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Leptanilloides sp. ���Male 4��� (unassociated to workers) Figs 8 A���C, 9A���B, 10A���B Material examined ECUADOR: 1 ♂, Zamora-Chinchipe Prov., Reserva Biol��gica San Francisco, entrance of the T1 trail, 1920 m, 3��58���44.09��� S, 79��05���8.26��� W, 16���29 Mar. 2011, Malaise trap, coll. Thibaut Delsinne and Tania Milena Arias-Penna (QCAZ, sc 4870101). GenBank accession number: KT601704. Measurements (in mm) and indices HW 0.34, HL 0.31, SL 0.24, LAII 0.07, LAIII 0.09, LAIV 0.09, LAXIII 0.17, EL 0.13, MaL 0.16, PrW 0.33, WL 0.66, MH 0.36, PL 0.12, PW 0.11, PPL 0.33, PPW 0.33, AivL 0.13, AivW 0.33, FFeL 0.36, FFeW 0.06, HFeL 0.49, HTL 0.4, CI 110.71, SI 70.97, PI 90.91, MI 54.24. Description HEAD. Broader than long, with large convex eye that occupies anterior half of side of head. Mandible slender, falcate, twisted at apex, overlapping at closure (Fig. 8A). Basal and masticatory margins strongly concave, separated by well-developed tooth. External margin straight along basal and medial lengths but strongly bent at apex. Mandible longer than eye length. Lateroclypeal tooth and hypostomal tooth lacking, clypeus short and transverse, with narrow clypeal lamella (apron). Antennal socket horizontal and exposed, located at anterior clypeal margin, margin not projecting anteriorly beyond ventral articulation with labrum. Antenna 13-segmented, each segment longer than wide, with second segment shortest. Scape of moderate length, 1.5 times longer than length of ultimate antennal segment. Scape length nearly four times length of second antennal segment. Lateral ocellus separated from median ocellus by its diameter. Distance between lateral ocelli similar to distance between median and lateral ocellus; as a result, ocelli forming equilateral triangle. MESOSOMA. Pronotum U-shaped in dorsal view and reduced anteromedially to thin horizontal strip, set below level of dorsally protruding mesonotum and triangular in lateral view, with pointed posterior apex directed towards wing base. Mesoscutum lacking notaulus. Parapsidal line not discerned, apparently absent. Axillae depressed, not meeting medially, connected by narrow furrow; tegula very small and inconspicuous. Mesopleuron lacking oblique transverse sulcus and hence not divided into anepisternum and katepisternum. Transcutal sulcus deeply impressed. Mesoscutum and mesoscutellum prominently bulging, in lateral view. Metapleural gland not discernable. Propodeum rounded in profile, with dorsal and declivous faces only poorly differentiated; dorsal surface somewhat shorter than declivous. Propodeal spiracle small, circular, positioned slightly above midheight of propodeum and slightly posterior to metanotum. Leg slender, mid tibia with one simple and hind tibia with one simple spur (observed under 50�� magnification), pretarsal claw lacking preapical tooth. WINGS (Fig. 9 A���B). With relatively well-developed venation (for Leptanilloides). Forewing with C present, tubular and pigmented. Sc+R approximated to wing margin, very narrow. Sc+R 1 in line with Sc+R, tubular. Pterostigma well-marked. M+Cu tubular and pigmented, curved towards posterior wing margin before division. Rs��f1 nebulous. M��f1 pigmented, tubular. Rs+M, Rs��f2, and Rs��f3 all joined, tubular and pigmented. 1r-rs absent. 2r-rs present, tubular and pigmented. Rs��f4 and Rs��f5 joined and not differentiated in absence of 2rs-m. Rs��f4&Rs��f5 partly tubular, then nebulous, terminating before wing margin. M and Cu diverging at cu-a. Free abscissa of M partly nebulous, then spectral, very weakly visible, joining to Rs+M&Rs��f2&Rs��f3. Abscissa of Cu joined, nebulous throughout most of length and continuing as spectral. Vein A tubular, joining cu-a at obtuse angle and confluent with Rs+M, apparently absent beyond cu-a. Posterior margin of fore wing with fold where hamuli attach, narrow, conspicuous. Hindwing with C present, narrow and faint. Anterior hindwing margin with small differentiated pigmentation area located in distal half of wing. Three hamuli originate in pigmented region. Sc+R present, nebulous, almost reaching third of wing length. Sc+R1 absent. Rs��f1&Rs��f2 partly nebulous, then spectral. Jugal lobe absent. METASOMA. Slender in lateral view, obovate in dorsal view, widest at abdominal segment V. Petiole (abdominal segment II) longer than high or wide, rectangular in dorsal view, with convex lateral margin, ovate in lateral view, and weakly constricted posteriorly, helcium thus apparently quite broad. Petiolar spiracle located on anterior third of segment, near anterodorsal extremity. Abdominal segment III three times larger than petiole at its maximum width, not developed as postpetiole nor separated from abdominal segment IV by marked constriction. Abdominal spiracle III located on anterior third of tergite. Abdominal segments II and III with tergosternal fusion. Abdominal segment IV and succeeding segments lacking tergosternal fusion. Spiracle present on anterior third of tergite IV. Abdominal segments V and VI not separated from succeeding segments by constrictions. Abdominal spiracles V and VI indiscernible. Abdominal tergite VIII (pygidium) small and simple but visible dorsally, not wholly covered by abdominal tergite VII. GENITALIA (Fig. 10 A���B). Pygostyle absent. Abdominal sternite IX (subgenital plate) with posterior margin broadly and deeply concave but not bifurcate. Basal ring present, not hypertrophied. Paramere relatively small, harpago rounded at apex; paramere shorter than petiole length. Volsella a simple, broad lobe, lacking differentiated cuspis. Aedeagus subequal in length to paramere and volsella, simple, narrow, distally spatulate. PILOSITY AND SCULPTURE. Integument mostly smooth and shiny, with scattered piligerous punctures. Pilosity common on most of body, suberect to decumbent. Color light brown, head and metasoma past abdominal segment III slightly darker. Appendages (antennae, mandibles, legs) lighter than body. Distribution Only known from