http://dx.doi.org/10.5852/eit.2015.12Q

www. europeani ournaloftaxonomv. eu

2015 • Boudinot B.E.

This work is licensed under a Creative Commons Attribution 3.0 License.

Research monograph

urn:lsid:zoobank.org:pub:54714320-5726-44CB-8FF5-60E0B984873D

Contributions to the knowledge of Formicidae (Hymenoptera,

Aculeata): a new diagnosis of the family, the first global male-based

key to subfamilies, and a treatment of early branching lineages

Brendon E. BOUDINOT

Department of Entomology/Nematology, Briggs Hall, Rm. 381

University of California, Davis, U.S.A.

[email protected]

urn:lsid:zoobank.org:author:919F03B0-60BA-4379-964D-A56EB582E16D

Table of contents

Abstract.2

Introduction .2

Material and Methods .3

Terminology.5

Figure abbreviations . 6

Measurements and indices.10

Repositories. 11

Results.12

Family Formicidae Latreille, 1809 . 12

Key to global subfamilies, based on extant males. 14

Treatments of focal taxa.17

Subfamily Amblyoponinae F orel, 1893. 17

Genus Apomyrma Brown, Gotwald Jr. & Fevieux, 1971.22

Apomyrma stygia Brown, Gotwald Jr. & Fevieux, 1971.24

Apomyrma CD01 . 24

Subfamily Feptanillinae Emery, 1910.29

Subfamily Martialinae Rabeling & Verhaagh, 2008 . 33

Genus Martialis Rabeling & Verhaagh, 2008.37

Martialis henreka Rabeling & Verhaagh, 2008 . 37

Shared apomorphies of the basal ants . 43

Brief global diagnoses of subfamilies, based on males.46

Subfamily Agroecornyrmecinae Carpenter, 1930 . 46

Subfamily Paraponerinae Emery, 1901.47

Subfamily Ponerinae Fepeletier de Saint-Fargeau, 1835 . 47

Subfamily Proceratiinae Emery, 1895 . 48

Subfamily Dorylinae Feach, 1815 . 48

Subfamily Myrmeciinae Emery, 1877 . 49

Subfamily Pseudomyrmecinae M.R. Smith, 1952 . 49

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European Journal of Taxonomy 120: 1-62 (2015)

Subfamily Aneuretinae Emery, 1913 . 49

Subfamily Dolichoderinae Forel, 1878 . 50

Subfamily Formicinae Latreille, 1809 . 51

Ectaheteromorph clade. 51

Subfamily Myrmicinae Fepeletier de Saint-Fargeau, 1835.56

Acknowledgments . 56

Fiterature.57

Abstract. The diagnosis of the Formicidae is revised, including five new, unreversed apomorphies, of

which one is a unique synapomorphy. The first global male-based key to all subfamilies is provided

and illustrated, and all ant subfamilies are diagnosed for males on a global scale for the first time.

Three lineages of “basal ants” are assessed in detail: the Amblyoponinae, Feptanillinae, and Martialinae.

The males of Martialis heureka (Martialinae) and Apomyrma (Amblyoponinae) are described. The

Martialinae and Feptanillinae are diagnosed based on males, and additional diagnostic traits for the male

of Amblyoponinae and worker of Martialis are provided. The placement of Scyphodon and Noonilla in

the Formicidae and Feptanillinae is confirmed. Morphological characters of the Amblyoponinae, the

Feptanillinae, and the Martialinae are contrasted, and potentially homologous apomorphies are signaled.

Keywords. Formicoidea, taxonomy, apomorphies, mesosoma morphology, coxal morphology

Boudinot B.E. 2015. Contributions to the knowledge of Fonnicidae (Hymenoptera, Aculeata): a new diagnosis of

the family, the first global male-based key to subfamilies, and a treatment of early branching lineages. European

Journal of Taxonomy 120: 1-62. http://dx.doi.org/10.5852/ejt.2015.120

Introduction

Ants are a globally diverse and dominant lineage of eusocial aculeates. As posited by Bolton (1994), “it

is a truism that [ants] occupy a position among the terrestrial invertebrates equivalent to that occupied

by our species among the vertebrates”. The higher taxonomy and internal phylogeny of the Formicidae

has stabilized significantly in the past few decades due to the active study of ant systematics at the macro

scale (Baroni Urbani et al. 1992; Bolton 1994, 2003; Moreau et al. 2006; Brady et al. 2006; Ward et

al. 2010; Schmidt 2013; Schmidt & Shattuck 2014; Brady et al. 2014; Ward et al. 2015), spurred by

the pioneering morphological studies of the family by Bolton (1990a, 1990b, 1990c, 1994, 2003). The

relationship of the Formicidae with other aculeate families is still rather liquid (see references in Ward

2014), but a recent phylogenomic study supports a sister-group relationship with the Apoidea (Johnson

et al. 2013). Bolton (1994, 2003) authoritatively diagnosed the family and its subfamilies and provided

worker-based keys to the subfamilies and genera (Bolton 1994).

Here, the diagnosis of the Formicidae is updated based on study of all adult castes. Five new unreversed

apomorphies are presented, one of which is a synapomorphy unique among the Hymenoptera. The

remainder of the paper focuses primarily on male ants. To date the vast majority of myrmecological

research has been based on workers due to their abundance and conspicuousness. The sexual dimorphism

of ants (and Hymenoptera in general) renders male-worker associations very challenging to make, and

has led to major taxonomic bias. Of the -12,800 valid extant ant species, males are described for only

-3,450, or 27% of the total, and males are unknown for almost a quarter of the genera. Very little

synthetic work has been done on males; indeed, bioregional male-based keys to subfamily are available

only for North America (Smith 1943), the Palearctic (Western Europe, Bernard 1967; Armenia,

Arakelian 1994; European Russia, ArnoFdi & Dlusslcy 1978; southern Siberia, Radchenko 1994; North

Korea, Radchenko 2005; there are several European country-specific treatments), Japan (Yoshimura &

Onoyama 2002), New Zealand (Brown 1958), and Madagascar (Yoshimura & Fisher 2007). To stimulate

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BOUDINOT B.E., Contributions to the Formicidae

research on and collection of male ants, and to ease the task of associating males and females, the first

global male-based key to subfamilies is presented.

All subfamilies keyed herein are provided with brief global male-based diagnoses. Three of these

subfamilies, Amblyoponinae, Leptanillinae, and Martialinae, are treated in more detail due to their

important phylogenetic position, as lineages near the base of the tree. The male of the “Martian

ant”, Martialis heureka —the sole representative of the Martialinae—is described and diagnosed. As

in the worker, the male of Martialis displays a unique mixture of pleisiomorphic and derived traits.

The discovery of the male of Martialis highlights the value of alternative myrmecological sampling

techniques, as only one undamaged worker of Martialis is known, while 25 males were found from one

jar of unsorted Amazonian Malaise trap residues. Another male is described herein, that of an Apomyrma

(Amblyoponinae) morphospecies. The male of Apomyrma was previously incompletely described from

pupae (Brown et al. 1971). The Teptanillinae are re-diagnosed, clarifying the boundaries of this clade,

and the “mysterious” male taxa Noonilla and Scyphodon are confirmed as leptanillines.

Material and Methods

Specimens were examined with several microscopes, but primarily with a Wild M5 stereo microscope

with 50 x maximum magnification. Stacked photomicrographs were captured and montaged via Auto-

Montage Pro (Synoptics Ttd., Cambridge, England) with a JVC KY-F57U camera mounted on a Leica

MZ 16A stereo microscope. All images were edited in Adobe Photoshop CS5, including those used from

Ant Web (2014) with permission, and all figures were compiled using Adobe Illustrator CS6 (Adobe

Systems Inc., California, U.S.A.). Dissections were carried out in watchglasses (Syracuse staining

dishes) filled with 95% ethanol and under either the Leica or the Wild microscopes mentioned above.

Genitalia were removed from the metasoma using size 1 entomology pins and size 3 forceps and were

imm obilized for imaging and examination with ethanol- imm ersed Blu-Taclc (Bostik, Indianapolis,

U.S.A.). Pleisiomorphic conditions were inferred from a synoptic examination of extant and extinct

Formicidae, and/or were understood from the work of Bolton (1990a, 1990b, 1990c, 2003). Taxonomic

catalog resources were from AntCat by Bolton (2014). Some specimens examined have alphanumeric

codes associated with them (i.e., CASENT#, INB#, INBIOCRI#, and UCRENT#) which uniquely

identify the specimens for databasing purposes.

Taxa examined

To ascertain specificity of characters provided in the diagnosis of the Formicidae (below), representatives

of all major aculeate lineages, and some “Parasitica” and “Symphyta”, were examined. The higher

classification of the Aculeata follows Pilgrim et al. (2008). While not all are listed, particular taxa

examined include:

“Symphyta”

Cephoidea: Cephidae.

Megalodontoidea: Pamphilidae (. Pamphiliuspacificus ).

Orussoidea: Orussidae ( Orussus sp.).

Tenthredinoidea: Argidae; Tenthredinidae.

“Parasitica”

Ceraphronoidea: Megaspilidae.

Chalcidoidea: Chalcididae; Encyrtidae; Eucharitidae; Eurytomidae; Perilampidae; Pteromalidae.

Cynipoidea: Cynipidae; Eucoliidae; Figitidae.

Evanioidea: Aulacidae; Gasteruptiidae.

Ichneumonoidea: Braconidae; Ichneumonidae.

Platygastroidea: Platygastridae; Scelionidae.

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European Journal of Taxonomy 120: 1-62 (2015)

Proctotrupoidea: Diapriidae; Proctotrupidae.

Trigonalyoidea: Trigonalyidae (Trigonalys melanoleuca).

Aculeata

Apoidea: Ampulicidae (Ampulex sp.); Crabronidae; Nyssonidae (Bembicinae, Bemb ix); Pamphredonidae;

Philanthidae (Philanthinae); Sphecidae (Ammophilinae, Sphecinae). Anthophila: Andrenidae; Apidae;

Megachilidae; Halictidae.

Chrysidoidea: Bethylidae; Chrysididae (Chrysidinae, Cleptinae); Dryinidae; Embolemidae (Embolemus

nearcticus); Plumariidae.

Formicoidea: Formicidae (see below).

Pompiloidea: Mutillidae (Sphaeropthalminae); Myrmosidae (Myrmosinae); Pompilidae (Pepsinae);

Sapygidae (Sapyginae).

Scolioidea: Bradynobaenidae (Bradynobaenus gayi ); Scoliidae ( Campsomerispilipes).

Tiphioidea: Sierolomorphidae (Sierelomorpha similis).

Thynnoidea: Chyphotidae (Chyphotes spp.); Thynnidae (Anthoboscinae, Anthobosca insularis).

Vespoidea: Rhopalosomatidae (Rhopalosoma nearcticum ); Vespidae (Eumeninae, Masarinae, Vespinae).

Ant taxa were examined and evaluated either from physical specimens or high quality images available

from Ant Web (2014). Taxa examined during this study cover slightly over 75% of the extant generic

diversity of the family, with males examined for 70% of the total. Parentheses indicate the number of

examined valid extant genera out of the total number of valid extant genera for each subfamily followed

by the total number of genera for which males were examined (x/y; z, where x = total examined, y =

total valid, z = male examined total). For each genus caste is indicated in brackets if only male (c?) or

female castes ($) were examined. Workers and gynes are treated together for the purposes of this work

as the female castes are overall more similar to each other than to males.

Formicidae (245/320; 225):

Feptanillinae (7/8; 6): Anomalomyrma [§], Leptanilla , Noonilla [c?], Phaulomyrma [f], Protanilla ,

Scyphodon [f], Yavnella [f].

Martialinae (1/1): Martialis.

Agroecomyrmecinae (2/2; 1): Ankylomyrma [§], Tatuidris.

Amblyoponinae (13/13; 10): Adetomyrma, Amblyopone , Apomyrma , Bannapone [§], Concoctio [$],

Myopopone , Mystrium , Onychomyrmex, Opamyrma [$], Paraprionopelta, Prionopelta, Stigmatomma ,

Xymmer.

Paraponerinae (1/1): Paraponera.

Ponerinae (32/47): Anochetus , Belonopelta , Brachyponera , Centromyrmex , Cryptopone, Diacamma,

Dinoponera , Dolioponera , Ectomomyrmex, Emeryopone, Harpegnathos, Hypoponera , Leptogenys,

Mayaponera , Megaponera , Mesoponera, Myopias , Neoponera, Odontomachus, Odontoponera ,

Ophthalmopone , Pachycondyla, Paltothyreus, Phrynoponera , Platythyrea , Plectroctena, P oner a,

Psalidomyrmex , Pseudoponera, Rasopone , Simopelta, Thaumatomyrmex.

Proceratiinae (3/3): Discothyrea, Probolomyrmex, Proceratium.

Dorylinae (18/18; 17): Acanthostichus , Aenictogiton , Aenictus, Amyrmex, most “ Cerapachys ” clades

[5 or cJ], Cheliomyrmex, Cylindromyrmex, Dorylus , Eciton , Labidus , Leptanilloides , Neivamyrmex,

Nomamyrmex, Simopone , Sphinctomyrmex sensu lato and .s’ms'z/ stricto , Tanipone , Vicinopone [y],

Myrmeciinae (2/2): Myrmecia , Nothomyrmecia.

Pseudomyrmicinae (3/3): Myrcidris, Pseudomyrmex, Tetraponera.

Aneuretinae (1/1): Aneuretus.

Dolichoderinae (20/28; 19): Anillidris [<$], Anonychomyrma [c?], Aptinoma [f], Arnoldius [$], Azteca,

Bothriomyrmex, Dolichoderus , Dorymyrmex, For elms, Iridomyrmex , Leptomyrmex , Linepithema,

Liometopum , Ochetellus, Papyrius, Philidris , Ravavy [f], Tapinoma , Technomyrmex , Turneria.

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BOUDINOT B.E., Contributions to the Formicidae

Formicinae (49/51; 41): Acropyga, Agraulomyrmex, Anoplolepis, Aphomomyrmex, Bajcaridris,

Brachymyrmex, Bregmatomyrma [$], Calomyrmex, Camponotus, Cataglyphis, Cladomyrma,

Echinopla, Euprenolepis [§], Forelophilus, Formica, Gesomyrmex [$], Gigantiops, Iberoformica

[5], Lasiophanes, Lasius, Lepisiota, Melophorus, Myrmecocystus, Myrmecorhynchus, Myrmelachista,

Myrmoteras, Notorious [fy], No to stigma, Nylanderia, Oecophylla, Overbeckia, Paraparatrechina,

Paratrechina, Petalomyrmex, Phasmomyrmex, Plagiolepis, Polyergus, Polyrhachis, Prenolepis,

Proformica, Prolasius, Pseudolasius, Pseudonotoncus [$], Rossomyrmex [$], Santschiella [$],

Stigmacros, Tapinolepis, Teratomyrmex [$], Zatania.

Ectaheteromorph clade (7/7; 6): Acanthoponera, Aulacopone [§], Ectatomma, Gnamptogenys,

Heteroponera, Rhytidoponera, Typhlomyrmex.

Myrmicinae (86/140; 83):

Myrmicini (2/2): Manica, Myrmica.

Pogonomyrmecini (2/2): Hylomyrma, Pogonomyrmex sensu stricto and angustus clade.

Stenammini (7/7): Aphaenogaster sensu stricto and phalangium clade, Goniomma, Messor sensu stricto,

Novomessor, Oxyopomyrmex, Stenamma, Veromessor.

Solenopsidini (13/20): Adelomyrmex, Bariamyrma, Cryptomyrmex, Dolopomyrmex, Kempfidris,

Megalomyrmex, Monomorium sensu stricto and groups of antarcticum, denticulatum, and latastei,

Myrmicaria, Oxyepoecus, Rogeria, Solenopsis, Stegomyrmex, Tropidomyrmex,

Attini sensu lato (38/45): Acanthognathus, Acromyrmex, Allomerus, Apterostigma, Atta, Basiceros,

Blepharidatta, Cephalotes, Cyatta, Cyphomyrmex sensu stricto and strigatus and wheeleri clades,

Daceton, Diaphoromyrmex, Eurhopalothrix, Kalathomyrmex, Lachnomyrmex, Lenomyrmex,

Microdaceton, Mycetagroicus, Mycetarotes, Mycetophylax, Mycetosoritis, Mycocepurus,

Myrmicocrypta, Ochetomyrmex, Octostruma, Paramycetophylax, Phalacromyrmex, Pheidole,

Procryptocerus, Protalaridris, Pseudoatta, Rhopalothrix, Sericomyrmex, Strumigenys, Talaridris,

Trachymyrmex, Tranopelta, Wasmannia.

Crematogastrini (28/64; 25): Acanthomyrmex, Atopomyrmex, Calyptomyrmex, Cardiocondyla,

Care bar a, Cataulacus, Crematogaster, Dacetinops, Formicoxenus, Harpagoxenus, Indomyrma,

Leptothorax, Melissotarsus, Meranoplus, Myrmecina, Nesomyrmex, Perissomyrmex [$], Podomyrma,

Pristomyrmex, Proatta [$], Recurvidris [§], Rhopalomastix, Temnothorax, Terataner, Tetramorium,

Trichomyrmex, Vollenhovia, Xenomyrmex.

Terminology

Terminology follows Harris (1979) for sculpture; Wilson (1955) for setational stature; Boudinot

(2013) for genitalia; Brown & Nutting’s abscissa-oriented nomenclature (1949) for wing venation;

Yoshimura & Fisher (2011) for cellular terminology with the modifications proposed in Boudinot et

al. (2013); Keller (2011) and Boudinot et al. (2013) for the head capsule; and Keller (2011) for sundry

morphological concepts, including helcial axiality (e.g., axial, supraaxial, and infraaxial). The wing

venation typification system presented by Ogata (1991) is used as valuable short-hand for venational

patterns.

Mesosomal terms are described here, as several of those used herein are not in standard use in the

majority of myrmecological works (Figs 1-2). These terms are preferred, as they refer to previously

unappreciated structures, and clarify interfamilial homologies. Terms explicitly relating to the alinota

(meso- and metanota) specifically apply to males and gynes, while terms of the meso- and metapecta

apply for all adult castes with two exceptions noted below. The mesonotum is comprised of the anterior

mesoscutum and posterior mesoscutellum which are divided by the transverse transscutal line. The

anterior mesoscutal margin often bears notauli, or anterior lateromedian sulci, which extend posteriorly

toward the transscutal line. Two fine posterior lateromedian sulci are present, termed the parapsidal

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European Journal of Taxonomy 120: 1-62 (2015)

lines , which extend anteriorly from the transscutal line. Bordering the posterolateral mesoscutal margins

are parascutal carinae, which separate the dorsal mesoscutal disc from the lateral preaxilla. Posterad

the transscutal line is the mesoscutellum, which is traversed by the scutoscutellar suture. This suture

divides the lateral mesoscutellar portions into the anterior axilla and posterior axillula , and anteriorly

delimits the mesoscutellar disc. Often the scutoscutellar suture is impressed, forming the scutoscutellar

sulcus. Bounding the axillula ventrolaterally is the mesoscutellar arm. In Hymenoptera, the meso-

and metapleurae are completely fused with their respective sterna; thus, the terms mesopectus and

metapectus are appropriate to use, although the margin between the metapleuron and metasternum

may be arbitrarily determined by presence of the coxal foramina. An oblique mesopleural furrow or

sulcus (Yoshimura & Fisher 2007) divides the mesopleural area into the lower katepisternum and upper

anepisternum. The term “oblique mesopleural sulcus” is preferred over “anapleural sulcus”, as the sulcus

has evolved several times independently in the Hymenoptera and is probably not homologous with the

sulcus corresponding to the anapleural suture joining the expanded katapleural and anapleural arches of

other pterygota. Above the anepisternum may occur a broad sulcus corresponding to the subalar area.

