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Infraspecific Categories of Apis Cerana: Morphometric, Allozymal and Mtdna Diversity H Infraspecific categories of Apis cerana: morphometric, allozymal and mtDNA diversity H. Randall Hepburn, Deborah Smith, Sarah Radloff, Gard Otis To cite this version: H. Randall Hepburn, Deborah Smith, Sarah Radloff, Gard Otis. Infraspecific categories of Apis cerana: morphometric, allozymal and mtDNA diversity. Apidologie, Springer Verlag, 2001, 32 (1), pp.3-23. 10.1051/apido:2001108. hal-00891755 HAL Id: hal-00891755 https://hal.archives-ouvertes.fr/hal-00891755 Submitted on 1 Jan 2001 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Apidologie 32 (2001) 3–23 3 © INRA/DIB-AGIB/EDP Sciences, 2001 Review article Infraspecific categories of Apis cerana: morphometric, allozymal and mtDNA diversity H. Randall HEPBURNa*, Deborah R. SMITHb, Sarah E. RADLOFFc, Gard W. OTISd a Department of Zoology and Entomology, Rhodes University, Grahamstown 6140, South Africa b Department of Entomology, University of Kansas, Lawrence, Kansas 66045, USA c Department of Statistics, Rhodes University, Grahamstown 6140, South Africa d Department of Environmental Biology, University of Guelph, Guelph, Ontario, N1G 2W1, Canada (Received 4 July 2000; accepted 29 September 2000) Abstract – An analysis of the infraspecific categories of Apis cerana was prepared from the relevant literature on taxonomy, morphometrics, allozyme polymorphism and mtDNA diversity. About 31 puta- tive biometric groups have been proposed and assigned to about eight equivocal “subspecies” and var- ious ecotypes. However nearly half of the area of distribution of A. cerana remains unexamined. Allozyme polymorphism is greatest in southeast and lowest in northern and western Asia. About four major mtDNA groups are discernable. There is a very low overall geographic congruity amongst the morphoclusters, allozyme polymorphs and mtDNA clusters. The greatest problems in resolving infraspecific categories in A. cerana are inadequate sampling, incompatible differences in sample sizes, character suites, sampling distance, confidence limits and range of geographical scales employed in different studies. Apis cerana / taxonomy / biogeography / Asia / honeybees 1. INTRODUCTION multivariate probability terms (Hepburn and Radloff, 1998; Ruttner, 1988). Contempo- The history of honeybee classification rary classification of honeybees stems from reflects a slow movement away from the multivariate methods of analysis originally fixed abstractions of the linnaean system to advanced by DuPraw (1964, 1965) and the analysis of population dynamics in substantially developed by Ruttner (1988), * Correspondence and reprints E-mail: [email protected] 4 H.R. Hepburn et al. Ruttner et al. (1978) and Daly (1991, 1992). 2. RESULTS AND DISCUSSION The last decade has been particularly fruit- ful in this regard. Ruttner (1988) completed The natural distribution of A. cerana the daunting task of providing the first mul- based on published citations with reference tivariate analytical attempt at a comprehen- to specific localities is shown in Figure 1. sive macroscale synthesis of honeybee clas- Literature consulted was predominantly that sification for the genus Apis. This was a recoverable from Apicultural Abstracts major impetus for subsequent mesoscale (1950–1999). Positions of localities (closed studies of honeybee morphometrics in circles) are only approximate because of A. mellifera L. (cf. Hepburn and Radloff, map scale; areas in which A. cerana has 1998) as well as for the analysis of allozymic been specifically sought but found absent and DNA diversity of honeybee populations are indicated with stars (Fig. 1). There are (Arias and Sheppard, 1996; Cornuet and several regions where A. cerana undoubt- Garnery, 1991; Smith, 1991; Smith et al., edly occurs but for which there is extremely 1991). sparse or no data at all (Afghanistan, much of India, Laos, Cambodia, Myanmar and In parallel with studies of A. mellifera, Sumatra) and other areas where it has the Ruttner monograph (1988) also opened recently been introduced (Papua New a new chapter in the study of the classifica- Guinea) but these are not considered. tion and biogeography of the honeybees of Asia. This is particularly evident in a recent spate of Asian regional studies in several 2.1. Morphometrics journals and monographs (Verma, 1990, 1992). The purpose of this communication is 2.1.1. Western Asia to report the results of a survey on the pub- lished literature to date on the infraspecific This region extends from the western bor- classification