farley, Dave General, Peter Hawkes, Ulrich Maschwitz, Dave Olson, Christian Peeters, Mike Sharkey,
Alberto Tinaut, Simon van Noort, and Alex Wild. We thank Barry Bolton, Brendon Boudinot, Jack Longino, and
an anonymous reviewer for comments on the manuscript. This work was supported in part by NSF grants EF-
0431330 (to Seán Brady, Ted Schultz, BLF and PSW), DEB-0842204 (to BLF and PSW), and DEB-1354996 (to
Jack Longino and PSW).
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