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Apis mellifera pomonella, a new honey bee subspeciesfrom Central Asia

Walter Sheppard, Marina Meixner

To cite this version:Walter Sheppard, Marina Meixner. Apis mellifera pomonella, a new honey bee subspecies from CentralAsia. Apidologie, Springer Verlag, 2003, 34 (4), pp.367-375. �10.1051/apido:2003037�. �hal-00891797�

367Apidologie 34 (2003) 367–375© INRA/DIB-AGIB/ EDP Sciences, 2003DOI: 10.1051/apido:2003037

Original article

Apis mellifera pomonella, a new honey bee subspecies from Central Asia

Walter S. SHEPPARDa, Marina D. MEIXNERa,b

a Department of Entomology, 166 FSHN Building, Washington State University, Pullman WA 99164-6382, USAb Institut für Bienenkunde, 2 Karl von Frisch Weg, 61440 Oberursel, Germany

(Received 16 August 2002; revised 7 December 2002; accepted 2 January 2003)

Abstract – Endemic honey bees of the Tien Shan Mountains in Central Asia are described as a newsubspecies, Apis mellifera pomonella, on the basis of morphometric analyses. Principal component anddiscriminant analysis of the morphological characters measured clearly place these bees into the orientalevolutionary branch of honey bees, but also show that they are distinct from the other subspecies in thislineage. The existence of this newly described honey bee subspecies extends the range of endemic A.mellifera more than 2000 km eastward than previously estimated. Sequence analysis of mitochondrial DNAplaces A. m. pomonella within the C mitochondrial lineage (a group that is inclusive of both C and Omorphological lineages). These findings support the conclusion that A. m. pomonella has a phylogeographichistory shared with subspecies from the eastern limit of the previously known range.

Apis mellifera pomonella / new subspecies / Central Asia / taxonomy

1. INTRODUCTION

The endemic distribution of the honey beeApis mellifera L. is generally considered toencompass Africa, Europe and portions ofwestern Asia. Across this range, variation inbehavior, morphology and genetic markerssupports an evolutionary history of the speciesthat includes differentiation into several majorphylogenetic lineages (Ruttner, 1988; Cornuetand Garnery, 1991; Garnery et al., 1992).Based primarily on morphological characters,more than two dozen subspecies have beendescribed within the lineages (Ruttner, 1992;Sheppard et al., 1997). These subspecies typi-cally exhibit reduced gene flow with othersuch groups due to water, mountain or desertbarriers and have been called “geographicraces”, to reflect their adaptation to specificgeographic areas (Ruttner, 1988).

Based on morphological similarities andsequence divergence among described subspe-cies, the speciation event that produced Apismellifera has been estimated to have occurredbetween 0.7 to 1.3 million years ago (Ruttner,1988; Cornuet and Garnery, 1991; Arias andSheppard, 1996). While Apis mellifera occu-pies a large geographic distribution allopatricfrom the rest of the genus, a number of theother Apis species reside sympatrically inAsia. Western Afghanistan is considered to bethe eastern limit of A. mellifera, with the clos-est proximity between Apis mellifera and itscongeners occurring somewhere in centralAfghanistan (Ruttner, 1988). Consistent withthe estimated antiquity of A. mellifera and thepaleoclimate of the region separating A. cer-ana and A. mellifera, Ruttner postulated thatan A. mellifera ancestor reached western Asiaabout one million years ago (ibid).

* Correspondence and reprintsE-mail: shepp@mail.wsu.edu

368 W.S. Sheppard, M.D. Meixner

While the relative youth of a million-year-old A. mellifera lineage has been supportedby molecular studies, cladogenesis of A. mel-lifera and A. cerana appears to have occurredat a much earlier time (6–9 mya) based onallozyme and DNA sequence differences(Sheppard and Berlocher, 1989; Cornuet andGarnery, 1991; Arias et al., 1996). The appar-ent discrepancy between the age of A. mellif-era subspecies and the A. mellifera/A. ceranacladogenesis suggests that alternative hypoth-eses for the origin of A. mellifera should beconsidered. These include the possibility thatA. cerana and A. mellifera are not related toeach other as sister taxa, rather they may be sorelated to as yet undescribed or extinct species.Unfortunately, perhaps the least studied areawithin the distribution of Apis is in the regionbetween the known distribution of Apis mellif-era and Apis cerana.

