Temnothorax wilsoni
Temnothorax wilsoni | |
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Scientific classification | |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Hymenoptera |
Family: | Formicidae |
Subfamily: | Myrmicinae |
Tribe: | Crematogastrini |
Genus: | Temnothorax |
Species group: | salvini |
Species subgroup: | pulchellus |
Species: | T. wilsoni |
Binomial name | |
Temnothorax wilsoni Prebus, 2021 |
Temnothorax wilsoni is known only from high elevations sites in Valle Nuevo National Park in the Dominican Republic with open Pinus occidentalis forests, where several nests were collected from under stones in the Cordillera Central. All nest collections contained only one dealate gyne, suggesting that this species may be monogynous. Other details of the biology of this species remain unknown but are probably similar to other members of the terricolous pan-Caribbean pulchellus group. (Prebus 2021)
Identification
Prebus (2021) – A member of the pulchellus group of the Temnothorax salvini clade. Temnothorax wilsoni can be separated from all other species in the salvini clade by the following character combination: anterior clypeal margin medially emarginate; in profile view, dorsum of mesosoma weakly convex; metanotal groove not impressed; propodeum not depressed below the level of the promesonotum; propodeum bearing standing setae dorsally; propodeal spines longer than the propodeal declivity, directed posteriorly, and weakly downcurved; in dorsal view, propodeal spines broadly approximated, the negative space between them "U" shaped; hind femora weakly to strongly incrassate; in dorsal view, apex of petiolar node narrower than the caudal cylinder of the petiole; petiolar node with four erect setae dorsally; postpetiole very broad: greater than or equal to 2.3 times the width of the petiole; dorsum of head smooth and shining; setae on head, mesosoma, waist segments and gaster erect, moderately long, sparse and blunt (never long and tapering);integument tricolored: predominantly testaceous yellow, with a dark brown head capsule; femora, postpetiole, and lower half of the mesosomal pleurites testaceous; gaster yellow, often with two posterolateral slightly darker spots on the first gastral tergite.
This species displays variation in coloration of the gaster within nest series, ranging from having two testaceous spots posterolaterally on the first gastral tergite to being completely yellow. This species may form a species complex with Temnothorax bahoruco, Temnothorax balaclava, and Temnothorax ciferrii, all of which have only been collected from allopatric populations in mountains and low-lying dry forest from the southern parts of Hispaniola. It is morphologically quite similar to T. bahoruco and T. balaclava, differing primarily in the conformation of the clypeus and in coloration.
Similar species: Temnothorax bahoruco, Temnothorax balaclava, Temnothorax ciferrii, Temnothorax harlequina, Temnothorax hippolyta, Temnothorax magnabulla, Temnothorax pulchellus, Temnothorax schwarzi, and Temnothorax terricola. The erect setae count of four on the dorsum of the petiolar node differentiates T. wilsoni from T. terricola and T. hippolyta, which have only two along the posterior margin. The presence of erect setae on the propodeum distinguishes T. wilsoni from T. pulchellus. The relatively small metapleural gland bulla of T. wilsoni, which reaches two thirds of the way to from the metacoxal insertion to the propodeal spiracle, distinguishes it from T. magnabulla, in which it extends three quarters of the way or more. Temnothorax wilsoni and T. harlequina can be separated most easily by the color of the integument: T. harlequina has a light yellow gaster which strongly contrasts with the darker integument of the rest of the body, whereas the gaster of T. wilsoni is always medium yellow, or bears dark spots; additionally, the dorsum of the mesosoma and waist segments of T. wilsoni are always testaceous yellow. The mesosoma is of T. wilsoni is less arched and more densely sculptured in comparison to T. harlequina Temnothorax wilsoni could also be confused with T. schwarzi due to the smooth and shining head capsule, but the relatively narrow apex of the petiolar node of T. wilsoni will separate the two. The medially emarginate anterior clypeal margin of T. wilsoni will differentiate it from T. balaclava and T. bahoruco
Keys including this Species
Distribution
Known from open, high-elevation Pinus occidentalis forest sites in the Cordillera Central.
Distribution based on Regional Taxon Lists
Neotropical Region: Dominican Republic (type locality).
Distribution based on AntMaps
Distribution based on AntWeb specimens
Check data from AntWeb
Countries Occupied
Number of countries occupied by this species based on AntWiki Regional Taxon Lists. In general, fewer countries occupied indicates a narrower range, while more countries indicates a more widespread species. |
Biology
Castes
Queen
Male
Nomenclature
The following information is derived from Barry Bolton's Online Catalogue of the Ants of the World.
