Bombus (Alpigenobombus) nobilis Friese, 1905

Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li & Orr, Michael C., 2023, Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae), European Journal of Taxonomy 892 (1), pp. 1-65 : 38-39

6. Bombus (Alpigenobombus) nobilis Friese, 1905 s. str.

Figs 5 , 61‒69 , 111

Bombus nobilis Friese, 1905: 513 .

Bombus (Nobilibombus) xizangensis Wang, 1979: 188 .

Bombus (Nobilibombus) chayaensis Wang, 1979: 189 .

Species-taxon concept and variation

The taxon concept of the species B. nobilis here agrees with the recent interpretation (Williams 2022a) that it is separate from the taxon concept of the species B. sikkimi (see the comments above on B. sikkimi ), based on: (1) our PTP analysis supports independent species-level coalescents in the COI gene ( Fig. 12 ); corroborated by (2) diagnostic morphological character states (see the keys).

The PTP and morphological results ( Fig. 12 , keys) support the interpretation that B. nobilis , B. sikkimi , and B. validus are separate species (Williams 2022a). The available COI-barcode-like sequences may all be low-divergence neonumts ( Fig. 11 ).

From the morphology and colour pattern it appears likely that the yellow-banded holotype queen of the taxon chayaensis (Fig. 63, from Chagyab, north-eastern Xizang) and the white-banded holotype worker of the taxon xizangensis (Fig. 62, from Zayü, southern Xizang) are conspecific with B. nobilis . A small collection from northern Myanmar (Adung valley, 1931) of both yellow-banded (NHMUK: 3 ♀♀ (2 queens, 1 worker), Figs 65–66) and white-banded (NHMUK: 2 ♀♀ (workers), Fig. 61) individuals matches this species in morphology.

Variation of B. nobilis is shown in the colour-pattern diagrams in Figs 61‒69. Bombus nobilis , with the yellow-banded colour pattern in the north (northern Hengduan) of its range (Figs 63–64, 69), appears to mimic the abundant B. (Melanobombus) friseanus Skorikov, 1933 ( Williams 2007: fig. 5j), whereas the white-and-yellow-banded colour pattern in the south (southern Hengduan: Figs 61–62, 67–68) appears to mimic the abundant B. (Ml.) prshewalskyi ( Williams 2007: fig. 5p).

Type material

Bombus nobilis Friese, 1905: 513 .

Type not found, but identity not in doubt ( Williams et al. 2009).

Bombus (Nobilibombus) xizangensis Wang, 1979: 188 .

Holotype by original designation: ♀ (worker) Zay ue , Xizang, China ( IOZ). Examined .

Bombus (Nobilibombus) chayaensis Wang, 1979: 189 .

Holotype by original designation: ♀ (queen) Chagyab , Xizang, China ( IOZ). Examined .

Morphological diagnosis

Female

Wings lightly clouded with brown with the veins dark brown, hair of medium length, oculo-malar area longer than broad, oculo-ocellar areas densely punctured with scattered medium-sized punctures with many small punctures between them, anterio-laterally to the lateral ocellus with large and small punctures sufficiently separated to have flat shining areas in between (cf. B. sikkimi , B. validus ); hair on the anterior dorsum of the thorax and of the side of the thorax and scutellum either golden yellow or grey-white, on the scutellum the yellow hair anteriorly scarcely divided in the middle by a triangle of black, T1 yellow, T2 either predominantly black or with large anterior lateral patches yellow.

Male

Wings lightly clouded with brown with the veins dark brown, hair of medium length, oculo-malar area longer than broad; genitalia ( Fig. 111 ) with the gonostylus short and distally near its midline axis deeply concavely rounded, its outer side about a half as long as its inner side with the two inner corners of the distal lobe rounded, the margin between them concave, penis-valve head weakly recurved, the recurved section about as long as broad at the base (cf. B. sikkimi , B. validus ); hair on the anterior dorsum of the thorax and of the side of the thorax and scutellum either golden yellow or grey-white, T1 either yellow or if nearly white then T2 anterio-laterally also nearly white, on the scutellum in the middle the yellow only shallowly divided by black.

