Widespread polytypic species or complexes of local species? Revising bumblebees of the subgenus Melanobombus world-wide (Hymenoptera, Apidae, Bombus)
Author
Williams, Paul H.
38A45E0C-02A8-407E-8E89-5162D454E9FE
Natural History Museum, Cromwell Road, London SW 7 5 BD, UK.
paw@nhm.ac.uk
Author
Altanchimeg, Dorjsuren
FB68F32E-4F6D-40C2-A921-20FBAD676D50
Institute of General and Experimental Biology, Peace Avenue 54 b, Ulaanbaatar 13330, Mongolia.
altanchimegd@mas.ac.mn
Author
Byvaltsev, Alexandr
B57BAD3E-9E42-4446-994E-4A45A738D404
Novosibirsk State University, ul. Pirogova 2, Novosibirsk, 630090 Russia.
byvam@yandex.ru
Author
Jonghe, Roland De
FC98CAB7-B2FF-4BEB-94FF-26F53D33CD04
Langstraat 105, B- 2260 Westerlo, Belgium.
roland.de.jonghe@telenet.be
Author
Jaffar, Saleem
77F70375-0A19-4D0E-A05A-987BB46543C0
South China Agricultural University, Guangzhou 510642, China.
saleemjaffar@stu.scau.edu.cn
Author
Japoshvili, George
CCC82B7C-A1E4-4D58-90A3-623116CBAE96
Agricultural University of Georgia, 240 Agmashenebli Alley, Tbilisi, Georgia.
g.japoshvili@agruni.edu.ge
Author
Kahono, Sih
F8513496-B409-434C-A182-4146232C89FA
Indonesian Institute of Sciences (LIPI), Jakarta, Indonesia.
sihkahono@gmail.com
Author
Liang, Huan
A99867E0-C686-4608-8DF7-0EDE8D2D57EC
Kunming Institute of Botany (Chinese Academy of Sciences), 132 Lanhei Road, Kunming, Yunnan 650201, China.
lianghuan@mail.kib.ac.cn
Author
Mei, Maurizio
82F344C7-B98A-462C-81E0-D6F3F02348D4
Università di Roma ‘ Sapienza’, Piazzale Valerio Massimo 6, Roma 00162, Italy.
maurizio.mei@uniroma1.it
Author
Monfared, Alireza
48CA77BA-8CF4-4812-89B1-696A11FEDE2D
Yasouj University, Zirtol, Yasouj, Iran.
alirezamonfared1@yahoo.com
Author
Nidup, Tshering
BE588EE1-5E2C-46CC-8907-CD344D88C869
Sherubtse College, Royal University of Bhutan, Trashigang, Bhutan.
tsheringnidup@sherubtse.edu.bt
Author
Raina, Rifat
48E5AE7A-D5DC-4549-94B7-FD8489D1EF9E
Zoological Survey of India, Pali Road, Jodhpur 342005, Rajasthan, India.
rifat72001@rediffmail.com
Author
Ren, Zongxin
27B9DD39-62A8-44D3-9D6A-E6C20D8AAA27
Kunming Institute of Botany (Chinese Academy of Sciences), 132 Lanhei Road, Kunming, Yunnan 650201, China.
renzongxin@mail.kib.ac.cn
Author
Thanoosing, Chawatat
6F4C150C-BC03-4F75-91A6-2A8AF6B5905C
Natural History Museum, Cromwell Road, London SW 7 5 BD, UK.
c.thanoosing@nhm.ac.uk
Author
Zhao, Yanhui
299C8EEA-699E-4B15-9BCD-9806E0E7EE63
Kunming Institute of Botany (Chinese Academy of Sciences), 132 Lanhei Road, Kunming, Yunnan 650201, China.
zhaoyanhui@mail.kib.ac.cn
Author
Orr, Michael C.
1E7F46C3-870E-460C-A611-BA1042ED99FB
Institute of Zoology (Chinese Academy of Sciences), 1 Beichen West Road, Chaoyang, Beijing 100101, China.
michael.christopher.orr@gmail.com
text
European Journal of Taxonomy
2020
2020-10-02
719
1
120
journal article
10.5852/ejt.2020.719.1107
7ca72f76-4fae-4305-8601-4662f4cd2b96
2118-9773
4064324
A4500016-C219-4353-B81C-5E0BB520547F
Bombus tanguticus
Morawitz, 1887
Figs 13
,
103–107
,
206
Bombus tanguticus
Morawitz, 1887: 200
.
Our PTP analysis (
Fig. 10
) of coalescents in the COI gene supports a single species
B. tanguticus
within the
tanguticus-
group as identified in the four gene species tree (
Figs 21–22
), corroborated by differences in morphology.