the sampling locality. Remarks The male (specimen code 4870101) collected in southern Ecuador is easily distinguished from any other leptanilloid males (either of Leptanilloides or Amyrmex species, males of all species of Asphinctanilloides being unknown) by the unique shape of its mandible. We did not give it a specific name because it is not associated with workers. However, we called it ���Male 4��� to indicate that it is different from the three males (���Male 1��� to ���Male 3���) described in Borowiec & Longino (2011), which are also without specific names. The wing venation of ���Male 4��� is relatively similar to that of ���Male 3��� in Borowiec & Longino (2011), except that the free abscissa of M on the forewing joins Rs+M&Rs��f2&Rs��f3 (not joining in ���Male 3���) and Sc+R1 is not visible on the hindwing (present as a short nebulous stub in ���Male 3���). ���Males 1 and 2��� of Borowiec & Longino (2011) have more reduced venation, as do the males of Amyrmex (Ward & Brady 2009). The forewing venation of L. mckennae and L. nubecula is also closely similar but, in these species, veins M and Cu diverge distal to crossvein cu-a (diverging at cu-a in ���Male 4���). Table 1 lists informative characters to separate leptanilloid males. DNA results The COI sequences of the two new species and ���Male 4��� differ significantly from each other (p-distances ranging from 18.8 to 21.7%) (Fig. 11). They also diverge (18���21.8%) from the other Leptanilloides DNA barcodes available in BOLD (6 specimens, 2 species: L. gracilis from the L. biconstricta speciesgroup and L. nubecula from the L. legionaria species-group). Our phylogenetic analyses constructed on the basis of the nuclear gene Wg support one clade consisting of L. prometea sp. nov., L. femoralis, L. gracilis and Amyrmex sp. and another clade composed of L. nubecula, L. nomada and L. mckennae. Both clades are supported by bootstrap values of 98% and 99% in the neighbour-joining tree, 97% and 100% in the most parsimonious tree, 83% and 100% in the tree with maximum likelihood and by posterior probabilities of 1 in the tree built with Bayesian inference (Fig. 12)., Published as part of Delsinne, Thibaut, Sonet, Gontran & Donoso, David A., 2015, Two new species of Leptanilloides Mann, 1823 (Formicidae: Dorylinae) from the Andes of southern Ecuador, pp. 1-35 in European Journal of Taxonomy 143 on pages 18-22, DOI: 10.5852/ejt.2015.143, http://zenodo.org/record/3787360, {"references":["Borowiec M. L. & Longino J. T. 2011. Three new species and reassessment of the rare Neotropical ant genus Leptanilloides (Hymenoptera, Formicidae, Leptanilloidinae). ZooKeys 133: 19 - 48. http: // dx. doi. org / 10.3897 / zookeys. 133.1479","Ward P. S & Brady S. G. 2009. Rediscovery of the ant genus Amyrmex Kusnezov (Hymenoptera: Formicidae) and its transfer from Dolichoderinae to Leptanilloidinae. Zootaxa 2063: 46 - 54."]}

Leptanilloides prometea Delsinne & Donoso sp. nov. urn:lsid:zoobank.org:act: CC756523-8025-4C13-8987-5F91A1E04B7E Figs 4 A–D, 5, 6A–F, 7A–C Diagnosis Leptanilloides prometea belongs to the L. biconstricta species-group and can be distinguished by the combination of the following characters: masticatory margin of mandible dentate but teeth extremely minute, blunt and irregularly shaped (Fig. 6C); lateroclypeal tooth present (Fig. 6C); head with piligerous foveolae separated by smooth interspaces equaling, on average, puncture diameters; fine, uninterrupted reticulation on mesopleuron, metapleuron and lateral side of petiole; flange over metapleural gland opening forming a short, blunt projection (Fig. 6D); and subpetiolar process relatively straight (not distinctly bulging), without posterior angle (Fig. 6E). Etymology Name in apposition, in honour of the Prometeo initiative of the Ecuadorian government that seeks to strengthen research and knowledge transference by getting national and international experts to work together. This project is aligned to the Ecuadorian “National Development Plan for Good Living”, which has the objective, among others, of guaranteeing the rights of Nature and promoting a healthy and sustainable environment. Furthermore, the species is the largest of the L. biconstricta species-group and its name nicely reminds us of ‘Prometheus’, the Titan in Greek mythology who brought fire, a symbol of enlightenment, to mankind. We hope that this new ant species will symbolize the promise of a bright future for Ecuadorian biodiversity. Type material Holotype ECUADOR: Worker, Zamora-Chinchipe Prov., Reserva Biológica San Francisco (RBSF), 2070 m, 3°58’ S, 79°05’ W, 13 May 2010, within 0.5 m 2 of leaf litter extracted with a mini-Winkler apparatus for 96 h, coll. Thibaut Delsinne and Tania Milena Arias-Penna (QCAZ, sc 4267803). Paratypes (n = 67) ECUADOR: Same data as holotype except that specimens were captured after 48 h of Winkler extraction: 30 workers, 96% ethanol (RBINS, General Inventory Number 33044: sc 4052302 [1 specimen with DNA data], sc 4052314 [1 specimen with DNA data], sc 4052316 [21 specimens without DNA data]; BMNH [1 specimen, sc 4052317]; CASC [1 specimen, sc 4052318]; ICN [1 specimen, sc 4052319]; JTLC [1 specimen, sc 4052320]; MCZC [1 specimen, sc 4052321]; MZSP [1 specimen, sc 4052322]; QCAZ [1 specimen, sc 4052323]); 3 workers, pinned (MUTPL, sc 4052310, 4052315; RBINS, sc 4052313); 2 workers, 96% ethanol after non-destructive DNA extraction (RBINS, sc 4052311, 4052312); 1 worker, gold-coated for SEM (RBINS, sc 4052301). – Same data as holotype except 21 Mar. 2010, within upper 5 cm of organic layer of a core (Ø 5 cm) extracted by heat using a modified high gradient extractor for 4 days: 28 workers, 96% ethanol (RBINS, sc 4060601); 2 workers, 96% ethanol after non-destructive DNA extraction (RBINS, sc 4060602, 4060603); 1 worker, pinned (MUTPL, sc 4060604). GenBank accession numbers: KT601698 – KT601703 and KT750331. Measurements (in mm) and indices Holotype (3 paratypes): HW 0.4 (0.4–0.44), HL 0.58 (0.61–0.62), SL 0.27 (0.29), PrW 0.27 (0.29), WL 