Posterodorsad this area is a ridge which is homologous with the mesepimeron ; the mesepimeron does

not extend anterad beyond the mesopleural wing process. Worker mesepimera are not differentiated and

should be considered lost due to fusion of the mesopleuron and mesonotum; similarly, the subalar area

is not developed in the worker caste. The metapleural spiracle may be covered by the supramesopleural

sclerite (Vilhelmsen et al. 2010), which for ease is referred to as the spiracular sclerite here. This

structure has previously been termed the “basalar lobe” (Deyrup & Cover 2004; MacGown et al. 2014)

and the “epimeral lobe” (Yoshimura & Fisher 2007). Finally, the metapleural area is divided into the

lower and upper metapleuron by a sulcus.

Terminology of the coxae is as follows (Fig. 3): Each coxa is divided into a basicoxa and disticoxa by a

basicoxal suture , which occurs near the coxal articulation with the mesosoma. Although not emphasized

here, each basicoxite bears lateral and medial coxal-pectal fossae , which articulate with the lateral

condyle of the pleuron and the medial condyle of the sternum , respectively. The disticoxite bears the

anterior and posterior trochanteral-coxal fossae with which the anterior and posterior trochanteral

condyles articulate. Two foramina occur on each coxa, a basicoxal and a disticoxal foramen ; both bear

intersegmental membranes. The disticoxal foramen is enclosed by the flexor (ancestrally lateral) and

extensor (ancestrally medial) margins.

Figure abbreviations

To maximize size of the figures and to reduce redundancy, abbreviations for figures 1, 2, 3, 9, and 12

are listed below.

Fig. 1: Anep = anepisternum. Ax = axilla, Axu = axillula, Ktep = katepisternum, Lmpl = lower

metapleuron, Mepm = mesepimeron, Msmtps = mesometapleural suture, Msnt = mesonotum, Mspp

mesopleural pit, Mssctla = mesoscutellar arm, Mssctld = mesoscutellar disc, Mssctm = mesoscutum,

Mtsctt = metascutellar trouch, Mtpds = metapleuropropodeal suture. Not = notaulus, Oms = oblique

mesopleural sulcus, PI = parapsidal line, Ppd = propodeum, Ppdl = propodeal lobe, Ppdsp = propodeal

spiracle, Prax = preaxilla, Prnt = pronotum, Prntl = pronotal lobe, Psc = parascutellar carina, Saa =

subalar area, Scscs = scutoscutellar suture, Spsc = spiracular sclerite, Sss = scutoscutellar sulcus, Tg =

tegulum, Tscl = transscutal line, Umpl = upper metapleuron.

Fig. 2: AsII= abdominal sternum II, Lcpmsp = lateral coxal articular process of the mesopleuron, Lcpmtp

= lateral coxal articular process of the metapleuron, Mcpmsp = medial coxal articular process of the

mesopectus, Mcpmtp = medial coxal articular process of the metapectus, Msd = mesodiscrimen, Msp =

mesopectus, Mscf= mesocoxal foramen, Mspfp = mesoprefurcal pit, Mtcf= metacoxal foramen, Mtp

metapectus. Mtpfp = metaprefurcal pit, Prn = pronotum, Vlmspl = ventrolateral mesopectal line.

6

BOUDINOT B E., Contributions to the Formicidae

Fig. 1. Mesosoma of Paraponera clavata male (Paraponerinae, Formicidae). A. Dorsal view. B. Dorsal

posterolateral oblique view. C. Lateral view. Scale bar = 1.0 mm. Abbreviations: see Material and

Methods.

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European Journal of Taxonomy 120: 1-62 (2015)

Fig. 2. Pterothoracic venter morphology of representative hymenopterans. A. Tenthredinidae, female.

B. Polistes (Vespidae), worker. C. Paraponera clavata gyne (Paraponerinae, Formicidae). Scale bars =

1.0 mm. Abbreviations: see Material and Methods.

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BOUDINOT B.E., Contributions to the Formicidae

A 1

Amdcm

Fig. 3. Coxal morphology of representative hymenopterans: right procoxae, left column; right mesocoxae,

middle column; right metacoxae, right column; top procoxa lateral view, bottom procoxa medial view;

top meso- and metacoxae anterior view, bottom meso- and metacoxae posterior view. A. Tenthredinidae.

B. Polistes sp., Vespidae. C. Paraponera clavata gyne (Paraponerinae, Formicidae). Scale bars =1.0 mm.

Abbreviations: see Material and Methods.

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European Journal of Taxonomy 120: 1-62 (2015)

Fig. 3: Aldcm = apicolateral disticoxal margin; Amdcm = Apicomedial disticoxal margin; Be = basicoxa;

Bcf= basicoxal foramen; Bcs = basicoxal suture; Dc = disticoxa; Dcf= disticoxal foramen; Dcs =

disticoxal suture; Tcfa = trochanteral-coxa fossa, anterior; Tcfp = trochanteral-coxa fossa, posterior; Tch

= trochanter. Green dot indicates concealed articulation.

Figs 9, 12: Wing abbreviations'. C = Costal vein. Sc = Subcostal vein, R = Radial vein, Rs = Radial

sector, M = Medial vein, Cu = Cubital vein, 1A = first Anal vein, CC = costal cell, BC = basal cell,

SMC = submarginal cell, MC1/MC = marginal cell 1, SBC = subbasal cell, SDC1/SDC = subdiscal cell

1. Genital abbreviations'. Bm = basimere; Bv = basivolsella; Cu = Cupula in Fig. 9D, 9F, and cuspis

in Fig. 9E, 9G (cuspis absent in Martialis)', Di = digitus; Fg = foramen genitale; Pm = paramere; Pv =

penisvalva; Sp = spiculum; Vc = valviceps; Vu = valvura.

Measurements and indices

Male morphometries practiced herein are improved by carefully attempting to render maximum

replicability. Specific improvements include measurement of antenna characters in medial view, and

determination that the anteriormost point of the axillae are superior landmarks for measuring mesoscutal

and mesoscutellar length, as the transscutal line may be difficult to ascertain in dorsal view. Abdominal

segment III and more-posterior segments were not measured due to the weak sclerotization of Martialis

males, which caused crumpling during the drying process. The wings are not measured, as accurate

metrics would require slide mounting. Numerous indices are supplied, as these calculations provide

proportionality information.

Male measurements

HL

HW1

HW2

MAL

MDL

SL

PDL

A3L

AAL

EL

EW

OOD

LOD

MOD

ML

Head Length, maximum length of head in full-face view from anterior clypeal margin to

posterior head margin between lateral ocelli, ignoring distance which ocelli project and

regardless of whether occipital carina or vertex is posteriormost

Head Width 1, maximum width of head excluding eyes in full-face view

Head Width 2, maximum width of head including eyes in full-face view

Malar Area Length, minimum distance between compound eye and lateral point of

mandibular insertion

Mandible Length, chord length of mandible from medial point of insertion to apex

Scape Length, maximum length of scape in medial view, excluding condylar neck

Pedicel Length, maximum length of pedicel in medial view

Ante nn omere 3 Length, maximum length of antennomere 3 in medial view

Apical Ante nn omere Length, maximum length of apicalmost ante nn omere in medial view

Eye Length, maximum diameter of eye with head positioned in profile view such that

anterior and posterior eye margins are in same plane of focus

Eye Width, maximum width of eye at an axis orthogonal to Eye Length with head

oriented as above

Ocular-Ocellus distance, minimum distance between lateral ocellus and compound eye

Lateral Ocellus Length, maximum diameter of lateral ocellus with head oriented such

that anterior and posterior lateral ocellus margins are in same plane of focus

Median Ocellus Length, maximum diameter of median ocellus in full-face view

Mesosoma Length, maximum diagonal length of mesosoma in profile view from

inflection point of anterior pronotal declivity (between pronotal neck and anteromedian

face) to propodeal lobe, or if propodeal lobe absent then to juncture of lateral and dorsal

margins of petiolar foramen

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BOUDINOT B.E., Contributions to the Formicidae

MLL

MLW

MTL

MTW

PFL

MFF

PTH

PTF

Mesoscutellum Fength, maximum length of mesoscutellum in dorsal view with anterior

mesoscutal and posterior mesoscutellar margins in same plane of focus, measured from

anteriormost points of axillae to posterior mesoscutellar margin

Mesoscutellum Width, maximum width of mesoscutellum with specimen oriented as for

MFF

Mesoscutum Fength, length of mesoscutum in dorsal view with specimen oriented as in

MFF, measured from anterior mesoscutal margin to anteriormost points of axillae

Mesoscutum Width, maximum width of mesoscutum measured with specimen oriented

as in MTF

Pro femur Fength, maximum length of pro femur in posterior view

Metafemur Fength. Maximum length of metafemur in anterior view

Petiole Height, dorsoventral height of petiole in profile view, from node dorsum to

ventral-most point orthogonal to Petiole Fength measurement

Petiole Fength, length of petiole in profile view along anteroposterior axis from

inflection point of petiolar presclerites (the articulatory surfaces) to posteriormost point

of posterior margin

Indices

Cl = Cephalic Index HW1/HF x 100

CS = Cephalic Size (HWl+HF)/2

SEI = Scape-Eye Index EF/SFx 100

SI = Scape Index SF/HWlx 100

El = Eye Index 1 EW/EFx 100

EYE = Eye Index 2 (EF+EW) /CSx 100

MDI = Mandible Index MDF/HFx 100

OBI = Ocular Bulge Index HWl/HW2x 100

OMI = Oculomandibular Index EF/MAFx 100

MNI = Mesonotum Index (MTF+MTW)/(MFF+MFW)

MTI = Mesoscutum Index MTW/MTF x 100

FI = F emora Index PFF/MFF x 100

PTI = Petiole Index PTH/PTF x 100

Repositories

Primary reference repositories for this work are as follows:

BEBC = Brendon E. Boudinot personal collection, Davis, California, U.S.A.

CASC = California Academy of Sciences Collection, San Francisco, California, U.S.A.

INPA = Instituto Nacional de Pesquisas da Amazonia, Manaus, Amazonas, Brazil

JTFC = John T. Fongino personal collection, Salt Fake City, Utah, U.S.A.

MCZC = Museum of Comparative Zoology, Cambridge, Massachusetts, U.S.A.

MZFU = Fund Zoological Museum, University of Fund, Sweden

MZSP = Museu de Zoologia da Universidade de Sao Paulo, Sao Paulo, Brazil

PSWC = Philip S. Ward personal collection. University of California, Davis, CA, U.S.A.

UCDC = R.M. Bohart Collection, University of California, Davis, U.S.A.

Additional repositories from which material was examined to construct the key to subfamilies

are as follows:

ABS = Archbold Biological Station, Fake Placid, Florida, U.S.A.

CMWC = Christopher M. Wilson personal collection, Sam Houston State University,

Huntsville, Texas, U.S.A.

11

European Journal of Taxonomy 120: 1-62 (2015)

DLMC

DZUP

FSCA

INBC

JKWC

LACM

MEM

MLBC

MMPC

SHSUE

SUOC

UCFC

UQCC

Danny L. McDonald personal collection, Sam Houston State University, Huntsville,

Texas, U.S.A.

Cole 9 ao Entomologica Pe. Jesus Santiago Moure, Curitiba, Brazil

Florida State Collection of Arthropods, Gainesville, Florida, U.S.A.

Instituto Nacional de Biodiversidad, Santo Domingo, Costa Rica

James K. Wetterer personal collection, Florida Atlantic University, Jupiter, Florida,

U.S.A.

Los Angeles County Museum of Natural History, California, U.S.A.

Mississippi Entomological Museum, Mississippi State University, Starkville,

Mississippi, U.S.A.

MarekL. Borowiec personal collection. University of California, Davis, California, U.S.A.

Matthew M. Prebus personal collection. University of California, Davis, California, U.S.A.

Sam Houston State University Entomology Collection, Huntsville, Texas, U.S.A.

Sam Noble Oklahoma Museum of Natural History Collection, University of Oklahoma,

Norman, Oklahoma, U.S.A.

Fullerton Collection, University of Central Florida, Orlando, Florida, U.S.A.

Universite du Quebec a Chicoutimi, Saguenay, Quebec, Canada

Results

Class Hexapoda Blainville, 1816

Order Hymenoptera Linnaeus, 1758

Suborder Apocrita Latreille, 1810

Infraorder Aculeata Latreille, 1802

Superfamily Formicoidea Latreille, 1804

Family Formicidae Latreille, 1809

Diagnosis

Aculeate Hymenoptera with the following apomorphies:

1. Eusocial, wingless worker caste present, colonies perennial (note 1).

2. Sexuals with synchronous nuptial flights (note 2)

3. Head capsule prognathous (worker, gyne) (note 1).

4. Infrabuccal sac present between labium and hypopharynx (note 1).

5. Antenna geniculate between long scape and funiculus (worker, gyne) (notes 1, 3).

6. Disticoxal foramen directed laterally and completely enclosing protrochanteral base, including

protrochanteral condyles, such that all disticoxal membrane concealed (all castes, Fig. 3C) (note 4).

7. All meso- and metacoxal cavities small, circular, monocondylic, ventrally-directed, and disticoxae

strongly produced laterally (all adult castes, Fig. 3C) (note 5).

8. Metapleural gland present (adult castes, but see note 6).

9. Propodeal spiracle located on lateral propodeal face distant from the anterodorsal propodeal comer,

often near propodeum midlength (all adult castes) (note 7).

10. Wings of alate gyne deciduous, being shed after copulation (note 1).

11. Forewing 3rs-m and 2m-cu absent (note 1).

12. Hindwing C not extending along anterior margin, even spectrally (note 8).

13. Hindwing basal/radial cell not produced distally (alate castes) (note 9).

14. Metasoma petiolate (abdominal segment II differentiated from segment II,I which is strongly

constricted between the pre- and postsclerites) (all castes), extremely rarely (~ 1 species) abdominal

segment III not constricted between pre- and postsclerites (notes 1 and 10).

12

BOUDINOT B.E., Contributions to the Formicidae

Additional, non-synapomorphic characters of value for diagnosis and identification include: Antenna

with 4-12 ante nn omeres (female) or 5-13 antennomeres (male) (note 11). Bulbus neck (= radicle) and

scape with common axis. Epicnemium extremely reduced, not visible in situ (note 12). Abdominal

segment II with sternum and tergum equally sclerotized. Pterostigma present or absent (note 13). Wing

venation variable, may be extremely reduced, with at minimum no closed cells (note 14). Jugal lobe

present or absent; abdominal sternum IX may be complex and modified apically (including prongs,

teeth, and lobes).

Notes

1. Noted as apomorphic by Bolton (2003).

2. Bolton (2003) indicated that “sexuals with mass nuptial flight” was an apomorphy of the F ormicidae.

Although mass flights do occur in several lineages of ants, it is not clear if the ancestral condition for

the Formicidae is to release large quantities of sexuals. The wording has been specifically rephrased

here to account for this uncertainty.

3. Males of many species have derived geniculate antennae with elongate scapes, including numerous

Myrmicinae, most Formicinae, and Tapinoma (Dolichoderinae). Most males, including poneroids

and numerous formicoids, however, have antennae which are not geniculate and have very short

scapes.

4. The procoxa of Formicidae is characteristically modified. The trochanteral foramen (situated apically

on the procoxa) is directed laterally and entirely enclosed, revealing no membrane in undamaged

specimens (Fig. 3C, left column, top row). Medially, the foramen is closed by an unfused seam of

the anterior and posterior apical coxal lobes, which completely surround the anterior trochanteral

process. The axis of coxal-trochanteral articulation, rather than being lateromedial as in Symphyta

(Fig. 3A), or rotated obliquely as in many Aculeata (Fig. 3B), is almost entirely anteroposterior. Feg

adduction and abduction occurs along this anteroposterior axis in more-or-less one plane of motion,

with the trochanter rotating within the closed disticoxal foramen. The coxae and their articulations

with the mesosoma and trochanters are poorly studied and show promise for valuable systematic

characters. Previous work on hymenopteran coxae include Johnson (1988), which solely focused on

the basicoxite and its musculature, Michener (1981), which focused on the meso- and metacoxae

of the Apoidea, and Vilhelmsen et al. (2010), which operationalized several coxal characters. This

character is unique to the Formicidae.

5. The meso- and metacoxal foramina are monocondylic, bearing only the medial coxal articular

processes and lacking the lateral coxal articular processes of the meso- and metapleurae. Fateral

condyles are lacking in the examined species of Chyphotinae, Bradynobaenidae s. str. , Mutillidae,

and Myrmosidae.

6. The metapleural gland, so distinctive of the female castes, is variably developed in males and has

been lost in various taxa.

7. The “high and far forward” placement of the propodeal spiracle remarked upon by Bolton (2003)

as a plesiomorphy for the Formicidae is actually an apomorphy for the family. In non-formicid

Aculeata (including Apoidea, Scoliidae, and Bradynobaenidae s. str.) the propodeal spiracle is

usually situated at the extreme anterodorsal corner of the propodeum, usually within a propodeal

spiracle length from the metanotum, and often on the dorsal propodeal face. Some Pompilidae and

Tiphiidae (Tiphiinae) have the spiracle situated more posteriorly. Although the propodeal spiracle of

'fSphecomyrma freyi is situated high—but laterally—and rather anteriorly (Wilson et al. 1967), it is

clearly not at the extreme of other Aculeates. Other '\Sphecomyrma species have more posteriorly

situated spiracles which are clearly situated laterally (Wilson 1985; Engel & Grimaldi 2005). The

potential male of "fSphecomyrma identified by Grimaldi et al. (1997) has a low spiracle situated at

about segment midlength.

8. Reduction of the hindwing costal vein occurs sporadically in other aculeate families.

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European Journal of Taxonomy 120: 1-62 (2015)

9. The basal/radial cell has been convergently reduced or lost in several ant subfamilies, and has been

lost in Mutillidae, Myrmosidae, Bradynobaenidae s. str. , and Chyphotidae. The generality of this

trait in these families was not evaluated.

10. The male of an unidentified Protanilla (Leptanillinae) from Thailand has secondarily lost petiolation,

where the third abdominal segment is no longer constricted between the pre- and postsclerites (Fig.

10A). These males are still recognizable as ants by the closed apical procoxal foramen, ventrally-

directed meso- and metacoxal cavities, and low and lateral propodeal spiracle. Other Pro tanilla species

(even in sympatry) retain the constriction, while yet others have petiolation of the third abdominal

segment (Fig. 10B). Some males of the Dolichoderinae (e.g., Azteca) and other unidentified males

of the Teptanillinae have very reduced petioles, but these are still distinctly differentiated from the

third abdominal segment and are slightly posteriorly constricted.

11. Antennomere count for males usually 13, less often 8-12 (count of 8 observed in Acropyga and

Stenamma; counts of 10+ more common). Ante nn omere counts may be extremely reduced in

inquilines, for example in Pheidole acutidens , which occasionally have an antennomere count of 5,

although this is variable infraspecifically, and indeed may vary between the left and right antennae.

12. Brothers (1975) contends that the form of the formicid epicnemium is unique, being highly reduced,

fused to and extending over the height of the mesepisternum, and obscured by the pronotum. This

putative homology was not evaluated in the present work.

13. The pterostigma is lost in most Teptanillinae, some myrmicine genera, and some species of

Leptomyrmex (Dolichoderinae).

