Phylogenetic relationships among bumble bees (Bombus Latr.): a reappraisal of morphological evidence

Abstract. Cladistic analyses of morphological characters of adult bumble bees show significantly higher congruence among characters of the male genitalia than among other characters of males and females. Phylogenetic interpretations of groups supported by the combined morphological evidence (even with minimal assumptions) include (1) that the earliest diverging species belong to the para-phyletic subgenus Mendacibombus , together with Bombus nevadensis and B.confusus ; (2) that, in contrast, species of the subgenus Bombus (sensu stricto) share a relatively recent common ancestor with B.pressus and B.rufipes ; and (3) that Psithyrus (the social parasites) is a monophyletic group, and the sister-group of B.persicus. For genus-group nomenclature a single genus Bombus (sensu lato) is recommended, to include Psithyrus as another subgenus. Names for sections and for groups of subgenera are synonymized with subgeneric names.     

Invasive Bombus terrestris (Hymenoptera: Apidae) parasitized by a flagellate (Euglenozoa: Kinetoplastea) and a neogregarine (Apicomplexa: Neogregarinorida)

The flagellate Crithidia bombi and the neogregarine Apicystis bombi have been found in individuals of Bombus terrestris, a Palaearctic species of bumble bee commercially reared and shipped worldwide for pollination services. B. terrestris has recently entered into the northwestern Patagonia region of Argentina from Chile, where it was introduced in 1998. Prevalence was 21.6% for C. bombi and 3.6% for A. bombi (n = 111). The pathogens were not detected in 441 bumble bees belonging to five of the eight known Argentine native species (Bombus atratus, Bombus morio, Bombus bellicosus, Bombus opifex, Bombus tucumanus) collected elsewhere in the country. Although the absence of natural occurrence of C. bombi and A. bombi in Argentine native bumble bees cannot be ascertained at present due to the limited surveys performed, it is important to report their detection in invasive B. terrestris. The invasion event is relatively recent and the accompanying pathogens are not species specific within the genus Bombus.     

管茎叶蝉属二新种（同翅目：叶蝉科：缘脊叶蝉亚科）

研究了英国卡底阜国立博物馆收藏的缘脊叶蝉亚科脊翅叶蝉族管茎叶蝉属的标本,记述2新种,绘制了特征图。研究标本保存在英国卡底阜国立博物馆。 1. 蒙卡管茎叶蝉Fistulatus monkoboensis, 新种 (图1) 　　体连翅长 ♂6.2 mm。 　　观察标本正模 ♂, BORNEO Sabah, Bukit Monkobo, 51°48′N, 116°58′E, 22-Ⅷ-1987, 1930 m. A. H. Kirk-Spriggs, Light trap sample, Upp. Montane forest summit, Bt. Monkobo, NMW Sabah (Borneo) Expedition, NWW. Z. 1987-094. 　　该新种因其独特的雄性外生殖器结构尾节侧瓣长方形,具一长尾节突腹向伸出,分布有数根大型刚毛而与本属内其它种相区别。 2. 雅典娜管茎叶蝉Fistulatus athena, 新种 (图2) 　　体连翅长 ♂6.2 mm。 　　观察标本正模 ♂, BORNEO Sabah, Bukit Monkobo, 51°48′N, 116°58′E, 18-Ⅷ-1987,1200 m. A. H. Kirk-Spriggs. Light trap sample, camp 1, Stunted hill forest, Bukit Monkobo. NMW Sabah(Borneo) Expedition, NMW. Z. 1987-094. 　　该新种与Fistulatus monkoboensis, sp.n. 在形态特征和雄性外生殖器结构上都相似,与后者主要区别如下 1) 前胸背板无小黑点;2) 尾节侧瓣侧突近端部突然端向收缩;3) 下生殖板基部阔,端部较细,指状;4) 阳基侧突端突长而钝圆,指状;5) 阳茎干端向收缩并更弯曲。     

Phylogenetic relationships among bumble bees (Bombus, latreille) inferred from mitochondrial cytochrome b and cytochrome oxidase I sequences

