Bombus incertus Morawitz, 1881

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 : 74-75

Bombus incertus Morawitz, 1881

Figs 15 , 123–124 , 195

Bombus incertus Morawitz, 1881: 229 .

Bombus insertus – Esmaili & Rastegar 1974: 25, incorrect subsequent spelling.

Bombus erzurumensis – Lecocq et al. 2015: supplementary table s1 (non Özbek, 1990: 209).

Vogt (1909: 49) initially considered the taxon incertus to be part (a ‘Form’) of the species B. lapidarius s. lat. although subsequently ( Vogt 1909: 52, 1911: 68) he regarded B. incertus as a species separate from B. lapidarius s. lat. Reinig (1935) also considered B. incertus to be a separate species.

Our PTP analysis ( Fig. 10 ) of coalescents in the COI gene supports three coalescents for candidate species within the sichelii- group as identified in the four gene species tree ( Figs 21–22 ): a west Asian B. incertus , which is strongly supported, and a widespread Eurasian B. sichelii s. lat., which is more weakly supported, and the Himalayan B. semenovianus . This is corroborated by differences in morphology that support the three as separate species (see the Diagnosis). The colour pattern of B. incertus appears to be particularly invariant.

Diagnosis

Females

Queens medium-sized body length 18–21 mm, workers 10–15 mm. Can be distinguished in West Asia by their combination of the hair of the face black, the hair of T2 posteriorly entirely bright white, the hindleg corbicular fringes predominantly black, and the red of T4–5 being a deep almost purplered rather than orange-red (cf. B. lapidarius , B. eriophorus , B. sichelii , B. alagesianus ). Queens of B. incertus are also larger than similarly coloured queens of B. sichelii from Iran and eastern Turkey, and can be distinguished by the complete absence of black hairs from T2 posteriorly and by the hindleg corbicular fringes without extensively orange hairs.

Males

Body length 12–15 mm. Can be distinguished in West Asia by their combination of the black band between the wing bases narrower than the anterior white band, the scutellum anterior margin with a few black hairs, T2 posteriorly entirely bright white, and the red of T4–7 being a deep almost purple-red rather than orange-red. Genitalia ( Fig. 195 ) with the gonostylus as long as broad, reduced as a rounded flat scale with the inner basal process reduced to a tooth (cf. rufipes- group, festivus- group, rufofasciatus - group); volsella with the inner distal corner broadly produced but without a narrow hook (cf. rufipesgroup, festivus- group, rufofasciatus -group); eye unenlarged relative to female eye.

Material examined

Holotype

? RUSSIA • ♀ (queen), holotype of Bombus incertus Morawitz, 1881 by monotypy; “Caucasien”; F. Morawitz leg.; not found (searched both ZIN and ZMUM), but identity as a species widespread in Turkey not in doubt.

Material sequenced (8 specimens)

TURKEY • 1 ♀ (queen); Lake Van; 38.4842° N, 42.2978° E; 18 Jun. 1988; K. Guichard leg.; BOLD seq: 1555C08; NHMUK: ML239 .

IRAN • 1 ♀ (worker); Tehran, Kamarbon ; 36.2162° N, 52.2928° E; 14 Aug. 2010; P. Abdoli leg.; NHMUK seq: IRANPW015; PW: ML11 • 1 ♀ (worker); Tehran, Kamarbon ; 36.2161° N, 52.2928° E; 14 Aug. 2010; P. Abdoli leg.; NHMUK: IRANPW016 ; PW: ML12 • 1 ♀ (worker); Kasra ; 38.3444° N, 47.6711° E; 6 Aug. 2009; A. Monfared leg.; NHMUK seq: IRANPW003; PW: ML13 • 1 ♀ (worker); Shahkuh, Gorgan ; 37.3698° N, 55.9065° E; 25 Sep. 2012; E. Spo leg.; NHMUK seq: NHMRC1; PW: ML14 • 1 ♀ (queen); Mazandaran, Ileka ; 36.2077° N, 51.4088° E; 9 May 2006; A. Taghavi leg.; BOLD seq: 1555C05; PW: ML236 • 1 ♀ (queen); Ardabil, Shabil ; 38.2809° N, 47.7182° E; 26 May 2006; A. Monfared leg.; BOLD seq: 1555C06; PW: ML237 • 1 ♀ (worker); Meshkinshahr, Walazir ; 38.3151° N, 47.7595° E; 4 Aug. 2005; A. Monfared leg.; BOLD seq: 1555C07; PW: ML238 .

