Temnothorax pergandei ( Emery, 1895 )

Prebus, Matthew M., 2021, Taxonomic revision of the Temnothorax salvini clade (Hymenoptera: Formicidae), with a key to the clades of New World Temnothorax, PeerJ (e 11514) 9, pp. 1-462 : 240-253

Temnothorax pergandei ( Emery, 1895)

Distribution: Fig. 127B ; worker, gyne & male: Fig. 129 ; worker variability: Fig. 130 ; gyne variability: Fig. 131 .

Leptothorax (Dichothorax) pergandei Emery, 1895: 323 , pl. 8, fig. 13. Syntype workers. Washington, D.C., U.S.A. One syntype worker here designated lectotype.

Leptothorax (Dichothorax) floridanus Emery, 1895: 323 . Syntype workers. Junior synonym of Temnothorax pergandei by Mackay, 1993: 289. One syntype here designated lectotype. Leptothorax (Dichothorax) pergandei Emery : Wheeler, 1903 b: 257: gyne and male. Leptothorax (Dichothorax) pergandei flavus Smith, 1929: 549 . Syntype workers. Adaton, Mississippi. Junior synonym of Temnothorax pergandei by Creighton, 1950: 260.

One syntype worker here designated lectotype.

Leptothorax (Dichothorax) manni Wesson, 1935: 208 . Syntype workers, gyne, and male. Nashville, Tennessee. Junior synonym of Temnothorax pergandei by Wesson, 1939: 180. One syntype worker here designated lectotype.

Leptothorax (Dichothorax) pergandei Emery : Wheeler & Wheeler, 1960: 23: larva.

Temnothorax pergandei (Emery) : Bolton, 2003: 272. First combination in Temnothorax .

Type material examined: Lectotype worker of Leptothorax (Dichothorax) pergandei : U.S.A.: District of Columbia: Washington, D.C.: September 1910, [no collector data], no. 54078 U.S. N.M. cotype, (USNMENT00922524) [ USNM].

Paralectotype workers of Leptothorax (Dichothorax) pergandei : same data as lectotype: 1 worker (USNMENT00922508) [ USNM] 1 worker (USNMENT00922509) [ USNM] 1 worker (USNMENT00922515) [ USNM] 1 worker (USNMENT00922517) [ USNM] 1 worker (USNMENT00922518) [ USNM] 1 worker (USNMENT00922510) [ USNM] 1 worker (USNMENT00922511) [ USNM] 1 worker (USNMENT00922522) [ USNM] 1 worker (USNMENT00922523) [ USNM] 1 worker (USNMENT00922525) [ USNM] 1 worker (USNMENT00532783) [ USNM] 1 worker (images of CASENT0904767 examined on antweb. org) [ MSNG] .

Lectotype worker of Leptothorax (Dichothorax) pergandei flavus : U.S.A.: Mississippi: Oktibbeha County: Adaton, [no collection date], M.R. Smith, cotype 52535 U.S. N.M., (USNMENT00921850, top specimen) [ USNM].

Paralectotype workers and gyne of Leptothorax (Dichothorax) pergandei flavus : same pin as lectotype (bottom specimen) [ USNM]; same data as lectotype: 1 dealate gyne (USNMENT00529209) [ USNM] 2 workers (USNMENT00529210) [ USNM] 3 workers, top missing gaster, middle missing head (USNMENT00921851) [ USNM] 3 workers (USNMENT00921852) [ USNM] 3 workers (USNMENT00921849) [ USNM] .

((( Lectotype worker of Leptothorax (Dichothorax) floridanus : U.S.A.: Florida: Highlands County: Ft. George, 18 July 1882, [no collector data], No. 37, No. 54079, U.S. N.M. cotype (USNMENT00532780) [ USNM].

Paralectotype workers of Leptothorax (Dichothorax) floridanus : same data as lectotype: 1 worker (USNMENT00921856) [ USNM] 1 worker (USNMENT00921857) [ USNM] 1 worker (images of CASENT0904768 examined on antweb.org) [ MSNG] .

Lectotype worker of Leptothorax (Dichothorax) manni : U.S.A.: Tennessee: Davidson County: Nashville, [no collection date], [no collector data] (top specimen on pin, missing gaster) (images of USNMENT00529639 examined on the Smithsonian National Museum of Natural History Department of Entomology Collections website) [ USNM].

Paralectotype gyne and male of Leptothorax (Dichothorax) manni : same pin as lectotype: 1 male, 1 dealate gyne (images of USNMENT0529639 examined on the Smithsonian National Museum of Natural History Department of Entomology Collections website) .

Non-type material examined: MEXICO: Hidalgo: 1.8 km NE Tezontepec , 20.20574 ° N 99.27112 ° W ± 50 m, 1,990 m, 18 July 2005, P. Hernández #05IIaTna311, xerophyll scrub, pitfall, 3 workers (CASENT0608344- CASENT0608346) [ JTLC] . Nuevo León: 33 mi east of General Bravo , 25.93661 ° N 98.674200 ° W, 245 m, 12 March 1953, W.S. Creighton, 4 workers (CASENT0759990) [ FSCA] .

