Bombus incertoides Vogt, 1911

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 : 87-88

Bombus incertoides Vogt, 1911 stat. rev.

Figs 16 , 153–156 , 201 , 212

Bombus separandus [subsp.] incertoïdes Vogt, 1911: 58 by indication of Vogt (1909): 61, not infrasubspecific after Skorikov 1923: 153 ( ICZN 1999: Art. 45.6.4.1).

Bombus keriensis (part) – Reinig 1935: 341. — Williams 1991: 96 (non Morawitz, 1887: 199).

Bombus incertoides was first treated as a separate species by Skorikov (1923), without explanation, but has since been treated as a part of B. keriensis s. lat. (the keriensis -complex) by Reinig (1935) and by Williams (1998).

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

From morphology, B. incertoides has enlarged male eyes and has a bright white banding pattern in the hair with (most strongly for the keriensis -complex) the thoracic dorsum with the posterior white band particularly narrowed by black hair anteriorly on the scutellum.

Bombus incertoides can co-occur locally with the closely similar (but locally yellow-banded) B. separandus , at least in the Altai mountains of Mongolia.

Diagnosis

Females

Queens medium-sized body length 18–21 mm, workers 10–12 mm. Can be distinguished within Mongolia by the hair of the face black, the pale hair anteriorly on the thoracic dorsum pure white, the hair on the side of the thorax in its lower half black, the anterior half of the scutellum black, and by the hair of T4–5 red (cf. B. sichelii , B. separandus ).

Males

Body length 12–15 mm. Can be distinguished within Mongolia by the hair of the face black. Genitalia ( Fig. 201 ) heavily sclerotised and dark, with the gonostylus longer than broad, its inner basal projection reduced to a 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); antenna short and eye slightly enlarged relative to female eye.

Material examined

Holotype

MONGOLIA • ♀ (queen), holotype of Bombus separandus incertoides Vogt, 1911 by monotypy ( Williams, 1991); “N.[orth]” (Leder’s material from northern Mongolia is likely to have come from

a site between Kyakhta and Ulaanbaatar: Williams et al. 2012a); “[18]92”; H. Leder leg.; RMNH (examined PW).

Material sequenced (9 specimens)

MONGOLIA • 1 ♀ (queen); Tov, Tuul river ; 47.8118° N, 106.6139° E; 24 Jun. 2005; D. Altanchimeg leg.; NHMUK seq: NHMSR2; DA: ML148 • 1 ♂; Khovd, Monkhkhairkhan ; 46.9676° N, 91.6496° E; 9 Aug. 2014; D. Altanchimeg leg.; NHMUK seq: NHMSR6; DA: ML173 • 1 ♀ (queen); Arkhangai, Bat-olziit ; 46.7974° N, 101.7551° E; 18 Jun. 2014; R. DeJonghe leg.; BOLD seq: 6880B06; RDJ: ML490 • 1 ♂; Arkhangai, Chuluut Khujirt Mts ; 47.4418° N, 100.3643° E; 7 Aug. 2018; R. DeJonghe leg.; BOLD seq: 6880B07; RDJ: ML491 • 1 ♂; same collection data as for preceding; BOLD seq: 6880B08; RDJ: ML492 • 1 ♀ (worker); Arkhangai, Chuluut Khujirt Mts ; 47.2697° N, 100.3387° E; 8 Aug. 2018; R. DeJonghe leg.; BOLD seq: 6880B09; RDJ: ML493 • 1 ♂; Arkhangai, Chuluut Khujirt Mts ; 47.2700° N, 100.3387° E; 9 Aug. 2018; R. DeJonghe leg.; BOLD seq: 6880B10; RDJ: ML494 • 1 ♂; same collection data as for preceding; BOLD seq: 6880B11; RDJ: ML495 • 1 ♀ (queen); Khovsgol, Khatgal ; 50.4815° N, 100.2170° E; 7 Jun. 2015; R. DeJonghe leg.; BOLD seq: 6880C04; RDJ: ML501 .

