Bombus makarjini 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 : 34-40

4. Bombus makarjini Skorikov

( Figs 27 , 49 , 59 )

< Bombus mendax > subsp. makarjini Skorikov 1910b:329 , type-locality citation (Cyrillic) ‘[ East Pamir ]’. Lectotype queen by present designation ZISP examined, (Cyrillic) ‘[Pamir]’ (Pamir, Tajikistan). Note 1.

[ Bombus mendax macarjini Skorikov ; Reinig 1930:102; Reinig 1932b:163, incorrect subsequent spellings.]

[ Bombus mendax marcajini Skorikov ; Reinig in Bischoff 1931:862 footnote, incorrect subsequent spelling.]

[ Bombus (Mendacibombus) macarjini Skorikov ; Reinig 1932a:258, incorrect subsequent spelling.]

[ Bombus marcajini Skorikov ; Reinig 1934:172, incorrect subsequent spelling.]

[ Bombus (Mendacibombus) makaryini Skorikov ; Panfilov 1957:236, incorrect subsequent spelling.]

Bombus (Mendacibombus) makarjini Skorikov ; Panfilov 1962:195; P.H. Williams 1991:15,40; P.H. Williams 1998:99.

Note 1 ( makarjini ). Skorikov’s original description of the taxon makarjini cites the type locality as east Pamir. The ZISP collection studied by Skorikov contains a queen that agrees closely with the original description and carries the labels: (1) white, printed ( Cryrillic ) ‘[[pass] Koytye-zyek , / [Pamir range] 2600 mt. / Makarin] 19VI09. ’; (2) white, printed (Cyrillic) ‘[k. Skorikova]’; (3) red, handwritten ‘ Lectotypus Bombus / mendax subsp. / makarjini Skor. / design. Podbolotsk. ’ (M. Podbolotskaya, unpublished); (4) green, printed ‘ Mendacibombus / MD# 3520 det. PHW’; (5) red, printed ‘ LECTOTYPE [female] / Bombus mendax ssp. / makarjini / Skorikov, 1910 / det. PH Williams 2012’; (6) white, printed ‘[female] Bombus / ( Mendacibombus ) / turkestanicus / det . PH Williams 2012’. This specimen, which lacks the last four joints of the left hindleg tarsus, 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.

Another queen was examined on loan from the ZISP collection in 1984 (MD# 737), with labels: (1) white, handwritten (Cyrillic) ‘[[Pass] Mas 4000 m / Pamir 30.VI. 0 9 / Makarin]’; (2) white, printed (Cyrillic) ‘[k.

Skorikova.] ’; (3) red, part handwritten ‘ Paralectotypus Bombus / mendax subsp. / makarjini Sko / design. Podbolotsk ’ (M. Podbolotskaya, unpublished). This specimen is interpreted as a syntype of the taxon makarjini and is designated here as a paralectotype of the taxon makarjini . This specimen resembles B. defector in the colour pattern of the hair, although B. defector is not known to occur in the Pamir. The specimen was unavailable for its morphology to be checked in this study.

Etymology. The species is named after A.G. Makarjin, who collected the lectotype specimen in the Pamir in 1909 (see also the type data for B. defector , B. marussinus ). He collaborated with A.G. Jakobson, the brother of the G.G. Jakobson (A. Byvaltsev, in litt.) who collected other Mendacibombus types in the Himalaya (see below on B. himalayanus , B. avinoviellus ).

Taxonomy and variation. The interpretation of this species is based here on DNA, on the form of the male genitalia, and on colour pattern. It is the least well characterized species in the subgenus because of a lack of material, especially of recent material that can be sequenced. Our concept disagrees with earlier concepts diagnosed originally in terms of the colour pattern of the hair (Skorikov, 19 10b). The species appears to be more variable and less easily diagnosed by colour pattern (cf. B. turkestanicus , B. margreiteri ) than was originally understood.