To further address this and other issuesrelated to the evolution of A. mellifera,improved sampling of honey bee populationsin regions of central Asia was undertaken. Wereport here the results of an investigationcharacterizing honey bees from the Tien ShanMountains of Kazakhstan, extending from1300–2000 km east of the previously knownrange of Apis mellifera.

2. MATERIALS AND METHODS

The Tien Shan Mountains lie in an east-westorientation and traverse some 1000 km from south-central Kazakhstan to western China. Adult honeybees were collected from three locations in the TienShan range: (1) in the western Tien Shan – twoapiaries in the village of Jabagly and 5 feralcolonies from the nearby Aksu-Jabagly NatureReserve (2) in the east-central Tien Shan, an apiarynear the village of Kurmetty and (3) an apiary nearthe village of Tujuk in the Ketman range (a branchof the eastern Tien Shan) some 100 km from theborder of China and 750 km from the Jabaglycollection. The sampling area in relation to thenatural range of A. mellifera and A. cerana is shownin Figure 1.

Feral colonies in the Jabagly reserve were foundat 2000 m and were either “caught swarms” inboxes (4) or a wild nest in a tree (1). In the Jabaglyand Kurmetty apiary locations, beekeepers reportedthat no imported honey bees were introduced intothe areas for more than 20 years. In the apiarysample taken near Tujuk, there was no recollectionby local villagers that bees had ever been introducedand the apiary was composed of the descendents ofwild swarms caught in previous years.

2.1. Morphometric analysis

Samples were preserved in 90% ethanol formorphometric analysis. A total of 31 samples were

Figure 1. Areas of distribution of Apis mellifera and A. cerana (adapted from Ruttner, 1988). The collectionlocations in Central Asia are indicated as A (Aksu-Jabagly), K (Kurmetty), and T (Tujuk).

A. m. pomonella, a new honey bee subspecies from Central Asia 369

subjected to morphometric analysis (15 each fromJabagly and Tujuk, 1 from Kurmetty). Fifteenworker bees per sample were dissected and meas-ured for 39 morphometric characters according toRuttner et al. (1978). Wing venation angles andcharacters of size were measured using a CCD cam-era combined with an on-screen measuring system(Meixner, 1994). Characters of pigmentation andpilosity were measured with a microscope and anocular micrometer. The statistical analysis of thedata was performed with SPSS for Windows andStatistica computer programs.

Multivariate statistical analyses were performedon the samples including all measured morphomet-ric characters except width of tomentum and darkstripe on tergum 4, length of proboscis, pigmenta-tion of tergite 2, and length of tergite 4, which wereexcluded due to missing data. Data of referencesamples for multivariate statistics were includedfrom the database of the Institut für Bienenkunde,Oberursel. The holotype has been deposited in theM.T. James Entomological Museum at WashingtonState University, Pullman, WA and paratypeshave been deposited in the M.T. James Museum andin the collection at the Institut für Bienenkunde,Oberursel, Germany.

2.2. Extraction of DNA, PCR amplification and sequencing

Total DNA was extracted from five samples(two each from Aksu-Jabagly and Tujuk, and onefrom Kurmetty) following procedures described inArias and Sheppard (1996). We used the polymer-ase chain reaction to amplify a mitochondrial DNAregion that encompasses the tRNA ILE and part ofthe ND2 gene. The primers ILE and L1 and reaction

conditions are described in Arias and Sheppard(1996). PCR products were run on 1.5% agarosegels, stained with ethidium bromide, and photo-graphed under UV illumination.

Amplification products were directly sequencedusing the cycle sequencing protocol (Craxton,1991) and an ABI 377 automated sequencer. Corre-sponding sequence data from other Apis melliferasubspecies (Arias and Sheppard, 1996) werealigned with 645 bp of amplified sequence from theKazakhstan samples using Clustal V (Higgins andSharp, 1988).

3. RESULTS

3.1. Morphometric analysis

To determine the position of the samplescollected from the Tien Shan in the morpho-logical tripod structure described by Ruttner(1988, 1992), an initial principal componentanalysis including reference samples from allsubspecies of A. mellifera was performed. InFigure 2, the result of this analysis is presentedin a three-dimensional plot. Along factor 1,mainly characters of size are included, whilefactors 2 and 3 contain many variables inde-pendent of size. Subspecies from Africa(diamonds) are positioned along factor 1, withsubspecies belonging to the other three line-ages (Ruttner, 1992) being more diverse infactors 2 and 3. The samples from the TienShan mountains (stars) form a compact clusterassociated with the Near Eastern lineages ofsubspecies.

