Three new species of Cataglyphis Foerster, 1850 (Hymenoptera, Formicidae) from Iran

Abstract Cataglyphis bazoftensissp. nov., C. fritillariaesp. nov., and C. dejdaranensissp. nov. are described from the Koohrang county of central-west Iran (Chaharmahal va Bakhtiari Province). All new species belong to the C. altisquamis complex and are characterized by presence of the thick, black, and decumbent setae on lateral and posterior surfaces of tibiae. Additionally, a key to Asian Cataglyphis was updated to accommodate the new species.


Introduction
Cataglyphis Foerster, 1850 is a moderately large ant genus comprising 94 valid species and 18 valid subspecies divided into nine species groups (Agosti 1990;Bolton 2020). Its representatives are distributed mostly in the Palearctic Region's semideserts and deserts (Agosti 1990;Brown 2000). However, Cataglyphis species were also recorded from other arid habitats, such as high altitude, mountain steppes, and forest steppes (Agosti 1990;Brown 2000). Members of the genus are among the commonest ants of the arid ecosystems of North Africa, Arabian Peninsula, and Central Asia, where they build crater nests directly in the ground (Agosti 1990;Collingwood and Agosti 1996;Brown 2000) and feed on dead insects (Lenoir et al. 2010). They have been reported as flower pollinators (Herrera et al. 1984) and as contributors in myrmecochorous seed dispersal (Hulme 1997;Boulay et al. 2007;Wehner 2020).
Many Cataglyphis species are polymorphic what causes challenges in their determination. Additionally, distinct polymorphism makes it necessary to prepare descriptions of new species based on nest samples. This approach should ensure comprehensive descriptions based on a detailed overview of infraspecific variability of studied samples. One should remember that genetic studies of some species did not confirm their homogeneity and suggested the presence of cryptic species and a high level of hybridization (Ionescu and Eyer 2016;Eyer et al. 2017).
Due to its location, geography, and predominance of open and arid habitats, Iran hosts the highest number of Cataglyphis worldwide (Paknia et al. 2008, Janicki et al. 2016. So far, there are 32 species of the genus known from the county, but some records need verification and confirmation (Paknia et al. 2008(Paknia et al. , 2009Moradloo et al. 2015;Rad et al. 2018). The present work is a contribution to the understanding of the Iranian Cataglyphis. We describe three new species of the C. altisquamis species group (sensu Agosti 1990): C. bazoftensis sp. nov., C. fritillariae sp. nov., and C. dejdaranensis sp. nov. based on the worker caste. The C. altisquamis species group is characterized by the following combination of characters in the worker caste: relatively large (WL up to 5 mm); body dull, uniformly yellow-black to black or bicolored with black gaster, petiole cuneiform or pseudo-nodiform, head finely reticulate with punctulate frons. The geographic range of this group extends from Portugal and Morocco to Central Asia. Members of this group were included in Radchenko's key (1998) to the Asian Cataglyphis. Herein, we modify the key to accommodate the new species. Additionally, we also included in the key C. asiriensis Collingwood, 1985, a member of the altisquamis species group known from Saudi Arabia, which was omitted by Radchenko (1998).

Materials and methods
Investigated specimens were collected from five sites in the Koohrang County, located in the northern part of the Chaharmahal va Bakhtiari Province of Iran. All sites were placed at altitude from 1738 to 2778 m. a.s.l. The county is surrounded by the Zagros Mountains, one of the two largest mountain ranges of Iran, and is characterized by an alpine climate. The only exception is the Bazoft region, which is warmer than other parts of the Koohrang County and covered by deciduous oak forests.
The dominant sampling method was a direct sampling (hand collecting). Individual specimens were collected on the ground or from nests under stones. All specimens were preserved in 75% EtOH. Photos were taken using a Nikon SMZ 1500 stereomicroscope, Nikon D5200 photo camera, and Helicon Focus software. All given label data are in the original spelling, only the geographic coordinates are given in decimal notation, instead of the degrees, minutes, seconds on the labels; a vertical bar (|) separates data on different rows and double vertical bars (||) separate labels. Type specimens' photographs are available online on AntWeb (www.AntWeb.org) and are accessible using the unique CASENT identifying specimen code.
Measurements: all measurements are given in mm.

