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Submitted on 1 Jan 1990
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Evidence of reproductive isolation confirms that
Apisandreniformis (Smith, 1858) is a separate species from
sympatric Apis florea (Fabricius, 1787)S Wongsiri, K
Limbipichai, P Tangkanasing, M Mardan, T Rinderer, Ha
Sylvester, G Koeniger, G Otis
To cite this version:S Wongsiri, K Limbipichai, P Tangkanasing,
M Mardan, T Rinderer, et al.. Evidence of reproductiveisolation
confirms that Apis andreniformis (Smith, 1858) is a separate
species from sympatric Apisflorea (Fabricius, 1787). Apidologie,
Springer Verlag, 1990, 21 (1), pp.47-52. �hal-00890810�
https://hal.archives-ouvertes.fr/hal-00890810https://hal.archives-ouvertes.fr
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Original article
Evidence of reproductive isolation confirms thatApis
andreniformis (Smith, 1858) is a separate
species from sympatric Apis florea (Fabricius, 1787)
S Wongsiri K Limbipichai P Tangkanasing M MardanT Rinderer HA
Sylvester G Koeniger G Otis
1 Bee Biology Research Unit, Faculty of Science,Chulalongkorn
University, Bangkok 10330, Thailand;
2 Department of Plant Protection, Universiti Pertanian
Malaysia,43400 Serdang, Selangor, Malaysia;
3 Honey-Bee Breeding, Genetics & Physiology Research1157 Ben
Hur Road, Baton Rouge, Louisiana 70820, USA;
4 Institut für Bienenkunde D 6370 Oberursel 1, FRG
(Received 7 September 1989; accepted 29 September 1989)
Summary — The species Apis andreniformis (Smith, 1858), the
small dwarf honey bee of South-east Asia, is recognized as a valid
biological species. This recognition is based on distinctive
endo-phallus characteristics in comparison with sympatric Apis
florea (Fabricius, 1787). Additionally, scan-ning electron
microscope images of drone basitarsi are presented, as are
preliminary comparisonsof wing venation.
Apis florea / Apis andreniformis / taxonomy / reproductive
isolation
INTRODUCTION
In 1984, our group collected dwarf honeybees in Thailand in the
province of Chan-taburi near the border with Kampuchea.Laboratory
examinations of worker beesfrom these collections revealed that
somebees had the species specific characteris-tics of Apis
andreniformis (Smith, 1858)and that others had the species
specific
characteristics of Apis florea (Fabricius,1787) that are
reported for worker bees(Maa, 1953).Wu and Kuang (1986, 1987)
reported
that secondary sex characteristics differedbetween drones of A
florea and A andre-formis. Specifically, both have a
furcatedbasitarsus, presumably modified to graspqueens during
mating (see Ruttner, 1988).The furcated basitarsus is quite
different in
*
Correspondence and reprints.
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the 2 groups. Following the report of Wuand Kuang, we
re-examined the sympatricpopulations in 4 Thai provinces, as well
asbees from East and West Malaysia.Drones and workers were
collected in1988 from nests of both A florea and A an-dreniformis
in the Thai provinces of Chan-taburi (Southeastern), Uthaithanee
(West-ern), Chiang Rai (Northern) andChumporn (Southern). A
andreniformiswas collected in 1989 in Malaysia.
Comparative studies of endophalli re-vealed evidence of
reproductive isolationbetween the 2 groups, and hence con-firmed
the biological validity of naming 2species of dwarf bees. In
addition, dronehind legs were examined using electronmicroscopy,
and worker bee wing venationwas studied.
Species-specific characteristics
Comparisons of the endophalli of A an-dreniformis and A florea
reveal numerousstructural characteristics which assure
complete reproductive isolation betweenthese species (fig 1). A
full or stage 9 ever-sion (Woyke and Ruttner, 1958; Ruttner etal,
1973) of an A andreniformis endophal-lus has a pair of bursal
cornua that fold for-ward once near their end; a pair of
uppercornua at the sides of the vestibulum,each of which has small
protrusions; ahairy field on the ventral surface that has 4separate
hairy patches distal to the pri-mary hairy area near the
vestibulum; an in-dented triangular hairy patch on the
dorsalsurface proximal to a fimbriate lobe; a fim-briate lobe
having 6 small protrusions; anda comparatively thick and straight
terminalportion extending distal to the fimbriatelobe.
