taxonID	type	description	language	source
9A5C0805FFAC0838F1A3FE6CCBB4F8E3.taxon	description	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, REL 2 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).	en	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 6 (1): 1-31, DOI: 10.1093/isd/ixab025, URL: http://dx.doi.org/10.1093/isd/ixab025
9A5C0805FFAD0821F203F922CB0AF966.taxon	materials_examined	Other material examined (AMNH, ANIC, BMNH, CASC, CHAH, CMNH, CNCC, CSCA, CUIC, EMEC, JTLC, LACM, MCZC, MNHN, MSNG, MZSP, PSWC, RAJC, SEMC, UCDC, UCRC, USNM, UTEP, UTIC). Mexico, Baja California: 10 km W Mexicali (Snelling, R. R.); 10 mi E Bahia San Quintín (Williams, F. X.); 19 mi S Ensenada (Timberlake); 28 km E Ensenada, 750 m (Ward, P. S.); 3.5 mi Arroyo Santo Tomas (Sleeper, E. L.); 5 km S San Quintin (Heydon, S. L.); La Rumorosa, 1,200 m (Ward, P. S.); Mexicali (Snelling, R. R.); Chihuahua: 16 mi W Gral. Trias, 1,770 m (Creighton, W. S.); 23 mi S Parral, 1,675 m (Creighton, W. S.); 3 mi S Encinillas, 1495 m (Creighton, W. S.); 34 mi S Parral, 1,770 m (Creighton, W. S.); 6 mi S Villa Matamoros (Gardner, R. C.; Kovacic, C. R.; Lorenzen, K.); Nogales Ranch, Sierra de en Medio, 1,525 m (Creighton, W. S.); Nogales Ranch, Sierra de en Medio, 1,585 m (Creighton, W. S.); Ojo del Cerro Chilicote (Townsend); Durango: 9 mi E Palmito (Alpert, G.); Villa Ocampo, 1,735 m (Creighton, W. S.); Nuevo León: China, 185 m (Creighton, W. S.); Sonora: 3 mi E Punta Cirios, 60 m (Johnson, R. A.); 30 mi SE Agua Prieta (Roth, V.); 4.8 mi S Cananea (Roth, V.); Nogales (Perkins, R. C. L.); United States, Arizona: Cochise Co.: 13 mi SW Apache (Rozen; Pavreau); 3 km SW Portal, 1,510 m (Ward, P. S.); 3 mi N Paradise, 1,525 m (Creighton, W. S.); 5 mi W Portal (Linsdale, D. D.); 7 km SE Sunnyside, 1670 m (Ward, P. S.); 9.5 km SW Portal, 1,775 m (Ward, P. S.); Carr Canyon, Huachuca Mtns., 1,645 m (Creighton, W. S.); Carr Canyon, Huachuca Mtns., 1,890 m (Obrien, C. W.); Cave Ck. Cyn., Chiricahua Mts., Herb Martyr Dam, 1,768 m (Hespenheide, H. A.); Cave Ck. Cyn., Chiricahua Mts., Sunny Flat, 1,554 m (Hespenheide, H. A.); Cave Creek Ranch, Chiricahua Mtns., 1,525 m (Wallace, G. E.); Cave Creek, Chiricahua Mts., 1,675 m (Kusche, J. A.); Chiricahua Mtns. (Knull, D. J.; Knull, J. N.); Chiricahua Mtns., 6 mi SE Camp Rucker (MacKay, W.); Chiricahua Mtns., Portal, 1,450 m (c. u.); Chiricahua Mtns., SWRS, 8 km W Portal, 1,650 m (Fisher, B. L.); Chiricahua Mtns., Tex Canyon, 14 km W Jct. Rt. 80 on FSR 74, 1,615 m (Ward, P. S.); Chiricahua Mtns., Tex Canyon, 8.6 mi W Jct. Rt. 80 on FSR 74, 1615 m (Saux, C.); Chiricahua National Monument Cpgrd., 1,645 m (Creighton, W. S.); Cochise Stronghold, Dragoon [as ‘ Dragon’] Mtns. (O’Brien, C. W.); Cochise Stronghold, Dragoon Mtns., 1,585 m (Creighton, W. S.); Coronado Peak, 2,020 m (Ward, P. S.); Cottonwood Canyon, Peloncillo Mtns., 1,465 m (Creighton, W. S.); Garden Canyon, Huachuca Mtns., 1,770 m (Creighton, W. S.); Garden Canyon, Huachuca Mtns., 1,770 m (Creighton, W. S.); Huachuca Mtns. (c. u.); Huachuca Mtns. [as ‘ Huachuca M. ’] (Knull, J. N.); Miller Canyon, Huachuca Mtns. (Creighton, W. S.); Paradise, Chiricahua Mts., 1,675 m (Kusche, J. A.); Portal (Alpert, G.); Portal (Raske, A. G.); Portal Mtn. (Nelson, J. W.); Portal, Cave Creek Ranch (Forister, G. W.); Ramsey Canyon [based on coordinates; no locality name cited], 1,700 m (Irwin; et al.); San Simon Rd., 4 mi NNW Portal, 1,400 m (Hespenheide, H. A.); Southwest Research Station, 8 km W Portal, 1,800 m (Alpert, G. D.); Southwestern Research Station, 1,680 m (Ward, P. S.); SW Res. Sta., Portal (Ewart, W. H.); Gila Co.: 18 km NNE Globe, 1,500 m (Ward, P. S.); Globe (Nutting); Jones Water Recr. Area, 17.3 mi N Globe (Schlinger, E. I.); Parker Ranch, Sixshooter Cyn. (Leech, H. B.; Green, J. W.); Graham Co.: Noon Creek, Pinaleña Mtns. (O’Brien, C. W.); Pinaleno Mts. (c. u.); Pinaleno Mts. [as ‘ Graham Mts., 3500 to 4500 ft. ’], 1,220 m (Bohart, R. M.); Post Canon, Pinaleno Mtns., 1,675 m (Wheeler, W. M.); La Paz: Colorado R., 1 mi S Ehrenberg (Smith, N. J.); Maricopa Co.: 0.5 mi N Pinnacle Peak, 745 m (Johnson, R. A.); Mohave Co.: Hualapai Mtns., s. of Kingman, 1,450 m (Schlinger, E.); Pima Co.: 3 km ESE Schuchuli (Aalbu, R. L.); Abra Wash, Growler Mtns., OCNM, 395 m (Creighton, W. S.); Alamo Cañon, Ajo Mtns., 670 m (Creighton, W. S.); Alamo Cañon, Ajo Mtns., 700 m (Creighton, W. S.); Alamo Cañon, Ajo Mtns., 760 m (Creighton, W. S.); Alamo Cañon, Ajo Mtns., 790 m (Creighton, W. S.); Brown Canyon, Baboquivari Mtns., 1,340 m (Creighton, W. S.); Forestry Cabin, Baboquivari Mtns., 1,065 m (Creighton, W. S.); Gates Pass, Tucson Mountain Park, 960 m (Ward, P. S.); Madera Canyon, Santa Rita Mts., 1,060 m (Schlinger, E. I.); Madera Cyn. (Nelson, G. H.); Organpipe (Melander, A. L.); Organpipe Cactus National Monument (Tinkham, E. R.); Sabino Basin, Sta. Catalina Mts. (c. u.); Sabino Cyn. (Vesterby, V. L.); San Miguel (Algert, E. D.); San Miguel (c. u.); Santa Rita Exp. Range (Rudgers, J. A.); Santa Rita Exp. Stn., nr. Green Valley, 1,360 m (Karban, R.); Sasabe (Oman); Tucson (Dietrich, A.; Dietrich, H.); Tucson (Knull, J.); Pima Co. or Pinal Co.: Santa Catalina Mts. (Crandall, R. H.); Sta Catalina Mtns. Fig. 4 – 6. Pseudomyrmex elongatulus group, workers, full-face dorsal view of head (a), and lateral profile of body (b). 4, P. apache, syntype, United States (CASENT 0902872); 5, P. arcanus, holotype, Mexico (CASENT 0863536); 6, P. championi, Mexico (CASENT 0246349). Images from AntWeb (www. antweb. org); photographers Zach (Ziv) Lieberman (4), Phil Ward (5), Leah Benuska (6). (Chrisman, M.); Pima Co. or Santa Cruz Co.: Catal Springs (Hubbard; Schwarz); Santa Rita Mtns. (Knull, J.); Santa Rita Mts. (Beamer, R. H.); Santa Rita Mts. (Bohart, R. M.); Santa Rita Mts. (Hubbard; Schwarz); Pinal Co.: 1.5 mi N Oak Flat Cpgrd., 1,235 m (Johnson, R. A.); 5 km NE Superior, 1,320 m (Ward, P. S.); Santa Cruz Co.: Canelo Pass, 1,615 m (Creighton, W. S.); Gooding Research Natural Area, Sycamore Canyon, 1,200 m (Ward, P. S.); Madera Canyon, Santa Rita Mtns. (Creighton, W. S.); Madera Cyn., 1,485 m (Kovac, C. R.); Madera Cyn., 1,485 m (Vesterby, V. L.); Madera Cyn. (Gilbert, A. J.; Smith, N. J.); Nogales (Burdine); Nogales (Knull, D. J.; Knull, J. N.); Pajarito Mtns., Sycamore Canyon, 1,220 m (Johnson, R. A.); Patagonia (Timberlake); Patagonia Mts., 2.6 mi W Harshaw (Leech, H. B.; Green, J. W.); Peña Blanca Springs, 1,130 m (Creighton, W. S.); Peña Blanca Springs (Byars, L. F.); Sweetwater, Santa Rita Mtns., 1,770 m (Creighton, W. S.); Sweetwater, Santa Rita Mtns., 1,830 m (Creighton, W. S.); Tumacacori Mtns. [as ‘ Tumacacori M. ’] (Knull, D. J.; Knull, J. N.); Walker Canyon, vic. Pena Blanca (Gilbert, A. J.; Smith, N. J.); Washington Camp [as ‘ Camp Washington, Patagonia Mts. ’] (c. u.); Yavapai Co.: Montezuma Castle, 977 m (Gadau, J. R.); Yuma Co.: Palm Cañon, Kofa Mtns., 670 m (Creighton, W. S.); Tacna (Hubbard, H. G.); California: Alameda Co.: Cedar Mtn. Rancho, Los Mochos Scout Cmp. (Gagne, W. C.); Butte Co.: 6 km N Feather Falls (town), 600 m (Ward, P. S.); Magalia, 730 m (Ward, P. S.); Colusa Co.: 1 km W Fouts Springs, 600 m (Ward, P. S.); 3 km W Fouts Springs, Fig. 7 – 9. Pseudomyrmex elongatulus group, workers and queens, full-face dorsal view of head (a), and lateral profile of body (b). 7, P. cognatus, holotype worker, Mexico (CASENT 0863537); 8, P. cognatus, dealate queen, Mexico (CASENT 0762787); 9, P. comitator, holotype queen, Mexico (JTLC 000010310). Images from AntWeb (www. antweb. org); photographer Phil Ward. 1,125 m (Ward, P. S.); 4 mi E Leesville (Levin, D. P.); Contra Costa Co.: 2 km NW Mount Diablo, 650 m (Ward, P. S.); Arroyo del Cerro, Mount Diablo, 375 m (Ward, P. S.); Knobcone Point, Mt. Diablo SP, 550 m (Ward, P. S.); Long Ridge, Mount Diablo State Park, 560 m (Ward, P. S.); El Dorado Co.: 14 km NW Shingle Springs, 340 m (Ward, P. S.); 15 km ENE Georgetown, 960 m (Ward, P. S.); 2 km ENE Georgetown, 880 m (Ward, P. S.); 9 km SW Pilot Hill, 340 m (Ward, P. S.); Fresno Co.: Prather, 470 m (Clark, J.); Glenn Co.: Open Ridge, 4 km N St. John Mountain, 990 m (Ward, P. S.); S end, Black Butte Lake, 150 m (Ward, P. S.); S end, Black Butte Lake, 150 m (Wild, A. L.); Imperial Co.: Andrade (Johnson, B.); El Centro (Paddock); Holtville (Snelling, R. R.); Seeley (Pineda, L.); Winterhaven (c. u.); Winterhaven (Westcott, R. L.); Lake Co.: 14 km ESE Middletown, 240 m (Ward, P. S.); 20 km ESE Lower Lake, 575 m (Fisher, B. L.); 20 km ESE Lower Lake, 575 m (Ward, P. S.); Borax Lake (Westcott, E. L.); Los Angeles Co.: 1.8 km ENE Topanga, 370 m (Ward, P. S.); 3 mi N Mt. Baldy, San Gabriel Mtns., 1,585 m (Weidert, E.); Altadena (c. u.); Eaton Canyon Park (Thompson, M. E.); Foothill, Pasadena (Sturtevant, A. H.); Glendale (Schlinger, E. I.); Griffith Park (Nisson); Los Angeles [as ‘ Ls Angls’] (Coquillett); Pasadena (c. u.); Pasadena (Hamsher, C. A.); S. Gabriel Mtns., 7 mi N Mt. Baldy, 1,585 m (Weidert, E.); San Gabriel Canyon (Flock, R. A.); San Pedro (Peterson, D. L.); Sister Elsie Peak (c. u.); SW of Valyermo, San Gabriel Mtns., 1,435 m (Henné, C. A.); Tanbark Flat (Bechtel, R. C.); Tanbark Flat (Bohart, R. M.); Tanbark Flat (Lawrence, T.); Tanbark Flat (MacSwain, J. W.); Tanbark Flat (Williams, F. X.); Los Angeles Co. or San Bernardino Co.: Mtns. near Claremont (Baker); Mariposa Co.: Mariposa (c. u.); Monterey Co.: 15 km SW Jolon, Fort Hunter Liggett Fig. 10 – 12. Pseudomyrmex elongatulus group, workers and queens, full-face dorsal view of head (a), lateral profile of body (b), and dorsal view of petiole, postpetiole and gaster (c). 