14. No closed cells are observed in some males of Teptanillinae and Myrmicinae.

Remarks

The Formicidae is an unequivocally monophyletic group, previously defined by Bolton (1994, 2003)

as eusocial, sexually dimorphic aculeate Hymenoptera bearing metapleural glands and geniculate

antennae, among other characters. Several previously unreported synapomorphies exist for the family,

including a suite of adaptations for terrestrial locomotion (characters 6 and 7). The “low and lateral”

propodeal spiracle placement may also be an adaptation for terrestrial locomotion, as it may reduce the

distance oxygen would need to diffuse to leg locomotor muscles. While this does not clarify whether

the ancestral ant was hypogaeic or epigaeic, it does indicate that terrestrial locomotion was a crucial

transition for the Formicidae, as these apomorphies are present in all adult castes of the family. Previous

diagnoses of the family (Brothers 1975; Gauld & Bolton 1988; Goulet & Huber 1993) were significantly

improved by Bolton (1994, 2003). Characters indicated in the family diagnosis by Bolton (2003) and

above will be valuable to evaluate for critical fossil taxa such as f Armania Dlussky and other fossils

assigned to the fArmaniidae whose relationship to the Formicidae is uncertain (see Dlussky 1975:

' \Archaeopone , f Dolichomyrma, ‘\Poneropterus , "fPseudarmania; Dlussky 1983: 1 \Armaniella ; Dlussky

1999: f Khetania; Dlussky et al. 2004: f Orapia; also see discussion in LaPolla et al. 2013).

Key to global subfamilies , based on extant males

Notes: It may be expected that some taxa will fail this key, as males of at least 70 genera are undescribed

and/or uncollected (see subfamily diagnoses for specific genera). This key treats alate males only,

because ergatoid males are identifiable similarly to the workers. Ergatoid males generally bear shorter

scapes and more antennomeres than the female, and are known for about a dozen genera.

1 Apicolateral corners of abdominal sternum IX pronged or toothed (even if minutely) (Fig. 4A-B)... 2

- Apicolateral corners of abdominal sternum IX lobed or rounded (Fig. 4C). 3

14

BOUDINOT B.E., Contributions to the Formicidae

2 Petiole hatchet-shaped in profile view, with distinct peduncle, and node with distinct anterodorsal

angle in profile view (Fig. 4D). Clypeus well-developed, with conspicuous convex median disc.

Pretarsal claws cleft. Pygostyles present. Neotropical. Paraponerinae ( Paraponera )

- Petiole nodiform, fusiform, subrectangular, or cylindrical in profile view, with or without a distinct

peduncle, and node without anterodorsal angle in profile view (Fig. 4E). Clypeus poorly developed,

more-or-less linear, without conspicuous convex median disc. Pretarsal claws not cleft; claws edentate

ortoothed. Pygostyles absent. Global.Dorylinae (part, excluding Leptanilloides genus group)

3 Wing venation reduced to extremely reduced, with at most only Sc+R+Rs, Rsfl, Mfl, M+Cu, and

lr-rs+Rsf4-6 tubular, at most only three closed cells present (costal, basal, subbasal) (Fig. 4F) and

propodeal lobes very inconspicuous or absent (Fig. 4G). Old World. Leptanillinae

- Wing venation more complete, often more than three closed cells present or propodeal lobes

conspicuous and present (Fig. 4H). Global.4

4 Abdominal segment III strongly reduced and differentiated from abdominal segment IV (Fig.

5A, B) and antennal toruli separated from anterior clypeal margin by at least one antennal socket

diameter and terminal abdominal tergum never produced as spine.5

- Abdominal segment III not reduced relative to abdominal segment IV or somewhat reduced,

but not differentiated from abdominal segment IV (Fig. 5C) or antennal toruli separated from

anterior clypeal margin by less than one antennal socket diameter or terminal abdominal tergum

posteriorly produced as spine. 8

5 Metatibia with 2 ventroapical spurs (anterior spur may be reduced in size). 6

- Metatibia with at most 1 ventroapical spur. 7

6 Jugal lobe present. Frontal carinae usually robust and conspicuous (Fig. 5D). Cuticle very thick and

usually coarsely sculptured. Australia.Myrmeciinae, part (Myrmeciini: Myrmecia)

- Jugal lobe absent. Frontal carinae fine, inconspicuous, or absent (Fig. 5E). Cuticle thin and usually

finely to not sculptured. New World; African, Asian, Australian.Pseudomyrmecinae

7 Abdominal tergum IV strongly and evenly convex in profile view and much longer than abdominal

sternum IV (Fig. 5F, black lines). Helcium supraaxial: Anteroposterior axis of helcium situated well

above anteroposterior axis of abdominal segment III postsclerites, such that poststernite with very

dorsoventrally tall anterior face relative to anterior face of posttergite (Fig. 5F, dark green lines

along anterior faces of abdominal segment III). Spiracle of abdominal tergum IV located in extreme

anteroventral corner (within at least five spiracular diameters).Agroecomy rmecinae ( Tatuidris )

- Abdominal tergum IV weakly or unevenly convex in profile view and about as long as

abdominal sternum IV (Fig. 5G, black lines). Helcium axial: Anteroposterior axis of helcium

situated at about midheight of abdominal segment III postsclerites, such that anterior faces

of postsclerites roughly equivalent, or anterior face of posttergite somewhat longer than that

of poststernite (Fig. 5G, green lines). Spiracle of abdominal tergum IV located distant from

antero ventral corner (distant by at least ten spiracular diameters). Myrmicinae

8 Abdominal segment IV with cinctus (constriction) between the pre- and postsclerite (Fig. 5H) or

jugal lobe present or oblique mesopleural sulcus absent or indistinct. 9

- Abdominal segment IV without a cinctus (Fig. 51) and jugal lobe absent and oblique

mesopleural sulcus always present. 19

9 Antennal toruli situated anteriorly, abutting, very nearly abutting, or overhanging anterior clypeal

margin; toruli less than one antennal socket diameter from anterior clypeal margin in frontal view

(Fig. 6A). 10

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European Journal of Taxonomy 120: 1-62 (2015)

- Antennal toruli situated posteriorly; toruli distant from anterior clypeal margin by at least one

antennal socket diameter in fiill-face view (Fig. 6B). 12

10 Abdominal segment IV with distinct cinctus between pre- and postsclerites. Oblique

mesopleural sulcus present (Fig. 6D) or absent. 11

Abdominal segment IV without cinctus between pre- and postsclerites. Oblique mesopleural

sulcus absent (Fig. 6C). Dorybnae (part, Leptanilloides genus group)

11 Oblique mesopleural sulcus present (Fig. 6D). Mandibles triangular.Proceratiinae (part)

Oblique mesopleural sulcus absent (Fig. 6C). Mandibles nearly linear.

. Ponerinae (Ponerini part, Dolioponera)

12 Mandibles triangular, worker-like, with distinct and elongate masticatory margin (Fig. 6E)... 13

Mandibles reduced, linear, spatulate, or falcate, without distinct masticatory margin (Fig. 6B, F-G)

... 16

13 Abdominal segment IV without cinctus between pre- and postsclerites (Fig. 5C). Petiolar

tergum and sternum completely fused in anterior third, without visible suture (Fig. 6H).

Mandibular teeth robust. Myrmeciinae, part (Prionomyrmecini: Nothomyrmecia)

Abdominal segment III with cinctus between pre- and postsclerites (Fig. 5H). Petiolar tergum

and sternum not insensibly fused in anterior third; if fused, suture visible along entire petiolar

length (Fig. 61). Mandibular teeth absent, fine, or robust. 14

14 Metatibia with one ventroapical spur or prora thin and anteriorly directed, extending beneath helcium

. 15

Metatibia with two ventroapical spurs and prora absent or thick and directed ventrally, not

extending beneath helcium. Ponerinae (Platythyreini: Platythyrea)

15 Crossvein lm-cu present, thus discal cell 1 closed (Fig. 7A). Mandibles dentate; at least two

teeth present on masticatory margin. Ectaheteromorph clade

Crossvein lm-cu absent, thus discal cell 1 open (Fig. 7B). Mandibles edentate or masticatory

margin produced apically as single tooth. Proceratiinae (part, Discothyrea )

16 Mandibles falcate (Fig. 6G) to narrowly linear (Fig. 11A-B). 17

Mandibles nub-like (Fig. 6F), spiniform (Fig. 6B), or spatulate (broad in profile view). 18

17 Petiolar tergum and sternum clearly delineated. Anterior clypeal margin with (Fig. 6G) or

without pegs. Abdominal segment III about same size as segment IV, metasoma after petiole

well-sclerotized. Global. Amblyoponinae (excluding Apomyrma)

Petiolar tergum and sternum smoothly fused (similar to Fig. 6H, but along entire petiolar

length; note that longitudinal line on petiole in Fig. 11C is a carina unassociated with sclerotic

margins). Anterior clypeal margin without pegs (Fig. 11A-B). Abdominal segment III slightly

reduced relative to segment IV, although metasoma after petiole weakly sclerotized (Fig. 11C).

Endemic to Amazon basin. Martialinae ( Martialis )

18 Mesosoma anteriorly elongated: Mesonotum almost twice as long as broad in dorsal view,

and lateral pronotal face longer in profile view than head in full-face view (Fig. 1C).

Pterostigma absent. Petiole broadly attached to abdominal segment III, node weak. Afrotropical

. Amblyoponinae (part, Apomyrma)

Mesosoma not anteriorly elongated: Mesonotum much less than twice as long as

broad in dorsal view, and lateral pronotal face in profile view shorter than to as

16

BOUDINOT B.E., Contributions to the Formicidae

long as head length in profile view (Fig. 7D). Pterostigma present or absent. Petiole

very narrowly attached to abdominal segment III, node strong, except Simopelta

(Neotropical). Ponerinae (Ponerini part)

19 Basimere strongly developed; distinct from and usually much larger than telomere. Telomere

restricted to posterior apex of basimere, not or only slightly extending anteroventrally beneath

basimere (Fig. 7E). Petiole narrowly or broadly attached to abdominal segment III. Masticatory

margin of mandible often finely serrate. Ante nn a with 11-13 antennomeres. Dolichoderinae

Basimere weakly developed; usually indistinct from and usually about the same size as telomere.

Telomere extending anteroventrally beneath basimere almost to base of paramere (Fig. 7F).

Petiole narrowly attached to abdominal segment III. Masticatory margin of mandible never

serrate. Ante nn a with 8-13 ante nn omeres. 20

20 Forewing venation nearly complete: Mf3-4 and 2rs-m present, thus submarginal cell 2 closed

(Fig. 7A, SMC2). Marginal cell 1 extremely long, at least one-third chord length of wing.

Petiolar peduncle long and slender; node short, dorsoventral height somewhat less than maximum

diameter of posterior petiolar foramen (Fig. 15C). Sri Fanka. Aneuretinae (Aneuretus)

Forewing venation reduced: Mf2-4 and 2rs-m absent, thus submarginal cell 2 open (Fig. 7B).

Marginal cell 1 length less than one-third chord length of wing. Petiolar peduncle short to absent

(Fig. 16B), not particularly slender when developed; node variable. Global. Formicinae

Treatments of focal taxa

Amblyoponinae Forel, 1893

Fig. 6G

Amblyoponinae Forel, 1893: 195 (as subfamily of Formicidae). Type genus: Amblyopone.

For taxonomic synopsis of Amblyoponinae, see AntCat (2014).

Synapomorphies

The Amblyoponinae was diagnosed by Bolton (2003) based primarily on the female castes, while more

recently Yoshimura & Fisher (2012a) diagnosed the males for the Malagasy region. Apomyrma and

Opamyrma violate most of these characters; these violations are noted below. Synapomorphies of the

Amblyoponinae from the two aforementioned resources are as follows, with respective pleisiomorphies

presented in brackets:

1. Dentiform clypeal setae present on anterior clypeal margin (all adult castes) (note 1). [Dentiform

clypeal setae absent.]

2. Metapleural gland orifice directed more-or-less posterodorsally (female castes) (note 2). [Metapleural

gland orifice directed laterally.]

3. Helcium supraaxial, thus petiole situated high on abdominal segment III, petiole without distinct

posterodorsal face and abdominal tergum III without distinct anterodorsal face (all adult castes)

(note 3). [Helcium infraaxial.]

4. Petiole very broadly attached to abdominal segment III (all adult castes) (note 4). [Petiole narrowly

attached to abdominal segment III.]

5. Helcial sternite very wide (all adult castes) (note 4). [Helcial sternite narrow.]

6. Abdominal segment IV tergosternal fusion present (all adult castes) (note 5). [Abdominal segment

IV tergosternal fusion absent.]

7. Basivolsella with ventroapical process, near bases of cuspis and digitus (male) (note 6). [Apical

process of basivolsella absent.]

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European Journal of Taxonomy 120: 1-62 (2015)

Fig. 4. Male morphology. A-C. Abdominal sternum IX in ventral view. B. Oblique. D-E. Petiole in

lateral view. F. Forewing in dorsal view. G-H. Propodeum in lateral view. — A. Aenictogiton indet.

(Zambia, CASENT0106126, M. Branstetter). B. Cerapachys “parasyscia” lineage (Kenya, B. Boudinot).

C. Emeryopone buttelreepeni (Thailand, CASENT0278779, B. Boudinot). D. Paraponera clavata

(?Panama, B. Boudinot). E. Cerapachys lividus (Madagascar, CASENT0138502, D. Raharinjanahary).

F. Phaulomyrma indet. (Thailand, UCRENT150358, A. Nobile). G. Leptanillinae indet. (Thailand,

CASENTO156249, B. Boudinot); arrow indicates dorsal margin of petiolar foramen. H. Adelomyrmex

dentivagans; arrow indicates propodeal lobe. All scale bars = 0.2 mm, except D-E =1.0 mm.

18

BOUDINOT B E., Contributions to the Formicidae

Fig. 5. Male morphology. A-C, F-I. Metasoma, lateral view. D-E. Head capsule, frontal view. —

A. Myrmica latifrons (U.S.A., CASENT0104816, A. Nobile). B. Pseudomyrmex indet. (Mexico,

CASENT0103327, A. Nobile). C. Nothomyrmecia macrops (Australia, CASENT0902784, Z.

Lieberman); note that abdominal segment III is reduced relative to, but not differentiated from

segment IV. D. Myrmecia pilosula (Australia, CASENT0902800, Z. Lieberman). E. Tetraponera

indet. (Madagascar, CASENT0053316, A. Nobile). F. Tatzuidris tatusia (Panama, CASENT0178870,

E. Prado). G. Acromyrmex volcanus (Costa Rica, INBIOCRI001283114, E. Ortega). H. Platythyrea

arthuri (Mayotte, CASENTO132466, E. Prado). I. Formicapallidefulva (U.S.A., CASENT0172882, A.

Nobile). Scale bars: A-F = 0.5 mm, G-I =1.0 mm.

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European Journal of Taxonomy 120: 1-62 (2015)

Fig. 6. Male morphology. A-B, E-G. Head capsule in frontal view. C-D. Mesosoma, lateral view. H-I.

Petiole, ventrolateral view. —A. Probolomyrmex indet. (Madagascar, CASENT0080551, A. Nobile).

B. Anochetus boltoni (Madagascar, CASENT0063847, A. Nobile). C. Leptanilloides gracilis

(Guatemala, CASENT0234561, M. Borowiec). D. Odontomachus simillimus (Seychelles,

CASENTO172666, A. Nobile). E. Platythyrea arthuri (Mayotte, CASENT0132466, E. Prado).

F. Protanilla indet. (Thailand, CASENTO 119776, A. Nobile). G. Stigmatomma indet. (Madagascar,

CASENT0007087, E. Prado). H. Nothomyrmecia macrops (Australia, B. Boudinot). I. Neoponera cf.

apicalis (Honduras, B. Boudinot). Scale bars: A, F-G= 0.1 mm, B = 0.2 mm, C = 0.25 mm, E, H-I =

0.5 mm, D = 1.0 mm.

20

BOUDINOT B.E., Contributions to the Formicidae

Fig. 7. A-B. Forewing. A. Ventral view, gyne. B. Dorsal view, male. C-D. Mesosoma, lateral view,

male. E-F. Paramere, lateral view, male. — A. Aneuretus simoni (Sri Fanka, CASENTO172259, A.

Nobile). B. Nylanderia vividula (U.S.A., CASENT0058918, A. Nobile). C. Apomyrma stygia (Central

African Republic, CASENT0086073, E. Prado). D. Anochetus boltoni (Madagascar, CASENT0063847,

A. Nobile). E. Dolichoderus validns (Costa Rica, INB0003662427, B. Boudinot). F. Formicapacifica

(U.S.A., JTFC000006350, B. Boudinot). Scale bars: A-C, E-F = 0.5 mm, D = 1.0 mm.

21

European Journal of Taxonomy 120: 1-62 (2015)

Notes on synapomorphies

1. Generally present in males, although these setae may be difficult to ascertain or absent in very small

species. Not present in any other extant ant taxon, although present in most f Sphecomyrminae.

Apomyrma and Opamyrma workers and Apomyrma males lack dentiform clypeal setae (the male

of Opamyrma is unknown); rather, the workers have dentiform setae on the labrum. The male of

Apomyrma has a very reduced labrum which lacks dentiform setae.

2. Males with or, more often, without metapleural gland orifice. The metapleural gland of Apomyrma

was not evaluated in this study due to insufficient magnification.

3. This corresponds to the third, fifth, and sixth amblyoponine synapomorphies of Bolton (2003).

Apomyrma workers and males have infraaxial helcia, while the worker of Opamyrma has an axial

helcium.

4. Petiole very narrowly attached in worker Apomyrma and Opamyrma. The petiolar conformation of

the male of Apomyrma, though broad, still differs from that observed in Amblyoponinae (see note 5).

5. Reversed in Adetomyrma female castes and variable in males (Yoshimura & Fisher 2012a).

6. Character from Yoshimura & Fisher (2012a) and confirmed here via dissection of Amblyopone ,

Myopopone, Onychomyrmex, Paraprionopelta , and New World Stigmatomma.

Comments

The supraaxial helcium serves to distinguish both female and male Amblyoponinae, Qxcluding Apomyrma

and Opamyrma , from the majority of the Formicidae. Besides occurrence in the amblyoponines, the

supraaxial state of the helcium is only developed in Acanthostichus (Dorylinae) and male Proceratium

(Proceratiinae), and wealdy in the workers of Martialis (Martialinae) and the male of Tatuidris

(Agroecomyrmecinae). Males of three amblyoponine genera ( Bannapone , Concoctio , and Opamyrma)

remain unknown.

The Amblyoponinae was only recently split from the Ponerinae sensu lato (Bolton 2003) and has been

recovered in all molecular phylogenies as poneroids, with uncertain relationship to the Proceratiinae and

the remainder of the group (Brady et al. 2006; Moreau & Bell 2013; Ward 2014). Two main clades are

recovered in the Amblyoponinae, termed the XMAS ( Xymmer , Myopias , Adetomyrma , Stigmatomma)

and OCP ( Onychomyrmex , Concoctio , Prionopelta) clades (Yoshimura & Fisher 2012a). Apomyrma , in

its original description (Brown et al. 1971), was proposed to be closely related to the Amblyoponinae

(then Amblyoponini), a contention supported by Wheeler & Wheeler (1985) and Holldobler & Wilson

(1990). Other authors demurred, placing the genus in its own tribe (Apomyrmini) in the Ponerinae sensu

lato (Dlussky & Fedoseeva 1988) and in the Teptanillinae (Bolton 1990b; Kugler 1992), and eventually

in its own subfamily, Apomyrminae (Baroni Urbani et al. 1992; Bolton 1994, 2003). The “ Apomyrma ”

sequences used by Saux et al. (2004) were contaminated (PS. Ward, pers. comm.), but fortuitously their

transfer of Apomyrma to the Amblyoponinae was supported by subsequent studies (Brady et al. 2006;

Moreau & Bell 2013). This classification is followed here.