We conducted a molecular study intending to derive an estimate of the relationships within the genus Bombus (bumble bees) by comparing the mitochondrial cytochrome b and cytochrome oxidase I (COI) genes from 19 species, spanning 10 of approximately 16 European subgenera and 3 subgenera from North and South America. Our trees differ from the most recent classifications of bumble bees. Although bootstrap values for deep branches are low, our sequences show significant data structure and low homoplasy, and all trees share some groups and patterns. In all cases, the subgenus Bombus s. str. clusters among the most derived bumble bees, contrary to other molecular studies. In all trees, B. funebris is the sister taxon of B. robustus, and in five of the six trees, B. wurflenii is the sister taxon to this clade. B. nevadensis is basal to the other species in the analysis of the cytochrome b gene, but appears to be among the most derived according to the analysis of the COI region. The species representing the subgenera Thoracobombus and Fervidobombus are consistently among the earliest diverged. Species that appear in very different positions in different trees are B. nevadensis, B. mesomelas, B. balteatus, and B. hyperboreus. All subgenera with two representatives in our analysis are apparently monophyletic except Fervidobombus, Melanobombus, and Pyrobombus. The groups formed by pocket makers and non-pocket makers within Bombus also appear to be paraphyletic, and therefore some subgenera may not accurately reflect phylogeny.     

The genus Nelcyndana Stål (Hemiptera, Cicadidae, Taphurini) with description of three new species from Borneo

The type species of Nelcyndana, Nelcyndana tener (Stål, 1870) from the Philippines, is redescribed and illustrated. The taxonomic position of the genus Nelcyndana in the tribe Taphurini is discussed. Three new species from Borneo are described and illustrated: Nelcyndana borneensis sp. n., Nelcyndana vantoli sp. n., and Nelcyndana mulu sp. n. Distributions maps for the three Borneo species are presented and a key for the identification of the four Asian species of the genus is provided.     

Review of the odd chrysidid genus Loboscelidia Westwood, 1874 (Hymenoptera, Chrysididae, Loboscelidiinae)

The chrysidid genus Loboscelidia is reviewed and 11 new species are described, including Loboscelidia cinnamonea (Borneo), Loboscelidia fulgens (Viet Nam), Loboscelidia fulva (Thailand), Loboscelidia incompleta (India), Loboscelidia kafae (Borneo), Loboscelidia laminata (Viet Nam), Loboscelidia meifungae (Borneo), Loboscelidia nasiformis (Thailand), Loboscelidia nitidula (Thailand), Loboscelidia pecki (Viet Nam), and Loboscelidia sisik (Borneo). A key to males of the species of Loboscelidia is given.     

Cyanoneuron (Rubiaceae) in Sulawesi? Reaffirmation of a Bornean endemic genus as a cautionary example on the use and interpretation of natural history specimen data

Cyanoneuron, a genus of five species in the Rubiaceae, was described from Borneo and Sulawesi. We show that the occurrence of the genus in Sulawesi was probably derived from careless specimen labeling, and that Cyanoneuron should be considered endemic only to Borneo according to present knowledge. Erroneous collection localities in natural history specimen data can lead to the wrong interpretation of species distribution patterns, so care should be taken to verify specimen records before they are used.     

Interspecific geographic distribution and variation of the pathogens Nosema bombi and Crithidia species in United States bumble bee populations