10, P. elongatulus, syntype worker, Mexico (CASENT 0902874); 11, P. elongatulus, queen, Mexico (CASENT 0762874); 12, P. ereptor, holotype queen, Mexico (CASENT 0863524). Images from AntWeb (www. antweb. org); photographers Zach (Ziv) Lieberman (10), Phil Ward (11, 12). MR, 490 m (Ward, P. S.); Hastings Res. (Shelford, V. E.); Indians Rd., Los Padres Natl. Forest, 720 m (Ward, P. S.); Napa Co.: 4 km E summit Mt. St. Helena, 450 m (Ward, P. S.); 5 km W Oakville, 560 m (Ward, P. S.); Bothe-Napa Valley St. Pk., 270 m (Ward, P. S.); Cedar Roughs, 10 km ESE Pope Valley, 610 m (Ward, P. S.); Mt. George, 8 km NE Napa, 320 m (Ward, P. S.); Mt. St. Helena (Buckett, J. S.); Nevada Co.: Empire Mine SHP, 780 m (Ward, P. S.); Orange Co.: Cleveland Nat. For., Chiguto Trail, below Blue Jay, 838 m (Strawn, A. J.); Irvine Park (Brown, K.); Laguna Beach (Culbertson); Limestone Canyon, El Toro Rd., 1.8 mi E Cooks Corner (Suarez, A. V.); Orange (Shanafelt, J. G.); Ortega Hwy, 4.3 mi NE Hwy 5 (Andersen, H.); San Clemente (Melander, A. L.); Santa Ana (McDonald, W.); Starr Ranch, 520 m (Ward, P. S.); Tonner Cyn. (MacKay, W. P.); Trabuco Cyn (Irwin, M. E.); Trabuco east (Eidert, E.); Riverside Co.: 6 mi S Valle Vista, Sec. 9, T. 6 S, R. 1 E, 716 m (Pinto, J.; Frommer, S.); Ag Ops citrus orchard, Riverside (c. u.); Blythe (Hardman, R. M.); Blythe (Van Duzee, E. P.); Cactus Spring Trail, between Hwy. 74 & Horsethief Cr., Deep Canyon area (Frommer, S.); Deep Canyon (MacKay, W. P.); Dripping Springs, Agua Tibia Mtns., 455 m (Creighton, W. S.); Hwy. 243, San Jacinto Mts., 915 m (Burnett, J.; Wasbauer, M.); Keen Camp, San Jacinto Mtns. (Shanafelt, J. G.); Palms to Pines Hwy. (Ross, E. S.); Pinyon Flat [as ‘ Pinon Flat’], San Jacinto Mts (Macdonald, R. L.); Pinyon Flat [as ‘ Pinon Flat’], San Jacinto Mts (Ross, E. S.); Poppet Flats (Clark, G.); Riverside (Clancy, D.); Riverside (Hokke, Z.); Riverside (Schlinger, E. I.); Riverside (Timberlake); Santa Rosa Plat. Res. (PEET survey); The Gavilan (Timberlake); White Water [as ‘ Whitewater’] (Melander, A. L.); Winchester (Icenogle, W.); San Benito Co.: San Benito Mtn., 1,320 m (Ward, P. S.); San Bernardino Co.: 2 mi N Yucaipa (Smith, N.; Smith, E.); Deep Canyon (‘ E. D. W. ’); Deep Canyon (Niilns, T. W.); Mtn. Home Canyon (Timberlake); nr. Cajon Pass (c. u.); Oak Glenn, 1,065 m (Smith, N.); Yucaipa (Harp, G.; Cronk, W.); Yucaipa (Harper, G.); San Diego Co.: ‘ San Diego Co. ’ (Van Duzee, E. P.); 5.2 mi NW Ramona, Hwy. 78 (Frommer, S.; et al.); 5.9 mi NE Ramona, Hwy. 78 (Frommer, S.; et al.); 7 mi S Jct. 76 & 395 (Musgrove, C. H.); Campo (c. u.); Chula Vista (Bonita Long), 90 m (Suarez, A. V.); Chula Vista (Terranova), 78 m (Suarez, A. V.); Elliott Reserve, 150 m (Suarez, A. V.); Elliott Reserve, 1 km E Miramar, 180 m (Ward, P. S.); Jacumba (Knull, D. J.; Knull, J. N.); La Jolla (Todd, E. L.); La Mesa (Mathers, R.); no specific locality (c. u. [Cook Colln.]); Pine Hills Lodge, 1,300 m (Wild, A. L.); Point Loma Naval Res. (Hawks; Bruyea); Poway (Sixtus; Bowers); San Diego (Blaisdell, F. E.); San Diego (Van Duzee, E. P.); Scripps Beach, La Jolla (Trepanier, M.); Torrey Pines S. P. (Trepanier, M.); Torrey Pines State Pk. (Faulkner; Brown); Torrey Pines State Reserve, 75 m (Carney, S. E.); Vista (Metcalf, H.); Witch Creek (Rude, P. A.); San Luis Obispo Co.: Cuesta Ridge, San Luis Obispo (Ball; Gilbert); La Panza Cp., 12 mi NE Pozo (Toschi, C. A.); Santa Lucia Range, 2 mi NW Cuesta Pass (Toschi, C. A.); Selby Campground, Carrizo Plain NM, 800 m (Ward, P. S.); San Mateo Co.: Jasper Ridge, 150 m (Ward, P. S.); Santa Barbara Co.: 1 km W Field Stn., Santa Cruz I., 240 m (Ward, P. S.); Can. del Medio, Santa Cruz I. (Schuster, R. O.; Toftner, E. C.); Cascada, Santa Cruz I., 110 m (Ward, P. S.); El Tigre Ridge, Santa Cruz I. (Waugaman, R. D.); N end Sedgwick Ranch, 610 m (Ward, P. S.); Ranger Peak, Los Padres N. F., 1,110 m (Ward, P. S.); Santa Cruz Island, Upper Maripro (Wetterer, J. K.); Santa Ynez Superstar Ranch (Schlinger, E. I.); Sedgwick Ranch, 740 m (Schlinger, E.); Santa Clara Co.: ‘ Santa Clara Co. ’ (c. u.); Shasta Co.: 1 km WNW Lamoine, 630 m (Ward, P. S.); Solano Co.: Cold Canyon, 300 m (Ward, P. S.); Cold Canyon, 420 m (Ward, P. S.); Cold Canyon, 600 m (Ward, P. S.); Cold Canyon, 19 km NNW Vacaville, 120 m (Ward, P. S.); Cold Canyon, 19 km NNW Vacaville, 95 m (Ward, P. S.); G. L. Stebbins Cold Canyon Reserve (French, L. D.); Pleasants Ridge, 18 km NNW Vacaville, 240 m (Ward, P. S.); Sonoma Co.: 2 km ENE Glen Ellen, 150 m (Ward, P. S.); 3 km NW Camp Meeker, 120 m (Ward, P. S.); 8 km NNW Cazadero, 300 m (Ward, P. S.); Annadel State Pk., 360 m (Ward, P. S.); Mt. Hood, 820 m (Ward, P. S.); Sugarloaf Ridge St. Pk., 640 m (Ward, P. S.); Stanislaus Co.: Del Puerto Canyon, 18 km WSW Patterson, 300 m (Ward, P. S.); Sutter Co.: 1 km W North Butte, Sutter Buttes, 200 m (Ward, P. S.); Tehama Co.: 26 km WSW Red Bluff, 240 m (Ward, P. S.); Tulare Co.: 15 km NE Three Rivers, 900 m (Ward, P. S.); 9 mi S Fairview (Rude, P.); Ash Mountain, Kaweah Power Station # 3 (Hannel, M.); Ash Mtn., Kaweah Power Sta. # 3 (Bezark, L. G.); Ash Mtn., Kaweah Power Sta. # 3 (Burdick, D. J.); Ash Mtn., Kaweah Power Sta. # 3 (Halstead, J. A.); Ash Mtn., Sequoia Natl. Pk., 915 m (Fitton, M. G.); Ash Mtn., Three Rivers (Haines, R. D.); Horse Creek Rd. (Brawner, O. L.); Strathmore (Snelling, R. R.); Three Rivers (Pierce, D.); Tulare (Snelling, R. R.); Visalia (Mankins, E. O.); Tuolomne Co.: Sonora (Addiego); Ventura Co.: Big Sycamore Cyn., Pt. Mugu St. Park (Nagano, C.); Matilija Crk., T 5 N, R 24 W, s. 22, 500 m (Longino, J.); Saticoy (Barrett, R. E.); Yolo Co.: 13 km W Rumsey, 650 m (Fisher, B. L.); 3 km NE Guinda, 410 m (Ward, P. S.); 4 km NW Rumsey, 150 m (Ward, P. S.); 6 km WNW Rumsey, 190 m (Ward, P. S.); Guinda, 150 m (Ward, P. S.); New Mexico: Dona Ana Co.: La Cueva, Organ Mts. (c. u.); Hidalgo Co.: Deer Creek, Diamond A Ranch, 1,500 m (Andersen, A. N.); Texas: Bexar Co.: San Antonio, 230 m (Ward, P. S.); San Antonio (Ward, P. S.); Cameron Co.: Esprza Rch., Brownsville (c. u.); Duval Co.: Freer (Rogers, R. R.); San Diego (c. u.); Goliad Co.: [no specific locality] (Mitchell, J. D.); Hidalgo Co.: Monte Alto, 20 m (Creighton, W. S.); La Salle Co.: Fowlerton, 90 m (Creighton, W. S.); Maverick Co.: El Indio [as ‘ El Indigo’] (Bixby, D. H.); Presidio Co.: Arsaca Canyon, Chinati Mtns., 1,465 m (Creighton, W. S.); Big Bend Ranch SP, Las Cruces Amarillas (Ubick, D.); Real Co.: Camp Wood (Ward, C. R.); Starr Co.: [no specific locality] (Knull, D. J.; Knull, J. N.); Travis Co.: Austin, 150 m (Ward, P. S.); Austin (Ward, P. S.); Austin, Zilker Park, Rollingwood Drive, 155 m (Wild, A. L.); Uvalde Co.: [no specific locality] (Knull, D. J.; Knull, J. N.); Uvalde (Bradley, J. C.). Worker measurements (n = 17). HW 0.88 – 1.00, HL 1.08 – 1.21, MFC 0.039 – 0.066, LHT 0.77 – 0.87, CI 0.80 – 0.84, FCI 0.042 – 0.070, REL 0.40 – 0.44, REL 2 0.48 – 0.53, FI 0.38 – 0.44, PLI 0.55 – 0.61, PWI 0.50 – 0.57, MSC 3 – 8. Worker Diagnosis. Medium-sized species (HW 0.88 – 1.00) with moderately elongate head (CI 0.80 – 0.84) (Fig. 4) and relatively short eyes (see REL and REL 2 values); frontal carinae separated by about basal scape width or slightly less; metanotal groove present but weakly impressed; in profile, dorsal face of propodeum almost flat and more or less differentiated from declivitous face, rounding into the latter, and the two faces subequal in length; petiole relatively short and high (PLI 0.55 – 0.61, PL / HL 0.48 – 0.54), with a slight anterior peduncle; profemur slender; hind leg relatively short (LHT / HL 0.69 – 0.73). Head subopaque, densely punctulate-coriarious. Standing pilosity sparse, absent from propodeum and (nearly always) the mesonotum (MSC 3 – 8). Uniformly light yellow- to orange-brown, the tip of gaster usually infuscated. Comments. P. apache is closely related to P. championi, and our UCE phylogeny recovers it nested phylogenetically within that species (Fig. 1). The main distinction between the two taxa rests on body color: workers of P. apache are concolorous yellow- or orange-brown, whereas those of P. championi are bicolored with at least the gaster and hind leg dark brown and contrasting with the lighter orange-brown mesosoma (other body parts are usually dark brown but in some individuals they are concolorous with the mesosoma). The distinction is a slight one, but where the two species are geographically adjacent to one another in Sonora and Chihuahua (Figs. 19 and 21) the color differences appear to be sustained. For the moment, we treat P. apache as a distinct species, presumably recently derived from P. championi, in a daughter and parent relationship that renders P. championi paraphyletic at most loci. Nevertheless, further study of populations in northwest Mexico is desirable; it might reveal evidence of gene flow between the two taxa. P. apache can also be confused with P. arcanus, a more distantly related species in the P. elongatulus group, whose workers are similarly colored (light yellow- to orange-brown) but have more elongate eyes and a more gently rounded propodeum and petiole (see under P. arcanus for more discussion). Distribution and Biology. Pseudomyrmex apache occurs widely across the southwestern United States from northern California to east Texas, and south into adjacent northern Mexico (Fig. 19). This species inhabits a variety of xeric habitats, including chaparral, coastal sage scrub, mesquite scrub, oak woodland, and oakpine-juniper woodland, nesting in dead branches of various woody plants, especially Arctostaphylos, Baccharis, Prosopis, and Quercus. Colonies are polydomous — occupying multiple dead branches within a tree or shrub — and they contain one to several inseminated queens (Ward 1985). For locality records with elevation, the range is 20 m to 2,020 m (mean 905 m; n = 145). In addition to the examined material listed above, we have seen one old worker labeled as coming from Washington, DC. (24 Aug 1919, leg. B. P. Currie) (USNM). We judge this to be either mislabeled or representing an accidental introduction.	en	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 6 (1): 1-31, DOI: 10.1093/isd/ixab025, URL: http://dx.doi.org/10.1093/isd/ixab025
