Bombus separandus Vogt, 1909

Williams, Paul H., Altanchimeg, Dorjsuren, Byvaltsev, Alexandr, Jonghe, Roland De, Jaffar, Saleem, Japoshvili, George, Kahono, Sih, Liang, Huan, Mei, Maurizio, Monfared, Alireza, Nidup, Tshering, Raina, Rifat, Ren, Zongxin, Thanoosing, Chawatat, Zhao, Yanhui & Orr, Michael C., 2020, Widespread polytypic species or complexes of local species? Revising bumblebees of the subgenus Melanobombus world-wide (Hymenoptera, Apidae, Bombus), European Journal of Taxonomy 719, pp. 1-120 : 93-96

publication ID

https://doi.org/10.5852/ejt.2020.719.1107

publication LSID

lsid:zoobank.org:pub:A4500016-C219-4353-B81C-5E0BB520547F

DOI

https://doi.org/10.5281/zenodo.4335562

persistent identifier

https://treatment.plazi.org/id/252087CA-1F1E-9566-FDA0-FE2ED89AF80E

treatment provided by

Valdenar

scientific name

Bombus separandus Vogt, 1909
status

stat. rev.

Bombus separandus Vogt, 1909 stat. rev.

Figs 16 View Figs 14‒16 , 173–180 View Figs 139–180 , 204 View Figs 199‒204 , 210 View Figs 209–210 , 212 View Figs 211–212

Bombus lapidarius Form [subsp.] separandus Vogt, 1909: 61.

Bombus lapidarius Form [subsp.] kohli Vogt, 1909: 61 (non Cockerell, 1906: 75 = B. morio (Swederus)). syn. nov.

Bombus kozlovi Skorikov, 1910a: 413, replacement name for kohli Vogt, 1909: 61. syn. nov.

Bombus lapidarius var. [subsp.] tenellus Friese, 1913: 86. syn. nov.

Bombus tenellus var. [not subsp.] alpivagus Richards, 1930: 639, but not infrasubspecific after Reinig 1935: 333 ( ICZN 1999: Art. 45.6.4.1). syn. nov.

Lapidariobombus alagesianus subsp. pamirus Skorikov, 1931: 226 (non Skorikov, 1931: 232 = B. oberti Morawitz, not B. difficillimus Skorikov as given by Skorikov). syn. nov.

Lapidariobombus alagesianus subsp. mongolicus Skorikov, 1931: 226 (non Friese, 1916: 110?= B. diversus Smith). syn. nov.

Bombus keriensis f.g. [subsp.] bucharicus Reinig, 1935: 340. syn. nov.

Bombus keriensis f.g. [subsp.] richardsi Reinig, 1935: 341 (non Frison, 1930: 6 = B. rufipes Lepeletier).

Bombus tenellus subsp. tibetensis S.-F. Wang, 1982: 439, replacement name for richardsi Reinig, 1935: 341. syn. nov.

Bombus kozlowi – Bischoff, 1936: 9, unjustified emendation.

Bombus keriensis (part) – Reinig 1935: 341. — Williams 1998: 134 (non Morawitz, 1887: 199).

Vogt (1909: 46) stated that his “Formen” are subpopulations, which can be considered equivalent to subspecies. Reinig (1935: 333) stated explicitly that his “f.g.” (forma geographica) are equivalent to subspecies. Vogt (1909: 61) described his taxon separandus as having the hair of the face and top of the head (“Gesicht und Stirn”) black and the underside of the thorax dark haired. He described his taxon kohli (later renamed kozlovi) as having the face with a few yellow hairs. However, Vogt (1911: 58) then characterised the taxon separandus as differing from the taxon keriensis only in having an absence of pale hairs on the face (he did not include the taxon kohli / kozlovi as part of the taxon separandus but did include the taxon incertoides as part of separandus). The taxon separandus was treated as a part of the species B. keriensis s. lat. by Reinig (1935) and by Williams (1998).

Our PTP analysis ( Fig. 10 View Fig ) of coalescents in the COI gene within the keriensis -complex supports six species including B. separandus, corroborated by differences in morphology. These species are also supported by the absence of a positive divergence-with-distance relationship among them ( Fig. 20 View Figs 17–20 ) (see Divergence and geographical distance, page 12).