Several bumble bee (Bombus) species in North America have undergone range reductions and rapid declines in relative abundance. Pathogens have been suggested as causal factors, however, baseline data on pathogen distributions in a large number of bumble bee species have not been available to test this hypothesis. In a nationwide survey of the US, nearly 10,000 specimens of 36 bumble bee species collected at 284 sites were evaluated for the presence and prevalence of two known Bombus pathogens, the microsporidium Nosema bombi and trypanosomes in the genus Crithidia. Prevalence of Crithidia was ≤10% for all host species examined but was recorded from 21% of surveyed sites. Crithidia was isolated from 15 of the 36 Bombus species screened, and were most commonly recovered from Bombus bifarius, Bombus bimaculatus, Bombus impatiens and Bombus mixtus. Nosema bombi was isolated from 22 of the 36 US Bombus species collected. Only one species with more than 50 sampled bees, Bombus appositus, was free of the pathogen; whereas, prevalence was highest in Bombus occidentalis and Bombus pensylvanicus, two species that are reportedly undergoing population declines in North America. A variant of a tetranucleotide repeat in the internal transcribed spacer (ITS) of the N. bombi rRNA gene, thus far not reported from European isolates, was isolated from ten US Bombus hosts, appearing in varying ratios in different host species. Given the genetic similarity of the rRNA gene of N. bombi sampled in Europe and North America to date, the presence of a unique isolate in US bumble could reveal one or more native North American strains and indicate that N. bombi is enzootic across the Holarctic Region, exhibiting some genetic isolation.     

Notes on the genus Bauhinia (Leguminosae-Caesalpinioideae) in SE Asia

During the revision of the genus Bauhinia for Flora Malesiana three undescribed species have been discovered, Bauhinia kostermansii sp. nov. and B. steenisii sp. nov. from Borneo, and B. franckii sp. nov. from the Malay Peninsula. Bauhinia chrysophylla has turned out to be a later homonym, for which reason Bauhinia aureifolia nom. nov. is proposed. The chromosome number of this species has been counted as 2n = 26. B. fabrilis comb, et stat. nov. is proposed for Phanera riedelii var. fabrilis.     

The identity of Siderobombyx and a new species of Xanthophytum (Rubiaceae)

The monotypic genus Sidernbombyx is reduced to Xanthophytum . A new species Xanthophytum bullatum from Borneo, Sarawak, Gunung Mulu National Park is described, the first species in the genus with bullate leaves. An overview of character states is given for the genus Xanthophytum . This covers aspects of vegetative, flower, fruit, seed and pollen morphology. It is shown that the genus Kajewskiellu is very close to Xanthophytum .     

大王马先蒿的传粉综合征状及其生物地理学意义

对云南西北部中甸和昆明黑龙潭的大王马先蒿（ＰｅｄｉｃｕｌａｒｉｓｒｅｘＣｌａｒｋｅＣ．Ｂ）的不同居群进行了传粉生物学观察。大王马先蒿花为黄色,花冠短管、无喙、具二齿、无蜜腺,是一种原始的花冠类型。该种由红腹熊蜂（ＢｏｍｂｕｓｆｒｉｓｅａｎｕｓＳｋｏｒｉｋｏｖ）和明亮熊蜂（ＢｏｍｂｕｓｌｕｃｏｒｕｍＬ．）的工蜂在直立的位置上,通过震动花粉背触式传粉;同时也观察到红腹熊蜂的工蜂在倒悬的位置上,通过震动花粉腹触式传粉。熊蜂的工蜂在大王马先蒿上的这两种传粉综合征状的发现,可能是仅存在于马先蒿原始类型中的一种特殊现象。     

Cytological variation and pathogenicity of the bumble bee parasite Nosema bombi (Microspora, Nosematidae)

In three field seasons, 2003-2005, bumble bees were collected in southern Sweden and eastern Denmark in search of microsporidian parasites. Of the 16 bumble bee species studied, microsporidia were found in Bombus hortorum, Bombus hypnorum, Bombus lapidarius, Bombus lucorum, Bombus pascuorum, Bombus pratorum, Bombus ruderarius, Bombus subterraneus and Bombus terrestris. Only one microsporidian species, Nosema bombi, was recorded. A microsporidium found in B. pratorum differed cytologically from microsporidia of the other host species. In the most frequently infected host, B. terrestris, the prevalence was 20.6%. Totally 1049 specimens were dissected. The light microscopic and ultrastructural cytology and pathology of N. bombi is described with focus on the variation recorded. Variation was especially prominent in the shape, size and coupling of spores, and in the length and arrangement of the polar filament. In four host species microsporidian infection was restricted to peripheral fat cells.     