0.71 (0.76–0.78), MH 0.23 (0.24), PL 0.22 (0.22–0.23), PW 0.11 (0.12), PPL 0.22 (0.22–0.23), PPW 0.2 (0.22–0.23), AivL 0.31 (0.31–0.33), AivW 0.37 (0.38–0.4), FFeL 0.38 (0.38–0.4), FFeW 0.11 (0.11), HFeL 0.44 (0.44–0.45), HTL 0.42 (0.44–0.46), CI 69.23 (65.45–71.43), SI 66.67 (66.25–72.22), PI 50.00 (52.5–53.66), MI 32.81 (31.43–32.36). Description Worker With the characters typical of the genus (see diagnosis and description in Brandão et al. 1999 and Borowiec & Longino 2011) and of the L. biconstricta species-group (see diagnosis of the group above). Other characters or differences are as follows: HEAD. Elongate and rectangular with lateral margin nearly straight and parallel. Posterior corner rounded. Posterior margin modestly convex, almost straight. Parafrontal ridge absent. Clypeal lamella strongly convex (Fig. 6C). Masticatory margin of mandible dentate but teeth extremely minute, blunt and irregularly shaped, difficult to discern even under magnifications of about 100× (Fig. 6C). Basal and masticatory margins united by a broad convexity. Basal margin faintly crenulate. Labial palp not readily visible but, at least, one-segmented, maxillary palp two-segmented. Hypostomal anterior border rounded and slightly projected outward but without forming a distinct tooth (Fig. 6A). Scape when laid back reaching about medial distance to posterior margin of head. MESOSOMA. Flange over metapleural gland opening conspicuous and posteriorly forming short, blunt projection, not surpassing propodeum declivity margin (Fig. 6D). Femur not conspicuously enlarged, relatively slender. Midtibia with one short (i.e., half size of foretibia strigil), pectinate spur (Fig. 6B), although pectination may be difficult to observe even at high magnification. Hindtibia with one broadly pectinate spur roughly as long as strigil. Metatibial gland present, visible at high magnification and good lighting as translucent oval area at apex of tibia, behind spur insertion (Fig. 4D). Metatibial gland pore plate observable in SEM (Fig. 7 A–C). METASOMA. Long and relatively slender. In dorsal view, petiole uniformly rectangular, twice as long as wide, as long as abdominal segment III (postpetiole). Anterior face concave, posterior face straight. In lateral view, petiole height approximately ¼ smaller than height of abdominal segment III. Petiolar tergite dome-shaped, with short and poorly differentiated anterior and posterior faces, maximum height situated in posterior half. Short tubulated portion present posteriorly. Petiolar spiracle inconspicuous, not in excavation, set near anterior rim of tergite, similar in form and slightly smaller in diameter than propodeal spiracle. Subpetiolar process with ventral margin relatively straight (not distinctly bulging), and without posterior angle (Fig. 6E). Anterior projection of subpetiolar process variable in shape, acute to rounded (Figs 5, 6E). Maximum height of petiolar sternite situated in its anterior half. In dorsal view, abdominal segment III (postpetiole) trapezoid, with straight, parallel anterior and posterior faces. Posterior face almost twice as long as anterior face. In lateral view, tergite evenly convex, without well-differentiated posterior face. Sternite evenly rounded, slightly bulging anteriorly. In dorsal view, abdominal segments IV–VI subequal in length. PILOSITY AND SCULPTURE. Mandible smooth and shiny, with few scattered piligerous punctures. Head with abundant deep piligerous punctures and smooth interspaces on average equaling puncture diameter, except on ventral side and front where punctures are sparser, separated by more than their diameter. Mesosoma and abdomen more finely and sparsely punctate. Fine reticulate sculpture present laterally on lower pronotum, entire mesopleuron, propodeum, and petiole. Body and appendages with abundant, short and subdecumbent to suberect hairs. Body color brownish to reddish, with head and mesosoma tending to be darker than petiole and gaster. Legs and antennae yellowish. Gyne Unknown. Male Unknown. Distribution Only known from the type locality. Biology Unknown. All the specimens were collected from a single reserve bordering the Podocarpus National Park on the eastern Andean slope of southern Ecuador. The habitat is an evergreen lower montane forest (Homeier et al. 2008) which is in nearly pristine condition. The forest harbours more than 300 tree species, with Lauraceae, Melastomataceae and Rubiaceae being the most species-rich families (Homeier et al. 2012). The mean annual temperature is c. 15 °C and mean annual precipitation is c. 2200 mm, with low seasonality (Bendix et al. 2008). Soils of the sampling area are cambisols, with a very thick (often> 50 cm) leaf litter layer (Homeier et al. 2012). Soil texture is sandy silt loam; proportion of sand, silt and clay is 41%, 52%, and 6%, respectively; mean pH = 3.2 ± 0.1 SD (n = 24 soil samples). Remarks Leptanilloides prometea sp. nov. is the largest species of the L. biconstricta species-group (Fig. 1). The most similar species in habitus and size is L. improvisa, but head of L. prometea sp. nov. is longer (Fig. 1) and its mandible has only minute and irregular teeth along the masticatory margin (teeth are conspicuous and regularly spaced in L. improvisa). Moreover, L. prometea sp. nov. can easily be distinguished from other species in the group by: the presence of a conspicuous lateroclypeal tooth (absent in L. caracola), the evenly rounded sternite of abdominal segment III (bulging anteriorly in L. biconstricta), the petiole being as long as the postpetiole (petiole longer than postpetiole in L. biconstricta), the petiolar sternite being higher in its anterior half (bulging medially in L. gracilis and L. femoralis), the head dorsum with foveolae separated by about their diameter (more densely foveolate in L. sculpturata, with foveolae separated by less than their diameter), and the short, blunt flange over the metapleural gland opening (long and sharply pointed in L. atlantica, L. copalinga and L. erinys).