Global distribution

(Mongolian mountain species) North Asia: MONGOLIA. (DA, NHMUK, RMNH, PW, RDJ.) This species is widespread but not common ( Fig. 212 ).

Behaviour

Expected to be food-plant generalists but no records. The male mate-searching behaviour is unknown.

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 139–180. 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 keriensis-group. 139. Queen, China-Sichuan. 140. Worker, China-Sichuan. 141. Queen, China-Sichuan. 142. Queen, China-Sichuan. 143. Worker, China-Xizang. 144. Queen, India-Kashmir. 145. Male, China-Qinghai. 146. Male, India-Kashmir. 147. Male, China-Gansu. 148. Queen, Armenia. 149. Queen, Iran. 150. Male, Georgia. 151. Worker, China-Qinghai. 152. Male, China-Qinghai. 153. Queen, Mongolia. 154. Worker, Mongolia. 155. Male, Mongolia. 156. Male, Mongolia. 157. Queen, China-Sichuan. 158. Worker, China-Qinghai. 159. Worker, China-Qinghai. 160. Male, China-Qinghai. 161. Male, China-Qinghai. 162. Male, China-Qinghai. 163. Queen, Pakistan. 164. Queen, Pakistan. 165. Queen, Pakistan. 166. Queen, Pakistan. 167. Queen, Afghanistan. 168. Queen, Pakistan. 169. Queen, Pakistan. 170. Queen, India-Kashmir. 171. Male, India-Kashmir. 172. Male, India-Kashmir. 173. Queen, China- Xinjiang. 174. Queen, Kyrgyzstan. 175. Queen, Kyrgyzstan. 176. Queen, Mongolia. 177. Queen, India- Kashmir. 178. Male, Kazakhstan. 179. Male, China-Xinjiang. 180. Male, Mongolia.

Figs 199‒204. 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. 199. Bombus alagesianus Reinig, 1930 stat. rev., Georgia. 200. B. tibeticus sp. nov., China-Qinghai. 201. B. incertoides Vogt, 1911 stat. rev., Mongolia. 202. B. qilianensis sp. nov., China-Qinghai. 203. B. keriensis Morawitz, 1887, India-Kashmir. 204. B. separandus Vogt, 1909 stat. rev., Kyrgyzstan. Scale bars = 1 mm.

Figs 211–212. Maps of sequenced samples showing the most frequent colour patterns locally. 211. The sichelii-complex, consisting of the single polytypic species B. sichelii Radoszkowski, 1859 (inset: B. sichelii worker, China-Neimenggu, photo PW). 212. The keriensis-complex, consisting of B. alagesianus Reinig, 1930 stat. rev. (blue spots), B. tibeticus sp. nov. (dark green spots), B. incertoides Vogt, 1911 stat. rev. (light green spots), B. qilianensis sp. nov. (yellow spots), B. keriensis Morawitz, 1887 (orange spots), and B. separandus Vogt, 1909 stat. rev. (red spots) (inset: B.qilianensis sp. nov. queen, China-Sichuan, photo PW). Maps projected onto a sphere in ArcGIS using World_Shaded_Relief basemap © 2014 ESRI and showing boundaries between countries as recognised by the UN.

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

Figs 17–20. Plots of (y-axis) pairwise proportion of genetic divergence between COI barcode region sequences against (x-axis) pairwise Great Circle geographical distance in km between sample sites with linear trend lines (black). 17. Genetic divergences within the candidate species for the sichelii-complex identified using Bayesian UF-PTP (Mantel r = 0.52). 18. Genetic divergences among the candidate species for the sichelii-complex identified using Bayesian UF-PTP (Mantel r = 0.21). 19. Genetic divergences within the candidate species for the keriensis-complex identified using Bayesian UF-PTP (Mantel r = 0.16). 20. Genetic divergences among the candidate species for the keriensis-complex identified using Bayesian UF-PTP (relationship not positive). For details of the measurements see the