Bombus turkestanicus Skorikov

Williams, Paul H., Huang, Jiaxing, Rasmont, Pierre & An, Jiandong, 2016, Early-diverging bumblebees from across the roof of the world: the high-mountain subgenus Mendacibombus revised from species’ gene coalescents and morphology (Hymenoptera, Apidae), Zootaxa 4204 (1), pp. 1-72 : 45-47

8. Bombus turkestanicus Skorikov

( Figs 17, 23 , 31 , 45, 52 , 63 )

[ Bombus mendax Gerstaecker ; Morawitz 1875:4, misidentification.]

< Bombus mendax > subsp. turkestanicus Skorikov 1910b:329 , type-locality citation (Cyrillic) ‘[Turkestan (Samarkand, Fergana, and Semirechensk Province )]’. Lectotype queen by present designation ZISP examined, (Cyrillic) ‘[Voru, Samarkand]’ (Alai, Tajikistan). Note 1.

Mendacibombus mendax subsp. turkestanicus ( Skorikov); Skorikov 1914 :124.

Mendacibombus turkestanicus ( Skorikov); Skorikov 1923 :149; Skorikov, 1931:215.

Bombus turkestanicus Skorikov ; Reinig 1934:172.

Bombus (Mendacibombus) turkestanus Skorikov ; Panfilov 1957:237; Panfilov 1962:195; P.H. Williams 1991:15, 41; S.- F. Wang & Yao 1996:303; P.H. Williams 1998:99; P.H. Williams 2011:28.

Note 1 ( turkestanicus ). Skorikov’s original description of the taxon turkestanicus cites the type locality as Samarkand , Fergana , and Semirechensk Province . The ZISP collection studied by Skorikov contains a queen that agrees with the original description and carries the labels: (1) white, printed ( Cryrillic ) ‘[ Voru , / Samarkands. obl. / Golbek] 20 VI 09 ’; (2) white, printed (Cyrillic) ‘[k. Skorikova]’; (3) red, handwritten ‘ Paralectotypus Bomb. / mendax subsp. / turkestanicus Skor. / design. Podbolotsk. ’ (M. Podbolotskaya, unpublished); (4) green, printed ‘ Mendacibombus / MD# 3525 det. PHW’; (5) red, printed ‘ LECTOTYPE [female] / Bombus mendax ssp. / turkestanicus / Skorikov, 1910 / det. PH Williams 2012’; (6) white, printed ‘[female] Bombus / ( Mendacibombus ) / turkestanicus / det . PH Williams 2012’. This specimen, which is complete, is regarded as one of Skorikov’s syntypes and is designated here as the lectotype in order to reduce uncertainty in the identity and application of the name.

A second queen collected at Voru by Golbek in 1909 (MD#320, NHM, sent by Skorikov as part of an exchange with the NHM in 1934), closely similar in morphology, is designated here as a paralectotype and interpreted as conspecific.

Etymology. The species is named after Russian Turkestan, which from 1867‒1918 was a Governorate-General (Krai) of the Russian Empire in Central Asia.

Taxonomy and variation. The interpretation of this species is based here on DNA, as well as on the form of the female labrum and of the male genitalia.

Skorikov (1910b) described females of the taxon turkestanicus s. str. from the Alai mountains (MD#3525) as having the corbicula framed with partly light rusty and partly greyish hairs, with the pale bands light yellow, and with the entire side of the thorax and the ventral side of the metasoma (S1‒4) light yellow. For the COI group of specimens with this B. turkestanicus colour pattern ( Fig. 13 ), the form of the female labrum is diagnostic ( Fig. 17 ).

Most specimens of this species are yellow-banded. For two queens from near the Kyrgyz Ata (river), Kyrgyzstan, one has a light-yellow-banded colour pattern (MD#325) and the other a cream-white-banded colour pattern (MD#324). A queen from Kyrgyzstan is white-banded and groups with B. turkestanicus by its COI sequence ( Fig. 14 : the white-banded individual labeled ‘white’ MD#1281 and the yellow-banded taxon turkestanicus s. str. MD#324). This material is interpreted as conspecific, as parts of the species B. turkestanicus s. l..

