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Morphometric Analysis and Biogeography of Apis Koschevnikovi Enderlein (1906) S

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Morphometric Analysis and Biogeography of Apis Koschevnikovi Enderlein (1906) S Morphometric analysis and biogeography of Apis koschevnikovi Enderlein (1906) S. Hadisoesilo, Rika Raffiudin, Wirian Susanti, Tri Atmowidi, Colleen Hepburn, Sarah E. Radloff, Stefan Fuchs, H. Randall Hepburn To cite this version: S. Hadisoesilo, Rika Raffiudin, Wirian Susanti, Tri Atmowidi, Colleen Hepburn, et al.. Morphometric analysis and biogeography of Apis koschevnikovi Enderlein (1906). Apidologie, Springer Verlag, 2008, 39 (5), pp.495-503. hal-00891922 HAL Id: hal-00891922 https://hal.archives-ouvertes.fr/hal-00891922 Submitted on 1 Jan 2008 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 39 (2008) 495–503 Available online at: c INRA/DIB-AGIB/ EDP Sciences, 2008 www.apidologie.org DOI: 10.1051/apido:2008029 Original article Morphometric analysis and biogeography of Apis koschevnikovi Enderlein (1906)* S. Hadisoesilo1,RikaRaffiudin2, Wirian Susanti2,TriAtmowidi2, Colleen Hepburn4, Sarah E. Radloff4,StefanFuchs5, H. Randall Hepburn3 1 Forest and Nature Conservation Res. and Dev. Centre, Jl. Gunung Batu, Bogor, Indonesia 2 Department of Biology, Fac. of Mathematics and Natural Sciences, Bogor Agricultural University, Jalan Raya Pajajaran, Bogor, Indonesia 3 Department of Zoology and Entomology, Rhodes University, Grahamstown, South Africa 4 Department of Statistics, Rhodes University, Grahamstown, South Africa 5 Institut fuer Bienenkunde (Polytechnische Gesellschaft), Fachbereich Biologie der J. W. Goethe-Universitaet Frankfurt am Main, Oberursel, Germany Received 12 July 2007 – Revised 3 March 2008 – Accepted 20 March 2008 Abstract – Multivariate morphometric analyses were performed on workers of Apis koschevnikovi from throughout their distribution in Malaysia, Borneo and Indonesia. Principal component analysis showed one morphocluster comprising bees from Kalimantan Indonesia, Sarawak, Sabah and the Malay Peninsula. The population is more homogeneous than A. cerana over the same geographical area, as seen from the average coefficient of variation in 12 characters in A. koschevnikovi (1.8%) compared to those same characters in A. cerana (4.3%). A. koschevnikovi is delimited to the tropical evergreen forest regions of Sumatera, Borneo, and the Malay Peninsula (Fig. 1). The altitudinal distributions show that A. koschevnikovi extends from sea level to about 1600 m. This significantly differs from A. nuluensis but not A. cerana. It appears that the range of A. koschevnikovi is diminishing because it is now either poorly represented or absent in several areas where it has been previously recorded. Apis koschevnikovi / morphometrics / distribution 1. INTRODUCTION Mathew, 1988; Tingek et al., 1988). However, A. koschevnikovi had indeed been widely col- Apis koschevnikovi was originally described lected in the Sundaland region of Southeast by Enderlein (1906) as “Apis indica vari- Asia during the interim as evidenced by col- ety koschevnikovi” and also by von Buttel- lections in various museums (Otis, 1997). Reepen (1906) as “Apis mellifica indica va- In a recent flurry of publications (cf. riety koschevnikovi”. However, authorship for Hepburn and Hepburn, 2007), it was soon es- this species has been formally assigned to En- tablished that A. koschevnikovi is a morphome- derlein (Engel, 1999). With the exceptions of trically distinct species (Tingek et al., 1988; Maa (1953) and Goetze (1964), there have Rinderer et al., 1989; Ruttner et al., 1989; been no other accounts of A. koschevnikovi Sulistianto, 1990), that is reproductively iso- from its original description until its rediscov- lated from (Koeniger et al., 1996), and dif- ery eight decades later in Borneo (Mathew and fers in both nuclear and mitochondrial DNA regions (Arias et al., 1996; Takahashi et al., Corresponding author: S.E. Radloff, 2002; Raffiudin and Crozier, 2007) from other s.radloff@ru.ac.za species of Apis with which it has a sympatric * Manuscript editor: Walter S. Sheppard distribution. Article published by EDP Sciences and available at http://www.apidologie.org or http://dx.doi.org/10.1051/apido:2008029 496 S. Hadisoesilo et al. Here we report the results of multivari- is Otis (1997). Geographical coordinates and ate analyses of morphometric variation in altitudes for identifiable localities in that this species over its entire known range of dataset, plus all localities recoverable from distribution to define population structure and all of the individual research papers (Hepburn provide