HFL
hind femur length; measured on dorsal side from trochanter to apex of femur; HL head length; measured in a straight line from mid-point of anterior clypeal margin to mid-point of posterior margin in full-face view; HW head width; measured in full-face view directly behind the eyes; PRL propodeum length; measured in lateral view, from metanotal groove to posterior-most point of propodeum; PRW propodeal width; maximum width of propodeum in dorsal view; PTH petiole height; the chord of ventral petiolar profile at node level is the reference line perpendicular to which the maximum height of petiole is measured, measured in lateral view; PTW petiole width; maximum width of the petiolar node in lateral view; PW pronotum width; maximum width of pronotum in dorsal view; SL scape length; maximum straight-line length of scape excluding the basal condylar bulb; WL Weber's length; measured as diagonal length from the anterior end of the neck shield to the posterior margin of the propodeal lobe.

Synoptic list of Cataglyphis of Iran
The list is created based on data from Paknia et al. (2008) and Janicki et al. (2016), while the species-group divisions follows Agosti (1990) and Radchenko and Paknia (2010).

Cataglyphis pallida species group
Cataglyphis emeryi (Karavaiev, 1911) First record from Iran: Radchenko (1998 Diagnosis. Bicolored body combined with thick, dense, black, decumbent setae on femora and tibiae resemble characters typical for species of the Cataglyphis setipes complex. Cataglyphis bazoftensis distinctly differs from them in the cuneiform petiolar node, which is distinctly nodiform among members of the setipes complex. At first glance, large species of the Cataglyphis bucharica complex, with cuneiform petiole, resemble C. bazoftensis, but most of these species differ in femora and tibiae lacking thick, black, decumbent setae. Members of the bucharica complex with setose femora and tibiae differ in the presence of black, erect setae on the posterior part of the head. Cataglyphis kurdistanica Pisarski is the most similar to C. bazoftensis, but differs in the presence of the major soldier caste, and posterior part of head and propodeum covered with long, black and erect setae. Cataglyphis altisquamis (André) and C. foreli (Ruzsky) differ from C. bazoftensis in major workers with reddish brown to brownish black head Head, mesosoma, and petiolar node red in the largest major workers; the percentage of brown in body coloration increases in smaller major workers with the smallest major workers reddish brown. Legs most often black or black with brownish black coxa; in the largest major workers coxa and trochanters mostly reddish brown to brown with darker brown spots of diffused borders; in the palest specimens femora brownish black with reddish iridescence, tibiae sometimes reddish brown apically, and tarsi reddish brown to brown. Antennal scapus red to red-brown; only in the darkest major workers brown; funicles darker than scapus, from brown to almost black, only in the palest specimens red-brown (Fig. 3). Head. Square; approximately 1.05× as long as wide; sides below eyes almost parallel, above eyes gently convex; posterior margin of head almost straight (Fig. 3). Anterior clypeal margin convex; without central impression; clypeal plate with three or four long black central setae and a pair of moderately long basal setae; anterior clypeal margin with a row of short black setae, and 8-10 long black setae, the longest as long as 0.75 length of clypeus; some workers have long setae often worn off or broken. Clypeus shiny and densely microreticulate; covered with very sparse and short, appressed pubescence. Eyes large and oval; approximately 1.3-1.4× as long as wide. Frontal carinae short; not extending beyond frontal lobes; interocular area with thin and shiny line and a pair of long black setae. Antennal fossa shallow; opalescent and densely microreticulated. Head shiny and densely microreticulate; mostly without appressed pubescence; only antennal fossa, posterior margin of head, and gular parts covered with sparse, short, adpressed hairs. Ocellar region with 2-4 moderately long and black setae; posterior angles without black setae; rest of frontal and lateral faces of head without erect setae; ventral side with a dozen yellowish to brown setae; interocular and ocular setae often broken. Antennal scape long; in frontal view almost straight, 1.3× as long as width of the head; base without tooth; apex slightly and gradually widening; funiculus long; pedicel elongated, approximately 0.82× as long as segments II and III combined, 1.6× as long as segment II (Fig. 3). Surface of scape densely microsculptured; distinctly to moderately shiny; covered with thick, moderately dense and decumbent setae. Mandibles rounded; basally smooth and shiny; apical ¾ length with deep grooves; surface shiny with several long yellow setae; masticatory margin with four large teeth. Mesosoma. Long; 2.4× as long as wide; metanotal groove shallow (Fig. 2). Pronotum convex on sides (Fig. 1). In lateral view promesonotum slightly arched in profile; propodeum