U.S.A.: Alabama: Colbert County: Cane Creek Canyon Nature Preserve, 34.62333 ° N 87.80278 ° W, 245 m, 2 June 2014, J.A. MacGown, upland hardwood forest, 1 worker ( MEM 243844) [ MEM]; Mobile County: Citronelle, 31.09046 ° N 88.22798 ° W, 95 m, 11 June 1924, W.S. Creighton, 3 workers (CASENT0867001) [ FSCA]. Arizona: Cochise County: 1 km S junction of Route 533 on State Line Road, 11 September 1997, 1,280 m, S.P. Cover#SPC4993, overgrazed desert with scattered Ephedra , mesquite and Acacia , bare hole entrance in Hilaria patch, fine sand w/ clay, 1 male & 1 worker (MCZENT00583595) [ MCZC]; Chiricahua Mountains, Cave Creek Canyon, 0.6 km N of Southwestern Research Station, 31.88883 ° N 109.20433 ° W, 1,740 m, 15 August 2001, S.P. Cover#SPC6341, oak-juniper woodland, nest in base of grass clump, 1 worker (CASENT0758667) [ UCDC]; Chiricahua Mountains, Paradise cemetery, 31.93167 ° N 109.20833 ° W, 1,682 m, 4 August 2001, R.A. Johnson#RAJ2539, juniper grassland, small tumulus nest, 3 workers (CASENT0869114) [ RAJC] 4 workers (CASENT0869115) [ RAJC]; Chiricahua Mountains, Tex Canyon, 31.68167 ° N 109.32 ° W, 1,625 m, 12 August 2002, R.A. Johnson#RAJ2913, oak-juniper-mesquite woodland, stray foragers, 4 workers (CASENT0869113) [ RAJC]; Santa Cruz County: Sonoita, 31.67833 ° N 39.11667 ° W, 1,490 m, S.L. Heydon, sweeping roadside, 1 worker & 1 dealate gyne (CASENT0731541) [ MMPC]; Patagonia Mtns. 0.4 mi N Jct FSR 135 on FSR 49, 31.43382 ° N 110.72094 ° W, 1,645 m, 31 August 1999, S.P. Cover#SPC5709, juniper and Emory oak woodland, 25’ tall on gentle W-facing slope, sandy loam, found when excavating 5,708, grassy open slope, 1 worker & 1 dealate gyne (MCZENT00583600) [ MCZC] 2 workers (MCZENT00583601) [ MCZC] 2 workers (MCZENT00583602) [ MCZC]; same data as previous, except: S.P. Cover#SPC5704, found when digging 5,705, at edge of grassy gap in sun, 1 worker & 1 dealate gyne (MCZENT00583603) [ MCZC] 2 workers (MCZENT00583604) [ MCZC] 2 workers (MCZENT00583605) [ MCZC] 2w (CASENT0869110) [ RAJC] 3w (CASENT0869111) [ RAJC] 3w (CASENT0869112) [ RAJC]; 1 mi S American Peak, 31.43333 ° N 110.72167 ° W, 1,625 m, 30 September 2005, R.A. Johnson#RAJ3598, oak-pinyon-juniper-grassland, in soil of excavated Trachymyrmex nest, 2 workers (CASENT0869116) [ RAJC]. Arkansas: Garland County: Hot Springs, 18 October 1932, D.E. Read, 1 dealate gyne & 2 workers (CASENT0758293) [ USNM]; Montgomery County: Ouchita Mountains, 2.4 km NW junction Route 27N on Route 270 (in Mt. Ida). 14 May1996, S.P. Cover#SPC4663, Quercus alba , Q. falcata , Ulmus alata , shortleaf pine, 18 m tall on gentle slope, rotten branch on litter surface in shade, 2 workers, 1 dealate gyne (MCZENT00583597) [ MCZC] 2 workers (MCZENT00583598) [ MCZC]; Polk County: Ouachita Mountains Biological Station, 34.45806 ° N 93.99889 ° W, 475 m, 23 June 2017, J.G. Hill, in glade mixed hardwood/pine forest, 1 worker ( MEM 244952) [ MEM]. Florida: Alachua County: Doyle Conner Bldg., Gainesville, 29.63499 ° N 82.37152 ° W, 30 m, 1 May 1974, F.W. Mead, blacklight trap, 1 male (CASENT0867165) [ FSCA]; Gainesville, 29.65163 ° N 82.32484 ° W, 30 m, 22 April 1971, F.W. Mead, blacklight trap, 1 male (CASENT0867167) [ FSCA]; Citrus County: Citrus Wildlife Management Area, 8.5 km W junction Route 41 on Route 44 on Sand Road 3 April 1995, S.P. Cover#SPC4390, open longleaf pine-turkey oak sandhill forest, to 15 m tall, rotten pine stub 5 cm in diameter in open sparsely grassy gap, pure sand, 1 worker & 1 dealate gyne (MCZENT00583599) [ MCZC]; Southern Ridge Sandhill, Archbold Biological Station, 21.18667 ° N 81.35200 ° W, 60 m 24 September 2010, S.P. Cover#SPC8411, pine-oak forest, nest in rotten stick in leaf litter, 1 worker (CASENT0758668) [ UCDC]; Columbia County: Osceola National Forest, 30.20198 ° N 82.45857 ° W, 50 m, 28 October 1976, J.R. Wiley, blacklight trap, 3 males (CASENT0867164) [ FSCA]; same data as previous, except: 12 December 1976, C. Ross, blacklight trap, 1 male (CASENT0867166) [ FSCA]; Liberty County: Torreya State Park, 30.56652 ° N 84.94780 ° W, 55 m, 30 August 1978, L.A. Stange, blacklight trap, 2 males (CASENT0867162) [ FSCA]; Marion County: 11 mi E of Okla. R., Ocala National Forest, 29.16676 ° N 81.79165 ° W, 45 m, 14 October 1959, H.A. Denmark, Pinus clausa , 14 workers & 1 dealate gyne (CASENT0759991- CASENT0759995) [ FSCA]; Juniper Springs, Ocala National Forest, 29.18387 ° N 81.712023 ° W, 5 m, 27 June 1960, H.A. Denmark, Pinus clausa , 1 worker (CASENT0759996) [ FSCA]; same