9A5C0805FFB50821F1A3F914CB06F8C6.taxon	description	Zoobank LSID: urn: lsid: zoobank. org: act: 5 CA 6 B 80 D-C 6 DA- 47 A 1 - 82 AD- 1721 A 990214 F	en	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 6 (1): 1-31, DOI: 10.1093/isd/ixab025, URL: http://dx.doi.org/10.1093/isd/ixab025
9A5C0805FFB50822F1A3F8ECC85AFB1E.taxon	materials_examined	Holotype Worker. MEXICO Michoacán: 20 km S Uruapán, 880 m, 19 ° 14 ′ N 102 ° 03 ′ W, 28 Dec 1987, ex dead twig of vine, tropical dry forest, P. S. Ward PSW 09293 (UNAM) (CASENT 0863536). Paratypes: series of workers, same data as holotype (CASC, CZUG, IEXA, MCZC, PSWC, UCDC, USNM). Other material examined (CASC, JTLC, LACM, MCZC, PSWC, RAJC, SEMC, UCDC, USNM). Mexico, Baja California: 19 km E El Arco, 400 m (Suarez, A. V.); 25 mi N El Arco (Ewart, W. H.); Palm Cyn., Angel de la Guardia Is. (Van Duzee, E. P.); Baja California Sur: 1 km NE La Burrera, 500 m (Ward, P. S.); 12 mi S Guillermo Prieto (Wasbauer, M.); 13 mi SW Guillermo Prieto (Phelps, B.; Phelps, M.); 19 mi SE El Cien (Andrews, F.; Faulkner, D.); 2 km WSW Todos Santos, 5 m (Ward, P. S.); 2.5 mi S Hwy. 1 on rd. to Pta Agua Verde, 325 m (Johnson, R. A.); 2.7 mi SE Valle Perdido (c. u. [Snelling?]); 28 – 29 km N Todos Santos, 275 m (Fisher, E.; Westcott, R.); 4.8 km N San Ignacio (Savary, W. E.; Mullinex, C. L.); 59 mi SE Guerrero Negro (Andrews, F.; Faulkner, D.); 7 mi NW Santa Rosalia, 260 m (Snelling, R. R.); 72 mi NW La Paz, 30 m (Snelling, R. R.); Bahia de Los Frailes (Ira LaRivers); Las Barracas (DeBach, P.); Playa Los Cerritos (Andrews, F.; Faulkner, D.); San Domingo (Ross; Bohart); San Jose del Cabo (c. u.); Triunfo (Michelbacher; Ross); Venancio (Michelbacher; Ross); Colima: 13 mi NE Comala, El Jabali (Ballmer, G.); 16 km NNE Comala, 1,280 m (Ward, P. S.); Jalisco: 20 mi W of Tecolotlán (Lipovsky, L. J.); 39 km N Colima, 970 m (MacKay, W. P.); Est. Biología Chamela, 100 m (Ward, P. S.); Mun. Zapotiltic, 24 km S Cd. Guzman, 1,095 m (MacDougal, J.); Tecolotlan (Villegas, B.); Michoacán: 20 km S Uruapán, 880 m (Ward, P. S.); Morelos: 7.3 mi S Yautepec, 1,066 m (Carney, L. B.); Nayarit: 13.9 mi E San Blas (Janzen, D. H.); 19.3 km S Rosamorada, 30 m (MacKay, W. P.); Isla Tres Marias, Maria Madre Id. Village (Keifer, H. H.); Oaxaca: Juquila, San Juan Mixtepec, 36 km W Puerto Escondido, 23 m (Kautz, S.); Puebla: 3 km SW El Salado, 850 m (da Silva, P. G.; Eager, T.); Sinaloa: 1 km NNE Chupaderos, Hwy 40, 400 m (Ward, P. S.); 23.5 mi SW El Palmito, mi 1704.8 (Janzen, D. H.); 4.2 mi S Villa Union on Hwy. 15 (Janzen, D. H.); 5 km E Concordia, 50 m (Ward, P. S.); Sonora: Rancho El Palmarito, 23 km E Alamos, 470 m (Irwin, M. E.); La Quintera (Timberlake). Worker measurements (n = 17). HW 0.82 – 0.97, HL 0.99 – 1.19, MFC 0.030 – 0.052, LHT 0.69 – 0.85, CI 0.80 – 0.85, FCI 0.036 – 0.055, REL 0.42 – 0.47, REL 2 0.52 – 0.58, FI 0.42 – 0.46, PLI 0.49 – 0.57, PWI 0.45 – 0.55, MSC 2 – 5. Worker Diagnosis. Small to medium-sized species (HW 0.82 – 0.97) with moderately elongate head (CI 0.80 – 0.85) and eyes (see REL and REL 2 values); frontal carinae separated by less than basal scape width; metanotal groove present but weakly impressed; in profile, dorsal face of propodeum rounding obtusely into declivitous face, the two faces subequal in length; petiole relatively slender (PLI 0.49 – 0.57, PL / HL 0.50 – 0.56), in profile the convex anterodorsal face of petiole ascending gradually to summit in the posterior half of node, then rounding into more rapidly descending posterior face (Fig. 5); profemur slender; hind leg relatively short (LHT / HL 0.68 – 0.72). Head sublucid, densely punctulate-coriarious to coriarious-imbricate. Standing pilosity sparse, absent from propodeum and mesonotum (MSC 2 – 5). Uniformly light yellow-brown to orange-brown. Comments. This species is characterized by a combination of light yellow-orange coloration, without additional maculation; moderately elongate eyes; weakly impressed metanotal groove (in the worker); and slender petiole. The workers of P. arcanus can be distinguished from those of other orange-yellow species in the P. elongatulus group as follows. Pseudomyrmex elongatulus has a better-developed metanotal groove, more elongate eyes (REL 0.47 – 0.53), petiole (PL / HL 0.57 – 0.61) and legs (LHT / HL 0.74 – 0.79), and some degree of infuscation on the gaster. Pseudomyrmex exoratus has a much more elongate head (CI 0.67 – 0.69) and a robust profemur (FI 0.46 – 0.49). Pseudomyrmex fasciatus essentially lacks a metanotal groove (as seen in in profile) and has conspicuous transverse maculation on the gaster. Pseudomyrmex apache has shorter eyes (on average), a more angular propodeum, and a more robust petiole, and the tip of the gaster is frequently darkened. Although they are not one another’s closest relatives (Fig. 1), P. apache and P. arcanus can be difficult to distinguish, the differences in propodeal and petiole shape being rather subtle. Bivariate plots of some measurements reveal two separate clouds of points (Fig. 3), but increased samples sizes might show some overlap. In our UCE phylogeny there is an early diverging sample of P. arcanus (D 1228) that is sister to all others (Fig. 1; Supp Fig. 4 [online only]). This is a southern outlier, from Oaxaca (Supp Table 2 [online only]), but we do not treat this as a separate species because 1) it is phenotypically similar to other samples of P. arcanus, 2) it is allopatric to the other populations, and 3) there are geographically intermediate populations from southern Mexico (Fig. 20) which we did not have an opportunity to sequence. More intensive sampling of such southern populations could be expected to reveal a paraphyletic series at the base of the P. arcanus tree, for which excisement of D 1228 would be arbitrary and ill-founded. Distribution and Biology. This species is found at low to medium elevations in western Mexico, from Baja California to Oaxaca. Recorded elevations vary from 5 m to 1,280 m (mean 474 m; n = 19). Habitats in which P. arcanus has been collected include coastal sand dunes, pasture, tropical dry forest, and mixed tropical / temperate mesic forest. Nests have been taken in a dead stalk of a woody Asteraceae (? Viguiera), dead stem of Cassia, dead twig of thorny vine, dead twig of vine, and dead twigs of unidentified woody plants. The records of ‘ Pseudomyrmex apache ’ cited in Ward (1985: 231) from Baja California refer to this species.	en	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 6 (1): 1-31, DOI: 10.1093/isd/ixab025, URL: http://dx.doi.org/10.1093/isd/ixab025
9A5C0805FFB60822F1A3FA82C985FA7B.taxon	description	Figs. 13 and 22 Zoobank LSID: urn: lsid: zoobank. org: act: B 01 C 7 CB 2 - 8 ACA- 47 D 3 - AE 4 D-E 4 C 4 F 4997 F 13	en	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 6 (1): 1-31, DOI: 10.1093/isd/ixab025, URL: http://dx.doi.org/10.1093/isd/ixab025
9A5C0805FFB60822F1A3FA82C985FA7B.taxon	materials_examined	Holotype Worker. MEXICO Jalisco: Est. Biología Chamela, 100 m, 19 ° 30 ′ N 105 ° 02 ′ W, 18 Dec 1987, ex dead twig of thorny vine, tropical dry forest, P. S. Ward PSW 09248 (UNAM) (CASENT 0863535). Paratypes: series of workers, same data as holotype (CASC, CZUG, IEXA, MCZC, PSWC, UCDC, USNM). Other material examined (EBCC, LACM, PSWC, UCDC). Mexico: Colima: Rio Tuxpan, 18 mi E Colima, 365 m (Dixon, J.; Heyer, R.); Jalisco: Chamela (Rodríguez, A.); Est. Biol. Chamela [as ‘ Búho’] (c. u.); Est. Biol. UNAM ‘ Chamela’ (Feener, D. H.); Est. Biología Chamela, 100 m (Ward, P. S.). Worker measurements (n = 6). HW 0.99 – 1.07, HL 1.18 – 1.32, MFC 0.040 – 0.058, LHT 0.83 – 0.92, CI 0.81 – 0.84, FCI 0.040 – 0.054, REL 0.44 – 0.46, REL 2 0.54 – 0.55, FI 0.41 – 0.46, PLI 0.54 – 0.57, PWI 0.47 – 0.51, MSC 25 – 39. Worker Diagnosis. Medium-sized species (HW 0.99 – 1.07) with moderately elongate head (CI 0.81 – 0.84) and eyes (see REL and REL 2 values); frontal carinae separated by slightly less than basal scape width; metanotal groove distinctly impressed; dorsal face of propodeum flat, elevated anteriorly above level of mesonotum, rounding into declivitous face, the two faces subequal in length; petiole relatively slender (PLI 0.54 – 0.57, PL / HL 0.54 – 0.56), in profile petiole with a slight anterior peduncle and with flat to convex anterodorsal face, ascending gradually to summit in the posterior half of node, then rounding into more rapidly descending posterior face; profemur slender; hind leg relatively short (LHT / HL 0.70 – 0.72). Head opaque, densely coriarious to coriarious-imbricate, punctulae very fine and inconspicuous. Standing pilosity common on head, mesosoma (including mesonotum and propodeum), petiole, postpetiole, and gaster (MSC 25 – 39, HTC 0, MTC 1 – 2). Body rather uniformly dark-brown, except scapes, mandibles, anterior quarter of head capsule, protibia, and protarsus, which are a contrasting light yellowish-brown. Comments. This species is notable for the conspicuous standing pilosity that covers the entire mesosoma dorsum (Fig. 13). Such pilosity is generally restricted to 1 – 3 pairs of setae on the worker pronotum in other members of the P. elongatulus group. Other salient features of the P. capillatus worker are the densely sculptured and opaque head, raised dorsal face of the propodeum, and unique color pattern (see Worker Diagnosis). Surprisingly, our UCE analyses show P. capillatus to be embedded phylogenetically within P. arcanus (Fig. 1), although the two are superficially quite different in appearance. Both species occur sympatrically at Estación Biología Chamela, the type locality of P. capillatus, but the Chamela sample of P. arcanus (D 1651) is not sister to the two P. capillatus samples. This suggests that P. arcanus, the more widespread of the two species, is in the process of achieving allelic monophyly relative to P. capillatus, but that this has not yet gone to completion. Distribution and Biology. Pseudomyrmex capillatus is thus far known only from Jalisco and Colima in western Mexico. In tropical dry forest at Estación Biología Chamela two nest series were collected by P. S. W., both in dead twigs of vines (the vines were 1 m apart, so the nests were likely part of a single colony), and a worker was collected foraging on low vegetation. Recorded elevations are 100 m and 365 m.	en	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 6 (1): 1-31, DOI: 10.1093/isd/ixab025, URL: http://dx.doi.org/10.1093/isd/ixab025
