Temnothorax subditivus ( Wheeler, 1903a )

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 : 428-440

Temnothorax subditivus ( Wheeler, 1903a)

Distribution: Fig. 165B ; worker, gyne & male: Fig. 167 ; variability: Fig. 168 .

Macromischa subditiva Wheeler, 1903a: 99 , fig. 5. Syntype workers. Texas, U.S.A.

One syntype worker here designated lectotype. Wheeler, 1931: 15 (Gyne and male of Macromischa flavitarsis Mann ), Smith, 1939: 503. (Male).

Macromischa laevissima Wheeler, 1911: 205 . Holotype worker. Veracruz, Mexico. Junior synonym of Temnothorax subditivus by Baroni Urbani, 1978: 512.

Macromischa (Macromischa) subditiva (Wheeler) : Mann, 1920: 408. First combination in Macromischa (Macromischa) .

Macromischa (Macromischa) flavitarsis Mann, 1920: 420 . Syntype workers and gynes. Guatemala. Junior synonym of Temnothorax subditivus by Baroni Urbani, 1978: 512. One syntype worker here designated lectotype.

Macromischa luciliae Mann, 1935: 35 , fig. 1. Syntype workers. Guatemala. Junior synonym of Temnothorax subditivus by Baroni Urbani, 1978: 512. One syntype worker here designated lectotype.

Leptothorax subditivus (Wheeler) : Baroni Urbani, 1978: 512. First combination in Leptothorax .

Temnothorax subditivus (Wheeler) : Bolton, 2003: 272. First combination in Temnothorax . Temnothorax mmp14 Prebus, 2017: 8. Phylogeny.

Type material examined: Lectotype worker of Macromischa subditiva : U.S.A.: Texas: Travis County: Austin, 15 May 1903, on bark of Salix nigra , Type No. A.M.N.H., M.C.Z. Type 2-1 16371 (MCZENT00016371, middle specimen on pin) [ MCZC].

Paralectotype workers of Macromischa subditiva : same pin as lectotype, 2 workers (top and bottom specimens on pin) [ MCZC]; same data as lectotype, 3 workers (MCZENT00578574) [ MCZC]; [no locality data], [no collection date], W.M. Wheeler, Type No. A.M.N.H., Type 16371, 1 worker (MCZENT00578573) [ MCZC] .

Lectotype worker of Macromischa flavitarsis : GUATEMALA: Sacatepéquez: Antigua , 24 December 1911, W.M. Wheeler, M.C.Z. Cotype 16368, (MCZENT00016368, top specimen on pin) [ MCZC].

Paralectotype workers and gynes of Macromischa flavitarsis : same pin as lectotype, (bottom specimen on pin) [ MCZC]; same data as lectotype, 3 alate gynes (MCZENT00534455) [ MCZC]; Guatemala City, 17 December 1911, W.M. Wheeler, 5 workers, (USNMENT00921848) [ USNM] 3 workers (USNMENT00921847) [ USNM] 5 workers (USNMENT00529208) [ USNM]; Antigua, 21 December 1911, M.C.Z. Cotype 16368, 3 workers (MCZENT00534456) [ MCZC]; Antigua, 20 December 1911, W.M. Wheeler, 3 workers (LACMENT323232) [ LACM] .

Lectotype worker of Macromischa luciliae : [no department], intercepted at Hololulu, Hawaii, U.S.A. [quarantine], 20 October 1932, on Oncidium splendidum roots from Guatemala cargo, Cotype 50802 U.S. N.M. (images of USNMENT00529613 examined on the Smithsonian Type Specimen Database).

Type material not examined: Macromischa laevissima holotype worker: MEXICO: Veracruz: La Buena Ventura, near Santa Rosa, A. Petrunkewitch [ AMNH] .

Non-type material examined: CAYMAN ISLANDS: Grand Cayman : 1 km up Mastic trail, 19.32000 ° N 81.19800 ° W, 5 m, 8 March 2008, #32, J.K. Wetterer, 1 worker (CASENT0868784) [ UCDC] 1 worker (CASENT0868474) [ MMPC] .