data as previous, except: 28 June 1960, H.A. Denmark, Pinus clausa , 1 worker (CASENT0867011) [ FSCA]; same data as previous, except: 26 November 1960, H.A. Denmark, Pinus clausa , 3 workers (CASENT0867004-CASENT0867006) [ FSCA]; same data as previous, except: 28 November 1960, H.A. Denmark, Pinus clausa , 1 worker (CASENT0867007) [ FSCA]; same data as previous, except: 21 December 1959, H.A. Denmark, Pinus clausa , 4 workers (CASENT0867008, CASENT0867009, CASENT0867016, CASENT0867017) [ FSCA]; same data as previous, except: 21 December 1959, H.A. Denmark, Pinus clausa , 2 workers (CASENT0867008, CASENT0867009) [ FSCA]; same data as previous, except: 20 May 1960, H.A. Denmark, Pinus clausa , 7 workers (CASENT0867010, CASENT0867012- CASENT0867015) [ FSCA]; Monroe County: Cape Sable, 25.27290 ° N 81.12147 ° W, 5 m, 4 April 1958, H.V. Weems, 1 worker (CASENT0867002) [ FSCA]; Little Torch Key, Torchwood Hammock, end of Pirates Rd., 24.65442 ° N 81.38783 ° W, 12 m, 13 March 2010, C.S. Moreau#CSM1380b, 1 worker (CASENT0759999) [ FSCA]; Saddlebunch Keys, 24.60515 ° N 81.59398 ° W, 5 m, 29 December 1959, H.V. Weems, on Flaveria linearis , 1 male (CASENT0759989) [ FSCA]; Okaloosa County: 4.5 mi NW Holt, A & M Res. Sta., Blackwater River State Forest, 30.73583 ° N 86.81628 ° W, 20 m, 1 September 1979, L.A. Stange, blacklight trap, 1 male (CASENT0867163) [ FSCA]; Fam Biol. Sta., 3 mi NW Holt, 30.73137 ° N 86.80135 ° W, 20 m, 8 August 1979, L.A. Stange, blacklight trap, 2 males (CASENT0867161) [ FSCA]; Orange County: Indian River City, Titusville Well Field, 0.2 km S Route 50, 28.53333 ° N 80.81483 ° W, 20 m, 1 March 2001, S.P. Cover#SPC6168, oak-sand pine scrub, hollow stick buried in litter, 1 worker (CASENT0758666) [ UCDC]; Wekiwa Springs State Park, 28.71065 ° N 81.46084 ° W, 20 m, 20 November 1993, Z. Prusak, sandhill, 1 dealate gyne & 2 workers (CASENT0867107) [ FSCA]; Osceola County: 7.4 mi S of Narcoossee, 28.20869 ° N 81.17210 ° W, 25 m, 1 February 1959, R.E. Woodruff, in malt trap, 2 workers (CASENT0867043) [ FSCA]; Putnam County: 29.62652 ° N 81.77870 ° W, 20 m, 9 August 1948, A. Van Pelt, P. palustris - Q. cinerea , 3 workers (CASENT0867018) [ FSCA]; same data as previous, except: 4 September 1948, A. Van Pelt, hydric hammock, 63 workers (CASENT0867019-CASENT0867039) [ FSCA]; same data as previous, except: 10 December 1948, A. Van Pelt, Turkey oak, 6 workers (CASENT0867040, CASENT0867041) [ FSCA]; same data as previous, except: 16 October 1948, A. Van Pelt, Q. spp., Leon, 14 workers & 1 dealate gyne (CASENT0867042, CASENT0867044-CASENT0867047, CASENT0867055, CASENT0867056) [ FSCA]; same data as previous, except: 11 October 1948, A. Van Pelt, P. palustris - Q. cinerea , 19 workers (CASENT0867048-CASENT0867054) [ FSCA]; same data as previous, except: 3 August 1948, A. Van Pelt, Pong pine-Fetterbush, 3 workers (CASENT0867058) [ FSCA]; same data as previous, except: 9 August 1948, A. Van Pelt, P. palustris - Q. cinerea , 32 workers (CASENT0867062-CASENT0867072) [ FSCA]; same data as previous, except: 3 September 1948, A. Van Pelt, Quercus spp. , 18 workers (CASENT0867073, CASENT0867080, CASENT0867082-CASENT0867085) [ FSCA]; same data as previous, except: 21 September 1948, A. Van Pelt, P. palustris - Q. cinerea , 25 workers (CASENT0867074- CASENT0867079, CASENT0867081, CASENT0867096, CASENT0867097) [ FSCA];]; same data as previous, except: 2 October 1948, A. Van Pelt, mesic hammock, 14 workers (CASENT0867087-CASENT0867091) [ FSCA]; same data as previous, except: 11 October 1948, A. Van Pelt, P. palustris - Q. cinerea , 19 workers (CASENT0867048-CASENT0867054) [ FSCA]; same data as previous, except: 28 October 1948, A. Van Pelt, P. palustris , 2 workers (CASENT0867059) [ FSCA]; same data as previous, except: 14 November 1948, A. Van Pelt, mesic hammock, 13 workers (CASENT0867098- CASENT0867102) [ FSCA]; same data as previous, except: 5 December 1948, A. Van Pelt, P. clausa - Q. spp., 5 workers (CASENT0867060-CASENT0867061) [ FSCA]; same data as previous, except: 23 April 1949, A. Van Pelt, P. palustris - Q. cinerea , 2 workers (CASENT0867057) [ FSCA]; same data as previous, except: 6 September 1949, A. Van Pelt, Q. cinerea , 11 workers & 1 dealate gyne (CASENT0867092-CASENT0867095) [ FSCA]; same data as previous, except: 17 January 1950, A. Van Pelt, P. serotina - Desmoth, 10 workers (CASENT0867086, CASENT0867103-CASENT0867105) [ FSCA]; Sarasota County: Myakka River State Park, 27.24028 ° N 82.31518 ° W, 5 m, 1 February 