Global distribution

(West Asian mountain species) West Asia: TURKEY, IRAN. (NHMUK, PW, YUY.) Contrary to some recent claims (S.-F. Wang 1985; S.-F. Wang & Yao 1996; Niu et al. 2018), this species does not occur in China. Examination of specimens in the IOZ collection labelled B. incertus shows these to be mostly misidentified B. separandus with a few B. keriensis from Xinjiang (examined PW). Bombus incertus is locally common.

Behaviour

Food-plant generalists ( Rasmont & Flagothier 1996). The male mate-searching behaviour is expected to resemble the patrolling of B. lapidarius .

Figures

Figs 14‒16. Maps of sequenced samples as in Figs 12–13. 14. The tanguticus-group and the lapidarius- group. 15. The sichelii-group. 16. The keriensis-group.

Figs 103–138. Simplified diagrams for the colour patterns of the hair on the dorsum for the species from the integrative analysis. The dorsum is divided into regions, each of which shows only the predominant or most apparent colour for that region using a simplified colour palette, with olive indicating a mixture of black and yellow hair, and grey indicating a mixture of black and white hair. The tanguticus-group. 103. Worker, China-Qinghai. 104. Worker, China-Qinghai. 105. Queen, China-Qinghai. 106. Queen, China-Xizang. 107. Queen, India-Kashmir. The lapidarius-group. 108. Queen, Russia-North Ossetia. 109. Worker, Azerbaijan. 110. Queen, Georgia. 111. Male, Russia-North Ossetia. 112. Male, Russia-North Ossetia. 113. Male, Russia-North Ossetia. 114. Male, Turkey. 115. Queen, Morocco. 116. Queen, Spain. 117. Queen, Spain. 118. Queen, UK. 119. Male, Spain. 120. Male, Spain. 121. Male, UK. 122. Male, UK. The sichelii-group. 123. Queen, Iran. 124. Male, Iran. 125. Queen, India-Kashmir. 126. Queen, India- Kashmir. 127. Male, India-Kashmir. 128. Male, India-Kashmir. 129. Queen, Iran. 130. Queen, Russia- Sakha. 131. Queen, China-Sichuan. 132. Worker, China-Sichuan. 133. Queen, Mongolia. 134. Worker, Mongolia. 135. Queen, Spain. 136. Worker, Austria. 137. Male, Turkey. 138. Male, Mongolia.

Figs 190‒198. Morphology of the male genitalia for species of the subgenus Melanobombus von Dalla Torre, 1880 from the dorsal aspect, anterior at the bottom of the image, posterior at the top. 190. Bombus friseanus Skorikov, 1933, China-Yunnan. 191. B pyrosoma Morawitz, 1890, China- Beijing. 192. B. formosellus (Frison, 1934), China-Taiwan. 193. B. eriophorus Klug, 1807, Russia- North Ossetia. 194. B. lapidarius (Linnaeus, 1758), UK. 195. B. incertus Morawitz, 1881, Turkey. 196. B. semenoviaus (Skorikov, 1914), India-Kashmir. 197. B. sichelii Radoszkowski, 1859, Austria. 198. B. ladakhensis Richards, 1928, China-Sichuan. Scale bars = 1 mm.

Fig. 10. October 2019 (Fig. 8) Bayesian PTP analysis of MrBayes tree of unique COI barcodes (UHF- PTP). Symbols as in Fig. 9.

Fig. 21. A sample of 10 000 species trees from among 100 million trees for species of the subgenus Melanobombus von Dalla Torre, 1880 reconstructed with *BEAST from gene trees for the COI, 16S, opsin and PEPCK genes, displayed in green using Densitree, with the summary ‘root canal’ superimposed as a thick blue line. The outgroup B. nobilis Friese, 1905 is shown.

Fig. 22. Most likely dated phylogenetic (ultrametric) tree for the species of the subgenus Melanobombus reconstructed von Dalla Torre, 1880 with *BEAST from trees for four genes (COI, 16S, PEPCK, opsin) with B. nobilis Friese, 1905 as the outgroup (not shown), estimated as the maximum-clade-credibility tree among a sample of 10 000 species trees with a 1% burn-in out of 100 million MCMC trees. Values above the nodes are Bayesian posterior probabilities showing support for groups. Values below the nodes are estimated dates of divergence in Ma (millions of years before the present) calibrated from a molecular estimate for the date of crown divergence within the subgenus Melanobombus. Grey bars show the 95% confidence limits on the estimated dates of divergence. Species groups discussed in the text are labelled in circles: rp = rufipes-group; fs = festivus-group; rf = rufofasciatus-group; tg = tanguticus-group; la = lapidarius-group; si = sichelii-group; and ke = keriensis-group.