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† Desyopone hereon sp. nov. Boudinot and Perrichot (Figures 1–5) ZooBank LSID: urn: lsid:zoobank.org:act: 9E345965-6AA1-42D8-A468-187BABBB38D2. Etymology. The specific epithet gratefully recognizes the Helmholtz-Zentrum Hereon, the research center which funds and operates the Imaging Beamline (P05) at DESY, thus making the present work possible. Holotype. Male (m), MAIG 6016 (Figures 1B, 2A,B, 3 and 4), deposited in the Museum of Amber Inclusions at University of Gda´nsk, Poland. Paratypes. 12 males (m). Synincluded with holotype in amber piece MAIG 6016 (Figures 1A,C,D, 2C,D and 5). Type locality. Exact locality unknown in the Bashilo river gorge near Weldiya, Semien Wollo Zone, Amhara Region, Ethiopia. Type horizon. A fine siltstone/mudstone of Early Miocene age (16–23 Ma). Material. Holotype and paratypes 1–4 are nearly complete, preserved without apparent distortion but with integument entirely covered by a white, opaque (bacterial?) coat for paratypes 1, 2, 4 (Figures 1A,C and 2A). Paratypes 5–12 mostly complete but variously preserved, more or less distorted by apparent dehydration (e.g., Figure 1D). Diagnosis. † Desyopone hereon is uniquely identifiable among all Ponerinae by the distinctly elongate petiolar peduncle and the enlarged, lobate mandibles. Description. Measurements: Holotype (paratypes) (taken from paratypes 1,2,4)—BL 3.11 (3.00–3.50), HW 0.57 (0.50–0.60), HWE 0.70 (0.66–0.70), SL 0.12 (0.10–0.12), EL 0.23, OLL 0.08 (0.08), OIL 0.16 (0.16), WL 1.25 (1.25), ML 0.52 (0.50–0.56), MW 0.49, FWL (2.75), PH 0.26 (0.30), PL ~0.40 (0.40), GL 1.20 (1.00–1.50), GW 0.62 (0.65–0.80), HWI 81 (76–86), SI 21 (20), OCI 50 (50), PI 154 (133). Head (Figures 2–4): Head capsule ovoid-elliptical in lateral view; in full-face view, posterior head margin broadly and evenly convex to compound eyes; oral region of head, i.e., the malar areas, clypeus, and mouthparts, narrower than distance between compound eyes; postgenal bridge short, about 2/5 the length of the head in full-face view as measured from the postocciput to the hypostoma; malar areas distinctly developed; clypeus medially bulging, laterally depressed. Compound eyes situated almost entirely in anterior half of head; eyes subspherical and relatively small, with their length being about 1/3 head length. Ocelli distant from compound eyes, with the lateral ocelli separated from the compound eyes by slightly more than one compound eye length. Antennal toruli located at about head midlength in full-face view, distinctly posterad the posterior clypeal margin. Antennal scapes short, just barely longer than wide, with their length distinctly Mesosoma (Figures 2 and 3): Pronotum short and simple but with distinct muscular convexity as seen in lateral and dorsal views; posterad the anteromedian pronotal lobe (“nuchal lobe”), pronotum in the form of a simple arch, without distinct dorsal and ventral surfaces. Propleurae widely emarginate posteromedially, together forming a broad arch for the prosternum. Prosternum with basisternum apparently arcuate anteriorly; prosternal process developed. Mesoscutum somewhat narrow, with the anteroposterior length slightly greater than the lateromedial width. Notauli developed, and Y-shaped; meeting in the posterior half of the mesoscutum. Parapsides developed, although indistinct. Scutoscutellar sulcus broad, with at least five cross-costae. Mesoscutellum simple, convex, longer than broad in full-face view. Oblique mesopleural sulcus developed. Spiracular lobe (ventrad wing insertions) absent. Mesopleural area divided into dorsal and ventral regions; both regions dorsoventrally taller than anteroposteriorly long. Mesosternal and metasternal regions without processes. Propodeum convex, without armature or distinct sculpturation. Propodeal lobes developed, weak. Legs: Mesotibiae with no spur, metatibiae with a single, pectinate spur. Pretarsal claws narrow and simple, without additional teeth. Arolia well-developed but not very large. Fore wings (Figures 2C,D and 5): Costal vein (C) present, complete. Rsf1 diverging from Sc+ R +Rs well proximad pterostigma, with Sc+ R abscissa about 1/3 pterostigma length. Pterostigma well-developed, long, and narrow, with its length> 5 × its width. Rf distad pterostigma tubular. M+Cu tubular. Mf1 diverging from M+Cu at or slightly distad crossvein cu-a; this abscissa very weakly curved and meeting Rf1 at a distinct oblique angle. Rs+M tubular. Rsf2–3 diverging from Rs+M proximad 1m-cu. Crossvein 1r-rs absent. Crossvein 2r-rs anterior juncture at 2/3 length of pterostigma; this crossvein meeting Rsf proximad crossvein 2rs-m, which is tubular (paratype 4 has 2rs-m duplicated on the right wing, see Figure 2D). Rsf4+ tubular, meeting Rf distally). Mf2 (=abscissa between Rs+M and 1m-cu) short, shorter than 2r-rs and 2rs-m. Mf3 similar in length to but distinctly shorter than Rsf2–3. Mf4+ tubular proximally, becoming nebulous distally, with this occurring at a distance that is about 3 × the length of 2rs-m. Cuf3 (=abscissa of Cu after 1m-cu) joined to 1A posteriorly; 1A is tubular for its entire length. Submarginal cells 1 and 2 similar in size and shape, but with 2 distinctly smaller than 1. Marginal cell 1 long and narrow, with a length that is about 1.5 × pterostigma length. Discal cell 1 subrhomboidal, its length slightly less than 2 × its anteroposterior width. Subdiscal cell closed. Hind wings (Figures 2C and 5): Wing with eight distal hamuli. Jugal lobe absent. C not distinctly developed. R splitting from Sc+ R +Rs distad crossvein 1rs-m; Rf incomplete, not reaching anterior wing margin. Rsf tubular for a distance that is about 2 × the length of Sc+ R +Rs distad 1rs-m. M+Cu splitting well distad crossvein cu-a. Mf1 meeting rs-m at a broad, oblique angle; Mf developed as a stud distad this juncture. Cuf developed distad the split of M+Cu, but exact condition uncertain. Crossvein cu-a long, slightly longer than the length of Mf1; this crossvein situated proximad split of M+Cu by about twice its length. Anal vein (A) tubular past its juncture with cu-a. Anal cell relatively long; M+Cu proximad cu-a distinctly longer than M+Cu distad cu-a. Metasoma (Figures 2B and 3A,D,E): Petiole nodiform and distinctly pedunculate, albeit without a marked inflection between its anterior portion and the anterior surface of the petiolar node; peduncle about 2/5 petiole length; petiolar node height about 3/5 entire petiole length; node broad and convex; posterior collar well-developed; tergosternal fusion absent, laterotergites present; sternum low and very weakly sinuate in lateral view, without an anteroventral (=subpetiolar) process, nor with a posterior process; posterior sternal margin distinctly notched. Helcium infraaxial (=below abdominal segment III midheight); helcial tergite broad and overlapping sternite laterally. Prora not distinctly developed. Gastral segments homonomous in appearance and gradually decreasing in length from abdominal segment III to VIII. Abdominal segment IV without cinctus (=constriction). Abdominal spiracles IV–VIII obscured by preceding tergites. Abdominal tergum VIII apparently simple. Abdominal sternum IX lobate and somewhat narrow, with a length that is about twice its width. Cerci (=pygostyles) developed. Genitalia (Figures 2B and 3A): Cupula present (only visible in µ- CT cross-sections). Gonopods longer than tall. Gonostyli broad proximally, indistinctly set off from gonocoxa, and lobate in appearance, being narrowly rounded apically. Lateropenites (=digiti) thickened apically, downcurved. Penites (=penisvalvae) apparently longer than tall, curving strongly to their narrowly lobate apices. Setation: Body with two primary hair classes: (1) short pubescence, which covers the head and all segments of the antennae, mesosoma, legs, and metasoma; (2) long hairs, which are sparse on all body regions, but are denser on the metasoma., Published as part of Boudinot, Brendon E., Richter, Adrian K., Hammel, Jörg U., Szwedo, Jacek, Bojarski, Błażej & Perrichot, Vincent, 2022, Genomic-Phenomic Reciprocal Illumination: Desyopone hereon gen. et sp. nov., an Exceptional Aneuretine-like Fossil Ant from Ethiopian Amber (Hymenoptera: Formicidae: Ponerinae), pp. 1-19 in Insects 73 (796) on pages 9-13, DOI: 10.3390/insects13090796, http://zenodo.org/record/7046409

Tribe Ponerini Lepeletier de Saint-Fargeau, 1835. Type genus. Ponera Latreille, 1804. Male diagnosis. In addition to the ponerine plesiomorphies, male Ponerini are distinguished by the following: (1) mandibles vestigial, with an enlarged mandalus, and being variably lobate, spatulate, spiniform, or nub-like (no exceptions known) [synapomorphy, homoplastic among Formicidae]; (2) face between antennal toruli not distinctly raised, thus toruli directed more-or-less dorsally; if the intertorular region is raised, then this region is grooved in appearance due to impression of the supraclypeal area (=frontal triangle) (Plectroctena and Psalidomyrmex with medial torular arches raised, but not face; Hagensia, Megaponera, Ophthalmopone, Simopelta, some Euponera, and some Odontomachus with intertorular region raised) [plesiomorphy]; (3) antennal toruli usually distant from posterior clypeal margin (some Brachyponera, many Leptogenys, Megaponera, and Ophthalmopone have toruli that are close to the clypeal margin) [plesiomorphy]; (4) meso- and metatibiae with two, one, or no spurs; (5) jugal lobe present or absent; (6) helcium distinctly infraaxial, i.e., situated well below the midheight of abdominal segment III (Simopelta is an exception due to softening and reduction in size of the metasoma) [synapomorphy, homoplastic among Formicidae]; (7) abdominal segment IV with or without cinctus; (8) cuticle usually not pruinose, being shiny and variably sculptured (Belonopelta, Hagensia, Megaponera, and Ophthalmopone are exceptions) [plesiomorphy]. Remarks. Definitive infraaxiality in males, i.e., with abdominal tergum III rising high above the petiole, is a strong diagnostic condition for Ponerinae, as this is an infrequent apomorphic condition at the subfamily level. It also occurs in Dolichoderinae and Formicinae, and to some extent in Myrmeciinae and various Myrmicinae. Discothyrea (Proceratiinae) may approach infraaxiality, but the third abdominal tergum is low., Published as part of Boudinot, Brendon E., Richter, Adrian K., Hammel, Jörg U., Szwedo, Jacek, Bojarski, Błażej & Perrichot, Vincent, 2022, Genomic-Phenomic Reciprocal Illumination: Desyopone hereon gen. et sp. nov., an Exceptional Aneuretine-like Fossil Ant from Ethiopian Amber (Hymenoptera: Formicidae: Ponerinae), pp. 1-19 in Insects 73 (796) on page 8, DOI: 10.3390/insects13090796, http://zenodo.org/record/7046409

Genus † Desyopone gen. nov. Boudinot and Perrichot Type species. † Desyopone hereon sp. nov., by present designation monotypy. ZooBank LSID: urn:lsid:zoobank.org:act: 7228E671-DF5E-44D1-ADE6-62FB6DB34220. Etymology. The genus name is a portmanteau of the traditional ponerine generic suffix, “-pone”, and the acronym for the Deutsches Elektronen-Synchrotron (DESY), whose storage ring and radiation beamline facilities were used to generate the phenomic data that were crucial for the correct identification of the new taxon. Diagnosis. † Desyopone has plesiomorphies 1–4 of Ponerinae and is identifiable as Ponerini at minimum due to the vestigial mandibles and infraaxial helcium. † Desyopone and Cryptopone are uniquely identified among all Ponerinae by: (1) subpetiolar process completely absent, with the poststernite low and nearly flat in profile. The new genus differs from the males of all known Cryptopone by the following: (2) head broader than long, excluding the compound eyes (vs. head narrower than long); (3) mandibles elongate (vs. short); (4) mandibles lobate (vs. spiniform); (5) mesospiracular sclerite evidently absent (vs. this sclerite present); (6) meso- and metatibiae with no spur and one spur, respectively (vs. two spurs each); and (7) petiolar peduncle long, about as long as node is tall (vs. peduncle short, considerable shorter than height of node). Remarks. The identity of Cryptopone is significantly clarified by the phenomic data from the new species and the phylogenomic revision of Branstetter and Longino [50]. Prior to this work, the diagnostic importance of the absent subpetiolar process was obscured by the inclusion of Wadeura guianensis in Cryptopone. Now it is clear that the absence of the subpetiolar process is a unique condition among extant Ponerinae that is shared between † Desyopone gen. nov. and Cryptopone, and thus constitutes a reasonable autapomorphy within the subfamily for the two genera. No known Cryptopone, however, matches the diagnostic character combination of † Desyopone gen. nov., with conditions 3–7 being apomorphic. Critically, the elongate peduncle of † D. hereon gen. et sp. nov. is nearly unique among Ponerinae; this condition is similarly derived in Harpegnathos and is approached by Dinoponera, some Odontomachus (e.g., O. chelifer, O. coquereli), and Platythyrea (although node at middle of segment rather than posterior)., Published as part of Boudinot, Brendon E., Richter, Adrian K., Hammel, Jörg U., Szwedo, Jacek, Bojarski, Błażej & Perrichot, Vincent, 2022, Genomic-Phenomic Reciprocal Illumination: Desyopone hereon gen. et sp. nov., an Exceptional Aneuretine-like Fossil Ant from Ethiopian Amber (Hymenoptera: Formicidae: Ponerinae), pp. 1-19 in Insects 73 (796) on page 8, DOI: 10.3390/insects13090796, http://zenodo.org/record/7046409, {"references":["50. Branstetter, M. G.; Longino, J. T. UCE phylogenomics of New World Cryptopone (Hymenoptera: Formicidae) elucidates genus boundaries, species boundaries, and the vicariant history of a temperate - tropical disjunction. Insect Syst. Divers. 2022, 6, 1 - 23. [CrossRef]"]}

Subfamily Aneuretinae Emery, 1913. Type genus. Aneuretus Emery, 1893. Diagnosis (all adult castes). All adults of Aneuretinae possess the following diagnostic plesiomorphies (1–6): (1) mandibles shovel-shaped (=“triangular”) [plesiomorphy of Poneroformicia, or at least Doryloformicia]; (2) meso- and metatibiae with one spur each [synapomorphy of clade Dolichoderomorpha]; (3) petiole with complete tergosternal fusion [synapomorphy of clade Dolichoderomorpha]; (4) petiole with an elongate anterior peduncle [possible synapomorphy of clade Myrmechoderines]; (5) helcium infraaxial [possible synapomorphy of clade Myrmechoderines]; and (6) abdominal segment IV unconstricted [synapomorphy of clade Dolichoderomorpha]. Males and queens of Aneuretinae share the following diagnostic traits (7–12): (7) near-complete fore wing venation, with only the subdiscal cell open; (8) crossvein 2rs-m furcal to post-furcal, i.e., 2r-sm meeting Rs at or distad 2r-rs; (9) well-prefurcal fore wing crossvein cu-a, i.e., cu-a meeting M+Cu proximad the split of M+Cu by more than one of its own lengths; (10) absence of free M in the hind wing after the juncture of rs-m and Mf1, i.e., the abscissa between Sc+ R +Rs and M+Cu linear, without a kink; (11) hind wing anal cell short, its length less than half that of the basal cell; and (12) absence of the hind wing jugal lobe. Males of Aneuretinae have the following diagnostic plesiomorphy (13): (13) genital gonocoxa and gonostylus not strongly differentiated in size, with the dorsal gonocoxal margin continuing more-or-less evenly to that of the gonostylus. Workers and queens of Aneuretinae share the following diagnostic plesiomorphies (14, 15): (14) mandible with biseriate dentition, i.e., with small teeth interspersed between large teeth [synapomorphy of Dolichoderomorpha]; (15) basal and masticatory margins of mandible not marked, i.e., these margins curving into one another, without a distinct angle [synapomorphy of Dolichoderomorpha]. Remarks. The operational paleontological definition of Aneuretinae has relied on character states 2, 4, 6, 7 (regardless of subdiscal cell state), and 12. With the explicit recognition of character state 3—which was previously indicated for the “formicomorph subfamilies” by Bolton [11] (p. 16)—it is possible to reject the placement of † Desyopone hereon gen. et sp. nov. from the Aneuretinae. The condition of helcial axiality is here reinterpreted from Bolton [11] (p. 18), who described the helcium of Aneuretinae as “high on [the] anterior face of abdominal segment III”, which is interpretable as supraaxial sensu Keller [46]. Although the helcial tergite of worker Aneuretus is dorsoventrally short, it can be seen that the helcium is at the ventralmost position of the sternum, which does not broaden. An axial helcium is confirmed for the Baltic amber taxa † Paraneuretus and † Protaneuretus as well. Wing venation was observable for Aneuretus and † Paraneuretus. Finally, we recognize the mandibular character states 14 and 15 as critical for the identification of Aneuretinae. The states of the mandibles have not been previously remarked upon, but along with the conformation of the clypeus (not defined here), they form the gestalt of the Aneuretinae and Dolichoderinae, which was likely used by Wheeler [7] to place † Paraneuretus and † Protaneuretus, although his justifications were not made explicit. Further refinement of the aneuretine diagnosis via comparative phenomics and traditional comparative morphology is highly desirable., Published as part of Boudinot, Brendon E., Richter, Adrian K., Hammel, Jörg U., Szwedo, Jacek, Bojarski, Błażej & Perrichot, Vincent, 2022, Genomic-Phenomic Reciprocal Illumination: Desyopone hereon gen. et sp. nov., an Exceptional Aneuretine-like Fossil Ant from Ethiopian Amber (Hymenoptera: Formicidae: Ponerinae), pp. 1-19 in Insects 73 (796) on pages 6-7, DOI: 10.3390/insects13090796, http://zenodo.org/record/7046409, {"references":["11. Bolton, B. Synopsis and classification of Formicidae. Mem. Amer. Entomol. Inst. 2003, 71, 1 - 370.","46. Keller, R. A. A phylogenetic analysis of ant morphology (Hymenoptera: Formicidae) with special reference to the poneromorph subfamilies. Bull. Am. Mus. Nat. Hist. 2011, 355, 1 - 90. [CrossRef]","7. Wheeler, W. M. The ants of the Baltic Amber. Schrift. Physikal. Okonom. Gesell. Konigsberg 1915, 55, 1 - 142."]}

Tribe Platythyreini Emery, 1901. Type genus. Platythyrea Roger, 1863. Male diagnosis. In addition to the ponerine plesiomorphies, male Platythyreini are distinguished by the following: (1) mandibles worker-like, shovel-shaped (=“triangular”) [plesiomorphy]; (2) face between antennal toruli distinctly raised, such that the antennal toruli are directed relatively laterad, and are situated dorsad a depression which may receive the antenna [synapomorphy, homoplastic in Ponerini]; (3) antennal toruli usually close to or indenting the posterior margin of clypeus, toruli never distant from the posterior margin by more than half of one of their diameters [apomorphy]; (4) meso- and metatibiae with two spurs each [plesiomorphy]; (5) jugal lobe present [plesiomorphy]; (6) helcium not distinctly infraaxial, being axial to more-or-less axial, i.e., at about or just slightly below the midheight of abdominal segment III [plesiomorphy]; (7) abdominal segment IV with cinctus [plesiomorphy]; (8) cuticle pruinose [synapomorphy, homoplastic in Ponerini and some Proceratiinae]. Remarks. Because male Platythyreini have never been explicitly diagnosed, we found it necessary to provide a diagnosis in order to confirm the identification of the fossils in question. We observe that the mandibular form, tibial spur count, and cuticular sculpture of male Platythyreini are sufficient for identification at the global scale., Published as part of Boudinot, Brendon E., Richter, Adrian K., Hammel, Jörg U., Szwedo, Jacek, Bojarski, Błażej & Perrichot, Vincent, 2022, Genomic-Phenomic Reciprocal Illumination: Desyopone hereon gen. et sp. nov., an Exceptional Aneuretine-like Fossil Ant from Ethiopian Amber (Hymenoptera: Formicidae: Ponerinae), pp. 1-19 in Insects 73 (796) on pages 7-8, DOI: 10.3390/insects13090796, http://zenodo.org/record/7046409

TECHNOMYRMEX SVOJTKAI PERRICHOT & ENGEL SP. NOV. (FIGS 1–3; SUPPORTING INFORMATION, FIGS S1, S 2) Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org:act: 565B4980-A80E-450B-A09B-603264A91F50. Type material: Holotype NHMW-N6976, a complete worker; in Early Miocene (16–23 Mya) amber from the North Shewa Zone, Amhara region, Ethiopia; housed in the Department of Mineralogy and Petrography, Naturhistorisches Museum Wien (NHMW, Austria). Paratypes MAIG 6020, two workers missing portions of legs; in Early Miocene (16–23 Mya) amber from the South Wollo Zone, Amhara region, Ethiopia; housed in the Museum of Amber Inclusions of the University of Gdansk (MAIG, Poland). Other material: MAIG 6020, one worker missing the head, portions of legs and gaster, but similar to the associated paratypes in its size and preserved structures, particularly the subcylindrical petiole, the shape and position of metathoracic and propodeal spiracles, thus considered here as conspecific; housed in the Museum of Amber Inclusions of the University of Gdansk (MAIG, Poland). Etymology: The specific epithet honours Mr Matthias Svojtka (University of Vienna) who discovered the fossil representing the holotype and generously made it available for study. Diagnosis: The species is uniquely identified within Technomyrmex by the following combination of worker characters: (1) anterior clypeal margin with a deep V-shaped median incision; (2) inner margins of incision continuously arched, with no marked angle with broadly semicircular anterolateral clypeal margins; (3) dorsum of head with a single pair of stiff setae situated above posterior level of eyes; (4) compound eyes situated around midlength of head (EPI 59), with about 40 ommatidia; (5) palp formula 6,4; (6) mandible with six conspicuous teeth apically on masticatory margin; (7) counting from the apex, first tooth longest, third and fifth teeth subequal in length, smaller than second and fourth, third to sixth teeth each separated by one denticle, sixth tooth followed by a continuous series of smaller teeth gradually decreasing in size along masticatory margin and basal angle, becoming denticles along entire basal margin; (8) the basal angle indistinct, continuously arched between margins; (9) mesosoma devoid of erect setae; (10) propodeal dorsum as long as declivitous face, their junction broadly rounded. Description: Holotype. Body integument minutely asperous, mostly glabrous except sparse stiff setae on anterior head margin and gaster. Head only slightly longer than wide. Compound eye oval, moderately large and feebly convex, its outer margin barely surpassing the outline of sides in full-face view, with 38–40 ommatidia. Ocelli absent. Frontal carinae well distant, closer to inner margin of eyes than to each other, not surpassing posterior margin of eyes and without frontal lobe, such that antennal sockets entirely exposed although directed laterad. Antenna 12-merous, scape surpassing vertexal margin by more than one-third its length; first and terminal funicular antennomeres subequal in length, nearly twice as long as broad; all other antennomeres cylindrical, only slightly longer than broad. Vertex with an erect seta (likely paired, but upper left side of head altered) longitudinally aligned with frontal carina, in profile situated at midlength between posterior margins of eye and head. Anterior clypeal margin bordered by seven stiff, short setae inclined ventrally; posterior clypeal margin semicircular, even with posterior level of toruli. Mesosoma: in profile, mesonotal dorsal outline feebly convex in its anterior third, flat and sloped in its posterior two thirds. Mesonotum and propodeum forming a distinct angle at metanotal groove. Metathoracic spiracles raised into distinct tubercules, conspicuously breaking outline of mesosoma in lateral view, distinctly anterior to metanotal groove. Dorsal surface of propodeum flat, declivitous surface convex. In profile, propodeal spiracle at junction of lateral and declivitous surfaces, slightly above midheight of sclerite. Dorsal surface of mesosoma apparently without erect setae. Legs: mesotibia and metatibia each with one spur, that of metatibia pectinate. Metasoma: petiole more than twice as long as wide, nearly as wide as high, broadest around its midlength; ventral and dorsal surfaces feebly convex; ventral surface without lobe, with two anterior longitudinal rows of three short setae. Gaster: abdominal tergite III largest, ventral surface of its anterior projection with a groove that accommodates entire petiole. Tergites III–VI with posterior margin bordered by a row of short appressed setae barely surpassing margin. Tergite VII in dorsal view small, trapezoidal. Paratypes and other specimens: As for the holotype, although incomplete or partially concealed due to preservation, but with posterior margin of head intact, not emarginate (MAIG 6020 paratypes); palpomeres clearly exposed (MAIG 6020 paratype 1), revealing a palp formula 6,4, with maxillary palp relatively long, exceeding hypostomal margin but not reaching occiput, and labial palp short, about one-third length of maxillary palp; mesonotum constricted and elongate (MAIG 6020 additional worker). Measurements and indices: Abbreviations are explained in Material and methods. Holotype: BL ~ 2.00, HL 0.50, HW 0.49, EL 0.15, FcW 0.18, SL 0.59, WL 0.77, PL 0.30, PW 0.13, PH 0.12, GL 0.80, GW 0.55, CI 98, OI 30, EPI 59, SI 129. Paratypes and additional worker: HL 0.52–0.54, HW 0.46, EL 0.11–0.15, SL 0.52, WL 0.77, PL 0.25–0.35, PW 0.20, SI 110. Note: Amber piece MAIG 6020 also contains a fourth, partially preserved worker, which displays a similar petiole and can probably be attributed to Technomyrmex. However, only the mesosoma and petiole are preserved, and it differs by a larger size (WL ~ 2.40 vs. 0.77; PL 0.77 vs. 0.25–0.35) and the presence of a dense, short pubescence covering all sclerites; it probably belongs to another species. Among the dolichoderines, the petiole reduced to a low segment without a node or scale and overhung by the first gastral segment, which is anteriorly grooved, is shared by Technomyrmex and Tapinoma Förster, 1850. The pygidium exposed in dorsal view, thus with five gastral tergites visible, as is seen on the new fossil, warrants placement in Technomyrmex (whereas the pygidium is reflexed ventrally, exposing only four gastral tergites in dorsal view in Tapinoma; see, e.g. Bolton, 2007; Fisher & Bolton, 2016). The new species is readily recognizable by its anterior clypeal margin with a deep V-shaped incision and broadly rounded lateral margins, the almost complete lack of erect setae on the dorsum of head, mesosoma and first gastral tergite (with the exception of a single pair of setae posterior to the frontal carinae), and the propodeal dorsum that is as long as, or even slightly longer than, the declivitous surface. In all other species of Technomyrmex with a deeply incised clypeal margin, the median notch is either U-shaped or semicircular, but never V-shaped. These and other features, such as the elongate mesosoma and metathoracic spiracles raised on tubercles, suggest that Technomyrmex svojtkai may belong to the bicolor group of species as defined by Bolton (2007)., Published as part of Perrichot, Vincent, Boudinot, Brendon E., Engel, Michael S., Fls, Xu, Chunpeng, Bojarski, Błażej & Szwedo, Jacek, 2022, Ants (Hymenoptera: Formicidae) from Miocene Ethiopian amber: filling gaps in the geological record of African terrestrial biota, pp. 775-791 in Zoological Journal of the Linnean Society 196 on pages 780-782, DOI: 10.1093/zoolinnean/zlac053, http://zenodo.org/record/7195057, {"references":["Bolton B. 2007. Taxonomy of the dolichoderine ant genus Technomyrmex Mayr (Hymenoptera: Formicidae) based on the worker cast. Contributions of the American Entomological Institute 35: 1 - 150.","Fisher BL, Bolton B. 2016. Ants of Africa and Madagascar: a guide to the genera. Berkeley: University of California Press."]}