1990, J. Longino#JTL2604, 2 workers (CASENT0867106) [ FSCA]; Seminole County: 21st District, Lot#172, 30.9613 ° N 84.85753 ° W, 35 m, 13 June 1953, P.B. Kannowski#PBK775, pine-oak, on sand, 3 workers (CASENT0867003) [ FSCA]; Walton County: 0.3 km S of the junction of AFB 211 on AFB 214 at Indigo Pond, Elgin Airforce Base, 30.69655 ° N 86.33200 ° W, 70 m, 12 June 2013, S.P. Cover#SPC8557, pine-oak forest, in rotten pine branch, 1 worker (CASENT0758669) [ UCDC]. Georgia: Clarke County, Athens, 05 April 1975, P. Decelles#750405-2, 1 dealate gyne & 2 workers (CASENT0758292) [ USNM]; Rabun County: Tallulah Gorge State Park, 34.73722 ° N 83.39306 ° W, 575 m, 5 June 2016, J.G. Hill, hardwood forest, mountain trail, 1 worker ( MEM 245145) [ MEM]; Whitfield County: Pinhoti trail, Mt. Sinai, 34.739 ° N 85.0167 ° W, 485 m, 30 March 2012, D. Booher, hardwood forest, SW facing slope, ex Carya nut, 1 worker (CASENT0758874) [ UGCA]. Illinois: Pope County: Herod, 37.58033 ° N 88.43616 ° W, 135 m, 6 November 1942, Ross & Sanders#INHS439.048, ground litter, 1 worker (CASENT0759704) [ INHS]; Union County: Giant City, 15 May 1968, R.E. Levy, 3 workers (CASENT0758308) [ UCDC]. Iowa: Boone County: Boone, 42.05971 ° N 93.88023 ° W, 345 m, 11 May 1941, Wm. Buren, 1 worker (CASENT0867168) [ FSCA]; Clayton County: Elkader, 23 June 1941, W.F. Buren, 2 males & 1 worker (CASENT0758291) [ USNM] 1 male & 1 alate gyne (CASENT0758229) [ USNM]; Mill County: Glenwood, 41.04694 ° N 95.742511 ° W, 310 m, June 1940, Wm. Buren, 1 worker (CASENT0867160) [ FSCA]. Kentucky: Madison County: Berea, 37.60083 ° N 84.29972 ° W, 295 m, 22 July 2013, D.M. Daugherty, Lindgren funnel trap baited with IPS lure, 1 worker ( MEM 245110) [ MEM]. Louisiana: Jefferson County: Jean Lafitte National Historical Park and Preserve, 29.76972 ° N 90.13 ° W, 5 m, 16 June 2016, X. Chen, on tallow root, 1 worker ( MEM 243841) [ MEM]. Mississippi: Winston County: Tombigbee National Forest, 33.21556 ° N 89.10417 ° W, 210 m, 6 June 2012, J.A. MacGown, sloped hardwood forest, in Carya glabra nut, 4 workers, 1 alate gyne, 1 male (CASENT0868827, CASENT0868956-CASENT0868960) [ MEM]. Missouri: Barry County: 4 miles WSW of Eagle Rock, 36.51972 ° N 93.82028 ° W, 405 m, 14 June 2007, L.E. Watrous, from glade soil, 1 worker ( MEM 244988) [ MEM]; Ozark Mtns. 14 km S junction Route 76/86 on Route 112 at FSR 1037, 19 May 1996. S.P. Cover#SPC4759, black oak, white oak forest 18–21 m tall, in Quercus borealis acorn on litter surface in light shade, 1 worker & 1 dealate gyne (MCZENT00583596) [ MCZC]. New Mexico: Hidalgo County: 1,234 m, 16 km N Rodeo, 31.9774 ° N 109.0356 ° W, G.W. Forister, 9 September 2010, 1 worker (CASENT0758313) [ UCDC]. North Carolina: Orange County: 9 km NNE Hillsborough, 36.15000 ° N 79.06667 ° W, 175 m, 22 July 1983, J. Longino#JTL22Jul83/stray, piedmont woodland, 3 workers (LACMENT141184-LACMENT141186) [ JTLC]; Stanly County: Morrow Mountain State Park, 9 October 2011, B. Guenard, hand collecting, 1 worker (CASENT0758261) [ MMPC] 1 worker (CASENT0732597) [ UCDC]; Wake County: park at turnoff on Tanager, 35.759970 ° N 78.688698 ° W, 2 June 2008, S.B. Menke#SBM_272, pitfall P8, 2 workers (CASENT0758310) [ UCDC]. Ohio: Adams: Abner Hollow Rd., Green Twp., 38.71561 ° N 83.43808 ° W, 170 m, 21 August 1997, G.A. Coovert, 2 workers (CASENT0759997, CASENT0759998) [ FSCA]. Tennessee: Davidson County: Hendersonville, 36.33587 ° N 86.56801 ° W, 175 m, 19 June 2011, B.E. Boudinot#BEB000439, suburban, lawn and fields, 1 worker (TESC00009756) [ BEBC]; McNairy County: Big Hill Pond State Park, 35.04417 ° N 88.73167 ° W, 150 m, 9 July 2012, J.G. Hill, sloped hardwood forest, 1 worker ( MEM 245132) [ MEM]. Texas: Brewster County: Chisos Mountains, The Basin, 1,540 m, 27 July 1970, P. Ward#3785-6, open vegetation pine-oak-juniper, ex honey bait, 3 workers (CASENT0915983) [ PSWC] 3 workers (CASENT0758355) [ PSWC] 2 workers (CASENT0758307) [ UCDC]; Cameron County: Laguna Atacosa National Wildlife Refuge, 5 m, 15 December 1984, P.S. Ward#7197-2, coastal mesquite scrub, ground forager (s), 2 workers (CASENT0758356) [ PSWC] 1 worker (CASENT0758309) [ PSWC]; Jeff Davis County: 38 km S Kent, 15 August 1996, L.A. Baptiste, sweeping vegetation, 1 worker (CASENT0758311) [ UCDC] 1 worker (CASENT0758312) [ UCDC]. Virginia: Fairfax County: Vienna, 38.89835 ° N 77.26515 ° W, 100 m, 22 June 1983, J. Longino#JTL22Jun83/ Vienna, waste ground in urban area, strays, 4 workers (LACMENT141180- LACMENT141183) [ JTLC].