- wilsoni. Temnothorax wilsoni Prebus, 2021: 337, figs. 133O, 149 (w.dq.m.) DOMINICAN REPUBLIC.
Unless otherwise noted the text for the remainder of this section is reported from the publication that includes the original description. Unless otherwise noted the text for the remainder of this section is reported from the publication that includes the original description.
Description
Worker
(n = 10): SL = 0.462-0.537 (0.505); FRS = 0.202-0.231 (0.213); CW = 0.548-0.628 (0.595); CWb = 0.512-0.579 (0.546); PoOC = 0.234-0.273 (0.251); CL = 0.596-0.671 (0.636); EL = 0.136-0.165 (0.150); EW = 0.104-0.133 (0.120); MD = 0.134-0.164 (0.148); WL = 0.692-0.781 (0.731); SPST = 0.226-0.289 (0.256); MPST = 0.219-0.274 (0.233); PEL = 0.293-0.348 (0.318); NOL = 0.158-0.187 (0.175); NOH = 0.102-0.125 (0.111); PEH = 0.190-0.231 (0.211); PPL = 0.171-0.209 (0.188); PPH = 0.212-0.245 (0.231); PW = 0.367-0.428 (0.393); SBPA = 0.147-0.187 (0.171); SPTI = 0.251-0.316 (0.287); PEW = 0.144-0.177 (0.161); PNW = 0.119-0.157 (0.141); PPW = 0.346-0.408 (0.376); HFL = 0.474-0.530 (0.504); HFWmax = 0.122-0.150 (0.136); HFWmin = 0.045-0.062 (0.050); CS = 0.810-0.915 (0.864); ES = 0.188-0.228 (0.210); SI = 88-98 (93); OI = 23-26 (24); CI = 84-88 (86); WLI = 129-138 (134); SBI = 29-34 (31); PSI = 29-39 (35); PWI = 225-241 (234); PLI = 146-187 (170); NI = 145-168 (157); PNWI = 77-96 (88); NLI = 52-61 (55); FI = 240-313 (270).
In full-face view, head subquadrate, longer than broad (CI 84-88). 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 emarginated medially. Antennal scapes moderately long: when fully retracted, just reaching the posterior margin of the head capsule (SI 88-98). Antennae 12-segmented; antennal club of composed of three segments, with the apical-most segment slightly longer than the preceding two in combination. Frontal carinae moderately long, extending past the antennal toruli by about two times the maximum width of the antennal scape. Compound eyes moderately protruding past the lateral margins of the head capsule. Lateral margin of head weakly 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 9 ommatidia in longest row. Pronotal declivity indistinct, neck and anterior face of pronotum forming a ~120° angle. Mesosoma weakly convex from where it joins the pronotal neck to the propodeal spines. Promesonotal suture extending from the posterior margin of the procoxal insertion only to the mesothoracic spiracle, which is moderately well 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 well developed, directed posterolaterally, and separated from the propodeal declivity by about three spiracle diameters. Propodeal spines well developed and moderately long (PSI 29-39), about as long as the propodeal declivity, tapering evenly from the base, weakly downcurved, and acute. Propodeal declivity weakly concave, forming a rounded ~100° angle with the base of the propodeal spines. Propodeal lobes rounded and weakly developed. Metapleural gland bulla moderately large, extending from the metacoxal insertion two thirds of the way to the propodeal spiracle. Petiole moderately long (PLI 146-187), without tubercles anterodorsally. Subpetiolar process in the form of a small, acute tooth; ventral margin of petiole weakly concave posterior to it. Petiolar peduncle short: petiolar node covering most of the petiolar dorsum. Petiolar node robust: transition between peduncle and node marked by a rounded angle of ~140°, resulting in a weakly concave anterior node face; anterior face forming a ~90° angle with the dorsal face, which is weakly convex; dorsal face rounding evenly into the short posterior face, which forms a ~110° angle with the caudal cylinder. Postpetiole evenly rounded anteriorly, flattened dorsally, and weakly lobed ventrally.
In dorsal view, humeri developed and distinct: evenly 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 broadly approximated basally and diverging apically, their apices separated from each other by about their length, the negative space between them "U" shaped. Petiolar peduncle with spiracles protruding past the lateral margins, but not noticeably constricted anterior to them. Petiolar node evenly ovular; node the same width as the peduncle, and evenly grading into the caudal cylinder, which is wider than the node. Postpetiole very broad (PWI 225-241) and campaniform, articulating with the nearly the entire anterior margin of the gaster. Anterior margin of the postpetiole convex and evenly rounds into the lateral margins, which diverge to the angulate posterior corners; posterior margin broadly concave. Metafemur weakly to strongly incrassate (FI 240-313).