Skorikov (1910b) described queens from the Pamir ( Tajikistan) with a colour pattern (corbicula framed with yellow, grey, or light rusty hairs) and with the ventral side of the metasoma (S1‒4) with black hair. The lectotype queen (MD#3520) has the pale bands greyish and scarcely cream. A queen with a similar greyish colour pattern, pale corbicular fringes, and black hairs on S2‒3 from Uzbekistan (but labelled ‘Tadzikistan’, MD#330) and another from Kazakhstan (but labelled ‘Kirgyzstan’, MD#4101) yielded COI sequences that differ from one another in just one base pair and which are strongly divergent from the other species ( Figs 11, 12 ). Queens from southern Xinjiang (MD#3563, 4108) failed to sequence. Another queen (MD#3562) from the same region gave a sequence of just 96 bp, although this part of the sequence does not include any known diagnostic positions for this species ( Table 5). Females show varying degrees of replacement of yellow hair with black on the lower side of the thorax near the midleg base, and on S2‒3. All of these specimens are currently indistinguishable from one another in any substantial morphological characters. Associated by us with these females are three males (MD#1240, 3519, 4403) with distinctive genitalia ( Fig. 27 ), one of which (MD#3519) has a COI sequence closely matching the COI sequences from females (MD#330, 4101). These males are broadly similar in the form of the genitalia to the genitalia ascribed to this species by Skorikov (1931: his fig. 18). Skorikov’s drawing differs in showing the gonostylus as slightly broader and with the sides slightly diverging in the proximal half (more like B. margreiteri ). However, Skorikov’s illustration also shows: that (1) the spatha is broad with sinusoidal rather than straight converging sides; and that (2) the penis valve has a pronounced inner shoulder. Therefore his illustration is likely to be of B. makarjini as understood here.

Diagnostic description. Wings nearly clear. Female hair colour pattern: generally black, but with pale hair (grey, cream-white, or yellow) over most of the face or at least in a large patch below the base of the antenna with variable amounts of black intermixed or covering the outer side of the face and above the antenna, in a small patch on the vertex of the head, in a transverse band anteriorly on the thoracic dorsum and extending laterally and ventrally from at least one third to two thirds of the way down the side of the thorax towards the midleg base (cf. B. turkestanicus ), in a transverse band posteriorly on the thoracic dorsum (scutellum; so the thoracic dorsum between the wing bases has the hair entirely black), 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 many obvious black hairs, with black hairs on S2‒3 or rarely S1‒4 (cf. B. turkestanicus ). Hindleg tibia with the corbicular fringes extensively yellow or grey (the hairs often orange in their basal half), the fringes with a few hairs black intermixed (cf. B. turkestanicus , 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 medium punctures, the lateral tubercles with only a few punctures (cf. B. margreiteri , B. defector ). Clypeus in its central half with very few punctures but some of them large (cf. B. margreiteri , B. defector ). Male morphology: genitalia ( Fig. 27 ) 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. Gonostylus brown ( Fig. 49 ), distally as thick as 0.25× its own breadth, the inner distal corner distinctly acute from both the inner and the dorsal aspects, in dorsal aspect long and narrow, 2× as long as its greatest breadth, the inner and outer lateral margins nearly parallel in the anterior proximal half. 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.

FIGURES 56‒67 . (Continued) FIGURES 56‒67 . (Continued) FIGURES 56‒67 . (Continued) Material examined. 8 queens 4 workers 3 males, from Afghanistan, China, Kazakhstan, Kyrgyzstan, Tajikistan, Uzbekistan ( Fig. 59 : IZB, OLL, RMNH, UMONS, ZISP), with 3 specimens sequenced (interpretable sequences listed in Figs. 11–13 ).

Habitat and distribution. Flower-rich alpine and subalpine grassland (not seen by us), at elevations 1550‒(3144)‒ 4500 m a.s.l.. A species of the Tian Shan and Pamir mountains (it is likely to occur in the Alai as well). There is overlap with B. marussinus in the Pamir, there is some overlap with B. turkestanicus in the Pamir and in the Tian Shan, and there is some overlap with B. defector and B. margreiteri in the Tian Shan, where the species may occur together.

Food plants. No records.

Behaviour. See Reinig (1930) and Reinig in Bischoff (1931), under the name B. mendax macarjini ; Panfilov 1962.

Figures

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.