Material sequenced in Fig. 12

CHINA • 1 ♀ (worker); Yunnan, Lijiang ; 27.0156° N, 100.1714° E; 12 Aug. 2017; H. Liang leg.; KIB seq: #2018064; KIB: AG#089 • 1 ♁; Xizang, Zhowagoin ; 28.3469° N, 98.0102° E; 3 Sep. 2014; Q.-T. Wu leg.; IOZ seq: FPW30; IOZ: AG#187 • 1 ♀ (worker); Sichuan, Queershan ; 29.5718° N, 98.1847° E; 5 Aug. 2018; Z. Ren et al. leg.; KIB seq: QESH201020; KIB: AG#143 • 1 ♀ (worker); Xizang, Dongdashan ; 29.5718° N, 98.1847° E; 21 Jul. 2018; Z. Ren et al. leg.; KIB seq: DDSM202013; KIB: AG#087 • 1 ♁; Xizang, Zhowagoin ; 28.3469° N, 98.0102° E; 2 Sep. 2014; Q.-T. Wu leg.; IOZ seq: FPW24; IOZ: AG#183 • Yunnan; IAR seq: HJF6; IAR: AG#090 .

Additional sequences in Fig. 10 and haplotype duplicates

CHINA • 1 ♀ (worker); Yunnan, Lijiang ; 27.0156° N, 100.1714° E; 2 Sep. 2012; Y. Zhao leg.; KIB seq: #218628241; KIB: AG#108 • 1 ♀ (worker); Xizang, Zhowagoin ; 28.3469° N, 98.0102° E; 3 Sep. 2014; Q.-T. Wu leg.; IOZ seq: FPW31; IOZ: AG#186 • 1 ♁; Xizang, Zhowagoin ; 28.4853° N, 97.3634° E; 30 Aug. 2014; Q.-T. Wu leg.; IOZ seq: FPW29; IOZ: AG#188 • 1 ♀ (worker); same collection data as for preceding but 31 Aug. 2014; IOZ seq: FPW28; IOZ: AG#182 • 1 ♀ (worker); Xizang, Zhowagoin ; 28.3469° N, 98.0102° E; 3 Sep. 2014; Q.-T. Wu leg.; IOZ seq: FPW26; IOZ: AG#189 • 1 ♀ (worker); same collection data as for preceding but 2 Sep. 2014; IOZ seq: FPW25; IOZ: AG#190 • 3 ♁♁; Xizang; 29.6114° N, 94.6441° E; 10 Aug. 2012; IAR seq: ZA3 ZA4 ZA5; IAR: AG#010 to AG#012 • 3 ♀♀ (workers); Xizang, Lulang ; 29.6414° N, 94.6982° E; 26 Jul. 2018; Z. Ren et al. leg.; KIB seq: SJLSM108001, SJLSM104001, SJLSM104002; KIB: AG#086 , AG#146 , AG#147 • 1 ♀ (worker); Xizang, Galonglashan ; 29.8219° N, 95.7110° E; 24 Jul. 2018; Z. Ren et al. leg.; KIB seq: GLLSL106001; KIB: AG#148 • 1 ♀ (worker); Yunnan, Lijiang ; 27.0156° N, 100.1714° E; 12 Aug. 2017; H. Liang leg.; KIB seq: #2018063; KIB: AG#103 • 1 ♀ (worker); Xizang, Markam ; 29.7432° N, 95.6753° E; 2 Sep. 2014; Q.-T. Wu leg.; IOZ seq: FPW32; IOZ: AG#191 .

Global distribution

Hengduan: China (Qinghai, Xizang, Yunnan, Sichuan, Gansu), Myanmar: IAR, IOZ, KIB, NHMUK, PW, SC, ZMHB.

This species is recorded at elevations of 2600‒4500 m in Sichuan, mostly above the tree line and into the subalpine zone (especially rich in Rhododendron species) ( Williams et al. 2009).

Behaviour

Male eye not obviously enlarged relative to female eye: males are expected to show ‘patrolling’ behaviour in search of mates ( Williams 1991).