The Pollination Syndrome of Pedicularis rex (Scrophulariaceae) and Its Biogeographic Significance

The pollination biology of different populations of Pedicularis rex Clarke C B. was studied in Zhongdian of Northwest Yunnan and in Kunming of central Yunnan, respectively. Seed production depends exclusively upon pollination by bmnblebees ( Bombus Latr. ). The flowers are yellow, with short-tubed, erostrate and nectarless corolla. Two species of the bumblebee workers, Bombus friseanus Skorikov and B. lucorum L. foraged in an upright (nototribic) position for pollen. Meanwhile, Bombus friseanus workers also foraged in an inverted (stemotribic) position on P. rex, vibrating pollen from the anthers within the galea. This pollination syndrome was previously reported in the North American species, e.g.P, labradorica which has the same corolla-type with P. rex. A comparison of the corolla types and pollination syndrome on the species of Pediadaris in East Asia and North America indicates that such a pollination syndrome may be only found in the primitive corolla type of Pedicularis. In spite of the separation in the geological history, P. rex and P.labradorica share the same corolla type and pollination syndrome. Further study is needed to elucidate the biogeography of the Pedicularis.     

Identification of Nosema bombi Fantham and Porter 1914 (Microsporidia) in Bombus impatiens and Bombus sandersoni from Great Smoky Mountains National Park (USA)

Ninety three bumble bees belonging to the genus Bombus, subgenus Pyrobombus (three Bombus vagans, seven Bombus bimaculatus, 17 B. sandersoni and 68 B. impatiens) from Great Smoky Mountains National Park were examined for microsporidia. Light microscopy of calcoflour and trichrome-stained smears, and PCR revealed infection with N. bombi in one specimen each of B. sandersoni and B. impatiens. Sizes and shapes of spores in both N. bombi isolates were similar to those described for European isolates of the microsporidium. A region of the rRNA gene from the B. impatiens isolate (1689 bp, accession GQ254295) aligned with homologous sequences from eight European isolates, with only three variable sites. Sequence variability of this region between novel isolates and the European ones was the same as among European isolates.     

<Emphasis Type="Italic">Psithyrus</Emphasis> females do possess wax glands

In behavioural experiments and histological investigations we could demonstrate that the previously widespread generalization that females of the social parasite genus Bombus subgenus Psithyrus do not possess wax glands is not justified. Wax pieces that accumulated on the abdomina of Bombus vestalis, B. sylvestris, and B. bohemicus females which were prevented from self-grooming provided the first evidence for wax production in these species. A histological analysis of epithelial tissues confirmed the existence of epithelial wax glands. Received 17 September 2007; revised 12 March and 2 July 2008; accepted 4 July 2008.     

Cross-species infection of deformed wing virus poses a new threat to pollinator conservation

The Deformed wing virus (family Iflaviridae, genus Iflavirus, DWV), one of the most prevalent and common viruses in honey bees, Apis mellifera L., is present in both laboratory-reared and wild populations of bumble bees, Bombus huntii Greene. Our studies showed that DWV infection spreads throughout the entire body of B. huntii and that the concentration of DWV is higher in workers than in males both collected in the field and reared in the laboratory, implying a possible association between the virus infection and foraging activities. Further results showed that gut tissue of B. huntii can support the replication of DWV, suggesting that B. huntii is a biological host for DWV, as are honey bees. Bumble bees and honey bees sometimes share nectar and pollen resources in the same field. The geographical proximity of two host species probably plays an important role in host range breadth of the virus.     

Proboscis length and resource utilization in two Uruguayan bumblebees: Bombus atratus Franklin and Bombus bellicosus Smith (Hymenoptera: Apidae)

Bumblebees (Bombus sp.) are eusocial insects with an annual life cycle whose use as pollinator of crops has gained great importance in modern agriculture. Several authors have found that resource use in Bombus species is usually based on the correlation between the proboscis length of the bumblebees and the corolla depth of the flowers. The aim of this study was to determine proboscis length of Bombus atratus and B. bellicosus, two Uruguayan bumblebees, and verify the resource exploitation testing two cultivated species, the red clover and the bird's foot trefoil. Bumblebee foraging activity was recorded in two culture conditions: in a red clover and bird's foot trefoil mixed meadow, and in contiguous plots of these legumes, and the proboscis length of collected foragers was determined. Both species of bumblebees exploited red clover and bird's foot trefoil although they did it in different proportions in all instances tested. The results indicated that the choice of the resources in B. atratus and B. bellicosus was influenced by their proboscis length. Bombus atratus has a longer proboscis and preferably visited red clover, possibly obtaining nectar easier and faster than B. bellicosus, which has a shorter proboscis. Bombus bellicosus used both resources without any clear preference.     

