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Pseudomyrmex ereptor sp. nov. Figs. 12 and 24 Zoobank LSID: urn:lsid:zoobank.org:act: 3F4584C3-EB0C-4C74- B067-01CBEDC8F4EB Holotype alate queen, MEXICO Veracruz: Los Tuxtlas, 10 km NNW Sontecomapan, 500 m, 18° 35 ′ N, 95° 05 ′ W, 21 Mar 1985, ex dead twig of liana [in nest of P. elongatulus], rainforest, P. S. Ward PSW07360 A (UNAM) (CASENT0863524). Other material examined. Known only from the holotype. Queen measurements (n = 1). HW 0.98, HL 1.25, MFC 0.058, LHT 0.76, CI 0.78, FCI 0.059, REL 0.44, REL2 0.56, FI 0.46, PLI 0.61, PWI 0.75, MSC 11. Queen Diagnosis. Small species (see HW, HL, and LHT measurements), with moderately elongate head (CI 0.78); upper surface of mandible smooth and shiny, with scattered punctures; eyes of moderate length (REL 0.44, REL2 0.56); frontal carinae separated by basal scape width; petiole relatively short (PLI 0.61, PL/HL 0.52) and broad (PWI 0.75, DPW/LHT 0.64), with stout, recurved anteroventral tooth; in profile, petiole with convex anterodorsal face ascending gradually to summit in posterior quarter of node, then rounding into steep posterior face; postpetiole very broad, ovorectangular in dorsal view (PPW/LHT 1.01); profemur robust (FI 0.46); hind leg short (LHT/HL 0.61). Head sublucid, densely but finely punctulate, the punctures separated by one to several diameters, the interspaces smooth and shiny, or (towards vertex) finely coriarious. Standing pilosity scattered on head, mesosoma (MSC 11), and gaster, short and sparse on petiole, postpetiole, and fourth abdominal (first gastric) tergite. Uniformly light orange-brown. Comments. The small size (HL 1.25, LHT 0.76), smooth and shiny mandibles, and very broad (ovorectangular) postpetiole (PPW/LHT 1.01) are distinctive features of P. ereptor that set it apart from the queen of its apparent host species, P. elongatulus, and from other species in the P. elongatulus group. Queens of P. elongatulus are larger (HL 1.37–1.42, LHT 0.90–0.94), with striate mandibles, and a slender, pyriform postpetiole as seen in dorsal view (PPW/LHT 0.78–0.83). Our UCE phylogeny shows that P. ereptor is sister to the entire assemblage of P. elongatulus samples (Fig. 1). It is well separated from an alate queen of P. elongatulus (D1985) collected in the same nest, thus confirming its status as a different species. Distribution and Biology. The only known specimen of P. ereptor — an alate queen—was collected in a nest of P. elongatulus in a dead twig attached to a liana, on Vereda Cima, Estación de Biología ‘Los Tuxtlas’, Veracruz. The nest (collection code PSW07360) contained numerous workers, alate queens, males, and brood of P. elongatulus. Pseudomyrmex ereptor apparently represents a workerless, inquiline parasite of P. elongatulus. The fact that only a single alate queen of P. ereptor was found in the nest suggests that she was a recent arrival. The absence of a dealate queen of P. elongatulus should not be over-interpreted; it could simply indicate that this was part of a larger polydomous colony. Pseudomyrmex colonies are often comprised of disjunct nests occupying several dead twigs on a plant., Published as part of Ward, Philip S. & Branstetter, Michael G., 2022, Species Paraphyly and Social Parasitism: Phylogenomics, Morphology, and Geography Clarify the Evolution of the Pseudomyrmex elongatulus Group (Hymenoptera:, pp. 1-31 in Insect Systematics and Diversity (AIFB) (AIFB) 6 (1) on pages 23-24, DOI: 10.1093/isd/ixab025, http://zenodo.org/record/7182517