Geographic range: All elevations, eastern U.S.A. as far north as Michigan, as far west as Arizona, and south as far as Hidalgo, Mexico ( Fig. 127B ).

Worker diagnosis: Temnothorax pergandei can be separated from all other species in the salvini clade by the following character combination: promesonotum evenly convex; metanotal groove deeply impressed and propodeum depressed below the level of the promesonotum; propodeal spines present and typically represented by small teeth; in profile view, postpetiole evenly convex dorsally, with a small flat area posteriorly; in dorsal view, petiolar node 1.1 to 1.3 times as broad as caudal cylinder; setae on head, mesosoma, legs, waist segments and gaster erect, moderately long, abundant and tapering; integument variously colored: may be uniformly dark brown, uniformly yellow, or bicolored.

Similar species: This striking species is difficult to confuse with any other species in Temnothorax , except the closely related T. bison sp. nov. The presence of propodeal teeth, the structure of the postpetiole, which bulges anterodorsally in T. bison sp. nov., and the relative width of the petiolar node in dorsal view (1.3 to 1.6 times as broad as the caudal cylinder in T. bison sp. nov.) will separate the two species.

Worker measurements & indices (n = 20): SL = 0.543 -0.708 (0.631); FRS = 0.213 -0.283 (0.248); CW = 0.643 –0.853 (0.726); CWb = 0.597 –0.784 (0.663); PoOC = 0.262 –0.307 (0.284); CL = 0.701 –0.860 (0.766); EL = 0.165 –0.227 (0.189); EW = 0.106 –0.158 (0.132); MD = 0.141 –0.199 (0.172); WL = 0.847 –1.135 (1.003); SPST = 0.109 –0.222 (0.177); MPST = 0.278 –0.399 (0.325); PEL = 0.346 –0.500 (0.403); NOL = 0.195 –0.272 (0.226); NOH = 0.099 –0.156 (0.128); PEH = 0.187 –0.275 (0.234); PPL = 0.171 –0.255 (0.224); PPH = 0.240 –0.344 (0.294); PW = 0.413 –0.557 (0.473); SBPA = 0.118 –0.228 (0.194); SPTI = 0.134 –0.264 (0.220); PEW = 0.158 –0.225 (0.191); PNW = 0.178 –0.262 (0.228); PPW = 0.257 –0.372 (0.322); HFL = 0.609 –0.798 (0.703); HFWmax = 0.137 –0.186 (0.160); HFWmin = 0.051 –0.068 (0.059); CS = 0.948 –1.214 (1.046); ES = 0.226 –0.306 (0.255); SI = 84–104 (95); OI = 23–26 (24); CI = 82–91 (87); WLI = 142–157 (151); SBI = 20–33 (29); PSI = 13–21 (18); PWI = 156–183 (169); PLI = 155–213 (181); NI = 144–245 (178); PNWI = 111–133 (120); NLI = 51–64 (56); FI = 239–315 (271).

Worker description: In full-face view, head subquadrate, longer than broad (CI 82–91). Mandibles densely striate but shining and armed with five teeth: the apical-most well developed and acute, followed by a less developed preapical tooth and three equally developed smaller teeth. Anterior clypeal margin weakly convex medially. Antennal scapes moderately long: when fully retracted, surpassing the posterior margin of the head capsule by about the maximum width of the scape (SI 84–04). Antennae 12-segmented; antennal club of composed of three segments, with the apical-most segment about as long as the preceding two in combination. Frontal carinae short and apices directed laterally, extending past the antennal toruli by about the maximum width of the antennal scape. Compound eyes moderately protruding past the lateral margins of the head capsule. Lateral margin of head convex, forming a continuous arc from the mandibular insertions to the posterior margin of the head. Posterior head margin flat but rounding evenly into the lateral margins.

In profile view, compound eyes ovular and moderately large (OI 23–26), with 16 ommatidia in longest row. Pronotal declivity indistinct, neck and anterior face of pronotum forming a ~120 ° angle. Mesosoma with promesonotum forming an even convexity; metanotal groove deeply impressed, distinctly separating the promesonotum from the propodeum, which is depressed below the level of the promesonotum and dorsally convex. Promesonotal suture extending from the posterior margin of the procoxal insertion to the mesothoracic spiracle, which is moderately well developed, then continuing to the dorsal surface as a very weak sulcus. Propodeal spiracle well developed, directed posterolaterally, and separated from the propodeal declivity by about three spiracle diameters. Propodeal spines poorly developed and short (PSI 13–21), in the form of small triangular teeth, angled posterodorsally. Propodeal declivity flat, forming a ~120 ° angle with the base of the propodeal spines. Propodeal lobes rounded and weakly developed. Metapleural gland bulla large, extending from the metacoxal insertion three quarters of the way to the propodeal spiracle. Petiole moderately long (PLI 155–213), without tubercles anterodorsally. Subpetiolar process in the form of a small, triangular, blunt tooth, which continues as a low carina to the caudal cylinder; ventral margin of petiole posterior to the subpetiolar process strongly concave. Petiolar peduncle moderately long: comprising about half of the petiole. Petiolar node robust and cuneiform: transition between peduncle and node marked by a rounded angle of ~130 °; anterior face forming a rounded ~90 ° angle with the posterior face; caudal cylinder long: about the as long as the maximum width of the antennal scape. Postpetiole weakly convex anteriorly, rounding evenly into the dorsal face, which bulges before flattening posteriorly; lobed ventrally.

In dorsal view, humeri weakly developed: evenly rounded and wider than the rest of the mesosoma, but not distinct from it; mesothoracic spiracles weakly protruding past the lateral margins of the mesosoma, visible as slight angles where the pronotum meets the mesonotum. Promesonotal suture visible as a weak sulcus. Metanotal strongly impressed, distinctly dividing the promesonotum from the propodeum. Propodeal spines very short, but broadly approximated basally and diverging apically, their apices separated from each other by about five times their length. Petiolar peduncle with spiracles protruding past the lateral margins, but not noticeably constricted anterior to them. Petiolar node, when viewed posterodorsally, trapezoidal and apically broadened, the apex flat; caudal cylinder narrower than the apex of the node. Postpetiole narrow (PWI 156–183) and subquadrate. Anterior margin of the postpetiole weakly convex and meeting the lateral margins at a ~95 ° angle; lateral margins parallel to the angulate posterior corners; posterior margin broadly concave. Metafemur moderately to strongly incrassate (FI 239–315).

Sculpture: median clypeal carina present, extending posteriorly to the level of the anterior margins of the antennal toruli, and flanked on either side by one weaker carina. Lateral clypeal lobes with additional, weaker carinae; ground sculpture smooth and shining on the posterior half of the median clypeal lobe, and weakly areolate anteriorly. Antennal scapes shining through weak coriarious ground sculpture. Cephalic dorsum predominantly weakly coriarious; concentric costulae surrounding the antennal insertions, extending from the apices of the frontal carinae, to the anterior margins of the lateral clypeal lobes. Lateral surfaces of head predominantly weakly coriarious, with strong costulae between the compound eye and the mandibular insertion. Ventral surface of head smooth and shining. Pronotal neck areolate. Lateral surface of the pronotum and the surface surrounding the propodeal spiracle weakly coriarious. Lateral face of the propodeum, meso- and metapleurae with strong costulae. Dorsal surface of promesonotum weakly coriarious; dorsal surface of propodeum areolate. Femora smooth and shining. Petiole shining ventrally through weak coriarious sculpture; dorsum of petiolar peduncle and lateral face of petiolar node with weak costulae; petiolar node otherwise weakly coriarious. Postpetiole weakly coriarious. First gastral tergite and sternite weakly coriarious, without spectral iridescence.