Leptanilloides copalinga Delsinne & Donoso sp. nov. urn:lsid:zoobank.org:act: 4324B13A-500B-464C-AE3F-62BCAF95A2CC Figs 2 A���F, 3A���F Diagnosis Leptanilloides copalinga sp. nov. belongs to the L. biconstricta species-group. It can be distinguished from other species of this group by the combination of the following characters: masticatory margin of mandible feebly dentate, with teeth extremely minute, blunt and irregularly shaped (Fig. 3A); lateroclypeal tooth present (difficult to observe in stereo microscopy even at high magnification due to the small size of the species but obvious in SEM; Fig 3A); head with piligerous foveolae separated by smooth interspaces equaling 2���3 diameters; reticulation on mesopleuron, metapleuron and lateral side of petiole superficial and interrupted; flange over metapleural gland opening conspicuous and sharply pointed posteriorly, resulting in a projection which surpasses propodeum declivity margin in lateral view (Fig. 3B); petiole slender in dorsal view, 52 ��� PI ��� 63 (Fig. 3D); and subpetiolar process straight (not distinctly bulging), with a rounded projection anteriorly (Fig. 3C). Etymology Name in apposition which refers to the Copalinga Private Reserve, the type locality, in honour of its owners Catherine Vits and Boudewijn de Roover to acknowledge their efforts for the conservation of Ecuadorian biodiversity. Type material Holotype ECUADOR: Worker, Zamora-Chinchipe Prov., Copalinga Private Reserve, 1510 m, 4��4���56.6��� S, 78��58���5.71��� W, 2 Apr. 2010, soil sample, coll. Thibaut Delsinne and Tania Milena Arias-Penna (QCAZ, sc 4006304). Paratypes ECUADOR: Same data as holotype: 1 worker, gold-coated for SEM (RBINS, sc 4006301); 1 pinned (MUTPL, sc 4006303); 1 worker, 96% ethanol (RBINS, sc 4006302, specimen with DNA data). GenBank accession number: KT601697. Measurements (in mm) and indices Holotype (paratype 4006303): HW 0.33 (0.34), HL 0.44 (0.44), SL 0.23 (0.21), PrW 0.2 (0.22), WL 0.56 (0.57), MH 0.17 (0.17), PL 0.17 (0.18), PW 0.09 (0.11), PPL 0.16 (0.18), PPW 0.16 (0.18), AivL 0.26 (0.26), AivW 0.3 (0.33), FFeL 0.23 (0.27), FFeW 0.09 (0.11), HFeL 0.29 (0.29), HTL 0.3 (0.29), CI 75 (76.25), SI 70 (60.66), PI 51.61 (62.5), MI 30 (30.39). Description Worker With the characters typical of the genus (see diagnosis and description in Brand��o et al. 1999 and Borowiec & Longino 2011) and of the L. biconstricta species-group (see diagnosis of the group above). Other characters or differences are as follows: HEAD. Elongate and rectangular with lateral margin nearly straight and parallel. Posterior corner rounded. Posterior margin modestly convex, almost straight. Parafrontal ridge absent. Clypeal lamella strongly convex (Fig. 3A). Basal and masticatory margins of mandible dentate but teeth extremely minute, blunt and irregularly shaped, difficult to discern even under magnifications of about 100�� (Fig. 3A). Basal and masticatory margins united by a broad convexity. Basal margin faintly crenulate. Labial and maxillary palps difficult to discern in situ but formula apparently 2,2. Hypostomal anterior border without distinct tooth. Scape when laid back failing to reach medial distance to posterior margin of head by nearly one maximum diameter. MESOSOMA. Flange over metapleural gland opening conspicuous and sharply pointed posteriorly, forming a projection which surpasses propodeum declivity margin (Fig. 3B). Femur not conspicuously enlarged, relatively slender. Midtibia with one short (i.e., half size of foretibia strigil), pectinate spur (Fig. 3F), although pectination may be difficult to observe even at high magnification. Hindtibia with one broadly pectinate spur roughly as long as strigil. Metatibial gland absent or very reduced and not visible even in SEM. METASOMA. Long and relatively slender. In dorsal view, petiole rectangular with lateral margin slightly convex, twice as long as wide (Fig. 3D), and as long as abdominal segment III (postpetiole). Anterior face concave, posterior face straight. In lateral view, petiole height approximately �� smaller than height of abdominal segment III. Petiolar tergite dome-shaped, with short anterior and posterior faces, maximum height situated in posterior half. Short tubulated portion present posteriorly. Petiolar spiracle inconspicuous, not in excavation, set near anterior rim of tergite, similar in form and slightly smaller in diameter than propodeal spiracle. Subpetiolar process with ventral margin straight to slightly concave, not distinctly bulging, anteriorly forming rounded projection (Fig. 3C). Maximum height of petiolar sternite situated medially. In dorsal view, abdominal segment III (postpetiole) trapezoid, with straight, parallel anterior and posterior faces. Posterior face twice as long as anterior face. In lateral view, tergite evenly convex, without well-differentiated posterior face. Sternite evenly rounded. In dorsal view, abdominal segments IV���VI subequal in length. PILOSITY AND SCULPTURE. Mandible smooth and shiny, with few, scattered piligerous punctures. Head with abundant deep piligerous foveolae and smooth interspaces on average equaling two or three puncture diameters. Mesosoma and abdomen more finely and sparsely punctate. Fine reticulate sculpture present laterally on lower pronotum, mesopleuron, propodeum and petiole. Dorsal part of pronotum, mesopleuron, propodeum and petiole smooth and shiny. Body and appendages with abundant, short and subdecumbent to suberect hairs. Body color yellow to reddish (Fig. 2), with head and mesosoma darker than petiole and gaster. Legs and antennae yellowish. Gyne Unknown. Male Unknown. Distribution Only known from the type locality. Biology Unknown. The type series was found by visual search in a 15��15�� 15 cm soil core inspected for ants during 20 person-minutes (the soil content was examined over a white plastic board using a headlamp to facilitate ant detection), which suggests subterranean habits as observed in other Leptanilloides species. The habitat is a relatively well-preserved evergreen lower montane forest. Soil texture is clay loam; proportion of sand, silt and clay is 44%, 22%, and 35%, respectively; mean pH = 4 �� 0.2 SD (n = 24 soil samples). Remarks This species belongs to the L. biconstricta species-group. The most similar species are L. biconstricta and L. atlantica, which also possess a long and sharply pointed flange over the metapleural gland opening and a straight (not bulging) petiolar sternite with a rounded anterior projection. However, these two species have a deeper constriction between the postpetiole and abdominal segment IV. Besides, L. copalinga sp. nov. has a petiole roughly as long as the postpetiole (longer in L. biconstricta) and its postpetiolar sternite is evenly rounded (distinctly bulging anteriorly in L. biconstricta). Finally, L. copalinga sp. nov. is larger than L. atlantica (Fig. 1). The other species of the L. biconstricta speciesgroup are easily distinguished from L. copalinga sp. nov. by, among other characters, the shape of their petiolar sternite (bulging medially in L. gracilis and L. femoralis; bulging anteriorly in L. erinys), the shape of the flange over the metapleural gland opening (short and rounded posteriorly in L. improvisa and L. prometea), the head sculpture (more densely foveolate in L. sculpturata) and the lateroclypeal tooth (absent in L. caracola)., Published as part of Delsinne, Thibaut, Sonet, Gontran & Donoso, David A., 2015, Two new species of Leptanilloides Mann, 1823 (Formicidae: Dorylinae) from the Andes of southern Ecuador, pp. 1-35 in European Journal of Taxonomy 143 on pages 8-12, DOI: 10.5852/ejt.2015.143, http://zenodo.org/record/3787360, {"references":["Brandao C. R. F., Diniz J. L. M., Agosti D. & Delabie J. H. C. 1999. Revision of the Neotropical ant subfamily Leptanilloidinae. Systematic Entomology 24: 17 - 36. http: // dx. doi. org / 10.1046 / j. 1365 - 3113.1999.00064. x","Borowiec M. L. & Longino J. T. 2011. Three new species and reassessment of the rare Neotropical ant genus Leptanilloides (Hymenoptera, Formicidae, Leptanilloidinae). ZooKeys 133: 19 - 48. http: // dx. doi. org / 10.3897 / zookeys. 133.1479"]}