Wadeura holmgreni Wheeler New Combination Figs. 13,14 Euponera (Trachymesopus) holmgreni Wheeler 1925:6. Holotype worker: Peru (N. Holmgren) [MCZC] (examined). Trachymesopus holmgreni: Kempf, 1960:424; Kempf, 1961:494, figs. 4,5,6 (redescription of worker). Pachycondyla holmgreni: Brown, in Bolton 1995:306; Mackay & Mackay, 2010:390. Cryptopone holmgreni: Schmidt and Shattuck 2014:185., Published as part of Branstetter, Michael G. & Longino, John T., 2022, UCE Phylogenomics of New World Cryptopone (Hymenoptera: Formicidae) Elucidates Genus Boundaries, Species Boundaries, and the Vicariant History of a, pp. 1-23 in Insect Systematics and Diversity (AIFB) (AIFB) 6 (1) on page 19, DOI: 10.1093/isd/ixab031, http://zenodo.org/record/7182481, {"references":["Wheeler, W. M. 1925. Neotropical ants in the collections of the Royal Museum of Stockholm. Ark. Zool. 17 A (8): 1 - 55.","Kempf, W. W. 1960. Miscellaneous studies on Neotropical ants (Hymenoptera, Formicidae). Stud. Entomol. (n. s.) 3: 417 - 466.","Kempf, W. W. 1961. A survey of the ants of the soil fauna in Surinam (Hymenoptera: Formicidae). Stud. Entomol. 4: 481 - 524.","Bolton, B. 1995. A New General Catalogue of the Ants of the World. Harvard University Press, Cambridge, MA.","MacKay, W. P., and E. MacKay. 2010. The Systematics and Biology of the New World Ants of the Genus Pachycondyla (Hymenoptera: Formicidae). Edwin Mellen Press, Lewiston, New York.","Schmidt, C. A., and S. O. Shattuck. 2014. The higher classification of the ant subfamily Ponerinae (Hymenoptera: Formicidae), with a review of ponerine ecology and behavior. Zootaxa 3817: 1 - 242."]}

Pseudomyrmex elongatulus Group Worker Diagnosis. Moderate in size (HW 0.82–1.21, LHT 0.68– 1.12), with a variably elongate head (CI 0.67–0.96), and medium to large eyes (REL 0.38–0.59, REL2 0.46–0.66). Masticatory margin of mandible with 5–6 teeth; mesial tooth on basal margin closer to apicobasal angle than to proximal tooth. Palp formula 6,4. Median clypeal lobe laterally rounded. Frontal carinae separated by basal scape width or less (FCI 0.02–0.07). Profemur relatively slender in most species (FI 0.37–0.49). Pronotal humeri and lateral margins of pronotum rounded. Metanotal groove varying from moderately impressed to absent. Basal and declivitous faces of propodeum weakly to moderately differentiated and usually subequal in length, in profile the juncture between the two gently rounded. Petiole relatively long and slender (PL/HL 0.48–0.69; EL/PL 0.72–0.90), much longer than high or wide (PLI 0.43–0.61, PWI 0.41–0.58); petiole with small anteroventral tooth; anterior peduncle of petiole weakly to moderately differentiated. Postpetiole broader than long, with small anteroventral tooth. Body sculpture punctulate-coriarious to coriarious-imbricate, the integument sublucid to opaque; dorsum of head never with extensive smooth, shiny interspaces; punctulae usually separated by their diameters or less. Standing pilosity generally sparse, nearly always absent from the extensor faces of tibiae, and usually present as 1–2 pairs on the pronotum, petiole, and postpetiole, while being absent from mesonotum and propodeum (MSC 2–8); one exceptional species with greater amounts of pilosity (MSC 23–28, HTC 0, MTC 1–2) including on the mesonotum and propodeum. Appressed pubescence dense but short on most of body, including head and abdominal tergite IV. Color varying from light yellow- or