Apomyrma Brown, Gotwald Jr. & Tevieux, 1971

Apomyrma Brown, Gotwald & Tevieux, 1971: 259. Type-species: Apomyrma stygia , by original

designation. Monotypic.

Apomorphies of Apomyrma

Note: Characters here indicated are apomorphic for the Formicidae generally, given Bolton’s (2003)

synthesis of plesiomorphies and novel observations.

22

BOUDINOT B.E., Contributions to the Formicidae

1. Raised clypeal disc between antennae lateromedially compressed, forming wedge-shaped process in

anterolateral view, posterolateral clypeal margins distant from antennal toruli (female castes) (note

1). [Clypeal disc broad, uncompressed.]

2. Axillae enlarged, meeting medially (male). [Axillae small, not meeting medially.]

3. Transverse sulcus posterior to helcial sternite present (female castes). [Transverse sulcus posterior

to helcial sternite absent.]

4. Abdominal segment III posttergites unfiised (all castes).

5. Spiracles of abdominal segment III enlarged and situated at extreme anterior margin of tergum in

profile view (female castes) (note 2). [Spiracle smaller, situated distant from anterior tergal margin

in profile view.]

6. Penisvalvae dorsally fused for most of length, and anterodorsally fused with basimeres (male).

[Penisvalvae unfused dorsally, unfused with basimere.]

7. Anterior base of penisvalvar lateral apodeme strongly produced laterally, forming a helmet- or

cowry-like bulbous structure. [Lateral apodeme nearly flush with to slightly raised from valviceps.]

Notes on apomorphies

1. Clypeus also lateromedially compressed in Leptanilla.

2. Although adduced as the sole synapomorphy for Apomyrma + Leptanillinae by Bolton (1990a), this

may be a convergence, as the third abdominal spiracle of Opamyrma is considerably posterior to the

anterior tergal margin in profile view.

Comments

The infraaxial helcium of the worker and male may be apomorphic, depending on placement of the

genus.

Male diagnosis

Male Apomyrma are recognizable by the combination of nub-like mandibles, anteriorly elongate

mesosoma, reduced wing venation (marginal, costal, discal, basal, subbasal cells closed; subdiscal cell 1

open; submarginal 1 closed or open; pterostigma absent; 2r-rs situated in basal half of wing), and small,

wedge-shaped petiole which is broadly and infraaxially attached to abdominal segment III.

1. Alate (Fig. 8B-C).

2. Mandalus somewhat enlarged, but clearly ringed by sclerotized mandibular cuticle in dorsal view

(Fig. 8A).

3. Mandibles strongly reduced, nub-like, lacking teeth (Fig. 8A).

4. Labrum strongly reduced, subrectangular.

5. Palpal formula 2,1, palps strongly reduced in size.

6. Ante nn a 13-merous; funiculus filiform.

7. Occipital carina absent (Fig. 8B-C).

8. Eyes situated anteriorly, malar area visible in profile view (Fig. 8B).

9. Oblique mesopleural sulcus absent (Fig. 8B).

10. Subalar sulcus broadly enlarged, larger than lower metapleural area (Fig. 8B).

11. Epimeron narrow, lamellar (Fig. 8B).

12. Metapleural spiracular plate absent (Fig. 8B).

13. Mesoscutum anteriorly elongated, with concomitant elongation of lateral pronotal face (Fig. 8B-C).

14. Notauli fine, shallowly impressed, nearly meeting at midline (Fig. 8B-C).

15. Axillae enlarged, meeting medially (Fig. 8C).

16. Scutoscutellar sulcus exceedingly fine (Fig. 8C).

17. Metapleural gland orifice conspicuous (Fig. 8B).

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European Journal of Taxonomy 120: 1-62 (2015)

18. Propodeum small, convex (Fig. 8B).

19. Propodeal spiracle small, circular, situated in anteroventral sector of lateral propodeal face (Fig. 8B).

20. Propodeal lobe absent (Fig. 8B).

21. Ventrolateral meso- and metapleural margins ecarinate (Fig. 8B).

22. Velum of calcar absent.

23. Forewing lacking membrane anterior to costal vein (Fig. 9A).

24. Forewing venation Ogata type IVa: Submarginal cell, marginal cell 1, and discal cell 1 closed;

subdiscal cell 1 open (Fig. 9A).

25. Forewing costal vein present, reaching 2r-rs (Fig. 9A).

26. Hindwing venation reduced, only R+Rs tubular (Fig. 9B).

27. Jugal lobe absent (Fig. 9B).

28. Petiole subsessile, conical, broadening posteriorly; posterior face weak; anterior and posterior

foramina oriented along main body axis (Fig. 8B).

29. Petiolar tergum lacking anterior parabolic carina (basipetiolar carina).

30. Subpetiolar process absent (Fig. 8B).

31. Helcium infraaxial (Fig. 8B), broad in dorsal view.

32. Prora of abdominal sternum III absent (Fig. 8B).

33. Abdominal segment III undifferentiated from IV (Fig. 8B).

34. Cinctus between abdominal segment IV pre- and postsclerites absent (Fig. 8B).

35. Abdominal tergum VIII posterior margin parabolic, unmodified (Fig. 8B).

36. Abdominal sternum VIII visible in situ.

37. Abdominal sternum IX unmodified; neither pronged nor toothed (Fig. 9C).

Distribution of Apomyrma

Afrotropical, confirmed from: Cote d’Ivoire, Ghana, Benin, Nigeria, Cameroon, Central African

Republic, Democratic Republic of the Congo (new record), and South Africa.

Apomyrma stygia Brown, Gotwald Jr. & Levieux, 1971

/V ___ _

Apomyrma stygia Brown, Gotwald Jr. & Levieux, 1971: 264, figs. 1—19 (worker, queen, male). COTE

D’IVOIRE, Lagunes: Station d’Ecologie de LAMTO, near Toumodi, 6°13’N 5°02’W, 75-120 m

elevation, 16 Apr. 1968 (J. Levieux) [UCDC paratype worker CASENT0260454 examined] [Original

label reads: “IVORY COAST Lamto, Toumodi, 16.iv.68 J. Levieux”].

Apomyrma CD01

Figs 7C, 8-9

Non-type material examined (male)

DEMOCRATIC REPUBLIC OF THE CONGO, Bandundu: Wamba, Kikongo Mission, 4°15’S 17°10’E,

350-500 m elevation, Jul. 2008, Malaise trap (T. Chapman), 20 Apr. 2008, forest Malaise trap (S.L.

Heydon & S.E. Stevenson), 30 Mar.-5 Apr. 2006, Malaise trap in riparian forest (S.L. Heydon & S.E.

Stevenson) and 24 Apr. 2006, Malaise trap in riparian forest (S.L. Heydon); Wamba, Nsheshe Forest NE

of Kikongo Mission, 21-28 Jul. 2008, Malaise trap in primary forest (T. Chapman). [Wamba is the name

of the river running to the east of the georeference point.]

Male description

Measurements (n=3). HL 0.36-0.41, HW1 0.32-0.38, HW2 0.45-0.51, MAL 0.06, MDL 0.03-0.04,

SL 0.06-0.07, PDL 0.05-0.06, A3L 0.06-0.08, AAL 0.11-0.14, EL 0.22-0.26, EW 0.18-0.21, OOD

0.09-0.10, LOD 0.06-0.07, MOD 0.06-0.08, ML 0.78-0.95, MLL 0.22-0.25, MLW 0.17-0.20, MTL

0.46-0.57, MTW 0.33-0.40, PFL 0.29-0.36, MFL 0.30-0.37, PTH 0.13-0.16, PTL 0.12-0.16.

24

BOUDINOT B.E., Contributions to the Formicidae

Indices. Cl 0.89-0.93, CS 0.34-0.40, SEI 356^127, SI 18.2-19.2, El 77.4-82.5, EYE 118-120, MDI

8.2-10.5, OBI 70.4-74.6, OMI 3.71-4.50, MNI 2.05-2.14, MTI 126-144, FI 93.6-98.9, PTI 97.9-

109.1. Small, mesosoma bulky and elongate; wings situated in posterior half of mesosoma (Fig. 8B).

Head (Fig. 8A). In full-face view head slightly longer than broad excluding eyes, broader than long

including eyes. Palpal formula 2,1; palps short, not reaching hypostomal margin. Stipes with carina

along medial margin. Labrum extremely reduced, very narrow and wealdy convex, glabrous; lacking

dentiform setae; lateral margins nearly contacting medial mandibular base. Mandibles strongly reduced,

nub-like, edentate; mandalus enlarged but still ringed by sclerite in dorsal view. Clypeus well-developed;

anterior margin weakly emarginate; medial clypeal portion maximum anteroposterior length between

1.5-2 maximum antennal socket diameters; posterior clypeal margin extending slightly between

antennal toruli. Supraclypeal area indistinct. Antennal toruli situated distant from anterior head margin;

anterior tentorial pits located anterior to lateral torular arches. Frons and ocellar area bulging. Occipital

carina absent, occiput obscured by vertex in full-face view. Compound eyes bulging; medial and

posterior margins weakly convex; compound eye slightly narrower posteriorly than anteriorly. Ocelli

small, situated distantly from compound eye. Hypostomal margin reduced, medial hypostoma narrowly

carinate. Antenna 13-merous; scape overall smaller than pedicel; pedicel subspherical, slightly shorter

than antennomere 3; funiculus filiform, not elongate, reaching propodeum when laid posteriorly against

mesosoma.

Mesosoma (Fig. 8B-C). Pronotal neck very short, discontinuous laterally with remainder of sclerite in

dorsal view; pronotum weakly musculated, anteromedian face linear and appressed to mesoscutum;

dorsoventral height of anteromedian pronotal face in profile view about % x mesoscutum height in

profile view; lateral pronotal face concave posterad procoxal insertion. Mesoscutum considerably longer

than broad in dorsal view (length 1.26-1.42 x width); anterior and posterolateral areas not swollen.

Notauli distinct, fine, not cross-ribbed; meeting or barely meeting at midlength, but not extending

to transscutal line. Parapsidal lines slightly longer than half mesoscutum length, slightly divergent.

Parascutal carinae fine, weakly sinuate, situated so low on mesoscutum that carinae completely obscured

by wing base. Scutoscutellar sulcus deeply impressed dorsally, V-shaped. Axillae enlarged, meeting

at body midline. Mesoscutellum at same height as mesoscutum, dorsal margin linear in profile view.

Metascutellum small, convex, not produced. Metanotal trough deep, long, elliptical, broader anteriorly.

Mesopectus lacking oblique sulcus; subalar furrow very large, about same size as petiole. Spiracular

sclerite absent. Power metapleural area offset by upper metapleural area by sulcus; upper metapleural

area anteroposteriorly narrow, convex, sclerite thinned and differentiated from lower metapleuron,

mesopectus, and propodeum. Metapleural gland orifice conspicuous, wide open, directed laterally,

slightly posteriorly. Propodeum parabolic in profile view, dorsal face shorter than and continuous with

posterior face; propodeal spiracle circular, small, situated anteroventrally. Propodeal lobe reduced to

small tubercle with dorsal, dorsoventrally-oriented carina; petiolar presclerites visible in profile view,

unobscured by propodeal lobe.

Metasoma (Fig. 8C). Petiole wedge-shaped, weakly nodiform in profile view; anteriormost portion of

petiolar tergum not offset by carina; petiolar tergum overlapping sternum; anterodorsal petiolar face

weakly concave, curving into shallow node in profile view; node with poorly-developed posterodorsal

face; petiolar sternum ventral margin convex in profile view, sternum with symmetrical curving

longitudinal carinae forming concentric long ellipses; petiolar sternum lacking process. Abdominal

segment III similar in size, shape, and sculpture to segment IV; helcium infraaxial, broad, presternite

not visible ventrad pretergite; prora absent, transverse sulcus ventrad helcial sternum absent (present in

worker). Abdominal terga IV-VIII and sterna IV-IX normally developed, not reduced or obscured in

situ. Abdominal sternum IX apical margin obtusely triangular, not pronged or toothed.

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European Journal of Taxonomy 120: 1-62 (2015)

Forewing (Fig. 9A, see note below). Tegulum reduced, elliptical, much longer than broad. Wings

hyaline, completely covered by fine setose layer. Pterostigma absent. Wing venation Ogata type IVa:

one submarginal cell and marginal cell 1 closed, lm-cu present, thus discal cell 1 closed. Costal vein

tubular, contacting Rf distally. Cells clustered in basal half of wing, with only marginal cell extending

into apical half. Rsfl less than half length of Mfl; Rsfl and Mfl parallel. Rs+M sinuate (see note on

wing venation below). 2r-rs slightly shorter than Mfl, directed posteroapically, oblique to long axis of

wing. Rsf4-6 tubular, meeting Rf, enclosing marginal cell. Mf4-6 present, tubular for a short distance

before disappearing. Crossvein cu-a tubular, situated distad Mfl. CuF slightly divergent with respect to

Mf4-6. 1A extending distad lm-cu but not enclosing subdiscal cell. Marginal and discal cells elongate;

length of each cell considerably greater than half their width.

Hindwing (Fig. 9B). Venation reduced, only R+Rs tubular. Four hamuli present.

Genitalia (Fig. 9C-H). Pygostyles absent. Abdominal sternum IX hexagonal in outline; spiculum some¬

what long, about four times longer than broad at base; anterior margins angle distinctly posterolaterally

from spiculum; anterolateral margins wealdy divergent; posterolateral margins extending to broadly

rounded apex posteromedially. Anterolateral sternal corners hooked anteriorly. Cupula dorsal and

ventral faces anteroposteriorly narrow; lateral face broad. Basimere ventromedially produced as lamina,

concealing basivolsella in ventral view. Basimere and telomere continuous in profile view; telomere

dorsoventrally narrow, dorsal telomeral margin concave, ventral margin convex, telomere weakly

upturned, apex setose and evenly rounded. Basivolsella lateromedially narrow, oriented dorsoventrally;

anterodorsal corner produced as dorsomedially-directed spur which abuts penisvalvar ventral margin

base in situ ; cuspis lobate, elongate, directed posterolaterad. Digitus elongate, arched, directed

posteromesad, apex hooked. Penisvalvae dorsomedially fused from base to almost 3 A ths their length,

and fused anterodorsally with basimeres; valvura short, linear; valviceps dorsolateral face convex,

lateral apodeme produced ventrolaterally, forming concave shell-like structure in which volsella may

rest; ventral margin strongly concave, raised dorsally and only slightly differentiated from concave

ventrolateral penisvalvar face; phallotreme situated apically, between dorsal penisvalvar margins where

penisvalvar dorsal margins curve ventrally.

Coloration. Body uniformly castaneous, legs yellowed.

S culpturation . Body weakly sculptured, mostly smooth and shining except for the following features:

head, mesonotum, metasoma posterad petiole, and legs covered with more-or-less even, somewhat

dense piligerous punctures; subalar sulcus wealdy longitudinally costulate.

Setation. All setae short to very short; head setae head coarse, decumbent to erect, evenly spaced,

apices slightly overlapping bases; mesoscutal setae similar in form and density to head, sparser mediad

parascutal carinae; mesoscutellar setae somewhat longer and more dilute than mesoscutal setae, but

otherwise similar, absent on lateral mesoscutellar face; metascutellum with few, coarse setae; pronotum

bearing comparatively fine setae anterolaterally, otherwise glabrous; mesopectus with comparatively

fine, long, and dilute setae; metapleuron and lateral propodeal face glabrous, posterior propodeal face

with setae similar to mesoscutellum; propleural and procoxal setae similar to head setae on anterior

surfaces; leg setae coarse, short, pale, appressed on most surfaces but decumbent to subdecumbent on

ventral femoral faces; only petiolar node bearing coarse setae; metasoma posterad petiole with setation

similar to head and mesoscutum.

Note

Interpretation of the abscissae enclosing submarginal cell 1 is ambiguous. The crossvein lm-cu may

be apically displaced, producing a second Rs+M abscissa which splits into Rsf2 and Mf2, or lm-cu

26

BOUDINOT B.E., Contributions to the Formicidae

Fig. 8 .Apomyrma CD01, male, photomicrographs (CASENT0086073, E. Prado). A. Head, frontal view.

B. Body, lateral view. C. Body, dorsal view. Scale bars: A= 0.2 mm, B-C = 0.5 mm.

27

European Journal of Taxonomy 120: 1-62 (2015)

? *

1 X

Bv Cu Di

H -

/

Vu , / 1 \

Pv^ vc

Fig. 9. Apomyrma CD01, male, photomicrographs. A. Forewing. B. Hindwing. C. Abdominal sternum

IX, ventral view. D. Genital capsule, dorsal view. E. Genital valves, slightly splayed and without cupula,

ventral view. F. Genital capsule, lateral view. G. Volsella and paramere, mesal view. H. Penisvalva in

situ , mesal view. Scale bars: A-B = 0.5 mm, C-H = 0.1 mm. Abbreviations: see Material and Methods.

28

BOUDINOT B.E., Contributions to the Formicidae

may be unmoved and Rsf2 may be absent. Without a transitional series it is currently not possible

to accurately determine these particular abscissal homologies, although the abscissa in question has a

thyridium (weakening), suggesting that it is 2rs-m (the latter case).

Discussion

The species-level identification of the males described here is uncertain, as unpublished molecular work

on Apomyrma has revealed more than one species in the genus (PS. Ward, unpubl. data). The male

described here, “ Apomyrma CD01”, differs from other male-based Apomyrma morphospecies by the

following characters: head capsule in full-face view shorter, compound eyes more strongly protuberant;

and ocelli large (equal in size to morphospecies Apomyrma CF01). The morphospecies probably differ in

morphometries, wing venation, and genitalic morphology, but these character systems were not evaluated

as no physical specimens of other morphospecies were available. It may be the case that of the specific

lineages of Apomyrma , the male of A. stygia will remain unknown until another nest collection is made.

A strong morphological case could be made to revive the Apomyrminae and remove Apomyrma from

the Amblyoponinae. Published molecular studies, however, recover Apomyrma within or sister to the

Amblyoponinae, but with weakly supported relationships with either the XMAS (Brady et al. 2006;

Moreau & Bell 2013) or the OCP clade (Moreau et al. 2006; Rabeling et al. 2008). Given these results,

Apomyrma is conservatively treated as an amblyoponine here, despite compelling morphological

characters suggesting a relationship with the Feptanillinae, Opamyrma , and even Martialis (see “Shared

apomorphies” section below). For further remarks on the similarities of Apomyrma to other taxa, see the

Discussion section of the Martialis heureka species account below.

Subfamily Leptanillinae Emery, 1910

Figs 4F, 6F, 10

Feptanillini Emery, 1910: 32. Type-genus: Leptanilla.

Note

All diagnoses in the references below pertain to the tribe Feptanillini; males of Anomalomyrmini undescribed.

Male references for subfamily

Wheeler 1910: 138 (diagnosis [Leptanilla ]); Wheeler & Wheeler 1930: 193 (diagnosis [Leptanilla ,

Phaulomyrma]); Morley 1939: 114 (morphology comments); Kutter 1948: 293 (diagnosis); Bernard

1967: 90 (diagnosis [Leptanilla]); Petersen 1968: 577 (generic diagnoses, discussion); Gotwald

1969: 97 (mouthparts morphology); Wheeler & Wheeler 1972: 37 (diagnosis); Baroni Urbani 1977:

430 (diagnosis, generic diagnoses); Bolton 1990b: 269 (diagnosis); Baroni Urbani et al. 1992: 316

(morphology); Ogata et al. 1995: 32 (diagnosis, genitalia); Bolton 2003: 39, 151 (diagnosis); Borowiec

et al. 2011: 11 (venation comments [Anomalomyrm ini ]).