COLOMBIA: La Guajira: Caribbean Region , 2 km S Rio Hacha, 18 January 1937, P. Hummelinck #294, 1 worker (MCZENT00583614) [ MCZC] ; Magdalena: Santa Marta 40 m, 11.250000 ° N 74.216667 ° W, 15 August 1985, P.S. Ward #7940, tropical thorn woodland, dead stalk of Manihot sp. , 1 alate gyne, 1 dealate gyne & 1 worker (CASENT0758338) [ PSWC] 1 male & 2 workers (CASENT0756100) [ PSWC]; Villa Culebra, near Bonda, 10 km E Santa Marta, October 1985, H.G. Müller, low vegetation, 2 workers (CASENT0758339) [ PSWC]; Bahia de Neguangue, Tayrona Pk., ca 25 km NE Santa Marta, 8 October 1985, H.G. Müller, low vegetation, 2 workers (CASENT0758340) [ PSWC]; same data as previous, 30 September 1985, H.G. Müller, 3 workers (CASENT0758344) [ PSWC]; Bahia Concha, Tayrona Pk., 10 km NE Santa Marta, 23 June 1985, H.G. Müller, ex vegetation, 3 workers (CASENT0758341) [ PSWC]; mouth of Rio Fundación, Cienaga Grande de Sta Marta , 10.716667 ° N 74.416667 ° W, 29 August 1985, H.G. Müller, 3 workers (CASENT0758342) [ PSWC]; Bahia de Cinto, Parque Nacional Tayrona, 30 km E Santa Marta 11.333333 ° N 74.066667 ° W, 14 April 1986, H.G. Müller, 1 worker (CASENT0758343) [ PSWC]; Punta de Betin , Santa Marta , October 1985, 11.250000 ° N 74.216667 ° W, H.G. Müller, low vegetation, 2 workers (CASENT0758345) [ PSWC] 1 alate gyne (CASENT0756101) [ PSWC]; Punta de Betin , Santa Marta , November-December 1985, 11.250000 ° N 74.216667 ° W, H.G. Müller, 2 workers (CASENT0758346) [ PSWC]; same data as previous, pan traps, 2 workers (CASENT0758347) [ PSWC]; Punta de Betin , Santa Marta , January-March 1986, 11.250000 ° N 74.216667 ° W, H.G. Müller, 2 workers (CASENT0758348) [ PSWC]; Bahia de Neguange, Parque Nacional Tayrona, 25 km NE Santa Marta 11.350000 ° N 74.083333 ° W, 16 June 1985, H.G. Müller, 1 worker (CASENT0758349) [ PSWC]; 2 km E Orihueca, 10.850000 ° N 74.150000 ° W, 20 m, 17 Aug 1985, J. Longino #845, 1 dealate gyne, 1 worker (LACMENT141164) [ UCDC] 1 worker (LACMENT141165) [ JTLC] 2 workers (LACMENT323176) [ LACM] 3 workers (CASENT0869131–3) [ MMPC] .

COSTA RICA: Guancaste: Isla San José, 10.866667 ° N 85.900000 ° W, 100 m, 22 October 1992, D.M. Olson, 3 workers (CASENT0758337) [ UCDC]; S. Naranjo ridge, Santa Rosa National Park , 10.8 ° N 85.68333 ° W, 100 m, 14 July 1985, J. Longino #JTL0461,xeric scrub on rocky ridge, nest in dead branch <0.5 m high, 1 dealate gyne, 1 male, 2 workers (INBIOCRI002281233–4) [ JTLC] 1 male & 3 workers (CASENT0869127–30) [ MMPC]; Paloverde National Park , August 1984, I.H. Rathet, ex Acacia thorn, 3 workers (LACMENT323177) [ LACM]; Puntarenas: Guacimal, road to Monteverde, 10.21667 ° N 84.85 ° W, 400 m, 20 July 1984, J. Longino #JTL20Jul84/2, roadside vegetation, strays from tree trunks, 1 worker (INBIOCRI002281232) [ JTLC]; La Pita, 10.16 ° N 84.90 ° W, 200 m, 13 July 1984, J. Longino #JTL13Jul84/pm, sweep sample, 1 worker (INBIOCRI002281229) [ LACM]; La Pita, road to Monteverde, 10.16667 ° N 84.91667 ° W, 120 m, 13 July 1984, J. Longino #JTL13 Jul 84/am, sweeping vegetation, 2 workers (INBIOCRI002281230, INBIOCRI002281231) [ JTLC] .

GUATEMALA: El Progreso : San Agustin Acasaguastlan, near road to San Cristobal Acasaguastlan, 14.91777 ° N 89.94683 ° W, 280 m, 18 June 2015, Z. Falin #GUAT1F15149, fogging Pereskia trunk/foliage, 1 worker (JTL-SV00859.3) [ JTLC]. Esquintla: Origin: San Jose , in quarantine, San Francisco , California, U.S.A., 18 April 1949, ex Odontoglossum grande , 1 worker (LACMENT323230) [ LACM]. Guatemala: Origin: Guatemala City , quarantine, San Francisco , California, U.S.A., 25 April 1947, ex Oncidium splendidum , 1 male (LACMENT323223) [ LACM] 1 dealate gyne (LACMENT323226) [ LACM] 1 worker (LACMENT323227) [ LACM]; [no locality data], in quarantine, San Francisco , California, U.S.A., 25 March 1947, ex Oncidium splendidum , 3 workers (MCZENT00581856) [ MCZC]. Totonicapan: Xeabaj I, road between antennae, 14.83126 ° N 91.40299 ° W, 3.180 m, 7 June 2015, Z. Falin #GUAT1F15049, under bark, Pinus stump, 1 worker (CASENT0636941) [ JTLC]. Zacapa: Zacapa, finca nr. Univ. Landivar campus, 14.99336 ° N 89.54607 ° W, 200 m, 21 June 2015, Z. Falin #GUAT1F15188, flight intercept trap, 1 worker (JTL-SV00862.3) [ JTLC] .

