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Generic Revision of the Ant Subfamily Dorylinae (Hymenoptera, Formicidae)

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Generic Revision of the Ant Subfamily Dorylinae (Hymenoptera, Formicidae) A peer-reviewed open-access journal ZooKeys 608: 1–280Generic (2016) revision of the ant subfamily Dorylinae (Hymenoptera, Formicidae)1 doi: 10.3897/zookeys.608.9427 MONOGRAPH http://zookeys.pensoft.net Launched to accelerate biodiversity research Generic revision of the ant subfamily Dorylinae (Hymenoptera, Formicidae) Marek L. Borowiec1 1 Department of Entomology and Nematology, One Shields Avenue, University of California at Davis, Davis, California, 95616, USA Corresponding author: Marek L. Borowiec ([email protected]) Academic editor: B. Fisher | Received 3 June 2016 | Accepted 17 July 2016 | Published 4 August 2016 http://zoobank.org/F865473C-0337-4FD2-915A-0E3DD2299E66 Citation: Borowiec ML (2016) Generic revision of the ant subfamily Dorylinae (Hymenoptera, Formicidae). ZooKeys 608: 1–280. doi: 10.3897/zookeys.608.9427 Abstract The generic classification of the ant subfamily Dorylinae is revised, with the aim of facilitating identi- fication of easily-diagnosable monophyletic genera. The new classification is based on recent molecular phylogenetic evidence and a critical reappraisal of doryline morphology. New keys and diagnoses based on workers and males are provided, along with reviews of natural history and phylogenetic relation- ships, distribution maps, and a list of valid species for each lineage. Twenty-eight genera (27 extant and 1 extinct) are recognized within the subfamily, an increase from 20 in the previous classification scheme. Species classified in the polyphyletic Cerapachys and Sphinctomyrmex prior to this publication are here distributed among 9 and 3 different genera, respectively. Amyrmex and Asphinctanilloides are synonymized under Leptanilloides and the currently recognized subgenera are synonymized for Dorylus. No tribal clas- sification is proposed for the subfamily, but several apparently monophyletic genus-groups are discussed. Valid generic names recognized here include: Acanthostichus (= Ctenopyga), Aenictogiton, Aenictus (= Par- aenictus, Typhlatta), Cerapachys (= Ceratopachys), Cheliomyrmex, Chrysapace gen. rev., Cylindromyrmex (= Holcoponera, Hypocylindromyrmex, Metacylindromyrmex), Dorylus (= Alaopone syn. n., Anomma syn. n., Cosmaecetes, Dichthadia syn. n., Rhogmus syn. n., Shuckardia, Sphecomyrmex, Sphegomyrmex, Typhlopone syn. n.), Eburopone gen. n., Eciton (= Camptognatha, Holopone, Mayromyrmex), Eusphinctus gen. rev., Labidus (= Nycteresia, Pseudodichthadia), Leptanilloides (= Amyrmex syn. n., Asphinctanilloides syn. n.), Lioponera gen. rev. (= Neophyracaces syn. n., Phyracaces syn. n.), Lividopone, Neivamyrmex (= Acama- tus, Woitkowskia), Neocerapachys gen. n., Nomamyrmex, Ooceraea gen. rev. (= Cysias syn. n.), Parasyscia gen. rev., †Procerapachys, Simopone, Sphinctomyrmex, Syscia gen. rev., Tanipone, Vicinopone, Yunodorylus gen. rev., Zasphinctus gen. rev. (= Aethiopopone syn. n., Nothosphinctus syn. n.). Keywords Taxonomy, systematics, morphology, dorylomorphs, doryline section, army ants Copyright Marek L. Borowiec. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 2 Marek L. Borowiec / ZooKeys 608: 1–280 (2016) Table of contents Preface ...................................................................................................................... 3 Material and methods............................................................................................... 3 Brief introduction to doryline ants ..................................................................... 5 History of taxonomic and phylogenetic research ................................................ 6 Current views on doryline classification and evolution ..................................... 11 Classification of the Dorylinae proposed here ...................................................14 Morphology ..................................................................................................... 16 Diagnostic characters of the worker ..................................................................18 Diagnostic characters of the male ..................................................................... 25 Diagnostic characters of the gyne ..................................................................... 26 Characters used to describe worker morphology ...............................................26 Characters used to describe male morphology .................................................. 33 Characters used to describe gyne morphology .................................................. 36 Key to the genera of doryline ants based on workers .........................................37 Provisional key to the genera of doryline ants based on males ...........................52 Taxonomic treatment of the genera of Dorylinae ....................................................66 Acanthostichus Mayr, 1887 ................................................................................66 Aenictogiton Emery, 1901b ............................................................................... 73 Aenictus Shuckard, 1840b................................................................................. 78 Cerapachys Smith, F., 1857 ............................................................................... 90 Cheliomyrmex Mayr, 1870 ................................................................................ 95 Chrysapace Crawley, 1924a, gen. rev. .............................................................. 101 Cylindromyrmex Mayr, 1870 ...........................................................................106 Dorylus Fabricius, 1793 .................................................................................. 112 Eburopone gen. n. ........................................................................................... 124 Eciton Latreille, 1804 ..................................................................................... 129 Eusphinctus Emery, 1893a, gen. rev. ................................................................138 Labidus Jurine, 1807 ...................................................................................... 144 Leptanilloides Mann, 1923.............................................................................. 150 Lioponera Mayr, 1879, gen. rev. ......................................................................156 Lividopone Fisher & Bolton, 2016 ..................................................................165 Neivamyrmex Borgmeier, 1940 ....................................................................... 170 Neocerapachys gen. n. ......................................................................................180 Nomamyrmex Borgmeier, 1936....................................................................... 185 Ooceraea Roger, 1862, gen. rev. ...................................................................... 191 Parasyscia Emery, 1882, gen. rev. .................................................................... 198 Procerapachys Wheeler, W. M. 1915b.............................................................. 205 Simopone Forel, 1891 ..................................................................................... 208 Sphinctomyrmex Mayr, 1866b .........................................................................214 Syscia Roger, 1861, gen. rev. ........................................................................... 219 Generic revision of the ant subfamily Dorylinae (Hymenoptera, Formicidae)3 Tanipone Bolton & Fisher, 2012 .....................................................................225 Vicinopone Bolton & Fisher, 2012 .................................................................. 230 Yunodorylus Xu, 2000, gen. rev. ...................................................................... 232 Zasphinctus Wheeler, W. M., 1918, gen. rev. .................................................. 237 Acknowledgments ................................................................................................ 243 References ............................................................................................................ 244 Appendix .............................................................................................................. 275 Preface The ant subfamily Dorylinae is a monophyletic group of predatory ants, occurring throughout most of the tropical and subtropical regions of the world, with an appreci- able number of species in warm temperate environments. The relatively few dorylines for which foraging biology is known usually prey on other ants or social insects, al- though notable exceptions occur and several of the charismatic ‘army ants’ evolved more generalized predatory habits. There are about 680 described species with an es- timate of the total diversity being at least 1,000. The diversity of both habits and morphology within the subfamily is high and nesting can be subterranean or arboreal, with colony sizes ranging from a few dozen to millions of workers. These workers vary from having well-developed compound eyes to being entirely blind, having very short to very long slender appendages, and with the cuticle varying from coarsely sculptured to polished and shiny, with dull or conspicuous coloration. Although numerous studies focusing on the biology of the few conspicuous species have been published, our overall knowledge of this clade is poor. A likely contributing
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