Figure 2. Positions of referencesamples from all recognized sub-species of A. mellifera and thesamples collected in the TienShan Mountains in a three-dimensional principal compo-nent analysis. Symbols representindividual colony samples.

370 W.S. Sheppard, M.D. Meixner

While this analysis provides a first impres-sion of the homogeneity of the Tien Shan beesand their position within the oriental branch ofthe morphometric structure of Apis mellifera,it does not provide much information about therelationships of the Tien Shan bees to sur-rounding subspecies. However, this resultdemonstrates that the bees collected in theTien Shan mountains are clearly not associ-ated with A. mellifera subspecies of the M or Clineage that might have been transported intothe area by migratory beekeeping practices inthe former Soviet Union.

Subsequently, a principal component anal-ysis including only the subspecies of theC- and O-branch was performed. In Figure 3,factor 1 of this analysis (accounting for 28% ofthe total variance) is plotted against factor 2(20%). In this analysis, the Tien Shan bees arealso clearly positioned among the lineage ofthe Near East, and not with the subspeciesfrom southeast Europe.

Figure 4 shows a discriminant analysis ofthe same data set as in Figure 3. In Figure 4,discriminant factor 2 (24% of total variance) isplotted against factor 1 (48%). This analysisfurther supports the conclusion that the honeybees from the Tien Shan are part of theOriental lineage of honey bee subspecies,without noticeable influence of commerciallyimportant subspecies from southeast Europe.

Within the Oriental branch, they form adistinctive cluster. Figure 5 shows the positionof the Tien Shan samples in a discriminantanalysis of the Oriental lineage. In thisdetailed analysis, the Tien Shan bees form acompact cluster that is distinct from all othersubspecies included, mainly due to charactersof wing venation in discriminant factor 2.

Based on the distinctiveness of the TienShan honey bee across its range from othersubspecies and the similarity of thepopulations sampled across more than 750 km,we conclude that the Tien Shan honey beedeserves subspecific rank and propose thename, Apis mellifera pomonella. The typelocality for A. m. pomonella in the Aksu rivervalley of Kazakhstan is shown in Figure 6.

3.2. Description of the Tien Shan honey bees

A. mellifera pomonella is a large bee, onlyslightly smaller than A. m. carnica and A. m.caucasica. It is in general very similar to A. m.anatoliaca, but has a broader abdomen andnarrower tomenta. It has comparatively broadtarsi, as described to be typical for the Easternlineage of honey bees by Ruttner (1988, 1992).Compared to its relatives A. m. caucasica andA. m. armeniaca, the proboscis of this bee isrelatively short.

Figure 3. Position of honey bee samples in a principal component analysis performed on subspeciesbelonging to the C- and O-lineage (Ruttner, 1988) of A. mellifera. Factor 1 is plotted against factor 2. Eachsymbol represents one colony sample.

A. m. pomonella, a new honey bee subspecies from Central Asia 371

Although the samples were collected inaltitudes up to 2000 m, the bees have compar-atively short cover hair on the abdomen,reaching a mean of only 0.27 mm. Their pig-mentation scores are intermediate, similar toA. m. anatoliaca. Mean values and standarddeviations of some morphometric charactersare summarized in Table I.

Beekeeping characteristics of A. m. pomo-nella appeared to be typical for a temperatemountain honey bee. Although only 25 colo-nies were opened for brood nest inspectionduring the month of June 2000, the bees werefound to be quite gentle and calm on the combsduring manipulation. A preliminary study of aJabagly village apiary in June 2000 revealedthat pollen forager activity (foragers returningto colonies with pollen loads) took place at atemperature threshold of 10.8 °C (Sheppard,unpublished data).

3.3. Sequence analysis of mitochondrial DNA

DNA sequences from the 5 samples werehighly similar, exhibiting 0.75% sequencedivergence. Sequences were deposited in Gen-bank under accession numbers AY135560,AY136622, AY136623, AY136624 andAY136625.

The mean sequence divergence between theTien Shan honey bees and other subspecies inthe C lineage was less than 1.0% (A. m.carnica, A. m. macedonica, A. m. caucasica,range 0.16%–1.09%), while the divergencesfrom the A-lineage and M-lineage were 1.52%(A. m. scutellata, range 1.10%–1.92%) and1.65% (A. m. mellifera, range 1.26%–2.08%),respectively.

Figure 4. Positions of the same samples as in Figure 3 in a discriminant analysis. Abscissa: discriminantfactor 1, ordinate: discriminant factor 2.

