Bombus (Alpigenobombus) kashmirensis Friese, 1909

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 : 33-35

3. Bombus (Alpigenobombus) kashmirensis Friese, 1909

Figs 2–3 , 35‒48 , 108

Bombus mastrucatus var. [subsp.] kashmirensis Friese, 1909 [September: Tkalců 1974]: 673. Note 1.

Bombus mastrucatus var. [subsp.] stramineus Friese, 1909 [September: Tkalců 1974]: 673.

Bombus tetrachromus Cockerell, 1909 [November: Tkalců 1974]: 397.

Alpigenobombus pulcherrimus Skorikov, 1914: 128 .

Bombus (Mastrucatobombus) mastrucatus subsp. meinertzhageni Richards, 1928: 335 .

Bombus mastrucatus var. [subsp.] kashmirensis ‒ redescribed by Friese & von Wagner 1910: 47.

Bombus mastrucatus var. [subsp.] stramineus ‒ redescribed by Friese & von Wagner 1910: 47.

Alpigenobombus pulcherrimus var. [not subsp.] albidocaudatus ‒ Skorikov 1914: 129, infrasubspecific.

NOTE 1. Acceptance of B. kashmirensis as the valid name in preference to B. stramineus follows Skorikov (1923) as the First Reviser ( ICZN 1999: Article 24).

Species-taxon concept and variation

The taxon concept of the species B. kashmirensis here agrees with the recent interpretation (Williams 2022a) that it is separate from the taxon concept of the species B. rainai (see the comments below on B. rainai ), 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 the divergent colour patterns of the taxa kashmirensis s. str., stramineus , and meinertzhageni are parts of a single species ( Williams 1991). The other named taxa ( tetrachromus , pulcherrimus ) appear to be re-descriptions of this species.

Variation in the colour-pattern diagrams of B. kashmirensis in Figs 35‒48 is arranged approximately from the Hengduan and outer Himalayan ranges (e.g., Pir Panjal: Figs 35‒37, 42‒44) to the trans-Himalaya (e.g., Zanskar: Figs 39‒41, 46‒48). Bombus kashmirensis , with the white-banded colour pattern in the south (Himalaya) and east (Hengduan) more mesic parts of its range, appears to mimic the abundant B. (Melanobombus) rufofasciatus Smith, 1852 , and B. (Ml.) prshewalskyi Morawitz, 1880 ( Williams 2007: fig. 5g), whereas the yellow-banded colour pattern in the north-west (trans-Himalayan mountains) more arid parts of its range appears to mimic the abundant B. (Ml.) keriensis Morawitz, 1887 , and similar species ( Williams 2007: fig. 5j).

Type material

Bombus mastrucatus var. [subsp.] kashmirensis Friese, 1909 [September: Tkalců 1974]: 673. Lectotype by designation of Tkalců 1974: ♀ (queen) Kashmir , India ( ZMHB). Examined .

Morphological diagnosis

Female

Wings nearly clear with veins dark brown, hair medium to long, oculo-malar area shorter than broad, clypeus adjacent to the labrum slightly raised in the centre so that the transverse anterior groove is interrupted, this central area with few small punctures and more shining than the surrounding areas, oculo-ocellar area in almost its outer half with large and medium punctures (cf. B. rainai ); hair of the side of the thorax in at least its upper half and the scutellum either white or yellow, hair of T5 orange with white tips.

Male

Wings nearly clear with veins dark brown, hair medium to long, oculo-malar area shorter than broad; genitalia ( Fig. 108 ) with the gonostylus long and distally near its midline axis convexly rounded, length on its outer side about a quarter as long as on its inner side with the two inner corners of the distal lobe rounded, the margin between them nearly straight, penis-valve head strongly recurved, the recurved hook much longer than broad, but broad and narrowing only just before the apex (cf. B. rainai ); male eye distinctly enlarged relative to female eye; hair of the side of the thorax either yellow or white with the pale hair reaching to the midleg bases, hair of T3 usually predominantly black, hair at the side of T5‒6 orange at the base but white-tipped.

Material sequenced in Fig. 12

CHINA • 1 ♀ (worker); Sichuan, Queer Mountain ; 31.8969° N, 99.1485° E; 5 Aug. 2018; Z. Ren et al. leg.; KIB seq: QESM302006; KIB: AG#083 • 1 ♀ (worker); Xizang, Ailashan ; 31.6142° N, 98.4803° E; 4 Aug. 2018; Z. Ren et al. leg.; KIB seq: ALSM104001; KIB: AG#085 • 1 ♀ (worker); Qinghai, Kunlun Pass road; 35.6917° N, 94.0533° E; 19 Aug. 2013; PW seq: PWK1; IAR: AG#084 .

PAKISTAN • 1 ♁; Baltistan [Olthingthang]; 34.7093° N, 76.1208° E; 16 Sep. 2016; S. Jaffar leg.; BOLD seq: 1555G02; PW: AG#082 .

Additional sequences in Fig. 10 and haplotype duplicates

CHINA • 1 ♀ (worker); Xizang, Yelanshan ; 30.1489° N, 97.3091° E; 22 Jul. 2018; Z. Ren et al. leg.; KIB seq: YLSH308001; KIB: AG#130 .

Global distribution

Himalaya and Hengduan: Pakistan, India (Kashmir, Himachal Pradesh, Uttarakhand, Sikkim), Nepal, China (Xizang, Qinghai, Sichuan, Gansu): IAR, IOZ, KIB, NHMUK, PW, RMNH, SC, ZMHB.

This species is recorded at elevations of 2400‒4800 m in the Himalaya and trans-Himalaya, and of 2272‒4700 m in the Hengduan mountains ( Williams 1991; Williams et al. 2009, 2010; An et al. 2014). In Kashmir, Sichuan, and Gansu, B. kashmirensis occurs primarily above the tree line and high into the alpine zone (queens have been seen to forage from flowers at 4000 m in Kashmir even when it was snowing heavily, PW pers. obs.) and even in the semi-arid trans-Himalaya ( Williams 1991; Williams et al. 2009).

Behaviour

Male eye slightly enlarged relative to female eye: males show ‘cruising’ behaviour when searching for mates in Kashmir ( 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. 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.