Genus Tatuidris Tatuidris Brown and Kempf 1968: 183. Type species: Tatuidris tatusia Brown and Kempf 1968: 187, by monotypy. Worker: HW = 0.56–1.10mm. Body short and compact. Color ferruginous to dark red. Integument thick and rigid. Body covered by hairs, which are variable in length and inclination. HEAD. Head shape pyriform, broadest behind. Maxillary palps one-jointed. Labial palps two-jointed. Labrum bilobed, broader than longer, capable of full reflexion over the buccal cavity. Mandibles opposing in most of their border (except in the tips of the masticatory margin). Masticatory margin with two blunt apical teeth overlapping at closure. Setae (mandibular brush) abundant, present on the ventral side of mandibles. Antenna 7 -segmented. Antennal club two segmented, well developed. Scape clavate, gently downcurved at base. Torulus with hypertrophied dorsal lobe and strongly curved downwards. Antennal scrobe present. Antennal socket and antennal scrobe confluent. Antenna socket apparatus sitting upside down on the roof of the expanded frontal lobe. Eyes present, small (REL= 5.41–11.48), located at the posterior apex of antennal scrobe. MESOSOMA. Promesonotal suture fused. Metapleural gland orifice round. Metapleural gland opening visible. Metapleural gland bulla separated from annulus of propodeal spiracle by more than the diameter of the spiracle. Katepisternal oblique groove absent. Lower mesopleuron with longitudinal costulae. Propodeum unarmed. Propodeal spiracle, in profile, located at about mid-length of sclerite. PETIOLE and POSTPETIOLE. Petiole short and sessile. Petiolar ventral process large and rounded. Petiole dorso-ventrally fused. Petiole broadly attached to postpetiole (abdominal segment III). Postpetiolar tergum and sternum overlapping at junction. Postpetiole in dorsal view wider in posterior half. GASTER. Articulation between postpetiole and gastral segment 1 (abdominal segment IV) broad. Postpetiolar postsclerites not set in a concavity or depression. Pretergite of first gastral segment with neck-like constriction. Stridulitrum present on first gastral segment. Limbus (i.e. anterior transverse cuticular ridge of the first gastral segment) absent. Suture between first gastral tergite and sternite anteriorly rounded. First gastral tergosternal suture strong, but not fused. Base of the first gastral sternum in profile rounded. First gastral sternite length is reduced, such that tergite is much larger than the sternite and strongly vaulted. First gastral tergum and sternum smooth or with scattered puncta. Sting present. LEGS. Mid and hind tibial spurs present. Gyne: HW = 1.28mm. Body short and compact, with exterior morphology and characters similar to workers. Body covered by hairs. Color yellow, paler than workers. Integument thick and rigid. HEAD. Head shape pyriform, broadest behind. Vertex straight, not concave. Labrum bilobed, broader than long, capable of full reflexion over the buccal cavity. Mandibles opposing in most of their border, except in the tips of the masticatory margin. Masticatory margin with two blunt apical teeth overlapping at closure. Mandibular setae present but less abundant than in workers. Antennal joints 7 -segmented. Antennal club two segmented, well developed. Scape clavate, gently downcurved at base. Antennal scrobe present. Antennal socket and antennal scrobe confluent. Antenna socket apparatus sitting upside down on the roof of the expanded frontal lobe. Eyes present, EL = 0.20mm, eyes larger than in workers (REL = 22.63), located laterally at posterior border of antennal scrobes. Lateral ocelli and median ocellus present. WINGS. WingL= 4.60mm, about 60 % longer than total body length. Forewing well developed, with costal cell, basal cell (radial), sub-basal cell (cubital), no vein present between sub-marginal cell 1 and sub-marginal cell 2, R 1 vein surrounding sub-marginal cell 3, discal cell 1 and discal cell 2 present, divided by cubital vein which extends a distance similar to the inferior edge of discal cell. Hindwing well developed, with Cu-a vein present. Basal cell completely surrounded by M-Cu and and rs-m+M veins. MESOSOMA. Promesonotal suture present, not fused. Scutellum broad. Anepisternum and katepisternum broad and shiny, not sculptured. Propodeum armed with a small posteriorly directed spine. Propodeal spiracle in profile at about one-diameter from posterior edge. Metapleural gland present, metapleural spiracle big, longer than broader, within a dorsally directed fold. PETIOLE and POSTPETIOLE. Petiole broadly attached to postpetiole (abdominal segment III). Postpetiole in lateral view much shorter than gaster (abdominal segment IV). GASTER. Shiny. Vaulted. Constriction between postpetiole and gaster present. Abdominal sternum IX simple, triangular in shape, without spines or lobes. Sting present. LEGS. Mid and hind tibia with pectinate spurs present. Male: HW = 0.88mm. Body compact, with exterior morphology (except head) similar to workers. Body covered by decumbent setae. Color dark. HEAD. Dorsum with scrabrous-strigate sculture. Lateral ocelli and median ocellus present. Antenna 12 -segmented. Antennal sockets exposed, not covered by frontal carinae, at mid-length from the anterior border of clypeus and the head posterior vertex margin. Antennal scrobes absent. Antennal carinae absent. Scape very short, about 1.3 times as long as pedicel. First flagellar segment relatively short, about the same length as pedicel, slightly curved at base. Antennal club absent, but apical segment is at least 2 times longer than preceding segment. Mandibles reduced, falcate, without differentiated masticatory and basal margins. Mandible edentate, with no visible apical tooth. Clypeus broad, with straight anterior margin. Clypeus does not extend to space between eyes. EL = 0.32mm, eyes larger than in workers (REL = 48.44) located at mid-length at lateral margin. WINGS. WingL = 3.6mm, about 50 % longer than total body length. Venation and cell composition of both fore- and hind-wings similar to that of gyne. MESOSOMA. Oblique mesopleural furrow close to but not reaching pronotum. Mesonotum notauli absent. Mesoscutum and mesoscutellum mostly shiny, with small foveae. Pronotum with rugae. PETIOLE and POSTPETIOLE. Constriction between petiole and postpetiole (abdominal segment III)present. Petiole and postpetiole similar in shape to worker petiole and postpetiole. Petiole broadly attached to postpetiole. Postpetiole, in lateral view, much shorter than gaster (abdominal segment IV). GASTER. Shiny. Vaulted. Constriction between postpetiole and gaster present. Abdominal sternum IX simple, triangular in shape, without spines or lobes. LEGS. Hind tibia with 1 pectinate spur.