In contrast, a full stage 9 eversion of anA florea endophallus
has a pair of bursalcornua that coil or generally fold twice, apair
of upper cornua at the sides of thevestibulum, each of which is
composed of3 joined protrusions; a continuous, non-indented hairy
field on the ventral surfaceextending from the vestibulum; a small
pro-trusion proximal to a small dorsal hairypatch; a fimbriate lobe
having 3 protru-sions and a comparatively thin and strong-ly curved
terminal portion extending distalto the fimbriate lobe.
Figure 1 illustrates these characters.The drawings are based on
direct observa-tions of several specimens as well as
theaccompanying photographs.
Figure 2 shows photographs of the basi-tarsal extensions
described by Wu andKuang (1986, 1987). Quite clearly, thethumb-like
extension is comparativelyshort in A andreniformis, and was
absentfrom the only drone of A andreniformis ex-amined from Sabah,
Malaysia. This exten-sion presumably provides a "clasper or-gan"
that fits on the queen’s hind tibiaduring mating (Ruttner, 1975).
However,whether or not this description of functionis accurate
awaits confirmation from obser-vations of matings. The wing
venation ofApis andreniformis workers is more similarto that of
Apis dorsata and Apis koschevni-kovi than it is to Apis cerana or
Apis florea.The cubital index of Apis andreniformis islarge (x for
colonies = 6.07, N = 11 colo-nies, 10 worker bees each; ranges =
5.03to 9.21 for colony averages and 3.5 to 13.8for individual bees)
while the cubital indexof A florea is comparatively small (x for
col-onies = 2.78, N = 6 colonies, 10 workerbees each; ranges = 2.53
to 3.14 for colo-ny averages and 1.93 to 5.27 for individual
bees).
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CONCLUSION
Apis andreniformis (Smith, 1858) is a validbiological species
which is found in south-east Asia and is sympatric with Apis
floreain much of Thailand. It is also found in theSouthern China
peninsula, Malaysia andBorneo but its exact distribution and
sym-patry with Apis florea are unknown. Equal-ly unknown are most
comparative aspectsof A andreniformis biology. Much that
waspublished concerning A florea may be Aandreniformis biology.
Extensive compara-tive studies are required to describe andproperly
attribute known dwarf honey beebiology to the correct species and
discoverwhat is unique to each species.
ACKNOWLEDGMENTS
In cooperation with the Louisiana AgriculturalExperiment
Station. Sincere appreciation is ex-pressed to Gerard Perrone for
his work in pre-paring the figures. We thank H V Daly, N Koeni-ger,
B Oldroyd and F Ruttner for suggestionsconcerning the manuscript. G
W Otis was sup-ported by a research leave grant from the
Uni-versity of Guelph, Guelph, Ontario. Financialsupport was
provided by the Research AffairsDivision of Chulalongkorn
University, Bangkok,Thailand.
Résumé — La preuve de l’isolementsexuel confirme qu’Apis
andreniformis(Smith, 1858) et Apis florea (Fabricius,1787) sont
deux espèces sympatriquesséparées. Apis andreniformis (Smith,1858)
et Apis florea (Fabricius, 1787) ontété trouvées en position de
sympatrie dansles régions thaïlandaises suivantes : Chan-taburi,
Uthaithanee, Chiang Rai et Chum-porn. L’examen de l’endophallus
desmâles de ces abeilles naines a révélé des
différences dans presque tous les détailsde structure (fig 1 ) :
dans les cornules ven-
trales (BC) et dorsales (UC), dans la sur-face pileuse ventrale
(HF) et dorsale(HDP), dans le lobe feuilleté (FL) et dansla
longueur et la forme de la partie termi-nale qui s’étend après le
lobe feuilleté (T).Ces différences sont la preuve de l’isole-ment
reproductif des deux espèces sympa-triques et donc, la preuve qu’il
s’agit biendans le cas présent de véritables espèces.Les
endophallus d’A andreniformis deThaïlande et de Malaysie sont
semblables.