MEXICO: Campeche: 10 km E Campeche , 28–29 August 1953, E.O. Wilson #141, 2 workers (MCZENT00581851) [ MCZC] ; Chiapas: 8 km SE Salto de Agua , 17.51615 ° N 92.30164 ° W ± 300 m, 100 m, 100 m, 16 June 2008, LLAMA#Go-A-08-2-04, 2 ° wet forest, beating vegetation, 1 worker (CASENT0609888) [ UCDC]; 6 km SE San Cristobal on Las Casas, 2,195 m, 10 August 1978, John E. Rawlins, 1 worker (MCZENT00581852) [ MCZC] ; Jalisco: Estación De Biolgía Chamela , 19.497778 ° N 105.044167 ° W, 135 m, 31 March–4 April 2012, D. Dubovikoff #CHUNAM1, 1 worker (CASENT0758297) [ UNAM] 1 worker (CASENT0758298) [ UNAM] 1 worker (CASENT0758299) [ UNAM] ; Michoacán: 8 km SW Tiquicheo , 425 m, 8 July 1970, E. Fisher & P. Sullivan, 1 dealate gyne (LACMENT323468) [ LACM] ; Nayarit: Maria Magdalena, Isla Tres Marias , 23 March 1964, R.R. Snelling, 1 worker (LACMENT323229) [ LACM] . Nuevo León: Chipinque Mesa, Monterrey , 1,280 m, 21 January 1952, W.S. Creighton, in Quercus fusiformis , 3 workers (LACMENT323234) [ LACM] 3 workers (CASENT0759986) [ FSCA] ; Tamaulipas: Antigue Morelos, 9 July 1969, S.&J. Peck #Ber167, Berlese of palm-thorn forest litter, 1 dealate gyne (MCZENT00583613) [ MCZC] ; Veracruz: Mirador, 16 March 1929, E. Skwarra #Z154a/ Sk. , Conostegia xalapensis , 3 workers (MCZENT00581853) [ MCZC]; same data as previous, except: 18 April 1929, E. Skwarra #Z196c/Sk., 1 dealate gyne (MCZENT00581854) [ MCZC]; same data as previous, except: 3 May 1929, E. Skwarra #Z333/Sk., Conostegia xalapensis , 3 males (MCZENT00581855) [ MCZC]; same data as previous, except: 4 May 29, E. Skwarra, 3 workers (LACMENT323231) [ LACM]; same data as previous, except: E. Skwarra #Z348/SK, 3 males & 2 alate queens (MCZENT00730929-30) [ MCZC] .

NICARAGUA: Estelí: 16 km N Estelí, 13.24082 ° N 86.35307 ° W ± 100 m, 870 m, 25 April 2011, J. Longino #JTL7420-s, tropical dry scrub, strays, 1 worker (CASENT0619627) [ JTLC]; same data as previous, except: J. Longino #JTL7421-s, tropical dry scrub, at bait, 3 workers (CASENT0619366-CASENT0619368) [ JTLC] .

PANAMA: Panamá, Juan Díaz, 1938, N.A. Weber #775, 2 workers (LACMENT323178) [ LACM] .

U.S.A.: Louisiana: Beauregard Parish: DeRidder, 24 June 1942, W.F. Buren, 3 workers (CASENT0867155) [ FSCA]; same data as previous, except: 10 July 1942, W.F. Buren, 1 alate gyne, 3 males, 8w (CASENT0867156-CASENT0867159) [ FSCA]; same data as previous, except: 3 September 1942, W.F. Buren, 3 workers (LACMENT323224) [ LACM]. Texas: Bastrop County: Bastrop State Park, 21 September 1979, P. Ward#3825-2, honey bait on ground, pine-oak forest, 1 worker (CASENT0758323) [ UCDC]; Bexar County: San Antonio, 28 April 1979, P.S. Ward#3298, foraging on low vegetation, 1 worker (CASENT0758324) [ UCDC]; Cameron County: Brownsville, 24 October 1942, [no collector data], 4 workers (LACMENT323225) [ LACM]; La Feria, Pandora stem, 16 April 1973, R.R. Snelling#RRS73-12, 3 workers (LACMENT323228) [ LACM]; La Feria, 26 January 1973, R.M. Duffield, 3 workers (CASENT0750538-CASENT0750540) [ FSCA]; Resaca de la Palma S.P., 25.99596 ° N 97.56622 ° W ± 50 m, 10 m, 5 November 2013, R. & C. Anderson# RSA 2013TX05, Celtis , ex sifted leaf litter, 1 worker (JTL713352) [ JTLC]; Sabal Palm Grove Sanctuary, 25.84994 ° N 97.41850 ° W ± 50 m, 10 m, 4 November 2013, R. & C. Anderson# RSA 2013TX02, hardwood forest, ex sifted leaf litter, 1 worker (CASENT0635918) [ JTLC]; In quarantine from Harlingen, San Bernadino, San Bernadino County, California, 12 November 1961, Young et al. California Department of Agriculture#61215-15, ex Washingtonia robusta , 3 workers (LACMENT323233) [ LACM]; Hardin County: 4.75 km E of US 69/287, 0.30 km N of FM 420, 15 September 1979, S.P. Lewis#SPL89, 1 worker (MCZENT00730931) [ MCZC]; 2.06 km W of Silsbee, 0.4 km N of Texas 327, 15 September 1979, S.P. Lewis#SPL97, 1 worker (MCZENT00730932) [ MCZC]; same data as previous, except: S.P. Lewis#SPL99, 1 worker (MCZENT00730933) [ MCZC]; Hidalgo County: Bentsen Rio Grande State Park 12–13 March 1980, semi-dry deciduous forest, P.S. Ward#3928, by water hole, on rotten log, 1 worker (CASENT0758328) [ UCDC]; Bentsen Rio Grande State Park, 26.18308 ° N 98.38243 ° W ± 50 m, 35 m, 5 November 2013, R. & C. Anderson# RSA 2013TX06, dry cedar elm, ex sifted leaf litter, 1 worker (JTL713359) [ JTLC]; Mission, 30 m, 15 December 1984, P.S. Ward#7185, roadside edge, ex dead twig of undetermined tree, 1 dealate gyne, 1 worker & 1 male (CASENT0915978) [ PSWC] 2 workers (CASENT0758327) [ UCDC] 1 worker (CASENT0758702) [ UCDC]; Progreso, 11 January 1966, W.S. Creighton, 1 dealate gyne, 3 workers (CASENT0759985) [ FSCA]; Santa Ana National Wildlife Refuge, 30 m, 14 December 1984, P.S. Ward#7167-2, subtropical thorn woodland, on low vegetation, 1 worker (CASENT0758326) [ UCDC]; same data as previous, except: 13 December 1984, J. Longino#JTL0047, mesic woodland, nest in dead stick, 1 worker (LACMENT141166) [ JTLC] 3 workers (CASENT0869120 -2) [ MMPC]; same data as previous, except: 14 December 1984, J. Longino#JTL0076, dead stalks of Arundo donax along riverbank, single-queen colony, 1 worker (JTL0002485) [ JTLC] 1 dealate gyne & 3 workers (CASENT0869123-6) [ MMPC]. Kinney County: Sycamore Creek at Hwy90, 29.38333 ° N 100.7 ° W, 300 m, 16 December 2004, J. Longino#JTL5400-s, live oak woodland, sifted leaf litter, 1 worker ( JTLC 000006433) [ JTLC]. Travis County: Austin, 15 April 1979, P.S. Ward#PSW3234, dead twig of Morus , 5 workers (CASENT0758322, CASENT0758336) [ UCDC]; Brackenridge Field Station, Austin 19 June 1978, D.H. Feener#Z715b, 1 male & 1 worker (MCZENT00581859) [ MCZC] same data as previous, Z714b, Z713 b, 1 alate gyne & 1 worker (MCZENT00581858) [ MCZC]; same data as previous, 21 August 1981, P.S. Ward#5133, on Ulmus trunk, semi-deciduous woodland, 2 workers (CASENT0758325) [ UCDC]; Walker County, Sam Houston National Forest, 18 km SW junction Route 30 on Farm Road 1791, 2 May 1993, S.P. Cover#3495, mature shortleaf-loblolly pine forest, 15–18 m tall with well-developed mixed deciduous understory to 12 m, in standing pine bark (stump remnant) in shade, 2 workers (MCZENT00581857) [ MCZC].