positioned lower than promesonotum, moderately convex in lateral view (Fig. 2). Mesosoma shiny and densely microreticulated; covered with sparse, short, appressed pubescence; lateral sides of pronotum and mesonotum almost hairless; anterior part of pronotum, posterior angles of both mesonotum and propodeum with indistinct vestiture. Pronotum and mesonotum without erect setae; propodeum without or with one or two short, black, erect setae. Petiole. Cuneiform; in lateral view almost triangular with very short peduncle. Anterior face in front of spiracle distinctly convex; posterior face almost flat; top of petiole in lateral view obtusely angulate, lacking erect setae; sometimes frontal face apically with a single short seta. In anterior or posterior view petiolar dorsum emarginated. Surface of petiole distinctly microreticulated; shiny to slightly opalescent. Gaster. Dull and distinctly microreticulate. Whole surface of gaster with indistinct, sparse, appressed pubescence; tergites I and II without erect setae; tergite III with 2-4 long, black, central setae placed close to anterior margin, and none or one seta at lateral margin; tergite IV in with 6-8 long black setae; setosity in older specimens usually broken (Fig. 2). Each of gastral sternites with three or four long, black, erect setae. Legs. Dorsal and lateral surfaces of femora and tibiae covered with thick, dense, black, decumbent setae; interspaces between setae hairless. Ventral surfaces of femora and tibiae with numerous long, black, suberect to erect, spiniform setae.
Minor worker ( Color. Head and mesosoma uniformly brown to black or brown with diffused redbrown parts; gaster, petiole, femora, and tibiae brown; trochanters and tarsi yellowbrown. Younger specimens often paler than older specimens, with large areas of body yellowish brown. Antennae bright brown, only in younger specimens mostly yellowish to yellowish brown (Figs 7,8). Head. More elongated than in major workers; 1.2× as long as wide; below eyes parallel sided, behind eyes regularly rounded; posterior margin of head convex. Anterior clypeal margin convex with shallow impression in central part. Eyes large and oval; 1.3-1.4× as long as wide. Sculpture and setation of head and legs same as in major worker. Mesosoma. Same as in major worker. Petiole. More conical than cuneiform; dorsum more rounded, anterior face slightly convex (Fig. 5). Gaster. Strongly microreticulated and dull. Tergites I and II without erect setae; tergite III with only a pair of black setae centrally; tergite IV with two long, and two short black setae; in older specimens, setae usually partially broken (Fig. 8). Each of gastral sternites with 2-4 long, black, erect setae.
Biology. Little known, nests were found under stones inside a deciduous, oak forest surrounded by a grazing area. All collecting sites were in alpine zone, from 1738 to 1886 m a.s.l.
Etymology. The species name bazoftensis is a feminine Latin adjective in the nominative case and refers to the Bazoft region of the Koohrang County, the type locality for this species.

Diagnosis.
Cataglyphis fritillariae belongs to a group of large species with welldeveloped and dull body sculpture. Femora and tibiae covered with thick, dense, black, decumbent setae cluster this species with the Cataglyphis setipes complex, while the shape of the petiolar node groups it with the C. altisquamis complex. At first glance, C. foreli appears similar to C. fritillariae, but it differs by the trapezoidal profile of petiole, larger eyes, and lack of thick, black, decumbent setae on femora and tibia. Cataglyphis kurdistanica Pisarski has similar petiole and black decumbent setae on legs, but it can be separated by the bicolored body and the presence of a soldier caste. Cataglyphis bucharica also appears similar to C. fritillariae, but it is readily recognized by reddish head and mesosoma, presence of numerous erect setae on propodeum, longer propodeal spiracle, and absence of thick, black, decumbent setae on femora. Cataglyphis asiriensis Collingwood, known from the Asir Mountains (Kingdom of Saudi Arabia), has a similar petiole shape and the legs covered with black decumbent setae but it differs from C. fritillariae by the presence of long, black, erect setae also present on the dorsal side of femora and tibiae, and more numerous black erect setae on the mesosoma. Cataglyphis dejdaranensis sp. nov. is the most similar to C. fritillariae, but it differs by the weakly sculptured gaster that has moderately shiny sides of gastral tergite I, petiolar node of major workers knob-shaped in profile, and more convex propodeum.  (Figs 9-11). Head. Square; approximately 1.13× as long as wide; sides below eyes almost parallel, above eyes gently convex, posterior margin almost straight (Fig. 11). Anterior clypeal margin convex; without central impression; with a row of short black setae, and eight additional long black setae, the longest approximately as long as 0.6 length of clypeus; clypeal plate with a pair of long and black setae close to basal margin, and two pairs of similar setae close to anterior margin; sometimes clypeal plate with 1-3 additional short setae. Clypeus opalescent and densely microreticulated; covered with very sparse, short and adpressed hairs. Eyes large and oval, 1.3-1.4× as long as wide. Frontal carinae short; not extending beyond frontal lobes; interocular area with thin shiny line and 1-3 long black setae placed along each side of the line. Antennal fossa shallow, opalescent, densely microreticulated. Head opalescent and densely microreticulated; covered with sparse, short, adpressed hairs (Fig. 11). Ocellar region with three or four moderately long and black setae and often additional two or three shorter black setae; a transverse row of 4-8 black setae present above ocelli; area behind eyes with three or four yellowish to brown short setae; rest of frontal and lateral faces of head without erect setae; ventral side of the head with a dozen white to brown setae. Antennal scape long; in frontal view straight; 1.3× as long as width of the head; from base to apex slightly and gradually widened; its base without tooth; funiculus long; pedicel elongated, approximately 0.96× as long as segments II+III combined and 1.9× as long as segment II (Fig. 11). Surface of scape densely microsculptured and opalescent; covered with thick, dense, decumbent setae. Mandibles rounded; basally smooth and shiny; apical ¾ length with deep grooves; surface shiny with several long, white to brown setae; masticatory margin with four large teeth. Mesosoma. Long; 2.4× as long as wide; metanotal groove shallow (Fig. 10). Pronotum convex on sides. In lateral view promesonotum slightly arched in profile; propodeum positioned lower than promesonotum, moderately convex in lateral view (Fig. 10). Mesosoma opalescent and densely microreticulated; covered with sparse, short and adpressed hairs. Pronotum with 2-5 moderately long and black setae medially, sometimes with one or two short black setae close to its anterior and posterior margins; mesonotum with one or two moderately long and black setae in front of spiracles, sometimes with one or two short, black setae medially and anteriorly; propodeum dorsally with 4-7 long and black setae, and often two or three additional short setae; older specimens sometimes with setae completely to partly broken (Fig. 10). Petiole. Trapezoidal in profile; its anterior face mostly flat with only basal part convex; posterior face straight to slightly concave; dorsum flat; peduncle very short. Surface opalescent and microreticulated; covered with sparse, short and adpressed hairs; top of knob with 3-6 moderately long, black, erect setae (Fig. 12). In anterior view petiolar dorsum with distinct emargination medially. Gaster. Dull and distinctly microreticulated. Whole surface of gaster with short, sparse, adpressed hairs; tergite I in anterior part with 2-4 long, black setae; tergite II without a pair of black setae anteriorly; tergite III with 2-4 long and black setae centrally; in older specimens, setae usually broken (Fig. 16). Each of gastral sternites with three or four long, black, erect setae. Legs. Dorsal and lateral surfaces of femora and tibiae covered with thick, dense, black, decumbent setae; no white, adpressed setae on surface between black setae. Ventral surfaces of femora and tibiae with numerous long, black, suberect to erect, spiniform setae (Fig. 14).
Minor worker ( Color. Uniformly yellowish brown to brownish black. Antennae and legs yellowish brown to bright brown (Figs 15, 16). Head. Almost square; 1.15-1.17× as long as wide; sides below eyes almost parallel, behind eyes regularly convex, posterior margin of head convex. Sculpture and setation of the head similar as in major worker but with lower number of long setae. Mesosoma. Long; 2.4-2.5× as long as wide; metanotal groove shallow. Pronotum convex on sides. In lateral view promesonotum slightly arched in profile; propodeum positioned lower than promesonotum, its dorsum and posterior side slightly convex (Fig. 16). Whole mesosoma opalescent and densely microreticulated (Figs 15, 16). Whole mesosoma covered with dense, short, adpressed hair; pronotum with additional two or three black and erect setae; mesonotum and propodeum with additional one or two black and erect setae. Petiole. In form of thick scale; its anterior surface slightly convex; apex rounded and posterior surface almost flat; surface microreticulated and covered by sparse, short, adpressed hairs; dorsum with 1-3 black, moderately elongated and erect setae (Fig. 13). Gaster. Dull and distinctly microreticulated; tergites I and II with up to two black and erect setae; tergite III without or with a pair of black and erect setae close to its anterior margin; tergite IV with 2-4 erect setae; sternites with two or three black and erect setae; whole surface of gaster with short, adpressed hairs (Figs 15, 16). Legs. Dorsal and lateral surfaces of femora and tibiae covered with thick, dense, black, decumbent setae; no white, adpressed setae on surface between black setae. Ventral surfaces of femora and tibiae with numerous long, black, suberect to erect, spiniform setae.