biogeographical information on and and Hepburn, 2007) and new localities re- distribution maps for A. koschevnikovi in rela- cently found by the present authors for A. tion to A. cerana and A. nuluensis with which koschevnikovi were used to prepare a new dis- it is sympatric. tribution map with GIS software (Fig. 1). The altitudes of all A. koschevnikovi localities and those of sympatric A. cerana (Hepburn and 2. METHODS AND MATERIALS Hepburn, 2006) and A. nuluensis (Fuchs et al., 1996) were obtained and analyzed for pattern 2.1. Honeybees differences. Although most characters of length are some 10–15% greater in worker honeybees of A. koschevnikovi than in A. cerana (Rinderer 2.2. Morphometrics et al., 1989; Sulistianto, 1990), these species may be confused in alcohol-preserved spec- Twenty-seven morphological characters of imens that do not show the natural reddish- worker bees related to size or angles of ve- yellow brightness of the former. Workers of A. nation were measured using the methods of koschevnikovi are one of four medium-sized Ruttner (1988) and the Ruttner parameters and bees and may be quickly distinguished from their Ruttner numbers are given in brackets as them by the cubital index which is 7.64 ± 1.40 follows: length of femur (5), length of tibia in A. koschevnikovi,3.74± 0.24 in A. cerana, (6), metatarus length (7), metatarsus width (8), 4.25 ± 0.47 in A. nigrocincta, and 3.77 ± 0.12 length of tergite 3 (9), length of tergite 4, (10), in A. nuluensis. length of sternite 3 (11), length of wax plate of The worker honeybees used in our study sternite 3, (12) width of wax plate of sternite derive from: (1) data from bees collected 3 (13), distance between wax plates, sternite 3 in Sabah, Borneo (Malaysia), and Sumatera (14), length of sternite 6 (15), width of sternite (Indonesia) from the database of the Insti- 6 (16), forewing length (17), forewing width tut für Bienenkunde at Oberursel, (Obeursel (18), cubital a (19), cubital b (20), wing angle database, n = 13 colonies); (2) new data from A4 (21), wing angle B4 (22), wing angle D7 bees collected from south Kalimantan Borneo (23), wing angle E9 (24), wing angle G18 (25), (Indonesia), (Hadisoesilo-Raffiudin database, wing angle I10 (26), wing angle I16 (27), wing n = 30 colonies), and from the Malay angle K19 (28), wing angle L13 (29), wing an- Peninsula and Sarawak Borneo (Malaysia), gle N23 (30), and wing angle O26 (31). The (Grahamstown database, n = 5 colonies). Col- Grahamstown database consists of the follow- lectively, honeybees from 48 colonies repre- ing twelve morphological characters (5), (6), senting 19 localities in Indonesia and Malaysia (7), (9), (10), (11), (12), (15), (17), B4 (22), were measured morphometrically and statisti- D7 (23) and G18 (25). cally analyzed. In addition, published morpho- metric data on A. koschevnikovi from Rinderer et al. (1989), Sulistianto (1990), and Fuchs 2.3. Data analysis et al. (1996) were included for a complete analysis of this species. Finally, to compare Because the morphometric databases were the average coefficient of variation for A. derived from measurements by different peo- koschevnikovi with that of A. cerana we used ple, to control for possible subjective er- the same A. cerana island database published ror, an analysis of the databases from by Radloff et al. (2005). north Borneo (Oberursel) and south Kali- The largest source of published locality mantan (Hadisoesilo-Raffiudin) was first per- data on the distribution of A. koschevnikovi formed, followed by an analysis of the Morphometric analysis of Apis koschevnikovi 497 Figure 1. Geographical distribution of A. koschevnikovi and A. cerana based on all published records. Localities are given in Table I and additional online material. combined Oberursel, Hadisoesilo-Raffiudin forests in Southeast Asia, A. koschevnikovi and Grahamstown databases. might have been expected to extend well Multivariate statistical analysis of the data into the mainland. However, numerous excur- included principal components, analysis of sions in tropical forests over the last decade variance and Levene’s F statistic procedures in Thailand, Myanmar, Cambodia and Viet- for testing heterogeneity of variances (John- nam failed to find the species (S. Hadisoesilo, son and Wichern, 2002). A Bonferroni ad- H.R. Hepburn, G.W. Otis, M. Pianchaeron, justment to the level of significance was used P.H. Thai, D.W. Roubik, S Wongsiri, all un- in order to ensure that the overall level of publ. data). It appears that A. koschevnikovi significance of the multiple comparison tests is probably restricted to the evergreen forests did not exceed α = 0.05.
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