RAVAVY MIAFINA FISHER, 2009 Measurements and indices: (N = 4, taken from CASENT 0078664, CASENT 0080308, CASENT0115570, CASENT0474633.) BL c. 3.33–4.46, HL 2.37–2.45, HW 1.84–1.91, HWE 2.13–2.25, EL 0.34–0.43, MSL 0.08–0.09, FcW 0.49–0.55, SL 0.074– 0.98, OLL 0.24–0.28, OIL 0.44–0.56, WL 1.30–1.67, ML 0.68–0.78, MW 0.81–0.92, FWL 3.85–4.24, PL 0.15–0.19, PW –, PH 0.28–0.31, GL 1.64–2.29, GW 0.58–0.70, CI 88–92, HWI 112–114 (122), MSI 28–32, SI 39–53, OI 14–18, EPI –, OCI 44–62, MI 104–129, PI 51–67., Published as part of Perrichot, Vincent, Boudinot, Brendon E., Engel, Michael S., Fls, Xu, Chunpeng, Bojarski, Błażej & Szwedo, Jacek, 2022, Ants (Hymenoptera: Formicidae) from Miocene Ethiopian amber: filling gaps in the geological record of African terrestrial biota, pp. 775-791 in Zoological Journal of the Linnean Society 196 on page 785, DOI: 10.1093/zoolinnean/zlac053, http://zenodo.org/record/7195057, {"references":["Fisher BL. 2009. Two new dolichoderine ant genera from Madagascar: Aptinoma gen. n. and Ravavy gen. n. (Hymenoptera: Formicidae). Zootaxa 2118: 37 - 52."]}

GENUS RAVAVY FISHER, 2009 Type species: Ravavy miafina Fisher, 2009: 47, figs 6, 7b., Published as part of Perrichot, Vincent, Boudinot, Brendon E., Engel, Michael S., Fls, Xu, Chunpeng, Bojarski, Błażej & Szwedo, Jacek, 2022, Ants (Hymenoptera: Formicidae) from Miocene Ethiopian amber: filling gaps in the geological record of African terrestrial biota, pp. 775-791 in Zoological Journal of the Linnean Society 196 on page 782, DOI: 10.1093/zoolinnean/zlac053, http://zenodo.org/record/7195057, {"references":["Fisher BL. 2009. Two new dolichoderine ant genera from Madagascar: Aptinoma gen. n. and Ravavy gen. n. (Hymenoptera: Formicidae). Zootaxa 2118: 37 - 52."]}

GENUS TECHNOMYRMEX MAYR, 1872 Type species: Technomyrmex strenuus Mayr, 1872: 147., Published as part of Perrichot, Vincent, Boudinot, Brendon E., Engel, Michael S., Fls, Xu, Chunpeng, Bojarski, Błażej & Szwedo, Jacek, 2022, Ants (Hymenoptera: Formicidae) from Miocene Ethiopian amber: filling gaps in the geological record of African terrestrial biota, pp. 775-791 in Zoological Journal of the Linnean Society 196 on page 780, DOI: 10.1093/zoolinnean/zlac053, http://zenodo.org/record/7195057, {"references":["Mayr G. 1872. Formicidae Borneensis collectae a J. Doria et O. Beccari in territorio Sarawak annis 1865 - 1867. Annali del Museo Civico di Storia Naturale 2: 133 - 155."]}

RAVAVY GOLDMANI BOUDINOT & PERRICHOT SP. NOV. (FIGS 4–5) Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org:act: 693FB2F0-817E-4BD6-B2E4-5585AFC99EA2. Type material: Holotype NIGP180512, male, housed in the Nanjing Institute of Geology and Palaeontology (NIGPAS, China); paratype IGR.ET2015/001a, male, housed in the Geological Museum of the University of Rennes (IGR, France). In Early Miocene (16–23 Mya) amber from the North Shewa Zone, Amhara region, Ethiopia. Etymology: The specific epithet honours Mr Yale Goldman (Bloomfield, Connecticut) who kindly facilitated access to the type specimens of the new species. Diagnosis: Because of the limited knowledge of male ants at the global scale, this diagnosis is broken into four parts in order to establish the identity of the treated specimens: identification at the subfamilial, tribal, generic and species levels. (I) Identifiable as Dolichoderinae by the following combination: (1) clypeal condyle large, rhomboidal (see note 1 below); (2) forewing cross-vein cu-a prefurcal, i.e. joining M+Cu proximad branching point of M and Cu; (3) helcium infraaxial; (4) abdominal segment III unpetiolated, i.e. not reduced in size relative to segment IV; (5) prora absent; (6) abdominal segment IV without cinctus; (7) gonostylus dorsoventrally short relative to gonocoxite, proximally constricted and not extending proximad gonocoxite (note 2); (8) basivolsellar process present (note 3). (II) Identifiable as Bothriomyrmecini by the following combination (note 4): (9) clypeus not extending posteriorly between toruli; (10) medial hypostomal lamina absent; (11) forewing cross-vein 2rs-m absent. (III) Identifiable as Ravavy by the following features, all of which are unique within Bothriomyrmecini (note 5): (12) mandible unidentate, spatulate with an apical spiniform tooth (vs. mandible multidentate, strapshaped to spiniform) (note 6); (13) palp formula 6,3 (vs. 6,4 or ≤ 4,3) (note 7); (14) pterostigma situated immediately proximad forewing midlength (vs. situated distad); (15) marginal cell extremely long, somewhat more than twice the length of submarginal cell 1; (16) Rf appendix, i.e. distalmost free abscissa of R, long, somewhat shorter than pterostigma; (17) discal cell subrectangular, longer proximodistally than wide anteroposteriorly and size small; (18) petiolar node broadly convex in profile view (vs. anteroposteriorly narrow and squamiform); (19) penite apices not downcurved, but rather being produced distally as long, linear processes. (IV) Distinguished from Ravavy miafina by the following (note 8): (20) smaller, BL ~ 2.57–2.34, WL ~ 0.43–0.46, FWL ~ 2.17–2.52 (vs. larger, BL ~3.33–4.46, WL ~ 1.30–1.67, FWL ~ 3.85–3.99); (21) head somewhat broader, CI ~ 122 (vs. narrower, CI ~112–114, although 122 in one specimen); (22) malar space relatively long, MSI ~ 6 and space distinctly longer than proximal width of mandible in lateral view (vs. relatively short, MSI ~ 28–32 and distinctly shorter than proximal mandibular width); (23) eye relatively long, OI ~ 40–39 (vs. eye relatively short, OI ~ 14–18); (24) ocelli relatively small, OCI ~ 38 (vs. 44–62); (25) antennomere III but not IV slightly kinked (vs. both antennomeres kinked); (26) mesoscutum relatively long, MI ~ 80 (vs. short, MI ~ 104–131); (27) scutoscutellar sulcus distinctly broad (vs. short, almost line-like); (28) petiole relatively long, PI ~ 80 (vs. short, PI ~ 51–67); (29) bristle-like setae apparently not developed on gastral sternites (vs. such setae present on all gastral sternites except the first); (30) abdominal sternum IX posterior margin broadly convex (vs. broadly emarginate); (31) gonostyli relatively long, length ~ 2–3 × height (vs. short, length ~ 1 × height); (32) gonostylar apices narrowly pointed (vs. broadly rounded); (33) ventral margin of penite more-or-less linear from base to apex (vs. base produced ventrally as a dorsoventrally long, anteroposteriorly narrow, serrate lobe); and (34) apicoventral penite processes short, length from proximodorsal inflection point ~ 1 × height (vs. long, length ~ 4 × height). Notes on diagnosis: 1. A large, rhomboidal clypeal condyle is an apparent synapomorphy of the Dolichoderomorpha, i.e. Dolichoderinae + Aneuretinae. The clypeal condyle of Formicinae and other subfamilies is variable in shape, often anteroposteriorly narrow. 2. The form of the gonopod is an apparent synapomorphy of Dolichoderinae as it is not shared with Aneuretinae. 3. Most dolichoderines have a posteroventral (ventroapical) basivolsellar process, which may be short and triangular to elongate and digitate. The process is absent in Azteca Forel, 1878 and Gracilidris Wild & Cuezzo, 2006 (sister taxa), Ochetellus Shattuck, 1992 and some Dolichoderus Lund, 1831. Formicines do not have a basivolsellar process; such a process is present convergently in the Amblyoponinae, which is a synapomorphy for that subfamily (Yoshimura & Fisher, 2012; Boudinot, 2015). 4. In their recent phylogenetic revision of the Dolichoderinae, Ward et al. (2010) provided reduction or loss of the medial hypostomal lamina as a synapomorphy of the Bothriomyrmecini that occurs in both sexes. Subsequently, Boudinot et al. (2016) recognized loss of forewing cross-vein 2rs-m as another diagnostic condition. Here, we propose the ‘short’ condition of the clypeus as an additional synapomorphy of the tribe. The clypeus in both males and females extends posteriorly between the antennal toruli in the majority of dolichoderine genera with only a few exceptions (Shattuck, 1992; Boudinot, 2015), such as the neotropicus and macro clades of Leptomyrmex Mayr, 1862 (Lucky & Ward, 2010; Boudinot et al., 2016; Barden et al., 2017). 5. Males and queens of the Bornean genus Loweriella Shattuck, 1992 are unknown in contrast to the other three bothriomyrmecine genera (Bothriomyrmex Emery, 1869a, Chronoxenus Santschi, 1919, Ravavy). Because Loweriella is the sister-group of Ravavy (Ward et al., 2010), a number of the diagnostic traits of the latter genus adduced here may be synapomorphies of the two genera. 6. Yoshimura & Fisher (2011) tentatively remarked that additional, vestigial denticles may be present in R. miafina and label a possible denticle on a dissected mandible imaged with a compound microscope. Because this apparent denticle is on the lateral mandibular margin, and as the apical tooth is a consistent feature of even ‘edentate’ mandibles (Boudinot et al., 2021), we interpret the mandible of R. miafina as strictly unidentate. 7. The 6,3 palpomere count is an apparent synapomorphy of Ravavy within the Bothriomyrmecini. Loweriella has a plesiomorphic 6,4 count, while Bothriomyrmex and Chronoxenus share a derived, reduced count which is ≤ 4,3 (Bolton, 2003; Fisher, 2009; Yoshimura & Fisher, 2011). 8. The fine anatomy and morphology of R. miafina is illustrated in Fisher (2009) and especiallyYoshimura & Fisher (2011). The latter work provided a revised diagnosis of this species plus a comparative analysis of genitalic form. Comparisons were made to the images in the literature, on AntWeb (2022), and to specimens examined at the California Academy of Sciences. Description: Integument uniformly imbricate, apparently with a more-or-less even vestiture of extremely short and appressed pubescence; bristlelike setae not visible on body with exception of tarsi, gastral tergites and gonostyli, those of gastral tergites sparse. Head roughly rectangular, narrow anterad eyes and broader posteriorly; malar space relatively broad, being longer than proximal width of mandible in lateral view; posterior head margin emarginate. Compound eye bulging, length slightly less than half head-length, subreniform with subtle emargination occurring on along ventral eye margin and with> 175 ommatidia but probably not more than 200 (complete count not possible). Ocelli small; lateral ocelli relatively wideset, separated from one another by somewhat more than three lateral ocellar lengths. Frontal carinae poorly developed. Antenna 13-merous, scape about as long as compound eye and about 3 × as long as pedicel; pedicel about twice as long as wide and slightly more than half the length of antennomere III; antennomere III slightly kinked at about midlength; antennomeres III–XIII subequal in length and about 4 × as long as wide. Ocellar sensilla (setae) not visible. Anterior clypeal margin more-orless linear; clypeal setation not observable; posterior clypeal margin convex, portion of epistomal sulcus anterad and between toruli linear, weakly concave. Mandibles spiniform, tapering to their unidentate apices, thus appearing wedge-shaped in dorsal view. Labrum deeply notched apically, sides distinctly narrowing apicomediad. Palp formula 6,3; maxillary palp relatively long, exceeding hypostomal margin but not occiput; labial palp short, about one-fourth length of maxillary palp. Prementum elongate, diamond shaped. Stipes without transverse carina. Medial hypostomal lamina absent. Pronotum short, strap-like. Mesoscutum relatively long, with maximum length greater than maximum width; notauli absent; parapsides diverging anteriorly; scutoscutellar sulcus distinctly broad; lateral mesopectal sulcus sinuate. Upper metapleural region distinct, about 4 × as long as broad; lower metapleural region indistinct. Propodeum without distinct dorsal and posterior surfaces; propodeal spiracle small, circular, set slightly below metaphragmal pit. Femora puny, thin; metafemur and metatibia slightly sinuate. Tibial spur formula 1,1. Arolia small, barely visible. Petiole elliptical in lateral view, with broadly convex tergum and sternum; node not squamiform. Abdominal tergum III with broad but shallow concavity anteriorly above helcium. Abdominal segments III–VII similar in size, unmodified. Abdominal sterna VII–VIII broadly emarginate posteriorly. Abdominal sternum IX posterior margin convex, simple. Cerci not visible. Cupula not visible. Gonocoxite length uncertain; gonocoxital apex distinctly offset from gonostylar base. Gonostylus elongate, triangular; length about 2–3 × width; apex narrowly pointed. Volsellae incompletely visible; basivolsellar process present, acute; cuspis absent; lateropenite (= digitus) apex directed ventrad at nearly right angle relative to body of volsella; dorsal margin of digitus almost lobate; digital apices narrowing, almost rod-like and curved laterad. Ventral margins of penites moreor-less linear from base to apex; dorsal margins broadly convex in lateral view; apical processes directed posterad and relatively short, length from proximodorsal inflection point ~ 1 × height. Fore and hind wing venation virtually identical to R. miafina. Jugal lobe absent. Measurements and indices: Dashes indicate metric could not be taken or calculated. Holotype: BL ~ 2.34, HL 0.54, HW 0.41, HWE 0.50, EL 0.21, MSL 0.09, FcW –, SL –, OLL –, OIL –, WL 0.91, ML 0.43, MW –, FWL 2.17, PL –, PW 0.09, PH 0.12, GL 0.86, GW 0.43, CI 92, HWI 122, MSI 6, SI –, OI 39, EPI –, OCI –, MI –, PI –. Paratype: BL ~ 2.57, HL 0.54, HW 0.41, HWE 0.48, MSL 0.09, EL 0.22, FcW 0.12, SL 0.22, OLL 0.04, OIL 0.011, WL 0.98, ML 0.46, MW 0.37, FWL 2.52, PL 0.12, PW –, PH 0.15, GL 0.93, GW 0.54, CI 90, HWI –, MSI 6, SI –, OI 40, EPI –, OCI 38, MI 80, PI 80. Note: For comparison, see the metrics for R. miafina provided below., Published as part of Perrichot, Vincent, Boudinot, Brendon E., Engel, Michael S., Fls, Xu, Chunpeng, Bojarski, Błażej & Szwedo, Jacek, 2022, Ants (Hymenoptera: Formicidae) from Miocene Ethiopian amber: filling gaps in the geological record of African terrestrial biota, pp. 775-791 in Zoological Journal of the Linnean Society 196 on pages 782-785, DOI: 10.1093/zoolinnean/zlac053, http://zenodo.org/record/7195057, {"references":["Forel A. 1878. Etudes myrmecologiques en 1878 (premiere partie) avec l'anatomie du gesier des fourmis. Bulletin de la Societe Vaudoise des Sciences Naturelles 15: 337 - 392.","Wild AL, Cuezzo F. 2006. Rediscovery of a fossil dolichoderine ant lineage (Hymenoptera: Formicidae: Dolichoderinae) and a description of a new genus from South America. Zootaxa 1142: 57 - 68.","Shattuck SO. 1992. Generic revision of the ant subfamily Dolichoderinae (Hymenoptera: Formicidae). Sociobiology 21: 1 - 181.","Lund PW. 1831. Lettre sur les habitudes de quelques fourmis du Bresil, adressee a M. Audouin. Annales des Sciences Naturelles 23: 113 - 138.","Yoshimura M, Fisher BL. 2012. A revision of male ants of the Malagasy Amblyoponinae (Hymenoptera: Formicidae) with resurrections of the genera Stigmatomma and Xymmer. PLoS One 7: e 33325.","Boudinot BE. 2015. 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. European Journal of Taxonomy 120: 1 - 62.","Ward PS, Brady SG, Fisher BL, Schultz TR. 2010. Phylogeny and biogeography of dolichoderine ants: effects of data partitioning and relict taxa on historical inference. Systematic Biology 59: 342 - 362.","Boudinot BE, Probst RS, Brandao CRF, Feitosa RM, Ward PS. 2016. Out of the Neotropics: newly discovered relictual species sheds light on the biogeographical history of spider ants (Leptomyrmex, Dolichoderinae, Formicidae). Systematic Entomology 41: 658 - 671.","Mayr G. 1862. Myrmecologische Studien. Verhandlungen der Kaiserlich-Koniglichen Zoologisch-Botanischen Gesellschaft in Wien 12: 649 - 776.","Lucky A, Ward PS. 2010. Taxonomic revision of the ant genus Leptomyrmex Mayr (Hymenoptera: Formicidae). Zootaxa 2688: 1 - 67.","Barden P, Boudinot BE, Lucky A. 2017. Where fossils dare and males matter: combined morphological and molecular analysis untangles the evolutionary history of the spider ant genus Leptomyrmex Mayr (Hymenoptera: Dolichoderinae). Invertebrate Systematics 31: 765 - 780.","Emery C. 1869 a. Descrizione di una nuova formica italiana. Annuario del Museo Zoologico della Reale Universita de Napoli 5: 117 - 118.","Santschi F. 1919. Fourmis du genre Bothriomyrmex Emery (Systematique et moeurs). Revue Zoologique Africaine 7: 201 - 224.","Yoshimura M, Fisher BL. 2011. A revision of male ants of the Malagasy region (Hymenoptera: Formicidae): key to genera of the subfamily Dolichoderinae. Zootaxa 2794: 1 - 34.","Boudinot BE, Moosdorf OTD, Beutel RG, Richter A. 2021. Anatomy and evolution of the head of Dorylus helvolus (Formicidae: Dorylinae): patterns of sex- and caste-limited traits in the sausagefly and the driver ant. Journal of Morphology 282: 1616 - 1658.","Bolton B. 2003. Synopsis and classification of Formicidae. Memoirs of the American Entomological Institute 71: 1 - 370.","Fisher BL. 2009. Two new dolichoderine ant genera from Madagascar: Aptinoma gen. n. and Ravavy gen. n. (Hymenoptera: Formicidae). Zootaxa 2118: 37 - 52."]}

Perrichot, Vincent, Boudinot, Brendon E., Engel, Michael S., Fls, Xu, Chunpeng, Bojarski, Błażej, Szwedo, Jacek (2022): Ants (Hymenoptera: Formicidae) from Miocene Ethiopian amber: filling gaps in the geological record of African terrestrial biota. Zoological Journal of the Linnean Society 196: 775-791, DOI: 10.1093/zoolinnean/zlac053

GENUS † GERONTOFORMICA NEL & PERRAULT, 2004 = † Sphecomyrmodes Engel & Grimaldi, 2005 junior syn.: Barden & Grimaldi (2016): 518. TAXONOMIC SYNOPSIS OF † GERONTOFORMICA I. Species group of cretacica (newly recognized) (Charentese amber) [Note 1]: 1. † G. cretacica Nel & Perrault, 2004 nomen dubium (new status) [Note 2] – Type species of † Gerontoformica Nel & Perrault, 2004 2. † G. occidentalis (Perrichot et al., 2008) II. Species group of gracilis (newly recognized) (Kachin amber): 3. † G. gracilis (Barden & Grimaldi, 2014) 4. † G. robusta (Barden & Grimaldi, 2014) 5. † G. spiralis (Barden & Grimaldi, 2014) [Note 3] 6. † G. subcuspis (Barden & Grimaldi, 2014) [Note 3] III. Species group of pilosa (Kachin amber): 7. † G. sternorhabda sp. nov. 8. † G. contega (Barden & Grimaldi, 2014) 9. † G. magna (Barden & Grimaldi, 2014) 10. † G. pilosa (Barden & Grimaldi, 2014) IV. Incertae sedis to species group within genus (Kachin amber): (11.) † G. orientalis (Engel & Grimaldi, 2005) nomen dubium (new status) [Note 4] – Type species of † Sphecomyrmodes Engel & Grimaldi, 2005 (12.) † G. rugosa (Barden & Grimaldi, 2014) nomen dubium (new status) [Note 5] (13.) † G. tendir (Barden & Grimaldi, 2014) nomen dubium (new status) [Notes 5, 6] Notes on classification Note 1: Under the label of ‘ orientalis group’, Boudinot et al. (2020b) previously united the cretacica and gracilis groups with the species † G. orientalis. The cretacica and gracilis groups are here divided based on the distinct forms of their petioles. The species † G. orientalis is of insufficient preservation for specieslevel identification (see Note 4 below). Note 2: The holotype of † Gerontoformica cretacica is poorly preserved, as noted by Barden & Grimaldi (2016), with obvious distortions and elongation of the scapes and flagella, and little detail of the body visible. Due to this unfortunate circumstance, we newly consider † G. cretacica to be a nomen dubium. It remains in the cretacica species group of † Gerontoformica, because it is the type species of the genus. However, it will be highly desirable to clarify the identity of this species through examination and description of more Charentese ants. At present, the only distinction between † G. cretacica and the co-eval † G. occidentalis that is not apparently affected by preservation is body size, with the former having a longer mesosoma than the latter (2.1 mm vs. 1.4 mm, respectively). Body size is, of course, highly variable among nestmates of crown ants, thus is a weak diagnostic trait when used in isolation and without the quantification of variation across conspecific individuals. Note 3: † Gerontoformica spiralis and † G. subcuspis were difficult to separate in the present study based on the available anatomical evidence and may be conspecific. Specifically, we observe that, in addition to conditions outlined in the key (see that section below), the two species are highly similar in the following conditions, which we only roughly characterize here: (1) proportions and fine details of the head, including frontal carina shape; (2) the degree of mesonotal, metanotal and propodeal convexity; (3) the width of separation between the meso- and metanota plus metanotum and propodeum; (4) the shape and proportions of the petiolar node; and (5) the form of the third abdominal (first ‘gastral’) segment. It is possible that † G. spiralis and † G. subcuspis represent the smaller and larger ranges of body size of a single species, with the former reported to have a total body length of 4.22–5.22 mm and the latter 5.35–5.76 mm (Barden & Grimaldi, 2014). Our focal uncertainties relate to the shapes and setational patterns of the tarsi of † G. subcuspis, and the form of the subpetiolar process and prora of † G. spiralis. A potential feature separating the two species is the distance of the toruli from the posterior clypeal margin, which appears to be narrower in † G. spiralis relative to † G. subcuspis, but this could be a visual artefact caused by the apparent light distortion in the holotype image of † G. spiralis. We recommend further re-evaluation of these two species, ideally using µ-CT and additional light photography to resolve the uncertainties of the tarsi, toruli and sternal processes of the metasoma. See also Note 1 on the diagnosis of † G. gracilis. Note 4: † Gerontoformica orientalis, the type species of † Sphecomyrmodes, is identifiable as † Gerontoformica relative to † Sphecomyrma Wilson & Brown, 1967 by the absence of the anteromedian clypeal process and presence of traction setae/ chaetae along the anterior clypeal margin, as recognized in the original description (Engel & Grimaldi, 2005) and illustrated in Boudinot et al. (2020b). However, in † Gerontoformica, the species † G. orientalis is unidentifiable due to poor preservation. No details of the head are clearly observable, except for the antenna, mandibles and anterior clypeal margin, while the mesosoma appears strongly distorted, the petiole is obscured, and the metasoma is mostly disarticulated. Boudinot et al. (2020b) placed † G. orientalis in the orientalis species group along with the type species of † Gerontoformica based on the absence of the cinctus of abdominal segment IV. As a more refined placement is not possible, we newly consider this species to be a nomen dubium. Note 5: † Gerontoformica rugosa and † G. tendir are newly considered as nomina dubia due to their poor preservation, being strongly desiccated and thus distorted. Both species appear to have some degree of abdominal segment III petiolation, as observed in the three confirmed members of the pilosa group, but it cannot be determined whether this is natural or exaggerated due to preservation. It is possible, but not yet determinable, that † G. rugosa is conspecific with † G. gracilis. That † G. rugosa or † G. tendir do have sculptured integument remains possible but requires substantiation via additional material of these species. We note that little surface texture variation has been explicitly documented among stem ants thus far. Note 6: † Gerontoformica tendir was defined by Barden & Grimaldi (2014) as having a medial clypeal lobe. This anteromedian lobe not only bears traction setae/chaetae, as previously observed, but is also lateromedially broader and proximodistally shorter than that of † Sphecomyrma. Given the poor preservation, it is possible that the apparent lobate form may be due to distortion of the amber matrix, as the lobe consists of the entire medial portion of the clypeus, which is distinct from the lateral clypeal lobes. Based on direct examination of the holotype, it appears that there is a transverse mesonotal carina in † G. tendir, but this also requires re-evaluation. Without additional specimens having an exaggerated and broadly, medially lobate clypeus, we remain uncertain about the identity of the species. A state of potential value for confirming the identity of † G. tendir from additional material is the absence of teeth on the pretarsal claws, as illustrated by Barden & Grimaldi (2014). Remarks The genus † Gerontoformica currently consists of 13 species, including the newly described † G. sternorhabda. The holotypes of nine of these species are sufficiently preserved for species-level identification, with eight of these being sufficiently defined given the potential synonymy of two (see Note 3 above). At the generic level, † Gerontoformica differs from † Sphecomyrma by presence of traction setae/chaetae along the anterior clypeal margin and absence of a narrow anteromedian clypeal lobe. These two conditions were used by Engel & Grimaldi (2005) to establish the genus † Sphecomyrmodes, which was synonymized under † Gerontoformica by Barden & Grimaldi (2016) after examination of the holotype of the type species of the latter taxon. Collectively, † Gerontoformica and † Sphecomyrmodes have been revised piecemeal after the former’s establishment by Nel et al. (2004). Specifically, Barden & Grimaldi (2014) added nine species of † Sphecomyrmodes, Barden & Grimaldi (2016) transferred all species of † Sphecomyrmodes to † Gerontoformica, and Boudinot et al. (2020b) moved one species to † Myanmyrma and recognized two morphologically defined groups of species in the genus. To understand the shifting boundaries of † Gerontoformica in the light of the present µ-CT-driven study, we detail the species-level history of the genus. Species attributed to the genus are distributed in Kachin amber (11 total) and Charentese amber (two total). The Charentese species, † G. cretacica and † G. occidentalis, were described four years apart by Nel et al. (2004) and Perrichot et al. (2008), respectively. Perrichot et al. described the smaller-bodied † G. occidentalis as † Sphecomyrmodes in comparison to the type species of that genus – † G. orientalis from Kachin amber – without comparison to † G. cretacica. Unfortunately, given the current status of morphological knowledge, the holotypes of both † G. orientalis and † G. cretacica are too poorly preserved to allow for confident specieslevel identification. However, both Charentese species uniquely share a distinct, nearly squamiform petiolar shape, with the node being anteroposteriorly narrow and dorsoventrally tall; this indicates that the Charentese species are closely related to one another, relative to the Kachin species. For this reason, we group † G. cretacica and † G. occidentalis together in the cretacica species group. It is possible that † G. cretacica and † G. occidentalis are conspecific, but any taxonomic action should wait for the accumulation and processing of more material from the Charentese formation. The identifiable Kachin species of † Gerontoformica, i.e. excluding † G. orientalis, † G. rugosa and † G. tendir, are evenly distributed in the G. gracilis and G. pilosa groups, with four species each. All species of the pilosa group are highly distinct in body form and setation, with † G. sternorhabda being an outlier, having the smallest body size and least pronounced constriction of the fourth abdominal segment, in addition to other defining features (see the species diagnosis below). In contrast, species of the newly recognized gracilis group are all largely similar to one another, without easily recognizable diagnostic structures. The most distinct Kachin species of the gracilis group is † G. robusta, which has a boxy mesosoma, with the meso- and metanota forming a nearly linear dorsal margin in lateral view. The other species of the gracilis group have a multi-humped profile due to the bulgelike development of the meso- and metanota and are otherwise similar to one another (see Note 1 on the diagnosis of † G. gracilis)., Published as part of Boudinot, Brendon E., Richter, Adrian, Katzke, Julian, Chaul, Júlio C. M., Keller, Roberto A., Economo, Evan P., Beutel, Rolf Georg & Yamamoto, Shûhei, 2022, Evidence for the evolution of eusociality in stem ants and a systematic revision of † Gerontoformica (Hymenoptera: Formicidae), pp. 1355-1389 in Zoological Journal of the Linnean Society 195 on pages 1361-1363, DOI: 10.1093/zoolinnean/zlab097, http://zenodo.org/record/6994456, {"references":["Nel A, Perrault G, Perrichot V, Neraudeau D. 2004. The oldest ant in the Lower Cretaceous amber of Charente- Maritime (SW France) (Insecta: Hymenoptera: Formicidae). Geologica Acta 2: 23 - 29.","Engel MS, Grimaldi DA. 2005. Primitive new ants in Cretaceous amber from Myanmar, New Jersey, and Canada (Hymenoptera: Formicidae). American Museum Novitates 3485: 1 - 23.","Barden P, Grimaldi D. 2016. Adaptive radiation in socially advanced stem-group ants from the Cretaceous. Current Biology 26: 1 - 7.","Perrichot V, Nel A, Neraudeau D, Lacau S, Guyot T. 2008. New fossil ants in French Cretaceous amber (Hymenoptera: Formicidae). Naturwissenschaften 95: 91 - 97.","Barden P, Grimaldi D. 2014. A diverse ant fauna from the mid-Cretaceous of Myanmar (Hymenoptera: Formicidae). PLoS One 9: e 93627.","Boudinot BE, Perrichot V, Chaul JCM. 2020 b. † Camelosphecia gen. nov., lost ant-wasp intermediates from the mid-Cretaceous (Hymenoptera, Formicoidea). ZooKeys 1005: 21 - 55."]}