Sculpture: median clypeal carina present, extending posteriorly nearly to the frontal triangle, and flanked on either side by two equally strong carinae. Lateral clypeal lobes with additional, weaker carinae; ground sculpture smooth and shining. Antennal scapes shining through weak areolate ground sculpture. Cephalic dorsum smooth and shining, but with coarse piligerous punctures; costulae flanking the frontal carinae. Lateral surfaces of head with weak areolate sculpture posterior to the compound eye, dense rugulose sculpture surrounding the compound eye, and weak rugae between the compound eye and the mandibular insertion. Ventral surface of head smooth and shining. Mesosoma with areolate sculpture on the pronotal neck. Lateral surfaces of the pronotum, mesopleurae, and anterior half of the lateral surface of propodeum areolate, but smooth and shining between the propodeal spiracle and the propodeal spines. Dorsal surface of mesosoma with weak costulae on the pronotum and lateral margins over areolate ground sculpture. Femora shining, with weak areolate sculpture on the distal third. Petiole smooth and shining ventrally, with areolate sculpture on all other surfaces; a weak carina present laterally, extending longitudinally from the petiolar spiracle to the caudal cylinder. Postpetiole smooth and shining, with weak areolate sculpture on the posterior quarter. 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 moderately abundant, erect, blunt-tipped setae, the longest of which are about the width of the compound eye. The head bears ~30, mesosoma ~18, petiole 4, postpetiole ~14, and first gastral tergite ~36 setae. Short, sparse pubescence present over the entire body, but difficult to detect against the lightly colored integument.
Color: predominantly testaceous yellow, with a dark brown head capsule. The femora, postpetiole, and lower half of the mesosomal pleurites are testaceous. Two slightly darker spots are present on the first gastral tergite posterolaterally.
Queen
(n = 2): SL = 0.559-0.565 (0.562); FRS = 0.267-0.268 (0.268); CW = 0.772-0.779 (0.776); CWb = 0.704-0.720 (0.712); PoOC = 0.271-0.272 (0.272); CL = 0.707-0.726 (0.717); EL = 0.234-0.239 (0.237); EW = 0.175-0.190 (0.183); MD = 0.135-0.142 (0.139); WL = 1.212-1.239 (1.226); SPST = 0.288-0.306 (0.297); MPST = 0.285-0.303 (0.294); PEL = 0.424-0.433 (0.429); NOL = 0.202-0.213 (0.208); NOH = 0.142-0.143 (0.143); PEH = 0.294-0.299 (0.297); PPL = 0.191-0.211 (0.201); PPH = 0.299-0.322 (0.311); PW = 0.748-0.838 (0.793); SBPA = 0.354-0.356 (0.355); SPTI = 0.333-0.367 (0.350); PEW = 0.199-0.207 (0.203); PNW = 0.221-0.234 (0.228); PPW = 0.487-0.535 (0.511); HFL = 0.646-0.654 (0.650); HFWmax = 0.135-0.141 (0.138); HFWmin = 0.052-0.058 (0.055); CS = 1.058-1.083 (1.070); ES = 0.327-0.329 (0.328); SI = 78-79 (79); OI = 30-31 (31); CI = 99-100 (99); WLI = 172-172 (172); SBI = 49-51 (50); PSI = 23-25 (24); PWI = 245-258 (252); PLI = 205-222 (214); NI = 141-150 (146); PNWI = 111-113 (112); NLI = 48-49 (48); FI = 243-260 (251).
In full-face view, head subquadrate, about as long as broad (CI 99-100). 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 weakly emarginated medially. Antennal scapes moderately long: when fully retracted, just reaching the posterior margin of the head capsule (SI 78-79). 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, 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 30-31), with 17 ommatidia in longest row. Mesoscutum rounded evenly anteriorly, covering the dorsal surface of the pronotum, and flat dorsally. Mesoscutellum slightly depressed below the level of the mesoscutum. 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 three spiracle diameters. Propodeal spines stout and well developed, but short (PSI 23-25), about as two thirds as long as the propodeal declivity, tapering evenly from the base, directed posteriorly, straight, and blunt. Propodeal declivity straight and flat, forming a rounded ~90° angle with the base of the propodeal spines. Propodeal lobes rounded and very weakly developed. Metapleural gland bulla moderately large, extending from the metacoxal insertion two thirds of the way to the propodeal spiracle. Petiole long (PLI 205-222), without tubercles anterodorsally. Subpetiolar process in the form of a small, very acute tooth, which grades evenly into the ventral margin of the petiole posteriorly. Petiolar peduncle short: petiolar node covering most of the petiolar dorsum. Petiolar node erect: transition between peduncle and node evenly rounded, resulting in a very slightly concave anterior node face; anterior face forming a very sharp ~80° angle with the dorsal face, which is short; dorsal face rounding evenly into the posterior face, which forms a ~100° angle with the caudal cylinder. Postpetiole evenly rounded anterodorsally, bulging slightly before it transitions into the flattened dorsal face; ventral surface weakly lobed.