Biology. Little known, in Dashte laleh a nest was found under rocks in a grazing area (Fig. 25). The site was located on a small plateau (3600 hectares), between 2100-2600 m above sea level, and in May was predominantly overgrown by Snake's head (Fritillaria imperialis L.). Other common plants recorded from this locality were Milkvetch (Astragalus spp.), Persian shallot (Allium stipitatum Regel), and kheshk (Daphne mucronata Royle). The species appears to be alpine, as all its collecting sites were placed on high altitude from 2143 to 2400 m.
Etymology. The species name fritillariae is named after the genitive singular case of the generic name of the Snake's head Fritillaria imperialis L., the dominant flower in Dashte laleh, the type locality of this ant species. Diagnosis. Cataglyphis dejdaranensis belongs to the group of large species with well-developed and dull body sculpture. Femora and tibiae covered with thick, dense, black, decumbent setae cluster this species with the Cataglyphis setipes complex, and the shape of the petiolar node groups it with the C. altisquamis complex. From all species of the C. setipes complex, C. dejdaranensis differs in a knob-shaped petiole, which is not forming a spherical node; from all species of the C. altisquamis complex, C. dejdaranensis differs in a weak microsculpture of gaster of which at least sides are visibly shiny. Cataglyphis dejdaranensis has the least sculpted gaster within all large species with well-developed body microsculpture. At first glance, C. foreli appears similar to C. dejdaranensis, but it differs in larger eyes, and femora and tibia lacking thick, black, decumbent setae. Cataglyphis kurdistanica Pisarski has similarly shaped petiole and legs with black decumbent setae, but it differs in bicolored body and presence of the soldier caste. Cataglyphis bucharica also appears similar to C. dejdaranensis but it differs in reddish head and mesosoma, numerous erect setae on propodeum, longer propodeal spiracle, and femora and tibia lacking thick, black, decumbent setae. Cataglyphis fritillariae sp. nov. is the most similar to C. dejdaranensis, but differs in strongly sculptured gaster, petiolar node of major workers trapezoidal in profile, and less convex propodeum. Head, mesosoma and gaster completely black or head anteriorly, and pronotum and mesosoma on lateral sides with indistinct brownish black spots of diffused borders. Legs completely black or black-brown with tarsi sometimes slightly paler than femora and tibiae. Antennae completely black (Figs 17-19, 23, 24). Head. Square; approximately 1.12× as long as wide; sides below eyes almost parallel, above eyes gently convex, posterior margin almost straight (Fig. 19). Anterior margin of the clypeus convex; without central impression; with a row of short black setae, and eight longer black setae, the longest as long as 0.6 length of clypeus; clypeal plate with a pair of long and black setae centrally and a pair of similar setae basally. Clypeus opalescent and densely microreticulated; covered with sparse, short and adpressed hairs. Eyes large and oval; approximately 1.2× as long as wide. Frontal carinae short, not extending beyond frontal lobes; interocular area with thin shiny line and two or three long black setae along each of its sides. Antennal fossa shallow, opalescent, densely microreticulated. Head opalescent and densely microreticulated; covered with sparse, short, adpressed hairs (Fig. 19). Ocellar region with group of 2-4 moderately long black setae; posterior angles with two or three long and 1-3 short black setae; rest of frontal and lateral faces of head without erect setae; ventral side of the head with a dozen white to brown setae. Antennal scape long; in frontal view straight; 1.1× as long as width of the head; its base without tooth; from base to apex slightly and gradually widening; funiculus long; pedicel elongated, approximately 0.85× as long as segments II and III combined and 1.7× as long as segment II (Fig. 19). Surface of scape densely microsculptured, opalescent; covered with thick, dense, decumbent setae. Mandibles rounded; basally smooth and shiny; apical half with deep grooves; surface shiny with several long white setae; masticatory margin with four large teeth. Mesosoma. Long, 2.2× as long as wide; metanotal groove shallow. Pronotum convex on sides. In lateral view promesonotum slightly