† GERONTOFORMICA GRACILIS (BARDEN & GRIMALDI, 2014) (FIGS 1A, B, 2A, B, D, E, G, H, J, K, 8, 9, 10, 11 (1’-2), 12(7-1); SUPPORTING INFORMATION, FIG. S3) † Sphecomyrmodes gracilis Barden & Grimaldi, 2014: 4–7, figs 2, 10B, 11C, D (wingless female, Kachin amber, JZC-Bu324A, AMNH). Combination in † Gerontoformica: Barden & Grimaldi (2016): 518, suppl. info. p. 16. Diagnosis (wingless female) Similarly identifiable as † G. sternorhabda to the genus † Gerontoformica (I–III above), including absence of the anteromedian lobate clypeal process [Note 1]. For the diagnosis and redescription, see Figures 2, 8 and 10 primarily. I. Among † Gerontoformica, with the following unique character : (1) proximal labial palpomere with a distinct, apicomedially situated lobate process (Fig. 7A, B) [Note 2]; II. Within † Gerontoformica, identifiable as a member of the gracilis species group : (2) mesoscutum without distinct, raised transverse carina separating mesoscutal and mesoscutellar regions (Fig. 10A); (3) tergum of petiolar node anteroposteriorly longer than dorsoventrally tall (high); and (4) abdominal segment IV without cinctus dividing the tergum and sternum into pre- and postsclerites; III. Within the gracilis species group, identified by the following : (5) petiole bun-like, with anteroposteriorly long node; vs. subsquamiform, with anteroposteriorly narrow node († G. cretacica and † G. occidentalis); (6) meso- and metanota bulging, with their dorsal silhouette bihumped; vs. meso- and metanota not bulging, with their dorsal silhouette forming an almost straight line († G.robusta);(7)transverse dorsal sulci of mesosoma broad, separating the meso- and metanota and the metanotum and propodeum by at least one tarsomere width; vs. these sulci not as broad, with the meso- and metanota and the metanotum and propodeum separated by less than one tarsomeral width († G. spiralis, † G. subcuspis) [Note 1]; and (8) propodeal spiracle covered anteriorly by an anterior flange which is directed posterolaterally; vs. spiracle not flanged († G. spiralis, † G. subcuspis; state uncertain for † G. robusta) [Note 3]. Notes on the diagnosis Note 1: † Gerontoformica gracilis is quite similar overall to † G. spiralis and † G. subcuspis. The primary structural distinction is the width of the dorsal transverse sulci that divide the mesonotal and metanotal regions, and the metanotal region and the propodeum, as well as development of an anterior flange around the propodeal spiracles. The character used by Barden & Grimaldi (2014) to distinguish † G. spiralis and † G. gracilis was the distance between the pro- and mesocoxae (their couplet 9); this is certainly a matter of preservation, as the prothorax of the holotype of † G. gracilis is elevated relative to the mesothorax, and the coxae are promoted anteriorly. Such pronotal elevation is possible in ant species with a mobile promesonotal articulation. The defining feature of † G. subcuspis provided by Barden & Grimaldi (2014) is the form of the subpetiolar process, which has an almost vertically oriented anterior margin in profile view. The process of † G. gracilis is more evenly rounded from posterior to anterior, while that of † G. spiralis is not visible. These distinctions should be re-evaluated in future study. See also Note 3 on the synopsis of † Gerontoformica classification above. Note 2: We do not know the distribution of the process of the proximal labial palpomere across † Gerontoformica, with the exception of its absence in † G. sternorhabda. Despite this, we note the development of these processes as they are unique to our knowledge of both extant and extinct species. Because of the difficulty of evaluating proximal palpomeres for ants in general, and especially for fossil ants, we strongly recommend the application of µ-CT methods to determine the phylogenetic extent of this obvious apomorphy. Note that it is possible that the holotype lacks this condition, as we discovered this character after our chance to directly examine the type specimen. Note 3: The anterior flange or hood of the propodeal spiracle is present in most † Gerontoformica examined, with the exception of † G. spiralis and † G. subcuspis. The flange could not be evaluated for † G. magna or † G. robusta. Measurements and indices Adult, specimen C-32: HWed = 0.71; HWev = 0.78; EWl = 0.17; HD = 0.62; ML = 1.49; PnLi = 0.59; PnWa = 0.18; MnL = 0.35; AIIILm = 0.52; AIIILl = 0.65; HPI = 0.99; HSI = 0.47; AIIILI = 0.80. Pupa, specimen C-31: HWed = 0.72; HWev = 0.71; EWl = 0.18; HD = 0.62; ML = 1.62; PnLi = 0.60; PnWa = 0.25; MnL = 0.34; AIIILm = 0.62; AIIILl = 0.73; HPI = 0.86; HSI = 0.44; AIIILI = 0.85. Redescription: adult Head: The head is narrow in facial view, i.e. it is lateromedially narrower than anteroposteriorly long as measured from the anterior clypeal margin to the apparent posterior head margin (Fig. 2K); in posterior and lateral view, the head is dorsoventrally broad, with the vertexal region dome-like; standing setae are most conspicuous on the clypeus. The compound eyes are situated in the posterior third of the head; they bulge laterally, breaking the silhouette of the lateral head margins in facial view; their height mesonotal carina; mspct, mesopectus; mtbt, metabasitarsus; mtcx, metacoxa; mtfm, metafemur; mtnt, metanotum; mtpfm, metaprefemur; mtplglvf, ventral flange of the metapleural gland; mtptc, metapretarsal claw; mttbsa, anterior spur of metatibia; mttbsp, posterior spur of metatibia; mttr, metatrochanter; pd, pedicel; pl, labial palp; pm, maxillary palp; pbt, probasitarsus; pnt, pronotum; ppd, propodeum; ppdsf, anterior flange of propodeal spiracle; pptc, propretarsal claw; pt, petiole; ptn, petiolar node; pts, petiolar sternum; ptsp, subpetiolar process; sc, scape; tgl, laterotergite; to, antennal torulus. above the surrounding surfaces of the cranium is comparatively high; their ommatidia count appears similar to that of † G. sternorhabda; they are apparently glabrous, i.e. lacking interstitial setation. The three ocelli are completely developed. The frontal carinae diverge posterolaterally toward but not reaching the compound eyes; they are circular in form, i.e. curving evenly from their anterior termini to their posterior termini which are directed anterolaterally, rather than posterolaterally; their anterior termini are distant from the epistomal line; the minimum distance between the frontal carinae is about 0.26× maximum head width as measured in full face view. The antennal scrobes are anteroposteriorly short and encircled by the frontal carina. The antennal toruli are distant from the posterior clypeal margin; they are in the form of a low, more-or-less even ring. The antennae are 12-merous. The scapes are somewhat flattened and curved; they are about four times as long as their maximum width; their length is somewhat more than half the width of the head, and somewhat less than half the length of the head; they lack standing setae along their shafts. The pedicels are slightly longer than twice their width; they are about one-third the length of the scapes, and about half the length of the third antennomere; they apparently lack standing setae. The flagellae are longer than the mesosoma and are simple, i.e. not thickening distally; they bear a range of standing and appressed setae; flagellomeres I are the longest, being somewhat more than four times as long as wide and about three fourths the length of the scape; flagellomeres II–IX are all longer than the pedicel; flagellomeres X are longer than each pair of flagellomeres from II to IX. The clypeus is about three times as wide lateromedially as long anteroposteriorly, with the length measured from the midpoints of the anterior and posterior clypeal margins and the width measured between the lateralmost points of the clypeus. The lateroclypeal areas are formed as lateral lobes. The medioclypeal area is anteriorly convex; its length at the midline of the head is about 0.21× head length also at midline, as measured in full-face view; it bears an array of standing (rather than appressed) setae on its disc and is margined anteriorly by chaetae. The mandibles are simple and apically bidentate. The maxillary palps are 6-merous (Figs 2B, H, 7A–D) [Note 1]; they are elongate, being almost as long as the head and longer than the scapes and mandibles; the proximal palpomeres are short compared to the others; the second palpomeres are dorsoventrally flattened and lobate apicomedially; the third through sixth palpomeres are long, thin, and cylindrical. The labial palps are 4-merous (Figs 2B, H, 7A–D) [Note 1]; they are short, being less than half the length of the maxillary palps, and with their individual lengths shorter than each of the maxillary palpomeres with the exception of the proximal maxillary ones; the proximal labial palpomere is narrow proximally and bears a distinct lobate process medially at about its apical third, with this process being about as long as wide; the second and third palpomeres are thickened medially; the fourth palpomeres are relatively more cylindrical. Mesosoma: The pronotum bears an anteromedian neck process, and lateral and posteromedian flanges; these flanges are not flared; the muscular node or ‘disc’ of the pronotum is hemispherical as observed in lateral view and almost elliptical in dorsal view, being distinctly longer than tall in lateral (profile) view, thus appearing narrow; setation was not observed on the pronotum [Note 2]. The pronotal lobes are well developed (Fig. 6E). The mesonotum is not divided into an anterior mesoscutal area and a posterior mesoscutellar area, i.e. the transverse mesonotal carina is not developed; the notum is convex in lateral view and is almost flattened along most of its length. The mesopectus is not clearly divided into dorsal and ventral areas; its dorsoventral height is almost 1.5× that of its anteroposterior length. The transverse mesonotal sulcus is anteroposteriorly long/broad. The metanotum is developed as a distinct and almost evenly convex bulge. The transverse metanotopropodeal sulcus is anteroposteriorly broad and continues ventrally toward the base of the mesocoxa, completely separating the lateral metapectal area from the lateral mesopectal area. The metapleural gland orifice is small and not remarkably hairy (Fig. 6F); it does not have a distinct bulla; it is margined dorsally and ventrally by flanges, including the ventral metapleural gland flange. The propodeum is rounded, with the dorsal and posterior margins curving broadly into one another in lateral view; its posterior surface is convex; it does not, apparently, bear standing setae. The propodeal spiracles are situated distant from the metanotum and below the dorsal propodeal margin as seen in lateral view, but they are located in the anterodorsal fourth of the sclerite. The propodeal lobes are apparently not developed. Legs: The legs are developed as expected for the Formicidae, with some notable characters and states. With the exception of the tibial apex and tarsi, they appear almost entirely glabrous. The state of the apical protibial foramina is uncertain. The mesoprefemora and metaprefemora are welldeveloped and are broader ventrally than dorsally. The protibia bears and anterior brush of dense suberect setae in their apical third, near the calcar. Each of the mesotibia and metatibia bear a pair of apicoventral spurs; the anterior tibial spurs are barbirulate; the posterior tibial spurs are simple. The calcar is apparently bifid apically, with one point being the apex of the elongate velum and the other point being a small array of hairs; two stout setae are developed posterior to the calcar. The plantar lobes of the tarsi are well developed, the ventral setation is sparse, and the apical row of chaetae is thinner. The fourth tarsomeres are only weakly notched distally, thus appearing cylindrical in dorsal view. The pretarsal claw teeth are well developed and located just past the midlength of their respective claws. The aroliae are well developed and comparatively large (Fig. 8D); they are nearly as long as the pretarsal claws. Metasoma: The petiole is nodiform and lacks tergosternal fusion between its postsclerites. The petiolar tergum is anteroposteriorly longer than dorsoventrally tall; it is asymmetrically convex, being somewhat longer anteriorly than posteriorly. The developmental state of the laterotergites is uncertain. The petiolar sternum is anteriorly flat; its main portion is at least twice as long as tall; it is narrowly convex in cross-section at its midpoint; it is weakly produced anteroventrally as a low, lobate subpetiolar process, which is shorter dorsoventrally than wide anteroposteriorly; posteriorly, the sternum is not distinctly concave or notched. The helcium is narrow relative to the third abdominal postsclerites; the helcial tergite conceals the helcial sternite in lateral view. The abdominal posttergite III is not constricted posteriorly and is not fused with the third poststernite; it is not necked or shouldered anteriorly, as the tergum evenly curves from its anterior base to posterior margin in lateral view. The abdominal tergosternal margin III is weakly curved, without distinct ‘shouldering’ as observed in various Formicinae and Dolichoderinae. The abdominal poststernite III is not constricted posteriorly, but is weakly angled lateromedially, and bears the prora anteroventrally. The prora is lip-like in lateral view, being anteroposteriorly short and moreor-less transverse in ventral view. The abdominal segments IV, V, and VI are not divided into pre- and postsclerites by a cinctus; they are homonomous in form, i.e. highly similar in shape, size and other qualities of appearance. The seventh abdominal tergum is somewhat dome-like. The seventh abdominal sternum is lateromedially cupped and narrowed distally. The sting is long and narrow. The third valvulae are digitate in form and highly exserted as preserved. Preservation: The specimen CASENT0741232 is exceptionally well preserved internally, particularly the head and mesosoma (Fig. 9). The metasoma is, however, less well preserved, and has some apparent fungal growth or decay bubbling emanating from between the third and fourth abdominal segments. There are several fractures around the specimen, including on the dorsal surface of the head, the ventral right side of the head, across the left side of the mesosoma, and across the petiole and the left side of the third and fourth abdominal segments. The right side of the face, opposite from where the scape is held, is indented, thus appearing to have a longitudinal scrobe (Fig. 10); this is definitely an artefact, as it is not symmetrically present on the left side of the face where the scape is preserved in a position that is distant from the head capsule. Similarly, the mesosoma is indented where the legs are in close proximity (Fig. 10). The left metatrochanter appears distorted in shape. Notes on description Note 1: The original description recognized 4-merous maxillary palps and did not state a labial palpomere count. A 6, 4 palp formula (sensu Bolton, 1994, 2003) was determined here based on direct examination of the holotype, and from our microphotographs and digital renders. As noted in the diagnosis, the process of the proximal labial palpomere is unique among † Gerontoformica, given our current knowledge. Note 2: We are not certain whether † G. gracilis is largely glabrous or whether it has appressed pubescence in various places. At the least we expect pubescence on inter-sclerite contact surfaces. Description: pupa The pupa (Figs 2A, D, G, J, 7, 10) is encased in a cocoon with a black meconium at the caudal end (Fig. 1A). Most structures of species-level identificatory value are incompletely developed, including the frontal carinae, perioral sclerites, mesosomal dorsum and anterior metasoma. The preservation of the specimen is fine externally but is poor internally, with the body cavity filled with a single, solid mass, despite apparent distinctions as seen from light microscopy. In overall appearance, the specimen is relatively bubbly or puffy looking, and has the propodeum crushed and incompletely differentiated from the petiole. Those metasomal segments which are posterior to the petiole are bulging, thus they appear slightly constricted as with a cinctus, but cincti are apparently absent. The metanotal and propodeal spiracles are visible externally, but those of the metasoma are not. Features of note include the following: The head is longer anteroposteriorly than broad lateromedially; the mandibles are contacting one another apically; the maxillolabial complex is exserted, with the palps and glossa clearly visible in ventral view; the maxillary palps are 6-merous; the labial palps are 4-merous; the antennae are directed caudally and reach the posterior margin of the third abdominal segment; the pronotum is longer than tall in lateral view; and the mesonotum lacks a transverse ridge, thus is not divided into anterior mesoscutal and posterior mesoscutellar regions. The pupa differs from the synincluded adults of both species as follows: The antennae appear wideset; the clypeus is anteroposteriorly longer, apparently with an additional band of cuticle along the anterior m a r g i n; t h e m a n d i b l e s a r e o b l i q u e l y o r i e n t e d relative to head length, converging anterad (vs. perpendicular to the long axis of the head at closure); and the labrum is apically notched and with distinct paramedian lobes. Remarks on the pupa Because little comparative work on pupal morphology has been done in Formicidae, it is basically unknown what the polarity of the ‘general features of note’ is within the family. Further, because there are no developmental series of anatomy available for pupal transformation, we cannot be certain of the stage that this specimen was preserved. The conditions listed above which differ between the pupa and adults are especially interesting, as they suggest that there are intermediate stages in the structural rearrangement of the body. For example, the wideset toruli and exceptionally long clypeus are hard to explain without an atlas of developmental transformation. We note the apically bilobate labrum, in particular, as the labra of the adults are apically rounded. At present we are unable to explain the labral difference, as there are no wo, Published as part of Boudinot, Brendon E., Richter, Adrian, Katzke, Julian, Chaul, Júlio C. M., Keller, Roberto A., Economo, Evan P., Beutel, Rolf Georg & Yamamoto, Shûhei, 2022, Evidence for the evolution of eusociality in stem ants and a systematic revision of † Gerontoformica (Hymenoptera: Formicidae), pp. 1355-1389 in Zoological Journal of the Linnean Society 195 on pages 1369-1377, DOI: 10.1093/zoolinnean/zlab097, http://zenodo.org/record/6994456, {"references":["Barden P, Grimaldi D. 2014. A diverse ant fauna from the mid-Cretaceous of Myanmar (Hymenoptera: Formicidae). PLoS One 9: e 93627.","Barden P, Grimaldi D. 2016. Adaptive radiation in socially advanced stem-group ants from the Cretaceous. Current Biology 26: 1 - 7.","Bolton B. 1994. Identification guide to the ant genera of the world. Cambridge: Harvard University Press, 222.","Bolton B. 2003. Synopsis and classification of Formicidae. Memoirs of the American Entomological Institute 71: 1 - 370.","Boudinot BE, Moosdorf OTD, Beutel RG, Richter A. 2021 b. Anatomy and evolution of the head of Dorylus helvolus (Formicidae: Dorylinae): patterns of sex- and castelimited traits in the sausagefly and the driver ant. Journal of Morphology 282: 1616 - 1658. Doi: 10.1002 / jmor. 21410."]}