From morphology, after examining>1000 specimens of the keriensis -group from across Asia and after comparing the distribution of states of several characters as part of this study (PW), Vogt’s character state of exclusively black hair on the face does appear to remain one of the most consistently diagnostic characters (in comparison with COI sequences) for B. separandus in the QTP and Central Asia, at least for queens (but see the comments on B. keriensis). Nonetheless, there are some individuals of the yellow-banded taxon kozlovi from Mongolia (part of B. separandus s. lat. according our COI-coalescent results, Fig. 10 View Fig ) that do have a few yellow hairs on the face, so that exclusively black hair on the face is not diagnostic in all parts of the range. Several aspects of the colour pattern vary strongly within the species: (1) B. separandus includes both yellow-banded and white-banded colour patterns; (2) the extent of the black hair between the wing bases varies; (3) the extent of the pale hair on the side and ventral area (leg bases) of the thorax varies; (4) the extent of the pale hair posteriorly on T3 varies (the ‘ciliation’ of authors); (5) the contrast of the pale posterior fringes on T4–5 varies. Many of these characters might appear to show a continuum of variation that is also continuous with the variation within B. keriensis s. str. ( Williams 1991), which might then appear consistent with the two taxa being parts of the same species ( Reinig 1935). However, as noted above, evidence from COI barcodes shows that the two genecoalescent groups differ most consistently in the absence of pale hairs on the face for B. separandus, supporting Vogt’s characterisation of the species, providing integrated morphological support for its status as a separate species. The overlapping colour variation of B. separandus but only in Mongolia (where B. keriensis s. str. does not occur) does not negate the fact that B. separandus is recognisable by morphological characters throughout its geographical range.

There are some queens of the yellow-banded taxon richardsi (later renamed tibetensis) from the northwestern QTP that have no pale hairs on the face, very few or no yellow hairs on the leg bases, and very few black hairs within the yellow bands of the thoracic dorsum. Many of the available specimens have failed to sequence, so the species’ diagnosis remains tentative. Particularly influential in the interpretation of these individuals is one yellow-banded queen with a reduced extent of pale hair on the side of the thorax from the Zanskar range (PW: ML311, Figs 177 View Figs 139–180 , 210 View Figs 209–210 ), which has produced a short sequence (382 bp) that nonetheless shows bases diagnostic for B. separandus compared to B. keriensis (base positions 267A, 286C, 301C, 306T, 339T, 363C, 423T, 498C, 529G, 540T, 541C, 556C). This is one of the specimens that most closely resemble some yellow-banded individuals of the candidate species keriensis s. str. also from Kashmir ( Figs 170 View Figs 139–180 , 209 View Figs 209–210 ) (the difference in the breadth of the black band between the wings is not diagnostic for these species).

Bombus separandus co-occurs locally with the similar B. incertoides in the Altai and Khangai mountains of Mongolia. Bombus separandus also co-occurs locally with the similar B. keriensis s. str. in the Zanskar mountains of Kashmir. Examination of specimens in the IOZ collection labelled B. incertus (S.-F. Wang 1985; S.-F. Wang & Yao 1996; Niu et al. 2018) shows that these are mostly misidentified B. separandus (with a few B. keriensis) from Xinjiang.

Diagnosis

Females

Queens medium-sized body length 17–20 mm, workers 9–14 mm. Can be distinguished in Central Asia and Mongolia by the combination of hair of the face usually without pale hairs (except in Mongolia where B. keriensis does not occur), the leg bases usually without pale hairs, T2 posteriorly entirely pale without black hairs, and T3 laterally usually without pale hairs (cf. B. sichelii, B. keriensis).

Males

Body length 10–15 mm. Can be distinguished reliably at present only by their COI sequence, although in the western Himalaya they may be distinguished by entirely black hair between the wing bases and hair of T3 entirely black. Genitalia ( Fig. 204 View Figs 199‒204 ) with the gonostylus shorter than broad, its inner basal projection reduced to a short stub (cf. rufipes- group, festivus- group, rufofasciatus -group); volsella with the inner distal corner broadly produced but without a narrow hook (cf. rufipes- group, festivus- group, rufofasciatus -group); eye unenlarged relative to female eye.

Material examined

Lectotype

CHINA • ♀ (queen), lectotype of Bombus lapidarius Form separandus Vogt, 1909 by designation of Williams (1991: 97); Xinjiang, Borohoro Shan, Tisilikau; RMNH (examined PW).