Figures

Figs 1‒9. Individuals of the subgenus Alpigenobombus Skorikov, 1914 (with photo credits). 1. Bombus mastrucatus Gerstaecker, 1869, worker Norway robbing (P. Haringsma). 2. B. kashmirensis Friese, 1909 s. str., worker China-Sichuan robbing (PW). 3. B. kashmirensis (taxon meinertzhageni Richards, 1928) worker India-Kashmir-Zanskar (PW).4. B. sikkimiFriese, 1918, worker India-Arunachal (MS). 5. B. nobilis Friese, 1905 s. str., worker China-Yunnan (ZR). 6. B. genalis Friese, 1918, worker India-Arunachal (MS). 7. B. breviceps Smith, 1852 (taxon dentatus Handlirsch, 1888) worker China- Yunnan (PW). 8. B. breviceps (taxon channicus Gribodo, 1892) worker Thailand (CT). 9. B. grahami (Frison, 1933) worker India-Arunachal (MS). Some images reversed.

Figs 19–105 (see pages 28–29). Simplified diagrams for the colour patterns of the hair on the dorsum for particular female (f) and male (m) specimens of the species from Fig. 12. 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.

Figs 106–115. Morphology of the male genitalia for species of the subgenus Alpigenobombus Skorikov, 1914, from the dorsal aspect, anterior at the bottom of the image, posterior at the top. 106. Bombus wurflenii Radoszkowski, 1860, Turkey. 107. B mastrucatus Gerstaecker, 1869, Austria. 108. B. kashmirensis Friese, 1909, India-Kashmir. 109. B. rainai Williams, 2022, India-Kashmir. 110. B. sikkimi Friese, 1918, Nepal. 111. B. nobilis Friese, 1905, China-Sichuan. 112. B. validus Friese, 1905, China-Gansu. 113. B. genalis Friese, 1918, China-Yunnan. 114. B. breviceps Smith, 1852, ChinaSichuan. 115. B. grahami (Frison, 1933) China-Sichuan (left penis-valve recurved hook missing). Scale bars = 1 mm.

Fig. 12. Interpretation of filtered MrBayes estimate of phylogeny as a metric tree (outgroup B. festivus Smith, 1861, not shown) for the subgenus Alpigenobombus Skorikov, 1914, from COI-like sequences from the sequences identified in black and orange in Fig. 11 together with Bayesian Poisson-tree-process (PTP) models re-applied for assessing support for species’ gene coalescents by maximum likelihood (for the 11 most likely candidate species). The scale bar is calibrated in substitutions per nucleotide site. Sequence labels and branch colours as in Fig. 10. Numbers above nodes are the Bayesian support values that all daughter haplotypes are parts of a single species. Grey spots show likely mitochondrial-to-nuclear transfers of the low-divergence numts accepted for estimating this tree. Asterisks mark sequences used as informal proxies for the type specimens of each of the taxon names in Table 2. To the right in grey are shown the interpretations of the PTP results as candidate species using the oldest available names for the species.

Fig. 11. Re-interpretation of sequences from Fig. 10, re-drawn with the percentage of nucleotides A and T at codon position 3 (AT3%) for each sequence given after the sequence label. Sequences are classified: (a) in purple – earlier-diverging groups of sequences without unique morphology that often have lower AT3% and shorter sequences that appear to duplicate species as pseudospecies in Fig. 10 – these sequences are interpreted as older, more divergent numts; (b) red – remaining sequences with low AT3% (<95%) – these are interpreted as younger, less divergent numts; (c) orange – sequences from the red group but with the highest AT3% available for some taxa, selected to retain these taxa in the analysis; (d) grey and brown – sequences that are duplicate haplotypes for the black and orange sequences above and often shorter; (e) black – likely orthologous sequences with high AT3% (≥ 95%) and unique haplotypes.

Fig. 10. Default interpretation of MrBayes estimate of phylogeny as a metric tree (outgroup B. festivus Smith, 1861, not shown) for the subgenus Alpigenobombus Skorikov, 1914, from COI-like sequences from GenBank and BOLD databases with additions from the authors, analysed with Bayesian Poissontree-process (PTP) models for assessing support for species’ gene coalescents by maximum likelihood (PTP scores are shown above branches: scores approaching 1, and where branches change from blue to red, indicates where the most likely species’ gene coalescents are identified for 17 candidate species). The scale bar is calibrated in substitutions per nucleotide site. Each sequence is labelled with: sequence length; a morphological taxon name; a code consisting of a sequence identifier from the project database and a specimen identifier from the online database; its country and (for larger countries) state or province of origin.