Colleen Hepburn

Population structure and classification of Apis cerana

Multivariate morphometric analyses of Apis cerana Fabricius, 1793 across its full geographical range were performed. Principal components plots did not reveal distinct morphoclusters. Further substructuring of the principal component plots could not initially be derived but only by introducing local labelling did it reveal six main morphoclusters. We apply geographically based common epithets to the morphoclusters and designate them as: as “Northern cerana”, “Himalayan cerana” “Indian plains cerana” “Indochinese cerana” “Philippine cerana” and “Indo-Malayan cerana”. A. cerana naturally occurs in climatic zones ranging from rainforest, savanna, steppe, grasslands and deciduous forest to taiga. The distributions of the morphoclusters are related to these physiographic and climatic factors. The taxonomy of A. cerana is formally revised and synonymous specific and infraspecific names summarized.ZusammenfassungIn diesem Artikel berichten wir über die Ergebnisse einer über das gesamte Verbreitungsgebiet reichenden multivariaten morphometrischen Analyse von Apis cerana und die statistisch definierten Morphokluster und Subklusterpopulationen innerhalb dieser. (1) Morphokluster I, „nördliche cerana“, erstreckt sich vom nördlichen Afghanistan und Pakistan durch das nordwestliche Indien über das südliche Tibet, das nördliche Myanmar, China und dann nordöstlich nach Korea bis zum fernöstlichen Russland und Japan; (2) Morphokluster II, „himalaya cerana“ schließt die Bienen des nördlichen Indien und einige Regionen des südlichen Tibet und Nepal ein. (3) Morphokluster III „indische Ebene cerana“ besiedelt die Ebenen des zentralen und südlichen Indien und Sri Lanka. (4) In Morphokluster IV, „indo-chinesische cerana“ gruppieren sich die Bienen von Myanmar, Nordthailand, Laos und Südvietnam; (5) Morphokluster V „philippinische cerana“ ist auf die Philippinen beschränkt; (6) Morphokluster VI, hier als „indo-malayische cerana“ bezeichnet, erstreckt sich von Südthailand über Malaysia und Indonesien. Wir stellen die Beziehungen der Morphokluster untereinander und ihre geophysikalischen und ökologischen Umgebungen dar und erstellen eine neue Verbreitungskarte auf Grudlage der gesamten über A. cerana publizierten Literatur (Hepburn and Hepburn, 2006). Einige Anmerkungen zu Genfluss und zeitlicher reproduktiver Isolation werden aus Daten zu Schwärmen und Wanderungen abgeleitet.Nach moderner taxonomischer Praxis sind keine der historisch verwendeten “formellen” lateinischen Namen für die Variationen von Apis passend oder legitim. Daher sollten die Namen unter den Nomenklaturregeln ihrer Zeit gültig erstellten Namen sowie auch die übrigen, überwiegend nach 1970 erstellten Namen als Synonyme angesehen werden. Eine detaillierte synonymische Zusammenstellung im standardisierten taxonomischen Format findet sich bei Engel (1999). Als Konsequenz dieser publizierten Synonyme sind Trinomen wie A. c. japonica oder A. c. cerana unter den Regeln der ICZN keine offiziellen Bestandteile der Klassifikation von Apis mehr. Alle früheren Unterarten sind damit außer Gebrauch. Im gleichen Sinne sind Namen der Morphokluster wie „himalaya cerana“ und „indo-malayische cerana“ wie wir sie hier gebrauchen nicht bindend, sie bieten Bienenwissenschaftlern aber eine weitere Möglichkeit, die zusammenhängenden Populationen von A. cerana sinnvoll und biologisch begründet zu unterteilen. Die ICZN Zusamenfassung der Taxonomie für A. cerana ist hier zusammen mit der formellen Synonymie der Unterartnamen und anderen Namen zusammengestellt, wie es von der Nomenklatur gefordert wird.Zuletzt soll angemerkt werden, dass A. cerana in das nordöstliche China, nach der Ambon Insel, Iran und papua-Neuguinea eingeführt wurde. Von dort hat sie Inseln in der Torresstraße besiedelt (Dunn, 1992) und in neuerer Zeit Neubritannien und die Salomoninseln erreicht (Anderson, 2005 — unpubl. data). A. cerana von Papua-Neuguinea sind morphologisch von denen aus Java nicht unterscheidbar. A. cerana wurde darüber hinaus mehrere Male in Darwin, Brisbane und nahe Perth, Australien aufgegriffen.