Figure 5. Positions of samples in a discriminantanalysis performed on honey bee subspeciesbelonging to the morphological O-lineage (Ruttner,1988) and the collection from the Tien Shanmountains.

372 W.S. Sheppard, M.D. Meixner

Inclusion of DNA sequence data from theTien Shan bees in a phylogenetic analysisusing either Parsimony or Maximum Likeli-hood methods did not change the overalltopology of the phylogenetic tree publishedin Arias and Sheppard (1996). The five sam-ples sequenced were incorporated into theC-branch subspecies group without improvingthe resolution within this branch (tree notshown).

4. DISCUSSION

The existence of this newly describedhoney bee subspecies extends the range ofendemic A. mellifera more than 2000 km east-ward than previously estimated. Further, it islikely that A.m. pomonella populations will befound at least to the eastern limit of the TienShan range in far western China (approxi-mately 100–150 km farther east than thepresent Tujuk samples). Evidence for suchwidespread distribution of A. m. pomonellacomes from the fact that samples from theeastern and western Tien Shan Mountainsform a tightly grouped morphological clusterin our analyses. This cluster is highly distinc-tive and separate from European and westAsian subspecies that were brought into agri-cultural areas of Uzbekistan and Kazakhstanfor managed beekeeping.

The finding of an endemic honey bee in thisarea is perhaps not surprising, given theimportance of the mountainous areas of Cen-tral Asia as a center of diversity for apples,

pears and apricots. The area of the Tien ShanMountains near Almaty (formerly known asAlma-Ata or “father of apples”), Kazakhstan,is the area of greatest genetic diversity for awild Malus species, M. sieversii, that is thepredominant ancestor of domesticated applevarieties (Hokanson et al., 1997). As a horti-cultural crop, apples are self-incompatible andtypically pollinated by honey bees on a com-mercial scale. Although a careful study of thepollinating insects of the wild apple forests ofKazakhstan is lacking, the sympatric distribu-tion of the new subspecies with wild applesthroughout the sampled areas of the Tien Shanin this study provide the basis for the trinomialepithet pomonella, derived from the Romandeity Pomona, protector of gardens and fruittrees.

Sequence analysis places A. m. pomonellaclearly within the C mitochondrial lineage(which is inclusive of both the C and Omorphological lineages, sensu Ruttner, 1988),a finding consistent with the morphometricresults. This supports the conclusion that A. m.pomonella has a phylogeographic historyshared with subspecies from the eastern limitof the previously known range. However,the question is raised as to whether A. m.pomonella represents an eastward expansionof the morphological O lineage that likelyoriginated somewhere in the region betweenIran and the Mediterranean Coast (Ruttner,1988; Franck et al., 2000). Alternatively, theTien Shan Mountains and unknown pointseastward may represent the area of origin and

Figure 6. Type location for A. m.pomonella in the Aksu rivervalley, approx. 30 km southwestof the village of Jabagly,Kazakhstan. Holotype was col-lected from a wild colony foundin a juniper tree on the slope inthe left side of the photograph.Latitude 42 20 70 N, Longitude70 22 39 E. View is from 2000 melevation.

A. m. pomonella, a new honey bee subspecies from Central Asia 373

Tab

le I

. C

ompa

rativ

e ch

arac

teri

stic

s of

A.

m.

mel

life

ra,

A.

m.

carn

ica,

A.

m.

ligu

stic

a, A

. m

. ca

ucas

ica,

A.

m.

arm

enia

ca,

A.

m.

med

a, A

. m

.an

atol

iaca

and

A.

m.

pom

onel

la.

Val

ues

are

mea

ns a

nd s

tand

ard

devi

atio

ns o

f sa

mpl

es,

each

sam

ple

repr

esen

ting

one

col

ony.

N =

num

ber

ofsa

mpl

ed c

olon

ies.

Mea

sure

men

ts o

f si

ze a

re i

n un

its

of 1

:100

mm

; w

ing

vena

tion

ang

les

are

in d

egre

es.

Cha

ract

ers

of p

igm

enta

tion

: 0

=co

mpl

etel

y da

rk, 9

= c

ompl

etel

y br

ight

(ye

llow

). M

easu

rem

ents

wer

e ta

ken

acco

rdin

g to

Rut

tner

(19

88).

mel

lifer

a(N

= 9

)ca

rnic

a(N

= 2

0)li

gust

ica

(N =

12)

cauc

asic

a(N

= 1

2)ar

men

iaca

(N

= 6

)m

eda

(N =

10)

anat

olia

ca(N

= 1

3)po

mon

ella

(N =

31)

hair

leng

th43

.47±

4.77

29.5

7±3.