Male diagnosis

Male Feptanillinae are recognizable by the combination of nub-like mandibles, extremely reduced wing

venation (three cells enclosed by tubular abscissae at most development: costal, basal, and subbasal

cells; no cells enclosed by tubular abscissae at least development), and absence or inconspicuousness

of propodeal lobes. Otherwise, male leptanillines are highly variable, often resembling “normal”

poneroids, although some males are so derived as to be difficult to intuitively ascribe to the Formicidae;

this modification includes even loss of abdominal segment II petiolation.

1. Mandibles strongly reduced , nub-like not meeting at head midline , or spatulate and hypertrophied

(Scyphodon) (note 1).

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European Journal of Taxonomy 120: 1-62 (2015)

2. Palpal formula 4,1 or 1,1 (note 2).

3. Clypeus usually strongly reduced such that antennal toruli situated anteriorly, separated from

anterior clypeal margin by much less than one torulus diameter; occasionally ( Yavnella , Protanilla ,

Noonilla ) antennal toruli situated about one torulus diameter from anterior clypeal margin (note 3).

4. Anterior clypeal margin without pegs.

5. Anterior tentorial pits usually situated lateral to lateral torular arch; occasionally ( Yavnella ) situated

anterolaterad torular arch.

6. Frontal carinae and lobes absent.

7. Ante nn a 13-merous; funiculus filiform to submoniliform.

8. Occipital carina absent.

9. Oblique mesopleural sulcus present, anterior terminus contacting posterolateral pronotal comer or

situated well ventral to pronotal corner.

10. Metapleural spiracular plate absent.

11. Propodeal lobes absent or very inconspicuous.

12. Metacoxal cavities closed.

13. Tibial spur formula 2s,2s; ls,2(ls,lp); ls,2s; ls,ls; 0,lp.

14. Metatarsus lacking posterolateral line of dense differentiated setae.

15. Pretarsal claws edentate.

16. Pterostigma usually strongly reduced, but may be enlarged ( Anomalomyrma , some Protanilla) (note 4).

17. Ogata wing venation type IVb (Fig. 4F), but would be type Illb should the maximum complement

of spectral veins be hypothetically tubular; at most 3 closed cells present (costal, basal, subbasal); at

the most extreme reduction only Sc+R+Rs and Rf present along anterior wing margin, with narrow

stretch of membrane present anterobasally (notes 5,6).

18. Hindwing venation reduced: all abscissae absent, or R+Rs tubular and short, or R+Rs and 1A tubular

and short.

19. Jugal lobe absent.

20. Petiolar laterotergite absent; tergum fused with sternum, suture visible.

21. Petiolar tergum not forming anteroventral collar around sternum.

22. Helcium axial or infraaxial.

23. Helcial sternite overlapped laterally by tergite, thus not visible in lateral view.

24. Abdominal segment III undifferentiated to somewhat constricted posteriorly to strongly differentiated

with posterior constriction, forming postpetiole.

25. Abdominal segment IV as long as or infrequently distinctly longer than following abdominal

segments; not vaulted.

26. Abdominal spiracles IV-VIII obscured by preceding tergites.

27. Pygostyles absent or present as extremely elongate rods.

28. Genitalia partially exserted; subject to extreme modification.

29. Basimere separated from telomere ventrally by corium or basimere and telomere fused.

30. Telomere highly variable; least modified telomeres are digitate to wedge-shaped; sometimes

telomere laminar.

31. Basivolsella lateromedially narrow in ventral view, occasionally extremely elongated.

32. Cuspis present or absent; when present usually lobate and otherwise unmodified.

33. Digitus highly variable; least modified digiti are elongate and arched.

34. Valviceps highly variable; almost always with lateral apodeme produced laterally (note 7).

Notes on diagnosis

1. Mandibles also reduced and nub-lilce in Ponerini (Ponerinae), Apomyrma (Amblyoponinae), and

some Myrmicinae (e.g., Acanthognathus, Daceton, the Adelomyrmex genus group, Myrmecina).

2. Some unidentified Protanilla males have higher palp counts, similar to workers. Future work should

establish the extent of palpomere count variation inter- and intragenerically.

30

BOUDINOT B.E., Contributions to the Formicidae

3. Anteroposteriorly broad clypeus is pleisiomorphic for Leptanillinae, and is present in Protanilla and

Yavnella.

4. The polarity of pterostigmal reduction is unclear, as Anomalomyrma and some Protanilla have an

enlarged pterostigma; it seems likely though that this is a secondary development.

5. Wing venation also reduced to the Ogata type IVb pattern convergently in other groups, including

some Myrmicinae ( Strumigenys , the Adelomyrmex genus group, and the inquiline Pheidole acutidens)

and some Proceratiinae ( Probolomyrmex ). These taxa may be distinguished from leptanillines by a

suite of characters, including presence of the propodeal lobe. Contrary to the sentiment of Ogata

et al. (1995) the reduced wing venation of the Teptanillinae is eminently valuable for diagnosis of

the subfamily. While certainly this reduction in venation may be driven by functional constraints,

the particular pattern occurring in the Teptanillinae is nearly globally unique. Although it has been

indicated that the forewing venation of Leptanilla is completely absent (Wheeler 1910; Bernard

1968; Wheeler & Wheeler 1972), no specimens were observed which had this state; at the least one

compound abscissa was present along the leading wing margin.

6. Detailed forewing abscissal development description: Costal vein present or absent, when absent

Sc+R+Rs very close to anterior wing margin (costal cell closed or open). Rsfl+Mfl tubular or

nebulous, indistinguishable from one another, or both abscissae absent (basal cell distally closed or

open). Rs+M usually absent, infrequently spectral; Rsf2+3 absent (submarginal cell 1 open). Rsf4+

tubular, continuous with 2r-rs which is directed posteroapically, ending before wing apex, or Rsf4+

and 2r-rs absent (marginal cell 1 open). M12+ absent or spectral and 2rs-m absent (submarginal cell

2 absent), lm-cu absent (discal cell 1 open). M+Cu nebulous or absent (basal cell closed posteriorly

or open). Cuf tubular to nebulous, short, or absent (subdiscal cell 1 absent). 1A tubular, partially

nebulous, or absent (subbasal cell closed or open).

7. One morphogroup of South East Asian male Leptanillinae has lateromedially compressed valviceps,

other morphogroups and genera have the lateral apodeme consistently laterally produced, and often

modified.

Taxa examined ($ = queen, S = male)

Anomalomyrma indet. [§: Indonesia]; Leptanilla africana Baroni Urbani [S'. Nigeria]; L. bifurcata

Kugler [S'. Israel]; L. islamica Baroni Urbani [S'. Yemen]; L. Israelis Kugler [S'. Israel]; L. miniscula

Santschi [S'. Tunisia]; L. swani Wheeler [S'. Australia: Queensland, W. Australia]; L. tanit Santschi [S'

Tunisia]; L. tenuis Santschi [S'. Tunisia]; L. GR01 [S' Greece]; L. GR02 [S'. Greece]; L. GR03 [S'.

Greece]; L. IL01 [S'. Israel]; L. TH01 [S'. Thailand]; L. TH02 [S'. Thailand]; L. TH03 [S: Thailand]; L.

TH04 [S'. Thailand]; L. TH05 [S'. Thailand]; L. TH06 [S: Thailand]; L. TH08 [S: Thailand]; L. TH10

[S'. Thailand]; L. ZA01 [S'. South Africa]; L. indet. [S'. Australia]; Aoo/fi//# BMNHO1 [S'. Ivory Coast];

Noonilla indet. [S'. Malaysia]; Phaulomyrma MM01 [S'. Myanmar]; Ph. TH01 [S'. Thailand]; Ph. indet.

[S'. Myanmar]; Protanilla TH01 [S'. Thailand]; Pr. TH02 [S'. Thailand]; Pr. TH03 [S'. Thailand]; Pr.

indet. [S' Indonesia]; Scyphodon cf. bruesi [S' Indonesia]; Yavnella argamani Kugler [S'. Israel]; Y

indica Kugler [S'. India]; Yavnella BEB001 [S'. Sri Lanka]; Leptanillinae ? Protanilla [S' Indonesia,

Malaysia]; Leptanillinae indet. [S'. Borneo, Democratic Republic of Congo, Indonesia, Malaysia, Papua

New Guinea, Thailand].

Distribution

Old World: Palearctic (Europe, Southern Asia, and Northern Africa), Affotropical, Australasian, and

Australian regions.

Discussion

The Leptanillinae, based on males, is defined by the apomorphies presented in the diagnosis above

(in italics). At present the subfamily is comprised of eight genera with the re-inclusion of the male-

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European Journal of Taxonomy 120: 1-62 (2015)

based taxa Noonilla Petersen, 1968 (stat. rev.) and Scyphodon Brues, 1925 (stat. rev.). Both TV. copiosa

Petersen, 1968 and S. anomalum Brues, 1925 are transferred to the Leptanillinae.

The Leptanillinae was first delimited as a tribe of Dorylinae by Emery (1910), but has consistently been

considered a distinct subfamily since Wheeler (1923), except for Bernard’s (1951) treatment of the

“Formicoidea”, in which the leptanillines were treated as a family. Opamyrma was described by Yamane

et al. (2008) who assigned the genus to the Amblyoponinae using the concept of Saux et al. (2004) for

Fig. 10. Representative males of Leptanillinae, lateral view A. Protanilla “TH01” (Thailand,

CASENT0119776, A. Nobile), arrow indicates loss of abdominal segment II petiolation. B. Protanilla

“TH03” (Thailand, CASENT0119791, E. Prado). C. Leptanilla swani (Australia, CASENT0172318,

A. Nobile). D. Protanilla sp. (Indonesia, CASENT0178838, A. Nobile), arrow indicates basolateral

basimeral process. E. Scyphodon sp. (Indonesia, MCZ155112w, A. Nobile). F. Noonilla sp., used with

permission from Petersen (1968). Scale bars: A, C, E-F = 0.2 mm, D = 0.5 mm, B = 1.0 mm.

32

BOUDINOT B.E., Contributions to the Formicidae

the subfamily, although they concomitantly referred to the genus as belonging to the Apomyrminae.

Discovery of the male of Opamyrma and the larvae and gynes of Opamyrma and Martialis are anticipated

to contribute to the resolution of the “basal ant problem”.

As noted by prior authors (e.g., BaroniUrbani 1977; Bolton 1990b; Ogata etal. 1995),theLeptanillinaeis

unfortunately subject to parallel taxonomies, with two genera known only from workers (. Anomalomyrma ,

Furcotanilla), four genera known only from males ( Noonilla , Phaulomyrma , Scyphodon , Yavnella), and

two genera known from both castes ( Leptanilla , Protanilla). The Leptanillinae is in need of generic

revision, especially given the highly variable morphologies of the males. As this is beyond the scope

of the present paper, the male of Protanilla , although recently identified by the Ant Tree of Life Team

(P.S. Ward et al. , unpubl. data; Ant Web 2014), is not described. Moreover, there is a spectacular and

perplexing diversity of leptanilline males from Southeast Asia, which await classification and association

with workers (Fig. 10).

Except for the mandibles of Scyphodon and the sexual characteristics of Noonilla, these two are “typical”

leptanillines (compare specimens in Fig. 10). They exhibit proposed formicid synapomorphies: the

prodisticoxal cavity is closed, the propodeal spiracle is situated low on the propodeum, and the metasoma

is petiolate. Placement of these genera in the Leptanillinae is supported by the presence of the following

apomorphies: mandibles nub-like to spatulate (secondarily hypertrophied in Scyphodon ); buccal

cavity reduced; medial hypostoma vestigial; clypeus strongly reduced; and Ogata venation type IVb.

The following characteristics support the placement: palpal formula 1,1; meso-segment of mesosoma

elongated anteriorly; and abdominal tergum VIII enlarged (not present in all material attributed to

Scyphodon ). Of the formicid pleisiomorphies present in these genera, the most diagnostically valuable

is absence of propodeal lobes.

Ogata et al. (1995) indicated several characters which were dubiously diagnostic for the Leptanillinae,

specifically including: reduced palp formula; elongated pronotum; fore femora shape; reduced venation;

metapleural gland absence; terminal abdominal segment reduction; cupula (= basal ring) absence;

and cuspis absent. It is agreed that palpal formula may be reduced in other Formicidae, but reduction

to 1,1 occurs in the Dorylinae ( Aenictogiton ), the Myrmicinae (e.g., Eurhopalothrix , Pheidole and

Tetramorium-cladQ inquilines, Strumigenys, Rhopalomastix ), and the Ponerinae (e.g., Hypoponera and

Simopelta). It is evident that Scyphodon and Noonilla are not closely related to the genera indicated

above. The elongate mesoscutum with concomitant posterior elongation of the pronotum is not present

in all Leptanillinae and also occurs in Apomyrma (Amblyoponinae), but is otherwise unique among the

Formicidae. The fore femoral modifications of Noonilla are autapomorphic, and are certainly sexual

characters. As discussed in note 5 of the Leptanillinae diagnosis, the reduced venation of the Leptanillinae

is highly diagnostic of the group. Certainly, of any of the dubious characters Ogata et al. indicated, the

metapleural gland absence is the least valuable and in no way supports contentions about relationships

in male ants. It is difficult to assess development of the cupula without dissecting the few specimens

of Noonilla and Scyphodon available, so evaluation of these characters is set aside for future studies.

Cuspides are present at least in some Protanilla males, and thus are not diagnostic for the Leptanillinae

on the whole, but absence may be apomorphic for the remainder of the subfamily.

Subfamily Martialinae Rabeling & Verhaagh 2008

Martialinae Rabeling & Verhaagh, in Rabeling et al. 2008: 14913. Type-genus: Martialis. Monotypic.

Apomorphies of Martialinae and Martialis

Note: Character states indicated here are generally apomorphic for the Formicidae given Bolton’s (2003)

synthesis of plesiomorphies (appendix 3; presented below in brackets) and novel observations. These

33

European Journal of Taxonomy 120: 1-62 (2015)

apomorphies may or may not be unique synapomorphies of M. heureka , but are of high diagnostic value.

Worker-based states may also apply to the gyne, which at present is unknown. Larvae are unknown.

1. Mandibles more-or-less linear, elongate, not crossing at rest (worker), mandibles narrowly linear

(male) (Fig. 11A-B) (note 1). [Mandibles triangular.]

2. Labrum with pair of trigger-hair-like setae at basolateral corners of sclerite, before margin curves

distally, and situated at about lateral margin of lateral torular arch in anteroventral view; these setae

directed dorsolaterally, contacting basal mandibular margin (worker) (note 2). [Labral trigger-hairs

absent.]

3. Clypeus strongly reduced (worker) (note 3). [Clypeus well-developed.]

4. Medial clypeal portion covered with dense patch of erect, linear setae projecting anteriorly (worker)

(note 3). [Dense setal patch absent.]

5. Antennal toruli exposed in dorsal view (worker) (note 4). [Antennal toruli partially concealed in

dorsal view.]

6. Antennal toruli situated at and projecting anteriorly beyond anterior clypeal margin (worker) (note

5). [Antennal toruli distant from anterior clypeal margin.]

7. Antennal toruli situated anterior to line drawn between anterior tentorial pits (worker) (note 5).

[Antennal toruli situated posterior to anterior tentorial pits.]

8. Antennal toruli dorsoventrally elongated (worker) (note 6). [Antennal toruli dorsoventrally short.]

9. Frons bulging medially posterior to posterior clypeal margin and between antennal toruli (worker).

[Frons not bulging anteromedially.]

10. Compound eyes absent (worker) (note 7). [Compound eyes present.]

11. Petiole completely tergosternally fused, without externally visible suture (worker, male) (note 8)

(Fig. 11C). [Petiole without tergosternal fusion.]

12. Abdominal segment III tergosternally fused (worker) (note 9).

13. Abdominal segment III reduced in size relative to segment IV, to which it is still broadly attached

(worker) (note 10). [Abdominal segment III not reduced in size relative to segment IV.]

14. Abdominal segment IV presclerite differentiated from postsclerite (worker) (note 11). [Abdominal

segment IV presclerite undifferentiated.]

Notes on diagnosis

1. Worker trait is character 1 of Rabeling et al. ’s (2008) generic diagnosis.

2. The labrum of the worker is lateromedially broad, with the apical half populated by somewhat dense

setae, while the basal half is glabrous except for a pair of elongate basolateral setae which are directed

apicolaterally, toward the basal mandibular margins. These setae are reminiscent of the trigger-hairs

of trap-jawed ants such as Strumigenys , the Daceton genus group, and the Odontomachus+Anochetus

clade (although the setae of the ponerines occur on the mandibles). At first blush it seems unlikely

that Martialis is a trap-jaw predator (Rabeling et al. 2008), as the most well-studied trap-jaw ants

( Strumigenys , Daceton genus group, Odontomachus ) all have mandibles, which are quite to very

close-set compared to Martialis. This does not preclude mandibular snapping, however, as Mystrium

(Moffett 1986; Gronenberg et al. 1998) and Protanilla (Holldobler & Wilson 1990) have mandibles

capable of rapid, forceful closure. While no apparent mandibular locking mechanism has been

identified for Martialis , a detailed study of its internal morphology has yet to be done. The putative

trigger setae of Martialis do not resemble the trigger setae found in Protanilla (Holldobler & Wilson

1990).

3. Characters 3 and 4 above were treated as character 2 in the subfamilial diagnosis of Rabeling et al.

(2008). Here they are considered independent.

4. Part of Rabeling et al. (2008) subfamilial character 5.

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BOUDINOT B.E., Contributions to the Formicidae

5. In addition to Martialis , the only ants with antennal toruli situated anterior to the anterior tentorial

pits are homoplastically derived in the proceratiine genera Probolomyrmex and Discothyrea (Keller

2011 ).

6. Rabeling et al. (2008) character 6 of the subfamilial diagnosis.

7. Character 3 of Rabeling et al. ’s (2008) subfamilial diagnosis.

8. Worker trait is character 13 of Rabeling et al. ’s (2008) subfamilial diagnosis

9. Part of Rabeling et al. (2008) character 14 of the subfamilial diagnosis; tergosternal fusion of

abdominal segment III may be an apomorphy of the poneroids.

10. Part of Rabeling et al. (2008) character 14 of the subfamilial diagnosis.

11. Generic diagnosis character 8 from Rabeling et al. (2008).

Additions to worker diagnosis of Rabeling et al. (2008)

Note: Characters indicated here are segregated from the “Apomorphies of Martialinae ...” section above

as they are of diagnostic value but are either plesiomorphies or of uncertain polarity, i.e., it is unclear

whether these traits are apomorphic.

1. Scape conspicuously curved.

2. Meso- and metacoxae very closely situated (metapleuron foreshortened).

3. Metatibia with potentially glandular patch of discolored cuticle posterobasad ventroapical spur (note

i).

4. Aroliae reduced.

5. Petiolar peduncle anteriorly delimited by parabolic carina.

6. Subpetiolar process present as small anteroventral denticle.

7. Helcium axial.

8. AIII prora carinate, transverse, lip-shaped.

9. Anterior and posterior margins of AIII postsclerites not parallel in profile view, posterior margin of

posttergite produced posteriorly.

10. Abdominal segments 4, 5, 6, and 7 about equal length.

Note

1. The potentially glandular patch of cuticle here identified is located on the apicoventral metatibial

surface basad the tibial spur, in a similar position to the confirmed metatibial glands of other

Formicidae. The cuticular patch was visible when backlit through the cleared leg of the holotype.

Unlike the remainder of the leg cuticle, this patch was clearly thick and opaque. Recorded as absent

by Baroni-Urbani et al. (1992), not recorded by Billen et al. (2013). Future specimens of Martialis

workers should be subjected to detailed SEM and TEM study.