orange-brown to dark brownish-black, often variously bicolored. Comments. In an earlier synopsis of Pseudomyrmex species groups (Ward 1989), the P. elongatulus group was subsumed under the P. pallens group. The two groups share a number of similar worker characteristics (6,4 palp formula, 5–6 teeth on the masticatory margin of the mandibles, and laterally rounded median clypeal lobe) but molecular studies have demonstrated that they are not closely related (Ward and Downie 2005, Chomicki et al. 2015). A recent worker-based key to Pseudomyrmex species groups (Ward 2017) separated the P.elongatulus group and P. pallens group primarily by geography: the species in the P. pallens group are almost wholly confined to South America, with just one species, P. pallens, extending north of Colombia as far as Costa Rica, while the P. elongatulus group occurs from southwestern United States to Costa Rica.Workers of the one overlapping species, P. pallens, can be distinguished from those of the P. elongatulus group by the combination of dense standing pilosity (MSC 13–28) and long eyes relative to petiole length (EL/PL 0.91–1.20)., Published as part of Ward, Philip S. & Branstetter, Michael G., 2022, Species Paraphyly and Social Parasitism: Phylogenomics, Morphology, and Geography Clarify the Evolution of the Pseudomyrmex elongatulus Group (Hymenoptera:, pp. 1-31 in Insect Systematics and Diversity (AIFB) (AIFB) 6 (1) on page 9, DOI: 10.1093/isd/ixab025, http://zenodo.org/record/7182517, {"references":["Ward, P. S. 1989. Systematic studies on pseudomyrmecine ants: revision of the Pseudomyrmex oculatus and P. subtilissimus species groups, with taxonomic comments on other species. Quaest. Entomol. 25: 393 - 468.","Ward, P. S., and D. A. Downie. 2005. The ant subfamily Pseudomyrmecinae (Hymenoptera: Formicidae): phylogeny and evolution of big-eyed arboreal ants. Syst. Entomol. 30: 310 - 335.","Chomicki, G., P. S. Ward, and S. Renner. 2015. Macroevolutionary assembly of ant / plant symbioses: Pseudomyrmex ants and their ant-housing plants in the Neotropics. Proc. R. Soc. B. 282: 20152200.","Ward, P. S. 2017. A review of the Pseudomyrmex ferrugineus and Pseudomyrmex goeldii species groups: acacia-ants and their relatives (Hymenoptera: Formicidae). Zootaxa. 4227: 524 - 542."]}

Wadeura Weber 1939 Revived Status Type species: Wadeura guianensis Weber, 1939:103; by original designation. See discussion of morphology under Cryptopone. Wadeura now contains four species: W. guianensis, W. holmgreni, W. holmgrenita sp. nov., and W. pauli. Phylogenetic information is available for three of the four known species, with W. pauli being sister to W. holmgrenita and W. guianensis. Wadeura holmgreni is very similar to W. holmgrenita and the two are likely sister species., Published as part of Branstetter, Michael G. & Longino, John T., 2022, UCE Phylogenomics of New World Cryptopone (Hymenoptera: Formicidae) Elucidates Genus Boundaries, Species Boundaries, and the Vicariant History of a, pp. 1-23 in Insect Systematics and Diversity (AIFB) (AIFB) 6 (1) on page 18, DOI: 10.1093/isd/ixab031, http://zenodo.org/record/7182481, {"references":["Weber, N. A. 1939. New ants of rare genera and a new genus of ponerine ants. Ann. Entomol. Soc. Am. 32: 91 - 104."]}

Branstetter, Michael G., Longino, John T. (2022): UCE Phylogenomics of New World Cryptopone (Hymenoptera: Formicidae) Elucidates Genus Boundaries, Species Boundaries, and the Vicariant History of a. Insect Systematics and Diversity (AIFB) 6 (1): 1-23, DOI: 10.1093/isd/ixab031, URL: http://dx.doi.org/10.1093/isd/ixab031