† GERONTOFORMICA STERNORHABDA SP. NOV. (FIGS 1C, D, 2C, F, I, L, 4– 6, 7E, F, 11 (1-2), 12(2-1, 2-3, 2-4), 13(3’-1); SUPPORTING INFORMATION, FIGS S1, S 2) Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org:act: E77C1A20-1398-465F-9607-AD11E97ADF1C Type material: Holotype. Wingless female (w). Myanmar, Kachin State: Hukawng Valley [CASENT 0741233 in an amber piece labelled AMNH SY-23 and deposited at AMNH]. Paratypes. Wingless females (w). Synincluded with holotype [CASENT0741234]; same locality as synincluded holo- and paratypes [UFV-LABECOL-009656, deposited in CELC]. Diagnosis (wingless female) Conforming to the diagnosis of the Formicidae (see: Boudinot et al. 2020a, b), including the following key conditions: (1) postgenal bridge elongated, thus head ‘prognathous’ (Fig. 2C); (2) cranial condyles of the anterior/ dorsal mandibular articulation enlarged (Fig. 5B); (3) toruli oriented dorsolaterally rather than simply dorsally (Fig. 5A); (4) procoxae elongate, about twice as long as wide (Fig. 2C); (5) prodisticoxal foramen ‘closed’ and protrochanter narrowly necked (Fig. 4F) [Note 1]; (6) meso- and metathoracicocoxal articulations ‘closed’, i.e. directed ventrally rather than laterally or ventrolaterally (Fig. 4F); (7) abdominal segment II completely petiolated (Fig. 4F); (8) subpetiolar process present (Fig. 4F); (9) prora present (Fig. 4F). I. Among Formicidae, identifiable as †Sphecomyrminae: (1) Mandibles simple and bidentate (Fig. 5A), without: (a) being strongly bowed and multidentate as in the † Camelomecia group, (b) the scythe-like blade as in the haidomyrmecine † Haidomyrmex clade, (c) projecting anteriorly with numerous teeth as in the haidomyrmecine † Aquilomyrmex clade, or (d) the strong torsion of † Zigrasimeciini (the state in † Boltonimecia is uncertain); (2) the antennal scrobes on the face do not extend all the way to the compound eye (Fig. 4B) (such scrobes observed in †Zigrasimeciinae, including † Boltonimecia) [Note 2]; (3) anterolateral corners of head not produced as robust triangles (Fig. 5A) (such corners observed in † Dilobops of the †Haidomyrmecinae); and (4) (a) scapes shorter than the width of the head (Fig. 5A) and (b) clypeus not extending posteriorly between the toruli (Fig. 2L) (such extension observed in † Brownimeciinae); II. Within †Sphecomyrminae, identifiable as † Gerontoformica : (5) anteromedian clypeal margin not produced as distinct medial lobe (Fig. 2L); vs. such a lobe present († Sphecomyrma, † G. tendir) [Note 3]; and (6) (a) mandibles short and fitting against clypeus when closed (Fig. 5A) and (b) metanotum developed (Fig. 2C); vs. mandibles elongate and metanotum not developed († Myanmyrma); III. Within † Gerontoformica, with the following unique condition : (7) the anteroventral (‘subpetiolar’) process of the petiolar sternum long, orthogonal to the longitudinal axis of the petiole, and more-or-less rod-like, i.e. with anterior and posterior margins parallel to subparallel (Fig. 2C); vs. short and triangular († G. gracilis, † G. occidentalis, † G. pilosa, † G. robusta, † G. rugosa, † G. spiralis, † G. subcuspis, † G. tendir) [Note 4]; IV. Within † Gerontoformica, identifiable as a member of the pilosa species group : (8) mesoscutum with distinct, raised transverse carina separating mesoscutal and mesoscutellar regions (Fig. 6B); vs. such a carina entirely absent or only poorly developed laterally, thus incomplete medially and not forming a distinct angle between the mesoscutal and mesoscutellar regions in profile view (cretacica, gracilis groups); (9) tergum of petiolar node anteroposteriorly longer than dorsoventrally tall (Fig. 2C); vs. petiolar node tergum taller than long (cretacica group); and (10) abdominal segment IV (metasomal III) with the cinctus distinct and impressed, i.e. divided into pre- and post-sclerites by a transverse sulcus (Fig. 5E); V. Within the pilosa species group, distinguished from all species by the following : (11) cinctus developed, but transverse sulci weakly impressed, thus pre- and postsclerites of abdominal segment IV not meeting at strongly oblique angle (Fig. 5E); vs. transverse sulci deeply impressed, thus pre- and postsclerites meeting at distinct oblique angle († G. contega, † G. magna, † G. pilosa); (12) head in full-face view broader lateromedially than long anteroposteriorly, excluding eyes (Fig. 2L); vs. head longer than broad († G. contega, † G. magna, † G. pilosa) [Note 5]; (13) pretarsal claws edentate (Fig. 5G, I, J); vs. each claw with a single tooth of variable location († G. contega, † G. magna, † G. pilosa) [Note 6]; and (14) body small, mesosoma length 1.5 († G. contega, † G. pilosa) and> 2.5 († G. magna); VI. Further distinguished by species in the pilosa group by the following : (15) anteromedian clypeal margin distinctly evenly curved to an ‘incision’ at the point of contact with the rounded lateroclypeal lobes (Fig. 2L); vs. anteromedian clypeal margin weakly convex, without distinct incision between the medioclypeus and lateroclypeal lobes, which are themselves anterolaterally angled († G. contega); (16) in profile, pronotum evenly curved, mesoscutum more-or-less aligned with mesoscutellum and metanotum, and propodeal dorsal and posterior surfaces curving into one another obliquely (Fig. 2C); vs. pronotum, mesonotum, and propodeum each shouldered in appearance, i.e. pronotum with strong anterior dorsolateral bulge, mesonotum with mesoscutal and mesoscutellar regions meeting at nearly a right angle, and propodeal dorsal and posterior surfaces also meeting at nearly a right angle († G. contega); (17) standing setation on body short, relatively sparse (with the exception of the propodeum and petiolar tergum) (Fig.5); vs. longer setation present († G. magna, † G. pilosa); and (18) anteroventral process of abdominal sternum III (metasomal II) robust but short anteroposteriorly (Fig. 2C); vs. prora anteroposteriorly long, sharkfin-like in form († G. pilosa) [Note 7]. Notes on the diagnosis Note 1: The right procoxa and protrochanter of the holotype are slightly disarticulated. However, the focal details to evaluate are the circular shape of the prodisticoxal foramen and the crank-like (curved) and thin proximal neck of the protrochanter, which can be evaluated from Figure 4F. Note 2: The only species of †Sphecomyrminae to have an elongate antennal scrobe is † G. contega. Although the scrobe of this species was observed via direct examination of the holotype, it remains a possibility that this is an artefact because other specimens of † Gerontoformica may have asymmetrical, sulcuslike distortions of the head corresponding to the position of the scrobe (see the ‘preservation’ section of the † G. gracilis description; Fig. 10). Conducting a µ-CT scan of the type specimen and/ or the accrual of additional specimens are necessary. N o t e 3: T h e a n t e r o m e d i a n c l y p e a l p r o c e s s o f † Sphecomyrma is lateromedially thin and at least as long as broad; it is distinctly bordered laterally by the medioclypeus, i.e. the clypeal ‘disc’ between the lateral clypeal lobes. In contrast, the entire medioclypeus of † G. tendir is apparently produced. See Note 5 of the ‘Notes on classification’ section above. Note 4: The form of the subpetiolar process is unknown for four species within the genus: † G. contega, † G. cretacica, † G. magna and † G. orientalis. Note 5: Although it is difficult to evaluate the head shape of † G. contega from the holotype due to cut of the amber matrix, the head does appear to be longer than broad. Reevaluation of this condition through µ-CT is recommended. Note 6: Teeth on the pretarsal claws have been recorded for most species of † Gerontoformica († G. contega, † G. gracilis, † G. magna, † G. occidentalis, † G. pilosa, † G. robusta, † G. spiralis and † G. subcuspis). The condition of having reduced teeth on the pretarsal claws, however, is shared with † G. cretacica and † G. tendir. Notably, Perrichot et al. (2008) recorded minute teeth on the pretarsal claws of † G. occidentalis, which suggests the need to re-evaluate the holotype of † G. cretacica. Note 7: The prora is clearly an important structural feature for distinguishing stem ants but is not clearly visible in most specimens, including † G. contega and † G. magna in the pilosa group, † G. cretacica and † G. orientalis in the cretacica / orientalis group, plus † G. rugosa and † G. tendir. It is extremely small and nearly absent in † G. gracilis and † G. occidentalis, and it is developed, but short and comparatively inconspicuous in † G. robusta, † G. spiralis, and † G. subcuspis. MEASUREMENTS AND INDICES Holotype, specimen C-33: HWed = 0.87; HWev = 0.95; EWl = 0.14; HD = 0.55; ML = 1.25; PnLi = 0.50; PnWa = 0.30; MnL = 0.27; AIIILm = 0.48; AIIILl = 0.53; HPI = 0.64; HSI = 0.69; AIIILI = 0.91. Paratype, specimen C-34: HWed = 0.94; HWev = 1.06; EWl = 0.20; HD = 0.66; ML = 1.39; PnLi = 0.45; PnWa = 0.27; MnL = 0.28; AIIILm = 0.50; AIIILl = 0.64; HPI = 0.70; HSI = 0.67; AIIILI = 0.78. UFV-LABECOL-009656: HWed = 0.79; HWev = 0.85; EWl = 0.11; HD = 0.46; ML = 1.05; PnLi = (~0.35–0.38); PnWa = –; MnL = 0.29; AIIILm = 0.4; AIIILl = 0.48; HPI = 0.58; HSI = 0.75; AIIILI = 0.83. Description Head: The head is broad in facial view, i. e. lateromedially wider than anteroposteriorly long from the anterior clypeal margin to the apparent posterior head margin (Fig. 2L); in posterior or lateral view, the head is dorsoventrally narrow, with the vertexal region not being particularly domed; standing setae are present on the vertexal and facial regions [Note 1]; there are short, decumbent setae distributed sparsely on the head capsule, a few longer, and suberect setae medially on the vertex near the ocelli. The compound eyes are situated in the posterior third of the head (Fig. 2L); they bulge laterally, breaking the silhouette of the lateral head margins in facial view; their height above the surrounding surfaces of the cranium is comparatively low; they comprise over 100 ommatidia, but not more than 200 (Fig. 4C) [Note 2]; they are apparently glabrous, i.e. lacking interstitial setation. The three ocelli are completely developed (Fig. 2L). The frontal carinae diverge posterolaterally, toward but not reaching the compound eyes (Figs 2L, 5A); they are sinuate in form, i.e. from their anterior margins they are directed relatively medially then bulge laterally before curving laterad; their anterior termini are close to the epistomal line, but not extending on to the clypeus; the minimum distance between the frontal carinae is about 0.29× maximum head width as measured in full face view. The antennal scrobes, or depressed contact surfaces of the face laterad the frontal carinae, are parallel in orientation relative to the frontal carinae and are apparently longer than wide (Figs 2L, 4B). The antennal toruli abut but do not indent the posterior clypeal margin [Note 3]; they are in the form of a low, more-or-less even ring. The antennae are 12-merous (Fig. 4B). The scapes are somewhat flattened and curved (Fig. 2C, F, I, L); they are about three to four times as long as wide (Fig. 2I); their length is about half the width of the head, and less than half the length of the head; they bear a vestiture of short subdecumbent to suberect setae. The pedicels are about twice as long as wide (Fig. 4F); they are about one-third the length of the scapes, and somewhat more than half the length of the third antennomere; their setation is similar to that of the scapes. The flagellae are longer than the mesosoma and are simple, i.e. not thickening distally (Fig. 4F); they bear a range of standing and appressed setae; flagellomeres I are the longest, being about four times as long as wide and more than half the length of the scapes; flagellomeres II–IX are about as long as the pedicel; flagellomeres X are longer than flagellomeres II–IX. The clypeus is about four times as wide (lateromedially) as long (anteroposteriorly), with the length measured from the midpoints of the anterior and posterior clypeal margins and the width measured between the lateralmost points of the clypeus (Fig. 2L). The lateroclypeal areas are formed as lateral lobes (Figs 2L, 5A). The medioclypeal area is anteriorly convex (Fig. 2L); its length at the midline of the head is about 0.21× head length also at midline, as measured in full-face view; it bears five or six long and flexuous setae that are situated near the anterior clypeal margin and anteriorly directed, consisting of one medial seta surrounded by a pair of setae and potentially a second, even more lateral pair [Note 4]; the anterior medioclypeal margin bears a row of chaetae. The mandibles are simple and apically bidentate (Fig. 5A). The maxillary palps are 5-merous (Figs 2I, 4A, 7E, F) [Notes 5, 6]; they are conspicuously short, with their total lengths shorter than the lengths of either the mandibles or scapes; with the exception of the apical palpomere, they are thick and bulging at about their midlengths; they are adorned with erect and appressed pilosity. The labial palps are 4-merous (Figs 2I, 4A, 7E, F) [Note 6]; they are short, being just over half the length of the maxillary palps; with the exception of the apical palpomere, they are more-orless conical and thickening toward their apices; they are adorned with erect and appressed pilosity; the proximal palpomere lacks a distinct process. Mesosoma: The pronotum bears an anteromedian neck process, and lateral and posteromedian flanges (Figs 2C, F, 5C, 6A); these flanges are flared in the paratype specimen C-34 (Fig. 4C) but not in the holotype or other paratype; the muscular node or ‘disc’ of the pronotum is almost spheroidal in shape, with the lateral margins strongly convex in dorsal view and the dorsal margin strongly convex in lateral view, and with an anteroposterior length approximating its dorsoventral height; pronotal setation is sparse, being represented by a few subdecumbent setae (Fig. 6A). The pronotal lobes are well developed (Fig. 6A) [Note 7]. The mesonotum is divided into an anterior mesoscutal area and a posterior mesoscutellar area by the transverse mesonotal carina (Figs 2C, F, 4C, F, 5C, 6B). The mesoscutal area is approximately in the form of a low saddle (i.e. is a low hyperbolic paraboloid), with a concave dorsal margin in lateral view, and with the anterior rim more upcurved than the posterior rim (Fig. 2C). The mesoscutellar area is convex, but sunken relative to the mesoscutal area (Fig. 2C). The mesopectus is divided into dorsal and ventral areas by a longitudinal sulcus (Fig. 2C); its dorsoventral height is about equal to its anteroposterior length. The dorsal mesopectal area is approximately rectangular in shape, being somewhat more than twice as long anteroposteriorly as tall dorsoventrally (Fig. 2C); its dorsoventral height is one-third the dorsoventral height of the lower mesopectal area as measured from the transverse sulcus directly ventrad to an imaginary line drawn parallel to the ventrolateral margins of the meso- and metapecta. The ventral mesopectal area is approximately triangular in shape, being broad along is dorsal margin and narrowing posteroventrally to its ventrolateral margin (Fig. 2C); its anteroposterior length along its dorsal margin is approximately equal to its dorsoventral height. The transverse mesometanotal sulcus is anteroposteriorly short/ thin (Fig. 2C). The metanotum is developed as a distinct bulge between the metanotal spiracles (Fig. 2C). The transverse metanotopropodeal sulcus is anteroposteriorly long/ broad and continues ventrally toward the base of the mesocoxa, completely separating the lateral metapectal area from the lateral mesopectal area (Fig. 2C). The metapleural gland orifice is large, hairy, and in the form of a broad subelliptical pit (Fig. 6B); it is margined dorsally by a bulge, the metapleural gland bulla, and ventrally by the ventral metapleural gland flange, which itself is a continuation of the sinuate ventrolateral carina of the metapectus and is spiniform and dorsally inclined. The propodeum is boxy (Figs 2C, 6B) [Note 8]; it bears standing setae and is the hairiest mesosomal region. The propodeal spiracles are situated distant from the metanotum, but near the dorsal propodeal margin in lateral view (Figs 2C, 5C); they are posteriorly to posterolaterally directed and protected anteriorly by the anterior flange of the propodeal spiracles. The propodeal lobes are apparently not developed. Legs: The legs are developed as expected for the Formicidae, with some notable characters and states. They are densely hairy, appearing shaggy, with the setae suberect (Fig. 4A, D, G–J). Apparently, the apical protibial foramina are open, i.e. without a bridge of sclerite dividing the calcar from the probasitarsus (Fig. 5G) [Note 9]. The mesoprefemora and metaprefemora are well developed and are broader ventrally than dorsally (Fig. 5D). The protibia bears and anterior brush of dense suberect setae in their apical third, near the calcar (Fig. 4A). Each of the mesotibia and metatibia bear a pair of apicoventral spurs (Figs 4I, H, 5H); the anterior tibial spurs are pectinate; the posterior tibial spurs are barbirulate to simple. The calcar is apparently bifid apically, with one point being the apex of the elongate velum and the other point a small array of hairs (Fig. 5G); two stout setae are developed (inserted) posterior to the calcar. The plantar lobes of the tarsi are not developed (absent), but the tarsomeres have a brush of dense ventral setae, and are apically margined by thick, coarse chaetae (Figs 4G–J, 5G–J) [Note 10]. The fourth tarsomeres of each leg are deeply notched distally, thus appearing V-shaped (probasitarsi) or arrowhead-shap, Published as part of Boudinot, Brendon E., Richter, Adrian, Katzke, Julian, Chaul, Júlio C. M., Keller, Roberto A., Economo, Evan P., Beutel, Rolf Georg & Yamamoto, Shûhei, 2022, Evidence for the evolution of eusociality in stem ants and a systematic revision of † Gerontoformica (Hymenoptera: Formicidae), pp. 1355-1389 in Zoological Journal of the Linnean Society 195 on pages 1364-1369, DOI: 10.1093/zoolinnean/zlab097, http://zenodo.org/record/6994456, {"references":["Boudinot BE, Khouri Z, Richter A, Van de Kamp T, Barden P, Perrichot V, Chaul JCM. 2020 a. Chapter II. The evolution of the ants: extinct ant sister-group illuminates eusociality origin and post-K / Pg persistence. In: Boudinot BE, Systematic and evolutionary morphology: case studies on Formicidae, Mesozoic Aculeata, and hexapodan genitalia. Ph. D. Thesis, University of California, Davis, 174 - 393.","Perrichot V, Nel A, Neraudeau D, Lacau S, Guyot T. 2008. New fossil ants in French Cretaceous amber (Hymenoptera: Formicidae). Naturwissenschaften 95: 91 - 97.","Bolton B. 2003. Synopsis and classification of Formicidae. Memoirs of the American Entomological Institute 71: 1 - 370.","Beutel RG, Gorb SN. 2001. Ultrastructure of attachment specializations of hexapods (Arthropoda): evolutionary patterns inferred from a revised ordinal phylogeny. Journal of Zoological Systematics & Evolution Research 39: 177 - 207.","Boudinot BE, Beutel RG, Gorb SN, Polilov AA. 2021 a. Functional diversity of attachment and grooming structures is retained in all but the smallest insects. Journal of Zoology 313: 99 - 113.","Wohrl T, Richter A, Guo S, Reinhardt L, Nowotny M, Blickhan R. 2021. Comparative analysis of a geometric and an adhesive righting strategy against toppling in inclined hexapodal locomotion. Journal of Experimental Biology 224: jeb 242677.","Bolton B. 1994. Identification guide to the ant genera of the world. Cambridge: Harvard University Press, 222."]}

Genus Metalasius gen.nov. ZooBank LSID. urn:lsid:zoobank.org:act:AC5A1489-C805- 4035-A1C0-3DDBC6E1206D. Type species. Lasius myrmidon Mei, 1998: 177, figs 1���11 (w.). Original designation. Species included. M. myrmidon (Mei, 1998) comb.nov., ��� M. pumilus (Mayr, 1868) comb.nov. Total clade definition (worker). 1. With characters of Lasiini. 2. Mandible with 6���8 teeth. 3. Palp formula 6,4. 4. Basal and masticatory mandibular margins meeting at a weakly oblique angle. 5. Clypeus modified for reception of labrum (specifically, clypeus with anterolateral notches; Note 1). 6. Frontalcarinae conspicuous, > 0.5 �� anteroposteriorantennal torulus diameter. 7. Frontal region of head, between antennal toruli strongly bulging, frontalcarinae raised above toruli (Note 2). 8. Antenna 12-merous. 9. Third antennomere broader than long (Note 3). 10. Compound eyes set at about head midlength. 11. Ocelli absent. 12. Dorsum of head completely without standing setae or with only a few small setae around each posterolateral head corner (Note 4). 13. Mesonotum, metanotum and/or propodeum discontinuous, metanotum undifferentiated medially. 14. Mesopleural anterodorsal margin, near posterolateral region of pronotum, inconspicuously bulging, this weak shoulder a very narrow groove which traverses the mesosternum. 15. Metapleural gland very small, orifice directed almost completely posteriorly (Note 5). 16. Propodeal spiracle situated in lower 2/3 of propodeum. 17. Legs entirely or almost entirely devoid of standing setae. 18. Petiole with raised squamiform node, without posterior elongation. 19. Abdominal tergum III vertical, without deep groove for receiving petiole; not concealing petiole when gaster raised (Note 6). 20. Tergosternal suture of abdominal segment III not raised dorsally before unfusing near spiracle, rather anterior margin of abdominal sternum III directed anteriorly away from helcium before narrowly curving posteriorly (Note 6). 21. Pubescence of abdominal terga VI and VII linear. Notes on definition: Note 1. Modification of the anterior clypeal margin for reception of the labrum is here interpreted as a synapomorphy of Metalasius plus the core Prenolepis genus group. The modification is most easily observed in anterior or anterodorsal view. In most Formicinae, the perceived anterior clypeal margin with the head in full-face view is usually a carina which runs across the clypeus from the lateral clypeal margins.This carina may be raised or otherwise modified; regardless, the carina itself or the anterior region of the clypeus is produced anteriorly, concealing the clypeolabral articulation. In the Prenolepis and Metalasius genus groups, the clypeolabral articulation is usually exposed or nearly exposed laterally by notches in the anterior clypeal carina, although it may be exposed medially where the anterior clypeal carina is absent. Note 2. See Note 1 for the Lasius genus group. Note 3. Surprisingly, this distinguishes members of the Metalasius genus group from most formicine genera, with the exception of some Myrmelachista, various plagiolepidines and the Lasius flavus group. Cladomyrma species have a cone-shaped third antennomere, which is usually longer than the diameter of its base (and often apex), except in C. dianeae Agosti et al. for which the length and basal diameter are subequal. Some Pseudolasius and Paraparatrechina approach having subequal diameter and length. Note 4. Mei (1998), in his diagnosis of Metalasius myrmidon, indicated that these standing setae were absent except for one specimen. Note 5. This state may be a synapomorphy for the clade consisting of Metalasius plus the Prenolepis genus group. It was not possible to evaluate this with confidence for ��� M. pumilus, but from the dorsal view of specimen SMFBE1226 on AntWeb, the gland does appear to be small. Note 6. These characters could not be evaluated with confidence for ��� M. pumilus with the available material or descriptions. Etymology. A combination of the Greek ���meta-��� (with, across, after) and ���l��sios��� (hairy), in reference to the former placement of the type species. Homophonously forming ���metal-Asius���, invoking the death of the Trojan leader Asius during the assault on the Archaean wall of Troy. Masculine. Comments. Although, we could not evaluate all of these characters for ��� M. pumilus, this species is consistently recovered as sister to M. myrmidon in combined analysis (Figs 4, S 7���S 9). Metalasius is most easily differentiated from both the Lasius and Prenolepis genus groups by the combination of mid-set eyes, the posteriorly directed and reduced metapleural gland orifice, raised frontal carinae and almost complete lack of standing setae on the head. In addition, both included species of Metalasius have broad third antennomeres and unusually small bodies relative to Lasius, being < 2 mm in total length (Mei, 1998; Dlussky, 2011)., Published as part of Boudinot, Brendon E., Borowiec, Marek L. & Prebus, Matthew M., 2022, Phylogeny, evolution, and classification of the ant genus Lasius, the tribe Lasiini and the subfamily Formicinae (Hymenoptera: Formicidae), pp. 113-151 in Systematic Entomology 47 on pages 140-141, DOI: 10.1111/syen.12522, http://zenodo.org/record/5975346, {"references":["Mei, M. (1998) Lasius (Cautolasius) myrmidon n. sp.: a new hypogaeic ant from Greece (Hymenoptera Formicidae). Bollettino della Societa Entomologica Italiana, 130, 177 - 182.","Mayr, G. (1868) Die Ameisen des baltischen Bernsteins. Beitrage zur Naturkunde Preussens, 1, 1 - 102.","Dlussky, G. M. (2011) The ants of the genus Lasius (hymenoptera, Formicidae) from late Eocene European ambers. Vestnik Zoologii, 45, 209 - 222."]}

Tribe Lasiini Ashmead, 1905 (Figs 6, 7, 8A���K, 9A���J, 10A���D) Type genus. Lasius. Included genus groups. Cladomyrma, Lasius group, Metalasius, Prenolepis group. Definition (worker). 1. With characters of Formicinae (see Bolton, 2003 and Note 1). 2. Mandible triangular, with 4���11 teeth; third tooth from apex reduced. 3. Dorsal mandibular groove, when present, running along lateral mandibular margin as seen in dorsal view (Note 2). 4. Palp formula usually 6,4, less often 3,4. 5. Frontal protuberance mediad antennal toruli with frontal carinae effaced, becoming broadly rounded posteriorly until continuous with face (Note 3). 6. Frontal protuberance raised above toruli or not. 7. Antennal toruli near or abutting posterior clypeal margin. 8. Antenna 12-, 11-, or 8-merous. 9. Compound eyes not enormous, that is, not longer than half the length of the head; eyes may be reduced to absent; long axis of eyes subparallel. 10. Ocelli present or absent (Note 4). 11. Metanotum differentiated or not. 12. Metapleural gland present, dorsal rim of metapleural gland curved inward. 13. Propodeal spiracle at or near posterolateral margin of propodeum. 14. Propodeal spiracle circular to elliptical, not slit-shaped. 15. Metacoxae wideset: Distance between mesocoxal bases less than between that between bases of metacoxae with mesosoma in ventral view and coxae oriented at right-angles to long axis of body. 16. Metatibiae without double row of ventral (inner) setae. 17. Petiolar foramen inprofile view low, not to barelyexceeding dorsal margin of metapleural gland, with or without dorsal margin or lip, but lip, when present, inconspicuous (Note 5). 18. Petiolar foramen in ventral view long, anterior margin exceeding metacoxal cavities anteriorly. 19. Petiolar node conspicuously shorter than propodeum, not reaching dorsal surface of propodeum (Note 6). 20. Petiolar apodeme (situated anteriorly on tergum) contiguous or nearly contiguous with petiolar node (Note 7). 21. Petiolar sternum U-shaped in cross-section (Note 8). 22. Abdominal segment III transverse sulcus absent. 23. Base of abdominal segment III with or without complete tergosternal fusion lateral to helcium, free sclerites commencing distantly up segment or near helcium; tergum III overhanging petiole or not. 24. Proventriculus sepalous (Note 9). Notes on definition: Note 1. The generic composition of the Lasiini has recently been reassessed following Blaimer et al. (2015) and Ward et al. (2016). The tribal definition proposed here is thus revised relative to Bolton (2003). It includes new characters and recognizes that several former plagiolepidines belong in the Lasiini. Note 2. Previous studies have not focused attention on the dorsal mandibular groove. Here it was observed that the groove, as seen with the mandible in dorsal view, runs along the outer margin of the mandible toward the mandibular apex in the Myrmelachistini, Lasiini and most Melophorini. This contrasts with the state observed in the ���formicoform radiation��� (i.e., the clade that includes Camponotini, Formicini and Melophorini, along with the monotypic tribes Gesomyrmecini, Gigantiopini, Oecophyllini, and Santschiellini; see the section ��� Kyromyrma, an ancestral formicine, under the section ��� Incertae sedis in the Formicinae��� below). When the groove is present in the ���formicoform radiation���, it is very close to the basal mandibular margin and is shortened, except in various plagiolepidines. The medial and shortened state may have arisen multiple times in the Formicini, Plagiolepidini, and Camponotini clade. Note 3. The ���frontal protuberance��� is the region between the antennal toruli, which is raised relative to the regions of the face laterad the toruli. To some degree, development of this condition accounts for the ���laterally directed��� torular condition of Formicidae (Boudinot et al., 2020). In other formicine tribes, including the Melophorini (with Prolasius as an exception), the frontal protuberance is carinate above the toruli (i. e., ���frontal carinae are present���), with these carinae continuing as a sharp ridge until their posterior terminus. Within the Lasiini, the frontal carinae are effaced; they are long in all groups except Cladomyrma, while in the Plagiolepidini the carinae are short in all genera except Anoplolepis. Note 4. Ocellus presence���absence is arelatively weak character as expression of ocelli may be variable for genera in which ocelli are observed, such as Lasius and Nylanderia. However, this statement is included as ocellus expression is a traditional character which is easy to evaluate and may have value for future works defining tribes wherein ocelli may be consistently present or absent. Within the Lasiini, ocelli are always absent in Cladomyrma, Euprenolepis Emery, and Pseudolasius Emery, variably present among Lasius, Paraparatrechina Donisthorpe, and Prenolepis Mayr species, and consistently present in Myrmecocystus, Paratrechina Motschoulsky (except P. kohli; see comments under Prenolepis genus group), and Zatania LaPolla et al. Note 5. The conformation of the dorsal region of the petiolar foramen is newly described here. There is complex variation of the form across the subfamily, but it appears at least that the form observed in the Lasiini is consistent; this form is also observed in the Myrmelachistini. In various lineages within the formicoform radiation, a raised and conspicuously carinate dorsal margin is observed. Note 6. Short petiolar nodes are also observed in the Myrmelachistini and Plagiolepidini (excluding Anoplolepis). The node height is variable in the Melophorini, being short in Prolasius and Myrmecorhynchus Andr��. Note 7. The apodeme is clearly separated from the node in most Melophorini, except Prolasius. Note 8. A U-shaped petiolar sternum was used by Bolton (2003) to diagnose the lasiine tribe group. This trait also occurs in the Myrmelachistini, Myrmoteratini Emery, Plagiolepidini, and four Melophorini (Lasiophanes, Prolasius, Stigmacros, Teratomyrmex); see ���Ancestral state estimation��� results above and Figure S15. Note 9. The proventriculus is weakly sepalous in Cladomyrma. The form of the proventriculus forms a natural division between the Plagiolepidini and the ���plagiolepidiform��� Prenolepis genus group, as observed by Emery (1925). Due to uncertainty about the polarity of the sepalous condition ��� and whether this form arose multiple times (Eisner, 1957; Agosti, 1990, 1991) ��� Bolton (2003) lumped the two plagiolepidiform clades, plus the Myrmelachistini, into the Plagiolepidini., Published as part of Boudinot, Brendon E., Borowiec, Marek L. & Prebus, Matthew M., 2022, Phylogeny, evolution, and classification of the ant genus Lasius, the tribe Lasiini and the subfamily Formicinae (Hymenoptera: Formicidae), pp. 113-151 in Systematic Entomology 47 on pages 128-132, DOI: 10.1111/syen.12522, http://zenodo.org/record/5975346, {"references":["Ashmead, W. H. (1905) A skeleton of a new arrangement of the families, subfamilies, tribes and genera of the ants, or the superfamily Formicoidea. The Canadian Entomologist, 37, 381 - 384.","Bolton, B. (2003) Synopsis and classification of Formicidae. Memoirs of the American Entomological Institute, 71, 1 - 370.","Blaimer, B. B., Brady, S. G., Schultz, T. R., Lloyd, M. W., Fisher, B. L. & Ward, P. S. (2015) Phylogenomic methods outperform traditional multi-locus approaches in resolving deep evolutionary history: a case study of formicine ants. BMC Evolutionary Biology, 15, 257.","Ward, P. S., Blaimer, B. B. & Fisher, B. L. (2016) A revised phylogenetic classification of the ant subfamily Formicinae (Hymenoptera: Formicidae), with resurrection of the genera Colobopsis and Dinoponera. Zootaxa, 4072, 343 - 357. https: // doi. org / 10.11646 / Zootaxa. 4072.3.4.","Boudinot, B. E., Perrichot, V. & Chaul, J. C. M. (2020) † Camelosphecia gen. nov., lost ant-wasp intermediates from the mid-Cretaceous (Hymenoptera, Formicidae). ZooKeys, 1005, 21 - 55.","Emery, C. (1925) Hymenoptera. Fam. Formicidae. Subfam. Formicinae. Genera Insectorum, 183, 1 - 302.","Eisner, T. (1957) A comparative morphological study of the proventriculus of ants (Hymenoptera: Formicidae). Bulletin of the Museum of Comparative Zoology, 8, 439 - 490.","Agosti, D. (1990) What makes the Formicini the Formicini? Actes Colloques Insectes Sociaux, 6, 295 - 303.","Agosti, D. (1991) Revision of the oriental ant genus Cladomyrma, with an outline of the higher classification of the Formicinae (Hymenoptera: Formicidae). Systematic Entomology, 16, 293 - 310."]}

Species ��� Metalasius pumilus (Mayr, 1868) comb.nov. Definition (worker). 1. With character states of Metalasius (Note 1). 2. Maxillary palps long, reaching occipital foramen (Note 2). 3. Compound eyes well-developed, with > 20 ommatidia. 4. Mesosomal dorsum devoid of setae. 5. Legs entirely devoid of standing setae. 6. Petiolar node weakly inclined anteriorly, node squamiform. Notes on definition: Note 1. Several characters could not be evaluated for ��� M. pumilus, including the ventromedial mandibular groove, palpomere proportions, clypeal profile, and lateral hypostomal carina. Note 2. From Dlussky (2011). Comments. We place ��� M. pumilus and M. myrmidon together based on the results of our phylogenetic analyses, and we interpret presence of the broad third antennomere and highly reduced cranial setation as synapomorphies of this clade. ��� Metalasius pumilus differs from M. myrmidon by the following: (i) compound eyes large; (ii) maxillary palps long, reaching occipital foramen, and (iii) standing setae completely absent from head dorsum and mesosoma. Although, we have not examined the neotype of ��� M. pumilus, designated by Dlussky (2011) and deposited in Muzeum Ziemi Polskiej Akademii Nauk in Warsaw, the specimen we have studied is unlikely to be misidentified as it exhibits unique diagnostic traits of the species among the Lasiini, let alone of the Baltic amber fauna, including absence of setae on the head dorsum, well-developed eyes, short and broad third antennomere, and very small body size (< 2 mm). Dlussky (2011) describes the eyes of both ��� L. schiefferdeckeri and ��� M. pumilus as ���shifted somewhat posteriorly so that the length of [the] gena [is] more than [that of the] maximum diameter of [the] eyes���. This very general statement is true of both species, however, the eyes of ��� L. schiefferdeckeri are distinctly set in the posterior head half whereas those of ��� M. pumilus and M. myrmidon are situated at head midlength, distinguishing them from all Lasius genus group members., Published as part of Boudinot, Brendon E., Borowiec, Marek L. & Prebus, Matthew M., 2022, Phylogeny, evolution, and classification of the ant genus Lasius, the tribe Lasiini and the subfamily Formicinae (Hymenoptera: Formicidae), pp. 113-151 in Systematic Entomology 47 on page 141, DOI: 10.1111/syen.12522, http://zenodo.org/record/5975346, {"references":["Mayr, G. (1868) Die Ameisen des baltischen Bernsteins. Beitrage zur Naturkunde Preussens, 1, 1 - 102.","Dlussky, G. M. (2011) The ants of the genus Lasius (hymenoptera, Formicidae) from late Eocene European ambers. Vestnik Zoologii, 45, 209 - 222."]}