Setae: antennal scapes and funiculi with long, subdecumbent pilosity. Dorsum of the head, pronotum, waist segments, and gaster with abundant, erect, flexuous, tapering setae, the longest of which are slightly longer than the length of the compound eye. The head bears ~36, mesosoma ~32, petiole 18, postpetiole ~28, and first gastral tergite ~66 setae. Pubescence is long, coarse, difficult to distinguish from the setae, and present over the entire body except for the ventral surface of the petiole.

Color: predominantly medium brown, with mandibles and tarsi testaceous yellow; masticatory margins of the mandibles dark brown.

Gyne measurements & indices (n = 9): SL = 0.633 –0.728 (0.680); FRS = 0.270 –0.327 (0.301); CW = 0.848 –0.963 (0.906); CWb = 0.755 –0.868 (0.816); PoOC = 0.289 –0.339 (0.315); CL = 0.821 –0.943 (0.880); EL = 0.244 –0.284 (0.267); EW = 0.173 –0.222 (0.198); MD = 0.142 –0.195 (0.171); WL = 1.405 –1.554 (1.488); SPST = 0.203 –0.274 (0.240); MPST = 0.352 –0.458 (0.407); PEL = 0.453 –0.515 (0.492); NOL = 0.212 –0.319 (0.262); NOH = 0.135 –0.181 (0.162); PEH = 0.293 –0.362 (0.328); PPL = 0.229 –0.291 (0.259); PPH = 0.370 –0.439 (0.406); PW = 0.786 –0.903 (0.835); SBPA = 0.341 –0.405 (0.377); SPTI = 0.307 –0.439 (0.370); PEW = 0.233 –0.277 (0.258); PNW = 0.256 –0.328 (0.295); PPW = 0.384 –0.465 (0.429); HFL = 0.793 –0.896 (0.852); HFWmax = 0.160 –0.185 (0.177); HFWmin = 0.063 –0.085 (0.072); CS = 1.166 –1.340 (1.256); ES = 0.343 –0.395 (0.366); SI = 79–89 (83); OI = 27–32 (29); CI = 90–96 (93); WLI = 178–188 (183); SBI = 41–49 (46); PSI = 14–18 (16); PWI = 161–170 (166); PLI = 175–223 (191); NI = 143–233 (163); PNWI = 107–119 (114); NLI = 46–63 (53); FI = 209–283 (248).

Gyne description: In full-face view, head subquadrate, slightly longer than broad (CI 90–96). Mandibles densely striate but shining and armed with five teeth: the apical-most well developed, followed by a less developed preapical tooth and three equally developed smaller teeth. Anterior clypeal margin flat medially. Antennal scapes moderately long: when fully retracted, just reaching the posterior margin of the head capsule (SI 79–89). Antennae 12-segmented; antennal club composed of three segments, with the apical-most segment as long as the preceding two in combination. Frontal carinae moderately long, directed laterally at their apices, and extending past the antennal toruli by about three times the maximum width of the antennal scape. Compound eyes moderately protruding past the lateral margins of the head capsule. Lateral margin of head evenly convex, converging from below the compound eyes to the mandibular insertions. Posterior head margin flat, rounding evenly into the lateral margins.

In profile view, compound eyes ovular and large (OI 27–32), with 20 ommatidia in longest row. Mesoscutum rounded evenly anteriorly, not quite covering the dorsal surface of the pronotum, and weakly convex dorsally. Mesoscutellum on the same level of the mesoscutum, and evenly convex dorsally. Posterior margin of metanotum extending slightly past the posterior margin of the mesoscutum. Propodeal spiracle well developed, directed posterolaterally, and separated from the propodeal declivity by about two and a half spiracle diameters. Propodeal spines short and stout (PSI 14–18), represented by posterodorsally directed, blunt, triangular teeth, the bases of which are attached to the propodeal declivity by flanges. Propodeal declivity straight and flat, forming a ~120 ° angle with the base of the propodeal spines. Propodeal lobes rounded and weakly developed. Metapleural gland bulla large, extending from the metacoxal insertion three quarters of the way to the propodeal spiracle. Petiole moderately long (175–223), without tubercles anterodorsally. Subpetiolar process in the form of a small, triangular, blunt tooth, which continues as a low carina to the caudal cylinder; ventral margin of petiole posterior to the subpetiolar process strongly concave. Petiolar peduncle moderately long: comprising about half the length of the petiole. Petiolar node robust and cuneiform: transition between peduncle and node marked by a rounded angle of ~130 °; anterior face forming a rounded ~90 ° angle with the posterior face; caudal cylinder long: about the as long as the maximum width of the antennal scape. Postpetiole weakly convex anteriorly, rounding evenly into the dorsal face, which bulges before flattening posteriorly; lobed ventrally.

In dorsal view, mesoscutum not fully covering pronotum anteriorly; humeri evenly rounded laterally. Propodeal teeth diverging apically, their apices separated from each other by about six times their length. Petiolar peduncle with spiracles protruding past the lateral margins, but not noticeably constricted anterior to them. Petiolar node, when viewed posterodorsally, trapezoidal and apically broadened, the apex weakly emarginate; caudal cylinder narrower than the apex of the node. Postpetiole narrow (PWI 161–170), and subquadrate. Anterior margin of postpetiole weakly convex, with corners marked by rounded angles as it transitions to the lateral margins, which are parallel to the angulate posterior corners; posterior margin broadly concave. Metafemur weakly to moderately incrassate (FI 209–283).