9A5C0805FFB60823F203FA69CE85FB06.taxon	materials_examined	Other material examined (AMNH, CASC, CUIC, CZUG, EMEC, IEXA, JTLC, KWJC, LACM, MCZC, MHNG, MNHN, MSNG, MZSP, OSAC, PSWC, SEMC, UCDC, UCRC, USNM). Mexico: Chihuahua: 6.5 km NW Batopilas, 1,550 m (Ward, P. S.); Mpio. Ocampo, 16 k E Basaseachic (Mackay, W.; Mackay, E.); Ciudad de México: Cd. de México, Jardín Botánico UNAM, 2,344 m (Dubovikoff); Colima: 16 km NNE Comala, 1,280 m (Ward, P. S.); Guanajuato: 5 mi S Salvatierra (Schlinger, E. I.); Guanajuato [as ‘ Guanaxuato’] (Dugès, E.); Guerrero: Xucumanatlan, 2,135 m (Smith, H. H.); Hidalgo: 11 mi W Jct. 45 & 85, 1,980 m (Scullen; Bolinger); 8 mi NE Jacala (Fisher, E. M.); 8 mi SW Jacala, 1,645 m (Snelling, R. R.); Jalisco: [no specific locality] (Krauss, N. L. H.); 1 km S El Fresnito (Rifkind, J.); 10 km S Autlán, 1,600 m (Ward, P. S.); 4 mi W Mazamitla, 2,070 m (Smith, R. F.); 7 mi S Manzamitla (Leech, H. B.); Guadalajara (c. u.); Lagos de Moreno, Sta. Rosa, 1,900 m (Vásquez, M.); Mascota, El Atajo, km. 15 camino a San Sebastian del O., 1,200 m (Vásquez, M.); Mixtlán, Cerro Chato, Carr. Ameca-Mascota, km. 48, 1,780 m (Vásquez, M.); Mun. Autlán de Navarro, SSE Autlán, 1,435 m (MacDougal, J.); San Ignacio, Cerro Gordo, 2,200 m (Navarrete, J. L.); Tepatitlán, carretera a Arandas, 1,950 m (Vásquez, M.); Tequila, Volcán de Tequila, 1,435 m (Pérez, D.); Zapopan, Santa Lucia, 1,642 m (Vásquez, M.); México: Valle de Bravo (Parker, F. D.); Michoacán: 12.3 mi E Morelia, 2,165 m (Snelling, R. R.); 15 – 20 mi W Jiquilpan (Leech, H. B.); 3 mi E Carapan (Parker, F. D.; Stange, L. A.); 8 km E Quiroga, 2,285 m (Evans, H. E.); Cotija (Villegas, B.); Morelia (Krauss, N. L. H.); Pátzcuaro [as ‘ Patzquero’] (Lipovsky, L. J.); Morelos: Chamilpa, 1,850 m (Quiroz, L.); Cuernavaca (Krauss, N. L. H.); Cuernavaca (Lipovsky, L. J.); Sto. Domingo, 1,900 m (Quiroz, L.); Tepotzlán (Olson, D. M.); Tlacotepec, Mpio. Zacualpan (Alemán, G.); Tlalnepantla, 2,030 m (Alemán, G.); Tres Marias [as ‘ Tres Maria’] (Pereira, P.); Zacualpan, 1,730 m (Alemán, G.); Oaxaca: 12 mi SE Oaxaca, 1,630 m (Scullen; Bolinger); 5 mi SE Oaxaca, 1,875 m (Scullen; Bolinger); 9 mi SE Nochistlan (Janzen, D. H.); Monte Alban (Malkin, B.); Monte Alban (Vaurie, P.; Vaurie, C.); Oaxaca (Malkin, B.); Puebla: Cacaloapan (Parker, F. D.); km. 275, Hwy. 150, NE of Chapulco (Cornell Univ. Mexico Field Party); Tehuacán (Janzen, D. H.); Sinaloa: 1.6 km NNE El Palmito, 2,130 m (Ward, P. S.); 6 mi NE Potrerillos (Parker, F. D.; Stange, L. A.); Potrerillos (Schlinger, E.); Sonora: La Quintera (Timberlake); Santa Barbara, near Alamos, 1,450 m (Franklin, K. A.); Veracruz: Acultzingo nr. Orizaba (Janzen, D. H.); Huatusco (Quiroz, L.). Worker measurements (n = 13). HW 0.82 – 1.05, HL 1.02 – 1.28, MFC 0.033 – 0.069, LHT 0.71 – 0.91, CI 0.80 – 0.87, FCI 0.040 – 0.071, REL 0.38 – 0.44, REL 2 0.46 – 0.55, FI 0.39 – 0.44, PLI 0.49 – 0.57, PWI 0.46 – 0.54, MSC 4 – 8. Worker Diagnosis. Small to medium-sized species (HW 0.82 – 1.05) with moderately elongate head (CI 0.80 – 0.87) (Fig. 6) and relatively short eyes (see REL and REL 2 values); frontal carinae separated by about basal scape width or less; metanotal groove present but weakly impressed; in profile, dorsal face of propodeum generally flat and more or less differentiated from declivitous face, rounding into the latter, and the two faces subequal in length; petiole moderately slender (PLI 0.49 – 0.57, PL / HL 0.52 – 0.56); in profile petiole with slight or no anterior peduncle, the anterodorsal face flat to convex, ascending gradually to summit in posterior half of node, then rounding into more steeply descending posterior face; profemur slender; hind leg relatively short (LHT / HL 0.69 – 0.75). Head opaque to subopaque, densely punctulate-coriarious, the punctulae on vertex usually separated by less than their diameters (but separated by more than their diameters in some populations, and the vertex correspondingly shinier). Standing pilosity sparse, absent from propodeum and mesonotum (MSC 4 – 8). Gaster, metacoxa, metafemur and metatibia dark brown, contrasting with light orange-brown mesosoma; head, first and second pairs of legs, petiole, and postpetiole varying from dark brown (usually) to much lighter and concolorous with mesosoma. Comments. The worker of this species is characterized by having a light-colored (orange-brown) mesosoma and contrasting dark gaster and hind leg. The head, other legs, petiole, and postpetiole also tend to be strongly infuscated, but in some samples (from Colima, Jalisco, and Chihuahua) they are lighter colored and even concolorous with the mesosoma. Workers of the closely related P. apache are uniformly orange-brown in color; see further discussion under that species. Pseudomyrmex championi has been confused with dark brown workers of a more distantly related species, P. cognatus, which occurs from southern Mexico to Nicaragua. The supposed descriptions of the larva (Wheeler and Bailey 1920, Wheeler and Wheeler 1956) and queen (Kempf 1961) of P. championi actually refer to P. cognatus. Workers of the two taxa can be distinguished as follows: P. championi is conspicuously bicolored, the dark gaster contrasting with the light orange-brown mesosoma, whereas P. cognatus is more uniformly dark brownish-black, with the mesosoma at most moderately lighter (and often dark on top); the metanotal groove is better developed (but still slight) in P. championi compared to P. cognatus; and the eye is generally shorter in P. championi than P. cognatus (REL 0.38 – 0.44 vs 0.43 – 0.48). The geographical distributions of P. championi and P. cognatus do not overlap (Figs. 21 and 22) but because they belong to different species complexes in the P. elongatulus group (Fig. 1), they cannot be treated as allopatric variants of the same species. Their similarities must reflect convergence or shared ancestral features. Distribution and Biology. Pseudomyrmex championi occurs in the Sierra Madre Occidental, Sierra Madre del Sur, and the intervening Transmexican Volcanic Belt (Fig. 21). Collection labels with habitat and / or elevation information document the occurrence of this species in oak woodland, oak-pine forest, old field / pasture, mixed tropical / temperate mesic forest, mid-montane dry forest, and yucca desert, at elevations ranging from 1,200 m to 2,344 m (mean 1,828 m; n = 28). Four nests of P. championi encountered by one of us (P. S. W.) were all from dead twigs of woody shrubs (one Asteraceae, one Verbenaceae, and two unidentified plants).	en	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 6 (1): 1-31, DOI: 10.1093/isd/ixab025, URL: http://dx.doi.org/10.1093/isd/ixab025
9A5C0805FFB70824F203FAB4CE77F940.taxon	description	Zoobank LSID: urn: lsid: zoobank. org: act: A 7 ABFF 2 F- 818 B- 4 FE 9 - B 143 - 380786 E 61 C 48	en	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 6 (1): 1-31, DOI: 10.1093/isd/ixab025, URL: http://dx.doi.org/10.1093/isd/ixab025
9A5C0805FFB70824F203FAB4CE77F940.taxon	materials_examined	Holotype Worker. MEXICO Chiapas: Tziscao, Lagos de Montebello, 1,500 m, 16 ° 05 ′ N 91 ° 41 ′ W, 21 Dec 1991, ex dead twig of vine, mixed tropical / temperate mesic forest, P. S. Ward PSW 11560 (UNAM) (CASENT 0863537). Paratypes: series of workers, 1 dealate queen, same data as holotype (CASC, CZUG, IEXA, JTLC, MCZC, PSWC, UCDC, USNM). Other material examined (AMNH, CASC, CEET, CNCC, CSCA, FSCA, JTLC, LACM, MCZC, MZLU, MZSP, PSWC, SEAN, SMPC, UCDC, USNM, UTIC, UVGC). El Salvador: Santa Ana: Monte Cristo (Cartwright, O. L.); Guatemala: Alta Verapaz: 1 km NE Santa Cruz Verapaz, 1,400 m (Ward, P. S.); 7 km SW Cobán, 1,460 m (Ward, P. S.); airport near Cobán, 1,320 m (Janzen, D. H.); Tactic, Finca Xilicá (Andrade, L.); Tactic, Finca Xilicá (Haeussler, A.); Baja Verapaz: 1 km SSW La Cumbre, 1,480 m (Ward, P. S.); 2 km N La Unión Barrios, 1,620 m (Ward, P. S.); 3 km ESE Purulhá, 1,620 m (Ward, P. S.); 4 km WSW Purulhá, 1,700 m (Ward, P. S.); Chimaltenango: km. 143 a Chimalte (Perez, R.); Escuintla: ex San José (in quarantine San Francisco, USA) (c. u.); Guatemala: ex Guatemala City (in quarantine San Francisco, USA) (c. u.); Guatemala City (Bloem, K.; Bloem, S.); Retalhuleu: Puente Samala, 3.8 mi NE San Felipe, Quezal-Retal. Rd. (Janzen, D. H.); Sacatepéquez: Antigua, 2,000 m (Ekis, G.); Antigua (Bloem, S.); Antigua (Krauss, N. L. H.); Parque Florencia, nr. San Miguel Milpas Altas, 1,900 m (Ward, P. S.); Santa Rosa: Barberena, El Naranjito (Perez, R.); Sololá: 1 km N San Andrés Semetabaj, 1,840 m (Ward, P. S.); 2 km ESE San Lucas Tolimán, 1,780 m (Wild, A. L.); 7 km SSE San Lucas Tolimán, 1,140 m (Ward, P. S.); Lake Atitlan (Wheeler, W. M.); Panajachel, 1,573 m (Stange, L. A.); San Lucas Toliman (Wheeler, W. M.); Tsanjuyo (c. u.); unknown: ex ‘ Guatemala’ (in quarantine San Francisco, CA, USA) (c. u.); Zacapa: 2 km SE La Unión, 1,550 m (LLAMA); San Lorenzo, 1,800 m (LeSage, L.); Honduras: Comayagua: 10 km E Comayagua, 1,730 m (LLAMA); 10 km E Comayagua, 2,000 m (Boudinot, B.); 9 km E Comayagua, 1,730 m (LLAMA); Cortés: P. N. Cusuco, 5 km N Buenos Aires (Hansson, C.); PN Cusuco, 1,220 m (LLAMA); Francisco Morazán: Cerro Uyuca, 2,000 m (Gupta, V.; Gupta, S.); Montañita [as ‘ Montañito’] (Hubbell, T. H.); Olancho: 