Donoso, David A. (2012): Additions to the taxonomy of the armadillo ants (Hymenoptera, Formicidae, Tatu i dri s). Zootaxa 3503: 61-81, DOI: 10.5281/zenodo.282479

Tatuidris tatusia Brown and Kempf 1968 Tatuidris tatusia Brown and Kempf 1968: 187, their figures 1���4. Holotype and Paratype workers: El Salvador, La Libertad, 2 mi S. Quetzaltepec, VII- 17-1961 (M.E. Irwin leg.) [holotype, UCDC] (image examined) [paratype, MCZC] (examined). Tatuidris kapasi Lacau and Groc, in Lacau et al. 2012: 2, their figures 1���6. Holotype worker: Guyane Francaise, Montagne de Kaw, 0 4 �� 38.21 �� N; 0 52 �� 17.36 �� W, Alt. 260 m., ix. 2008 (S. Groc, A. Dejean, and B. Corbara leg) [CPDC] (image examined). n. syn. Worker, male and gyne diagnosis: With same characters as in the genus description (Figures 1���4). Pilosity variability. Currently, four striking pilosity patterns are known to occur within Tatuidris collections (Figure 5). Pilosity pattern A (Figure 6) consists of a mix of both long flexuous and short appressed setae. This is the most common pilosity pattern and the one that most resembles the type specimens from El Salvador and the gyne from Otongachi, Ecuador. Pilosity pattern B (Figure 7) is characterized by very short, fully appressed, and regular spaced setae arrayed homogeneously and equidistantly on the head, mesosoma, petiole, postpetiole and gaster. Pilosity pattern C (Figure 8) is characterized by dense lanose-looking setae. Pilosity pattern D (Figure 9) consists of short and uniform decumbent (strongly inclined but not fully appressed) setae scattered throughout the body. DNA Barcodes. In total, 28 sequences (20 of them with full, 658 bp, length) of the barcode gene were recovered from BOLD and included in the analyses (Table 1). Among these sequences, 120 out of the 658 basepairs (18.23 %) were variable. In general, specimens (excluding colony duplicates) presented very high levels of intraspecific pairwise differences (average = 5.36 %, min = 0%, max= 15.08 %). The upper values of intraspecific pairwise differences encountered here are well above the 2 % usually recovered for species differences in DNA barcode literature in general (Ratnasingham and Hebert 2007), and above the percentages of average intraspecific variability reported previously for ants (e.g. between the 1.67 % and 6.37 % encountered for Anochetus and Odontomachus in Madagascar by Fisher and Smith 2008; see also Smith and Fisher 2009, Jansen et al. 2009). Pilosity patterns present among individuals (excluding pilosity pattern ���C��� for which no sequence was available) were not in similar clusters as depicted on the NJ tree (Figure 10). Instead, four different clusters (groups) with strong geographic structure were observed: G 1, Mexico + Honduras + Guatemala; G 2, Costa Rica + Nicaragua; G 3, Ecuador (West of Andes); and G 4, Ecuador (East of Andes/Amazon Basin) (Figure 10). Furthermore, across this geographic range, pairwise barcode divergence among specimens was significantly related to distance (R 2 = 0.37, pSize. Specimens of T. tatusia are small (average WbL = 0.62mm), but specimens can vary greatly in size, with larger specimens being twice as large as the smaller ones (min WbL = 0.45 mm, max WbL = 0.90 mm). Size variability within trap catches (possibly same colonies) may be considerable. For example, workers from one collection catch in Nicaragua (collection series MGB # 1179) varied 30 % in size (WbL from 0.65 to 0.85 mm). It is still unclear whether intra-colony size variation is due to the presence of morphological worker castes (e.g. minor and major castes) or continuous size variability. The PCA analysis revealed that most variability among specimens is related to size (proportion of variance explained by PC- 1 = 0.915), with PC- 2, PC- 3 and PC- 4 (e.g. shape) explaining little (0.033, 0.021 and 0.011, respectively) of total variation (Table 2). Eye length loaded positively and contrasted against all other variables, in PC 2. Tibia length and tibia width loaded negatively and contrasted against most other variables, in PC 3. Size variability summarized in this analysis was not related to pilosity patterns (Figure 12). In general no PC correlated with pilosity patterns. Hand drawings courtesy of Natalie Clay (Figure 12). Sculpture. The strength and depth of all sculpture patterns is accentuated in larger sizes. Collections from Nicaragua also tend to present more accentuated sculpture patterns. The head dorsum is usually smooth and shining, except for the area below eyes, which presents longitudinal carinae. The head vertex is covered with transverse carinulae. The lateral surface of the mandible is smooth and shining except for longitudinal superficial striae on the side that vary in depth. The antennal scape is shagreened and superficially areolate. The surface of the ventrolateral part of the pronotum varies strongly across specimens, from smooth and shining to strongly striate, or carinulate. The dorsum of the mesosoma has concentric carinulae and sometimes is slightly punctate. The mesopleuron is smooth and shining except for punctuations and areolae on the ventral margin. The propodeal declivity is smooth with fine transverse striae. The petiole and postpetiole are dorso-laterally strigulate. The gaster is mostly smooth and shiny but sometimes finely and sparsely strigulate. Distribution. The genus Tatuidris is restricted to the Neotropics, but it has an ample distribution that spans from northern Mexico to central Brazil, French Guiana (Lacau et al. 2012) and Amazonian Peru (Figure 13). No collections are known from the Caribbean, Gal��pagos, or other islands. Most specimens and collections are currently known to occur in localities west of the Andes, with more collections tending to occur towards Central America and Mexico. Most collections come from mountainside (pre-montane) areas at mid elevations (usually 800���1200m of altitude). Collections