arched in profile, propodeum positioned lower than promesonotum; distinctly convex in lateral view (Figs 24,25). Mesosoma opalescent and densely microreticulated; covered with sparse, short and adpressed hairs. Pronotum posteriorly with 2-4 moderately long and black setae; and anteriorly with 2-4 short black setae; sometimes setae broken or missing; mesonotum with up to two moderately long and black setae close to the anterior margin and usually two black setae close to the median groove; propodeum apically with 1-4 short, black setae; sometimes mesonotal and propodeal setae broken. Petiole. In form of knob; its anterior face distinctly convex; posterior face only slightly convex and dorsum regularly rounded; peduncle short. Surface opalescent and densely microreticulated; covered with dense, short, adpressed hairs; dorsum of knob with 3-5 very short, black, erect setae. Gaster. Finely microreticulated and moderately shiny; gastral segment I and lateral sides of remaining segments appear distinctly shinier than remaining parts of gaster. Whole surface of gaster with short, sparse, adpressed hairs; tergites I and II without erect setae; tergite III with one or two long black setae centrally and close to anterior margin, and up to one setae at lateral margin; tergite IV in younger specimens with four long black setae on each side; in older specimens setae usually broken (Fig. 24). Each of gastral sternites with three or four long, black, erect setae. Legs. Dorsal and lateral surfaces of femora and tibiae covered with thick, dense, black decumbent setae; no white adpressed setae on surface between black setae. Ventral surfaces of femora and tibiae with numerous long, black, suberect to erect setae.  Color. Slightly paler than major workers; head, mesosoma and gaster mostly brown to dark brown; upper part of head, propodeum and gaster usually blackish brown but never black; transition from brown to blackish brown diffused. Antennae and legs brown to almost black, legs brown to dark brown (Figs 23, 24). Body. Morphological characters similar to these of major worker except petiole which appears as a very thick scale; approximately 2× as high as broad, with anterior surface only slightly convex and posterior surface flat, and with only rudiment of peduncle (Fig. 21). Dorsal surface of gastral tergites I and II without erect setae; tergite III without or with only a pair of erect setae; tergite IV with two or four setae. Sternites setose as in major worker. Legs generally similarly setose as in major workers except fewer suberect and erect black setae on ventral sides of femora and tibiae. Head slightly longer and scape slightly shorter than in major workers.

Cataglyphis dejdaranensis
Biology. Little known. On the locality Cheri, specimens were collected in a mountain grazing area overgrown by grass, and on the locality Dejdaran Valley ants were found on mountain pastures with scant vegetation (Fig. 26). Nests were located under large stones. Both collecting sites were placed on high altitude: 2319 m and 2778 m.
Etymology. The species name dejdaranensis is a feminine Latin adjective in the nominative case and refers to Dejdaran Valley, where one of the specimens of this species was collected.

Comments
All three new Iranian species belong to the group of large, polymorphic Cataglyphis. Based on their morphology, they should be assigned to the Cataglyphis altisquamis group, sensu Agosti (1990). Herein, we present a modified version of the key to the Asian Cataglyphis (Radchenko 1998). The key was modified to accommodate the three new species and C. asiriensis, which was not included in the original version. We also decided not to refer to figures available in Radchenko's paper but provide photographs of type specimens of species which are included in the key (except photographs of C. oxiana Arnol'di and C. piligera Arnol'di that were not available).
In the key to the Asian Cataglyphis altisquamis species group proposed by Radchenko (1998)  Gaster dull (Figs 9, 15). Petiole in major worker trapezoidal in profile (Fig. 12). Propodeal dorsum distinctly longer than its declivity in profile. Petiole high, narrow-cuneiform, with weakly convex anterior surface, about as high as propodeum. Head and mesosoma red to red-brown (posterior half of thorax sometimes dark brown); gaster dark brown to black. Mountains of Uzbekistan and Tajik