Comments

An attempt was made to discern the palpal count of the holotype worker, but the labrum is partially

reflexed over the maxillolabial complex. It seems as if there are two maxillary palpomeres, but this could

not be confirmed in any specimen orientation or lighting. The labial palps were not visible. Rabeling et

al. (2008) recorded the propodeal lobes as absent in the worker; after comparative study of the male and

worker it is apparent that weakly developed carinae are present in the area associated with propodeal

lobes. These carinae are not obvious in perfect profile view. Their homology with propodeal lobes is

uncertain.

Male diagnosis

Uniquely identified among the global fauna by the following character combination: mandibles linear,

barely meeting at head midlength; clypeus reduced, with antennal toruli situated less than 1 antennal

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European Journal of Taxonomy 120: 1-62 (2015)

socket distance from anterior clypeal margin; notauli present; wing venation somewhat reduced, Ogata

type IVa (five cells enclosed by tubular abscissae: costal, basal, subbasal, submarginal, and marginal cells

closed); jugal lobe absent; petiolar tergum and sternum clearly fused; posterior petiolar foramen raised

dorsad anterior foramen; helcium axial; abdominal segment III reduced relative to and differentiated

from segment IV; abdominal segment IV with cinctus impressed yet indistinctly margined; pygostyles

absent; and genitalia small, relatively unmodified.

1. Alate.

2. Mandalus enlarged (Fig. 11 A-B).

3. Mandibles linear, lateral and medial margins wealdy tapered to apex; barely meeting at head midline

(Fig. 11 A-B).

4. Palpal formula 2,1.

5. Clypeus reduced, greatest anteroposterior length about 1.5 times antennal socket diameter; antennal

toruli separated from anterior clypeal margin by less than 1 antennal socket diameter (Fig. 11 A-B).

6. Anterior clypeal margin without pegs (Fig. 11 A-B).

7. Anterior tentorial pits situated posteromediad antennal toruli (Fig. 11B).

8. Frontal carinae and lobes absent (Fig. 11 A-B).

9. Ante nn a 13-merous; funiculus filiform.

10. Occipital carina reduced, not enclosing occiput.

11. Notauli present, meeting medially and extending posteriorly to transscutal line (Fig. 11C-D).

12. Scutoscutellar sulcus not ribbed (Fig. 11D).

13. Oblique mesopleural sulcus present, anterior terminus separated slightly ventrad posterolateral

pronotal corner (Fig. 11C).

14. Metapleural spiracular sclerite absent (Fig. 11C).

15. Propodeal lobes present, weakly developed (Fig. 11C).

16. Metacoxal cavities narrowly closed.

17. Tibial spur formula lp,lp.

18. Pretarsal claws edentate.

19. Pterostigma indistinct, nearly absent (Fig. 12A).

20. Ogata forewing venation type IVa (Fig. 12A): Submarginal cell 1+2 closed by Rs+M+Mf2-3; 2rs-m

present, Mf4-6 absent; marginal cell 1 closed; lm-cu absent, thus discal cell 1 open; subdiscal cell

1 open.

21. Hindwing venation reduced (Fig. 12B): Only R+Rs and 1A tubular.

22. Jugal lobe absent (Fig. 12B).

23. Petiole nodiform, with indistinct posterior face; posteriorforamen raiseddorsally above longitudinal

petiolar axis (note 1) (Fig. 11C).

24. Petiolar tergum with anterior parabolic carina (basipetiolar carina).

25. Petiolar tergum and sternum fused; laterotergite absent (Fig. 11C).

26. Petiolar tergum not forming anteroventral collar around sternum.

27. Helcium axial, broad (Fig. 11C).

28. Helcial sternite projecting ventrad lateral tergite margins.

29. Abdominal segment III weakly reduced relative to and differentiated from segment IV (Fig. 11C).

30. Prora of abdominal sternum III weak, transversely parabolic (Fig. 11C).

31. Abdominal tergum IV neither vaulted nor elongated relative to following segments (Fig. 11C).

32. Abdominal spiracle 4 exposed, 5-8 concealed by preceding tergites (Fig. 11C).

33. Abdominal sternum IX acutely triangular, apex nearly pointed; neither pronged nor toothed (Fig. 12C).

34. Pygostyles absent.

35. Genitalia small, partially exserted (Fig. 11C).

36. Cupula anteroposteriorly narrow along all faces (Fig. 12D-F).

37. Basimere continuous with telomere (Fig. 12D-H).

36

BOUDINOT B.E., Contributions to the Formicidae

38. Telomere short, digitate, extending anteroventrad beneath basimere (Fig. 12F).

39. Cuspis absent (Fig. 12E).

40. Valviceps dorsomedially fused, lobate, longer than tall, apex downtumed, two sides forming tube

(Fig. 12D, H).

Note

1. The posterior petiolar foramen is convergently raised above the anterior petiolar foramen in Tatuidris

(Agroecomyrmecinae).

Genus Martialis Rabeling & Verhaagh, 2008

Martialis Rabeling & Verhaagh, in Rabeling et al. 2008: 14914. BRAZIF. Type-species: Martialis

heureka , by original designation. Monotypic.

Martialis heureka Rabeling & Verhaagh, 2008

Figs 11-12

Martialis heureka Rabeling & Verhaagh, in Rabeling et al. 2008: 14914, figs. 1-2 (worker). BRAZIF,

Amazonas: Manaus, Headquarters of Empresa Brasileira de Pesquisa Agropecuaria (EMBRAPA)

Amazonia Ocidental, kilometer 28 highway AM 010, 2°53’S, 59°59’W, elev. 40-50 m, 9 May 2003, ex

leaf litter at dusk, primary tropical lowland rainforest. (C. Rabeling) [MZSP],

Male description

Measurements (n=3). HE 0.35-0.42, HW1 0.35-0.40, HW2 0.44-0.49, MAE 0.04-0.06, MDF 0.10-

0.11, SF 0.20-0.23, PDF 0.10-0.13, A3F 0.17-0.21, AAF 0.17-0.21, EF 0.17-0.19, EW 0.14-0.17,

OOD 0.14-0.15, FOD 0.04-0.05, MOD 0.04-0.05, ME 0.64-0.78, MFF 0.16-0.18, MEW 0.17-0.20,

MTF 0.29-0.37, MTW 0.36-0.45, PFE 0.39-0.48, MFF 0.43-0.53, PTH 0.16-0.19, PTE 0.20-0.23.

Indices. Cl 0.94-1.02, CS 0.35-0.41, SEI 83.6-86.2, SI 55.6-58.4, El 83.0-85.6, EYE 88.1-88.7, MI

26.7-27.8, OBI 80.1-82.8, OMI 3.46^1.74, MNI 2.00-2.26, MTI 79.9-83.2, FI 88.3-91.3, PTI 76.9-

83.2. Small, but body variable in overall size (Fig. 11C).

Head (Fig. 11A-B). In full-face view head about as broad as long excluding eyes, broader than long

including eyes. Palpal formula 2,1; palps short, not reaching hypostomal margin. Stipes simple, lacking

carinae on medial surface. Fabrum very small, medially emarginate, setose; lateral margins distant

from mandibular bases by somewhat less than maximum lateromedial labrum length; labrum lacking

basolateral trigger setae observable in workers. Mandibles linear, narrow; lateral and medial margins

weakly tapering to apex; masticatory mandibular margin reduced, bidentate; apical tooth asymmetrical,

larger than symmetrical basal tooth; mandalus enlarged, diameter equal to maximum mandible width.

Clypeus reduced; anterior margin broadly emarginate; medial clypeal portion maximum anteroposterior

length about 1.5 maximum antennal socket diameters; posterior clypeal margin produced between

antennal toruli. Supraclypeal area arc-shaped, anteroposteriorly longer than maximum antennal socket

diameter. Antennal toruli situated anteriorly, with anteriormost portion of torular arch anterad anterior

tentorial pit. Frons and ocellar area bulging. Occipital carina present, weakly developed, obscured

in full-face view by vertex, not enclosing occiput. Compound eyes bulging strongly; medial margin

weakly convex; posterior margin wealdy emarginate; compound eye narrower dorsally than ventrally.

Ocelli small, situated distant from compound eye. Hypostomal margin reduced, lacking lamina. Antenna

13-merous; scape longer than maximum compound eye diameter and slightly more than 2 x pedicel

length; pedicel cylindrical, long, about 4 Z> x antennomere 3 length; funiculus filiform, elongate, reaching

metasoma when laid against mesosoma.

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European Journal of Taxonomy 120: 1-62 (2015)

Mesosoma (Fig. 11C-D). Pronotal neck continuous with remainder of sclerite in dorsal view; main portion

of pronotum swollen, muscular; anteromedian pronotal face convex in profile view, short, dorsoventral

height of pronotum from pronotal neck about l A x mesoscutum height in profile view; lateral pronotal

face concave. Mesoscutum broader than long in dorsal view (length 0.80-0.83 x width); anterior and

posterolateral areas swollen. Notauli distinct, crossribbed, meeting at body midline, not extending to

transscutal line although narrow longitudinal line present from notauli to transscutal line. Parapsidal lines

impressed, slightly divergent. Parascutal carinae nearly linear; weakly sinuate. Scutoscutellar sulcus

unimpressed. Axillae small and widely situated. Mesoscutellum high and convex in profile view; not

modified. Metascutellum small, lateromedial width slightly less than one half anteroposterior length; in

profile view metascutellum strongly produced. Metanotal trough deep, small, circular. Mesopectus with

oblique longitudinal sulcus, anterior terminus of sulcus nearly contacting posterolateral pronotal corner.

Spiracular sclerite inconspicuous. Lower metapleural area strongly offset from upper metapleural area

by deep, broad, margined sulcus. Metapleural gland orifice occluded; presence of internal metapleural

gland not visible through metapleural sclerite. Propodeum parabolic in profile view, dorsal face about

as long as and continuous with posterior face; propodeal spiracle circular, small; propodeal lobe wealdy

developed, carinate, clearly visible in anterolateral oblique view.

Metasoma (Fig. 11C). Petiole nodiform, pedunculate; anteriormost portion of petiolar tergum offset

by parabolic carina; petiolar tergum and sternum fused, longitudinal lateral carinae not suggestive of

suture; petiolar node shallow, in profile view anterodorsal face nearly linear, dorsum weakly convex,

posterior face very weak; petiolar sternum linear for most of length, posteriorly narrowed, ventral

petiolar surface with paired diverging carinulae; subpetiolar process absent. Abdominal segment III

slightly reduced and differentiated from segment IV; helcium axial, sternal presclerite visible in profile

view, not obscured by tergal presclerite; abdominal posttergite and poststernite III not fused; abdominal

sternum III prora present as anterolateral bosses subtending helcium, anteromedian area of sternum

concave. Abdominal terga IV-VIII and abdominal sterna IV-IX normally developed, not reduced or

obscured in situ. Abdominal tergum VIII posterior margin unmodified. Abdominal sternum IX apically

ligulate, narrow, posterior margin very narrowly convex, nearly triangular.

Forewing (Fig. 12A). Tegulum reduced, subrectangular, longer than broad. Wings weakly infuscated,

completely covered in fine setose layer. Pterostigma poorly-developed, only anterior enclosing abscissa

tubular. Wing venation Ogata type IVa: Submarginal cell 1+2 and marginal cell 1 closed, lm-cu absent,

thus discal cell 1 open. Costal vein tubular to pterostigma. Rsfl slightly more than Vi x length of and

meeting Mfl obliquely. Rs+M continuous with undifferentiated Mf2-3 until meeting very short 2rs-

m. Rsf2+3 absent. 2r-rs very long, longer than combined lengths of Rsfl and Mfl; 2r-rs directed

posteroapically, not orthogonal with anterior wing margin. Rsf4-6 tubular to Rf, enclosing marginal

cell. Mf4-6 absent. Crossvein cu-a incompletely tubular, situated basad Mfl. Cuf divergent with respect

to Rs+M+Mf2+3. 1A extending only slightly beyond cu-a, not enclosing subdiscal cell 1.

Hindwing (Fig. 12B). Hindwing venation reduced, only R+Rs and 1A tubular; R not reaching anterior

wing margin; 1A short, weakly indicated. Three hamuli present. Claval region poorly developed.

Genitalia (Fig. 12C-H). Pygostyles absent. Abdominal sternum IX spiculum short; anterior margin linear,

curving posterolaterally near lateral margins; lateral margins short, slightly divergent; posterolateral

margins wealdy concave, tapering strongly to acute, narrowly rounded apex. Cupula dorsal and lateral

faces about as broad as telomeral base; lateral face narrowing ventrally to narrow bar-like ventral face.

Basimere and telomere more-or-less continuous, basimere weakly shouldered dorsomedially anterad

telomeral base; dorsomedian margins of basimeres parallel for about half length of paramere; telomere

acutely triangular in profile view; basimere and telomere with ventrolateral layer of posteroventrally-

directed setae. Basivolsella lateromedially broad; base transversely connected with basimere; cuspis

38

BOUDINOT B.E., Contributions to the Formicidae

absent; digitus clavate, apex swollen and directed ventrally; digital stem short. Valvura short, linear,

directed anteriorly and situated at about % valviceps height; valviceps linear, dorsoventrally short;

valviceps dorsomedially fused for most of length; in profile view dorsal valviceps margin weakly

convex, ventral margin concave, edentate, valviceps apex weakly convex, subrectangular, produced

ventrally; dorsolateral valviceps face convex, margined by lateral apodeme which extends almost to apex

before curving ventrad and contacting ventral margin; ventrolateral valviceps face concave; phallotreme

situated at aedeagal apex, sclerotic aperture formed by valviceps circular.

Coloration. Body a lm ost uniformly brown to brownish yellow; extremities slightly lighter colored.

Sc ulpturation . Body wealdy sculptured overall; head with fine piligerous punctae; mesonotal piligerous

punctae coarser, posterolateral mesoscutal area above parascutal carina roughened; dorsomedian

scutoscutellar area finely and densely anteroposteriorly striate; mesoscutellum weakly roughened;

metascutellum with fine transverse carina subtending posteriorly produced portion of disc; mesopectus

and metapleuron smooth, shining, slightly rough; propodeum finely striate, striae extending from anterior

margin down along lateral faces, dorsal propodeal face weakly rugose, posterior face mostly smooth;

petiole mostly smooth and shining, with lateral longitudinal carinulae; abdominal segment III mostly

smooth and shining; abdominal segments posterior to segment III weakly sclerotized.

Setation. Head, median pronotal portion including pronotal neck, mesonotum, and procoxae covered

by dense layer of somewhat short, uniform, weakly curved, erect to suberect setae, longer setae present

on these areas very sparsely; clypeus lacking clypeal brush of worker, although setal layer denser than

on remainder of head capsule; setae sparse and subdecumbent to nearly appressed on pronotal lateral

face, mesopectus, metapleuron, and propodeum; setae somewhat denser on metasoma, but not as dense

as on head and mesonotal dorsum; petiolar setae elongate, linear, setae on remaining segments shorter

and curved; setae on legs, including meso- and metacoxae, about as dense as on metasoma, mostly

subdecumbent with a few longer suberect setae present.

Distribution

Neotropical: Known only from the Amazon basin near Manaus (Amazonas, Brazil).

Discussion

Described from a single stray worker from the Amazon just north of Manaus, Brazil, Martialis heureka

Rabeling & Verhaagh, 2008 is one of the most significant taxa in the Formicidae described in recent years.

Displaying a bizarre mixture of pleisiomorphic and autapomorphic traits, the species was attributed to its

own subfamily, the Martialinae. This decision was supported by further morphological study (Brandao

et al. 2010) and multi-locus molecular phylogenetic reconstruction (Rabeling et al. 2008; although see

Kiick et al. 2011). Rabeling et al. (2008) recovered Martialis as the sister to all remaining extant ants

including the Old World subfamily Feptanillinae, while a reanalysis by Kuck et al. (2011) found the

converse. Given this debate and the mysterious biology of the species, Martialis is of high interest. Here

the male of Martialis is described for the first time based on material from the Biological Dynamics of

Forest Fragments Project (BDFFP [English], PDBFF [Portuguese]) study region, about 50 km north of

the type locality.

The male of Martialis differs from the worker by standard intercaste dimorphism, e.g., eyes well-

developed, ocelli and notauli present, alate, flight sclerites developed, mesosoma musculated for flight,

but the male also differs notably in several specific characters: mandibles far shorter, reduced relative

to worker; labral trigger setae absent; clypeal brush absent; antennal toruli situated more anteriorly;

forelegs weak; metatibial gland absent; petiolar node weakly developed; helcium broader; abdominal

tergum and sternum III of equal length; cinctus between pre- and postsclerites IV not developed. On

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European Journal of Taxonomy 120: 1-62 (2015)

the other hand, the male of Martialis displays numerous similarities with the worker, such as linear

mandibles, reduced palpal count (although worker count unconfirmed, but certainly less than 3,3),

clypeus poorly developed, anterior tentorial pits posteriorly-situated, antennal toruli well-developed

(but not quite as cup-like as in worker), scapes long, pedicel elongate, frontal carinae absent, propodeal

lobes wealdy developed (contra the initial worker diagnosis), tibial spur formula 1,1, basipetiolar carina

present, petiolar tergum and sternum fused, abdominal segment III differentiated from IV, and sculpture

and setation remarkably similar, although the somatic sclerites of the male are generally less strongly

sclerotized. The two castes are similar in several other specifics, but this brief list captures most of the

significant features.

Specimens were examined from collecting events in January, February, April, and October of 1985.

The BDFFP study region displays weak seasonality of rainfall and day length, with the months of

June through December roughly representing the “dry season”, July through September being the driest

(Bierregaard Jr. et al. 2001). It is possible that Martialis flights occur year-round, although the sampling

of BDFFP material examined for this study is too small to confidently assert the flight phenology. While

the range of Martialis has only been extended by about 50 km by the discovery of the male, the quantity

of males recovered exceeds that of workers by an order of magnitude (25 males vs. 3 workers). Thus,

Martialis may be recovered via alates more readily than workers. The use of Malaise traps and flight

intercept traps should be encouraged for studies of ant diversity, particularly for species with cryptic

habits. Although our knowledge of flight phenology is poor, tropical rainforests may be particularly

amenable to these studies due to the relatively more year-round flights of Neotropical (Kaspari et al.

2001a, 2001b) than of Nearctic ants (Dunn et al. 2007).

As the Manaus region is considered the ecological “crossroads” of the Amazon where a high proportion

of species ranges overlap (Bierregaard Jr. et al. 2001), it will be valuable to sample for Martialis in

other Amazonian regions. Moreover, it is of interest whether the fragmented populations of Martialis in

the BDFFP plots have survived the intervening 30 years. This may not be the case, as ant communities

have been observed to hemorrhage in BDFFP study plots (Vasconcelos et al. 2001), although hypogaeic

ants may be less sensitive to habitat changes than epigaeic ants. The placement of the Malaise samples

relative to the edges of the study plots is unknown. As well, the gyne of Martialis remains unknown. It

is possible that this caste will be ergatoid, but the presence of alate gynes cannot be ruled out. In general,

the male of a given ant species is more frequently collected via Malaise traps than females; thus it is

possible that although only males were encountered, alate gynes could still be present. Regardless, the

natural history of Martialis will be fascinating to uncover.

Material examined

Holotype worker examined at MZSP Specimen was cleared by non-destructive extraction of DNA,

which allowed for examination of internal characters. All males were examined from the following

collecting events at the Fazenda Esteio study area of the Biological Dynamics of Forest Fragments

Project in Amazonas, Brazil, with an elevation of about 90 ± 10 m, collected by Bert Klein (WWF)

via Malaise trap in 1985: Plot 1112 “Cidade Powell”, 2.38692° S, 59.87494° W, ± 100 m, 26 Feb.