9 km N Catacamas, 1,270 m (LLAMA); 9 km N Catacamas, 1,330 m (LLAMA); 9 km N Catacamas, 1,340 m (LLAMA); 9 km NNW La Unión, 1,460 m (Ward, P. S.); 9 km NNW La Unión, 1,470 m (Ward, P. S.); Parq. Nac. La Muralla, 1,450 m (Ward, P. S.); Parq. Nac. La Muralla, 1,470 m (Ward, P. S.); PN La Muralla, 1,440 m (LLAMA); PN La Muralla, 1,450 m (LLAMA); PN La Muralla, 1,460 m (LLAMA); PN La Muralla, 1,480 m (LLAMA); PN La Muralla, 1,490 m (LLAMA); Mexico: Chiapas: 1 km E Tulanca (Pérez, L.); 2 km SE Custepec, 1,520 m (LLAMA); 3 km E San Cristóbal (Jones, R. W.); 36 mi N inters. Hwy. 190 × Tux. Gut. - Vil. Her. Rd., km 58 (Janzen, D. H.); 5 km E Rayón, 1,700 m (Ward, P. S.); km 44 Tuxtla Gutierrez to San Cristobal, near Chiapa de Corzo (Whitacre, D. F.); Laguna Montebello, 1,430 m (O’Brien, C. W.; O’Brien, L.; Marshall); Río Yashanal (Girón, M.); San Miguel, Mpio. El Bosque, 1,105 m (Philpott, S. M.); Tziscao, Lagos de Montebello, 1,500 m (Ward, P. S.); Tzuluwitz, Mpio. San Juan Cancuc, 1,310 m (Philpott, S. M.); Yevalchen, Mpio. Tumbala, 1,350 m (Philpott, S. M.); Zinacantan, 1,900 m (Ekis, G.); Nicaragua: Jinotega: 5 km [from] Jinotega, Los Pinares (Maes; Pineda); PN Cerro Saslaya, 1,110 m (LLAMA); Santa Maria de Ostuma (Maes; Pineda); Matagalpa: Fuente Pura (Maes, J. M.; et al.); Hotel Selva Negra, km 139 N of Matagalpa, 1,200 m (Kugler, C.; Hahn, J.); Reserva Miraflor, 1,369 m (MacKay, W.); Santa Martha, 8 km N Matagalpa, 1,219 m (Ross, E. S.); Selva Negra, 1,285 m (MacKay, W.); Nueva Segovia: 9 km NW Jalapa, 1,410 m (LLAMA); 9 km NW Jalapa, 1,470 m (LLAMA). Worker measurements (n = 14). HW 0.93 – 1.06, HL 1.07 – 1.22, MFC 0.040 – 0.057, LHT 0.82 – 0.94, CI 0.82 – 0.88, FCI 0.039 – 0.058, REL 0.45 – 0.48, REL 2 0.52 – 0.56, FI 0.40 – 0.44, PLI 0.49 – 0.57, PWI 0.44 – 0.52, MSC 2 – 5. Worker Diagnosis. Medium-sized species (HW 0.93 – 1.06) with moderately elongate head (CI 0.82 – 0.88) (Fig. 7) and relatively large eyes (see REL and REL 2 values); frontal carinae separated by less than basal scape width; metanotal groove weakly impressed, in profile barely discernable; dorsal face of propodeum flat and more or less differentiated from declivitous face, rounding into the latter, and the two faces subequal in length; petiole moderately slender (PLI 0.49 – 0.57, PL / HL 0.53 – 0.59); in profile petiole with slight or no anterior peduncle, the anterodorsal face flat to convex, ascending gradually to summit in posterior half of node, then rounding into more steeply descending posterior face; profemur slender; hind leg relatively short (LHT / HL 0.71 – 0.77). Head subopaque, densely punctulate-coriarious, the punctulae on vertex usually separated by less than their diameters. Standing pilosity sparse, absent from propodeum and mesonotum (MSC 2 – 5). Head, postpetiole, gaster and legs dark brown, mesosoma and petiole varying from concolorous to lighter medium-brown, not strongly contrasting, however, with rest of body (mesosoma sometimes darkened dorsally); mandibles, antennae, and distal portions of legs tending to be a lighter medium-brown to yellowish-brown. Comments. This is a widespread and rather common Mesoamerican species that was previously confused with P. championi, although the two are not sister taxa — in fact, UCE data demonstrate that they belong to different complexes within the P. elongatulus group (Fig. 1). Pseudomyrmex cognatus can be recognized by the combination of dark brown coloration of most of the body (mesosoma often lighter but not contrastingly so); densely punctulate-coriarious head; relatively large eyes (worker REL 0.45 – 0.48, queen REL 0.42 – 0.46); and weakly developed metanotal groove (in the worker). For distinctions between P. cognatus and P. championi, see under the latter species. Workers from a single nest collection from Parque Florencia, Sacatepéquez, Guatemala (PSW 15039) have more widely separated frontal carinae (MFC 0.065 – 0.070; n = 3), shorter eyes (REL 0.43 – 0.45), and a broader and shorter petiole (PWI 0.56 – 0.58, PL / HL 0.50 – 0.53). In the UCE phylogeny this sample is sister to all others of P. cognatus (Fig. 1). We have no other P. cognatus workers from this site, i. e., no sympatric association with more ‘ typical’ P. cognatus that would confirm PSW 15039 as a distinct species. Hence there is some ambiguity about the status of these divergent individuals. For the moment we treat them as conspecific with P. cognatus, while noting that further study might support their treatment as a different species. Distribution and Biology. Pseudomyrmex cognatus ranges from Chiapas, Mexico to Nicaragua, and has been recorded from mixed tropical / temperate mesic forest, montane rainforest, montane rainforest edge, oak-pine forest, mixed pine-mesophyll forest, oak cloud forest, roadside, and old field / pasture, at elevations ranging from 1,105 m to 2,000 m (mean 1,507 m; n = 47). Nests have been collected from dead twigs of various woody plants, including Asteraceae, Fabaceae, Orchidaceae, and Pinus; one was collected from a dead sedge culm.	en	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 6 (1): 1-31, DOI: 10.1093/isd/ixab025, URL: http://dx.doi.org/10.1093/isd/ixab025
9A5C0805FFB00825F203F974CB45F920.taxon	description	LSID: urn: lsid: zoobank. org: act: 4 BAA 462 E- 31 A 4 - 4799 - BEF 6 - D 7 BC 23 B 346 F 0	en	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 6 (1): 1-31, DOI: 10.1093/isd/ixab025, URL: http://dx.doi.org/10.1093/isd/ixab025
9A5C0805FFB00825F203F974CB45F920.taxon	materials_examined	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 JTL 6094 (UNAM) (JTLC 000010310). 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 JTL 6093 - 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; REL 2 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, REL 2 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 punctulate-coriarious. 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.	en	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 6 (1): 1-31, DOI: 10.1093/isd/ixab025, URL: http://dx.doi.org/10.1093/isd/ixab025
9A5C0805FFB10826F1A3F951CE4CFBB6.taxon	materials_examined	Other material examined (AMNH, BMNH, CASC, CEET, CHAH, CSCA, CUIC, CZUG, FSCA, IEXA, JTLC, LACM, MCZC, MNHN, MZSP, PSWC, SEMC, SMPC, UCDC, UCRC, USNM, UTEP, UWEM). Belize: Cayo: Belmopan (Krauss, N. L. H.); Chiabul Forest, 5 km SE Millionario (Beard, J.); Chiabul Forest, San Pastor (Beard, J.); El Cayo (Krauss, N. L. H.); Las Cuevas, 5 km SE Millionario (Beard, J.); Las Cuevas, 5 km SE Millionario (Lyal, C.; Hollis, D.); Orange Walk: Rio Bravo Conserv. Area, La Milpa Res. Stn. (Davis, L. R.); Guatemala: Escuintla: 6.6 mi NE Escuintla (Janzen, D. H.); Guatemala: ex Guatemala City (in quarantine San Francisco, USA) (c. u.); Peten: Tikal Mayan ruins, 183 m (Ross, E. S.); Petén: Tikal (Hubbell, T. H.); unknown: ex ‘ Guatemala’ (in quarantine San Francisco, CA, USA) (c. u.); Mexico: Chiapas: 8 km SE Salto de Agua, 100 m (LLAMA); 8 km SE Salto de Agua, 60 m (LLAMA); 8 km SE Salto de Agua, 70 m (LLAMA); Lago Metzabok, 560 m (LLAMA); Lago Metzabok, 570 m (LLAMA); Lago Metzabok, 575 m (LLAMA); Laguna Bélgica (Girón, M.); Nahá, 860 m (LLAMA); Nahá, 950 m (LLAMA); Palenque ruins, 170 m (Cox, D. J.); Simojovel to Santo Domingo (Chemsak, J. A.); Tonalá [as ‘ Tonola’] (Petrunkewitch); Yevalchen, Mpio. Tumbala, 1,335 m (Philpott, S. M.); Morelos: Huautla (Quiroz, L.); Oaxaca: 11 – 17 mi W Tehuantepec (Janzen, D. H.); 22 mi S Jesus Carranza (Graber, J. W.); 5 mi E Temascal (Janzen, D. H.); Café Carlota, 950 m (Gray, K. W.); Temascal (Janzen, D. H.); Uluapan, 4 km NE Ayautla, 410 m (Longino, J.); Uluapan, 4 km NE San Bartolomé Ayautla, 420 m (ADMAC); Uluapan, 4 km NE San Bartolomé Ayautla, 470 m (ADMAC); Uluapan, 4 km NE San Bartolomé Ayautla, 500 m (ADMAC); Uluapan, 4 km NE San Bartolomé Ayautla, 510 m (ADMAC); Uluapan, 4 km NE San Bartolomé Ayautla, 620 m (ADMAC); Puebla: Villa Juarez (intercepted at Brownsville, Texas) (Heinrich; Jackson); San Luis Potosí: 13 km N Tamazunchale, 290 m (MacKay, W.); 84 km W Cd. Valle, 645 m (MacKay, W.); Ciudad Valles, Carr. al Naranjo, Salto Micos, 168 m (Vásquez, M.); El Salto, 520 m (Carney, L.); Tamasopo, Cascadas El Paraiso, 400 m (Vásquez, M.); Tamazunchale, 120 m (Creighton, W. S.); Tamazunchale, 245 m (Creighton, W. S.); Tamazunchale (Ross, W. S.); Tamazunchale (intercepted at Brownsville, Texas) (Heinrich); Tamazunchale (intercepted at Laredo, Texas) (c. u.); Tamazunchale (POE: Brownsville, Texas) (Arsego); Xilitla, Arroyo La Conchita, 431 m (Vásquez, M.); Tabasco: 13 km W border Chis., Rt. 186 (MacKay, W.); 3 mi W Cardenas (Janzen, D. H.); Cardenas (Janzen, D. H.); Teapa (Shenefelt, R. D.); Teapa (Smith, H. H.); Tamaulipas: 10 km W El Encino, 510 m (Ward, P. S.); Gomez Farias (MacKay, W.); Tampico (Locke); Veracruz: 10 mi E Conejos (Janzen, D. H.); 10 mi W Veracruz (Bohart, G. E.); 11 km NNE Sontecomapan, 175 m (Ward, P. S.); 15 km SW Paso del Toro, 50 m (Ward, P. S.); 29.5 mi NW Tuxpam (Janzen, D. H.); 3 km NW La Tinaja, 50 m (MacKay, W.); 6 km NW Tecolapán, 55 m (Kautz, S.; Eilmus, S.); 8 km NNE Soteapan, 1,010 m (Ward, P. S.); 8 km NNE Soteapan, 970 m (Ward, P. S.); 9 km NE San Andrés Tuxtla, 965 m (Ward, P. S.); 9 km NNW Sontecomapan, 20 m (Ward, P. S.); Apazapan, 347 m (Quiroz, L.); Apazapan (Sivinski, J.); Camaron (Skwarra, E.); Catemaco Hills, 8 mi S Angel (Janzen, D. H.); Cordoba (Bohart, G. E.); Córdoba, Cuauhtémoc (Navarrete, J. L.); Cotaxtla Exp. Stn., Cotaxtla (Janzen, D. H.); Est. Biol. ‘ Los Tuxtlas’, nr. San Andres Tuxtla (Ibarra M., G.); Est. Biol. de Los Tuxtlas (Hespenheide, H. A.); Est. Biol. La Mancha, 20 m (ADMAC); Est. Biol. Los Tuxtlas, 110 m (ADMAC); Est. Biol. Los Tuxtlas, 140 m (ADMAC); Est. Biol. Los Tuxtlas, 160 m (ADMAC); Est. Biol. Los Tuxtlas, 170 m (ADMAC); Est. Biol. Los Tuxtlas, 180 m (ADMAC); Est. Biol. Los Tuxtlas, 450 m (ADMAC); Estación