7427

.92±

2.35

33.2

2±1.

9632

.53±

1.55

27.3

9±3.

1728

.86±

2.98

26.8

5±1.

95

prob

osci

s60

7.15

±13

.12

(N =

4)

640.

87±7

.51

(N =

18)

635.

94±

7.86

706.

64±1

5.06

664.

57±

7.01

628.

11±

8.87

(N

= 8

)64

3.62

±17

.64

(N =

12)

641.

16±

17.9

6

body

siz

e46

3.02

±9.

5845

3.44

±6.2

543

6.93

±7.

845

6.62

±7.

7644

9.85

±7.

3443

2.9±

3.59

444.

98±

7.32

449.

49±

14.0

4

cubi

tal i

ndex

1.82

±0.

22.

69±

0.18

2.52

±0.

122.

14±

0.12

2.61

±0.

162.

5±0.

232.

25±

0.22

2.24

±0.

20

ster

nite

6 in

dex

77.9

9±2.

9283

.81±

1.2

84.1

4±1.

4982

.14±

1.14

81.6

0±1.

6681

.12±

2.29

83.7

8±2.

0478

.18±

3.03

(N =

29)

tars

us in

dex

54.8

6±1.

6455

.31±

1.37

55.2

4±0.

6857

.81±

0.75

57.1

7±0.

8256

.34±

1.66

57.3

6±0.

9256

.78±

1.35

pigm

enta

tion

terg

ite 2

2.1±

0.74

1.39

±0.

95(N

= 1

5)7.

87±

0.4

3.92

±0.

278.

78±

0.28

8.59

±0.

465.

17±

1.18

(N =

7)

5.65

±1.

12

tom

entu

m in

dex

1.63

±0.

572.

07±

0.26

2.2±

0.54

2.76

±0.

212.

7±0.

172.

46±

0.62

2.2±

0.29

2.05

±0.

39

374 W.S. Sheppard, M.D. Meixner

thus the other subspecies in the O lineagewould represent westward expansion from thehomeland of A. m. pomonella.

The taxonomic classification of honey beesbelow the species level reflects an interpreta-tion of the extent of accumulated geneticdifferences between putative “subspecies” rel-ative to the distribution of overall within-spe-cies variation (Hepburn and Radloff, 1997;Sheppard, 1997). As a further complication,there are examples of gene flow betweendescribed subspecies. This phenomenon hasbeen studied and described in terms of geneticintrogression and hybridization using a varietyof molecular markers (Badino et al., 1984,1988; Cornuet et al., 1986; Smith et al., 1991;Meixner et al., 1993; Hepburn et al., 1998;Franck et al., 2000). Notable examples includenortheastern and northwestern Italy, wheregenetic introgression of A. m. ligustica occurswith neighboring A. m. carnica and A. m. mel-lifera, respectively. Evaluation of honey beepopulations from these and other areas ofintrogression with adequate multivariate mor-phometric analysis or with suitable geneticmarkers, can discern the hybrid background.Based on the tight morphological clustering ofthe Tien Shan honey bee from across the sam-pled range and the distance from other knownsubspecies, we do not consider it likely thatthere is widespread gene flow between A. m.pomonella and any of the western Asiansubspecies in the O lineage.

The description of this new honey beesubspecies is not directly consequential to theissue of sister taxon status between A.mellifera and A. cerana. Further studies ofwild honey bee populations in other regions ofcentral Asia, especially in areas closer to thenatural range of A. cerana, are needed todevelop an adequate understanding of thisaspect of Apis phylogeography.

ACKNOWLEDGEMENTS

We thank Drs. Tom Unruh, Roman Jashenko,Alex Kreutzberg, Vitaliy Kastsheev, MutatbekChildebaev, and Mr. Shufar Zurazaev for assistancein the field. Mr. Jamie Strange and Ms. DianaJohnson provided technical assistance. We are inde-bted to the Institut für Bienenkunde (Oberursel,Germany) for morphological reference data. TheWashington State Tree Fruit Research Commissionand the Washington State Beekeepers Associationprovided financial assistance.