(5 specimens) and 1 Oct. (1 specimen), 1 hectare Amazonian rainforest fragment; plot 1208 “Cidade

Powell”, 2.37204° S, 59.87252° W, ± 250 m, 22 Oct. (1 specimen), 10 hectare Amazonian rainforest

fragment; plot 1301 “Florestal”, 2.38897° S, 59.85012° W, ± 500 m, 23 Jan. (4 specimens), 24 Apr.

(7 specimens), and 2 Oct. (7 specimens), 100 hectare Amazonian rainforest fragment. (Note: data

extrapolated from Bierregaard Jr. et al. 2001, and PDBFF & INPA-SI 2014; latitude and longitude

recorded from Google Earth with error estimates to account for uncertainty of exact plot location; half

the male material examined remains at INPA.)

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BOUDINOT B.E., Contributions to the Formicidae

Fig. 11. Martialis heureka Rabeling & Verhaagh, 2008, male, photomicrographs. A. Head, frontal view.

B. Head, anteroventral oblique view. C. Body, lateral view. D. Body, dorsal view. Scale bars: A-B = 0.2

mm, C-D = 0.5 in in

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European Journal of Taxonomy 120: 1-62 (2015)

Fig. 12. Martialis heureka Rabeling & Verhaagh, 2008, male, wing photomicrographs and genitalia

illustrations, genital membranes not shown. A. Forewing. B. Hindwing. C. Abdominal sternum IX,

ventral view. D. Genital capsule, dorsal view. E. Genital capsule, ventral view. F. Genital capsule, lateral

view. G. Volsella and paramere, mesal view. H. Penisvalva in situ , mesal view. Scale bars: A-B = 0.5

mm, C-H = 0.1 mm. Abbreviations: see Material and Methods.

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BOUDINOT B.E., Contributions to the Formicidae

Shared apomorphies of the basal ants

Of particular interest for the “basal ant” problem (Brady et al. 2006; Rabeling et al. 2008; Ktick et al. 2011)

is the relationship of the Leptanillinae, Martialinae, and the Amblyoponinae. While the relationships

wit hin the formicoid clade (Dorylinae, myrmeciomorphs, dolichoderomorphs, Formicinae, ectahetero-

morphs, Myrmicinae) have crystallized in the past decade (Moreau et al. 2006; Brady et al. 2006, 2014;

Ward et al. 2015), the relationships of the “poneroids” (Agroecomyrmecinae, Amblyoponinae, Parapo-

nerinae, Ponerinae, and Proceratiinae) are still unresolved (Ward 2014). The poneroids may constitute a

clade or a grade, depending on placement of the root on the ant tree of life (Brady et al. 2006).

The dilemma of rooting the ant tree of life was highlighted by the discovery of M. heureka , which

has been recovered as sister to the remainder of the extant Formicidae (Rabeling et al. 2008), a result

contested by Ktick et al. (2011), who recovered Feptanillinae as sister to the extant Formicidae. These

results not only disagree in topology, but may be subject to the biases of long-branch attraction and

CG-bias (Ward 2014). As summarized by Ward (2014), one of the major questions of ant systematics

is whether Martialis and/or the Feptanillinae are sister to the extant Formicidae or whether they are

highly derived poneroids. Unpublished molecular phylogenetic analyses by PS. Ward (discussed in

Ward 2014), in which the outgroups are excluded, recover Martialis and the Feptanillinae as part of

a bipartition comprising part of the poneroids; moreover, these analyses recover Opamyrma as sister

to the Feptanillinae. Considerable uncertainty thus exists, even with molecular data. The present work

seeks, in part, to render this problem more tractable by providing novel morphological characters that

are shared by Martialis, Apomyrma, Opamyrma , the Feptanillinae, and the Amblyoponinae.

The male of Martialis is more generalized morphologically than most Feptanillinae, although some

Protanilla display a mosaic of generalized and specialized characteristics. Martialis males differ

from all known male Feptanillinae by the following characters: mandibles linear, meeting at head

midline, bidentate apically; antennal toruli situated posterad anterior portion of antennal torulus; lower

metapleuron dorsoventrally longer than anteroposteriorly broad; metanotal trough pit-like (vs. oblong);

propodeal lobe present; first submarginal cell enclosed by tubular abscissa; marginal cell 1 closed; petiole

pedunculate; genitalia more generalized. The worker of Martialis is superficially similar to Protanilla

and Anomalomyrma. The natural history of all three of these taxa is virtually unknown.

Male Martialis , in comparison with those of the Amblyoponinae, excluding Apomyrma , differ by lacking

several amblyoponine apomorphies, such as the anteroventral petiolar tergum collar, modified peg-like

setae on clypeus and labrum. Martialis further differs from the Amblyoponinae without Apomyrma

by the following characters: mandibles linear (rather than curved and subfalcate); clypeus reduced;

antennal toruli strongly developed, conspicuous (vs. inconspicuous); metascutellar trough pit-like;

petiolar tergum and sternum fused; helcium axial; and helcial stemite projecting ventrad helcial tergite

in profile view.

The male of the amblyoponine Apomyrma , however, displays several characters that are on the one hand

very leptanilline in nature and on the other are similar to Martialis. Apomyrma has reduced, nub-like

mandibles, lacks propodeal lobes, and has an infraaxial petiole, as in the Feptanillinae. Additionally, the

pro- and mesonotum is elongated, similar to most of the Feptanillini. The petiole of Apomyrma is unlike

other Amblyoponinae, as it is infraaxial and lacks the anterior tergal collar, but the petiole differs from

Martialis in being tergosternally unfused. Unlike Martialis, Apomyrma has a distinct metapleural gland

orifice and short, robust legs. Martialis is easily distinguished from Apomyrma. The male of Opamyrma

is unknown, but would inevitably be valuable to describe and compare.

Are the workers of Apomyrma, Opamyrma, and the Feptanillinae convergently similar due to subterranean

habits? Nothing is known yet of the habits of Martialis. The workers are superficially similar to the

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European Journal of Taxonomy 120: 1-62 (2015)

Sphecomyrminae in mesosoma and metasoma form relative to the Leptanillinae and Amblyoponinae,

although several sphecomyrmine genera have peg-like setae as in the Amblyoponinae. Martialis males

are superficially similar to males tentatively identified as Sphecomyrma (Grimaldi et al. 1997), differing

mainly in having antennae situated anteriorly, reduced venation, tergostemal petiolar fusion, a tibial spur

formula of 1,1, and having abdominal segment III reduced. The extinct genus Baikurus differs similarly,

except the mandibles are curved and palps are longer; the petiole is not visible in the specimen illustrated

by Grimaldi et al. (1997).

Below is presented a list of apparent morphological apomorphies shared among the Amblyoponinae

(excluding Apomyrma and Opamyrma), Martialinae, Apomyrma, Opamyrma , and Leptanillinae, with

groups presented by increasing qualitative similarity. Pleisiomorphic conditions are presented in brackets

next to the respective apomorphic conditions.

Apomorphies shared among all five taxa:

1. Compound eyes reduced (worker) (note 1). [Compound eyes not reduced in most Formicidae.]

2. Petiole anteriorly tergostemally fused (worker, gyne) (note 2). [Petiole anteriorly tergostemally

unfused in Formicidae.]

Notes:

F Proposed by Ward (1994) as a putative synapomorphy of the Amblyoponinae, Leptanillinae, and

Apomyrma , but prone to homoplasy.

2. Ward (1994) proposed this as a putative synapomorphy of the Amblyoponinae, Leptanillinae, and

Apomyrma.

Apomorphies shared by Martialis , Opamyrma , Apomyrma , and the Leptanillinae:

1. Frontal carinae lost (worker, gyne) (note 1). [Frontal carinae present, lobe-like in Amblyoponinae

and other poneroids, excepting Proceratiinae.]

2. Antennal toruli directed dorsally or anterodorsally rather than laterally (female castes) (note 1).

[Antennal toruli directed more-or-less laterally in Amblyoponinae and other poneroids, excepting

Proceratiinae.]

3. Compound eyes completely absent (worker). [Compound eyes present in most Formicidae.]

4. Occiput enclosed by occipital carina (worker, gyne) (note 2). [Occipital carina not enclosing occiput

in other Formicidae.]

5. Occiput enlarged, such that it is visible in full-face view (worker, gyne). [Occiput smaller, not visible

in full-face view in other Formicidae.]

6. Parascutal carina situated very low on mesoscutum, almost completely obscured by wing base and

thus inconspicuous (male). [Parascutal carina raised on mesoscutum, near height of mesoscutal

dorsum, only partially obscured by wing base, conspicuous; in Formicidae.]

7. Spiracular sclerite absent, metapleural spiracle unconcealed (male) (note 3). [Spiracular sclerite

present, concealing metapleural spiracle; in Formicidae.]

8. Pygostyles absent (male) (note 4). [Pygostyles present for Formicidae; see note below.]

9. Valviceps linear to arched, with concave ventral and convex dorsal margin in ectal view (male).

[Valviceps more-or-less elliptical, with both dorsal and ventral margins convex in ectal view in other

poneroids.]

10. Valviceps lateral apodeme evenly linear, extending almost to valviceps apex before curving ventrad,

delimiting convex dorsolateral and concave ventrolateral face (male). [Lateral apodeme sinuate,

wavy, extending to valviceps apex but not curving ventrad; dorsolateral and ventrolateral faces more

or less flat in other poneroids.]

11. Valviceps ventral margin edentate (male) (note 5). [Valviceps ventral margin dentate in other

poneroids.]

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BOUDINOT B.E., Contributions to the Formicidae

12. Valviceps dorsomedially fused (male). [Valviceps unfiised and articulating dorsomedially in most

other Formicidae.]

13. Characters of the sting apparatus (worker, gyne) (note 6).

Notes:

1. Also observed in Dorylinae.

2. The genus Apomyrma is an exception as no occipital carina occurs.

3. The spiracular sclerite has been lost in various lineages of Formicidae, including the entire

Myrmicinae and Tatiddris.

4. Pygostyles have been lost in numerous lineages of Formicidae and thus constitute a relatively weak

character. Within the poneroid group, the Agroecomyrmecinae, Ponerinae, and Paraponerinae retain

pygostyles; in the Proceratiinae, only Probolomyrmex has lost pygostyles. The Amblyoponinae are

somewhat more complicated, with loss of pygostyles occurring at least twice: one or more times

in the XMAS clade (. Myopopone , Mystrium + Xymmer), and once outside of the XMAS clade.

Pygostyles are retained in the OCP clade and in Stigmatomma and Adetomyrma of the XMAS clade,

thus spanning the root of the Amblyoponinae. In formicoids, loss of the pygostyles has been adduced

as an apomorphy of the Dorylinae (Bolton 1990c). The bizarre, elongate, filamentous structures of

some unassociated leptanilline males are, amazingly, appendages of the basimere (Fig. 10D).

5. Absence of ventral valviceps teeth has evolved several times, including the amblyoponine genus

Adetomyrma (Yoshimura & Fisher 2012b).

6. Kugler (1992) indicated as synapomorphic a set of sting apparatus characters applying to Apomyrma ,

Protanilla , and Leptanilla ; these characters also seem to occur in Martialis. These characters are,

in short, lack of medial lobe on quadrate plate, fulcral arm short and lacking lateral extensions.

Notably, the anal plate of Apomyrma and Protanilla is absent and no anal plate was found for

Martialis (Brandao et al. 2010); this may, however, be an artefact of the poor preservation of the

Martialis specimen examined by Brandao et al., and waits to be confirmed for Protanilla (Kugler

1992). The present author admittedly has insufficient expertise with the sting apparatus to critically

evaluate these characters, however. Future researchers are encouraged to examine the sting apparati

of the poneroids, as knowledge of this group is poor (Kugler 1992) and as understanding the sting

apparatus might assist in providing a more detailed picture of the evolution of the early branching

ant lineages.

Apomorphies shared by Opamyrma , Apomyrma , and the Feptanillinae:

F Fateral bases of mandibles set deep in pits, suggestive of a trap-jaw mechanism (female castes).

[Pleurostoma posterad mandibular insertion without a deep pit for reception of mandible; mandibles

without trap-jaw mechanism in other poneroids, excepting Mystrium.]

2. Clypeus raised dorsally, distinctly margined posteriorly (worker, gyne). [Clypeus flush with frons,

lacking posterior margination in other Formicidae.]

3. Entire clypeus anteriorly produced (not just median portion), with rectangular anterolateral margins

(worker, gyne) (note 1). [Clypeus not produced anteriorly; anterolateral corner obtuse in other

Formicidae.]

4. Fabrum with peg-lilce setae (worker, gyne) (note 2). [Fabrum without dentiform setae in other

Formicidae, excepting Amblyopone.]

5. Thickened setae present on ventromedial mandibular face (worker, gyne). [Thin setae present on

ventromedial mandibular face in other poneroids, with some exceptions.]

6. Male mandibles strongly reduced, nub-like (male) (note 3). [Mandibles long, falcate to linear in

Amblyoponinae and Martialis.]

7. Basivolsella lateromedially narrow in ventral view (male) (note 4). [Basivolsella lateromedially

broad in ventral view in other poneroids, excepting various proceratiine species.]

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European Journal of Taxonomy 120: 1-62 (2015)

8. Digital stem elongate and strongly arched (male) (note 4). [Digital stem short, linear to wealdy

curved in other Formicidae.]

9. Larva elongate, slender, and club-shaped (“leptanilloid” in terminology of Wheeler & Wheeler

1976) (note 5). [Larva pogonomyrmecoid, myrmecioid, or platythyreoid in other poneroids; see

Wheeler & Wheeler 1976 for de fini tion of larval forms.]

Notes:

1. The clypeus of Leptanilla is strongly produced as a subrectangular median process in addition to the

anterior clypeal migration.

2. Polymorphic in Leptanillinae: present at least in Anomalomyrmini. Also present in Amblyopone,

Onychomyrmex.

3. Mandibles secondarily elongated in Noonilla and enlarged in Scyphodon.

4. The genitalia of Leptanillinae are subject to extreme modification, but this state is visible in species

with more-generalized genitalia. The male of Opamyrma is unknown.

5. May apply to Martial is: larval caste unknown for Martialis and Opamyrma.

Apomorphies shared by Opamyrma and Apomyrma:

1. Promesonotal flexion extreme (worker, gyne?) (note 1). [Promesonotal articulation flexible, but not

enhanced.]

2. Propleurae bulging in profile view (worker, gyne?) (note 2). [Propleurae flat, only weakly produced

anteriorly beyond pronotum, if at all.]

3. Scapes clavate (worker, gyne?).

4. Petiolar tergum extending anteroventrally and fusing medially, forming collar (worker, gyne?) (note

3).

Notes:

1. Promesonotal flexion is enhanced in other Formicidae, including some Leptanilloides (Dorylinae),

and the Leptanillini.

2. Also present in Leptanillini, Xymmer (Amblyoponinae), and Leptanilloides.

3. The tergal-tergal fusion “collar” around the petiolar base is not present in the male of Apomyrma ,

whereas such a collar is present in both the male and worker of Xymmer.

Brief global diagnoses of subfamilies , based on males

Note

The treatment of the subfamilies below follows the current systematic classification of the Formicidae

(Bolton 2003; Brady etal. 2006,2014; Ward etal. 2015). The subfamilies are organized by the systematic

results of Brady et al. (2006) and Ward et al. (2015), with the “poneroids” in alphabetical order first,

followed by the formicoid clade comprised of the Dorylinae, myrmeciomorph clade (Myrmeciinae,

Pseudomyrmecinae), dolichoderomorph clade (Aneuretinae, Dolichoderinae), Fonnicinae,

ectaheteromorph clade (Ectatomminae, Heteroponerinae), and Myrmicinae.

Subfamily Agroecomyrmecinae Carpenter, 1930

Figs 5F, 13A-B

Diagnosis

Uniquely identified globally by petiolation of abdominal segment III (post petiole) and supraaxial

helcium. Furthermore, the petiolar comformation seems to be globally unique, with the posterior

petiolar foramen raised completely dorsad the anterior foramen. Identification may be confirmed with

the following combination of characters: mandibles reduced, edentate; antennal toruli situated distant

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BOUDINOT B.E., Contributions to the Formicidae

from anterior clypeal margin; antenna 12-merous; meso- and metatibia with one ventroapical spur each;

forewing with five closed cells; jugal lobe absent; petiolar tergum and sternum distinct; abdominal

segment IV pre- and postsclerites separated by cinctus; abdominal tergum IV vaulted; abdominal tergum

VIII not spiniform; abdominal sternum IX apex rounded.

Comments

Two extant genera are known in the Agroecornyrmecinae, the Neotropical Tatuidris and Afrotropical

Ankylomyrma (Ward et al. 2015). The male of Ankylomyrma is unknown and will be a notable discovery.

Subfamily Paraponerinae Emery, 1901

Figs 1, 2C, 3C, 4D, 13C-D

Diagnosis

The hatchet-shaped petiole (Fig. 4D) and the morphology of abdominal sternum IX are both globally

unique among the Formicidae. The ninth abdominal sternum of Paraponera is strongly produced

posteriorly as an apically bidentate linear process. These characters may be supplemented by the following

combination: mandibles triangular, unidentate; clypeus well-developed, antennal toruli situated distant

from anterior clypeal margin; antenna 13-merous; meso- and metatibiae with two ventroapical spurs

each; eight closed cells present on forewing; jugal lobe present; petiolar tergum and sternum distinct;

abdominal segment IV pre- and postsclerites separated by cinctus; abdominal tergum IV not vaulted;

abdominal tergum VIII not spiniform.

Comment

One species of the Paraponerinae is extant, Paraponera clavata. This species dwells in rainforests and

is known from Honduras through Central America into tropical South America.

Subfamily Ponerinae Fepeletier de Saint-Fargeau, 1835

Figs 4C, 5H, 6B, D-E, I, 7D, 13E-F

Diagnosis

Ponerinae share the following characters: antennal toruli situated well-posterad anterior clypeal margin

(except Dolioponera ); at least 4 closed cells present on forewing ( Dolioponera with 3); propodeal

lobes usually present; jugal lobe usually present; petiolar tergum and sternum distinct; cinctus between

abdominal pre- and posttergites IV usually present; and abdominal sternum IX unpronged and edentate.

Three final sets of characters are required for identification: 1) (Platythyreini) mandibles triangular, tibial

spur formula 2,2; 2) (Ponerini) mandibles spatulate, linear, or nub-like and mesonotum not anteriorly

elongated; and 3) (Ponerini, Dolioponera) forewing with three closed cells, propodeal lobes present,

antennal toruli situated at anterior extreme of head, oblique mesopleural sulcus absent, and cinctus

present. The eighth abdominal tergum of male Ponerinae may be spiniform, a unique state among the

Formicidae, but this character is not present in all genera and may be interspecifically variable. The spur

formula of Ponerini is variable.

Comments

The Ponerinae is global in distribution and has recently been provided a molecular phylogeny (Schmidt

2013) and a global generic revision (Schmidt & Shattuck 2014), which did not treat males; the subfamily

is now comprised of 47 valid genera. Males are unknown for 9 genera (. Asphinctopone , Austroponera ,

Boloponera, Feroponera , Fisheropone , Iroponera, Loboponera , Odontoponera , Promyopias); the males

of Belonopelta , Emeryopone. Myopias and Simopelta will be described in forthcoming publications (B.

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European Journal of Taxonomy 120: 1-62 (2015)

Boudinot in prep, and R.S. Probst et al. in prep.). The male of Dolioponera was discovered by the author

during the review period of the present work.

