Bombus rufipes Lepeletier, 1835

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 : 48-49

Bombus rufipes Lepeletier, 1835

Figs 12 , 35–37 , 182

Bombus rufipes Lepeletier de Saint-Fargeau, [1835] : 473.

Bombus flavipes Handlirsch, 1888: 225 .

Bombus rufipes var. [subsp.] obscuripes Friese, 1914: 10.

Bombus rufipes var. [subsp.] intermissus Friese, 1918: 516.

Bremus rufipes var. [subsp.] richardsi Frison, 1930: 6.

Bombus rufipes and B. eximius are now considered parts of the subgenus Melanobombus (see the comments on B. eximius ).

Our PTP analysis ( Fig. 10 ) of coalescents in the COI gene within the rufipes- group supports two species B. eximius and B. rufipes , corroborated by differences in morphology.

From morphology, the black wings, black hair of the male face, and the matching form of the male genital volsella and gonostylus support as conspecific the individuals from Sumatra, which usually (not always) have the mid and hind tibiae orange (taxon obscuripes), together with the individuals from Java, which more usually have the mid and hind tibiae predominantly black (taxon rufipes s. str.). The gonostylus interior proximal process is less strongly bifid for the male from Sumatra. For the male labelled Borneo, the black wings, black facial hair, and form of the volsella and gonostylus all match B. rufipes . For this male, the hair of the mid and hind basitarsi is orange, although the hair of the mid tibia is black and the hair of the hind tibia is orange at the base and black at the tips. A few individuals from Sumatra and Peninsular Malaysia have the hair of the scutellum brown or orange (taxon richardsi). A queen of ‘ostindischen’ origin is described as having T5–6 orange-red (taxon flavipes , not seen).

Diagnosis

Females

Queens large body length 24–26 mm, workers 14–18 mm. Can be distinguished by their combination of the hair of the metasoma entirely black with the wings nearly black with the veins black (cf. B. eximius ), the hair and integument of mid and hind tibiae and of all barsitarsi either orange or black. The labral lamella has the anterior edge broad and nearly straight (cf. B. (Megabombus) senex Vollenhoven, 1873 , B. (Megabombus) melanopoda Cockerell, 1910 , from Sumatra).

Males

Body length 15–18 mm. Can be distinguished by their combination of the hair of the face black with the metasoma entirely black with the wings nearly black with the veins black (cf. B. eximius ), and the thoracic dorsum anteriorly black (cf. B. festivus ). Genitalia ( Fig. 182 ) with the gonostylus reduced to a transverse ‘S’-shaped band (cf. non- rufipes -group), the proximal inner projection broadened distally and bifid (cf. B. eximius ); volsella projecting beyond gonostylus by ca 5 × its breadth at its midpoint (cf. B. eximius ), without an obvious inner distal process or hook (cf. non- rufipes -group); penis valve head with the outer flange greatly expanded as a triangular projection (cf. non- rufipes -group); eye unenlarged relative to female eye.

Material examined

Holotype

INDONESIA • ♀ (worker), holotype of Bombus rufipes Lepeletier, 1835 by monotypy; Java; OUMNH (examined PW from photographs). The original description refers to a single worker with no evidence that the author had additional specimens in front of him, so this worker can be recognised as the holotype

by monotypy. Recognising this specimen as the holotype invalidates the neotype designation by Frison (1930: 11).

Material sequenced (2 specimens)

INDONESIA • 2 ♂♂; East Java, Mt Argopuro ; 7.9756° S, 113.5193° E; 5 Mar. 2010; BOLD seq: 1555E01, 1555E02; SK: ML249, ML250 .

Global distribution

(Southeast Asian mountain species primarily in the southern islands) Southeast Asia: MALAYSIA: Peninsular Malaysia, INDONESIA: Sumatra, Java,? Kalimantan,? Sulawesi. (NHMUK, NHMW, PW, OUMNH, SK.)

It was reported previously (by Williams in Starr 1989) that there is a male labelled “Bandjar/ Borneo” (= Banjar?) in the NHMUK collection (examined PW). The only other bumblebee species recorded apparently from Borneo is the holotype of B. folsomi ( Frison, 1923) , which is labelled “Kina Bala/ N. Borneo” (= Gunung Kinabalu, Sabah). However, the latter specimen appears to be a mislabelled queen of B. (Pyrobombus) ephippiatus Say, 1837 , probably originating from Costa Rica or Panama ( Starr 1989; Williams 1998, examined PW).