Genus ��� Sussudio gen.nov. Type and included species. ��� P. boreus Wheeler, 1915. Original designation. ZooBank LSID. http://zoobank.org/urn:lsid:zoobank.org:act: BFF515E7-9966-443F-8A34-555A4B41906B Definition (worker). 1. With characters of the Formicinae (Note 1). 2. Cranium size variable, width somewhat broader than to about twice that of mesosoma (Note 2). 3. Mandible triangular, with 7���8 teeth. 4. Dorsal mandibular groove extending along lateral mandibular margin in dorsal view. 5. Palps reduced, not reaching midlength of postgenal bridge. 6. Frontal carinae raised above antennal toruli. 7. Antennal toruli abutting posterior clypeal margin. 8. Antennomere count unknown (Note 3). 9. Antennomere III longer than broad. 10. Compound eyes situated posterior to head midlength. 11. Compound eyes not enormous and long axes subparallel. 12. Ocelli minutely present or absent. 13. Promesonotum domed, raised well above propodeum. 14. Metanotal groove well-developed. 15. Petiolar foramen in profile view clearly raised and margined with conspicuous thickened carina or lip. 16. Petiolar node squamiform, tall; dorsoventral height equal to or possibly exceeding dorsum of propodeum. 17. Abdominal segment III without raised tergosternal fusion. Notes on definition: Note 1. Several characters were not possible to evaluate. These include the following: Reduction of third tooth from mandibular apex, palp formula, the metapleural gland, ventral length of the petiolar foramen, relative separation of the meso- and metacoxae, presence or absence of setal double-row on ventral tibial surfaces, petiolar apodeme conformation, cross-sectional shape of the petiolar sternum, and presence or absence of the posthelcial sulcus. Note 2. The massive head of ��� P. boreus led Wheeler (1915) to erroneously assume that the specimens were majors of Pseudolasius; Wheeler did note the posteriorly set eyes of ��� P. boreus, one of the distinctions between it and Pseudolasius, but vacillated based on the insufficient knowledge of the latter genus at that time. Note 3. The examined specimens are missing their terminal antennomeres, and no previous descriptions of ��� P. boreus include an antennomere count. Etymology. Asimplification of ���pseudo- Pseudolasius ��� inspired by the musician Phil Collins. Masculine. Comments. ��� Sussudio boreus comb.nov. was previously placed in the Lasiini (Emery, 1925, Dlussky & Fedoseeva 1988, Bolton, 1994), and has been considered to be a member of the genus Pseudolasius (Wheeler, 1915; LaPolla & Dlussky, 2010). Based on an examination of a syntype (GZG-BST04646) and a nontype (NHMW1984-31-211) worker, as well as the original description from Wheeler (1915), we reject both placements. Neither specimen can be mistaken due to the unique combination of characters they display, including setation, high petiolar nodes, massive crania, and mesosomal form. ��� Sussudio boreus does not display the modifications of the third abdominal segment characteristic of the Prenolepis genus group, also rejecting its original placement in Pseudolasius (see diagnostic Note 2 above). ��� Sussudio is excluded from the Lasiini overall by two additional characters: (i) The petiolar node, which is completely vertical and very tall dorsoventrally, is as tall as or possibly taller than the propodeum; and (ii) the raised and conspicuously lipped petiolar foramen. No lasiine, extant or extinct, has such a tall petiolar node, nor do any species display the petiolar foramen conformation observed in ��� Sussudio. We cannot confidently place ��� Sussudio in any extant tribe, therefore, we consider ��� Sussudio incertae sedis in the Formicinae. As with other fossil taxa, which are of uncertain placement in the Formicinae, micro-CT scanning could be used to evaluate the form of the helcium and third abdominal sternum., Published as part of Boudinot, Brendon E., Borowiec, Marek L. & Prebus, Matthew M., 2022, Phylogeny, evolution, and classification of the ant genus Lasius, the tribe Lasiini and the subfamily Formicinae (Hymenoptera: Formicidae), pp. 113-151 in Systematic Entomology 47 on pages 142-143, DOI: 10.1111/syen.12522, http://zenodo.org/record/5975346, {"references":["Wheeler, W. M. (1915 1914) The ants of the Baltic Amber. Schriften der Physikalisch-Okonomischen Gesellschaft zu Konigsberg, 55, 1 - 142.","Emery, C. (1925) Hymenoptera. Fam. Formicidae. Subfam. Formicinae. Genera Insectorum, 183, 1 - 302.","Dlussky, G. M. & Fedoseeva, E. B. (1988) Origin and early stages of evolution in ants [in Russian]. Pp. 70 - 144 in: Ponomarenko, A. G. (ed.) Cretaceous Biocenotic Crisis and Insect Evolution. [in Russian.]. Moskva: Nauka, 232 pp.","Bolton, B. (1994) Identification Guide to the Ant Genera of the World, p. 222. Harvard University Press, Cambridge.","LaPolla, J. S. & Dlussky, G. M. (2010) Review of fossil Prenolepis genus-group species (Hymenoptera: Formicidae). Proceedings of the Entomological Society of Washington, 112, 258 - 273."]}

Species Metalasius myrmidon (Mei, 1998) comb.nov. Definition (worker). 1. With character states of Metalasius. 2. Dorsal mandibular groove absent (Note 1). 3. Ventromedial base of mandible without trough or impression (Note 2). 4. Maxillary palps short, exceeding hypostomal margin but not reaching postgenal bridge midlength. 5. Maxillary palpomere 3 longest, 4 shorter but as long as 5 and 6 together (Note 3). 6. Clypeus, in lateral view, convex and weakly bulging anteriorly (Note 3). 7. Anterior tentorial pit situated lateral to midlength of the epistomal suture. 8. Lateral hypostomal carina absent (Note 4). 9. Compound eyes absent, reduced or vestigial, with at most 9 ommatidia (Note 5). 10. Propodeal spiracle situated distinctly in lower half of propodeum (Note 6). 11. Legs almost entirely devoid of standing setae (Note 3). 12. Petiolar node weakly inclined anteriorly, node well-developed, squamiform (Note 7). Notes on definition: Note 1. The dorsal mandibular groove is discernable in all examined Lasius and Prenolepis genus group taxa. Note 2. Presence of a trough on the ventromedial base of the mandible is a newly detected synapomorphy of the core Prenolepis genus group. The impression is enhanced when the ventromedial mandibular margin is carinate and/or produced medially, and best seen when the mandibles are open and with the head in lateral anteroventral view. The trough may be reduced or absent in some species. Note 3. Previously included in the original diagnosis of the species M. myrmidon by Mei (1998). Note 4. The lateral hypostoma is usually delimited by a carina, which is discontinuous with the medial hypostomal lamina. Among formicines, the lateral hypostomal carina is absent only in Acropyga (some species) and Brachymyrmex, both genera outside of Lasiini. Note 5. The specimens, which were available for examination had 5���8 ommatidia; the maximum ommatidium count is from Mei (1998, p. 178). Note 6. Alowered propodeal spiracle appears sporadically replicated in only a few Prenolepis genus group members. Note 7. The petiolar node is strongly inclined anteriorly in the core Prenolepis genus group. Comments. Metalasius myrmidon is uniquely identified among the Lasiini by absence of the dorsal mandibular groove and lateral hypostomal carina, short and broad third antennomere, mid-set compound eyes (when present), which are reduced to at most 9 ommatidia and near complete absence of standing setae on the head. High magnification may be required to evaluate the lateral hypostoma., Published as part of Boudinot, Brendon E., Borowiec, Marek L. & Prebus, Matthew M., 2022, Phylogeny, evolution, and classification of the ant genus Lasius, the tribe Lasiini and the subfamily Formicinae (Hymenoptera: Formicidae), pp. 113-151 in Systematic Entomology 47 on page 141, DOI: 10.1111/syen.12522, http://zenodo.org/record/5975346, {"references":["Mei, M. (1998) Lasius (Cautolasius) myrmidon n. sp.: a new hypogaeic ant from Greece (Hymenoptera Formicidae). Bollettino della Societa Entomologica Italiana, 130, 177 - 182."]}

Genus Lasius Fabricius, 1804 = Donisthorpea Morice and Durrant, 1915. = Acanthomyops Mayr, 1862 syn.rev. = Austrolasius Faber, 1967 syn.nov. = Cautolasius Wilson, 1955 syn.nov. = Chthonolasius Ruzsky, 1912 syn.nov. = Dendrolasius Ruzsky, 1912 syn.nov. Type species. Formica nigra Linnaeus, 1758 (= L. niger). Subgeneric classification remarks. The modern body of taxonomic work on Lasius was initiated by Wilson���s revision of the genus (Wilson, 1955), which was classified into four subgenera at the time: Cautolasius, Chthonolasius, Dendrolasius, and Lasius s. str. In this work, Wilson provided a phylogeny of Lasius (Fig. 1A), but this treatment was an intuitive account of what he considered trends in the evolution of the morphology and biogeography of the genus. Subsequently, the subgenus Austrolasius was erected for a few socially parasitic species (Faber, 1967), andthe former genus Acanthomyops was included in Lasius as the sixth subgenus (Ward, 2005). The first phylogenetic inference based on molecular data was presented by Hasegawa (1998), who used COI to investigate relationships of four Lasius species (phylogeny not figured here). Since then, two major attempts at resolving the phylogeny were presented by Janda et al. (2004) and Maruyama et al. (2008) (Fig. 1B, C). Both studies used a combination of morphological characters and molecular data, including mitochondrial markers (COI, COII, tRNA-Leu, 16S). Amore recent effort focused on the phylogeny of European species related to L. niger (Lasius s. str.) and included nuclear genes LW Rh and wg in addition to 16S and COI (Talavera et al., 2015). To date, the monophyly of the subgenera has not been questioned except for the nominotypical subgenus (Janda etal., 2004; Maruyama etal., 2008). Recently, Seifert (2020, p. 21) stated that there is clear justification for elevating the subgenera to generic status. Such an action would considerably complicate the classification of the Lasius genus group because of the robustly supported paraphyly of the subgenera that we have uncovered here (Figs 1D, 2, S 1, Table 4). Specifically, in order to retain monophyly at the genus rank, four new genera would need to be erected for the species groups of brunneus, nearcticus, atopus, and pallitarsis. An additional issue would be the placement of the species which have not been sequenced, particularly those of the niger species group and, for example, the recently described L. brevipalpus Seifert, which is incertae sedis in the niger clade. The strongest morphological reorganization at the generic or subgeneric levels would be to recognize the reciprocally monophyletic niger and flavus clades as Lasius and Acanthomyops, respectively, but we refrain from doing so here (for our rationale, see ���Species group classification of Lasius ��� below). Comments on extant species. We determine that one species, Lasius escamole Reza should be excluded from Lasius and considered a junior synonym of the dolichoderine Liometopum apiculatum Mayr syn.nov. Reza (1925) described L. escamole in the context of a cultural study on the eponymous traditional Mexican dish known tobe made from the larvae of L. apiculatum (Hoey-Chamberlain et al., 2013). Although Reza���s description and illustrations are extremely vague, it is possible to see details that point to a dolichoderine identity. In the figures of the original description, the mandibles have long masticatory margin and small, sharp, even denticles, the ventral metasoma is shown as a slit-like anal opening rather than a formicine-like acidopore, and various figures display the fine, dense, appressed pilosity characteristic of Liometopum Mayr, but no erect setae as expected for Lasius. Many species have been added to Lasius since Wilson���s revision, mostly in Europe and Mediterranean, while North American taxa have largely remained untreated except for a thorough revision of Acanthomyops (Wing, 1968a, 1968b). Careful research has revealed multiple Palaearctic Lasius species that show only subtle morphological differentiation from close relatives (Seifert, 1983, 1990, 1991, 1992, 2020; Schlick-Steiner et al., 2003). There is no reason to believe that North America does not harbour a diverse fauna of such ���cryptic species���. For example, the question of the putative Holarctically distributed Lasius species was resolved by Sch��r et al. (2018) who elevated to species rank the American representatives of L. alienus, L. flavus, and L. umbratus Nylander, recognizing the following revived taxa, in order: L. americanus Emery, L. brevicornis Emery, and L. aphidicola (Walsh). Renewed focus on the Nearctic fauna is necessary, as is expanded sequencing at the global scale. Comments on extinct species. Without having scored the Baltic Lasius fossils other than ��� L. schiefferdeckeri, that is, ��� L. punctulatus Mayr and ��� L. nemorivagus Wheeler, we are unable to quantitatively address their placement. Historically, ��� L. punctulatus and ��� L. schiefferdeckeri were considered to be members of Lasius s. str. (Wilson, 1955; Dlussky, 2011), while the queen-based ��� L. nemorivagus was placed in Chthonolasius (Wilson, 1955) later to be implicitly considered incertae sedis in the genus (Dlussky, 2011). Our combined-evidence dating analyses recover ��� L. schiefferdeckeri as sister to or within the Lasius genus group (Figs 4, S 7���S 9). As the specific relationship of the fossil to the extant species of the Lasius genus group is uncertain, we conservatively consider the fossil incertae sedis in Lasius. There remains the possibility that ��� L. schiefferdeckeri is ancestral to the extant niger clade and is indicative of low rates of phenotypic transformation, as suggested by Mayr (1868), Wheeler (1915), and Wilson (1955). The placement of ��� L. schiefferdeckeri may be refined in future study by scoring characters which are explicitly derived from comparison of the brunneus and niger groups within the niger clade. The two differentiating traits proposed for the brunneus and niger groups are, on average, < 8 mandibular teeth in the brunneus group (Seifert, 1992; some niger group species with < 8), and presence of a subapical cleft in the mandibles of brunneus group males (Wilson, 1955). While ��� L. schiefferdeckeri demonstrates both a tooth count of < 8, and presence of a subapical cleft, the latter character is probably plesiomorphic of the Lasius genus group, and the polarity of the former is uncertain. Notably, Wilson (1955) observed that the male mandibles of ��� L. schiefferdeckeri are observed to vary from the ��� brunneus form��� to the derived ��� niger form���. With these three traits in mind, it does seem reasonable that ��� L. schiefferdeckeri is stem to or directly ancestral to the niger clade. Note on biology. Despite the interest in this genus, however, basic natural history remains unknown for many species, including the morphologically aberrant L. atopus and the species we sequence here, ��� L. nr. atopus ���. Only a handful of recently published studies have addressed the behaviour of some of the more rarely encountered species (e.g., Raczkowski & Luque, 2011), indicating that more effort is needed to elucidate the biology of Lasius., Published as part of Boudinot, Brendon E., Borowiec, Marek L. & Prebus, Matthew M., 2022, Phylogeny, evolution, and classification of the ant genus Lasius, the tribe Lasiini and the subfamily Formicinae (Hymenoptera: Formicidae), pp. 113-151 in Systematic Entomology 47 on pages 135-136, DOI: 10.1111/syen.12522, http://zenodo.org/record/5975346, {"references":["Fabricius, J. C. (1804) Systema Piezatorum Secundum Ordines, Genera, Species, Adjectis Synonymis, Locis, Observationibus, Descriptionibus. Brunswick: C. Reichard. xiv + 15 - 439 + 30 pp.","Faber, W. (1967) Beitrage zur Kenntnis sozialparasitischer Ameisen. I. Lasius (Austrolasius n. sg.) reginae n. sp., eine neue temporar sozialparasitische Erdameise aus Osterreich (Hym. Formicidae). Pflanzenschultz Berichte, 36, 73 - 107.","Wilson, E. O. (1955) A monographic revision of the ant genus Lasius. Bulletin of the Museum of Comparative Zoology, 113, 1 - 201.","Linnaeus, C. (1758) Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Holmiae [= Stockholm]: L. Salvii, 824 pp.","Ward, P. S. (2005) A synoptic review of the ants of California (Hymenoptera: Formicidae). Zootaxa, 936, 1 - 68.","Hasegawa, E. (1998) Phylogeny and host-parasite relationships in social parasitism in Lasius ants. Entomological Science, 1, 133 - 135.","Janda, M., Folkova, D. & Zrzavy, J. (2004) Phylogeny of Lasius ants based on mitochondrial DNA and morphology, and the evolution of social parasitism in the Lasiini (Hymenoptera: Formicidae). Molecular Phylogenetics and Evolution, 33, 595 - 614.","Maruyama, M., Steiner, F. M., Stauffer, C., Akino, T., Crozier, R. H. & Schlick-Steiner, B. C. (2008) A DNA and morphology based phylogenetic framework of the ant genus Lasius with hypotheses for the evolution of social parasitism and fungiculture. BMC Evolutionary Biology, 8, 1 - 15. Article 237. https: // doi. org / 10.1186 / 1471 - 2148 - 8 - 237.","Talavera, G., Espadaler, X. & Vila, R. (2015 2014) Discovered just before extinction? The first endemic ant from the Balearic Islands (Lasius balearicus sp. nov.) is endangered by climate change. Journal of Biogeography, 42, 589 - 601. https: // doi. org / 10.1111 / jbi. 12438.","Seifert, B. (2020) A taxonomic revision of the Palaearctic members of the subgenus Lasius s. str. (Hymenoptera, Formicidae). Soil Organisms, 92, 15 - 86.","Reza, A. (1925) Recursos alimenticios de Mexico de origen animal poco conocidos. Memorias y Revista de la Sociedad Cientifica \" Antonio Alzate \", 44, 1 - 22.","Hoey-Chamberlain, R., Rust, M. K. & Klotz, J. H. (2013) A review of the biology, ecology and behavior of velvety tree ants of North America. Sociobiology, 60, 1 - 10.","Wing, M. W. (1968 a) Taxonomic revision of the Nearctic genus Acanthomyops (Hymenoptera: Formicidae). Memoirs of the Cornell University Agricultural Experiment Station, 405, 1 - 173.","Wing, M. W. (1968 b) A taxonomic revision of the Nearctic genus Acanthomyops Mayr (Hymenoptera: Formicidae). [Abstract.]. Dissertation Abstracts. B. Sciences and Engineering, 28, 3934.","Seifert, B. (1983) The taxonomical and ecological status of Lasius myops Forel (Hymenoptera, Formicidae) and first description of its males. Abhandlungen und Berichte des Naturkundemuseums Gorlitz, 57, 1 - 16.","Seifert, B. (1990) Supplementation to the revision of European species of the ant subgenus Chthonolasius Ruzsky, 1913 (Hymenoptera, Formicidae). Doriana, Supplemento agli Annali del Museo Civico di Storia Naturale \" G. Doria \". GEN, 6, 1 - 13.","Seifert, B. (1991) Lasius platythorax n. sp., a widespread sibling species of Lasius niger (Hymenoptera: Formicidae). Entomologia Generalis, 16, 69 - 81.","Seifert, B. (1992) A taxonomic revision of the Palaearctic members of the ant subgenus Lasius s. str. (Hymenoptera: Formicidae). Abhandlungen und Berichte des Naturkundemuseums Gorlitz, 66, 1 - 67.","Schlick-Steiner, B. C., Steiner, F. M., Schodl, S. & Seifert, B. (2003) Lasius austriacus sp. n., a central European ant related to the invasive species Lasius neglectus. Sociobiology, 41, 725 - 736.","Schar, S., Talavera, G., Espadaler, X., Rana, J. D., Anderson, A. A., Cover, S. P. & Vila, R. (2018) Do Holarctic ant species exist? Trans-Beringian dispersal and homoplasy in the Formicidae. Journal of Biogeography, 45, 1 - 12. https: // doi. org / 10.1111 / jbi. 13380.","Dlussky, G. M. (2011) The ants of the genus Lasius (hymenoptera, Formicidae) from late Eocene European ambers. Vestnik Zoologii, 45, 209 - 222.","Mayr, G. (1868) Die Ameisen des baltischen Bernsteins. Beitrage zur Naturkunde Preussens, 1, 1 - 102.","Wheeler, W. M. (1915 1914) The ants of the Baltic Amber. Schriften der Physikalisch-Okonomischen Gesellschaft zu Konigsberg, 55, 1 - 142.","Raczkowski, J. M. & Luque, G. M. (2011) Colony founding and social parasitism in Lasius (Acanthomyops). Insectes Sociaux, 58, 237 - 244."]}

On the identity of ��� Protrechina Wilson: Wilson (1985) described a genus putatively close to Paratrechina sensu lato (��� Prenolepis genus group) from mid-Eocene Claiborne amber (Arkansas, 40.4���37.2 Ma; Saunders et al., 1974). This genus, ��� Protrechina, supposedly differs from Paratrechina s. l., Lepisiota Santschi and Brachymyrmex Mayr ��� among other, unstated formicines ��� by the absence of standing macrosetae on the mesosomal dorsum, a state similar to that observed for Metalasius gen.nov. as noted below. The genus has been variably treated as a lasiine (Bolton, 1994, 1995; LaPolla & Dlussky, 2010), a ���prenolepidine��� (H��lldobler & Wilson, 1990), or as incertae sedis in the subfamily where it remains at present (Wilson, 1985; Dlussky & Fedoseeva 1988; Bolton, 2003; Ward et al., 2016). The type specimen of ��� Protrechina carpenteri Wilson, at the Museum of Comparative Zoology (Harvard), should be reexamined to evaluate its tribal, and perhaps generic, placement. This would be particularly valuable given the approximately mid-Eocene origin of the Prenolepis genus group here inferred (Fig. 4; Table 5); such a study will be facilitated by use of micro-CT (e.g., Hita-Garcia et al., 2017; Barden et al., 2017b; Boudinot et al., in press), given the poor condition of the specimen reported by LaPolla & Dlussky (2010)., Published as part of Boudinot, Brendon E., Borowiec, Marek L. & Prebus, Matthew M., 2022, Phylogeny, evolution, and classification of the ant genus Lasius, the tribe Lasiini and the subfamily Formicinae (Hymenoptera: Formicidae), pp. 113-151 in Systematic Entomology 47 on page 133, DOI: 10.1111/syen.12522, http://zenodo.org/record/5975346, {"references":["Wilson, E. O. (1985) Ants from the Cretaceous and Eocene amber of North America. Psyche, 92, 205 - 216.","Saunders, W. B., Mapes, R. H., Carpenter, F. M. & Elsik, W. C. (1974) Fossiliferous amber from the Eocene (Claiborne) of the Gulf coastal plain. GSA Bulletin, 85, 979 - 984.","Bolton, B. (1994) Identification Guide to the Ant Genera of the World, p. 222. Harvard University Press, Cambridge.","Bolton, B. (1995) A New General Catalogue of the Ants of the World, p. 504. Harvard University Press, Cambridge.","LaPolla, J. S. & Dlussky, G. M. (2010) Review of fossil Prenolepis genus-group species (Hymenoptera: Formicidae). Proceedings of the Entomological Society of Washington, 112, 258 - 273.","Holldobler, B. & Wilson, E. O. (1990) The Ants. Harvard University Press, Cambridge, Massachusetts. xii + 732 pp.","Dlussky, G. M. & Fedoseeva, E. B. (1988) Origin and early stages of evolution in ants [in Russian]. Pp. 70 - 144 in: Ponomarenko, A. G. (ed.) Cretaceous Biocenotic Crisis and Insect Evolution. [in Russian.]. Moskva: Nauka, 232 pp.","Bolton, B. (2003) Synopsis and classification of Formicidae. Memoirs of the American Entomological Institute, 71, 1 - 370.","Ward, P. S., Blaimer, B. B. & Fisher, B. L. (2016) A revised phylogenetic classification of the ant subfamily Formicinae (Hymenoptera: Formicidae), with resurrection of the genera Colobopsis and Dinoponera. Zootaxa, 4072, 343 - 357. https: // doi. org / 10.11646 / Zootaxa. 4072.3.4.","Hita-Garcia, F., Fischer, G., Liu, C., Audisio, T. L. & Economo, E. P. (2017) Next-generation morphological character discovery and evaluation: an X-ray micro-CT enhanced revision of the ant genus Zasphinctus Wheeler (Hymenoptera: Formicidae: Dorylinae) in the Afrotropics. ZooKeys, 693, 33 - 93.","Barden, P., Herhold, H. W. & Grimaldi, D. A. (2017 b) A new genus of hell ants from the Cretaceous (Hymenoptera: Formicidae: Haidomyrmecini) with a novel head structure. Systematic Entomology, 42, 837 - 846."]}

Exclusion of ��� Glaphyromyrmex from the Lasiini: The Baltic amber formicine, ��� Glaphyromyrmex Wheeler, was placed in the Formicini until recently (Wheeler, 1915, Donisthorpe, 1943, Dlussky, 1967, Dlussky & Fedoseeva 1988, Bolton, 1994), when Dlussky (2008) transferred the genus to the Lasiini. Placement of ��� Glaphyromyrmex in the Lasiini is counterintuitive given presence of a ventral double-row of setae on its metatibiae, a state which does not occur in any lasiine. Intuitive placement of ��� Glaphyromyrmex based on morphology is challenging, however. ��� Glaphyromyrmex differs from members of all formicine tribes in which double seta rows occur (Formicini, Camponotini, some Melophorini). Specifically, ��� Glaphyromyrmex differs from the Formicini and most Camponotini in eye position (eyes set at about head midlength), from the Camponotini in having antennal toruli which abut the posterior clypeal margin (vs widely separated), and from the Melophorini, which have well-defined dorsal and ventral flaps surrounding the metapleural gland, a conformation that is apparently absent in ��� Glaphyromyrmex. Recognizing these differences, however, we transfer ��� Glaphyromyrmex back to the Formicini (tribal transfer) based on our combined analyses (Fig. 4) and recommend revised study of the fine-scale external anatomy of the fossil taxon., Published as part of Boudinot, Brendon E., Borowiec, Marek L. & Prebus, Matthew M., 2022, Phylogeny, evolution, and classification of the ant genus Lasius, the tribe Lasiini and the subfamily Formicinae (Hymenoptera: Formicidae), pp. 113-151 in Systematic Entomology 47 on page 132, DOI: 10.1111/syen.12522, http://zenodo.org/record/5975346, {"references":["Wheeler, W. M. (1915 1914) The ants of the Baltic Amber. Schriften der Physikalisch-Okonomischen Gesellschaft zu Konigsberg, 55, 1 - 142.","Donisthorpe, H. (1943) A list of the type-species of the genera and subgenera of the Formicidae. [part]. Annals and Magazine of Natural History, 10, 617 - 688.","Dlussky, G. M. (1967) Ants of the genus Formica from the Baltic Amber. Paleontological Journal, 1, 69 - 77.","Dlussky, G. M. & Fedoseeva, E. B. (1988) Origin and early stages of evolution in ants [in Russian]. Pp. 70 - 144 in: Ponomarenko, A. G. (ed.) Cretaceous Biocenotic Crisis and Insect Evolution. [in Russian.]. Moskva: Nauka, 232 pp.","Bolton, B. (1994) Identification Guide to the Ant Genera of the World, p. 222. Harvard University Press, Cambridge.","Dlussky, G. M. (2008) Ants of the tribe Formicini (hymenoptera, Formicidae) from late Eocene amber of Europe. Paleontological Journal, 42, 500 - 513."]}

