Bombus sichelii Radoszkowski, 1860

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 : 76-79

Bombus sichelii Radoszkowski, 1860

Figs 15 , 129–138 , 197 , 211

Bombus SICHELII Radoszkowski, 1860: 481 .

Bombus alticola Kriechbaumer, 1873: 339 .

Bombus lapidarius subsp. nigritulus Friese, 1905: 518 .

Bombus lapidarius subsp. albidulus Friese, 1905: 518 .

Bombus sicheli Form [subsp.] uniens Vogt, 1909: 62.

Bombus noster Skorikov, 1910b: 578 .

Bombus sicheli var. geogr . [subsp.] czerskiae Vogt, 1911: 59.

Bombus sicheli var. geogr . [subsp.] rehbinderianus Vogt, 1911: 60.

Bombus sicheli var. geogr . [subsp.] cazurroi Vogt, 1911: 60.

Lapidariobombus verticiflavus Skorikov, 1931: 225 .

Bombus sicheli f.g. [subsp.] daghestanicus Reinig, 1935: 345 (non Radoszkowski, 1877: vii = B. sylvarum (Linnaeus)) .

Bombus jeholensis Yasumatsu, 1935: 45 .

Bombus sicheli subsp. chinganicus Reinig, 1936: 6 .

Pyrobombus sicheli subsp. flavissimus Tkalců, 1975: 177 , replacement name for latofasciatus Vogt, 1909: 49.

Pyrobombus erzurumensis Özbek, 1990: 209 .

Bombus Sicheli – Radoszkowski 1877: 213, incorrect subsequent spelling.

Since Vogt (1909), B. sichelii s. lat. has generally been considered as a species separate from B. lapidarius s. lat. Vogt (1909: 46) stated that his “Formen” are subpopulations, which could be considered equivalent to subspecies. Reinig (1935: 333) stated explicitly that his “f.g.” (forma geographica) are equivalent to subspecies. These taxa are considered here to be parts of a sichelii -complex.

The taxon erzurumensis was described more recently as a separate species from north-eastern Turkey, although we have seen it also from Iran. Within the sichelii -complex it has an especially narrow band of black hair posteriorly on T2. It was first recognised as likely to be conspecific with B. sichelii s. lat. by Williams (1998).

Our PTP analysis ( Fig. 10 ) supports two coalescents in the COI gene, for a west Asian B. incertus , which is strongly supported, and a widespread Eurasian B. sichelii s. lat. Although the support for a COI coalescent for B. sichelii s. lat. is relatively weak, it is the best supported in this part of the tree, so no alternative interpretation of the member taxa is better supported as species. Corroboration from a positive divergence-with-distance relationship both within ( Fig. 17 ) and among ( Fig. 18 ) the component lineages (Fig. 9) is accepted as additional support for B. sichelii s. lat. as a single broadly distributed species in its long accepted sense.

From morphology, B. sichelii s. lat. has a colour pattern of the hair of T2 with black hairs posteriorly that is consistently distinct from B. incertus .

Diagnosis

Females

Queens medium-sized body length 17–20 mm, workers 9–13 mm, Can be distinguished by usually having at least some pale hair on the face at the base of the antenna (cf. B. separandus , B. incertus ), T2 posteriorly usually has at least a narrow line of black hair (cf. B. keriensis , B. separandus , B. alagesianus , B. incertus etc.), and the hindleg corbicular fringes have some extensively orange hairs. Some individuals from eastern Turkey and Iran (the taxon erzurumensis ) resemble B. incertus closely because they have very few black hairs posteriorly on T2 and few or no pale hair on the face, but the queens of B. sichelii are smaller than B. incertus , the red of T4–5 is more orange-red, and the hindleg corbicular fringes have some extensively orange hairs.

Males

Body length 11–15 mm. Can be distinguished in Europe and West Asia by their combination of yellow or white hair posteriorly on the thoracic dorsum, T2 with yellow or white but posteriorly with some black, and T4–7 a pale red. Genitalia ( Fig. 197 ) 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. rufipes- group, festivus- group, rufofasciatus -group); eye unenlarged relative to female eye.