Bibliography of the giant honeybees, Apis dorsata Fabricius (1793) and Apis laboriosa F. Smith (1871)

This bibliography of the published literature on Apis dorsata and A. laboriosa was compiled from 988 references written by 934 authors published in 290 periodicals, conference proceedings, theses, reports and books covering the period 1793–2005. The literature shows a balance between the applied aspects of beekeeping and basic honeybee biology for A. dorsata; but for A. laboriosa it reflects more basic biology.

Morphometric analysis and biogeography of Apis koschevnikovi Enderlein (1906)

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.ZusammenfassungAn Arbeiterinnen von Apis koschevnikovi aus deren gesamten Verbreitungsgebiet in Südostasien wurden multivariate morphometrische Analysen durchgeführt. Siebenundzwanzig morphologische Merkmale der Arbeiterinnen wurden nach der Methode von Ruttner (1988) vermessen, die verwendeten Merkmale enthielten Größenmessungen und Flügelwinkel. Hieraus wurden acht Hauptkomponenten abgeleitet, die insgesamt 78,7 % der Variation in den Daten repräsentierten. Die multivariaten Analysen der Proben von A. koschevnikovi zeigten klar, dass die Art aus einem einzigen anhand von nur 12 morphologischen Merkmalen abgrenzbaren Morphokluster zusammengesetzt ist.Die Hauptkomponentenanalyse zeigte einen die Bienen von Sumatra, Borneo und der malayischen Halbinsel enthaltenden Morphokluster (Abb. 2 und 3). Die Population ist einheitlicher als die von A. cerana in dem gleichen Verbreitungsgebiet. Dies kann aus dem Vergleich der mittleren Varianzkoeffizienten von 12 gleichen Merkmalen ersehen werden, der bei A. koschevnikovi 1,8 %, bei A. cerana aber 4,3 % beträgt.Die Höhenverteilung zeigt, dass von den 102 erfassten Fundorten von A. koschevnikovi 96 % niedriger als 1200 m und 4 % zwischen 1200 und 2700 m lagen (Tab. I und zusätzliches elektronisches Onlinematerial zur Verteilung von A. koschevnikovi auf Grundlage aller publizierter Nachweise). Diese Höhenverteilung ist nicht unterschiedlich von der sympatrischen A. cerana (χ2 = 6,9, df = 3, P = 0,0764). Sie ist aber signifikant verschieden von der Höhenverteilung der geographisch sympatrischen A. nuluensis, für die bisher nur drei Fundorte bekannt sind, die alle über 3000 m liegen (χ2 = 104,0, df = 3, P < 0,0001).In zahlreichen Exkursionen im tropischen Regenwald über die letzten 10 Jahre in Thailand, Myanmar, Kambodscha and Vietnam konnte die Art A. koschevnikovi nicht nachgewiesen werden, ihr Vorkommen ist auf die Region des immergrünen Regenwaldes von Sundaland begrenzt (Abb. 1). Anscheinend ist das Verbreitungsgebiet von A. koschevnikovi im Schwinden, da diese in einigen Gebieten, in denen sie früher gefunden wurde, nun nur geringfügig vertreten ist oder vollständig fehlt.

Determining spatio-temporal distribution of bee forage species of Al-Baha region based on ground inventorying supported with GIS applications and Remote Sensed Satellite Image analysis

In arid zones, the shortage of bee forage is critical and usually compels beekeepers to move their colonies in search of better forages. Identifying and mapping the spatiotemporal distribution of the bee forages over given area is important for better management of bee colonies. In this study honey bee plants in the target areas were inventoried following, ground inventory work supported with GIS applications. The study was conducted on 85 large plots of 50 × 50 m each. At each plot, data on species name, height, base diameter, crown height, crown diameter has been taken for each plant with their respective geographical positions. The data were stored, and processed using Trimble GPS supported with ArcGIS10 software program. The data were used to estimate the relative frequency, density, abundance and species diversity, species important value index and apicultural value of the species. In addition, Remotely Sensed Satellite Image of the area was obtained and processed using Hopfield Artificial Neural Network techniques. During the study, 182 species from 49 plant families were identified as bee forages of the target area. From the total number of species; shrubs, herbs and trees were accounting for 61%, 27.67%, and 11.53% respectively. Of which Ziziphus spina-christi, Acacia tortilis, Acacia origina, Acacia asak, Lavandula dentata, and Hypoestes forskaolii were the major nectar source plants of the area in their degree of importance. The average vegetation cover values of the study areas were low (<30%) with low Shannon’s species diversity indices (H′) of 0.5–1.52 for different sites. Based on the eco-climatological factors and the variations in their flowering period, these major bee forage species were found to form eight distinct spatiotemporal categories which allow beekeepers to migrate their colonies to exploit the resources at different seasons and place. The Remote Sensed Satellite Image analysis confirmed the spatial distribution of the bee forage resources as determined by the ground inventory work. An integrated approach, combining the ground inventory work with GIS and satellite image processing techniques could be an important tool for characterizing and mapping the available bee forage resources leading to their efficient and sustainable utilization.