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 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., ��� Paraneuretus and ��� Protaneuretus from Baltic amber (37���42 My; Wheeler 1915; LaPolla et al. 2013), while others are less certain, i.e., ��� Pityomyrmex (also from Baltic amber, Wheeler 1915; placed in Aneuretinae by Dlussky & Rasnisyn 2009) and ��� Aneuretellus (Sakhalin amber, 56���59 My; Dlussky 1988; LaPolla et al. 2013). The impression-fossil taxa ��� Britaneuretus (see Antropov et al. 2014) and ��� 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 ��� Cananeuretus (78���19 My, Canadian amber; Engel & Grimaldi 2005; LaPolla et al. 2013) with Aneuretus simoni are uncertain. The description and illustration of ��� 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 ��� Burmomyrma to the Aneuretinae is weak, especially given that Aneuretus has complete (���ancestral���) wing venation while ��� Burmomyrma lacks almost all vein abscissae. Placement of ��� 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 ��� 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 ��� 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., Published as part of Boudinot, Brendon E., 2015, 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, pp. 1-62 in European Journal of Taxonomy 120 on pages 49-50, DOI: 10.5852/ejt.2015.120, http://zenodo.org/record/3780152, {"references":["LaPolla J. S., Dlussky G. M. & Perrichot V. 2013. Ants and the fossil record. Annual Review of Entomology 58: 609 - 630. http: // dx. doi. org / 10.1146 / annurev-ento- 120710 - 100600","Brady S. G., Schultz T. R., Fisher B. L. & Ward P. S. 2006. Evaluating alternative hypotheses for the early evolution and diversification of ants. Proceedings of the National Academy of Sciences of the United States of America 103: 18172 - 18177. http: // dx. doi. org / 10.1073 / pnas. 0605858103","Dlussky G. M. & Rasnitsyn A. P. 2009. Ants (Insecta: Vespida: Formicidae) in the Upper Eocene amber of central and eastern Europe. Paleontological Journal 43: 1024 - 1042.","Wheeler W. M. 1915 (1914). The ants of the Baltic Amber. Schriften der Physikalisch-Okonomischen Gesellschaft zu Konigsberg 55: 1 - 142.","Antropov A. V., Belokobylskij S. A., Compton S. G., Dlussky G. M., Khalaim A. I., Kolyada V. A., Kozlov M. A., Perfilieva K. S. & Rasnitsyn A. P. 2014. The wasps, bees and ants (Insecta: Vespida = Hymenoptera) from the insect limestone (Late Eocene) of the Isle of Wight, UK. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 104: 335 - 446.","Dlussky G. M. 1996. Ants (Hymenoptera: Formicidae) from Burmese amber. Paleontological Journal 30: 449 - 454.","Engel M. S. & Grimaldi D. A. 2005. Primitive new ants in Cretaceous amber from Myanmar, New Jersey, and Canada (Hymenoptera: Formicidae). American Museum Novitates 3485: 1 - 23.","McKellar R. C., Glasier J. R. N. & Engel M. S. 2013. New ants (Hymenoptera: Formicidae: Dolichoderinae) from Canadian Late Cretaceous amber. Bulletin of Geosciences 88: 583 - 594. http: // dx. doi. org / 10.3140 / bull. geosci. 1425","Ward P. S., Brady S. G., Fisher B. L. & Schultz T. R. 2010. Phylogeny and biogeography of dolichoderine ants: effects of data partitioning and relict taxa on historical inference. Systematic Biology 59: 342 - 362. http: // dx. doi. org / 10.1093 / sysbio / syq 012","Wilson E. O., Eisner T., Wheeler G. C. & Wheeler J. 1956. Aneuretus simoni Emery, a major link in ant evolution. Bulletin of the Museum of Comparative Zoology 115: 81 - 99.","Bolton B. 2003. Synopsis and classification of Formicidae. Memoirs of the American Entomological Institute 71: 1 - 370."]}

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.] 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, CASENT0132466, E. Prado). I. Formica pallidefulva (U.S.A., CASENT0172882, A Nobile). Scale bars: A���F = 0.5 mm, G���I = 1.0 mm. . 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 ��� 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, excluding 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 weakly 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 H��lldobler & Wilson (1990). Other authors demurred, placing the genus in its own tribe (Apomyrmini) in the Ponerinae sensu lato (Dlussky & Fedoseeva 1988) and in the Leptanillinae (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 (P.S. 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., Published as part of Boudinot, Brendon E., 2015, 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, pp. 1-62 in European Journal of Taxonomy 120 on pages 17-22, DOI: 10.5852/ejt.2015.120, http://zenodo.org/record/3780152, {"references":["Forel A. 1893. Sur la classification de la famille des Formicides, avec remarques synonymiques. Annales de la Societe Entomologique de Belgique 37: 161 - 167.","Bolton B. 2003. Synopsis and classification of Formicidae. Memoirs of the American Entomological Institute 71: 1 - 370.","Yoshimura M. & Fisher B. L. 2012 a. A revision of male ants of the Malagasy Amblyoponinae (Hymenoptera: Formicidae) with resurrections of the genera Stigmatomma and Xymmer. PLoS ONE 7: e 3325. http: // dx. doi. org / 10.1371 / journal. pone. 0033325","Brady S. G., Schultz T. R., Fisher B. L. & Ward P. S. 2006. Evaluating alternative hypotheses for the early evolution and diversification of ants. Proceedings of the National Academy of Sciences of the United States of America 103: 18172 - 18177. http: // dx. doi. org / 10.1073 / pnas. 0605858103","Ward P. S. 2014. The phylogeny and evolution of ants. Annual Review of Ecology, Evolution, and Systematics 45: 2.1 - 2.21. http: // dx. doi. org / 10.1146 / annurev-ecolsys- 120213 - 091824","Brown W. L. Jr., Gotwald W. H. Jr. & Levieux J. 1971 [1970]. A new genus of ponerine ants from West Africa (Hymenoptera: Formicidae) with ecological notes. Psyche 77: 259 - 275.","Wheeler G. C. & Wheeler J. 1985. A simplified conspectus of the Formicidae. Transactions of the American Entomological Society 111: 255 - 264.","Holldobler B. & Wilson E. O. 1990. The Ants. Harvard University Press, Cambridge.","Dlussky G. M. & Fedoseeva E. B. 1988. Origin and early stages of evolution in ants. [In Russian] In: Ponomarenko A. G. (ed.) Cretaceous Biocenotic Crisis and Insect Evolution: 70 - 144. Nauka, Moskva.","Bolton B. 1990 b. The higher classification of the ant subfamily Leptanillinae (Hymenoptera: Formicidae). Systematic Entomology 15: 267 - 282. http: // dx. doi. org / 10.1111 / j. 1365 - 3113.1990. tb 00063. x","Kugler C. 1992. Stings of ants of the Leptanillinae (Hymenoptera: Formicidae). Psyche 99: 103 - 115.","Baroni Urbani C., Bolton B. & Ward P. S. 1992. The internal phylogeny of ants (Hymenoptera: Formicidae). Systematic Entomology 17: 301 - 329. http: // dx. doi. org / 10.1111 / j. 1365 - 3113.1992. tb 00553. x","Bolton B. 1994. Identification Guide to the Ant Genera of the World. Harvard University Press, Cambridge.","Saux C., Fisher B. L. & Spicer G. S. 2004. Dracula ant phylogeny as inferred by nuclear 28 S rDNA sequences and implications for ant systematics (Hymenoptera: Formicidae: Amblyoponinae). Molecular Phylogenetics and Evolution 33: 457 - 468. http: // dx. doi. org / 10.1016 / j. ympev. 2004.06.017"]}

Apomyrma stygia Brown, Gotwald Jr. & L��vieux, 1971 Apomyrma stygia Brown, Gotwald Jr. & L��vieux, 1971: 264, figs. 1���19 (worker, queen, male). C��TE D���IVOIRE, Lagunes: Station d���Ecologie de LAMTO, near Toumodi, 6��13���N 5��02���W, 75���120 m elevation, 16 Apr. 1968 (J. L��vieux) [UCDC paratype worker CASENT0260454 examined] [Original label reads: ��� IVORY COAST Lamto, Toumodi, 16.iv.68 J. Levieux���]., Published as part of Boudinot, Brendon E., 2015, 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, pp. 1-62 in European Journal of Taxonomy 120 on page 24, DOI: 10.5852/ejt.2015.120, http://zenodo.org/record/3780152

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 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. 11 A���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 (H��lldobler & 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 (H��lldobler & 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. 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 1). 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 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 weakly 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. Antenna 13-merous; funiculus filiform. 10. Occipital carina reduced, not enclosing occiput. 11. Notauli present, meeting medially and extending posteriorly to transscutal line (Fig. 11 C���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 1p, 1p. 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; 1m-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; posterior foramen raised dorsally 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. 12 D���F). 37. Basimere continuous with telomere (Fig. 12 D���H). 38. Telomere short, digitate, extending anteroventrad beneath basimere (Fig. 12F). 39. Cuspis absent (Fig. 12E). 40. Valviceps dorsomedially fused, lobate, longer than tall, apex downturned, two sides forming tube (Fig. 12D, H). Note 1. The posterior petiolar foramen is convergently raised above the anterior petiolar foramen in Tatuidris (Agroecomyrmecinae)., Published as part of Boudinot, Brendon E., 2015, 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, pp. 1-62 in European Journal of Taxonomy 120 on pages 33-37, DOI: 10.5852/ejt.2015.120, http://zenodo.org/record/3780152, {"references":["Rabeling C., Brown J. M. & Verhaagh M. 2008. Newly discovered sister lineage sheds light on early ant evolution. Proceedings of the National Academy of Sciences B 105: 14913 - 14917. http: // dx. doi. org / 10.1073 / pnas. 0806187105","Bolton B. 2003. Synopsis and classification of Formicidae. Memoirs of the American Entomological Institute 71: 1 - 370.","Moffett M. W. 1986. Mandibles that snap: notes on the ant Mystrium camillae Emery. Biotropica 18: 361 - 362.","Gronenberg W., Holldobler B. & Alpert G. D. 1998. Jaws that snap: control of mandible movements in the ant Mystrium. Journal of Insect Physiology 44: 241 - 253. http: // dx. doi. org / 10.1016 / S 0022 - 1910 (97) 00145 - 5","Holldobler B. & Wilson E. O. 1990. The Ants. Harvard University Press, Cambridge.","Keller R. A. 2011. A phylogenetic analysis of ant morphology (Hymenoptera: Formicidae) with special reference to the poneromorph subfamilies. Bulletin of the American Museum of Natural History 355: 1 - 90.","Billen J., Bauweleers E., Hashim R. & Ito F. 2013. Survey of the exocrine system of Protanilla wallacei (Hymenoptera: Formicidae). Arthropod Structure & Development 42: 173 - 183. http: // dx. doi. org / 10.1016 / j. asd. 2013.01.001"]}

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. INDICES. CI 0.89���0.93, CS 0.34���0.40, SEI 356���427, SI 18.2���19.2, EI 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 weakly 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. 8 B���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. Lower 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. 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, 1m-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. Rsf1 less than half length of Mf1; Rsf1 and Mf1 parallel. Rs+M sinuate (see note on wing venation below). 2r-rs slightly shorter than Mf1, 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 Mf1. CuF slightly divergent with respect to Mf4���6. 1A extending distad 1m-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. 9 C���H). Pygostyles absent. Abdominal sternum IX hexagonal in outline; spiculum somewhat long, about four times longer than broad at base; anterior margins angle distinctly posterolaterally from spiculum; anterolateral margins weakly 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 3/4 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. SCULPTURATION. 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 weakly 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 1m-cu may be apically displaced, producing a second Rs+M abscissa which splits into Rsf2 and Mf2, or 1m-cu 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 (P.S. 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 morphometrics, 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 Leptanillinae, 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., Published as part of Boudinot, Brendon E., 2015, 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, pp. 1-62 in European Journal of Taxonomy 120 on pages 24-29, DOI: 10.5852/ejt.2015.120, http://zenodo.org/record/3780152, {"references":["Brady S. G., Schultz T. R., Fisher B. L. & Ward P. S. 2006. Evaluating alternative hypotheses for the early evolution and diversification of ants. Proceedings of the National Academy of Sciences of the United States of America 103: 18172 - 18177. http: // dx. doi. org / 10.1073 / pnas. 0605858103","Moreau C. S., Bell C. D., Vila R., Archibald S. B. & Pierce N. E. 2006. Phylogeny of the ants: diversification in the age of angiosperms. Science 312: 101 - 104. http: // dx. doi. org / 10.1126 / science. 1124891","Rabeling C., Brown J. M. & Verhaagh M. 2008. Newly discovered sister lineage sheds light on early ant evolution. Proceedings of the National Academy of Sciences B 105: 14913 - 14917. http: // dx. doi. org / 10.1073 / pnas. 0806187105"]}

Subfamily Formicinae Latreille, 1809 Figs 5I, 7B, F, 16 A���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., Published as part of Boudinot, Brendon E., 2015, 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, pp. 1-62 in European Journal of Taxonomy 120 on page 51, DOI: 10.5852/ejt.2015.120, http://zenodo.org/record/3780152

Subfamily Leptanillinae Emery, 1910 Figs 4F, 6F, 10 Leptanillini Emery, 1910: 32. Type-genus: Leptanilla. Note All diagnoses in the references below pertain to the tribe Leptanillini; 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 [Anomalomyrmini]). Male diagnosis Male Leptanillinae 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). 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. Antenna 13-merous; funiculus filiform to submoniliform. 8. Occipital carina absent. 9. Oblique mesopleural sulcus present, anterior terminus contacting posterolateral pronotal corner 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; 1s,2(1s,1p); 1s,2s; 1s,1s; 0,1p. 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 IIIb 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-like 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. 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 Leptanillinae 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 Leptanillinae 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). Rsf1+Mf1 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). Mf2+ absent or spectral and 2rs-m absent (submarginal cell 2 absent). 1m-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 (&female; = queen, &male; = male) Anomalomyrma indet. [&female;: Indonesia]; Leptanilla africana Baroni Urbani [&male;: Nigeria]; L. bifurcata Kugler [&male;: Israel]; L. islamica Baroni Urbani [&male;: Yemen]; L. israelis Kugler [&male;: Israel]; L. miniscula Santschi [&male;: Tunisia]; L. swani Wheeler [&male;: Australia: Queensland, W. Australia]; L. tanit Santschi [&male;: Tunisia]; L. tenuis Santschi [&male;: Tunisia]; L. GR01 [&male;: Greece]; L. GR02 [&male;: Greece]; L. GR03 [&male;: Greece]; L. IL01 [&male;: Israel]; L. TH01 [&male;: Thailand]; L. TH02 [&male;: Thailand]; L. TH03 [&male;: Thailand]; L. TH04 [&male;: Thailand]; L. TH05 [&male;: Thailand]; L. TH06 [&male;: Thailand]; L. TH08 [&male;: Thailand]; L. TH10 [&male;: Thailand]; L. ZA01 [&male;: South Africa]; L. indet. [&male;: Australia]; Noonilla BMNH01 [&male;: Ivory Coast]; Noonilla indet. [&male;: Malaysia]; Phaulomyrma MM01 [&male;: Myanmar]; Ph. TH01 [&male;: Thailand]; Ph. indet. [&male;: Myanmar]; Protanilla TH01 [&male;: Thailand]; Pr. TH02 [&male;: Thailand]; Pr. TH03 [&male;: Thailand]; Pr. indet. [&male;: Indonesia]; Scyphodon cf. bruesi [&male;: Indonesia]; Yavnella argamani Kugler [&male;: Israel]; Y. indica Kugler [&male;: India]; Yavnella BEB001 [&male;: Sri Lanka]; Leptanillinae? Protanilla [&male;: Indonesia, Malaysia]; Leptanillinae indet. [&male;: Borneo, Democratic Republic of Congo, Indonesia, Malaysia, Papua New Guinea, Thailand]. Distribution Old World: Palearctic (Europe, Southern Asia, and Northern Africa), Afrotropical, 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- based taxa Noonilla Petersen, 1968 (stat. rev.) and Scyphodon Brues, 1925 (stat. rev.). Both N. 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 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., Baroni Urbani 1977; Bolton 1990b; Ogata et al. 1995), the Leptanillinae is 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; AntWeb 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 -clade 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., Published as part of Boudinot, Brendon E., 2015, 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, pp. 1-62 in European Journal of Taxonomy 120 on pages 29-33, DOI: 10.5852/ejt.2015.120, http://zenodo.org/record/3780152, {"references":["Emery C. 1910. Hymenoptera. Fam. Formicidae. Subfam. Dorylinae. Genera Insectorum 102: 1 - 34. http: // www. biodiversitylibrary. org / bibliography / 45481 / summary","Wheeler W. M. 1910. Ants: Their Structure, Development and Behavior. Columbia University Press, New York.","Wheeler G. C. & Wheeler E. W. 1930. Two new ants from Java. Psyche 37: 193 - 201.","Morley B. D. W. 1939. The phylogeny of the Cerapachyinae, Dorylinae, and Leptanillinae (Hym. Formicidae). Bulletin de la Societe Entomologique de France 44: 114 - 118.","Kutter H. 1948. Beitrag zur Kenntnis der Leptanillinae Hym. Formicidae. Eine neue Ameisengattung aus Sud-Indien. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 21: 286 - 295.","Bernard F. 1967 [1968]. Faune de l'Europe et du Bassin Mediterraneen. 3. Les fourmis Hymenoptera Formicidae d'Europe occidentale et septentrionale. Masson, Paris.","Petersen B. 1968. Some novelties in presumed males of Leptanillinae (Hym., Formicidae). Entomologiske Meddelelser 36: 577 - 598.","Gotwald W. H. Jr. 1969. Comparative morphological studies of the ants, with particular reference to the mouthparts (Hymenoptera: Formicidae). Cornell University Agricultural Experiment Station 408: 3 - 150.","Wheeler G. C. & Wheeler J. 1972. The subfamilies of Formicidae. Proceedings of the Entomological Society of Washington 74: 35 - 45.","Baroni Urbani C. 1977. Materiali per una revisione della sottofamiglia Leptanillinae Emery Hymenoptera: Formicidae. Entomologica Basiliensia 2: 427 - 488.","Bolton B. 1990 b. The higher classification of the ant subfamily Leptanillinae (Hymenoptera: Formicidae). Systematic Entomology 15: 267 - 282. http: // dx. doi. org / 10.1111 / j. 1365 - 3113.1990. tb 00063. x","Baroni Urbani C., Bolton B. & Ward P. S. 1992. The internal phylogeny of ants (Hymenoptera: Formicidae). Systematic Entomology 17: 301 - 329. http: // dx. doi. org / 10.1111 / j. 1365 - 3113.1992. tb 00553. x","Ogata K., Terayama M. & Masuko K. 1995. The ant genus Leptanilla: discovery of the worker-associated male of L. japonica, and a description of a new species from Taiwan (Hymenoptera: Formicidae: Leptanillinae). Systematic Entomology 20: 27 - 34. http: // dx. doi. org / 10.1111 / j. 1365 - 3113.1995. tb 00081. x","Bolton B. 2003. Synopsis and classification of Formicidae. Memoirs of the American Entomological Institute 71: 1 - 370.","Borowiec M. L., Schulz A., Alpert G. D. & Banar P. 2011. Discovery of the worker caste and descriptions of two new species of Anomalomyrma (Hymenoptera: Formicidae: Leptanillinae). with unique abdominal morphology. Zootaxa 2810: 1 - 14.","Wheeler W. M. 1923. Social life among the insects. New York, Harcourt, Brace and Co.","Bernard F. 1951. Hymenopteres, super-famille des Formicoidea Ashmead 1905. In: Grasse P. P. (ed) Traite de Zoologie. Tome X. Fasc. II: 997 - 1104. Masson et Cie, Paris.","Yamane S., Bui T. V. & Eguchi K. 2008. Opamyrma hungvuong, a new genus and species of ant related to Apomyrma (Hymenoptera: Formicidae: Amblyoponinae). Zootaxa 1767: 55 - 63.","Saux C., Fisher B. L. & Spicer G. S. 2004. Dracula ant phylogeny as inferred by nuclear 28 S rDNA sequences and implications for ant systematics (Hymenoptera: Formicidae: Amblyoponinae). Molecular Phylogenetics and Evolution 33: 457 - 468. http: // dx. doi. org / 10.1016 / j. ympev. 2004.06.017"]}

Subfamily Dorylinae Leach, 1815 Figs 5 A���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. 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.)., Published as part of Boudinot, Brendon E., 2015, 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, pp. 1-62 in European Journal of Taxonomy 120 on pages 48-49, DOI: 10.5852/ejt.2015.120, http://zenodo.org/record/3780152, {"references":["Brady S. G., Schultz T. R., Fisher B. L. & Ward P. S. 2014. The rise of army ants and their relatives: diversification of specialized predatory doryline ants. BMC Evolutionary Biology 14: 93. http: // dx. doi. org / 10.1186 / 1471 - 2148 - 14 - 93.","Bolton B. 2003. Synopsis and classification of Formicidae. Memoirs of the American Entomological Institute 71: 1 - 370."]}

Subfamily Agroecomyrmecinae Carpenter, 1930 Figs 5F, 13 A���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 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 Agroecomyrmecinae, the Neotropical Tatuidris and Afrotropical Ankylomyrma (Ward et al. 2015). The male of Ankylomyrma is unknown and will be a notable discovery., Published as part of Boudinot, Brendon E., 2015, 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, pp. 1-62 in European Journal of Taxonomy 120 on pages 46-47, DOI: 10.5852/ejt.2015.120, http://zenodo.org/record/3780152, {"references":["Ward P. S., Brady S. G., Fisher B. L. & Schultz T. R. 2015. The evolution of myrmicine ants: Phylogeny and biogeography of a hyperdiverse ant clade (Hymenoptera: Formicidae). Systematic Entomology 40: 61 - 81. http: // dx. doi. org / 10.1111 / syen. 12090"]}

Subfamily Paraponerinae Emery, 1901 Figs 1, 2C, 3C, 4D, 13 C���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., Published as part of Boudinot, Brendon E., 2015, 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, pp. 1-62 in European Journal of Taxonomy 120 on page 47, DOI: 10.5852/ejt.2015.120, http://zenodo.org/record/3780152

Subfamily Myrmeciinae Emery, 1877 Figs 5 C���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., Published as part of Boudinot, Brendon E., 2015, 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, pp. 1-62 in European Journal of Taxonomy 120 on page 49, DOI: 10.5852/ejt.2015.120, http://zenodo.org/record/3780152, {"references":["Ward P. S. & Brady S. G. 2007. Phylogeny and biogeography of the ant subfamily Myrmeciinae (Hymenoptera: Formicidae). Invertebrate Systematics 17: 361 - 386. http: // dx. doi. org / 10.1071 / IS 02046"]}

Subfamily Ponerinae Lepeletier 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. 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 definition 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.)., Published as part of Boudinot, Brendon E., 2015, 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, pp. 1-62 in European Journal of Taxonomy 120 on pages 47-48, DOI: 10.5852/ejt.2015.120, http://zenodo.org/record/3780152, {"references":["Schmidt C. 2013. Molecular phylogenetics of ponerine ants (Hymenoptera: Formicidae: Ponerinae). Zootaxa 3647: 201 - 250. http: // dx. doi. org / 10.11646 / zootaxa. 3647.2.1","Schmidt C. A. & Shattuck S. O. 2014. The higher classification of the ant subfamily Ponerinae (Hymenoptera: Formicidae), with a review of ponerine ecology and behavior. Zootaxa 3817: 1 - 242. http: // dx. doi. org / 10.11646 / zootaxa. 3817.1.1","Bolton B. 2003. Synopsis and classification of Formicidae. Memoirs of the American Entomological Institute 71: 1 - 370.","Yoshimura M. & Fisher B. L. 2007. A revision of male ants of the Malagasy region (Hymenoptera: Formicidae): key to subfamilies and treatment of the genera of Ponerinae. Zootaxa 1654: 21 - 40.","Yoshimura M. & Onoyama K. 2002. Male-based keys to the subfamilies and genera of Japanese ants (Hymenoptera: Formicidae). Entomological Science 5: 421 - 443."]}

Subfamily Myrmicinae Lepeletier de Saint-Fargeau, 1835 Figs 4H, 5A, G, 16 D���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., Published as part of Boudinot, Brendon E., 2015, 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, pp. 1-62 in European Journal of Taxonomy 120 on page 56, DOI: 10.5852/ejt.2015.120, http://zenodo.org/record/3780152, {"references":["Ward P. S., Brady S. G., Fisher B. L. & Schultz T. R. 2015. The evolution of myrmicine ants: Phylogeny and biogeography of a hyperdiverse ant clade (Hymenoptera: Formicidae). Systematic Entomology 40: 61 - 81. http: // dx. doi. org / 10.1111 / syen. 12090"]}

Subfamily Dolichoderinae Forel, 1878 Figs 7E, 15 D���E Diagnosis 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; 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 of the 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.)., Published as part of Boudinot, Brendon E., 2015, 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, pp. 1-62 in European Journal of Taxonomy 120 on pages 50-51, DOI: 10.5852/ejt.2015.120, http://zenodo.org/record/3780152, {"references":["Yoshimura M. & Fisher B. L. 2011. A revision of male ants of the Malagasy region (Hymenoptera: Formicidae): Key to genera of the subfamily Dolichoderinae. Zootaxa 2794: 1 - 34.","Czechowski W., Radchenko A., Czechowska W. & Vepsalainen K. 2012. The ants of Poland with reference to the myrmecofauna of Europe. Fauna Poloniae 4. Warsaw, Natura Optima Dux Foundation.","Shattuck S. O. 1992. Generic revision of the ant subfamily Dolichoderinae (Hymenoptera: Formicidae). Sociobiology 21: 1 - 181."]}