VENEZUELA: Lara: Quibor, Jimenez , 8 July 1979, R.W. Brooks, A.A. Grigarick, J. McLaughlin, R.O. Schuster, 1 male (CASENT0758329) [ UCDC] . Zulia: Maracaibo vic., 1941, R.G. Wesson, dry thorn scrub, 6 workers (MCZENT00730934–6) [ MCZC] .

Geographic range: all elevations, southern U.S.A. to Colombia & Venezuela, including the Cayman Islands ( Fig. 165B ).

Worker diagnosis: Temnothorax subditivus can be separated from all other species in the salvini clade by the following character combination: posterior margin of head convex; antennal scape moderately long, surpassing the posterior margin of the head by about the maximum width of the antennal scape; in profile view, dorsum of mesosoma strongly convex; pronotal declivity not marked by an angle as it transitions from the anterior face of the pronotum to the dorsal face; propodeum not depressed below the level of the promesonotum; propodeal spines as long as or longer than the propodeal declivity, directed posterodorsally, and straight; hind femora moderately to strongly incrassate; petiolar node node robust and erect, squamiform, and not leaning posteriorly over the caudal cylinder of the petiole; postpetiole moderately broad: greater than 1.7 times the width of the caudal cylinder of the petiole; setae on head, mesosoma, waist segments and gaster erect, moderately long, sparse and blunt (never long and tapering).

Similar species: Temnothorax subditivus is extremely variable across its vast range, and may be confused with T. goniops , T. pilicornis sp. nov., T. tenuisculptus , and members of the augusti and pastinifer groups. It can be separated from all of the above, except the pastinifer group, by the extremely squamiform petiolar node, which in dorsal view is much broader than the caudal cylinder of the petiole. When viewed dorsally, the closely approximated propodeal spines, the bases of which are united by a transverse carina, will distinguish T. subditivus from all members of the pastinifer group.

Worker measurements & indices (n = 15): SL = 0.528 –0.704 (0.610); FRS = 0.184 –0.237 (0.209); CW = 0.569 –0.712 (0.645); CWb = 0.525 –0.657 (0.592); PoOC = 0.257 –0.322 (0.285); CL = 0.644 –0.810 (0.717); EL = 0.153 –0.198 (0.172); EW = 0.117 –0.146 (0.130); MD = 0.138 –0.189 (0.164); WL = 0.745 –1.003 (0.836); SPST = 0.247 –0.368 (0.296); MPST = 0.255 –0.318 (0.283); PEL = 0.301 –0.417 (0.346); NOL = 0.168 –0.240 (0.207); NOH = 0.100 –0.174 (0.142); PEH = 0.201 –0.277 (0.243); PPL = 0.120 –0.166 (0.144); PPH = 0.148 –0.206 (0.183); PW = 0.361 –0.470 (0.419); SBPA = 0.117 –0.167 (0.144); SPTI = 0.179 –0.300 (0.247); PEW = 0.124 –0.163 (0.143); PNW = 0.212 –0.298 (0.251); PPW = 0.246–0.32 (0.285); HFL = 0.574 –0.837 (0.670); HFWmax = 0.138 –0.208 (0.168); HFWmin = 0.049 –0.066 (0.055); CS = 0.848 –1.062 (0.951); ES = 0.213 –0.269 (0.237); SI = 95–109 (103); OI = 24–26 (25); CI = 79–86 (83); WLI = 134–153 (141); SBI = 21–28 (24); PSI = 30–45 (35); PWI = 179–216 (199); PLI = 218–273 (241); NI = 117–210 (148); PNWI = 160–208 (175); NLI = 53–70 (60); FI = 261–359 (304).