de Biología Los Tuxtlas, 115 m (Ward, P. S.); Estación de Biología Los Tuxtlas, 150 m (Ward, P. S.); Estación de Biología Los Tuxtlas, 170 m (Ward, P. S.); Estación de Biología Los Tuxtlas, 450 m (Ward, P. S.); Fortin, 950 m (Ward, P. S.); Fortín de las Flores (Edwards, G. B.); Fortín de las Flores - Sumidero, Planta de la Cerveceria, 840 m (Weems, Jr., H. V.); La Mancha (c. u.); Las Hamacas, 17 km N Santiago Tuxtla (Wilson, E. O.); Los Tuxtlas, 140 m (Kautz, S.; Heil, M.); Los Tuxtlas (Jeanne, R. L.); Los Tuxtlas, 10 km NNW Sontecomapan, 200 m (Ward, P. S.); Los Tuxtlas, 10 km NNW Sontecomapan, 500 m (Ward, P. S.); Mirador (Skwarra); Pueblo Nuevo, nr. Tetzonapa (Cornell Univ. Mexico Field Party); Pueblo Nuevo, nr. Tetzonapa (Wilson, E. O.); Rio Tonto (Janzen, D. H.); Ruiz Cortínez, 12 km NE San Andrés Tuxtla, 1,050 m (ADMAC); Sa. Teoviscocla, nr. Cuichapa (Cornell Univ. Mexico Field Party); San Andrés Tuxtla (Janzen, D. H.); St. Lucrecia (Knab, F.); St. Lucrecia (Mann, W. M.); Tuxpango-Orizaba (Gillogly, A. R.); Vera Cruz (Townsend); Veracruz (Krauss, N. L. H.); unknown: ‘ Mexico’ (c. u.); ‘ Mexique’ (Smith, H. H.); United States: Florida: Palm Beach Co.: MacArthur Beach SP (Deyrup, M.). Worker measurements (n = 14). HW 0.93 – 1.09, HL 1.03 – 1.32, MFC 0.034 – 0.045, LHT 0.80 – 0.97, CI 0.81 – 0.90, FCI 0.033 – 0.046, REL 0.47 – 0.53, REL 2 0.54 – 0.61, FI 0.43 – 0.47, PLI 0.44 – 0.52, PWI 0.41 – 0.48, MSC 2 – 4. Worker Diagnosis. Medium-sized species (HW 0.93 – 1.06, LHT 0.80 – 0.97); head only moderately elongate (CI 0.81 – 0.90) and with rounded posterolateral corners in full-face view (Fig. 10); eyes relatively large (see REL and REL 2 values); frontal carinae separated by less than basal scape width; metanotal groove strongly impressed, conspicuous in profile; dorsal face of propodeum flat, rounding into declivitous face, the two subequal in length; petiole slender, elongate-triangular in profile (PLI 0.47 – 0.53, PL / HL 0.57 – 0.61, PL / LHT 0.75 – 0.80); petiole with slight anterior peduncle, in profile the anterodorsal face flat to convex, ascending gradually to summit in posterior quarter of node, then rounding into steeply descending posterior face; profemur moderately robust (FI 0.43 – 0.47); hind leg moderately long (LHT / HL 0.74 – 0.79). Head subopaque to sublucid, densely punctulate-coriarious, the punctures becoming less dense on vertex (separated by their diameters or more). Standing pilosity sparse, absent from propodeum and mesonotum (MSC 2 – 4). Typically light yellow-brown to orange-brown, with darker brown anterolateral patches on abdominal tergite 4 (first gastric tergite) and brown transverse bands on abdominal tergites 5 – 7 (gastric tergites 2 – 4); in some samples from Guatemala and Belize head and mesosoma more infuscated, and gaster tending to be more uniformly medium brown. Comments. Distinctive features of this species are the moderately elongate head with rounded posterolateral corners, large eyes (worker REL 0.47 – 0.53, queen REL 0.44 – 0.46), conspicuous metanotal groove in the worker (Fig. 10), and slender elongate-triangular petiole (worker PLI 0.44 – 0.52, queen PLI 0.45 – 0.51). Pseudomyrmex elongatulus is typically light yellow-brown to orange-brown with darker transverse maculation on the gaster, but some specimens from Guatemala and Belize are darker brown overall. Leaving aside P. ereptor (see Pseudomyrmex ereptor sp. nov.), the closest relative of P. elongatulus is P. exoratus (Fig. 1), easily told apart by its much more elongate head (worker CI 0.67 – 0.69, vs 0.81 – 0.90 in P. elongatulus). Distribution and Biology. This species is widely distributed in eastern Mexico, from Tamaulipas to Chiapas, with outlying populations in Guatemala and Belize. The record from Morelos needs to be confirmed since it lies outside the expected range of this species. It may represent a transient introduction. There are records of P. elongatulus being intercepted at U. S. ports of entry in Texas and California, from shipments originating in Puebla, San Luis Potosí, and Guatemala (see list of material examined, above), indicating that this species has a propensity to be transported by human commerce. A single worker collected recently by Mark Deyrup in a natural setting in south Florida evidently represents an accidental introduction. It is unclear if P. elongatulus is established in that state; a second visit to the site failed to locate additional workers (Deyrup, personal communication). P. elongatulus has been recorded from lowland rainforest, rainforest edge, second-growth rainforest, mixed tropical / temperate mesic forest, shaded coffee, tropical dry forest, and roadside habitats, at elevations ranging from 20 m to 1355 m (mean 417 m; n = 53). Nests are found in dead twigs of various plants. Specific nest site records include dead stalk of grass, dead twig of liana, and dead twigs / stems of Cecropia, Heliocarpus appendiculatus, Gliricidia sepium, Mimosa, Orchidaceae, woody Asteraceae, and unidentified woody plants.	en	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 6 (1): 1-31, DOI: 10.1093/isd/ixab025, URL: http://dx.doi.org/10.1093/isd/ixab025
9A5C0805FFB20827F203FB2AC8BBFC4B.taxon	description	Zoobank LSID: urn: lsid: zoobank. org: act: 3 F 4584 C 3 - EB 0 C- 4 C 74 - B 067 - 01 CBEDC 8 F 4 EB 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 PSW 07360 A (UNAM) (CASENT 0863524). 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, REL 2 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, REL 2 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 (D 1985) 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 PSW 07360) 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.	en	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 6 (1): 1-31, DOI: 10.1093/isd/ixab025, URL: http://dx.doi.org/10.1093/isd/ixab025
9A5C0805FFB30827F1A3FC79C9ABFC00.taxon	description	Figs. 14 and 24 Zoobank LSID: urn: lsid: zoobank. org: act: 8 D 3 D 17 CF- 131 E- 4 E 3 F- 9 C 67 - B 9 B 67 E 475 B 1 D	en	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 6 (1): 1-31, DOI: 10.1093/isd/ixab025, URL: http://dx.doi.org/10.1093/isd/ixab025
9A5C0805FFB30827F1A3FC79C9ABFC00.taxon	materials_examined	Holotype Worker. MEXICO Oaxaca: Temascal, 25 m, 2 Feb 1964, at house, D. H. Janzen (UNAM) (CASENT 0863539). Paratypes: series of workers, same data as holotype (CASC, CZUG, IEXA, JTLC, LACM, MCZC, PSWC, UCDC, USNM). Other material examined (LACM, MZLU, PSWC, UCDC). Honduras: Atlántida: Lancetilla, Tela (Cave, R.); Mexico: Chiapas: Palenque ruins, 170 m (Cox, D. J.); Oaxaca: 5 mi E Temascal (Janzen, D. H.); Temascal, 25 m (Janzen, D. H.); Temascal (Janzen, D. H.). Worker measurements (n = 6). HW 0.83 – 0.86, HL 1.19 – 1.28, MFC 0.043 – 0.049, LHT 0.78 – 0.84, CI 0.67 – 0.69, FCI 0.050 – 0.058, REL 0.42 – 0.45, REL 2 0.62 – 0.66, FI 0.46 – 0.49, PLI 0.52 – 0.56, PWI 0.49 – 0.55, MSC 3 – 4. Worker Diagnosis. Medium-sized species (LHT 0.78 – 0.84), with notably elongate head (CI 0.67 – 0.69) and eyes (see REL and REL 2 values); frontal carinae separated by basal scape width or slightly less; ocelli prominent (Fig. 14); metanotal groove conspicuously impressed; dorsal face of propodeum rounding insensibly into declivitous face, the two subequal in length; petiole relatively slender (PLI 0.52 – 0.56, PL / HL 0.51 – 0.52), in profile the weakly convex anterodorsal face of petiole ascending gradually to summit in the posterior half of node, then rounding into steeper posterior face; profemur relatively robust (FI 0.46 – 0.49); hind leg short, relative to head length (LHT / HL 0.65 – 0.67). Head sublucid, anterior half densely punctulate with shiny interspaces, becoming sparsely punctulate-coriarious on vertex. Standing pilosity sparse, absent from propodeum and mesonotum (MSC 3 – 4). Light orange-brown, with dark brown anterolateral patches on abdominal tergite 4 (first gastric tergite) and a transverse band on abdominal tergite 6 (gastric tergite 3); distal half of anterior face of metafemur weakly infuscated. Comments. This light orange-brown species is easily recognized by its exceptionally elongate head (worker CI 0.67 – 0.69, queen CI 0.55 – 0.59), which is unlike that of any other species in the P. elongatulus group. Pseudomyrmex exoratus also has a rather robust profemur (worker FI 0.46 – 0.49, queen FI 0.47 – 0.49) and distinctive maculation on the gaster: dark anterolateral patches on abdominal tergite 4 and a transverse band on abdominal tergite 6. Our UCE phylogeny shows P. exoratus to be closely related to the more common and widespread species, P. elongatulus (Fig. 1). Both species occur sympatrically at the type locality of P. exoratus. Distribution and Biology. Pseudomyrmex exoratus is known from only three low-elevation localities in southern Mexico (in Oaxaca and Chiapas) and one collection from Honduras. Janzen collected two series at Temascal, Oaxaca, labeled 2. ii. 1964 II and 2. ii. 1964 IV (the latter including alate queens and males), plus a single queen on 10 Jan 1964, for which the label states ‘ flew into car, at house’. Little is known about the biology of this species although it is presumed to nest in dead twigs, like most other members of the P. elongatulus group. The robust profemur suggests that it might occupy rather hard dead wood.	en	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 6 (1): 1-31, DOI: 10.1093/isd/ixab025, URL: http://dx.doi.org/10.1093/isd/ixab025
9A5C0805FFB30828F203FBBAC8E3F870.taxon	description	Figs. 15 and 20 Zoobank LSID: urn: lsid: zoobank. org: act: 52 CB 59 B 3 - D 99 C- 4 E 09 - 8 A 83 - 522 B 3 E 21714 F Holotype Worker. COSTA RICA Puntarenas: 2 km SSE Monteverde, 1,230 m, 10.289 06 − 84.80304 ± 3 m, 5 Jan 2018, ex dead twig of vine, montane rainforest edge, P. S. Ward PSW 18011 (MUCR) (CASENT 0863540). Paratypes: series of workers, 1 dealate queen, same data as holotype (CASC, JTLC, LACM, MCZC, PSWC, UCDC, UNAM, USNM). Other material examined (AMNH, INBC, JTLC, LACM, MCZC, MIZA, MUCR, PSWC, SMPC, UCDC, USNM).	en	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 6 (1): 1-31, DOI: 10.1093/isd/ixab025, URL: http://dx.doi.org/10.1093/isd/ixab025