Diagnostic description. Wings nearly clear. Female hair colour pattern: generally black, but with pale hair (yellow or cream-white) over most of the face (with variable amounts of black intermixed on the outer side of the face and above the antenna), in a usually small patch on the vertex of the head, in a transverse band anteriorly on the thoracic dorsum (occasionally with black hairs intermixed posteriorly) and extending laterally and ventrally all the way to the midleg base (cf. B. makarjini ), in a transverse band posteriorly on the thoracic dorsum (scutellum; so the thoracic dorsum between the wing bases has the hair entirely black in a narrow band), on T1‒2, although T2 with a few black hairs intermixed along the posterior margin, orange hair on T3 as a posterior fringe, and throughout on T4‒6, T6 without obvious black hairs, S1‒6 entirely pale (cf. B. makarjini ). Hindleg tibia with the corbicular fringes usually nearly entirely yellow or grey, the long hairs in the fringes sometimes orange close to the base and with a few short orange-brown bristles along their inner edges, only rarely with a few long dark hairs intermixed ( Fig. 23 ) (cf. B. makarjini , B. margreiteri , B. defector ). Female morphology: labrum with the basal depression narrow, the transverse ridge just broader medially than the basal depression, in the median third subsiding slightly but not clearly or abruptly interrupted, with many scattered punctures, the lateral tubercles with only a few punctures ( Fig. 17 ) (cf. B. margreiteri , B. defector ). Clypeus in its central half with many widely spaced small and large punctures (cf. B. margreiteri , B. defector ). Male morphology: genitalia ( Fig. 31 ) with the volsella distally rounded (finger-shaped) and curled back dorsally but not anteriorly; volsella at its broadest near the midpoint of its length, the dorsal surface just distal to this point without a raised curved ridge just inside the inner margin; volsella with the apex narrowed, narrower than the adjacent penis-valve head. Penis-valve inner shoulder located at Ĺ 0.5× the length of the penis valve from the distal end to the broadest point of the spatha; penis valve proximal to the outer shoulder <2× as broad as the penis-valve head; penis-valve breadth just proximal to the penis-valve head 0.11× the length of the penis valve distal to the broadest point of the spatha.

Material examined. 21 queens 87 workers 27 males, from Afghanistan, China, Kazakhstan, Kyrgyzstan, and Tajikistan ( Fig. 63 : AMNH, IAR, IZB, KUK, NHM, NME, OLL, PW, RMNH, SMNS, UMONS, ZISP), with 26 specimens sequenced (interpretable sequences listed in Figs. 11–13 ).

Habitat and distribution. Flower-rich alpine and subalpine grassland, at elevations 950‒(2146)‒ 4350 m a.s.l.. A species of the Tian Shan, Alai, Pamir, and Hindu Kush mountains. Compared to B. defector , the distribution of B. turkestanicus extends less far to the east, overlaps broadly in the mid part of the range (and the two species often occur together), but extends further to the south in the Pamir and Hindu Kush, where it reaches higher elevations. Bombus turkestanicus overlaps with B. margreiteri in the Tian Shan, although the two seldom occur together. There is some overlap of B. turkestanicus with B. marussinus in the Pamir and Hindu Kush, although the two seldom occur together.

Food plants. Williams (2011). Behaviour. No records.

Figures

FIGURE 13. Assigning names to species from an estimate of phylogeny for 130 COI sequences of the subgenus Mendacibombus. Based on a BEAST analysis of the 30 unique haplotypes (Fig. 11), with the remaining sequences interpolated back into the tree with 0 branch lengths from their matching samples (where a sequence matches more than one longer sequence within a species in the Collapse results, it is added after the first matching sequence, with the addition of the two short-sequence yellow unbanded specimens from Pakistan, MD # 4058, 4059). The sequence labels follow the format used in Figs 11, 12. Lineages accepted as species from the GMYC models (Fig. 11) are shown as thick branches, the most recent common ancestors of each species are shown with a black spot, and branches within species are shown as thin branches. Asterisks mark samples used as informal proxies for the type specimens of each of the taxon names (Table 1). The proxy sample for the type specimen for the oldest available name (the valid name) for each species (from Fig. 11) is marked in bold. The interpretations of the species with their valid names are shown in grey to the right.