Worker description: In full-face view, head subquadrate, slightly longer than broad (CI 79–86). Mandibles densely, finely 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 evenly convex medially. Antennal scapes moderately long: when fully retracted, surpassing the posterior margin of the head capsule by slightly more than the maximum width of the scape (SI 95–109). Antennae 12-segmented; antennal club of composed of three segments, with the apical-most segment longer than the preceding two in combination. Frontal carinae long, extending past the antennal toruli by about three and a half times the maximum width of the antennal scape. Compound eyes moderately protruding past the lateral margins of the head capsule. Lateral margin of head moderately convex, forming a continuous arc from the mandibular insertions to the posterior margin of the head. Posterior head margin convex, rounding evenly into the lateral margins.

In profile view, compound eyes ovular and moderately large (OI 24–26), with 14 ommatidia in longest row. Pronotal declivity indistinct, but neck and anterior face of pronotum forming a ~120 ° angle. Mesosoma predominantly evenly convex from where it joins the pronotal neck to the propodeal spines; propodeum very weakly depressed, so that the dorsal margin dips slightly anterior to the base of the propodeal spines. Promesonotal suture extending from the posterior margin of the procoxal insertion only to the mesothoracic spiracle, which is weakly developed. Metanotal groove visible as a disruption of the sculpture laterally from where it arises between the mid- and hind coxae to where it ends in the poorly developed metathoracic spiracle, which is nearly indistinguishable against the ground sculpture. Propodeal spiracle poorly developed, directed posterolaterally, and separated from the propodeal declivity by about five spiracle diameters. Propodeal spines well developed and moderately long (PSI 30–45), about as long as the propodeal declivity, flared at the base, straight, directed posterodorsally, and acute. Propodeal declivity flat, forming a rounded ~110 ° angle with the base of the propodeal spines. Propodeal lobes rounded and weakly developed. Metapleural gland bulla small, extending from the metacoxal insertion halfway to the propodeal spiracle. Petiole long (PLI 218–273), without tubercles anterodorsally. Subpetiolar process in the form of a small, acute tooth, the ventral face of the petiole weakly bulging below the petiolar node. Petiolar peduncle very long: comprising about three quarters of the petiole. Petiolar node squamiform: node arising abruptly from the petiolar peduncle, forming a rounded ~100 ° angle with it; anterior face rounding evenly into the very short dorsal face, which is strongly convex; dorsal face rounding evenly into the posterior face, which forms a ~90 ° angle with the caudal cylinder. Postpetiole evenly rounded anteriorly, flattened dorsally, and lobed ventrally. Sting extremely well developed: longer than the first gastral sternite.

In dorsal view, humeri well developed and distinct: rounded and wider than the rest of the mesosoma; mesothoracic spiracles weakly protruding past the lateral margins of the mesosoma, visible as slight angles where the pronotum meets the mesonotum. Metanotal groove absent: mesonotum and propodeum completely fused and lateral margins converging evenly to the bases of the propodeal spines. Propodeal spines narrowly approximated basally and joined by a transverse welt; diverging apically, their apices separated from each other by about their length, the negative space between them “V” shaped. Petiolar peduncle with spiracles slightly protruding past the lateral margins. Petiolar node squamiform: transversely ovular when viewed dorsally; node much wider than the peduncle, and a little less than twice as broad as the caudal cylinder. Postpetiole narrow to moderately broad (PWI 179-216) and campaniform, articulating with most of the anterior margin of the gaster, but leaving angulate corners of the gaster exposed on each side. Anterior margin of the postpetiole weakly convex, nearly flat, and evenly rounding into the lateral margins, which diverge slightly to the angulate posterior corners; posterior margin broadly concave. Metafemur moderately to strongly incrassate (FI 261–359).

Sculpture: median clypeal carina present, extending posteriorly to the level of the antennal toruli, and flanked on either side by two equally strong carinae. Lateral clypeal lobes with additional, weaker carinae; ground sculpture shining through weak costulae. Antennal scapes dully shining through weak areolate ground sculpture. Cephalic dorsum predominantly with weak costae, but with indications of weak areolae laterally; median strip and posterior half of the head smooth and shining. Lateral surfaces shallowly areolate, but with stronger rugae surrounding the compound eye, and between the eye and the mandibular insertion. Ventral surface of head smooth and shining, with weak costulae. Mesosoma with areolate sculpture on the pronotal neck. Lateral faces of mesosoma with weak areolae and costulae, which become stronger on the lateral face of the propodeum and meso- and metapleuron. Propodeal declivity shining through shallow areolae. Dorsal surface of pronotum with very weak costulae and areolae, which becomes stronger laterally; the border between the pronotum and mesonotum indicated by a transverse strip of stronger sculpture. Remainder of the mesosoma smooth and shining medially, but with areolate sculpture laterally and with a transverse strip of areolate sculpture directly anterior to the base of the propodeal spines. Femora weakly shining through shallow areolate sculpture. Petiole and postpetiole predominantly smooth and shining; dorsum of the postpetiole with a strip of shallow areolate sculpture along the posterior border. Gaster smooth and shining, without spectral iridescence.