Résumé – Apis mellifera pomonella, nouvellesous-espèce de l’Abeille domestique en Asie cen-trale. Les abeilles domestiques endémiques desMonts Tien Shan d’Asie centrale sont décritescomme nouvelle sous-espèce, Apis mellifera pomo-nella, sur la base d’analyses multivariées descaractères morphométriques. Les analyses morpho-métriques ont porté sur 31 échantillons d’abeillesprovenant de lieux situés dans les zones est et ouestdes Monts Tien Shan (Fig. 1). Les analyses en com-posantes principales et les analyses discriminantesprouvent que ces échantillons ne sont pas associésaux sous-espèces d’A. mellifera qui auraient puêtre importées dans la région par les pratiquesd’apiculture transhumante dans l’ancienne UnionSoviétique. Elles forment au contraire un groupedistinct qui se rattache à la branche évolutive orien-tale des abeilles domestiques (Figs. 2 à 5). Cesrésultats étendent l’aire de répartition d’A. melliferade plus de 2000 km vers l’est par rapport aux précé-dentes estimations. A. m. pomonella est une grosseabeille, seulement un peu plus petite qu’A. m. cau-casica. Elle se caractérise par un large abdomen etde larges tarses, deux caractères typiques des sous-espèces de la lignée orientale de l’Abeille domesti-que. Contrairement à A. m. caucasica, les abeillesdes Monts Tien Shan ont un proboscis plutôt court.Le tableau I donne les moyennes et les écarts-typesde certains caractères morphologiques d’A. m.pomonella, ainsi que ceux des sous-espèces envi-ronnantes. Les données des séquences mitochon-driales confirment aussi qu’A. m. pomonellapartage une histoire évolutive avec la lignée orien-tale et du sud-est de l’Europe, mais ne permettentpas de résoudre l’origine de cette sous-espèce.L’aire de répartition de cette nouvelle sous-espèced’A. mellifera en Asie centrale est considéréecomme étant le centre de la diversité génétique pourles pommes et d’autres espèces d’arbres fruitiers,d’où le nom proposé pour cette nouvelle sous-espèce : Apis mellifera pomonella.

Apis mellifera pomonella / Asie centrale / sous-espèce nouvelle / taxonomie

Zusammenfassung – Apis mellifera pomonella,eine neue Unterart der Honigbiene in Mittel-asien. Honigbienen aus dem Tien Shan Gebirge inZentralasien werden auf der Grundlage von multi-variaten Analysen morphologischer Eigenschaftenals neue Unterart Apis mellifera pomonella be-schrieben. Insgesamt wurden 31 Bienenproben vonSammelorten im westlichen und östlichen TienShan Gebirge morphometrisch analysiert. DieErgebnisse der Hauptkomponentenanalyse undDiskriminanzanalyse zeigten, dass diese Probennicht mit anderen Unterarten von A. mellifera asso-ziiert sind, die durch kommerzielle Wanderimkereiin der früheren Sowjetunion in diese Gegendgebracht worden sein könnten. Statt dessen werdensie klar in die orientalische Evolutionslinie der

A. m. pomonella, a new honey bee subspecies from Central Asia 375

Honigbienen eingeordnet und erweitern damit dasVerbreitungsgebiet von Apis mellifera um mehr als2000 km weiter nach Osten als zuvor angenommen.A. m. pomonella ist eine grosse Biene, nur unwe-sentlich kleiner als A. m. caucasica. Sie kann weiterdurch ein breites Abdomen und breite Tarsengekennzeichnet werden, beides für die orientalischeLinie typische Eigenschaften. Im Gegensatz zu A.m. caucasica haben die Bienen des Tien ShanGebirges jedoch einen relativ kurzen Rüssel. Mit-telwerte und Standardabweichungen einiger mor-phologischer Charaktere von A. m. pomonellaund benachbarter Unterarten sind in Tabelle Izusammengefasst. Sequenzdaten aus dem mito-chondrialen Genom unterstützen ebenfalls dieAnnahme einer gemeinsamen Evolutionsge-schichte von A. m. pomonella und der südosteuro-päischen und orientalischen Evolutionsline derHonigbienen, lösen jedoch die mögliche Herkunftdieser neuen Unterart nicht weiter auf. Das Verbreitungsgebiet der neuen Unterart vonApis mellifera in Zentralasien wird als Zentrum dergenetischen Vielfalt für Äpfel und andere Arten vonBaumfrüchten betrachtet. Als Name für die neueUnterart wird deshalb Apis mellifera pomonellavorgeschlagen.

Apis mellifera pomonella / Mittelasien / neueUnterart / Taxonomie

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