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Position of the Red Honey bee, Apis koschevnikovi(Buttel-Reepen 1906), within the Genus Apis

F. Ruttner, D. Kauhausen, N. Koeniger

To cite this version:F. Ruttner, D. Kauhausen, N. Koeniger. Position of the Red Honey bee, Apis koschevnikovi (Buttel-Reepen 1906), within the Genus Apis. Apidologie, Springer Verlag, 1989, 20 (5), pp.395-404. �hal-00890794�

Original article

Position of the Red Honey bee, Apis koschevnikovi(Buttel-Reepen 1906), within the Genus Apis

F. Ruttner D. Kauhausen 2 N. Koeniger

1 Universität Frankfurt, Institut für Bienenkunde (Polytechn. Gesellschaft), D-6370 Oberursel, FRG2 Bayerische Landesanstalt für Bienenzucht, D-8520 Erlangen, FRG

(received 19 December 1989, accepted 29 March 1989)

Summary — The "Red Bee" of Borneo, described in 1988 by Koeniger et al. and Tingek et al. as aseparate species, was first named by H. v. Buttel-Reepen in 1906. The correct name, therefore, is

Apis koschevnikovi Buttel-Reepen 1906 and not Apis vechti Maa 1953. By multivariate analysis(PCA and DA) Apis koschevnikovi can clearly be separated from the 2 other cavity-nesting Apis spe-cies. Although A. koschevnikovi is very similar to A. cerana phenotypically, it is much larger thanbees of this species from a similar geographic latitude, corresponding in size to equatorial popula-tions of A. mellifera from Africa.

Apis— taxonomy — Apis koschevnikovi - morphometrics — Borneo

THE CORRECT SCIENTIFIC NAME

In the spring of 1988, Koeniger et al. andTingek et al. published data on the taxo-nomic status of a special honey bee fromS.E. Asia as a true species which they de-termined as Apis vechti (Maa, 1953). Butthis was not the first description of this

honey bee, very conspicuous by its red-dish legs and hair, and by its size. Buttel-Reepen (1906) found several specimensof this kind in the Museum of Natural His-

tory in Berlin which were labelled as

"Cameroon", and one as "N. Borneo". Hedismissed the possibility as unlikely, andnamed this cerana-type honey bee "Apismellifica indica-koschevnikovi " [in Buttel-Reepen’s taxonomic system A. indica (=A. cerana) was taken as a subspecies ofA. mellifera. The prefix "indica" was usedby Buttel-Reepen to show the relationship

of his new form, but elsewhere in thesame work he referred to it as koschevni-kovi. There is no doubt but that koschev-nikovi by itself is the name for this bee].To him, the specimens at the Berlin Mu-seum were evidence that A. cerana alsooccurs on the African continent.

About 50 years later (1953) Maa de-scribed a number of specimens of the"Red Bee", all collected in Borneo, in histaxonomic revision of the genus Apis.Maa considered the "Red Bee" to be a

separate species and gave it a new

name, Apis vechfi (after the entomologistJ. Van der Vecht then of the Bogor Mu-seum, now Putten, The Netherlands). Heeven noticed a certain geographic vari-ability sufficient to establish 2 subspecies("vechti

" and "linda"). Maa did not re-

evaluate the specimens of the Berlin Mu-seum and proposed retaining the name

&dquo;A. koschevnikovi &dquo; for the supposedly Af-rican &dquo;Red Bee&dquo;.

The present situation is as follows :

1) &dquo;Apis vechti &dquo; is a true species: it differsfrom A. cerana in the endophallus (Tingeket al., 1988) and time of drone flight (= re-productive isolation; Koeniger et aL, 1988).It is sympatric with A. cerana indica.

2) There is no evidence yet that a cerana-type honey bee exists on the African conti-nent. Mislabelling of the specimens at theBerlin Museum is the most probable expla-nation for the present confusing situation.Consequently, 2 names are probably usedfor the same bee of the same area. After

consulting Dr Charles Michener in this

matter, we asked Dr Frank Koch, Museumfur Naturkunde (Humboldt Universität, Ber-lin) to re-examine the specimens de-scribed by Buttel-Reepen in 1906 (whichhave miraculously survived the last 83

years) to confirm their cerana-type charac-ters. The following criteria were usedwhich discriminate best between A. ceranaand A. mellifera :

i) prolongation of radial vein of hind wing(&dquo;indica vein&dquo;) - present;ii) tomentum on last tergite-present;iii) number of hamuli on hind wing - 17 to19.