Material examined

Lectotype

RUSSIA • ♀ (queen), lectotype of Bombus sichelii Radoszkowski, 1860 by designation of Tkalců (1974b); Kamchatka; Dittmar leg.; ZMHB (examined PW).

Material sequenced (46 specimens, includes some sequences from Lecocq et al. 2015)

SPAIN • 1 ♂; Catalonia, Cerbi ; 42.646° N, 1.152° E; 16 Sep. 1993; C. Schmid-Egger leg.; BOLD seq: BCZSMHYM02963; ZSM: ML327 • 1 ♂; same collection data as for preceding; BOLD seq: BCZSMHYM02964; ZSM: ML328 .

SWITZERLAND • 1 ♀ (worker); 46.2210° N, 7.6203° E; 22 Jul. 1999; P. Williams leg.; BOLD seq: 1555H11; PW: ML313 • 1 ♀ (worker); Ticino, Lago Sambuco ; 46.466° N, 8.648° E; 28 Jul. 1993; C. Schmid-Egger leg.; BOLD seq: BCZSMHYM02961; ZSM: ML326 .

AUSTRIA • 1 spec.; M̹hl; 47.5014° N, 10.7361° E; D. Michez leg.; GenBank seq: KM458067 ; UMONS: ML285 .

FRANCE • 1 spec.; Col d’Allos; 44.1733° N, 6.3544° E; O. Poncheau leg.; GenBank seq: KM458070 ; UMONS: ML288 • 1 spec.; Gavarnie; 42.7119° N, 0.0075° W; D. Michez leg.; GenBank seq: KM458072 ; UMONS: ML289 .

RUSSIA • 1 ♀ (worker); Chuvashiya, Severniy ; 56.2899° N, 47.1931° E; 24 Jun. 2010; A. Lashtukhin leg.; BOLD seq: SHMEL-E11; SDM ML331 • 1 spec.; Buryatia, Mondy ; 51.4103° N, 100.5936° E; T. DeMeulemeester leg.; GenBank seq: KC915894 ; UMONS: ML290 • 1 spec.; same collection data as for preceding; GenBank seq: KC915896 ; UMONS: ML291 • 1 ♀ (worker); Tuva, 31 km Erzin ; 50.0366° N, 95.4348° E; 18 Jul. 2014; A. Lelej leg.; BOLD seq: 1555F11; NSUN: ML268 • 1 ♀ (worker); Sakhalin, south Schmidta Peninsula ; 53.8874° N, 142.7429° E; 13 Aug. 2003; J. Jobe leg.; BOLD seq: 1552F06; SEMC: ML189 • 1 ♀ (queen); Kamchatka; 53.7887° N, 159.3172° E; BOLD seq: 6879D08; FSCV: ML217 .

TURKEY • 1 ♀ (worker); 41.2008° N, 42.5283° E; BOLD seq: 6879A04; PW: ML188 • 1 spec.; Zigana Geçidi; 40.6353° N, 39.4044° E; P. Rasmont leg.; GenBank seq: KM458068 ; UMONS: ML286 • 1 spec.; same collection data as for preceding; GenBank seq: KM458069 ; UMONS: ML287 .

IRAN • 1 ♀ (queen); 38.2974° N, 48.0360° E; 26 May 2006; NHMUK seq: IRANPW022; PW: ML15 • 1 ♀ (worker); 38.2724° N, 47.6886° E; 29 May 2015; NHMUK seq: IRANPW020; PW: ML16 • 1 ♀ (worker); 38.2717° N, 47.8515° E; 31 May 2014; NHMUK seq: IRANPW023; PW: ML17 .

KYRGYZSTAN • 1 ♀ (queen); Santash ; 42.7389° N, 79° E; 28 Jun. 2016; R. DeJonghe leg.; BOLD seq: 1555F03; RDJ: ML260 .