Material sequenced (25 specimens)

MONGOLIA • 1 ♀ (queen); Bulgan; 47.4257° N, 103.6940° E; 22 Jun. 2013; NHMUK seq: NHMSR1; PW: ML147 GoogleMaps 1 ♀ (worker); Bayanolgiy , Bulgan; 46.9333° N, 91.1667° E; 26 Jul. 1992; Culverwell leg.; BOLD seq: 6877B11; PW: ML151 GoogleMaps 1 ♀ (queen); Khenteyn Nuruu, Terelj ; 47.9905° N, 107.3719° E; 16 Jun. 2015; R. DeJonghe leg.; BOLD seq: 6877A08; RJ: ML333 GoogleMaps 1 ♀ (queen); Khenteyn Nuruu, Terelj ; 47.9956° N, 107.3303° E; 17 Jun. 2015; R. DeJonghe leg.; BOLD seq: 6880B05; RDJ: ML489 GoogleMaps 1 ♀ (worker); Arkhangai, Chuluut Khujirt Mts ; 47.5750° N, 101.1092° E; 11 Aug. 2018; R. DeJonghe leg.; BOLD seq: 6880B12; RDJ: ML497 GoogleMaps .

RUSSIA • 1 ♀ (worker); Altai, Ulagansky , 6 km NE of Aktash; 50.3158° N, 87.7036° E; 6–7 Jul. 2017; S. Knyazev leg.; BOLD seq: 1555F07; PW: ML264 GoogleMaps 1 ♀ (worker); Altai, Ulagansky , 8 km NE of Aktash; 50.3160° N, 87.7253° E; 5–6 Jul. 2017; S. Knyazev leg.; BOLD seq: 1555F08; PW: ML265 GoogleMaps 1 ♀ (worker); Tuva, 27 km S of Erzin ; 50.0049° N, 95.1719° E; 11–12 Jul. 2014; A. Lelej leg.; BOLD seq: 1555F09; PW: ML266 GoogleMaps 1 ♀ (worker); Tuva, 25 km of Erzin ; 50.1611° N, 95.4778° E; 14–15 Jul. 2014; A. Lelej leg.; BOLD seq: 1555F10; PW: ML267 GoogleMaps 1 ♀ (worker); Tuva, 31 km of Erzin ; 50.0366° N, 95.4348° E; 18 Jul. 2014; A. Lelej leg.; BOLD seq: 1555F12; PW: ML269 GoogleMaps 1 ♀ (worker); same collection data as for preceding; 16 Jul. 2014; A. Lelej leg.; BOLD seq: 1555G01; PW: ML270 GoogleMaps .

KYRGYZSTAN • 1 ♀ (queen); Jalalabad , Jylamysh; 42.6522° N, 74.4812° E; 27 May 2009; L. Best leg.; BOLD seq: 1552H03; PW: ML156 GoogleMaps 1 ♀ (queen); Oshakaya , Kyzyl-Suu; 39.4761° N, 71.8933° E; 18 Jul. 1998; H. Rausch leg.; BOLD seq: 1552A08; OLML: ML191 View Materials GoogleMaps 1 ♀ (worker); Chong Aksuu Valley nr Karakol lake; 42.8386° N, 77.3889° E; 26 Jun. 2016; R. DeJonghe leg.; BOLD seq: 1555F04; RJ: ML261 GoogleMaps 1 ♀ (worker); same collection data as for preceding; BOLD seq: 1555F05; RJ: ML262 GoogleMaps 1 ♀ (worker); same collection data as for preceding; BOLD seq: 1555F06; RJ: ML263 GoogleMaps 1 ♀ (queen); Kungey Ala-Too , Chong Aksuu nr Karakol lake; 42.8397° N, 77.37° E; 25 Jun. 2016; R. DeJonghe leg.; BOLD seq: 6877A09; RJ: ML336 GoogleMaps 1 ♀ (queen); Terskey Ala-Too , Turgon-Aksuu river; 42.4183° N, 78.9467° E; 2 Jul. 2016; R. DeJonghe leg.; BOLD seq: 6877A11; RJ: ML341 GoogleMaps 1 ♀ (queen); same collection data as for preceding; BOLD seq: 6877A10; RJ: ML342 GoogleMaps 1 ♀ (queen); Terskey Ala-Too , Turgon-Aksuu river; 42.4019° N, 79.0486° E; 3 Jul. 2016; R. DeJonghe leg.; BOLD seq: 6877A12; RJ: ML343 GoogleMaps 1 ♀ (queen); Terskey Ala-Too , Saruu, Juktuu river; 42.0947° N, 77.9617° E; 6 Jul. 2016; R. DeJonghe leg.; BOLD seq: 6877B01; RJ: ML348 GoogleMaps 1 ♀ (queen); Santash ; 42.7389° N, 79° E; 29 Jun. 2016; R. DeJonghe leg.; BOLD seq: 6877B02; RJ: ML356 GoogleMaps 1 ♀ (queen); same collection data as for preceding; BOLD seq: 6877B03; RJ: ML357 GoogleMaps .