Sculpture: median clypeal carina present, extending from the anterior margin nearly to the level of the anterior margins of the antennal insertions; flanked by weaker, indistinct carinae near the lateral margins of median clypeal lobe. Lateral clypeal lobes with additional weaker carinae; ground sculpture smooth and shining on the median clypeal lobe medially but becoming weakly areolate-coriarious laterally. Antennal scapes weakly coriarious. Cephalic dorsum predominantly costulate over smooth and shining sculpture, with moderately strong striae flanking the frontal carinae; a median strip of smooth sculpture extends from the frontal triangle to the median ocellus, then expands to encompass the middle of the ocellar triangle and extends to the posterior head margin; concentric costulae surrounding the antennal insertions, extending from the frontal carinae to the anterior margin of the lateral clypeal lobes. Lateral surfaces of head with weak coriarious sculpture posterior to the compound eye, denser rugose sculpture surrounding the compound eye, extending between the compound eye and the mandibular insertion. Ventral surface of head weakly costulate. Pronotal neck areolate. Lateral face of the pronotum weakly coriarious, with costulae on the ventral third. Anepisternum smooth and shining on its anterior half, but weakly coriarious on its posterior half; katepisternum predominantly weakly coriarious. Metapleuron and lateral face of propodeum with costulae. Propodeal declivity transversely coriarious. Mesoscutum with predominantly longitudinal coriarious sculpture, with a patch of transverse, finely costulate sculpture anteromedially, and a shallow sulcus posteromedially. Mesoscutellum smooth and shining medially, surrounded by weak coriarious sculpture. Femora smooth and shining. Petiole shining ventrally through weak coriarious sculpture; dorsum of petiolar peduncle and lateral face of petiolar node with weak costulae; petiolar node otherwise weakly coriarious. Postpetiole coriarious, but with sculpture weaker anteromedially. First gastral tergite and sternite weakly coriarious, without spectral iridescence.

Setae: antennal scapes and funiculi with long, subdecumbent pilosity. Dorsum of the head, pronotum, waist segments, and gaster with abundant, erect, flexuous, tapering setae, the longest of which are about the width of the compound eye. Pubescence is long, coarse, difficult to distinguish from the setae, and present over the entire body except for the ventral surface of the petiole.

Color: predominantly medium brown, with mandibles, genae, antennal scapes, antennal funiculus, pronotum, propodeum, tibiae, tarsi and venter of petiole testaceous yellow; masticatory margins of the mandibles dark brown.

Male measurements & indices (n = 2): SL = 0.261 –0.324 (0.293); FRS = 0.133 –0.151 (0.142); CW = 0.552 –0.592 (0.572); CWb = 0.466 –0.475 (0.471); PoOC = 0.251 –0.266 (0.259); CL = 0.561 –0.603 (0.582); EL = 0.240 –0.267 (0.254); EW = 0.196 –0.209 (0.203); MD = 0.038 –0.041 (0.040); WL = 1.062 –1.169 (1.116); SPST = 0.188 –0.216 (0.202); MPST = 0.310 –0.378 (0.344); PEL = 0.334 –0.335 (0.335); NOL = 0.194 –0.258 (0.226); NOH = 0.076 –0.078 (0.077); PEH = 0.160 –0.166 (0.163); PPL = 0.202 –0.215 (0.209); PPH = 0.216 –0.260 (0.238); PW = 0.600 –0.616 (0.608); SBPA = 0.224 –0.257 (0.241); SPTI = 0.224 –0.257 (0.241); PEW = 0.163 –0.182 (0.173); PNW = 0.142 –0.197 (0.170); PPW = 0.255 –0.268 (0.262); HFL = 0.819 –0.891 (0.855); HFWmax = 0.093 –0.098 (0.096); HFWmin = 0.051; CS = 0.747 –0.777 (0.762); ES = 0.338 –0.372 (0.355); SI = 56–68 (62); OI = 45–48 (47); CI = 79–83 (81); WLI = 228–246 (237); SBI = 48–54 (51); PSI = 18; PWI = 140–164 (152); PLI = 156–165 (161); NI = 249–339 (294); PNWI = 87–108 (98); NLI = 58–77 (68); FI = 182–192 (187).

Male description: In full-face view, head elongate and subovate, longer than broad (CI 79–83). Mandibles very weakly striate, shining, and armed with five teeth: the apical-most well developed, followed by a smaller preapical tooth and three roughly equally developed smaller teeth. Anterior clypeal margin entire and weakly convex. Antennal scapes short: when fully retracted, failing to reach the posterior margin of the head capsule by about two times the maximum width of the scape (SI 56–68). Antennae 13-segmented; antennal club composed of four segments, with the apical-most segment slightly longer than the preceding two in combination. Frontal carinae short, extending past the antennal toruli by two times the maximum width of the antennal scape. Compound eyes strongly protruding past the lateral margins of the head capsule. Margin between the anterior margin of the compound eye and the mandibular insertions straight. Posterior head margin flat convex, evenly rounding into the lateral margins.

In profile view, compound eyes ovular and large (OI 45–48), with 30 ommatidia in the longest row. Mesoscutum bulging anteriorly, covering the dorsal surface of the pronotum, and evenly convex dorsally. Mesoscutellum on the same level of the mesoscutum and evenly rounded dorsally. Posterior margin of metanotum extending beyond the posterior margin of scutellum. Propodeal spiracle well developed, directed laterally, and separated from the propodeal declivity by about four spiracle diameters. Propodeal spines absent but indicated by angulate flanges on the dorsal and declivitous faces of the propodeum. Propodeal lobes rounded and weakly developed. Metapleural gland bulla large, extending three quarters of the way between the insertion of the metacoxa and the propodeal spiracle. Petiole moderately long (PLI 156–165), without anterodorsally. Subpetiolar process absent; ventral margin of petiole concave. Petiolar peduncle short: comprising about a third of the total length of the petiole. Petiolar node low and cuneiform, the convergence of the anterior and posterior faces marked by a rounded angle. Postpetiole evenly rounded anterodorsally, flattened dorsally, and with a lobed, concave ventral surface.

In dorsal view, mesoscutum covering pronotum anteriorly, humeri not visible. Petiolar peduncle with spiracles strongly protruding past the lateral margins, the peduncle broadened where they arise. Petiolar node slightly wider than the peduncle; petiole narrowing posterior to the node, before widening again to the caudal cylinder, which is about the same width as the node. Postpetiole narrow (PWI 140–164) and campaniform. Anterior margin of postpetiole convex, with the anterior corners evenly rounding into the lateral margins, which are parallel to the angulate posterior corners; posterior margin of postpetiole broadly concave. Metafemur not incrassate (FI 182–192).