In dorsal view, mesoscutum covering pronotum anteriorly, but humeri visible laterally as rounded sclerites. Propodeal spines weakly diverging apically, their apices separated from each other by about one and a half times their length. Petiolar peduncle with spiracles not protruding past the lateral margins. Petiolar node widest anteriorly, and emarginated anterodorsally; lateral faces converge to the indistinct posterior face. Petiolar node slightly narrower than the peduncle, and evenly grading into the caudal cylinder, which is about the same width as the node. Postpetiole very broad (PWI 245-258), anteroposteriorly compressed, and campaniform, articulating with most of the anterior margin of the gaster, leaving small, angulate margins on each side exposed. Anterior margin of postpetiole weakly convex, with corners marked by rounded angles as it transitions to the lateral margins, which evenly diverge to the angulate posterior corners; posterior margin broadly concave. Metafemur weakly to moderately incrassate (FI 243-260).
Sculpture: median clypeal carina present, extending from the anterior margin nearly to frontal triangle, and flanked by weaker, indistinct carinae; lateral margins of median clypeal lobe with two carinae that are as strong as the medial carina. Lateral clypeal lobes with additional weaker carinae; ground sculpture smooth and shining. Antennal scapes smooth and shining. Cephalic dorsum mostly smooth and shining, with costulae flanking the frontal carinae, and a median carina extending from the frontal triangle nearly to the median ocellus; weak concentric costulae surrounding the antennal insertions. Lateral surfaces of head with weak areolate sculpture posterior to the compound eye, dense rugulose sculpture surrounding the compound eye, and rugae between the compound eye and the mandibular insertion. Ventral surface of head with weak rugulae. Pronotal neck areolate. Pronotum with weak areolate ground sculpture arranged into longitudinal rows and separated by superficial costulae. Anepisternum and katepisternum shining on their anterior halves, transitioning into weak costulae posteriorly. Propodeum with stronger costulae laterally. Propodeal declivity weakly areolate. Mesoscutum with costulae over weak areolate ground sculpture surrounding a smooth and shining central strip which extends over half of the sclerite from the anterior margin; smooth and shining patches laterally. Mesoscutellum smooth and shining medially, surrounded by weak costulae and areolae. Femora smooth and shining, with traces of weak areolate sculpture distally. Petiole with weak areolate sculpture laterally, and on the dorsoposterior surface of the node. Dorsal surface of the peduncle and the anterior face of the node shining through weak sculpture. Postpetiole smooth and shining anteriorly, areolate laterally and on the posterior half. Gaster smooth and shining, without faint traces of spectral iridescence. Surface of the first gastral sternite smooth and shining.
Setae: antennal scapes and funiculi with short, decumbent pilosity. Dorsum of the head, pronotum, waist segments, and gaster with moderately abundant, erect, blunt-tipped setae, the longest of which are about a third of the width of the compound eye. Short, sparse pubescence present over the entire body, but difficult to detect against the lightly colored integument.
Color: head capsule, anepisternum, propodeum, mesoscutellum, waist segments (except petiolar node), posterior half of the first gastral tergite and the following tergites dark brown. The mandibles, antennae, pronotum, katepisternum, mesoscutum, propodeal spines, petiolar node, and femora testaceous. Postpetiole and basal half of first gastral tergite yellow.
Male
(n = 1): SL = 0.329; FRS = 0.143; CW = 0.563; CWb = 0.494; PoOC = 0.204; CL = 0.514; EL = 0.248; EW = 0.172; MD = 0.038; WL = 0.926; SPST = 0; MPST = 0.231; PEL = 0.322; NOL = 0.225; NOH = 0.055; PEH = 0.154; PPL = 0.180; PPH = 0.197; PW = 0.501; SBPA = n/a; SPTI = n/a; PEW = 0.143; PNW = 0.155; PPW = 0.301; HFL = 0.681; HFWmax = 0.058; HFWmin = 0.036; CS = 0.751; ES = 0.334; SI = 67; OI = 44; CI = 96; WLI = 187; SBI = n/a; PSI = n/a; PWI = 210; PLI = 179; NI = 409; PNWI = 108; NLI = 70; FI = 161.