from lowland Amazonia are few. Natural history. Little is known about the biology of the ant genus Tatuidris and, until recently, no observations of live specimens were registered. Details of a first collection event of a small live colony (3 workers and 4 gynes) by Dr. Thibaut Delsinne (pers. comm.) in a mid-elevation forest in southeastern Ecuador suggest that Tatuidris may well be a highly specialized predator, as colonies kept in captivity did not accept any food item offered to them. Food items rejected by the ants included live and dead termites, millipedes, mites, various insect parts, sugar water, tuna, biscuits, live and dead fruit flies (Drosophila), live springtails, live myriapods (Chilopoda and Diplopoda), live and dead Diplura, small live spiders, small live pseudoscorpions, one small snail, uncooked hen egg (i.e. piece of cotton wool soaked with fresh whisked hen egg; Brown 1977), ant larvae (Gnamptogenys sp.), and live ant workers (Cyphomyrmex sp., Brachymyrmex sp.). Potential food items (arthropods) for Tatuidris were taken from soil samples and Winkler samples (following Silva and Brand��o 2010) collected at the site where Tatuidris was a priori determined to be present. Further observations by T. Delsinne suggest that T. tatusia may be a sit-and-wait predator, as ���both workers and gynes moved very slowly and were very clumsy. They often remained motionless during several tens of seconds or even several minutes when disturbed (either by my handling or by the contact with another arthropod). It is difficult to see them as powerful predators!��� (pers. comm.). These observations were mainly performed at night, suggesting that T. tatusia may be nocturnal, a hypothesis also supported by collection patterns. For example, in the R��o Toachi forest of Ecuador T. tatusia specimens tend to fall in pitfall traps, instead of Winkler sacs (Donoso and Ram��n 2009). Because pitfall traps usually work 24 -h, but Winkler sacs generally uses litter sifted during the day, then ants with nocturnal habits may be underrepresented in Winkler samples. The small eyes of Tatudris species provide further support for this hypothesis. Other. The relative position of eyes is highly variable within the species. For example, eye location ranges from being completely within the antennal scrobes to completely outside the scrobes (Figure 1 b). In some cases (specimen J.Longino# 2088 -S) the eye itself is located outside the antennal scrobe, but the eye���s fossa is well marked and confluent with the antennal scrobe. In most specimens, the antennal carina bifurcates from the antennal scrobes and lies straight above the eyes. However, in specimens from Nicaragua (MGB # 1179 colony collection), a strongly impressed antennal carina is present. In these specimens about 40 % of the eye���s area lies within the antennal scrobes. In the gyne, only ~ 1 / 6 of the eye lies within the antennal scrobes. A depression sometimes forms in the integument in the sides of the propodeum, below the propodeal spiracle and above the metapleural gland. The depth of this depression varies among specimens and tends to be deepest in larger specimens. Justification for the synonymy. A comprehensive analysis of the description of T. kapasi suggests that the morphological characters proposed by Lacau and Groc (2012) to separate T. kapasi from T. tatusia lie within the continuous variability encountered in Tatuidris specimens across the Neotropics. Two considerations were taken. First, the description of T. kapasi relies on a relatively large specimen where sculpture, body profile and body proportions are expected to vary the most. Diagnostic characters of T. kapasi (i.e. occipital border more concave, clypeus with the free margin nearly straight and laterally concave, dorsum of mesosoma mostly sculptured with concentric rugulae and carinulae, and dorsum of the petiolar and postpetiolar nodes with superficial concentric rugulae and carinulae) are all present to some degree in larger specimens among the collections reviewed in the present study (Figures 6���9). Another diagnostic character of T. kapasi was denser pilosity without long setae (similar to Morph D), but setal patterns did not correlate with any morphology- or molecular-based grouping in this study. Second, the type locality of T. kapasi lies at a range limit of the genus, where morphological extremes might be expected. Worker measurements (in mm) and indices: [average (min���max) of 10��� 58 specimens]: AScL 0.46 (0.31, 0.67); AScW 0.24 (0.18, 0.36); CIx 129.03 (117.07, 137.93); EL 0.05 (0.03, 0.08); FL 0.43 (0.31, 0.70); FW 0.11 (0.08, 0.17); HL 0.59 (0.43, 0.88); HW 0.76 (0.56, 1.10); IAD 0.36 (0.25, 0.54); PL 0.16 (0.10, 0.24); PpL 0.16 (0.10, 0.25); PW 0.25 (0.18, 0.37); PpW 0.36 (0.26, 0.53); PPpIx 68.92 (58.14, 80.00); PrW 0.52 (0.38, 0.77); TiL 0.35 (0.27, 0.52); TiW 0.10 (0.06, 0.17); WbL 0.62 (0.45, 0.89); SL 0.42 (0.32, 0.51); SW 0.13 (0.11, 0.17); SI 329.82 (300.00, 360.00); REL 9.02 (5.41, 11.48). Gyne measurements (in mm) and indices: (n= 1) AScL 0.47; AScW 0.15; EL 0.20; FFS 0.09; FL 0.77; FW 0.20; HL 0.88; HW 1.28; IAD 0.56; Pl 0.22; PPL 0.28; PPW 0.72; PW 0.45; TL 0.69; TW 0.20; WingL 4.60; WbL 1.53; REL 22.63. Male measurements (in mm) and indices: (n= 1) EL 0.32; FFS 0.13; HL 0.66; HW 0.88; IAD 0.21; ScL 0.11; WingL 3.6; WbL 1.22; FL 0.9; REL 48.44. Specimens examined: Elements in this list follow this order: COUNTRY, State or Province, Locality, elevation, coordinates, collector (collection code, habitat, method) [specimen code], [REPOSITORY]. BELIZE, 2.5 millas S Belmopan, S&J Peck (B- 242, Limestone forest, ex: Berlese) [MCZC]. Caves Branch, S&J Peck (B- 248, Hi Canopi Forest, ex: Berlese [MCZC]. BRAZIL, Amazonas, Manaus, Universidade do Amazonas, 03��05' 36 ''S; 59 �� 57 ' 52 ''W, E Pereira Franken (Terra Firme: PLATO, ex: Pitfall) [INPA]. COLOMBIA, Magdalena, El Campano, 1300m, 11 ��07'N; 74 ��06'W, Phillip