9A5C0805FFB30828F203FBBAC8E3F870.taxon	materials_examined	Costa Rica: Alajuela: Rio Peñas Blancas, 950 m (Longino, J.); Zarcero, 1,600 m (Solis, A.); Cartago: Agua Caliente, nr. Cartago (Wheeler, W. M.); Nombre Dulce, vivero Linda Vista, 1,300 m (Hanson, P.); Heredia: Monte Allegro, Sto. Domingo de Heredia (Perfecto, I.); Puntarenas: 2 km SSE Monteverde, 1,230 m (Ward, P. S.); 2 km SSW Monteverde, 1,245 m (Ward, P. S.); 3 km SE Monteverde, 1,200 m (Longino, J.); 3 km SSE Monteverde, 1,100 m (Longino, J.); Monteverde, 1,100 m (Ward, P. S.); Monteverde, 1,200 m (Cover, S.); Monteverde, 1,200 m (Ward, P. S.); Monteverde, 1,400 m (Longino, J.); Monteverde (Daly, H.); San Luis de Monteverde, 1,100 m (LaPierre, L.); San Luis Valley, 1,090 m (Sumnicht, T. P.; Longino, J. T.); Sta. Elena, 1,300 m (Longino, J.); vic. Alto San Luis, 1,100 m (Hovore, F. T.); San José: San Pedro de Montes de Oca (Ballou, C. H.); El Salvador: La Libertad: Ecoparque El Espino (Mendoza, E.); Guatemala: Chimaltenango: El Amparo (c. u.); Escuintla: 6.6 mi NE Escuintla (Janzen, D. H.); Guatemala: ex Guatemala City (in quarantine San Francisco, USA) (c. u.); Santa Rosa: El Piño (Sivinski, J.); unknown: ex ‘ Guatemala’ (intercepted at Hoboken, NJ, USA) (c. u.); Honduras: Atlántida: 9 km S Yaruca, 950 m (Ward, P. S.); Cortés: Lago Yojoa, N end (Brown, W. L.); PN Cusuco, 1,210 m (LLAMA); PN Cusuco, 1,330 m (LLAMA); Mexico: Chiapas: 15 mi NW Ocozocoautla, 790 m (Newton, A.); Finca Hamburgo, Río Tepuzapa, 1,030 m (Andresen, D.); Finca Irlanda, 975 m (Philpott, S. M.); Nahá, 860 m (LLAMA); Nahá, 950 m (LLAMA); Sierra Morena, 1,380 m (LLAMA); Nicaragua: Matagalpa: RN Cerro Musún, 1,090 m (LLAMA); RN Cerro Musún, 750 m (LLAMA); RN Cerro Musún, 880 m (LLAMA); RN Cerro Musún, 900 m (LLAMA). Worker measurements (n = 12). HW 0.84 – 0.94, HL 1.05 – 1.15, MFC 0.019 – 0.042, LHT 0.74 – 0.83, CI 0.79 – 0.86, FCI 0.023 – 0.045, REL 0.45 – 0.48, REL 2 0.54 – 0.59, FI 0.41 – 0.47, PLI 0.46 – 0.58, PWI 0.42 – 0.50, MSC 2 – 5. Worker Diagnosis. Small to medium-sized species (HW 0.84 – 0.94) with moderately elongate head (CI 0.79 – 0.86) (Fig. 15) and relatively large eyes (REL 0.45 – 0.48); frontal carinae separated by less than basal scape width; metanotal groove weakly impressed, in profile barely discernable; dorsal face of propodeum flat, rounding into declivitous face, and the two subequal in length; petiole relatively slender (PLI 0.46 – 0.58, PL / HL 0.54 – 0.60); in profile, anterodorsal face of petiole flat to convex, ascending gradually to summit in posterior half of node, then rounding into more steeply descending posterior face; profemur generally slender (FI 0.41 – 0.47); hind leg relatively short (LHT / HL 0.67 - 0.76). Head subopaque to sublucid, densely punctulate-coriarious, the punctulae on vertex separated by less than their diameters. Standing pilosity sparse, absent from propodeum and mesonotum (MSC 2 – 5). Light yellow- to orange-brown, with dark brown anterolateral patches on abdominal tergite 4 (first gastric tergite), sometimes joined medially, and conspicuous transverse bands on abdominal tergites 5 and 6 (gastric tergites 2 and 3); distal half of metafemur weakly (and variably) infuscated. Comments. Among species in the P. elongatulus group that are predominantly yellow-orange to light orange-brown in color, this species can be recognized by the combination of relatively large eyes (worker REL 0.45 – 0.48, queen REL 0.45 – 0.46), moderately elongate head (worker CI 0.79 – 0.86, queen CI 0.72 – 0.76), weakly developed metanotal groove in the worker, and conspicuous transverse maculation on abdominal tergites 4 – 6 (gastric tergites 1 – 3). Our phylogenomic data reveal that P. fasciatus is sister to P. cognatus (including its phylogenetically embedded inquiline, P. comitator) (Fig. 1). Pseudomyrmex fasciatus and P. cognatus are broadly sympatric and quite different in appearance, with workers and queens of P. cognatus being brown to brownish-black in color. Distribution and Biology. Pseudomyrmex fasciatus occurs from Chiapas, Mexico to Costa Rica. Recorded habitats include montane rainforest edge, montane wet forest, montane moist forest, and second-growth forest edge, at elevations ranging from 750 m to 1,600 m (mean 1,115 m; n = 28). Nests have been collected in dead twigs of various woody plants, including vines and orchids. Two series of specimens intercepted in quarantine at San Francisco, from Guatemala, are labeled ‘ ex Oncidium wentworthianum ’ and ‘ ex Epidendrum atropurpurium ’, respectively. A third interception, at Hoboken, New Jersey was from an Oncidium stem.	en	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 6 (1): 1-31, DOI: 10.1093/isd/ixab025, URL: http://dx.doi.org/10.1093/isd/ixab025
9A5C0805FFBC0829F203FF39CAA2FDC6.taxon	description	Figs. 16 and 26 Zoobank LSID: urn: lsid: zoobank. org: act: A 03034 A 9 - FADF- 4 AD 5 - 9990 - 6 E 13516 BD 6 A 2	en	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 6 (1): 1-31, DOI: 10.1093/isd/ixab025, URL: http://dx.doi.org/10.1093/isd/ixab025
9A5C0805FFBC0829F203FF39CAA2FDC6.taxon	materials_examined	Holotype Worker. COSTA RICA Alajuela: Río Peñas Blancas, 800 m, 10 ° 19 ′ N, 84 ° 43 ′ W, 22 – 25 May 1990, ex live stems Inga sapling, wet forest clearing, J. Longino JTL 2694 (MUCR) (CASENT 0863541). Paratypes: series of workers, same data as holotype (CASC, JTLC, LACM, MCZC, PSWC, UCDC, UNAM, USNM). Other material examined (ANSP, BMNH, GCSC, JTLC, LACM, PSWC, UCDC). Costa Rica: Alajuela: Peñas Blancas, 940 m (Longino, J.); Rio Peñas Blancas, 800 m (Longino, J. T.); Rio Peñas Blancas, 950 m (Longino, J.); Cartago: 4 km E Moravia, 1,200 m (Longino, J.); La Fuente, e. slope of volcano of Turrialba, 1,200 m (Alfaro, A.); Turrialba, km. 22, to Limón (King, A. B. S.); Heredia: Rara Avis, 730 m (Rifkind; Gum). Worker measurements (n = 8). HW 1.03 – 1.20, HL 1.18 – 1.41, MFC 0.034 – 0.053, LHT 0.96 – 1.09, CI 0.85 – 0.89, FCI 0.031 – 0.047, REL 0.48 – 0.51, REL 2 0.55 – 0.59, FI 0.41 – 0.45, PLI 0.44 – 0.53, PWI 0.42 – 0.50, MSC 2 – 5. Worker Diagnosis. Relatively large species (HW 1.03 – 1.20, LHT 0.96 – 1.09); head only moderately elongate (CI 0.85 – 0.89) and with rounded posterolateral corners in full-face view (Fig. 16); eyes relatively large (REL 0.48 – 0.51); frontal carinae separated by less than basal scape width; metanotal groove strongly impressed, conspicuous in profile; dorsal face of propodeum generally convex in profile, rounding into declivitous face, the two subequal in length; petiole slender, elongate-triangular in profile (PLI 0.44 - 0.53, PL / HL 0.58 – 0.60, PL / LHT 0.71 – 0.79); petiole with slight anterior peduncle, in profile the anterodorsal face flat to convex, ascending gradually to summit in posterior third of node, then rounding into more steeply descending posterior face; profemur relatively slender (FI 0.41 – 0.45); hind leg relatively long (LHT / HL 0.76 – 0.82). Head subopaque, densely punctulate-coriarious, the punctures becoming less dense on vertex (separated by their diameters or more). Standing pilosity sparse, absent from propodeum and mesonotum (MSC 2 – 5). Head, dorsum of propodeum, metafemur, and gaster dark brown, other parts of body with variably lighter coloration (medium brown to yellow-brown). Comments. Pseudomyrmex nimbus can be recognized by the combination of large size (worker HW 1.03 – 1.20; queen HW 1.15 – 1.18), well-developed metanotal groove (in worker), and predominantly dark coloration. The relatively long legs (worker LHT / HL 0.76 – 0.82, queen LHT / HL 0.70 – 0.72) and elongate eyes (worker REL 0.48 – 0.51; queen REL 0.48) are also notable. This species is in the P. elongatulus complex (within the P. elongatulus group), and is a distant sister to the trio of P. exoratus, P. elongatulus, and P. ereptor (Fig. 1). Distribution and Biology. Pseudomyrmex nimbus is known only from several cloud forest sites in Costa Rica, at elevations of 730 m to 1,200 m. The following are Jack Longino’s field notes on the type series (JTL 2694): ‘ Workers abundant on small Inga sapling. I dissected the entire tree, finding 3 nests excavated in live stems (there was also stem-borer damage here and there, so the ants could have been invading pre-existing cavities). I never found reproductives. Either the queen was little different from workers and escaped, or the colony was queenless, or there was more of the colony off the Inga tree. ’ Another nest series (JTL / 5 Jul 84 / 1017) was collected ‘ inside branch of still live but recently felled Inga ’ and included workers, alate queens, and a male. A third collection by Longino (JTL 0900) was taken in a live Cecropia insignis sapling. Thus, the available data suggests that this species is a live-stem nester, a habit not seen in the majority of Pseudomyrmex species (Ward 1991). There is no evidence that P. nimbus keeps trophobionts (hemipterans) in these nests.	en	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 6 (1): 1-31, DOI: 10.1093/isd/ixab025, URL: http://dx.doi.org/10.1093/isd/ixab025
9A5C0805FFBD082AF1A3FDF1C847FEBB.taxon	description	Figs. 17 and 25	en	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 6 (1): 1-31, DOI: 10.1093/isd/ixab025, URL: http://dx.doi.org/10.1093/isd/ixab025