Sont jointes en outre des photos aumicroscope électronique de
l’extension ba-sitarsale caractéristique des pattes anté-rieures
des mâles des deux espèces(fig 2). Les photographies de l’aile
anté-rieure des 2 espèces montrent clairementl’index cubital élevé,
caractéristique d’A an-dreniformis (x = 6,07) et celui petit,
carac-téristique d’A florea (x = 2,78) (fig 3).
La répartition précise d’A andreniformisaussi bien que sa
biologie sont encore in-connues. Des études comparatives
sontnécessaires, car dans le passé certainscaractères d’A
andreniformis ont certaine-ment été attribués à tort à A
florea.
Apis florea / Apis andreniformis / systé-matique / isolement
sexuel
Zusammenfassung — Der Beweis fürdie reproduktive Isolierung
bestätigt,dass Apis andreniformis (Smith, 1858)eine von der
sympatrischen Apis florea(Fabricius, 1787) getrennte Art ist.
Apisandreniformis (Smith, 1858) und Apisflorea (Fabricius, 1787)
wurden infolgenden Provinzen von Thailandsympatrisch nachgewiesen :
Chantaburi,Uthaithanee, Chiang Rai und Chumporn.Untersuchungen des
Endophallus derDrohnen beider Zwergbienen ergaben beifolgenden
strukturellen EinzelheitenUnterschiede (Abb 1) : bei den
ventralen(BC) und dorsalen (UC) Hörnchen
-
(Cornua), beim ventralen Haarfeld (HF),beim dorsalen Haarfeld
(HDP), beimFederanhang (FL) und in der Länge undForm des Endteils
nach dem Federanhang(T). Diese Unterschiede sind Beweise füreine
reproduktive Isolation zweier sympa-trischer Formen, also Beweise,
dass essich hierbei um echte Arten handelt. Die
Endophalli von A andreniformis vonThailand und Malaysia sind
ähnlich.
Zusätzlich wurden elektronenmikros-
kopische Fotos über den basitarsalenFortsatz (Klammerorgan) des
Hinterbeinsder Drohnen von beiden Arten beigefügt(Abb 2).
Fotografien der Vorderflügelbeider Arten zeigen deutlich den
charak-teristisch grossen Kubital-Index von Aandreniformis (x =
6,07) und den charak-teristisch kleinen Kubital-Index von Aflorea
(x = 2,78) (Abb 3).
Sowohl die genaue Verbreitung von Aandreniformis als auch ihre
Biologie sindnoch unbekannt. Vergleichende Unter-suchungen sind
nötig, da in der Vergan-genheit sicher einige Eigenschaften der
A.andreniformis fälschlich A. florea zuge-ordnet wurden.
Apis florea / Apis adreniformis /Taxonomie / reproduktive
Isolierung
REFERENCES
Fabricius JC (1787) Mantissa Insectorum Vol IProft. Hafniae
Maa T (1953) An inquiry into the systematics ofthe tribus
Apidini or honey bees Hym. Treu-bia 21, 525-640
Ruttner F (1975) Ein metatarsaler Haftapparatbei den Drohnen der
Gattung Apis (Hyme-noptera: Apidae). Entomol Germanica 2, 22-29
Ruttner F (1988) Biogeography and taxonomy ofhoney bees.
Springer Verlag, Berlin NY
Ruttner F, Woyke J, Koeniger N (1973) Repro-duction in Apis
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Res12, 21-34
Smith F (1858) Catalogue of the Hymenopter-ous insects collected
at Sarawak, Borneo;Mount Ophir, Malakka; and at Singapore,Wallace
AR. J Proc Linn Soc London Zool 2,42-130
Woyke J, Ruttner H (1958) An anatomical studyon the mating
process in the honeybee. BeeWorld 39, 3-18
Wu Y, Kuang B (1986) A study of the genus Mic-rapis (Apidae).
Zoological Research 7, 99-102, In Chinese
Wu Y, Kuang B (1986) Two species of smallhoneybee — A study of
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