Unlike the female castes, which have the lateral torular arch fused with the frontal carina (Bolton 2003),

no single character was found to distinguish male Ponerinae from other subfamilies. The Platythyreini

and Ponerini share numerous characteristics but are easier to key separately due to the informatively

variable development of male mandibles. The Ponerini themselves are challenging to key as the males

of some genera are highly derived, such as Simopelta , which lacks the mesopleural sulcus and the

cinctus of abdominal segment III, and Dolioponera , which also lacks the mesopleural sulcus and has

the antennal toruli situated near the anterior head margin. A polythetic de fini tion of the Ponerini is thus

required. Fortunately, some of the derived character states of the Ponerini also serve to distinguish them

from the “generalized” ants, including the Formicinae and Dolichoderinae.

Yoshimura & Fisher (2007) and previously Yoshimura & Onoyama (2002) used the scutoscutellar

sulcus and conformation of the mesopleural sulcus to diagnose the Ponerinae for the Malagasy and

Japanese regions, respectively. These structures are variably developed on the global level, with several

genera presenting unsculptured scutoscutellar sulci, and as noted above the mesopleural sulcus is not

always present. In general, presence of the abdominal segment III cinctus is more stable than these sulci.

Understanding the generic boundaries of male Ponerinae will be most difficult in the Afrotropics, where

the most genera occur and where the least number of genera have males described. A key to the New

World genera, including Ponerinae, is in the works (B. Boudinot, in prep.).

Subfamily Proceratiinae Emery, 1895

Figs 6A, 14A-B

Diagnosis

All proceratiine genera share the following characters which are required for identification: oblique

mesopleural sulcus present; mesotibia with one or no apicoventral spurs, metatibia with one apicoventral

spur; propodeal lobe present; three to five closed cells present on forewing; jugal lobe absent; petiolar

tergum and sternum distinct; abdominal sternum IX unpronged and edentate. Two conditional sets of

characters are required in conjunction with those indicated above: 1) if mandibles triangular then antennal

toruli situated well-posterad anterior clypeal margin and crossvein lm-cu absent; and 2) if mandibles

reduced then antennal toruli situated at or produced anterad anterior clypeal margin; lm-cu may be

present or absent. Additionally, proceratiine males may or may not have vaulted fourth abdominal terga,

and the eighth abdominal tergum is never spiniform.

Comments

Yoshimura & Fisher’s (2009) key to the Malagasy Proceratiinae has global applicability. The three

genera of Proceratiinae, Discothyrea , Probolomyrmex , and Proceratium, seem to be well-defined taxa.

Subfamily Dorylinae Teach, 1815

Figs 5A-B, E, 6C, 14C-D

Diagnosis

Most Dorylinae are uniquely identified by the bidentate or pronged ninth abdominal sternum, lack of

pygostyles, and poorly developed clypeus. Males of the Leptanilloides genus group are highly derived

and are identifiable by the following combination of characters: antennal toruli abutting or very nearly

abutting anterior clypeal margin; oblique mesopleural sulcus absent; four closed cells present on

forewing; cinctus between abdominal pre- and postsclerites IV absent.

48

BOUDINOT B.E., Contributions to the Formicidae

Comments

The males of the Dorylinae have a long history of treatment due to the conspicuousness and remarkable

morphology of several constituent genera ( Dorylus , Aenictus , Anictogiton, the Eciton genus group).

Indeed, the first male-based ant taxon described was Dorylus helvolus (L., 1764). The concept of the

Dorylinae has shifted greatly over the past two hundred years; a very recent molecular phylogeny (Brady

et al. 2014) has redefined the Dorylinae in a broad sense, including the formerly accepted subfamilies (as

of Bolton 2003) Aenictinae, Aenictogitoninae, Cerapachyinae, Ecitoninae, and Leptanilloidinae, which

themselves include several family-level synonyms. Little further will be said of the Dorylinae here as a

generic revision of the subfamily is being prepared which will treat both males and females, and which

will significantly clarify the generic limits of this diverse subfamily (M.L. Boroweic, in prep.).

Subfamily Myrmeciinae Emery, 1877

Figs 5C-D, 6H, 14E-F

Diagnosis

Uniquely identified by the combination of petiolation of abdominal segment III (Myrmeciini), retention

of the jugal lobe and of two ventroapical spurs on each meso- and metatibia, and complete fusion of the

petiolar tergum and sternum anteriorly (Prionomyrmecini). The third abdominal segment of the male of

Nothomyrmecia (Prionomyrmecini) is incompletely petiolated, although it is still recognizable by the

other states indicated above.

Comment

Extant Myrmeciinae are restricted to Australia and New Caledonia, and are comprised of two monogeneric

tribes, Myrmeciini (Myrmecia) and Prionomyrmecini ( Nothomyrmecia ). Ward & Brady (2007) provided

keys to the extant and extinct genera of Myrmeciinae, including males, although the two extant genera

may also be separated by the key presented above.

Subfamily Pseudomyrmecinae M R. Smith, 1952

Figs 5B, E, 15A-B

Diagnosis

Pseudomyrmecine males are uniquely identified by the absence of the jugal lobes in combination with the

following character combination: cuticle soft, flexible, wealdy-sculptured; frontal carinae inconspicuous

or absent; meso- and metatibia each with two ventroapical spurs; more than three forewing cells closed;

abdominal segment III petiolated; abdominal sternum IX unpronged.

Comments

The Pseudomyrmecinae is comprised of three genera, the widespread New World Pseudomyrmex, the

Amazon endemic Myrcidris , and the Old World genus Tetraponera. Ward (1990) provided a male-based

key to the genera for the subfamily.

Subfamily Aneuretinae Emery, 1913

Figs 7A, 15C

Diagnosis

The male of Aneuretus simoni is uniquely identified by the exceedingly long and thin petiolar peduncle

and the unpetiolated third abdominal segment. The species is further identified by the following

combination of characters: oblique mesopleural sulcus present; seven closed cells present on forewing;

jugal lobe absent; abdominal segment IV without cinctus between pre- and postsclerites; abdominal

49

European Journal of Taxonomy 120: 1-62 (2015)

sternum IX unpronged and edentate; telomere extending anteroventrad basimere. Additional characters

for distinguishing A. simoni from the Dolichoderinae and Formicinae are indicated in couplets 19 and

20 above.

Comments

The sole extant member of the Aneuretinae, A. simoni , is restricted to Sri Lanka, and is the survivor

of a lineage which has a somewhat diverse fossil record (LaPolla et al. 2013). The subfamily is of

considerable interest as it is sister to the Dolichoderinae (Brady et al. 2006). Eight fossil genera are

ascribed to the Aneuretinae based on the work of several authors (e.g., Dlussky & Rasnitsyn 2009).

Some taxa, based on workers, are definitely members of the Aneuretinae, i.e., f Paraneuretus and

f Protaneuretus from Baltic amber (37^12 My; Wheeler 1915; LaPolla et al. 2013), while others are less

certain, i.e., f Pityomyrmex (also from Baltic amber, Wheeler 1915; placed in Aneuretinae by Dlussky

& Rasnisyn 2009) and f Aneuretellus (Sakhalin amber, 56-59 My; Dlussky 1988; LaPolla et al. 2013).

The impression-fossil taxa '\Britaneuretus (see Antropov et al. 2014) and f Mianeuretus (see Carpenter

1930) may not be members of the Aneuretinae. Because of the occurrence of definitive aneuretines in

Baltic amber, it will be critical to carefully study the reproductives occurring in these fossils to determine

whether any may be placed in the Aneuretinae.

Two fossil “aneuretine” taxa are worth discussing specifically. The affinities of ^Burmomyrma (—98 My,

Burmese amber; Dlussky 1996; LaPolla et al. 2013) and "fCananeuretus (78-19 My, Canadian amber;

Engel & Grimaldi 2005; LaPolla et al. 2013) with Aneuretus simoni are uncertain. The description and

illustration of f Burmomyrma in Dlussky (1996) provide no characters which support a relationship

of the fossil taxon with Aneuretus ; the diagnosis includes one extreme autapomorphy and several

characters which are pleisiomorphic for the family or are broadly shared among several subfamilies.

The character combination indicated by Dlussky (1996) to assign f Burmomyrma to the Aneuretinae is

weak, especially given that Aneuretus has complete (“ancestral”) wing venation while f Burmomyrma

lacks almost all vein abscissae. Placement of f Burmomyrma within the Leptanillinae, and indeed other

aculeate hymenopteran families, cannot be ruled out. No taxonomic action is taken here, however.

' \Cananeuretus , on the other hand, cannot be so easily considered distantly related to the Aneuretinae. The

Grassy Lake deposit of Canadian amber includes representatives of the Sphecomyrminae, Ectatomminae,

and critically, the Dolichoderinae (LaPolla et al. 2013). While the placement of the fossil dolichoderine

f Chronomyrmex (see McKellar et al. 2013) in the Leptomyrmecini (sensu Ward et al. 2010) is debatable,

co-occurrence of these subfamilies in this deposit suggests the placement of f Cananeuretus is plausible.

As the diagnosis of the Aneuretinae provided here and previously (Wilson et al. 1956; Bolton 2003)

is based largely on pleisiomorphic characters, other characters should be considered. For example,

future studies of Canadian amber should be sensitive to specific traits occurring in Aneuretus and extant

Dolichoderinae. Aneuretus shares, among other characters, a deep median notch on the anterior clypeal

margin and fine serrations intercalated among larger denticles on the masticatory mandibular margin,

both of which occur in the Tapinomini, the tribe sister to the remaining Dolichoderinae (Ward et al.

2010). Reconsideration of the fossil record of Aneuretinae will be valuable for improving our concepts

of both the Aneuretinae and Dolichoderinae.

Diagnosis

Subfamily Dolichoderinae Ford, 1878

Figs 7E, 15D-E

The Dolichoderinae are uniquely identified by the telomere, which is strongly reduced and does not

extend anteroventrad the basimere. Males of the subfamily are further identified by the following

combination of characters: oblique mesopleural sulcus present; seven or fewer closed cells present on

forewing; jugal lobe absent; petiolar peduncle short or absent; abdominal segment III unpetiolated;

50

BOUDINOT B.E., Contributions to the Formicidae

abdominal segment IV without cinctus between pre- and postsclerites; abdominal sternum IX unpronged

and edentate. Additional characters for distinguishing males of the Dolichoderinae from the Aneuretinae

and Formicinae are indicated in couplets 19 and 20 of the key above.

Comments

The Dolichoderinae is one ofthe major ant lineages, with over 700 described species distributed in 28 valid

genera. Males are unknown for four genera (. Ecphorella , Gracilidris , Loweriella , Nebothriomyrmex). One

of the historically intractable problems of myrmecology has been the separation of male Dolichoderinae

and Formicinae. Here, the telomeral character described by Yoshimura & Fisher (2011) is confirmed

as a diagnostic synapomorphy of the subfamily on a global scale, while the antennal torulus to clypeus

distance character was first described to the author’s knowledge in Czechowski et al. (2012). Several

other characters were found to distinguish male Dolichoderinae and Formicinae, as indicated in the key

and this diagnosis. Genera of Dolichoderinae have been keyed globally by Shattuck (1992), although

this key stands in need of updating. A key to the New World dolichoderine genera is in preparation (B.

Boudinot, in prep.).

Subfamily Formicinae Fatreille, 1809

Figs 51, 7B, F, 16A-B

Diagnosis

The Formicinae are uniquely identified by the following combination of characters: mandibles never

serrate; antennal toruli usually situated posterad posterior clypeal margin; antenna 8-13-merous; oblique

mesopleural sulcus present; at most six closed cells present on forewing; jugal lobe absent; petiolar

peduncle short to absent; petiole narrowly attached to abdominal segment III; abdominal segment III

unpetiolated; abdominal segment IV without cinctus between pre- and postsclerites; abdominal sternum

IX unpronged and edentate.

Comments

In terms of both number of described species (-3,000) and genera (51), the Formicinae is one of the

most diverse lineages of ants. Genera of the Formicinae are relatively easily delimitable based on males

(B. Boudinot, in prep.), but little work has been done to render males identifiable. Males are unknown

or at least undescribed for seven genera ( Agraulomyrmex , Alloformica , Bregmatomyrmex , Forelophilus,

Pseudonotoncus , Santschiella , Teratomyrmex), and the identity of Echinopla and Phasmomyrmex is

uncertain.

Ectaheteromorph clade

Fig. 16B-C

Diagnosis

Male ectaheteromorphs are uniquely identified by the following combination of characters: mandibles

triangular, multidentate; antennal toruli situated posterad anterior clypeal margin; tibial spur formula

1,1 or 2,2 (if 2,2 then prora anteriorly directed); crossvein lm-cu present; abdominal segment III

unpetiolated; cinctus present between the pre- and postsclerites of abdominal segment IV; abdominal

sternum IX unpronged and edentate. The jugal lobes may be present or absent.

Comments

No nomothetic (single unique) character separates the males of Ectatomminae Emery, 1895 and

Heteroponerinae Bolton, 2003, which are better distinguished genus-by-genus. In brief, Typhlomyrmex

may be distinguished by the scapes, which are longer than the compound eye; Acanthoponera males have

51

European Journal of Taxonomy 120: 1-62 (2015)

Fig. 13. Male representatives of three subfamilies. A, C, E. Frontal view. B, D, F. Lateral view. —

A-B. Tatuidris tatusia, Agroecomyrmecinae (Panama, CASENTO178870, E. Prado). C-D. Paraponera

clavata , Paraponerinae (Guyana, CASENT0902407, R. Perry). E-F. Pseudoponera stigma , Ponerinae

(Paraguay, CASENTO 178182, A. Nobile). Scale bars: A= 0.1 mm, B = 0.5 mm, C, F = 1.0 mm, D = 2.0

mm, E = 0.2 mm.

52

BOUDINOT B.E., Contributions to the Formicidae

Fig. 14. Male representatives of three subfamilies. A, C, E. Frontal view. B, D, F. Fateral view. —

A-B. Proceratium creek, Proceratiinae (U.S.A., CASENT010441, A. Nobile). C-D. Acanthostichus,

Dorylinae (French Guiana, CASENT0056970, A. Nobile). E-F. Myrmecia chasei, Myrmeciinae

(Australia, CASENT0903663, W. Ericson). Scale bars: A, C = 0.2 mm, B, E = 0.5 mm, D = 1.0 mm, E

= 2.0 mm.

53

European Journal of Taxonomy 120: 1-62 (2015)

Fig. 15. Male representatives of three subfamilies. A, D. Frontal view. B-C, E. Lateral view. — A-B.

Pseudomyrmex holmgreni , Pseudomyrmecinae (Paraguay, CASENTO173758, A. Nobile). C. Aneuretus

simoni, Aneuretinae, used with permission from Wilson et al. (1956). D-E. Technomyrmex difficilis ,

Dolichoderinae (Madagascar, CASENT0049968, A. Nobile). Scale bars: A, D = 0.2 mm, B = 1.0 mm,

E = 0.5 mm, no scale available for C.

54

BOUDINOT B.E., Contributions to the Formicidae

Fig. 16. Male representatives of three subfamilies. A-B. Formica wheeleri , Formicinae (U.S.A.,

CASENTO173024, A. Nobile). C-D. Rhytidoponera , Ectatomminae, ectaheteromorph clade (Australia,

CASENT0004610, A. Nobile). E-F. Pogonomyrmex rastratus (Argentina, CASENTO 172673, A.

Nobile). Scale bars: A, C = 0.5 mm, B, D, F = 1.0 mm, E = 0.2 mm.

55

European Journal of Taxonomy 120: 1-62 (2015)

long maxillary palps, which almost reach the postocciput; Ectatomma (Neotropical) and Rhytidoponera

(Australasian) are large and have jugal lobes. Heteroponera and Gnamptogenys are distinguishable

globally only using conditional statements. Characters have been found to do this, and will be described

in a forthcoming publication on the New World genera (B. Boudinot in prep.). The male of Aulacopone

is unknown.

Subfamily Myrmicinae Lepeletier de Saint-Fargeau, 1835

Figs 4H, 5A, G, 16D-E

Diagnosis

Male Myrmicinae are uniquely identified by the strongly petiolated third abdominal segment (postpetiole),

axial helcium, 1,1 maximum ventroapical tibial spur count, unvaulted abdominal tergum IV, and

presence of propodeal lobes. All myrmicines lack jugal lobes and have posteriorly-situated antennal

toruli, but are highly variable otherwise: mandibles fully-developed to nub-like; antenna 8-13-merous;

forewing with (0)1-8 eight closed cells; and petiole sessile to long-pedunculate. Some myrmicines, e.g.,

Adelomyrmex and Acanthognathus , have extremely reduced wing venation similar to Leptanillinae; all

myrmicine taxa examined during this study with reduced wing venation have conspicuous propodeal

lobes, differentiating them easily from Leptanillinae despite secondary petiolation of abdominal segment

III in some leptanillines.

Comments

Of all the ant subfamilies, the Myrmicinae will be the grand challenge to understand with respect to

males. At the time of writing, 139 valid genera and 6,500 valid species are described. Males of at least 30

genera are unknown, but as generic delimitation is still very active in the Myrmicinae some uncertainty

exists for this number. Based on a study of the New World genera (B. Boudinot, in prep.), distinctions

between genera may be weak and in many cases genera will have to be keyed multiple times due to

variability. The recent subfamily-wide phylogeny of Ward et al. (2015) will contribute significantly to

improving the classification of the Myrmicinae.

Acknowledgments

First I want to extend my gratitude to Dr Philip S. Ward (UCD) for his advice, for his support for my

Brazil museum trip (which precipitated the present work), for his critical review of the manuscript, and

valuable discussions of this research through its various phases. I want to also extend my gratitude to

Dr. John (Jack) T. Longino (University of Utah) for diverting my attention to entomology, critically

encouraging my interest in the Formicidae, and providing a valuable review of this manuscript. This

work would not be possible without Barry Bolton’s extensive and influential literature. I want to thank

my Brazilian colleagues, who supported my trip to Brazil in several functions. At the INPA, I thank Dr.

Marcio Oliveira, Thiago Mahlmann, and Itanna Fernandes; at the MZSP, I thank Dr. C. Roberto (Beto)

F. Brandao, Rodolfo da S. Probst, Thiago S. R. da Silva, Monica Ullyssea, and Livia Prado; and at the

DZUP, I thank Dr. Rodrigo M. Feitosa, Gabriela Camacho, and the rest of the Feitosa lab. At UC Davis,

I thank Dr. Lynn Kimsey for a valuable loan and discussion of non-formicid Aculeata; Marek Borowiec

for use of Leptanillinae on loan from Lund, for lending books on the European myrmecofauna, and for

providing discussion of the “basal ant problem”; and Drs. Steve Heydon and Lynn Kimsey for helping

discover non-formicid literature. Thanks to Dr. Brian Fisher and Michele Esposito for Ant Web; Stan

Blum, Barry Bolton, and the AntCat community for online taxonomic catalog resources. An anonymous

reviewer provided some useful comments. This work was supported by National Science Foundation

grant DEB-1354739 (Project ADMAC), a UC Davis Jastro grant, and an Ernst Mayr grant from the

MCZ for the project “Male ants of Central America”, which has expanded to “Male ants of the New

World”.

56

BOUDINOT B.E., Contributions to the Formicidae

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Manuscript received: 4 November 2014

Manuscript accepted: 27 January 2015

Published on: 20 April 2015

Topic editor: Koen Martens

Desk editor: Kristiaan Hoedemakers

Printed versions of all papers are also deposited in the libraries of the institutes that are members of

the EJT consortium: Museum National d’Histoire Naturelle, Paris, France; Botanic Garden Meise,

Belgium; Royal Museum for Central Africa, Tervuren, Belgium; Natural History Museum, London,

United Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium; Natural History

Museum of Denmark, Copenhagen, De nm ark.

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