Bibliography of the Asian Species of Honeybees

This bibliography of the literature on species of honeybees native to Asia is an extension of the recently published bibliographies on Apis andreniformis (Hepburn and Hepburn 2009), Apis cerana (Hepburn and Hepburn 2006), Apis dorsata and Apis laboriosa (Hepburn and Hepburn 2007a), Apis florea (Hepburn and Hepburn 2005) and Apis koschevnikovi (Hepburn and Hepburn 2007b). It was compiled from nearly 3,550 references, written by about 3,000 authors and published in over 700 different periodicals, conference proceedings, theses, reports, books and patents covering the period 1787–2009. The bibliography was prepared by obtaining all references to Asian honeybees published in Zoological Record (1864–2003) and Apicultural Abstracts (1950–2004) as well as the species-specific bibliographies of other workers cited in the Hepburn bibliographies. Subsequently, copies of most of the original publications were obtained, and the references cited therein were traced in continuous iterations into the past until no new references were found. For the period 2005–2009, particular use was made of the website Google Scholar, which is supported by various other citation websites, the most reliable of which are the British Library (BL Direct), US National Library of Medicine and the National Institutes of Health (http://www.pumed.gov), Institut de l’Information Scientifique et Technique of the Centre National de la Recherche Scientifique (INIST-CNRS) and Ingenta, a division of Publishing Technology plc (http://www.ingentaconnect.com). Recent references in Chinese are available at the website Wanfang Data, an affiliate of the Chinese Ministry of Science and Technology, but unfortunately, the citations are incomplete. Oddly enough, Google often gives more download choices for citations of the same article than does Google Scholar. In some cases, incomplete citations (particularly in Chinese) are given because complete information could not always be obtained, and they are marked with an asterisk. Similarly, several citations were

Bibliography of Apis andreniformis Smith, 1858

Commentary – This bibliography of the literature on A. andreniformis was compiled from 122 references, written by 59 authors and published in 48 different periodicals, conference proceedings, theses, reports and books covering the period 1865–1 August 2008. The literature is entirely confined to basic biology. Of an estimated 4000 publications on Asian Apis, that of A. andreniformis constitutes only 3%. That over 98% of all publications on this species have appeared over the past 25 years attests to a growing interest in A. andreniformis. Accessing this small literature is facilitated by their capture on the website Google Scholar c ©. Virtually all of them are in English. Given the widespread occurrence of absconding and migration in A. andreniformis, it is perhaps surprising that these phenomena have not been investigated more intensively. It is difficult to draw many conclusions about human demography and publications on A. andreniformis, but numerically, the literature is derived mainly from studies in Thailand and China and in western institutions. The bibliography was prepared by obtaining all references to Asian honeybees published in Apicultural Abstracts (1950–2005) and Zoological Record (1864–2003). Subsequently, copies of the original publications were obtained and the references cited in these works were traced in continuous iterations into the past until no new references were found. Each reference was entered in a computer database and keywords for subject matter and countries for all publications were noted. On completion of the database the original 254 keywords were compressed into 36 more inclusive keyword categories.

Bibliography of Apis koschevnikovi Enderlein (1906)

Commentary – Apis koschevnikovi was originally described a century ago independently by both Enderlein and von Buttel-Reepen in 1906. With the two exceptions of Maa (1953) and Goetze (1964), there is no other mention of A. koschevnikovi in the apicultural literature until its rediscovery in Borneo some 80 years later (Mathew and Mathew, 1988; Tingek et al., 1988). These reports provided the stimulus for subsequent research and the resulting publications, which constitute 96% of the total 102 for A. koschevnikovi. Some 93% of these publications are in the English language and because they have appeared within the last 20 years, they are readily accessible as well. Our knowledge of even the basic biology of this species is very rudimentary and all of the publications on this species are directed to this end.