Ward (# 7891 - 9, montane rainforest, ex: Sifted litter, leaf mold, rotten log) [QCAZ]. COSTA RICA, Rincon de la Vieja, Pailas, 1000m, 10.7887, ��� 85.349, A Smith (10 COSTA-0629 and 10 COSTA-0630, Primary Forest) [AlexSmith]. Alajuela, Casa Eladio, Rio Pe��as Blancas, 800m, 10 �� 19 ' N; 84 �� 43 ' W, J Longino (Wet forest, ex: Sifted leaf litter) [INBIOCRI002280066, INBIOCRI002280068 and INBIOCRI002280065] [JTLC]. Alajuela, El Aleman, Rio Pe��as Blancas, 6,5 Km E Monteverde, 940m, 10 �� 18 ' N; 84 �� 45 ' W, J Longino (#0861-s, Wet forest, ex: Sifted leaf litter) [INBIOCRI002280059] [JTLC]. Heredia, 16km SSE La Virgen, 1100m, 10 �� 16 'N; 84 ��05'W [INB0003214087, INB0003214014, INB0003214104, INB0003214131, INB0003214232 and INB0003212497] [INBC]. Heredia, La Selva Biological Station-1, 50m, M Molina and H Garcia (AMI- 1 -W-006-05) [JTLC]. Heredia, La Selva Biological Station-2, 50m, 10.433, ��� 84.0167, J Longino (All ants from Louisa Stark (OTS 89 - 1 student) study, ex: Litter and ground-nesting) [INBIOCRI001281194 and INBIOCRI001281181] [JTLC]. Heredia, La Selva Biological Station-2, 09��09' N; 079�� 51 ' W, Michael Kaspari (45196.01 and 45000.6) [MEK]. Puntarenas, Rio San Luis, 850m, 10 �� 17 ' N; 84 �� 48 ' W, J. Longino (Moist forest, ex: Sifted leaf litter on ground) [INBIOCRI002280062 and INBIOCRI002280063] [JTLC]. ECUADOR, Cotopaxi, 19 km ENE La Man��, 1100m, 00�� 53 ' S; 79 ��03' W, Phillip Ward (# 11418 - 6, Second-Grown Rainforest, ex: Sifted leaf litter and logs) [QCAZ]. Pichincha, R.B. Maquipucuna, 1200m, 00��07'00''N; 78 �� 38 '06W, R Anderson (# 99-208 - 6 and # 99-208, Montane evergreen forest) [CASENT0423526 and CASENT0001968] [CASC]. Pichincha, Uni��n del Toachi-Otongachi, 850m, 00�� 21 '05"S; 78 �� 57 ' 10 "W, Donoso & Vieira (Bosque Secundario, ex: Pitfall) [18 workers and 1 gyne, MEKOU 12079 - 84 and MEKOU 12103 - 4] [QCAZ]. Zamora, Bombuscaro Numex, 950m, ��� 4.114972 S; ��� 78.96794 W, M Leponce (SPM _ 33796, Evergreen premontane rainforest, ex: Winkler sample) [QCAZ]. Zamora, Copalinga Private Reserve, 1000m, ��� 4.0912222 S; ��� 78.96069 W, T Delsinne and T. Arias-Penna (SPM _ 4130219, Secondary evergreen premontane rainforest, ex: Winkler sample) [QCAZ]. EL SALVADOR [Paratype and Holotype], 2 mi. S Quetzaltepec, ME Erwin [MCZ Paratype 31577] [MCZC]. GUATEMALA, Peten, Parq. Nac. Tikal, 270m, 17.232 N; ��� 89.623 W (Tropical Moist Forest) [CASC]. HONDURAS, Comayagua, PN Cerro Azul Meambar, 1120m, 14.887136; ��� 87.89974 (LLAMA #Wa-C- 04- 1-15, Ridgetop cloud forest, ex: Sifted leaf litter) [CASENT 0615399] [CASC]. MEXICO, Chiapas, 860m, 16.980 N; ��� 91.586 W, MG Branstetter (# 856, Mesophil forest) [QCAZ]. Chiapas, 12 mi NW Ocozocoautla, 400m, A Newton (ex: Berlese) [MCZC]. Chiapas, 6 kmSW Ocosingo, 1400m, 16.867221 N; ��� 92.0787132 W, R Anderson (# 91���116, Forest litter, ex: Berlese) [CASENT 0603397] [INBC]. Chiapas, Lago Metzabok, 575m, 17.124, ��� 91.636, MG Branstetter (Lowland wet forest) [CASC]. Oaxaca, Mirador Grande, 990m, 17.89844, ��� 96.36253, MG Branstetter (# 1405) [QCAZ]. Tamaulipas, El Cielo, 870m, 23.276, ��� 99.276, MG Branstetter (# 1465) [QCAZ]. Ve r a c r u z, Los Tuxtlas, Ejido-L��pez Mateos, 50m, 18 �� 24 ' 56 ''N; 94 �� 56 ' 53 ''W, P Rojas (LM 7 S-H 1, LM 42 S-H 4, LM 42 S-H 5 and LM 21 A-H 3, Selva alta perennifolia, ex: Winkler) [IEXM]. Veracruz, Los Tuxtlas, Ejido-L��pez Mateos, Ejido 2, 50m, 18 �� 24 ' 56 ''N; 94 �� 56 ' 53 ''W, P Rojas (LM 42 S- H 3, Selva alta perennifolia, ex: Winkler) [IEXM]. Veracruz, Los Tuxtlas, Volc��n S. M. Pajapan, 510m, 18 �� 16 '00''N; 94 �� 46 ' 71 ''W, A Cartas (847 d, Selva mediana subperennifolia, ex: Berlese) [IEXM]. NICARAGUA, Matagalpa, RN El Mus��n, 4.8km NNW Rio Blanco, 1170m, 12 �� 58.4 'N; 085�� 14 'W, M.G.Branstetter (# 1179, Mesic forest, ex: Sifted leaf litter) [QCAZ]. PANAMA, Chiriqui, 20,4 Km North San Felix, R. Anderson (# 17768 _ 1, Wet mountain forest, ex: Litter sample) [WEMC]. Chiriqui, Alto Lino, 3800m, HG Real [Male, CASENT 0102681] [CASC]. Chiriqui, La Fortuna, Finca La Suisse, 1200m, R. Anderson (# 17838, # 17787, # 17788, # 17789, # 17790, # 17838 and # 17839 [WEMC]. PERU, Cuzco, La convenci��n Province, 4 km S Camisea River. Campamento Cashiriari- 2, Plot 1, 579m, 11 �� 51 ' 51.3 ''S; 72 �� 46 ' 45.6 ''W, J Santisteban et al. (# 38, Primary Rainforest, hilly terrain, ex: Winkler Trap) [MUSM-ENT 0201599/ANTWEB-CASENT 0178882] [MUSM]., Published as part of Donoso, David A., 2012, Additions to the taxonomy of the armadillo ants (Hymenoptera, Formicidae, Tatu i dri s), pp. 61-81 in Zootaxa 3503 on pages 69-75, DOI: 10.5281/zenodo.282479, {"references":["Brown, W. L. & Kempf, W. W. (1968) Tatuidris, a remarkable new genus of Formicidae (Hymenoptera). Psyche, 74, 183 - 190.","Lacau, S, Groc, S., Dejean, A. de Oliveira, M. L. & Delabie J. C. H. (2012) Tatuidris kapasi sp. nov.: a new armadillo ant from French Guiana (Formicidae: Agroecomyrmecinae). Psyche, 2012, Article ID 926089, 6 pages, doi: 10.1155 / 2012 / 926089","Ratnasingham, S. & Hebert, P. D. N. (2007) BOLD: The Barcode of Life Data System (www. barcodinglife. org). Molecular Ecology Notes, 7, 355 - 364.","Fisher, B. L. and Smith, M. A. (2008) A revision of malagasy species of Anochetus Mayr and Odontomachus Latreille (Hymenoptera: Formicidae). PLoS ONE, 3, e 1787.","Smith, M. A. & Fisher, B. L. (2009) Invasions, DNA barcodes, and rapid biodiversity assessment using ants of Mauritius. Frontiers in Zoology, 6, 31.","Jansen, G., Savolainen, R. & Vepsalainen, K. (2009) DNA barcoding as a heuristic tool for classifying undescribed Nearctic Myrmica ants (Hymenoptera: Formicidae). Zoologica Scripta, 38, 527 - 536.","Brown, W. L. (1977) An aberrant new genus of myrmicine ant from Madagascar. Psyche, 84, 218 - 224.","Silva, R. R. & Brandao, C. R. F. (2010) Morphological patterns and community organization in leaf-litter ant assemblages. Ecological Monographs, 80, 107 - 124"]}