MONGOLIA • 1 ♀ (worker); Ovorhangay, Karakorum ; 47.1667° N, 102.8333° E; 13–16 Aug. 1990; C. Cockell leg.; BOLD seq: 6874D04; PW: ML18 • 1 ♀ (worker); Hovsgol, Tsagaan Nur ; 50.601° N, 101.523° E; 16 Jul. 2005; J. Gelhaus leg.; BOLD seq: 1550G02; SEMC: ML215 • 1 ♀ (worker); Dalbay ; 51.025° N, 100.766° E; 20 Jul. 2007; D. Song leg.; BOLD seq: 9808C12; PCYU: ML332 • 1 ♀ (worker); Dalbay ; 51.0279° N, 100.7667° E; 4 Aug. 2007; D. Song leg.; BOLD seq: 9808E01; PW: ML218 • 1 ♀ (worker); Khovsgol Nuur ; 51.0628° N, 100.7322° E; 21 Jul. 2004; D. Sheppard leg.; NHMUK seq: PW28 ; PW: ML324 .

CHINA – Neimenggu Province • 1 ♀ (worker); Moerdaga ; 51.3349° N, 120.6784° E; 18 Aug. 2009; P. Williams leg.; NHMUK seq: CT465; PW: ML465 • 1 ♀ (worker); nr. Huanggangliang ; 43.5423° N, 117.6233° E; 24 Aug. 2010; P. Williams leg.; BOLD seq: 1555B12; PW: ML231 . – Shanxi Province • 1 ♀ (worker); Yingxian ; 39.4008° N, 113.4167° E; 8 Sep. 2007; P. Williams leg.; BOLD seq: 1555C01; PW: ML232 . – Gansu Province • 1 ♀ (worker); Pinnan ; 34.3281° N, 105.6278° E; 30 Aug. 2009; P. Williams leg.; BOLD seq: 1555H01; PW: ML303 . – Sichuan Province • 1 ♀ (worker); Hongyuan ; 32.6403° N, 102.3286° E; 4 Aug. 2002; P. Williams leg.; BOLD seq: 1555H02; PW: ML304 • 16 specs; Hongyuan ; 32.3282° N, 102.4543° E; Y. Dong leg.; YD seq: DYX53.1, DYX53.2, DYY17, DYX61.1, DYX1.1, DYX18.1, DYX30.2, DYX34.1, DYX11.2, DYZX2, DYX64, DYX1.3, DYX13.1, DYX27.1, DYX13.2, DYX69.1; YD: ML443 to ML458 .

Global distribution

(Widespread Palaearctic species, in steppes in the north and in mountains in the south) Europe: SPAIN, FRANCE, SWITZERLAND, AUSTRIA. – West Asia: TURKEY, IRAN. – North Asia: KYRGYZSTAN, MONGOLIA, RUSSIA: European Russia, North Caucasus, Ural, West Siberia, Altai Mountains, East Siberia, Far East. – East Asia: CHINA: Xizang, Sichuan, Gansu, Ningxia, Shanxi, Hebei, Neimenggu, Liaoning, Jilin, Heilongjiang; N KOREA, S. KOREA. (FSCV, IAR, IOZ, NHMUK, PW, SC, SEMC, YUY, ZMHB.) The species is widespread ( Fig. 210 ) and often abundant.

Behaviour

Food-plant generalists ( Rasmont 1988; Neumayer & Paulus 1999; Williams et al. 2009; An et al. 2014). 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.

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

Figs 209–210. Images of queens of two cryptic species from around the western Qinghai-Tibetan plateau. 209. Bombus keriensis Morawitz, 1887 from Mt Apharwat (4000 m a.s.l.) in the Pir Panjal mountains (ML405). 210. B. separandus Vogt, 1909 stat. rev. from Nimaling plain (4800 m a.s.l.) in the Zanskar mountains (ML311). Viewed from the left lateral aspect.