Pseudomyrmex comitator sp. nov. Figs. 9 and 24 LSID: urn:lsid:zoobank.org:act: 4BAA462E-31A4-4799-BEF6- D7BC23B346F0 P. championi nr; Chomicki et al. 2015: 4. Placement in molecular phylogeny. Holotype dealate queen, MEXICO Chiapas: 29 km E La Trinitaria, 1,520 m, 16° 06 ′ N 91° 46 ′ W, 21 Jul 2007, ex nest of P. cognatus in dead stick, pine oak scrubby forest, J. Longino JTL6094 (UNAM) (JTLC000010310). Paratype dealate queen, MEXICO Chiapas: Lagos de Montebello, 1,520 m, 16° 08 ′ N 91° 44 ′ W, 21 Jul 2007, on ground, pine oak Liquidambar forest, J. Longino JTL6093-s (UCDC) (JTLC 000010349). Other material examined. Known only from the type material. Queen measurements (n = 2, with holotype measurement given first). HW 0.81, 0.77; HL 1.01, 1.00; MFC 0.038, 0.046; LHT 0.70, 0.69; CI 0.80, 0.77; FCI 0.047, 0.060; REL 0.40, 0.42; REL2 0.51, 0.54; FI 0.40, 0.43; PLI 0.67, 0.83; PWI 0.56, 0.80; MSC 4, 7. Queen Diagnosis. Small species (see HW, HL, and LHT measurements), with elongate head (CI 0.77–0.80); upper surface of mandible finely reticulate with scattered punctures, lacking striae; eyes relatively short (REL 0.40–0.42, REL2 0.51–0.54); frontal carinae separated by slightly less than basal scape width; petiole stout and short, the height and width of petiole about 0.8× the length (less so in the paratype); in profile, petiole with flat to convex anterodorsal face, ascending to summit in posterior quarter of node, then rounding into more steeply descending posterior face; profemur slender; hind leg relatively short (LHT /HL 0.69). Head sublucid, densely punctulatecoriarious. Standing pilosity sparse and short on most of body (MSC 4–7). Dark brown to brownish-black, head and gaster darker than rest of body. Comments. This species, known only from two dealate queens, bears some resemblance to P. cognatus, but differs in ways that suggest it represents a workerless inquiline (see also notes on Distribution and Biology below). The P. comitator queens are notably smaller than those of P. cognatus (HW 0.77–0.81, HL 1.00–1.01, vs HW 0.99– 1.11, HL 1.28–1.42 in queens of P. cognatus), lack striae on the mandibles, have reduced mesosomal pilosity (MSC 4–7 vs 15–22 in P. cognatus queens), and possess an oddly swollen and foreshortened petiole (PLI 0.67–0.83, vs 0.45–0.53 in P. cognatus queens) (Figs. 8 and 9). The two specimens of P. comitator do not show the same degree of modification of the petiole (compare PLI and PWI values), with the holotype being more extreme, indicating some instability in the expression of this character. Both individuals of P. comitator were sequenced; they are sister taxa in our UCE tree and embedded phylogenetically within the putative host species (Fig. 1). They are more closely related to P. cognatus populations from Honduras and Nicaragua, however, than to samples from Chiapas. Distribution and Biology. Pseudomyrmex comitator is an apparent workerless social parasite of P. cognatus, and is currently known only from two adjacent sites in high-elevation forest (1,520 m) of Chiapas, Mexico. The holotype was collected by Jack Longino in a nest of P. cognatus that contained workers, alate males, alate queens, and brood (larvae, prepupae) of the presumed host. Only a single dealate queen of P. comitator was found in the nest. The paratype queen was encountered as a stray on the ground, in a nearby locality.