Martialis heureka Rabeling & Verhaagh, 2008 Figs 11���12 Martialis heureka Rabeling & Verhaagh, in Rabeling et al. 2008: 14914, figs. 1���2 (worker). BRAZIL, Amazonas: Manaus, Headquarters of Empresa Brasileira de Pesquisa Agropecu��ria (EMBRAPA) Amaz��nia 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). HL 0.35���0.42, HW1 0.35���0.40, HW2 0.44���0.49, MAL 0.04���0.06, MDL 0.10��� 0.11, SL 0.20���0.23, PDL 0.10���0.13, A3L 0.17���0.21, AAL 0.17���0.21, EL 0.17���0.19, EW 0.14���0.17, OOD 0.14���0.15, LOD 0.04���0.05, MOD 0.04���0.05, ML 0.64���0.78, MLL 0.16���0.18, MLW 0.17���0.20, MTL 0.29���0.37, MTW 0.36���0.45, PFL 0.39���0.48, MFL 0.43���0.53, PTH 0.16���0.19, PTL 0.20���0.23. INDICES. CI 0.94���1.02, CS 0.35���0.41, SEI 83.6���86.2, SI 55.6���58.4, EI 83.0���85. 6, EYE 88.1���88.7, MI 26.7���27.8, OBI 80.1���82.8, OMI 3.46���4.74, 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. 11 A���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. Labrum 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 weakly 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 4/5 x antennomere 3 length; funiculus filiform, elongate, reaching metasoma when laid against mesosoma. MESOSOMA (Fig. 11 C���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 1/3 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 weakly 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, 1m-cu absent, thus discal cell 1 open. Costal vein tubular to pterostigma. Rsf1 slightly more than 1/2 x length of and meeting Mf1 obliquely. Rs+M continuous with undifferentiated Mf2-3 until meeting very short 2rsm. Rsf2+3 absent. 2r-rs very long, longer than combined lengths of Rsf1 and Mf1; 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 Mf1. 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. 12 C���H). Pygostyles absent.Abdominal sternum IX spiculum short; anterior margin linear, curving posterolaterally near lateral margins; lateral margins short, slightly divergent; posterolateral margins weakly 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 posteroventrallydirected setae. Basivolsella lateromedially broad; base transversely connected with basimere; cuspis absent; digitus clavate, apex swollen and directed ventrally; digital stem short. Valvura short, linear, directed anteriorly and situated at about 2/3 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 almost uniformly brown to brownish yellow; extremities slightly lighter colored. SCULPTURATION. Body weakly 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 (Brand��o et al. 2010) and multi-locus molecular phylogenetic reconstruction (Rabeling et al. 2008; although see K��ck et al. 2011). Rabeling et al. (2008) recovered Martialis as the sister to all remaining extant ants including the Old World subfamily Leptanillinae, while a reanalysis by K��ck 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 welldeveloped, 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 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 weakly 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.) Shared apomorphies of the basal ants Of particular interest for the ���basal ant��� problem (Brady et al. 2006; Rabeling et al. 2008; K��ck et al. 2011) is the relationship of the Leptanillinae, Martialinae, and the Amblyoponinae. While the relationships within the formicoid clade (Dorylinae, myrmeciomorphs, dolichoderomorphs, Formicinae, ectaheteromorphs, 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, Paraponerinae, 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 K��ck et al. (2011), who recovered Leptanillinae 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 Leptanillinae are sister to the extant Formicidae or whether they are highly derived poneroids. Unpublished molecular phylogenetic analyses by P.S. Ward (discussed in Ward 2014), in which the outgroups are excluded, recover Martialis and the Leptanillinae as part of a bipartition comprising part of the poneroids; moreover, these analyses recover Opamyrma as sister to the Leptanillinae. 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 Leptanillinae, and the Amblyoponinae. The male of Martialis is more generalized morphologically than most Leptanillinae, although some Protanilla display a mosaic of generalized and specialized characteristics. Martialis males differ from all known male Leptanillinae 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, Published as part of Boudinot, Brendon E., 2015, 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, pp. 1-62 in European Journal of Taxonomy 120 on pages 37-46, DOI: 10.5852/ejt.2015.120, http://zenodo.org/record/3780152, {"references":["Rabeling C., Brown J. M. & Verhaagh M. 2008. Newly discovered sister lineage sheds light on early ant evolution. Proceedings of the National Academy of Sciences B 105: 14913 - 14917. http: // dx. doi. org / 10.1073 / pnas. 0806187105","Brandao C. R. F., Diniz J. L. M. & Feitosa R. S. M. 2010. The venom apparatus and other morphological characters of the ant Martialis heureka (Hymenoptera, Formicidae, Martialinae). Papeis Avulsos de Zoologia 50: 413 - 423.","Kuck P., Hita Garcia F., Misof B. & Meusemann K. 2011. Improved phylogenetic analyses corroborate a plausible position of Martialis heureka in the ant tree of life. PLoS ONE 6: e 21031. http: // dx. doi. org / 10.1371 / journal. pone. 0021031","Kaspari M., Pickering J., Longino J. T. & Windsor D. 2001 a. The phenology of a Neotropical ant assemblage: Evidence for continuous and overlapping production. Behavioral Ecology and Sociobiology 50: 382 - 390. http: // dx. doi. org / 10.1007 / s 002650100378","Kaspari M., Pickering J. & Windsor D. 2001 b. The reproductive flight phenology of a Neotropical ant assemblage. Ecological Entomology 26: 245 - 257. http: // dx. doi. org / 10.1046 / j. 1365 - 2311.2001.00320. x","Dunn R. R., Parker C. R., Geraghty M. & Sanders N. J. 2007. Reproductive phenologies in a diverse temperate ant fauna. Ecological Entomology 32: 135 - 142. http: // dx. doi. org / 10.1111 / j. 1365 - 2311.2006.00839. x","Vasconcelos H. L., Carvalho K. S. & Delabie J. H. C. 2001. Landscape modifications and ant communities. In: Bierregaard Jr., Gascon C., Lovejoy T. E. & Mesquita R. C. G. (eds) Lessons from Amazonia: The Ecology and Conservation of a Fragmented Forest: 199 - 207. Yale University Press, New Haven.","Brady S. G., Schultz T. R., Fisher B. L. & Ward P. S. 2006. Evaluating alternative hypotheses for the early evolution and diversification of ants. Proceedings of the National Academy of Sciences of the United States of America 103: 18172 - 18177. http: // dx. doi. org / 10.1073 / pnas. 0605858103","Moreau C. S., Bell C. D., Vila R., Archibald S. B. & Pierce N. E. 2006. Phylogeny of the ants: diversification in the age of angiosperms. Science 312: 101 - 104. http: // dx. doi. org / 10.1126 / science. 1124891","Brady S. G., Schultz T. R., Fisher B. L. & Ward P. S. 2014. The rise of army ants and their relatives: diversification of specialized predatory doryline ants. BMC Evolutionary Biology 14: 93. http: // dx. doi. org / 10.1186 / 1471 - 2148 - 14 - 93.","Ward P. S., Brady S. G., Fisher B. L. & Schultz T. R. 2015. The evolution of myrmicine ants: Phylogeny and biogeography of a hyperdiverse ant clade (Hymenoptera: Formicidae). Systematic Entomology 40: 61 - 81. http: // dx. doi. org / 10.1111 / syen. 12090","Ward P. S. 2014. The phylogeny and evolution of ants. Annual Review of Ecology, Evolution, and Systematics 45: 2.1 - 2.21. http: // dx. doi. org / 10.1146 / annurev-ecolsys- 120213 - 091824","Grimaldi D., Agosti D. & Carpenter J. M. 1997. New and rediscovered primitive ants (Hymenoptera: Formicidae) in Cretaceous amber from New Jersey, and their phylogenetic relationships. American Museum Novitates 3208: 1 - 43.","Bolton B. 1990 c. Army ants reassessed: the phylogeny and classification of the doryline section (Hymenoptera, Formicidae). Journal of Natural History 24: 1339 - 1364. http: // dx. doi. org / 10.1080 / 00222939000770811","Yoshimura M. & Fisher B. L. 2012 b. A revision of the Malagasy endemic genus Adetomyrma (Hymenoptera: Formicidae: Amblyoponinae). Zootaxa 3341: 1 - 31.","Kugler C. 1992. Stings of ants of the Leptanillinae (Hymenoptera: Formicidae). Psyche 99: 103 - 115.","Wheeler G. C. & Wheeler E. W. 1976. Ant larvae: Review and synthesis. Memoirs of the Entomological Society of Washington 7: 1 - 108.","Bolton B. 2003. Synopsis and classification of Formicidae. Memoirs of the American Entomological Institute 71: 1 - 370."]}

Subfamily Pseudomyrmecinae M.R. Smith, 1952 Figs 5B, E, 15 A���B Diagnosis Pseudomyrmecine males are uniquely identified by the absence of the jugal lobes in combination with the following character combination: cuticle soft, flexible, weakly-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., Published as part of Boudinot, Brendon E., 2015, 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, pp. 1-62 in European Journal of Taxonomy 120 on page 49, DOI: 10.5852/ejt.2015.120, http://zenodo.org/record/3780152, {"references":["Ward P. S. 1990. The ant subfamily Pseudomyrmecinae (Hymenoptera: Formicidae): generic revision and relationship to other formicids. Systematic Entomology 15: 449 - 489. http: // dx. doi. org / 10.1111 / j. 1365 - 3113.1990. tb 00077. x"]}

Rhopalothrix orbis Taylor, 1968 Rhopalothrix orbis Taylor, 1968: 336, figs. 1���3. Holotype, worker: Australia, Queensland: Tamborine Mountains, north side near Curtis Falls, Berlese funnel sample, leafmould, rainforest, 8.v. 1953 (T. E. Woodward) [ANIC] (not examined)., Published as part of Longino, John T. & Boudinot, Brendon E., 2013, New species of Central American Rhopalothrix Mayr, 1870 (Hymenoptera, Formicidae), pp. 301-324 in Zootaxa 3616 (4) on page 315, DOI: 10.11646/zootaxa.3616.4.1, http://zenodo.org/record/220287

Rhopalothrix kusnezovi Brown & Kempf, 1960 Rhopalothrix kusnezovi Brown & Kempf, 1960: 238, fig. 60. Holotype, queen: Argentina, Tucum��n (N. Kusnezov, no. 10068) [IFML] (not examined)., Published as part of Longino, John T. & Boudinot, Brendon E., 2013, New species of Central American Rhopalothrix Mayr, 1870 (Hymenoptera, Formicidae), pp. 301-324 in Zootaxa 3616 (4) on page 312, DOI: 10.11646/zootaxa.3616.4.1, http://zenodo.org/record/220287

Meranoplus sylvarius Boudinot & Fisher sp. n. (Figs. 10, 23, 28, 29, 57 ���62, 66) Holotype worker, MADAGASCAR: Fianarantsoa, Vevembe, 22 �� 47.46 ��� S, 47 �� 10.91 ��� E, 600 m, rainforest transition to montane forest, collection code BLF14131, 23��� 24 Apr 2006 (B.L. Fisher et al.) (CASC: CASENT017529). Paratype gyne with same data as holotype worker (CASC: CASENT0317567). Paratype workers with same data as holotype worker ( BMNH: CASENT0317527; CASC: CASENT0317536, CASENT0317566, CASENT0070611, CASENT0317528, CASENT0317540���CASENT0317542, CASENT0317557���CASENT0317559, CASENT0317544, CASENT0317547, CASENT0317548, CASENT0317548���CASENT0317552, CASENT0317554���CASENT0317556; MHNG: CASENT0317538; MCZ: CASENT0317543; USNM: CASENT0317549; NHMB: CASENT0317553). Description Worker (holotype in parentheses). HL 1.03���1.13 (1.06), HW 0.90���1.02 (0.96), HLA 0.37���0.41 (0.39), CW 0.36���0.38 (0.38), CDD 0.09���0.11 (0.11), SL 0.73���0.80 (0.78), EL 0.22���0.23 (0.23), EW 0.15���0.16 (0.16), PML 0.72���0.80 (0.76), PWA 0.88���0.98 (0.94), PWP 0.47���0.53 (0.48), SPL 0.19���0.22 (0.21), WL 1.04���1.17 (1.12), PTL 0.23���0.25 (0.25), PTH 0.43���0.49 (0.46), PPL 0.23���0.24 (0.24), PPH 0.40���0.45 (0.42), ATW 1.23���1.34 (1.32), ATL 1.27���1.48 (1.42), CI 87.4���90.6 (90.1), SI 77.7���81.3 (81.1), OMI 58.3���59.6 (59.4), CDI 23.9 ���- 31.2 (27.3), SEI 324���355 (336), PMI 122.7���124.4 (122.8), PWI 51.5���55.3 (51.5), CS 1.0��� 1.1 (1.0), EYE 37.7���40.4 (38.5) (7 measured). Chocolate brown to almost black. Head longer than broad (CI 77���82). Mandibles with nearly parallel striation. Face with dilute costae which veer away from posterior margin of head; area anterior to nuchal carina without rugae or areolation. Area around eyes costate; costae in malar area somewhat rugose. Frontal carinae comparatively narrow, not concealing margin of eyes in full-face view. Scrobal carina weakly developed; anterior half bulging. Anterior margin of clypeus with weakly bilobed lamina; lobes close-set (CDI 24���31). Middle portion of clypeus shining and weakly costate; lateral thirds of middle portion with about two costae; middle third without costae. Eyes large (EYE 35���41). Maximum length of eye distinctly less than �� length of malar area (OMI 55���60). Face with erect to subdecumbent setae of various lengths. Setae ventrally and anterior to the eye mostly subdecumbent. Promesonotal shield longer than broad (PMI 122���125); posterolateral corners close-set (PWI 51���55). Dorsum of promesonotum with dilute costae, mostly concentrated in anterior half; interstices with setae raised on tiny nodules. Promesonotal shield setae decumbent to suberect, of mixed lengths. Lateral face of pronotum with weak rugae; setae raised on nodules. Lateral face of mesonotum smooth and shining. Katepi-anepisternal suture present; mesopleural-mesonotal suture present; mesopleural-metapleural suture absent to present. Mesopleuron smooth and strongly shining, with very dilute (1 to 4), very weak indications of raised sculpture other than in posteroventral area. Metapleural lobes well-developed. Propodeal spines thornlike; dorsomedian carina absent; dorsolateral carina extends dorsolaterally to promesonotum. Posterior face of propodeum smooth and shining; very weak costae may be present in top third, which otherwise has setae raised on nodules. Petiole cuneate in profile, without pedicel. Anteroventral process of petiole a simple denticle. Postpetiole nodiform, about as tall as long (PTI 50���55). Sub-post-petiolar process well-developed, with a strong inflection between posterior half of sternum. Dorsal face of postpetiole with weak to indistinct rugae; lateral face with weak to indistinct rugae; posterior face smooth and shining except for nodulose setae and transverse costae near extreme base. Fourth abdominal tergum (ATIV) comparatively small (ATW/WL 1.15���1.23, ATL/HL 1.20���1.33). ATIV shining, with fine areolate sculpture between stellate setiferous punctures; smooth near base of tergum. Setae of ATIV of mixed lengths; shortest subdecumbent and longer than width of metafemora; longest setae about twice as long as shortest. Queen (paratype). HL 1.22, HW 1.17, HLA 0.41, CW 0.44, CDD 0.13, SL 0.82, EL 0.29, EW 0.22, SPL 0.20, WL 1.61, PTL 0.31, PTH 0.56, PPL 0.29, PPH 0.56, ATW 1.82, ATL 1.89, CI 95.2, SI 70.6, OMI 71.4, CDI 29.1, SEI 285, CS 1.2, EYE 42.4. Color: chocolate brown. Large (WL 1.61). Head longer than broad (CI 95). Mandibles with more-or-less parallel striae. Face with weakly rugose costae which continue to the posterior margin of the head. Area above eye areolate; behind and beneath eye costate. Scrobal carina weakly bulging in anterior half. Anterior margin of clypeus with two welldeveloped, close-set laminar lobes (CDI 29). Middle portion of clypeus with strong costae in lateral thirds; middle third without sculpture. Eyes large (EYE 42). Maximum eye length about �� length of malar area (OMI 71). Scapes long (SI 71). Face with erect to subdecumbent setae; longest setae about 1.5 length of shortest setae. Mesosoma longer than tall. Anterolateral corners of pronotum angled; dorsolateral margin of pronotum ecarinate. Mesosomal dorsum with little sculpture: pronotum smooth medially, rugose laterally; scutum smooth and shining in anterior half, with dilute rugae in posterior half; scutellum with transverse rugae branching more frequently in lateral portions. Mesosomal dorsum with subdecumbent to erect setae of mixed lengths. Lateral face of pronotum rugose anteriorly; with setae raised on strong nodules. Katepisternum mostly smooth and shining; without costae extending longitudinally margin to margin. Anteroventral portion of anepisternum smooth and shining; dorsal portion with longitudinal rugae, interstices roughened. Metapleuron roughened with rugae; area anteroventral to propodeal spiracle smooth and shining. Propodeal spines thorn-like, long (SPL 0.20). Meso and metatibial spurs absent. Wings unknown. Petiole cuneate in profile; subpetiolar process dentiform. Postpetiole nodiform; sternal process large. Dorsal and lateral faces of postpetiole rugose; posterior face roughened near base and with dilute longitudinal costae. Forth abdominal tergum (ATIV) with sides convex; nearly as broad as long (ATW/GL 0.96). ATIV with fine, weak areolate sculpture between stellate setiferous punctures; weak, dilute costae present in basal region. ATIV with subdecumbent short setae; suberect setae about 1.5-2 times as long as shorter setae. Diagnosis Worker. The absence of meso- and metatibial spurs is unique in the Malagasy fauna. This new species may be separated from M. radamae by the absence of a dorsomedial costa extending from the base of the propodeal spine to the promesonotum. Gyne. The combination of stellate setiferous punctures, comparatively large size (WL 1.61), dark chocolatebrown color, malar area about �� the length of the eye (OMI 71.4), long scapes (SI 70.6), uniquely identify gynes of this species in the Malagasy fauna. Comments The type series of M. sylvarius is designated from a large nest collection from the rainforest/montane forest of Vevembe, the only known locale of this species. Restriction to moist forest habitat is unique for Malagasy Meranoplus. Although not collected in sympatry with M. radamae, M. sylvarius is morphologically distinct enough to support the hypothesis of reproductive isolation. The lack of meso- and metatibial spurs is unique in the Malagasy Meranoplus fauna. Although M. radamae shows much geographic variation in its sculpturation, the trends of sculpture reduction across body parts never move in synchrony to such extremes, suggesting that the almost complete reduction of sculpture on specimens of M. sylvarius would be highly unlikely under circumstances of gene flow., Published as part of Boudinot, Brendon E. & Fisher, Brian L., 2013, A taxonomic revision of the Meranoplus F. Smith of Madagascar (Hymenoptera: Formicidae: Myrmicinae) with keys to species and diagnosis of the males, pp. 301-339 in Zootaxa 3635 (4) on pages 333-337, DOI: 10.11646/zootaxa.3635.4.1, http://zenodo.org/record/247717

Rhopalothrix triumphalis Longino & Boudinot, sp. nov. (Figs 1 A, 2 B, 3 F, 14, 16) Type material. Holotype, worker: MEXICO, Chiapas: 2.8 km ESE Custepec, 15.72078 -92.93925 ± 50 m, 1800 m, 17 Jul 2007, mixed hardwood forest, ex sifted leaf litter (R.S. Anderson# 2007 -017) [CAS, unique specimen identifier CASENT0602067]. Paratypes (workers): same data as holotype but 18 Jul 2007, liquidambar forest (R.S. Anderson# 2007 -018) [ECOSCE, CASENT0601766]; 1.8 km SE Custepec, 15.72198 -92.95037 ± 50 m, 1530 m, mixed liquidambar forest (R.S. Anderson# 2007 -020) [MCZC, CASENT0601887]; 2 km SE Custepec, 15.72298 -92.94493 ± 50 m, 1650 m, ridgetop oak forest (R.S. Anderson# 2007 -021) [USNM, CASENT0601965]; 2.8 km SE Custepec, 15.72260 -92.93995 ± 50 m, 1840 m, oak forest (R.S. Anderson# 2007 -022) [UNAM, CASENT0603013]. Geographic range. Mexico (Chiapas). Diagnosis. Anterior labral lobe bilobed, with lateral lobule longer than medial lobule; masticatory margin of mandible with two teeth; squamiform setae of first gastral tergite abundant, elongate, 4 × or 5 × longer than wide; HW 0.57–0.65. Description. Worker. HW 0.57–0.65 (n= 12); mandible with two teeth on masticatory margin, second tooth from base largest; subapical tooth with minute reclinate denticle at base; subapical tooth about twice as long as apical tooth; intercalary teeth prominent, one closest to apical tooth about half as long as apical tooth; labrum trapezoidal, anterior margin bilobed, lateral lobule triangular, longer than medial lobule, medial lobules rounded, flanking semicircular median notch; arcuate promesonotal groove and metanotal groove moderately impressed; propodeal tooth variable, obtuse, right angled, or acute, infradental lamella evenly and shallowly concave; squamiform setae abundant on first gastral tergite, uniformly covering entire tergite; gastral setae relatively long and thin, 4–5 × longer than wide, with elongate stem below widened apex. Queen. HW 0.69; mandible and labrum similar to worker; face shape similar to worker but with less strongly developed grooves and ridges; compound eye longer than maximum width of scape; ocelli small, cuticle adjacent to ocelli marked with black pigment spots on evenly light brown background; shape of propodeal tooth, infradental lamella, petiole and postpetiole similar to worker; katepisternum and anepisternum large, convex, separated by broad U-shaped groove; layer of sparse, long, appressed pubescence covers mandible, face, scapes, legs, dorsal mesosoma and metasoma; abundant stiff erect setae on face, anterior edge of scape, side of head, dorsal mesosoma, dorsal gaster. The male is unknown. Biology. This species occurs in cloud forest habitats, from 1360–2140 m elevation. All specimens are from Winkler or Berlese samples of sifted leaf litter. It is sympatric with R. megisthmica on the slopes of Volcán Tacaná in Chiapas. At the type locality it was moderately abundant, occurring in 17 of 100 miniWinkler samples. Comments. Rhopalothrix triumphalis, R. atitlanica, and R andersoni share a distinctive labrum shape with bilobed anterolateral margin, with the lateral lobule longer and more triangular than the medial lobule. The three form a geographic replacement series, with no known zones of sympatry. An isolated specimen from eastern Guatemala (Petén: 13 km NW Machaquilá, 16.44173 -89.53527 ± 26m, 390m, 28 May 2009, LLAMA#Wm-B- 06- 1 -05, CASENT0614342) matches the size and morphology of R. triumphalis, differing only in the disposition of squamiform setae on the gaster. Instead of uniformly covering the first gastral tergite, they cover only the posterior 3 / 4. This specimen is left unidentified in this report, pending further understanding of the group. Etymology. Referring to the type locality, El Triunfo Biosphere Reserve in the Sierra Madre de Chiapas.