A new pollination system: dung-beetle pollination discovered in Orchidantha inouei (Lowiaceae, Zingiberales) in Sarawak, Malaysia

Lowiaceae, a family of the Zingiberales, comprise 11 species in the single genus Orchidantha. Here we present the first report on the pollination of Lowiaceae and describe a new system of dung-beetle pollination from Sarawak, Borneo. Orchidantha inouei has a zygomorphic flower located just above the ground. Observations revealed that the plant is visited frequently and is pollinated by scarabaeid dung beetles, mainly members of the genus Onthophagus. All four species of Onthophagus collected on O. inouei have also been caught using traps baited with dung or carrion in Borneo. Onthophagus was presumably attracted to the dung-like odor of the flower. Pollination of O. inouei is different from other examples of beetle pollination in that its flower provides neither reward nor protected space. Dung beetles are excellent at following a particular dung scent. Orchidantha is the only genus that includes species lacking floral nectar. It is interesting that this deception pollination using dung beetles was found in Zingiberales, in which all known species have mutual and specialized relationships with their long-distance, but costly, pollinators-bees, birds, and bats.     

Distribution and flower visitation records of bumblebees in Lebanon (Hymenoptera: Apidae)

Summary

West Palearctic bumblebees are common wildflowers and crop pollinators that are well studied in their central and northern distribution ranges, but fewer information is available on their southern distribution areas. Lebanon falls on the southern limit of their distribution and no published information is available on the local bumblebees. Our study aims to produce a data baseline of the local bumblebee species. In order to do so we grouped available old records of bumblebees in Lebanon with recent author collections and produced preliminary distribution maps. We listed four species: Bombus terrestris, B. argillaceus, B. niveatus vorticosus and B. melanurus. Preliminary distribution shows that Bombus terrestris and B. argillaceus are widespread and have a large foraging range, whereas B. niveatus vorticosus and B. melanurus have a restricted distribution to altitudes above 1800 m with a smaller foraging range. The male cephalic labial gland secretions analysis of local Bombus terrestris specimens provides preliminary evidence that the local subspecies could be Bombus terrestris calabricus. Therefore, we highlight the importance of regulating foreign Bombus terrestris subspecies importation for agriculture purposes, as well as monitoring B. niveatus vorticosus and B. melanurus that are rendered vulnerable by their isolated populations.     

First Record of the Gekkonid Genus Cnemaspis Strauch, 1887 from Gunung Mulu National Park,Northern Sarawak,East Malaysia may Represent an Undescribed Species

We provide confirmed photographic evidence for the previously overlooked occurrence of the polyphyletic Asian gecko genus Cnemaspis from Gunung Mulu National Park, the world-renowned UNESCO natural heritage site in northern Sarawak, East Malaysia. This new record from Sarawak province represents a remarkable range extension for Cnemaspis cf. kendallii by 550 km to the northeast and denotes the most northern occurrence of the genus in Borneo. Our new finding makes it very likely that these gekkonid lizards also inhabit appropriate limestone karst habitats in adjacent Sabah, Brunei, and Kalimantan. Given the visible differences in the Mulu specimen compared to those from the remaining distribution range on Borneo and the Malaysian Peninsula together with the fact that numerous Cnemaspis species are restricted to small areas, it seems plausible that another undescribed, rather cryptic and possibly locally endemic Bornean species is involved. Lastly, the new record contributes to the importance of the Mulu National Park as a major conservation area in East Malaysia of international concerns.