Setae: antennal scapes and funiculi with short, decumbent pilosity. Dorsum of the head, pronotum, waist segments, and gaster with abundant, erect, blunt-tipped setae, the longest of which are about the width of the compound eye. The head dorsum bears ~44, mesosoma ~34, petiole 10, postpetiole ~18, and first gastral tergite ~82 setae. Finer, blunt setae are present on the ventral surface of the head, propleuron, procoxae, and ventral surface of the gaster. Short, sparse pubescence present over the entire body, but is difficult to detect against the dense sculpture.

Color: Predominantly testaceous brown; tarsi and sting yellow.

Gyne measurements & indices (n = 3): SL = 0.603 –0.703 (0.644); FRS = 0.242 –0.265 (0.253); CW = 0.732 –0.812 (0.782); CWb = 0.673 –0.752 (0.722); PoOC = 0.292–0.33 (0.305); CL = 0.765 –0.838 (0.795); EL = 0.225 –0.254 (0.244); EW = 0.166 –0.196 (0.182); MD = 0.165 –0.176 (0.170); WL = 1.187 –1.320 (1.251); SPST = 0.307 –0.402 (0.345); MPST = 0.345 –0.379 (0.367); PEL = 0.390 –0.440 (0.420); NOL = 0.209 –0.288 (0.243); NOH = 0.181 –0.195 (0.188); PEH = 0.323 –0.342 (0.331); PPL = 0.180 –0.224 (0.206); PPH = 0.269 –0.305 (0.290); PW = 0.671 –0.820 (0.741); SBPA = 0.339 –0.408 (0.369); SPTI = 0.349 –0.406 (0.370); PEW = 0.185 –0.243 (0.212); PNW = 0.297 –0.323 (0.307); PPW = 0.403 –0.441 (0.416); HFL = 0.706 –0.800 (0.740); HFWmax = 0.146 –0.168 (0.157); HFWmin = 0.051 –0.059 (0.055); CS = 1.056 –1.160 (1.120); ES = 0.308 –0.352 (0.335); SI = 83–95 (89); OI = 29–30 (30); CI = 88–96 (91); WLI = 166–178 (173); SBI = 48–55 (51); PSI = 25–30 (28); PWI = 166–218 (199); PLI = 192–217 (204); NI = 115–148 (129); PNWI = 133–163 (146); NLI = 53–67 (58); FI = 264–300 (284).

Gyne description: In full-face view, head subquadrate, slightly longer than broad (CI 88–96). Mandibles densely striate, dull, and armed with five teeth: the apical-most well developed, followed by a four roughly equally developed smaller teeth. Anterior clypeal margin very slightly emarginate medially. Antennal scapes moderately long: when fully retracted, just reaching the posterior margin of the head capsule (SI 83–95). Antennae 12-segmented; antennal club of composed of three segments, with the apical-most segment as long as than the preceding two in combination. Frontal carinae long, extending past the antennal toruli by about four 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 evenly to the mandibular insertions. Posterior margin of head weakly convex, rounding evenly into the lateral margins.

In profile view, compound eyes ovular and large (OI 29–30), with 18 ommatidia in longest row. Mesoscutum rounded evenly anteriorly, covering the dorsal surface of the pronotum, and flat dorsally. Mesoscutellum on the same level as the mesoscutum; posterior margin of metanotum extending past the posterior margin of the mesoscutellum. Propodeal spiracle weakly developed, directed posterolaterally, and separated from the propodeal declivity by about five spiracle diameters. Propodeal spines stout and well developed, but short (PSI 25–30), about half as long as the propodeal declivity, flared at the base, weakly downcurved, directed posteriorly, and acute. Propodeal declivity straight and flat, forming a rounded ~100 ° angle with the base of the propodeal spines. Propodeal lobes rounded and very weakly developed. Metapleural gland bulla small, extending from the metacoxal insertion halfway to the propodeal spiracle. Petiole long (PLI 192–217), without tubercles anterodorsally. Subpetiolar process in the form of a weakly developed, acute tooth, which grades evenly into the ventral margin of the petiole posteriorly. Petiolar peduncle very long: comprising about two thirds of the total length of the petiole. Petiolar node squamiform: node arising abruptly from the petiolar peduncle, forming a rounded ~120 ° angle with it; anterior face rounding evenly into the very short dorsal face, which is strongly convex; dorsal face rounding evenly into the posterior face, which forms a ~90 ° angle with the caudal cylinder. Postpetiole evenly rounded anteriorly, dorsal face bulging anterodorsally before flattening posteriorly; ventral surface lobed.