of A. cerana Fabr. throughout its ders of Afghanistan to the north and Pak- entire natural range of some 30 million km2. istan to the south at about longitude 60°, The sympatric occurrence of other Apis thence eastwards below the Himalayan species with A. cerana in southeastern Asia mountain range and across the Indian sub- raises a very undesirable spectre: some pre- continent to Myanmar at about longitude vious “A. cerana” literature could well be 94° (Fig. 2). The extent and quality of infor- contaminated through the inadvertent inclu- mation on the honeybee populations of this 2 sion of data derived from species other than area (about 4 million km ) is extremely vari- A. cerana. The likelihood of detecting such able and ranges from anecdotal descriptions errors seems remote at best. We have to full multivariate statistical analyses of included the results of morphometric stud- morphometric characters. ies as well as those on allozymic and DNA The only information on the classifica- diversity. In the end we present a current tion of the honeybees of Afghanistan and portrait of putative infraspecific categories Pakistan (Fig. 2, area 1) are those of Maa of A. cerana. There were no formal “Mate- (1953) and Ruttner (1988, 1992) who con- rials and Methods” for this study, solely cluded, respectively, that these bees could analyses of the published literature cited in not be morphologically or morphometri- the references. It should be noted that the cally discriminated from those of neigh- approach taken was one of trying to resolve bouring China (classically regarded as distinct groups of A. cerana independently A. cerana cerana (Ruttner et al., 1989). of “correcting” their complex taxonomic However, extremely few samples from this history in terms of the International Code area were available to Ruttner. Further pos- of Zoological Nomenclature (Engel, 1999). sible discrimination of these honeybee Infraspecific categories of Apis cerana 5 Figure 1. Known distribution of A. cerana = (circles); reported absence = (stars). populations is suggested by a report of two well as population structure for the honey- different “kinds” of bees from Afghanistan bees of this region is blurred because of fun- even though the identifications made at the damental incompatibilities in the method- time subsequently proved incorrect (Schnei- ologies and forms of analysis employed in der and Djalal, 1970). Thus, virtually noth- studies published to date (Fig. 2, areas 2–3, ing is known about population structure of 7–11, 14–16). This becomes particularly the honeybees of Afghanistan and Pakistan. evident from comparisons of three impor- Of greater interest, nothing is yet known of tant publications based on the Indian sub- the details of honeybees in eastern Iran continent. Firstly, Kshirsagar (1983) pro- which is geographically the closest point of posed some seven ecotypes for the bees of India vis-a-vis the more usually accepted contact between A. cerana and A. mellifera “hills” and “plains” varieties or ecotypes (Ruttner, 1988; Ruttner et al., 2000). (Kapil, 1956; Narayanan et al., 1960, The honeybees of the sub-Himalayan and 1961a, b; Ruttner, 1988). Secondly, there is Indian regions have been intensively inves- a series of papers by Verma and colleagues tigated in recent years. However, the current (Mattu and Verma, 1983a, b; 1984a, b; picture of biometric groups and ecotypes as Sihanuntavong et al., 1999; Singh et al., 6 H.R. Hepburn et al. Figure 2. Distributional areas of putatively distinct subspecies, biometric groups and/or ecotypes of A. cerana. A. cerana cerana: 1. Eastern Afghanistan and northern Pakistan (by inference only); 2. Kashmir; 3. Himachal Pradesh; 4. China (with biotypes/ecotypes a = Yunnan, b = Guangdong- Guangxi, c = Hunan, d = northern, e = Changbei Shan, f = unspecified, g = Taiwan); 5. Korea; 6. Ussuria. A. cerana himalayana: 7. Nepal Terai plains; 8. Nepal midlands; 9. Himalayas; 10. Brahmaputra; 11. Manipur, Mizoram and Nagaland. A. cerana skorikovi: (possibly = A. cer- ana cerana) 12. Tibet. A. cerana abaensis: 13. Central China. A. cerana indica: 14. Uttar Pradesh; 15. Orissa; 16. Southern India; 17. Sri Lanka (with montane, lowland and Anuradhadpura ecotypes); 18. Yunnan and possibly northern Myanmar; 19. Northern Thailand; 20. Southern Thailand and continental Malaysia; 21. Phuket Island; 22. Samui Island; 23. Sumatra (northern half by inference only), Java, Borneo, Lombok, Bali, Flores and most of Sulawesi; 24. Southern Sulawesi; 25. Timor; 26. Sabah. A. cerana hainanensis: 27. Hainan Island
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