Wadeura pauli Fernandes & Delabie (New Combination) (Figs. 13,14) Cryptopone pauli Fernandes & Delabie, 2019:409. Holotype worker: Brazil, Rondônia, Floresta Nacional do Jamari (Flona Jamari), 63°01’24“W 9°09’40”S, Serra da Onça, 29.v.2018, in soil sample (D.C. Castro) [INPA] (not examined). Sequenced specimen: Guyana, Potaro-Siparuni: 8km WNW Chenapau, 5.00851 −59.64133 ± 20 m, 510 m, 13 Oct 2015, premontane rainforest, ex sifted leaf litter (M. G. Branstetter #MGB2118). Measurements. Worker HW 1.41 (holotype), 1.46 (sequenced specimen). Comments. This species is by far the largest in the genus. It is now known from six specimens: the holotype and three paratypes from Rondônia, the sequenced specimen from Guyana, and a specimen from Rio Trombetas, Pará, Brazil (imaged specimen ANTWEB1041391 on AntWeb). The sequenced specimen has a tiny compound eye, raising the possibility that it is an ergatoid queen. Wadeura holgreni and W. holmgrenita have both completely eyeless workers and worker-like individuals with tiny compound eyes. The latter are potentially ergatoid queens. Fisheropone hartwigi (Arnold) New Combination Cryptopone hartwigi Arnold, 1948:213, figs. 2, 2a, 2b. Syntype worker: South Africa, Pretoria (E.K. Hartwig) [SAMC] (not examined). Combination in Hypoponera: Taylor, 1967:12. Combination in Cryptopone, status as species: Bolton, 1995: 166. Comments. This species is transferred based on molecular evidence and no attempt is made to provide morphological evidence or to revise the diagnosis of the genus Fisheropone. The transfer serves mainly to exclude the species from Cryptopone., Published as part of Branstetter, Michael G. & Longino, John T., 2022, UCE Phylogenomics of New World Cryptopone (Hymenoptera: Formicidae) Elucidates Genus Boundaries, Species Boundaries, and the Vicariant History of a, pp. 1-23 in Insect Systematics and Diversity (AIFB) (AIFB) 6 (1) on page 20, DOI: 10.1093/isd/ixab031, http://zenodo.org/record/7182481, {"references":["Fernandes, I. O., and J. H. Delabie. 2019. A new species of Cryptopone Emery (Hymenoptera: Formicidae: Ponerinae) from Brazil with observations on the genus and a key for new word species. Sociobiology 66: 408 - 413.","Arnold, G. 1948. New species of African Hymenoptera. No. 8. Occas. Pap. Natl. Mus. South. Rhod. 2: 213 - 250.","Taylor, R. W. 1967. A monographic revision of the ant genus Ponera Latreille (Hymenoptera: Formicidae). Pac. Insects Monogr. 13: 1 - 112.","Bolton, B. 1995. A New General Catalogue of the Ants of the World. Harvard University Press, Cambridge, MA."]}

Centromyrmex brachycola (Roger) Ponera brachycola Roger, 1861:5. Combination in Centromyrmex: Emery, 1890a:74; Emery, 1890b:40 (footnote); Kempf 1967:405. Pachycondyla mirabilis Mackay & Mackay, 2010:466, figs. 5, 161, 586-588. Holotype worker: Bolivia, Rosario on Lake Rocagua (W. M. Mann) [LACM] (examined). Combination in Cryptopone: Schmidt and Shattuck 2014:185. New Synonymy. Comments. Kempf (1967) reviewed the Neotropical Centromyrmex. Under C. brachycola he discussed a series from Bolivia, Rosario on Lake Rocagua, collected by Mann. Kempf referred to Mann’s account of finding this colony in a termite mound (Mann 1934:189). Parts of this series are in multiple collections, including LACM. Mackay and Mackay observed specimens of