PAKISTAN • 1 ♀ (worker); Chitral, Khyber-Pakhtunkhwa, Tirich Valley ; 36.3150° N, 71.9824° E; 8 Aug. 1984; W. Budenburg leg.; BOLD seq: 6877C02; PW: ML383 GoogleMaps .

INDIA • 1 ♀ (queen); Kashmir , Zanskar, Nimaling Plain; 33.7927° N, 77.5881° E; 15 Jul. 1980; P. Williams leg.; BOLD seq: 1555H10; PW: ML311 GoogleMaps .

Global distribution

(Central Asian and Mongolian mountain species extending into the northwestern corner of the QTP) North Asia: MONGOLIA, RUSSIA: Tuva, Altai Mountains. – Central Asia: KAZAKHSTAN, KYRGYZSTAN, TAJIKISTAN. – East Asia: CHINA: Xinjiang. – Himalaya: PAKISTAN, INDIA: Kashmir. ( IAR, IOZ, NHMUK, OLML, PW, RDJ, RMNH, ZIN, ZMHB.). This species is widespread and can be common ( Fig. 212 View Figs 211–212 ).

Behaviour

Very few food-plant records but expected to be generalists ( Caragana versicolor in Williams 1991). The male mate-searching behaviour is expected to resemble the patrolling of B. keriensis s. str.

RMNH

National Museum of Natural History, Naturalis

PW

Paleontological Collections

NHMUK

Natural History Museum, London

OLML

Oberösterreichisches Landesmuseum

ZIN

Russian Academy of Sciences, Zoological Institute, Zoological Museum

Kingdom

Animalia

Phylum

Arthropoda

Class

Insecta

Order

Hymenoptera

Family

Apidae

Loc

Bombus separandus Vogt, 1909

Williams, Paul H., Altanchimeg, Dorjsuren, Byvaltsev, Alexandr, Jonghe, Roland De, Jaffar, Saleem, Japoshvili, George, Kahono, Sih, Liang, Huan, Mei, Maurizio, Monfared, Alireza, Nidup, Tshering, Raina, Rifat, Ren, Zongxin, Thanoosing, Chawatat, Zhao, Yanhui & Orr, Michael C. 2020
2020
Loc

Bombus tenellus subsp. tibetensis

Wang S. - F. 1982: 439
Reinig W. F. 1935: 341
1982
Loc

Bombus kozlowi

Bischoff H. 1936: 9
1936
Loc

Bombus keriensis

Reinig W. F. 1935: 340
1935
Loc

Bombus keriensis

Reinig W. F. 1935: 341
Frison T. H. 1930: 6
1935
Loc

Bombus keriensis

Williams P. H. 1998: 134
Reinig W. F. 1935: 341
Morawitz F. F. 1887: 199
1935
Loc

Lapidariobombus alagesianus subsp. pamirus

Skorikov A. S. 1931: 226
Skorikov A. S. 1931: 232
1931
Loc

Lapidariobombus alagesianus subsp. mongolicus

Skorikov A. S. 1931: 226
Friese H. 1916: 110
1931
Loc

Bombus tenellus

Reinig W. F. 1935: 333
Richards O. W. 1930: 639
1930
Loc

Bombus lapidarius

Friese H. 1913: 86
1913
Loc

Bombus kozlovi

Skorikov A. S. 1910: 413
Vogt O. 1909: 61
1910
Loc

Bombus lapidarius

Vogt O. 1909: 61
1909
Loc

Vogt O. 1909: 61
Cockerell T. D. A. 1906: 75
1909