The NHMUK “Bandjar/ Borneo” specimen of B. rufipes reached this collection from B. Pittioni’s collection that was assembled in Austria (the specimen carries a det. label from F. Maidl, who was at Vienna’s NHMW museum, and a label “Pittioni Coll./Turner Bequest/ B.M. 1954–79 ”). The NHMW collection has a further three workers and two males with the same locality data in the same handwriting and with the same identification labels. Of nine localities named ‘Bandjar’ identified in Malaysia and Indonesia from US military gazetteers ( USBGN 1968, 1970), one is in Borneo and six are in Java. Frison (1930) interpreted ‘Bandjar’ as referring to Java for a queen, two workers, and the seven males of B. rufipes loaned to him from the ‘Zoological Museum, Buitenzorg, Java’ (= Bogor) collection. Two further workers of B. rufipes without locality labels are listed by Frison that were loaned to him by Maidl from the NHMW. Handlirsch (1891) wrote that all but one of the B. rufipes then in the NHMW collection were from eastern Java (for the other, see below). The occurrence of B. rufipes on Borneo, perhaps on the higher Maratus mountains of south-eastern Kalimantan near Banjar (across the Java Sea from Java and Sumatra), remains possible, but would need to be confirmed.

Two other records appear to extend the known range. One is a queen in the NHMUK from “ Selangor / Bukit Kutu” (Peninsular Malaysia, examined). There is no obvious reason to doubt this record.

Another is a queen in the NHMW collection labelled “N. Celebes / 1908” (= northern Sulawesi, examined PW). The occurrence of B. rufipes in northern Sulawesi would need to be confirmed because it is ca 2000 km from the nearest sites with confirmed records for B. rufipes on Java (see the Discussion). The species is widely distributed but not common in collections.

Behaviour

Some aspects of the behaviour of this species have been described ( Michener & Amir 1977; Kato et al. 1992).

Figures

Figs 12‒13. Maps of sequenced samples. 12. The rufipes-group and festivus-group. 13. The rufofasciatus- group as recognised as species from the UHF-PTP analysis in Fig. 10. Keys to the coloured symbols are shown on the left (in some cases symbols on the map for one species may overlie symbols for another). Maps projected onto a sphere in ArcGIS using World_Shaded_Relief basemap © 2014 ESRI and showing boundaries between countries as recognised by the UN.

Figs 25–63. 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 rufipes-group. 25. Queen, China-Taiwan. 26. Queen, China-Guangdong. 27. Worker, China-Sichuan. 28. Worker, Thailand. 29. Queen, Nepal. 30. Male, China-Sichuan. 31. Male, China-Sichuan. 32. Male, Thailand. 33. Male, Nepal. 34. Male, Nepal. 35. Queen, Indonesia-Sumatra. 36. Queen, Indonesia-Java. 37. Male, Indonesia-Java. The festivus-group. 38. Queen, Nepal. 39. Queen, China-Sichuan. 40. Worker, China- Sichuan. 41. Worker, China-Xizang. 42. Worker, China-Yunnan. 43. Male, Nepal. 44. Male, China- Yunnan. 45. Male, China-Yunnan. The rufofasciatus-group. 46. Queen, India-Kashmir. 47. Worker, India-Kashmir. 48. Male, India-Kashmir. 49. Male, India-Kashmir. 50. Queen, India-Arunachal Pradesh. 51. Queen, Bhutan. 52. Worker, Bhutan. 53. Worker, Bhutan. 54. Male, Bhutan. 55. Queen, India-Kashmir. 56. Queen, India-Kashmir. 57. Worker, India-Kashmir. 58. Worker, India-Kashmir. 59. Worker, India- Kashmir. 60. Worker, Pakistan. 61. Male, India-Kashmir. 62. Male, India-Kashmir. 63. Male, Nepal.

Figs 181‒189. 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. 181. Bombus eximius Smith, 1852, China-Guangxi. 182. B. rufipes Lepeletier, 1835, Indonesia- Java. 183. B. festivus Smith, 1861, Nepal. 184. B. simillimus Smith, 1852, India-Himachal Pradesh. 185. B. miniatus Bingham, 1897, Bhutan. 186. B. eurythorax Wang, 1982 stat. rev., India-Kashmir. 187. B. prshewalskyi Morawitz, 1880 stat. rev., China-Sichuan. 188. B. rufofasciatus Smith, 1852, India- Kashmir. 189. B. richardsiellus (Tkalců, 1968), Burma (left volsella distal part missing). 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.