9A5C0805FFBD082AF1A3FDF1C847FEBB.taxon	materials_examined	Other material examined (AMNH, CASC, CEET, CHAH, CSCA, EMEC, GKMC, IEXA, JTLC, LACM, MCZC, MNHN, MSNG, MZLU, MZSP, NHMW, PSWC, SMPC, STDC, UCDC, UCRC, USNM, UVGC). Belize: Belize: Belize (c. u.); Big Falls Road (Mosser, G. K.); Monkey Bay W. S. (Mosser, G. K.); Cayo: El Cayo (Krauss, N. L. H.); nr. Macal River, 10 mi S Augustine (Reiskind, J.); nr. Teakettle Bank, Pooks Hill, 85 m (Bartlett, C. R.); Pine Mtn. Ridge, Rubber Camp, Mecal River, 415 m (Alpert, G. D.); Stann Creek: Cockscomb Basin Jaguar Preserve, 414 m (Alpert, G. D.); Mama Noots, Bocawina, 16 km W Dangriga, 83 m (Keller, M. F.); Guatemala: Alta Verapaz: Cacao, Trece Aguas (Barber; Schwarz); Region of Sebal, headwtrs Rio Pasión. (Hubbs; van der Schalie); Guatemala: Mixco (Mann, W. M.); Izabal: 16 km ESE Morales, 440 m (LLAMA); 16 km ESE Morales, 530 m (LLAMA); Bobos [as ‘ Bobas’] (Mann, W. M.); Lago Izabal, 1.5 km NE El Estor (Janzen, D. H.); Livingston (Barber; Schwarz); Los Amates (Kellerman); Parque Arqueológico, 84 m (Mackay, W.; Mackay, E.); Polochic R. (Barber; Schwarz); Petén: 13 km NW Machaquilá, 375 m (LLAMA); Sololá: Tsanjuyo (c. u.); Zacapa: Gualán, Finca La Cartuchera (Vasquez, J.); Honduras: Comayagua: PN Cerro Azul Meambar, 750 m (LLAMA); PN Cerro Azul Meambar, 770 m (LLAMA); Cortés: Lago de Yojoa, 625 m (Krauss, N. L. H.); P. N. Cusuco, 5 km N Buenos Aires (Hansson, C.); Yoro: Pico Pijol, 2,200 m (Hansson, C.); Mexico: Chiapas: 21 km SW Salto de Agua, 180 m (LLAMA); 4 mi S Simojovel (Schlinger, E. I.); 4 mi SW Simojovel (Bechtel, R. C.; Schlinger, E. I.); Lago Metzabok, 560 m (LLAMA); Laguna Bélgica (Girón, M.); Nahá, 860 m (LLAMA); Palenque Ruins (Goodnight, C. J.); Plan Paredon, Mpio. Simojovel, 990 m (Philpott, S. M.); Playón de la Gloria, 160 m (LLAMA); Playón de la Gloria, 170 m (Cox, D. J.); Playón de la Gloria, 180 m (LLAMA); Oaxaca: Uluapan, 4 km NE San Bartolomé Ayautla, 400 m (ADMAC); Uluapan, 4 km NE San Bartolomé Ayautla, 420 m (ADMAC); Puebla: Cascada Las Hamacas, 8 km ENE Cuetzalan, 290 m (ADMAC); Quintana Roo: Rives Caraibes, entre Puerto Juares et Touloum (Carayon, J.); Tabasco: 3 mi W Cardenas (Janzen, D. H.); Teapa (Smith, H. H.); Veracruz: 10 km N Sontecomapan, 10 m (Ward, P. S.); 4 mi N Catemaco (Fisher, E. M.); 6 mi NE Catemaco, 455 m (Newton, A.); 8 mi NW Sontecomapan, 150 m (Newton, A.); Est. Biol. de Los Tuxtlas (Hespenheide, H. A.); Est. Biol. Los Tuxtlas, 110 m (ADMAC); Est. Biol. Los Tuxtlas, 120 m (ADMAC); Est. Biol. Los Tuxtlas, 140 m (ADMAC); Est. Biol. Los Tuxtlas, 150 m (ADMAC); Est. Biol. Los Tuxtlas, 160 m (ADMAC); Est. Biol. Los Tuxtlas, 170 m (ADMAC); Est. Biol. Los Tuxtlas, 180 m (ADMAC); Est. Los Tuxtlas (Quiroz, L.); Estación de Biología Los Tuxtlas, 115 m (Ward, P. S.); Estación de Biología Los Tuxtlas, 150 m (Ward, P. S.); La Buenaventura (Petrunkewitch); Lago Catemaco (Gillogly, A. R.); Las Hamacas, 17 km N Santiago Tuxtla (Wilson, E. O.); Los Tuxtlas, 160 m (Kautz, S.; Heil, M.); Los Tuxtlas (Jeanne, R. L.); Los Tuxtlas Res. Stn. (Sivinski, J.); Los Tuxtlas, 10 km NNW Sontecomapan, 200 m (Ward, P. S.); Playa Escondita, nr. San Andreas Tuxtlas (Meyer, D.); Pueblo Nuevo, nr. Tetzonapa (Wilson, E. O.); St. Lucrecia (Mann, W. M.); Yucatán: env. Uxmal (Carayon, J.). Worker measurements (n = 10). HW 1.12 – 1.21, HL 1.19 – 1.35, FCI 0.038 – 0.055, LHT 0.96 – 1.12, CI 0.90 – 0.96, FCI 0.032 – 0.047, REL 0.54 – 0.59, REL 2 0.59 – 0.62, FI 0.37 – 0.41, PLI 0.43 – 0.49, PWI 0.43 – 0.49, MSC 2 – 4. Worker Diagnosis. Relatively large species (HW 1.12 – 1.21, LHT 0.96 – 1.12) with broad head and large eyes (see REL and REL 2 values); frontal carinae separated by less than basal scape width; pronotum laterally submarginate, flattened, and with a weak longitudinal median depression; metanotal groove obsolete to absent; in profile propodeum short and high, the dorsal face shorter than, and rounding insensibly into, the declivitous face; petiole elongate and gracile, with a well differentiated anterior peduncle (Fig. 17) (PLI 0.43 – 0.49, PL / HL 0.64 – 0.69); profemur slender; hind leg relatively long (LHT / HL 0.80 – 0.84). Head varying from sublucid to (in more southern populations) subopaque; anterior half of head between eyes densely punctulate-coriarious, punctures becoming sparser on posterior half (separated by more than their diameters), and with intervening reticulate-coriarious sculpture of varying reflectance. Standing pilosity sparse, confined on the mesosoma dorsum to the pronotum (MSC 2 - 4). Head dark blackish-brown, remainder of body varying from contrastingly yellow-orange (except for transverse infuscated bands on abdominal tergites 4 – 6) to medium brown, appendages generally lighter. Comments. Pseudomyrmex salvini is a distinctive species, sister to all other members of the P. elongatulus group (Fig. 1). It is easily recognized by its large size, elongate eyes (more than one-half head length), broad head (worker CI 0.90 – 0.96, queen CI 0.81 – 0.84), short high propodeum (in the worker), and elongate petiole with well-developed anterior peduncle (Fig. 17). In populations from Veracruz the head contrasts strikingly with the lighter colored remainder of the body; farther southward and eastward the mesosoma and metasoma become increasingly infuscated and the contrast is muted. Distribution and Biology. This species is confined to southeastern Mexico, Guatemala, Belize, and Honduras, where it occurs in or at the edge of mesic forest, at elevations ranging from 10 m to 990 m (mean 316 m; n = 35). One worker, from Pico Pijol, Honduras (leg. C. Hansson) (MZLU), has the elevation given as ‘ 2200 m’, but this is likely to be an error. Habitats recorded on specimen labels include littoral vegetation, rainforest, rainforest edge, mesophil forest, and roadside. Among the material examined by us, there were only four explicit nest series: ‘ ex dead twig of liana’, ‘ ex dead weed stalk’, ‘ ex dead twig’, and ‘ nest in twig’. Most collections consist of workers or dealate queens foraging on low vegetation. At Los Tuxtlas, Veracruz one of us (P. S. W.) observed a salticid spider, apparently the darkcephalic morph of Synemosyna decipiens (Cambridge) (see Cutler 1985: 87), mimicking workers of P. salvini.	en	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 6 (1): 1-31, DOI: 10.1093/isd/ixab025, URL: http://dx.doi.org/10.1093/isd/ixab025
9A5C0805FFBE082AF203FF1CCEC6FE63.taxon	distribution	Distribution and Biology. Pseudomyrmex veracruzensis is known only from two adjacent sites in the Sierra de los Tuxtlas, Veracruz. Collections include two nest series from dead twigs of Heliocarpus, and isolated workers foraging on low vegetation, all at the edge of montane rainforest (1,010 m – 1,055 m).	en	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 6 (1): 1-31, DOI: 10.1093/isd/ixab025, URL: http://dx.doi.org/10.1093/isd/ixab025
9A5C0805FFBE082AF1A3FE29CFC6FF4B.taxon	description	Figs. 18 and 26 Zoobank LSID: urn: lsid: zoobank. org: act: BB 9 D 186 E- 1313 - 44 F 6 - A 9 A 6 - F 63 AB 60653 B 2 Holotype Worker. MEXICO Veracruz: Ruiz Cortínez, 13 km NE San Andrés Tuxtla, 1,055 m, 18.53387 – 95.13839 ± 4 m, 5 Jun 2016, ex dead twig of Heliocarpus, montane rainforest edge, P. S. Ward PSW 17618 (UNAM) (CASENT 0863542). Paratypes: series of workers, same data as holotype (CASC, CZUG, IEXA, JTLC, MCZC, PSWC, UCDC, USNM). Other material examined (PSWC, UCDC).	en	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 6 (1): 1-31, DOI: 10.1093/isd/ixab025, URL: http://dx.doi.org/10.1093/isd/ixab025
9A5C0805FFBE082AF1A3FE29CFC6FF4B.taxon	materials_examined	Mexico: Veracruz: 10 km NNE San Andrés Tuxtla, 1,010 m (Ward, P. S.); Ruiz Cortínez, 13 km NE San Andrés Tuxtla, 1,050 m (Ward, P. S.); Ruiz Cortínez, 13 km NE San Andrés Tuxtla, 1,055 m (Ward, P. S.). Worker measurements (n = 8). HW 0.97 – 1.07, HL 1.13 – 1.30, MFC 0.035 – 0.050, LHT 0.84 – 0.94, CI 0.81 – 0.86, FCI 0.034 – 0.048, REL 0.46 – 0.48, REL 2 0.55 – 0.58, FI 0.42 – 0.45, PLI 0.44 – 0.49, PWI 0.43 – 0.48, MSC 2 - 5. Worker Diagnosis. Medium-sized species (HW 0.97 – 1.07) with moderately elongate head (CI 0.81 – 0.86) and eyes (REL 0.46 – 0.48); frontal carinae separated by less than basal scape width; metanotal groove moderately impressed, evident in profile; dorsal face of propodeum more or less flat, rounding into declivitous face, the two subequal in length; petiole slender, elongate (PLI 0.44 – 0.49, PL / HL 0.57 – 0.60, PL / LHT 0.76 – 0.82); petiole with slight anterior peduncle, in profile the anterodorsal face flat to convex, ascending gradually to summit in posterior half of node, then rounding into more steeply descending posterior face; profemur relatively slender (FI 0.42 – 0.45); hind leg relatively short (LHT / HL 0.71 – 0.76). Head subopaque, densely punctulate-coriarious, the punctures becoming less dense on vertex (separated by their diameters or more). Standing pilosity sparse, absent from propodeum and mesonotum (MSC 2 – 5). Medium ferrugineus-brown, with darker and rather diffuse transverse bands on anterior half of abdominal tergites 4 – 6 (gastric tergites 1 – 3); metafemur and, to a lesser extent, mesofemur infuscated. Comments. Key features of this species are the ferrugineus-brown body, with darker transverse bands on the gaster; relatively large size (worker HW 0.97 – 1.07); and low, elongate petiole (worker PL 0.65 – 0.77, worker PLI 0.44 – 0.49). Workers of P. veracruzensis are superficially similar to those of the Costa Rican species, P. nimbus, but can be distinguished by their shorter legs (LHT 0.84 – 0.94 and LHT / HL 0.71 – 0.76 vs LHT 0.96 – 1.09 and LHT / HL 0.76 – 0.82 in P. nimbus), smaller eyes (REL 0.46 – 0.48 vs 0.48 – 0.51 in P. nimbus), less strongly impressed metanotal groove, differently shaped petiole (compare Figs. 16 and 18), and lighter coloration. An index combining PL, LHT and HW also separates the two species: (PL / LHT) / HW 0.74 – 0.80 versus 0.62 – 0.71 in Pseudomyrmex nimbus. P. veracruzensis is somewhat isolated phylogenetically, as sister to	en	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 6 (1): 1-31, DOI: 10.1093/isd/ixab025, URL: http://dx.doi.org/10.1093/isd/ixab025