In full-face view, head subovate, slightly longer than broad (CI 96). 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 moderately long: when fully retracted, just reaching the posterior margin of the head capsule (SI 67). Antennae 13-segmented; antennal club composed of four segments, with the apical-most segment as long as 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. Lateral margin of head convex, margin between the anterior margin of the compound eye and the mandibular insertions straight. Posterior head margin evenly convex, rounding evenly into the lateral margins.
In profile view, compound eyes ovular and large (OI 44), with 22 ommatidia in the longest row. Mesoscutum bulging anteriorly, covering the dorsal surface of the pronotum, flat dorsally, and rounded posteriorly. Mesoscutellum depressed slightly below the level of the mesoscutum; posterior margin even with the posterior margin of the metanotum, not overhanging it. Propodeal spiracle well developed, directed posterolaterally, and separated from the propodeal declivity by about three spiracle diameters. Propodeal spines absent, indicated by weak carinae on the dorsal and declivitous faces of the propodeum, which converge into a rounded angle. Propodeal lobes rounded and weakly developed. Metapleural gland bulla small, extending halfway between the insertion of the metacoxa and the propodeal spiracle. Petiole moderately long (PLI 179), without tubercles anterodorsally. Subpetiolar process absent. Petiolar peduncle short: comprising less than a quarter of the total length of the petiole. Petiolar node low and rounded, the convergence of the anterior and dorsal 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 barely visible laterally as slivers of rounded sclerites. Petiolar peduncle with spiracles slightly protruding past the lateral margins, the peduncle broadened where they arise. Petiolar node 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 moderately broad (PWI 210) and campaniform, articulating with most of the anterior margin of the gaster, leaving small angulate margins on each side exposed. Anterior margin of postpetiole convex, with the anterior corners evenly rounding into the lateral margins, which evenly diverge to the angulate posterior corners; posterior margin of postpetiole broadly concave. Metafemur not incrassate (FI 161).
Sculpture: median clypeal carina present but very weak and rounded, flanked by additional weak carinae over smooth and shining ground sculpture. Antennal scapes smooth and shining. Head areolate, with weak costulae flanking the frontal carinae and a central strip of weak sculpture extending from the frontal triangle to the median ocellus. Lateral surfaces of pronotum, anepisternum, katepisternum, and metasternum smooth and shining. Propodeum weakly areolate laterally, nearly smooth on the dorsum and declivity. Dorsally, mesoscutum weakly costulate along the Mayrian furrows, otherwise smooth and shining. Mesoscutellum costulate, but sculpture weaker medially. Femora smooth and shining, with traces weak areolate sculpture distally. Petiole with shallow areolate sculpture ventrally, the dorsal surface of the node smooth and shining. Dorsal surface of postpetiole shining, with traces of strigulae, especially on the posterior quarter. Gaster smooth and shining.
Setae: antennal scapes and funiculi with short, decumbent pilosity. Dorsum of the head, pronotum, waist segments, and gaster with moderately abundant, erect, blunt-tipped setae, the longest of which are about a quarter of the width of the compound eye. Short, sparse pubescence present over the entire body, but difficult to detect against the integument.
Color: predominantly dark brown, with mandibles, antennae, legs, and genitalia yellow. Pronotum, wing bases, coxae, and gastral segments (excluding the first) testaceous.
Type Material
Holotype worker: DOMINICAN REPUBLIC: La Vega: Valle Nuevo Scientific Reserve, 18°46′41″N 70°38′22″W / 18.77800°N 70.63950°W ± 5 m, 2,255 m, 3 December 2003, S.P. Cover #SPCDR-53, pine woodland, nest under rock (MCZENT00583608, top specimen on pin) Museum of Comparative Zoology.
Paratype workers and gyne: same pin as holotype (bottom specimen on pin) [MCZC]; same data as holotype, 2 workers (MCZENT00583609) [MCZC], 1 worker (CASENT0756092) Museo Nacional de Historia Natural "Prof. Eugenio de Jesús Marcano", 1 dealate gyne (MCZENT00583610) [MCZC].
Etymology
Patronym, named in honor of E.O. Wilson, a naturalist and myrmecologist of unmeasurable influence, whose expedition to Hispaniola in 2003 recovered all of the known specimens of Temnothorax wilsoni used in this study.
References
- Cantone S. 2018. Winged Ants, The queen. Dichotomous key to genera of winged female ants in the World. The Wings of Ants: morphological and systematic relationships (self-published).
- Prebus, M.M. 2021. Taxonomic revision of the Temnothorax salvini clade (Hymenoptera: Formicidae), with a key to the clades of New World Temnothorax. PeerJ 9, e11514 (doi:10.7717/peerj.11514).