Sculpture: median clypeal carina absent. Lateral clypeal lobes with weak, indistinct carinae, ground sculpture weakly coriarious. Antennal scapes smooth and shining. Head predominantly weakly areolate-coriarious, with a central strip of smooth and shining sculpture extending from the frontal triangle to the median ocellus. Lateral surface of head weakly coriarious. Ventral surface of head smooth and shining anteriorly, transversely rugulose posteriorly. Pronotal neck areolate. Lateral surfaces of pronotum, anepisternum, katepisternum, predominantly smooth and shining, but with anterior third of pronotum densely rugose. Metapleuron and lateral face of propodeum longitudinally rugose. Dorsally, mesoscutum weakly coriarious, the Mayrian furrows moderately well impressed. Mesoscutellum smooth and shining medially, becoming coriarious laterally. Femora smooth and shining. Petiole costulate ventrally; lateral faces of the peduncle and node longitudinally rugose, the posterodorsal surface of the petiole smooth and shining. Dorsal surface of postpetiole shining. First gastral tergite and sternite weakly coriarious.

Setae: antennal scapes and funiculi with long, subdecumbent pilosity. Dorsum of the head, pronotum, waist segments, and gaster with abundant, erect, flexuous, tapering setae, the longest of which are about half the width of the compound eye. Pubescence is long, coarse, difficult to distinguish from the setae, and present over the entire body except for the ventral surface of the petiole.

Color: predominantly medium brown, with mandibles, genae, antennal scapes, antennal funiculus, and tarsi testaceous yellow; masticatory margins of the mandibles dark brown.

Etymology: Patronym, in honor of Theodore Pergande, a myrmecologist who sent his extensive collections to Forel and Emery.

Comments: Temnothorax pergandei is often found in open, sparsely vegetated habitats throughout its range. With its rapid movements and constricted mesosoma, T. pergandei presents itself in the field as distinctly Pheidole -like, often foraging for small arthropods such as aphids and heteropterans during the diurnal hours ( Wheeler, 1903b). Wheeler observed that workers of T. pergandei are territorial, showing aggression toward workers from other colonies. It often nests directly in the ground, with a very indistinct nest entrance that is typically only discoverable by following foraging workers back to it, much like other ground nesting Temnothorax . This is by no means the rule, though: nests have been collected under the bark of pine stumps ( Smith, 1929), in rotten sticks on the ground (this study), or under moss mats and grass roots ( Wheeler, 1903b). Emery (1895) states that Pergande collected T. pergandei in association with a Monomorium minimum Buckley nest, but this observation appears to be spurious; most collections of T. pergandei have been found apart from other ant species. Nests are small and monogynous, with 75 to 250 workers ( Smith, 1929, Wesson, 1935; Heinze, Hoelldobler & Trenkle, 1995). Records of mating flights have been made from late spring in the vicinity of Austin, Texas, U.S.A. ( Wheeler, 1903b), and early summer near Portal, Arizona, U.S.A. ( Heinze, Hoelldobler & Trenkle, 1995). Gynes attract males with secretions from the poison gland, and typically mate with males only once in swarming flights ( Heinze, Hoelldobler & Trenkle, 1995). Temnothorax pergandei is a highly morphologically variable species with a broad range, with a great deal of diversity in integument coloration, sculpturation, and propodeal spine length ( Fig. 130 & Fig. 131 ). These qualities have led to a profusion of names, with the morphologically similar but geographically distant T. floridanu s appearing alongside the original description of T. pergandei ( Emery, 1895) . However, little sense has been made of the different forms, as there is often a great deal of intranidal and geographic variation ( Creighton, 1950). Mackay (1993) synonymized T. floridanus under T. pergandei following a morphometric analysis of specimens throughout the range of both. I have found that the yellow form, formerly the subspecies T. pergandei flavus , is paraphyletic with respect to the typical dark form, as is the case with bicolored specimens. Most of the genetic structure that I have observed in the phylogeny of T. pergandei is explained by geography, with distinct Eastern, Central Great Plains, Northwestern and Southwestern clades, with the Eastern clade corresponding roughly to the Ozarks, the Appalachians and everything east of them. Whether these clades indicate distinct species is a question left to further analysis, but I cannot yet discern a distinct morphological pattern that supports separation.

Figures

Figure 127 Geographical and elevational distribution of the pergandei group. (A) Temnothorax bison sp. nov. (B) T. pergandei. The colored scale to the left of each map represents elevation in meters. Points in black represent the species named in each subfigure, while points in red represent other members of the species group. Type localities are represented by triangles; if present, types of synonyms are named; non-type localities are represented by circles. Bounding box in inset map shows location of main map. Full-size DOI: 10.7717/peerj.11514/fig-127

Figure 129 Temnothorax pergandei. (A–C) Worker (CASENT0758291). (A) Full-face view. (B) Profile view. (C) Dorsal view. (D–G) Gyne (CASENT0758229). (D) Full-face view. (E) Profile view. (F) Dorsal view. (G) Wings. (H–K) Male (CASENT0758229). (H) Full-face view. (I) Profile view. (J) Dorsal view. (K) Wings. Scale bars 0.2 mm. Full-size DOI: 10.7717/peerj.11514/fig-129

Figure 130 Comparison of Temnothorax pergandei worker morphology. (A–C) Missouri, U.S.A. MCZENT00583596). (A) Full-face view. (B) Profile view. (C) Dorsal view. (D–F) Mississippi, U.S.A. USNMENT00921850). (D) Full-face view. (E) Profile view. (F) Dorsal view. (G–I) Arizona, U.S.A. CASENT0172989). (G) Full-face view. (H) Profile view. (I) Dorsal view. Scale bars 0.2 mm. Full-size DOI: 10.7717/peerj.11514/fig-130

Figure 131 Comparison of Temnothorax pergandei gyne morphology. (A–C) Missouri, U.S.A. (MCZENT00583596). (A) Full-face view. (B) Profile view. (C) Dorsal view. (D–F) Mississippi, U.S.A. (USNMENT00529209). (D) Full-face view. (E) Profile view. (F) Dorsal view. (G–I) Arizona, U.S.A. (CASENT0172990). (G) Full-face view. (H) Profile view. (I) Dorsal view. Scale bars 0.2 mm. Full-size DOI: 10.7717/peerj.11514/fig-131