this series at LACM and mistakenly described it as the new species Pachycondyla mirabilis, close to W. gilva and W. holmgreni., Published as part of Branstetter, Michael G. & Longino, John T., 2022, UCE Phylogenomics of New World Cryptopone (Hymenoptera: Formicidae) Elucidates Genus Boundaries, Species Boundaries, and the Vicariant History of a, pp. 1-23 in Insect Systematics and Diversity (AIFB) (AIFB) 6 (1) on page 21, DOI: 10.1093/isd/ixab031, http://zenodo.org/record/7182481, {"references":["Roger, J. 1861. Die Ponera-artigen Ameisen (Schluss). Berl. Entomol. Z. 5: 1 - 54.","Emery, C. 1890 a. Voyage de M. E. Simon au Venezuela (Decembre 1887 - Avril 1888). Formicides. Ann. Soc. Entomol. France 10 (6): 55 - 76.","Emery, C. 1890 b. Studii sulle formiche della fauna neotropica. Bull. Soc. Entomol. Ital. 22: 38 - 80, Tav. V-IX.","Kempf, W. W. 1967 (1966). A synopsis of the Neotropical ants of the genus Centromyrmex Mayr (Hymenoptera: Formicidae). Stud. Entomol. 9: 401 - 410.","MacKay, W. P., and E. MacKay. 2010. The Systematics and Biology of the New World Ants of the Genus Pachycondyla (Hymenoptera: Formicidae). Edwin Mellen Press, Lewiston, New York.","Schmidt, C. A., and S. O. Shattuck. 2014. The higher classification of the ant subfamily Ponerinae (Hymenoptera: Formicidae), with a review of ponerine ecology and behavior. Zootaxa 3817: 1 - 242.","Mann, W. M. 1934. Stalking ants, savage and civilized. Natl. Geogr. Mag. 66: 171 - 192."]}

Ward, Philip S., Branstetter, Michael G. (2022): Species Paraphyly and Social Parasitism: Phylogenomics, Morphology, and Geography Clarify the Evolution of the Pseudomyrmex elongatulus Group (Hymenoptera:. Insect Systematics and Diversity (AIFB) 6 (1): 1-31, DOI: 10.1093/isd/ixab025, URL: http://dx.doi.org/10.1093/isd/ixab025

Wadeura holmgrenita, New Species (Figs. 13,14,15) (Zoobank LSID: urn:lsid:zoobank.org:act: 7E5A70E9-2F48-4200-9 EA0-4544436F5DA5) Type material. Holotype worker: Peru, Madre de Dios: Estación Biológica Villa Carmen, 12.8753 -71.41095 ± 300 m, 650 m, 21 Oct 2015, rainforest, in soil (B.L.Fisher, F.A.Esteves # BLF 31580) [MUSM, unique specimen identifier CASENT0637779]. Paratypes, same data as holotype [1 worker, CAS, CASENT0370848; 1 worker, MCZC, CASENT0370849; 1 worker, MZSP, CASENT0370850]; same data except collection number BLF 31570 [1 worker, UCDC, CASENT0370863; 1 worker, JTLC, CASENT0370864]; same data except collection number BLF 31572 [1 worker, USNM, CASENT0370865]., Published as part of Branstetter, Michael G. & Longino, John T., 2022, UCE Phylogenomics of New World Cryptopone (Hymenoptera: Formicidae) Elucidates Genus Boundaries, Species Boundaries, and the Vicariant History of a, pp. 1-23 in Insect Systematics and Diversity (AIFB) (AIFB) 6 (1) on page 20, DOI: 10.1093/isd/ixab031, http://zenodo.org/record/7182481

Pseudomyrmex arcanus sp. nov. Figs. 5 and 20 Zoobank LSID: urn:lsid:zoobank.org:act: 5CA6B80D-C6DA-47A1- 82AD-1721A990214F, Published as part of Ward, Philip S. & Branstetter, Michael G., 2022, Species Paraphyly and Social Parasitism: Phylogenomics, Morphology, and Geography Clarify the Evolution of the Pseudomyrmex elongatulus Group (Hymenoptera:, pp. 1-31 in Insect Systematics and Diversity (AIFB) (AIFB) 6 (1) on page 18, DOI: 10.1093/isd/ixab025, http://zenodo.org/record/7182517