In dorsal view, mesoscutum covering pronotum anteriorly, but humeri visible laterally as rounded sclerites. Propodeal spines diverging basally, but parallel distally, their apices separated from each other by about two and a half times their length. Petiolar peduncle with spiracles protruding past the lateral margins, the peduncle broadened where they arise. Petiolar node squamiform: transversely oval when viewed dorsally; node much wider than the peduncle, and a about one and a half times as broad as the caudal cylinder. Postpetiole narrow to moderately broad (PWI 166–218), anteroposteriorly compressed, and subquadrate, articulating with most of the anterior margin of the gaster, leaving small, angulate margins on each side of the gaster exposed. Anterior margin of postpetiole very weakly convex, nearly flat, with corners evenly rounding into the lateral margins, which evenly diverge to the angulate posterior corners; posterior margin weakly concave.

Metafemur moderately incrassate (FI 264–300).

Sculpture: median clypeal carina present, extending nearly to the frontal triangle, and flanked by additional weaker carinae over weakly areolate ground sculpture. Antennal scapes areolate and weakly shining. Cephalic dorsum densely costate, over weak areolate sculpture. Lateral surface of head with denser rugae forming whorls around the compound eye and forming transverse reticulations between the compound eye and the mandibular insertions. Ventral surface of head weakly costulate. Pronotal neck areolate. Pronotum, propodeum, and posterior two thirds of the anepisternum and katepisternum sculptured like the cephalic dorsum; anterior thirds of anepisternum and katepisternum smooth and shining. Propodeal declivity shallowly areolate. Mesoscutum costulate. Mesoscutellum costulate, with a patch of smooth and shining sculpture posteromedially. Femora shining through weak areolate sculpture. Petiole weakly areolate, with the anterior face of the node smooth and shining, and the dorsum transversely striate. Dorsal surface of postpetiole shallowly areolate. Gaster smooth and shining, only faint traces of spectral iridescence on the first gastral tergite and sternite.

Setae: antennal scapes and funiculi with short, decumbent pilosity. Dorsum of the head, pronotum, waist segments, and gaster with abundant, erect, blunt-tipped setae, the longest of which are about a third of the width of the compound eye. Finer, blunt setae are present on the ventral surface of the head, propleuron, procoxae, and ventral surface of the gaster. Short, sparse pubescence present over the entire body, but is difficult to detect against the dense sculpture.

Color: predominantly testaceous brown; tarsi yellow.

Male measurements & indices (n = 3): SL = 0.368 –0.421 (0.402); FRS = 0.107 –0.142 (0.126); CW = 0.505 –0.537 (0.519); CWb = 0.439 –0.473 (0.455); PoOC = 0.219 –0.234 (0.225); CL = 0.503 –0.529 (0.514); EL = 0.217 –0.228 (0.222); EW = 0.175 –0.191 (0.184); MD = 0.033 –0.051 (0.040); WL = 0.810 –0.877 (0.841); SPST = n/a; MPST = 0.268 –0.295 (0.280); PEL = 0.273 –0.289 (0.282); NOL = 0.151 –0.186 (0.167); NOH = 0.039 –0.072 (0.056); PEH = 0.134 –0.161 (0.145); PPL = 0.123 –0.154 (0.140); PPH = 0.170 –0.181 (0.174); PW = 0.511 –0.558 (0.537); SBPA = n/a; SPTI = n/a; PEW = 0.131 –0.184 (0.149); PNW = 0.157 –0.219 (0.182); PPW = 0.248 –0.276 (0.261); HFL = 0.596 –0.679 (0.634); HFWmax = 0.076 –0.090 (0.083); HFWmin = 0.044 –0.053 (0.048); CS = 0.694 –0.738 (0.712); ES = 0.310 –0.321 (0.314); SI = 81–96 (89); OI = 42–46 (44); CI = 86–90 (88); WLI = 179–190 (185); SBI = n/a; PSI = n/a; PWI = 150–195 (178); PLI = 188–222 (203); NI = 258–387 (313); PNWI = 119–127 (122); NLI = 55–64 (59); FI = 158–205 (175).

Male description: In full-face view, head subglobular, slightly longer than broad (CI 86–90). Mandibles smooth and 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 crenulae. Anterior clypeal margin evenly rounded medially. Antennal scapes moderately long: when fully retracted, just reaching the posterior margin of the head capsule (SI 81–96). Antennae 13-segmented; antennal club of composed of three segments, with the apical-most segment as long as than the preceding three in combination. Frontal carinae long, extending past the antennal toruli by about four times the maximum width of the antennal scape. Compound eyes strongly protruding past the lateral margins of the head capsule. Posterior margin of head strongly convex, not differentiated from lateral margins.

In profile view, compound eyes ovular and very large (OI 42–46), with 22 ommatidia in longest row. Mesoscutum rounded evenly anteriorly, covering the dorsal surface of the pronotum, and flat dorsally. Mesoscutellum inflated and much higher than the mesoscutum; posterior margin of mesoscutellum overhanging the posterior margin of the metanotum. Propodeal spiracle weakly developed, directed posterolaterally, and separated from the propodeal declivity by about four spiracle diameters. Propodeal spines absent. Propodeum evenly rounded. Propodeal lobes rounded and very weakly developed. Metapleural gland bulla small, extending from the metacoxal insertion halfway to the propodeal spiracle. Petiole long (PLI 188–222), without tubercles anterodorsally. Subpetiolar process absent; ventral margin of petiole weakly bulging medially. Petiolar peduncle very long: comprising about two thirds of the total length of the petiole. Petiolar node low and cuneiform: node evenly grading into the petiolar peduncle anteriorly; anterior face meeting the posterior face at a ~90 ° angle. Postpetiole evenly rounded anteriorly, flattened dorsally; ventral surface concave.