Bombus tanguticus
has a distinctive morphology and has been placed in a separate subgenus,
Tanguticobombus
(
Pittioni 1939b
), although this was synonymised with the subgenus
Melanobombus
by
Richards (1968)
. A close relationship with other species of the subgenus
Melanobombus
is supported by an analysis of COI barcodes by
Williams (2018)
, which like the tree for the COI gene here (
Fig. 10
), placed this species in the
lapidarius
-group. In contrast, when the species tree was estimated from four gene trees that include slower-evolving nuclear genes that can provide more reliable support for older relationships,
B. tanguticus
is placed in its own
tanguticus
-group next to the
lapidarius
-group (
Fig. 21
). Female morphology is inconclusive, because the labrum has a very narrow median groove and the posterior ocello-ocular areas are relatively unpunctured, which support grouping
B. tanguticus
with
B. lapidarius
. In contrast, the more densely hairy hind corbiculum and elongate hind basitarsus (as well as large queen body size) support a closer relationship of
B. tanguticus
with
B. simillimus
. Resolving this question might be possible with information from other genes, but it is very likely that the morphology of the male genitalia, once the male is described, will be diagnostic among the two species groups. This is one of the few bumblebee species world-wide for which the male still remains unknown (
Williams 2018
).
The typical colour pattern of this species is yellow banded, but a worker from
Qinghai
(ML219) has a white-banded colour pattern, as confirmed from their identical COI barcode sequences (Fig. 9).
Diagnosis
Females
Queens very large body length
22–28 mm
, workers
12–13 mm
. Can be distinguished in the QTP by their combination of labral lateral tubercles medially narrowing and projecting as small pointed pyramids, the median longitudinal furrow between them deep and unusually narrow, 0.2× as broad as the labrum, forming a triangular notch (broader for co-occurring
B. tibeticus
,
B. (Sibiricobombus) semenovi
Morawitz, 1887
,
B. (Sibiricobombus) asiaticus
Morawitz, 1875
s. lat.
); mandible lacking a distinct sulcus obliquus, the posterior notch (incisura) anterior to the posterior tooth very weak or absent; clypeus with many large punctures especially in the dorsal half, the apical impressions unusually deep; antennal segment 3 (A3) short, nearly equal in length to A5, and only 1.5 × as long as A4 (substantially longer for co-occurring
B. tibeticus
,
B. semenovi
,
B. asiaticus
s. lat.
); the mid basitarsus with the posterior distal corner acute; hind tibia with the outer (corbicular) surface weakly reticulate and shining, for the queens uniformly covered (even in the centre) with dense short black fine branched hairs (these can become abraded for older queens and are few or absent in workers); hind basitarsus long and narrow, nearly 4× as long as its greatest breadth, the posterior edge almost straight in its distal 4⁄5, the posterior distal corner acutely produced (basitarsus shorter and broader for co-occurring
B. tibeticus
,
B. semenovi
,
B. asiaticus
s. lat.
), the outer surface with the short hair black with only short side branches so that some of the shining outer surface is visible between them (cf. dense overlapping pale hair for
B. tibeticus
); T6 posteriorly with a subapical rounded (nearly circular) boss, the apex for workers truncated or for queens deeply and narrowly divided medially (T6 undivided for queens of co-occurring
B. tibeticus
,
B. semenovi
,
B. asiaticus
s. lat.
).
Males
Unknown.
Material examined
Lectotype
CHINA
•
♀
(queen),
lectotype
of
Bombus tanguticus
Morawitz, 1887
by designation of
Williams (2018)
;
Qinghai
, “Bassin des gelb.[en] Flusses” (= source area of the Huang He, probably near Zhaling Hu:
Williams 2018
); [1884]; [N. Przhevalsky leg.];
ZIN
(examined
PW
).
Material sequenced
(
3 specimens
)
CHINA
–
Qinghai Province
•
1 ♀
(worker);
Xiangpishan
;
36.7589° N
,
99.6041° E
;
30 Aug. 2010
;
P. Williams
leg.; BOLD seq: 6875A06;
PW
:
ML9
•
1 ♀
(worker);
Kunlun Pass
;
35.7258° N
,
94.1626° E
;
13 Aug. 2013
;
P. Williams
leg.;
NHMUK
seq: NHMRC2;
PW
:
ML219
. –
Xizang Province
•
1 ♀
(queen);
Kamba county
(Z715
80 km
);
28.2193° N
,
88.7942° E
;
10 Jul. 2018
;
Q.-T. Wu
leg.;
IOZ
seq: BB11;
IOZ
:
ML566
.
Global distribution
(Qinghai-Tibetan-Plateau species)
East Asia
:
CHINA
:
Xizang
,
Qinghai
. –
Himalaya
:
INDIA
: Kashmir,
Sikkim
. (IAR, IOZ, NHMUK, PW, ZIN.) This is one of the highest recorded bumblebees in
the world, from an elevation of ca
5640 m
a.s.l. (
Williams 2018
: along with
B. tibeticus
). The species is widely distributed but generally very rare (
Fig. 206
). There are data available for only 12 queens and
2 workers
, including the two recent queens from Kamba county,
Xizang
, collected in 2018 (IOZ, examined PW).
Behaviour
Expected to be food-plant generalists but very few records known (
Williams 2018
). Males of
B. tanguticus
are expected to be flying during mid to late August, either visiting flowers or searching for mates in semi-arid habitats at elevations above
4500 m
a.s.l. (
Williams 2018
). Males of the mountain species
B. keriensis
have been found patrolling near high ridges towards the upper end of the species’ elevational range (above
4000 m
a.s.l.) in Kashmir (
Williams 1991
). Similar ‘hill-topping’ near high mountain ridges towards the upper end of its elevational range in the QTP is likely for
B. tanguticus
.