Meranoplus radamae Forel, 1891 (Figs. 9, 14, 16 ���18, 20, 22, 24, 30, 32, 48 ���56, 65) Meranoplus radamae Forel, 1891: 148, pl. 4, fig. 10. Lectotype worker, here designated, MADAGASCAR: Imerina (Central Madagascar) (M. Hildebrandt) (MHNG: CASENT0101239) [examined]. Description Worker. HL 0.86���1.04, HW 0.79���0.92, HLA 0.32���0.38, CW 0.29���0.36, CDD 0.08���0.10, SL 0.60���0.72, EL 0.18���0.22, EW 0.13���0.16, PML 0.58���0.76, PWA 0.73���0.92, PWP 0.43���0.55, SPL 0.16���0.22, WL 0.86 ��� 1.1.05, PTL 0.18���0.25, PTH 0.38���0.46, PPL 0.19���0.26, PPH 0.32���0.42, ATW 1.01���1.24, ATL 1.09���1.40, CI 88.7���93.8, SI 70.5���79.4, OMI 55.9���62.7, CDI 22.4���34.2, SEI 290���350, PMI 121.3���134.3, PWI 54.0��� 62.5, CS 0.8 ���1.0, EYE 36.2���40.5 (19 measured). Coloration variable: dark orange, orange brown or yellowish brown with yellow abdominal segments IV���VII. Head longer than broad (CI 92���95). Mandibles striate. Face longitudinally rugose to dilutely costate; area anterior to nuchal carina areolate to rugose. Area around eyes areolate to rugose or costate beneath and behind. Frontal carinae concealing dorsal margin of eyes in full face view. Scrobal carina weak to nearly absent. Anterior margin of clypeus with weakly bilobed lamina; lobes relatively close set (CDI 22���36). Middle portion of clypeus costate to rugose laterally; smooth and shining medially. Setae on face subdecumbent to erect; of mixed lengths; longest setae about twice as long as shortest. Promesonotal shield longer than broad (PMI 120���134). Lateral margins of promesonotum tapering to midline posteriorly; posterolateral denticles comparatively close-set (PWI 54���63). Dorsum of promesonotum areolate to dilutely rugose. Setae on promesonotal shield subdecumbent to erect; of mixed lengths; longest setae about 1.25 as long as shortest. Lateral face of pronotum rugose anteriorly; posterior portion shining, variably smooth to weakly areolate; setiferous nodules mostly hidden by sculpture. Lateral face of mesonotum smooth and shining to somewhat roughened. Anapleural and mesometapleural sutures present. Mesopleuron shining; with or without weak areolate sculpture or costae on katepisternum. Metapleuron shining with or without weak areolation; smooth anterodorsal metapleural bulla. Lateral face of propodeum shining, roughened with rugae dorsal to spiracle. Propodeal spines long, thorn-shaped (0.16���0.22), with both dorsolateral and dorsomedian costae reaching notopropodeal suture. Posterior face of propodeum with costae in dorsal region. Petiole cuneate in profile although sometimes with bulge posteriorly near apex; without pedicel. Subpetiolar process dentiform. Anterior and posterior faces of petiole smooth and shining. Postpetiole nodiform; without distinct posterodorsal angle. Dorsal face of postpetiole areolate to rugose; lateral face rugose; posterior face areolate to rugose. Fourth abdominal tergite (ATIV) can be small (ATW/WL 1.13���1.33, ATL/WL 1.15���1.48). ATIV base variably rugose and rough to shining between setiferous punctures. ATIV with fine areolate sculpture, which may be weak, between stellate setiferous punctures. ATIV setae of mixed lengths; short setae subdecumbent; long setae suberect; longest setae 1.25���1.5 times as long as shortest setae. Queen. HL 1.09���1.15, HW 1.05���1.10, HLA 0.35���0.38, CW 0.37���0.41, CDD 0.10���0.13, SL 0.70���0.74, EL 0.26���0.29, EW 0.21, SPL 0.19���0.23, WL 1.39���1.49, PTL 0.27���0.31, PTH 0.47���0.49, PPL 0.24���0.30, PPH 0.47���0.52, ATW 1.47���1.58, ATL 1.68���1.92, CI 92.1���96.1, SI 68.0��� 70.3, OMI 72.6���78.5, CDI 25.9���32.8, SEI 253���283, CS 1.1, EYE 43.3���44.8 (6 measured). Light brown to pale yellowish orange. Small (WL 1.4���1.5). Head longer than broad (CI 92���97). Mandibles striate; ridges may be evenly curved or wavy. Face areolate to longitudinally rugose; area posterior to ocelli areolate to transversely rugose. Area above and below eyes areolate; areolate to rugose behind eyes. Scrobal carina distinct to indistinct; broadest in anterior half. Anterior margin of clypeus with bilobed lamina; lobes relatively close-set (CDI 26���33). Middle portion of clypeus rugose to costate laterally, smooth medially; lateral ridges may meet anteriorly. Eyes large (EYE 43���53). Maximum eye length �� length of malar area (OMI 73���79). Scapes long (SI 67���70). Face with setae of mixed lengths; longest setae 1.25���2 times as long as shortest setae. Pronotum with anterolateral angles; dorsolaterally ecarinate. Pronotal dorsum variably areolate to rugose. Scutum smooth to rugose on anterior half; always with some rugae on posterior half. Scutellum areolate to longitudinally rugose. Lateral face of pronotum areolate to sub-areolate on anterior portion; shining, roughened, sometimes with rugae on posterior portion. Katepisternum smooth and shining, with longitudinal costae often extending from anapleural suture to mesometapleural suture. Anepisternum shining ventrally; rugose dorsally. Metapleuron and lateral face of propodeum rugose, with variable shining area anterodorsal to metapleural bulla. Propodeal spines long, thorn-like (SPL 0.19���0.23). Wings as in Meranoplus male diagnosis. Petiole cuneate in profile, without pedicel. Postpetiole nodiform, without distinct posterodorsal angle. Subpostpetiolar process well-developed; robust to thin. Dorsal face of postpetiole areolate to rugose; lateral face rugose-costate; posterior face areolate with interstices punctate to longitudinally rugose. Fourth abdominal tergum (ATIV) with sides convex in dorsal view; long (ATL/WL 1.20���1.29). Base of ATIV rough to smooth, with variably developed rugae; setiferous punctures strong near base, increasing in stellation posteriorly; ATIV with weak fine areolation between setiferous punctures. ATIV setae of mixed lengths: short setae subdecumbent, shortest setae about �� length of longest setae; long setae erect, varying in density. Male. HL 0.52���0.63, HW 0.55���0.60, HLA 0.06���0.09, SL 0.15���0.18, EL 0.24���0.26, EW 0.19���0.23, WL 0.70���1.20, PTL 0.19���0.25, PTH 0.20���0.22, PPL 0.15���0.19, PPH 0.19���0.22, ATW 0.79���1.01, ATL 0.83���0.96, CI 96.3���106.4, SI 27.0��� 29.7, OMI 301���428, SEI 61���69, CS 0.5���0.6, EYE 75.9���80.5 (3 measured). Body piceous brown, appendages light brown. Small (WL 0.7���1.2, CS 0.5���0.6). Head longer than broad or broader than long (CI 96���106). Mandibles variably developed: short and spatulate with edentate masticatory margin to longer, bladelike, with apical tooth. Frontal carinae distinct to indistinct. In full-face view, posterior margin of head not raised above ocelli greater than distance between lateral ocelli. Head variably rugose with foveate sculpture. Scapes short (SI 27���30). Eyes large (EYE 76���81), greater than three times as long as malar area (OMI 300���430). Ocelli about the size of the antennal socket. Majority of setae on body shorter than eye length, subdecumbent to erect, uniformly straight to evenly curved. Notauli distinct; may continue to posterior margin of scutum after converging. Parapsidal lines distinct, impressed or not. Scutum smooth and shining, albeit undulating with coarse setiferous punctures. Scutellum shining at least medially; roughened laterally. Katepisternum smooth and shining. Anepisternum smooth and shining; posterior half variably rough and rugose to smooth and shining. Lower half of metapleuron with a shining patch of variable size; dorsal half rugose. Posterior face of propodeum with longitudinal costae laterally; variably foveate to smooth. Petiole and postpetiole shape variable (PTI 87���114, PPI 72���95). Petiole and postpetiole sessile; apex of petiolar node node acute to rounded; postpetiole subglobose. Petiolar sternum occasionally with setae. Postpetiolar sternum setose. Fourth abdominal tergum shining, with weak, fine areolate sculpture and with stellate setiferous punctures. Fourth abdominal sternum shining. Foramen genitale constricted; basal ring broadening posteriorly; basimere longer than tall; basimere dorsomedian margin folded ventrally; basimere ventromedian margin dorsally curved, cupping the volsella; telomere arrowhead-shaped with a ventral membrane basally; setae of cuspis numerous, subequal in length to the apex of the telomere in ventral view; basal setose region of cuspis triangular; ventral margin of cuspis edentate, evenly curved; digitus long, thin, narrowing to apex which is evenly ventrally curved; valviceps of penisvalva strongly rounded with an apicoventral lobe; valvura apex subrectangular, narrow; ventral margin of penisvalva slightly concave to straight in basal half, otherwise straight and serrate, with teeth short and pointed anteriorly. Diagnosis Worker. Workers of this species are immediately differentiable from M. mayri by the absence of promesonotal spines. The following characters diagnose this species within the M. nanus group: posterolateral denticles close-set (PWI Gyne. Within the Malagasy Meranoplus fauna, gynes of M. radamae may be identified by the following characters: stellate setiferous punctures present on fourth abdominal tergum; propodeal spines thorn-like; smooth area behind eyes interrupted by rugae, or ruga behind eye terminates at ventrolateral mandibular condyle; eye length about �� length of malar area (OMI mean 75); never bicolored nor dark brown. Male. Males of M. radamae may be separated from those of M. mayri by the presence of the following characters: posterior margin of head not visible in full face view; small (WL 0.70���1.20); eyes large (EYE 76���81); maximum eye length greater than or equal to three times length of malar area (OMI 301���428); head comparatively small (CS 0.5���0.6); apex of penisvalva evenly rounded; penisvalva with apicoventral lobe; ventral margin of penisvalva serrate, without long spine-like teeth. Comments Meranoplus radamae is restricted to the grasslands and woodlands of the High Plateau, and the spiny forests of southern Madagascar. Specimens are known from 370���1550 m above sea level, and nests have been collected under stones in grassland and from ground nests. A few collections have been made in southern rainforest sites, but these may represent local adaptation to disturbed habitats. It is interesting to note that the southern rainforest collections of M. radamae, at R.S. Kalambatitra and Mount Vasiha, have reduced sculpturation on the face and promesonotal shield; these characters are represented in extreme form by the distinct M. sylvarius described here. Material examined MADAGASCAR: Antananarivo, Ambatolaona, 18 �� 55.68 ��� S, 47.52.97�� E, 1382 m, urban/garden, coll���n codes BLF 16883, BLF16899, 19Feb 2007 (B.L. Fisher et al.); Antananarivo, Angavokely, 18 �� 56 ��� S, 47 �� 45 ��� E, 10 Feb 1992 (B. Pettersson); 29.5 km WNW Tolanaro, Vasiha Mt., 24 �� 55 ��� 37 ��� S, 46 �� 44 ��� 49 ��� E, 300 m, sifted leaf litter in rainforest, 6���7 Feb 1995 (K.C. Emberton et al.); Antananarivo, Ankalalahana, 19.00659 �� S, 47.11220 �� E, 1375 m, Uapaca woodland, coll���n codes BLF 26350, BLF 26357, BLF 26360, BLF 26366, BLF 2637, BLF26391, 29��� 31 Mar 2010 (B.L. Fisher et al.); Antananarivo, Antsahadinta, 19 ��00.76��� S, 47 �� 27.40 ��� E, 1403 m, urban garden, coll���n codes BLF 17519, BLF17522, 8May 2007 (B.L. Fisher et al.); Antananarivo, Arivonimamo, Navoatra, 18.97799 �� S, 47.11929 �� E, 1373 m, Uapaca woodland, coll���n code ARA915, 6��� 8 May 2010 (A. Ravelomanana); Antananarivo, Manalalondo, Antaponimanadala, 19.25531 �� S, 47.17710 �� E, 1984 m, savanna grassland, coll���n code ARA1053, 13��� 16 May 2010 (A. Ravelomanana); Fianarantsoa, Ambositra, Antapia, 20.71971 �� S, 47.08685 �� E, 1495 m, Uapaca woodland, coll���n code ARA600, 3��� 5 Feb 2010 (A. Ravelomanana); Fianarantsoa, Ambositra, Mampiarika, 20.73456 �� S, 47.08355 �� E, 1480 m, Uapaca woodland, coll���n code ARA 468, Feb 2010 (A. Ravelomanana); Fianarantsoa, Ambositra, Mampiarika, 20.73568 �� S, 47.08384 �� E, 1464 m, Uapaca woodland, coll���n code ARA469, 1��� 3 Feb 2010 (A. Ravelomanana); Fianarantsoa, For��t Atsirakambiaty 7.6 km 285 �� WNW Itremo, 20 �� 35.6 ��� S, 46 �� 33.8 ��� E, 1550 m, under stone in grassland, coll���n codes BLF 7206, BLF7226, 22��� 26 Jan 2003 (B.L. Fisher et al.); Fianarantsoa, Itremo, Ampangabe, 20.6119 �� S, 46.60688 �� E, 1414 m, savanna woodland, coll���n code ARA791, 21��� 23 Mar 2010 (A. Ravelomanana); Fianarantsoa, Itremo, Antohatsahomby, 20.55436 �� S, 46.58438 �� E, 1640 m, Uapaca woodland, coll���n code ARA611, 18Mar 2010 (A. Ravelomanana); Mahajanga, Ambohimanga, 18 �� 45.67 ��� S, 47.33.86��� E, 1361 m, disturbed montane rainforest, coll���n code BLF17326, 26Apr 2007 (B.L. Fisher et al.); Toliara, R.S. Kalambatitra, 23.4185 �� S, 46.4583 �� E, 1365 m, grassland, coll���n codes BLF 21500, BLF 21538, BLF21535, 8Feb 2009 (B.L. Fisher et al.); Toliara, For��t Vohidava 88.9 km N Amboasary, 24 �� 14 ��� 26 ��� S, 46 �� 17 ��� 16 ��� E, 500 m, coll���n codes BLF 15846, BLF 15847, BLF 15849, BLF 15850, BLF 15692, BLF15738, 9Dec 2006 (B.L. Fisher et al.); Toliara, Vohibasia Forest, 59 km E Sakaraha, 22 �� 28 ��� S, 44 �� 51 ��� E, 750 m, sifted litter in tropical dry forest, coll���n code BLF1310, 13Jan 1996 (B.L. Fisher et al.)., Published as part of Boudinot, Brendon E. & Fisher, Brian L., 2013, A taxonomic revision of the Meranoplus F. Smith of Madagascar (Hymenoptera: Formicidae: Myrmicinae) with keys to species and diagnosis of the males, pp. 301-339 in Zootaxa 3635 (4) on pages 327-331, DOI: 10.11646/zootaxa.3635.4.1, http://zenodo.org/record/247717

Rhopalothrix subspatulata Longino & Boudinot, sp. nov. (Figs 1 D, 2 D, 3 B, 4, 12, 16) Type material. Holotype worker: COSTA RICA, Heredia: La Selva Biological Station, 10.43333 -84.01667 ± 1.5 km, 50 m, 1 Feb 1994, rainforest, ex Berlese of litter and soil (ALAS #B/05/ 381) [INBio, unique specimen identifier INBIOCRI001259541]. Paratypes (workers): same data as holotype [LACM, INBIOCRI001259538; MCZC, INBIOCRI001259539; USNM, INBIOCRI001259540; CAS, INBIOCRI001259542]; same data but 15 Apr 1993 (ALAS #B/05/053) [EAPZ, INBIOCRI002281441]. Geographic range. Costa Rica, Nicaragua. Diagnosis. Sharing with R. nubilosa and R. weberi a characteristic labrum shape: anterior margin of labrum with two long, bluntly triangular lobes, sinus between them deep, length of anterolateral lobe longer than or about equal to distance from base of sinus to transverse carina at base of labrum; worker hardly differing from R. weberi, being slightly larger (HW> 0.40) and with larger mandibular teeth; queen differing from R. weberi in stronger facial concavity and carina medial to compound eye; worker differing from R. nubilosa in smaller size ( Description. Worker. HW 0.42–0.49 (n= 14); mandible with two closely-spaced short triangular teeth at base, a smaller tooth about mid-distance between basal teeth and base of subapical tooth, reclinate denticle at base of subapical tooth absent, apical tooth short, about 1 / 3 × length of subapical tooth, intercalary teeth minute; labrum about as long as broad, with two long, bluntly triangular lobes, sinus between them deep, length of anterolateral lobe longer than or about equal to distance from base of sinus to transverse carina at base of labrum; erect setae on leading edge of scape stiff but narrow, hardly clavate, unlike the squamiform setae typical of many other species; arcuate promesonotal groove and metanotal groove moderately impressed; propodeal tooth right angled, infradental lamella thin, evenly and shallowly concave; first gastral tergite with four squamiform setae on posterior margin, two at mid-disk. Queen. HW 0.51–0.53 (n= 2); mandible and labrum similar to worker; face shape similar to worker but with large, shallow, circular concavity medial to compound eye, separated from eye by distinct slightly elevated carina that partially covers eye in full face view; compound eye longer than maximum width of scape; ocelli small, cuticle adjacent to ocelli marked with black pigment spots on evenly light brown background; shape of propodeal tooth, infradental lamella, petiole and postpetiole similar to worker; katepisternum and anepisternum large, convex, separated by thin groove; pubescence layer of abundant, short, curved setae covers mandible, face, scapes, legs, dorsal mesosoma and metasoma; abundant stiff erect setae on face, anterior edge of scape, side of head, dorsal mesosoma, dorsal gaster. Fore wing: pterostigma placed about 3 / 5 the length of the costal margin; veins Sc+R, M+Cu, 1 A, and crossvein 2 rs-m tubular; 2 rs-m terminating posteriorly as a node; veins Rs, M, and Cu nebulous; Rs+M spectral; vannal region vestigial; only basal and submarginal 1 cells enclosed by tubular or nebular veins. Hind wing: only veins C+Sc+R and 1 A tubular, and only for a very short length of the remigium; veins Sc+R, M+Cu, R, Rs and crossvein 2 rs-m nebulous; no cells are closed; vannal region vestigial. The male is unknown. Biology. This species occurs in lowland rainforest, from 50–800 m elevation. Of the approximately 50 workers known, one was collected in a Malaise trap sample and the rest were in Berlese or Winkler samples. The Winkler samples were of sifted litter and rotten wood on the forest floor; the Berlese samples were cores of forest floor litter and about 5–10 cm of the mineral soil beneath. At La Selva Biological Station, 3 % of 208 Berlese samples and 2 % of 640 miniWinkler samples contained workers. Given that each miniWinkler sample covered an area about 60 times greater than a Berlese sample (1 m 2 versus 165 cm 2), R. subspatulata was far more abundant in Berlese samples. This suggests that R. subspatulata, and perhaps Rhopalothrix in general, are subterranean, nesting and foraging in mineral soil, and perhaps only rarely venturing up into the litter layer. The pale color and greatly reduced eyes also suggest subterranean habits. Alate queens were collected in three of 50 canopy fogging samples from the ALAS project at La Selva (http:// viceroy.eeb.uconn.edu/ ALAS / ALAS.html): 8 October and 10 November 1994 and 28 December 1999 (mid to late wet season). This suggests that the species mates above ground and has normally dispersing queens. Etymology. Referring to the sparse spatulate setae on the gaster.

Rhopalothrix isthmica (Weber, 1941) (Figs 1 A, 2 C, 3 D, 8, 16) Geographic range. Panama, Honduras, Guatemala. Description. Worker. HW 0.58���0.68 (n= 13); mandible with three teeth on masticatory margin, middle tooth largest; subapical tooth with distinct reclinate denticle at base; subapical tooth about twice as long as apical tooth; intercalary teeth prominent, one closest to apical tooth about half as long as apical tooth; labrum trapezoidal, anterior lobes triangular, inner margins of lobes shallowly sloping to semicircular median notch; propodeal tooth acute to right-angled, infradental lamella evenly and shallowly concave; squamiform setae abundant on first gastral tergite, either uniformly covering entire tergite or covering varying extent of posterior portion, at least posterior half with abundant setae. The queen and male are unknown. Biology. This species occurs in moderately seasonal to aseasonal wet forest, from sea level to 2000 m elevation. All recent specimens are from Winkler or Berlese samples of sifted leaf litter. It is rare for ant species to occur across such a broad elevational spectrum, and given the considerable variability in setal pattern and wide geographic range, it may comprise multiple allopatric populations with unknown degrees of genetic divergence. In Honduras it was a rare cloud forest species at Comayagua and La Muralla and a rare lowland species along the north Caribbean coast. At La Union, in the mountains southeast of Zacapa, Guatemala, it was a rare species sympatric with the much more abundant R. megisthmica. In cloud forest on the slopes of Volc��n Atitl��n in western Guatemala it was abundant���occurring in 29 of 100 miniWinkler samples���and sympatric with the smaller R. atitlanica., Published as part of Longino, John T. & Boudinot, Brendon E., 2013, New species of Central American Rhopalothrix Mayr, 1870 (Hymenoptera, Formicidae), pp. 301-324 in Zootaxa 3616 (4) on pages 311-312, DOI: 10.11646/zootaxa.3616.4.1, http://zenodo.org/record/220287

Rhopalothrix apertor Longino & Boudinot, sp. nov. (Figs 1 F, 2 B, 3 A, 6, 16) Type material. Holotype, worker: COSTA RICA, Heredia: 7 km SW Pto Viejo, 10.40389 -84.03944 �� 500 m, 160 m, 4 Mar 2005, mature wet forest, ex sifted leaf litter (TEAM#AMI- 2 -W-033-01) [CAS, unique specimen identifier CASENT0629589]. Paratypes (workers): same data as holotype [USNM, CASENT0629588; MCZC, INB0003667720]. Geographic range. Costa Rica. Diagnosis. Masticatory margin of mandible dominated by a single, blunt, peg-like tooth; tooth at base of subapical tooth, instead of being the small reclinate denticle typical of other species, is a distinct recurved tooth, directed posteriorly; first gastral tergite largely devoid of setae, with one pair of squamiform setae at posterolateral margins; first gastral sternite with pronounced median keel, this keel weak to absent in other species. Description. Worker. HW 0.54���0.74 (n= 6); masticatory margin of mandible with single large blunt, in some almost capitate, tooth at about mid-length, a tiny denticle proximad, base of subapical tooth with prominent recurved acute tooth, directed posteriorly, subapical tooth shorter than width of mandible at base, about twice as long as apical tooth, only one intercalary tooth present, outer margin of mandible broadly flattened at base; labrum about as long as broad, with two long, bluntly rounded anterior lobes, sinus between them deep, length of anterolateral lobe longer than or about equal to distance from base of sinus to transverse carina at base of labrum; arcuate promesonotal groove and metanotal groove strongly impressed; propodeal tooth small, at about midlength of posterior face of propodeum, infradental lamella very narrow; first gastral tergite largely devoid of setae, with one pair of squamiform setae at posterolateral margins; first gastral sternite with pronounced median keel. The queen and male are unknown. Comments. Workers of this species fall into two distinct size classes. Five specimens have HW 0.54���0.58. Four of these are from La Selva Biological Station (50���150 m elevation), all from different samples, and one is from a 500 m elevation site on the Barva Transect above La Selva. A series of three specimens from one miniWinkler sample (and thus probably from the same colony) have HW 0.74, longer and relatively thinner mesotibiae, and a more robust flattened mandibular base. These are from immediately adjacent to La Selva, at 160 m elevation. There is the potential that they are two cryptic species. The holotype and paratype were chosen from the one series of larger workers. Biology. This species occurs in lowland rainforest, from 150���500 m elevation. All specimens are from Winkler samples of sifted leaf litter. It is rare: it occurred in three of 208 Project ALAS Berlese samples, and three of over 1500 TEAM project miniWinkler samples. Etymology. The mandible looks like a bottle opener., Published as part of Longino, John T. & Boudinot, Brendon E., 2013, New species of Central American Rhopalothrix Mayr, 1870 (Hymenoptera, Formicidae), pp. 301-324 in Zootaxa 3616 (4) on pages 309-310, DOI: 10.11646/zootaxa.3616.4.1, http://zenodo.org/record/220287

Rhopalothrix megisthmica Longino & Boudinot, sp. nov. (Figs 1 B, 2 C, 3 F, 9, 16) Type material. Holotype, worker. Guatemala, Zacapa: 2 km SE La Union, 14.94460 ���89.27726 �� 57 m, 1550 m, 12 May 2009, cloud forest, ex sifted leaf litter (LLAMA Wm-B- 03- 1 -05) [CAS, unique specimen identifier CASENT0612564]. Paratypes (workers): same data as holotype but 14.94665 -89.27593 �� 50 m (LLAMA#Wa-B- 03- 1 -04) [USNM, CASENT0614540]; 14.94677 -89.27585 �� 50 m (LLAMA#Wa-B- 03- 1 -07) [MCZC, CASENT0614546]; 14.94723 -89.27707 �� 50 m (LLAMA#Wa-B- 03- 1-45) [UNAM, CASENT0612503]; 14.95372 -89.27618 �� 50 m, 1430 m (LLAMA#Wa-B- 03- 2-31) [UVGC, CASENT0629572; ECOSCE, CASENT0629573]; 3.5 km SE La Union, 14.95 -89.27 (unknown error), 1500 m, 4 Jun 1991 (R.S. Anderson# RSA 91 -050) [LACM, CASENT0603567; EAPZ, CASENT0603699]. Geographic range. Guatemala, Mexico. Diagnosis. Differing from R. isthmica in larger size (HW> 0.73 versus Description. Worker. HW 0.73���0.83 (n= 12); mandible with two or three teeth on masticatory margin (can vary within individuals, with two teeth on one mandible and three teeth on the other), second tooth from base largest; subapical tooth with distinct reclinate denticle at base; subapical tooth about twice as long as apical tooth; intercalary teeth prominent, one closest to apical tooth about half as long as apical tooth; labrum trapezoidal, anterior lobes triangular, inner margins of lobes shallowly sloping to semicircular median notch; metanotal groove moderately to strongly impressed; propodeal tooth usually large and acute (shorter and obtuse on one specimen), infradental lamella evenly and shallowly concave; squamiform setae abundant on first gastral tergite, either uniformly covering entire tergite or covering at least 3 / 4 of posterior portion. The queen and male are unknown. Biology. This species occurs in cloud forest habitats, from 1400���2000 m elevation. All specimens are from Winkler or Berlese samples of sifted leaf litter. It is sympatric with R. isthmica at the type locality and with R. triumphalis on the slopes of Volc��n Tacan�� in Chiapas. At the type locality it was moderately abundant, occurring in 24 of 100 miniWinkler samples. Etymology. Referring to its similarity to R. isthmica, differing mainly in larger size., Published as part of Longino, John T. & Boudinot, Brendon E., 2013, New species of Central American Rhopalothrix Mayr, 1870 (Hymenoptera, Formicidae), pp. 301-324 in Zootaxa 3616 (4) on pages 313-314, DOI: 10.11646/zootaxa.3616.4.1, http://zenodo.org/record/220287

Rhopalothrix nubilosa Longino & Boudinot, sp. nov. (Figs 1 D, 2 D, 3 C, 10, 16) Type material. Holotype, worker: COSTA RICA, Heredia: 10 km NE Vara Blanca, 10.23696 -84.11983 �� 125 m, 1500 m, 9 Mar 2005, montane wet forest, second growth, vegetation only about 4 m high, ex sifted leaf litter (ALAS # 15 /WF/02) [CAS, unique specimen identifier CASENT0629594]. Paratypes (workers): same data as holotype [JTLC, CASENT0629593]; same data but 10.23684 -84.11909 �� 50 m, mature forest (ALAS # 15 /WF/02/ 09) [INBio, INB0003666702]; 10.23754 -84.12001 �� 50 m (ALAS # 15 /WF/02/ 43) [INBio, INB0003667096]; 10.23771 -84.11998 �� 50 m (ALAS # 15 /WF/02/ 47) [INBio, INB0003667132]; 10.23243 -84.11620 �� 50 m (ALAS # 15 /WF/04/ 15) [INBio, INB0003668070]; 10.23216 -84.11618 �� 50 m (ALAS # 15 /WF/04/ 21) [INBio, INB0003668120]. Geographic range. Costa Rica. Diagnosis. Sharing with R. subspatulata and R. weberi a characteristic labrum shape: anterior margin of labrum with two long, bluntly triangular lobes, sinus between them deep, length of anterolateral lobe longer than or about equal to distance from base of sinus to transverse carina at base of labrum; differing from both in larger size (HW = 0.57 versus R. weberi in subapical tooth longer than apical tooth (about same length in R. weberi); differing from R. subspatulata in larger number of squamiform setae on first gastral tergite (about 12 versus about 6). Description. Worker. HW 0.57 (n= 1); mandible with two closely-spaced short triangular teeth at base, a smaller tooth about mid-distance between basal teeth and base of subapical tooth, reclinate denticle at base of subapical tooth minute but present, apical tooth half the length of subapical tooth, intercalary teeth minute; labrum about as long as broad, with two long, bluntly triangular lobes, sinus between them deep, length of anterolateral lobe longer than or about equal to distance from base of sinus to transverse carina at base of labrum; erect setae on leading edge of scape moderately clavate, longest on basal bend, becoming shorter and thinner toward apex; arcuate promesonotal groove and metanotal groove moderately impressed; propodeal tooth right angled, infradental lamella evenly and shallowly concave; first gastral tergite with 6���8 squamiform setae on posterior margin, a similar number distributed between posterior border and midlength of tergite. The male is unknown. Biology. This species occurs in cloud forest, at 1500 m elevation. It is known from one site on the Barva Transect in Costa Rica, where it occurred in 5 of 200 miniWinkler samples. Comments. This is a slightly more robust version of R. subspatulata. Among several other species of Rhopalothrix considered in this report, this degree of difference would be considered intraspecific. However, the sharply parapatric distribution on the Barva transect and the low variability among the many lowland collections of R. subspatulata suggest a separate montane species. Etymology. Referring to the cloud forest habitat., Published as part of Longino, John T. & Boudinot, Brendon E., 2013, New species of Central American Rhopalothrix Mayr, 1870 (Hymenoptera, Formicidae), pp. 301-324 in Zootaxa 3616 (4) on pages 314-315, DOI: 10.11646/zootaxa.3616.4.1, http://zenodo.org/record/220287

Rhopalothrix weberi Brown & Kempf, 1960 (Figs 1 E, 2 E, 3 C, 15, 16) Rhopalothrix weberi Brown & Kempf, 1960: 234. Holotype, worker: Cuba, Las Villas: Casa Harvard near Central Soledad, 24 July 1933, in red clay from under stones and grass roots which had been run through a Berlese funnel (N.A. Weber) [MCZC] (not examined). Replacement name for Heptastruma wheeleri Weber, 1934: 55. [Junior secondary homonym of Rhopalothrix wheeleri Mann, 1922: 43.]. Geographic range. Cuba, Honduras to southern Mexico. Description. Worker (based on two workers from Tikal National Park, Guatemala). HW 0.38���0.39 (n= 2); mandible with two closely-spaced, very short teeth at base, these basal teeth with broad confluent bases, either with distinct denticulate apices or completely confluent and forming a low lamina at base, a small denticle about middistance between basal teeth and base of subapical tooth, reclinate denticle at base of subapical tooth absent, subapical tooth shorter than width of mandible at base, only slightly longer than subapical tooth, intercalary teeth present but minute; labrum about as long as broad, with two long, bluntly triangular lobes, sinus between them deep, length of anterolateral lobe longer than or about equal to distance from base of sinus to transverse carina at base of labrum; anterior clypeal margin concave; erect setae on leading edge of scape stiff but narrow, hardly clavate, unlike the squamiform setae typical of many other species; arcuate promesonotal groove and metanotal groove weakly impressed; propodeal tooth right angled, infradental lamella thin, evenly and shallowly concave; first gastral tergite with sparse squamiform setae on posterior half, one specimen with five setae on posterior margin and three anteriorly, one specimen with seven and five, respectively. Queen. HW 0.40���0.54 (n= 5); mandible and labrum similar to worker; face shape similar to worker but with grooves and ridges more shallowly impressed; compound eye shorter than maximum width of scape, with 5���6 facets across longest axis; ocelli small, cuticle adjacent to ocelli marked with black pigment spots on evenly light brown background; shape of propodeal tooth, petiole and postpetiole similar to worker; shape of infradental lamella variable (see Comments); katepisternum and anepisternum large, convex, separated by broad groove; pubescence layer of abundant, short, curved setae covers mandible, face, scapes, legs, dorsal mesosoma and metasoma; abundant stiff erect setae on face, anterior edge of scape, side of head, dorsal mesosoma, dorsal gaster. The male is unknown. Biology. This species occurs in lowland wet to moist forest, from 200���575 m elevation. The recent collections are all from Winkler samples of sifted litter and rotten wood on the forest floor. Among the Project LLAMA specimens, seven are dealate queens and only two are workers, mostly from separate samples. This could reflect subterranean habits of the workers, with new queens dispersing up into the litter layer. Comments. Judging from the descriptions of Weber (1934) and Brown & Kempf (1960), and the geographic proximity of the Pet��n region to Cuba, we tentatively assign to R. weberi material from eastern Chiapas, across the Pet��n region, and south to north coastal Honduras. Quiroz-Robledo & Valenzuela-Gonz��lez (2010) identified a worker from Veracruz as R. weberi, which, based on the figure in the publication, also appears to belong here. Four queens from Tikal National Park and vicinity are very uniform, with HW 0.40���0.41 and the propodeal infradental lamella thin and very like the workers from this locality. One queen from near La Ceiba in Honduras has HW 0.44 and the infradental lamella is more expanded, forming two convexities below the tooth, separated by a narrow notch. Two queens from the Metzabok Reserve in eastern Chiapas are distinctly different. One of them has HW 0.44 and an infradental lamella like the queens from Guatemala. The other queen has HW 0.54 and infradental lamella like the queen from Honduras. Thus there is the potential for multiple cryptic species in R. weberi. LaPolla et al. (2006) report R. weberi from Guyana, but this identification is incorrect. The single specimen (examined) cannot be clearly assigned to genus: it has the habitus of the R. isthmica clade, but the mandible and labrum are like species in the genus Octostruma Forel, 1912., Published as part of Longino, John T. & Boudinot, Brendon E., 2013, New species of Central American Rhopalothrix Mayr, 1870 (Hymenoptera, Formicidae), pp. 301-324 in Zootaxa 3616 (4) on pages 321-323, DOI: 10.11646/zootaxa.3616.4.1, http://zenodo.org/record/220287