In dorsal view, mesoscutum covering pronotum anteriorly; humeri barely visible laterally. Propodeal spines absent. Petiolar peduncle with spiracles protruding past the lateral margins, the peduncle broadened where they arise. Petiolar node cuneiform: dorsum flat when viewed posterodorsally, and narrower than the caudal cylinder. Postpetiole narrow (PWI 150–195) and campaniform, articulating with most of the anterior margin of the gaster, leaving angulate margins on each side of the gaster exposed. Anterior margin of postpetiole weakly concave, nearly flat, with corners evenly rounding into the lateral margins, which evenly diverge to the angulate posterior corners; posterior margin weakly concave. Metafemur not incrassate (FI 158–205).

Sculpture: median clypeal carina present, extending nearly the level of the antennal toruli, and flanked by additional weaker carinae over smooth ground sculpture. Antennal scapes weakly areolate and shining. Cephalic dorsum densely areolate, with costulae radiating from the frontal triangle. Lateral surface of head with denser rugae forming whorls around the compound eye. Ventral surface of head weakly areolate. Pronotal neck weakly areolate. Pronotum smooth and shining. Anterior half of the katepisternum and lateral portions of the anepisternum smooth and shining, otherwise areolate. Metapleuron areolate. Lateral face of propodeum areolate; propodeal declivity smooth and shining. Median strip of mesoscutum costulate over weak areolate sculpture; smooth and shining laterally. Mesoscutellum smooth and shining medially; lateral faces costulate over weak areolate sculpture. Femora shining through weak areolate sculpture, which becomes stronger distally. Petiole weakly areolate, with the anterior face of the node smooth and shining. Dorsal surface of postpetiole smooth and shining, with posterior quarter shallowly areolate. Gaster smooth and shining, without spectral iridescence.

Setae: antennal scapes and funiculi with short, decumbent pilosity. Dorsum of the head, pronotum, waist segments, and gaster with abundant, erect, blunt-tipped setae, the longest of which are about a third of the width of the compound eye. Finer, blunt setae are present on the ventral surface of the gaster. Short, sparse pubescence present over the entire body, but is difficult to detect against the sculpture.

Color: predominantly testaceous brown; mandibles, antennae, coxae, tarsi, and genitalia yellow.

Etymology: Systematic, from the Latin ‘subditivus’ (= spurious, counterfeit). W.M. Wheeler originally assigned this species to Macromischa with reservations; the species epithet is likely a reflection of his doubts.

Comments: Temnothorax subditivus has the largest range of any salvini clade species, which encompasses much of southern North America, all of Mesoamerica, parts of northern South America, and the Cayman Islands. Temnothorax subditivus is also highly morphologically variable, with forms that are uniformly dark brown to uniformly yellow, densely sculptured to mostly smooth and shining ( Fig. 168 ). Temnothorax subditivus inhabits dry forests and disturbed habitats from sea level to high elevations, and is often found nesting in hollow, dead vegetation, under the bark of trees, or in epiphytes, but may also be found in rock crevices or abandoned termite burrows (this study; Achury, Chacón de Ulloa & Arcila, 2008; Creighton, 1965; Escalante-Jiménez & Vasquez-Bolaños, 2015; Gutiérrez-Martínez & Naranjo, 2016; Pérez-Sánchez, Lattke & Viloria Petit, 2012; Rojas, Fragoso & Mackay, 2014; Wheeler, 1903a). Colonies are small, with 12 to 145 workers, and monogynous ( Creighton, 1965). They are apparently trophic generalists, like many Temnothorax for which food preference is known, but favour insects ( Creighton, 1965). The male pupae bear bizarre appendages with unknown function ( Schumann, 1992). Temnothorax subditivus is most closely related to T. politus of the Sonoran Desert. The high degree of morphological variability and broad range have led taxonomists to assign a large number of names to this species, but the phylogeny of this species shows a great deal of geographical structure, with a pectinate series from the northern part of the range to the southern part (Prebus in prep.). This pattern could be attributed to either isolation by distance or to a recent radiation of closely related species. Increased sampling along with species delimitation analysis across the range of T. subditivus should elucidate whether this taxon represents a broadly distributed morphologically and ecologically variable species or a species complex.

Figures

Figure 165 Geographical and elevational distribution of the subditivus group. (A) Temnothorax politus (B) T. subditivus. 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-165

Figure 167 Temnothorax suditivus. (A–C) Worker (CASENT0758340). (A) Full-face view. (B) Profile view. (C) Dorsal view. (D–G) Gyne (CASENT0758338). (D) Full-face view. (E) Profile view. (F) Dorsal view. (G) Wings. (H–K) Male (CASENT0915978). (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-167

Figure 168 Morphological variability in Temnothorax suditivus workers. (A–C) Estelí, Nicaragua (CASENT0619368). (A) Full-face view. (B) Profile view. (C) Dorsal view. (D–F) Jalisco, Mexico (CASENT0758299). (D) Full-face view. (E) Profile view. (F) Dorsal view. (G–I) Grand Cayman, Cayman Islands (CASENT0868474). (G) Full-face view. (H) Profile view. (I) Dorsal view. Scale bars 0.2 mm. Full-size DOI: 10.7717/peerj.11514/fig-168