FIGURE 14. Assessing morphological characters by assessing groups of species within the subgenus Mendacibombus from three characters of hair colour (characters 1 ‒ 3) and 13 characters of male genital morphology (characters 4 ‒ 16) by maximum parsimony analysis with TNT. Numbers above the branches of the tree are the character numbers from Table 6; solid squares show unique character-state changes; and open squares show parallel or reverse character-state changes.

FIGURES 15 ‒ 23. Morphology of the female (queen) head from the ventral aspect (labrum and mandibles): 15, B. marussinus queen, Pakistan MD # 482; 16, B. avinoviellus queen, India MD # 598; 17, B. turkestanicus queen, Tajikistan MD # 320; 18, B. defector queen, Kyrgyzstan MD # 1080; 19, B. margreiteri queen, Mongolia MD # 319; 20, B. himalayanus queen, India MD # 453. Morphology of the female (queen) left hindleg tibia from the left lateral aspect, dorsal at the top of the image, anterior to the left: 21, B. defector, China MD # 1251; 22, B. margreiteri, Mongolia MD # 319; 23, B. turkestanicus, Kyrgyzstan MD # 324.

FIGURES 24 ‒ 35. Morphology of the male genitalia for species of the subgenus Mendacibombus from the dorsal aspect, anterior to the left of the image, posterior to the right: 24, B. superbus, China MD # 339; 25, B. waltoni, China MD # 216; 26, B. convexus, China MD # 215; 27, B. makarjini, Kazakhstan MD # 1240; 28, B. marussinus, Pakistan MD # 408; 29, B. avinoviellus, India MD # 404; 30, B. himalayanus, India MD # 426; 31, B. turkestanicus, China MD # 351; 32, B. defector, Kyrgyzstan MD # 367; 33, B. margreiteri, China MD # 294; 34, B. handlirschianus, Russia MD # 835; 35, B. mendax, Spain MD # 689.

FIGURES 36 ‒ 55. Morphology of parts of the male genitalia from the dorsal aspect of the left side, anterior to the left of the image, posterior to the right, white bars indicating morphological characters referred to in the key to males: 36, B. waltoni; 37, B. mendax; 38, B. superbus; 39, B. convexus; 40, B. waltoni; 41, B. convexus; 42, B. defector; 43, B. mendax; 44, B. mendax; 45, B. turkestanicus; 46, B. avinoviellus; 47, B. mendax; 48, B. mendax; 49, B. makarjini; 50, B. handlirschianus; 51, B. himalayanus; 52, B. turkestanicus; 53, B. himalayanus; 54, B. himalayanus; 55, B. marussinus.

FIGURES 56 ‒ 67. Maps of the global distributions of species of the subgenus Mendacibombus from the material examined, with grey spots for specimen records and black spots for the samples with COI sequences (each species is unknown from regions outside the map boundaries shown); grey crosses for records of all species of Mendacibombus combined. Relief map with hill shading, Cartesian orthonormal projection, grey lines show borders to national administration according to UN maps, north at the top of the map. Image created in ArcGIS using World _ Shaded _ Relief basemap © 2014 Esri.

FIGURES 11 ‒ 12. Recognising species from an estimate of phylogeny for 30 unique COI haplotypes of the subgenus Mendacibombus by BEAST, using GMYC models and the PTP procedure. From the GMYC models (11), the single threshold T is shown by the vertical grey bar so that the intersecting lineages are interpreted as indicating 12 separate species (black spots at coalescent nodes; outgroups not shown). Values below the nodes of the maximum clade-credibility tree are Bayesian posterior probabilities showing branch support (groups with values <0.8 are considered unreliable) from the BEAST analysis with two billion MCMC generations and a 1 % burn-in. The x - scale axis is calibrated in substitutions per nucleotide site. From the PTP procedure (12), the Bayesian PTP solution with highest support also indicates 12 species. Values above the nodes are PTP Bayesian support values that all daughter haplotypes are parts of a single species; values below the nodes are Bayesian posterior probabilities showing branch support. The scale bar is calibrated in substitutions per nucleotide site. Each unique haplotype is represented by one of the longest available sample sequences, labelled with: the sequence length in number of base pairs, a taxon name or name for an unpublished colour pattern (light / dark / yellow / white / pale), and then a code that consists of a specimen identifier (MD #) from the project database and (after the hyphen) a sample identifier from BOLD or from GenBank, followed with its geographic origin.