Thus there is no doubt that all of Buttel-

Reepen’s specimens are cerana-type andthat they were not collected in Cameroonbut most likely in Borneo. According to therules of priority, the name &dquo;A. vechti &dquo; is re-dundant and the correct scientific name ofthe Red Honey bee has to be Apis ko-schevnikovi (Buttel-Reepen, 1906).

Here it should be noted that no typesimilar to the &dquo;Red Bee&dquo; is among the vari-ous honeybees collected by Wallace inNorth Borneo and described by Smith

(1858; 1865); he lists A. dorsata and a va-riety of this species, A. testacea, A. indicawith the variety perrottetii and A. andreni-

formis, all clearly different from A. koschev-nikovi. These findings raise the question ofthe position of this species between theother 4 known recent Apis species.

MATERIAL AND METHODS

For the morphometric analysis of the &dquo;Red Bee&dquo;,two sets of data were used.

Phenetic analysis including fossil hon-eybees

Characters of 12 individuals of A. koschevnikoviand 12 of A. cerana indica, both from the regionof Tenom, Sabah, N. Borneo, were analysed to-gether with data on individual honey bees asused in an earlier analysis (Ruttner et al., 1986).The origin of these bees was the following :1) A. armbrusteri— Schwabische Alb, Upper Mi-ocene, 10 specimens2) A. dorsata - Pakistan, 10 specimens3) A. florea- Pakistan, 10 specimens4) A. cerana cerana - Afghanistan, 10 speci-mens

5) A. mellifera— Iran, 10 specimens.The characters used were 16 wing venation

angles; no characters of size were included. Themultivariate statistical methods were factor anddiscriminant analyses (PCA, DA). Both methodsgave basically identical results. Finally, ellipsesof confidence were calculated (Cornuet, 1982).

Taxonomic-morphomefric analysis

In these analyses all 34 characters were usedas in the standard method of honey bee mor-phometry (Ruttner et al., 1978; Ruttner, 1988);not individual bees but the means of samples of15-20 bees were taken as units. Five samplesof A. koschevnikovi from N. Borneo and 1 sam-

ple from Sumatra were analysed together withsamples of A. cerana of various origins and oftropical populations (Arabia, E. Africa) of A. mel-lifera.

Voucher specimens of A. koschevnikovi havebeen placed for permanent preservation in theHoneybee Taxonomy Collection of the Institutfur Bienenkunde Oberursel, University of Frank-furt.

RESULTS

Phenetic analysis

As was to be expected, the positions of theclusters are identical with those of an earli-er investigation of fossil and known recentspecies : the fossil A. armbrusteri and A.dorsata are represented as a common

cluster, with only slight differentiation ofthe 2 species (Figs. 1 and 2). A. koschev-nikovi appears together with the 2 othercavity-nesting species as 3 separate clus-ters without overlapping, A. cerana in thecentre and A. mellifera at the periphery.The cluster of A. koschevnikovi is found indirect contact with that of A. cerana, butdistant from the mellifera cluster. The cera-na samples from Borneo are at the centreof the general cerana cluster (Fig. 2B). Inthe graph showing the ellipses of confi-dence (95%, Fig. 2), the K cluster heavilyoverlaps the C cluster but not the M clus-ter.

Taxonomic analysis

The samples of A. cerana are arranged in2 separate groups (Fig. 3). While the popu-

lations with small bees of the subspeciesA. cerana indica from S. India, Sri Lanka,Thailand, Indonesia, etc. appear to the left,the samples of the large A. cerana ceranafrom Afghanistan, Pakistan, N. India, Chi-na, and those of A. cerana japonica are tothe right. The clusters of the tropical melli-fera populations are at great distance inthe far right corner.

The cluster of the 5 A. koschevnikovi

samples (15 bees each) from Borneo isfound between these 2 groups (Fig. 3),close to that of A. cerana cerana, the clos-est samples being the populations fromS.W. China (Yunnan, Fig. 3). When ellip-ses of confidence are calculated, A. kos-chevnikovi overlaps with A. c. japonica inthe graph of factor 1/factor 2, but it is well

separated in the graphs of factors 1/3 (Fig.4). In this analysis (1/2) the distance of theA. koschevnikovi centroid to the centroid ofA. c. cerana is less than the distance be-

tween the centroids of A.c. cerana and A.c. indica.

If the group A. koschevnikovi-A. ceranais analysed by itself, that is including exclu-sively samples of these 2 species, the ce-rana subspecies appear in one compact,only slightly subdivided cluster, while thekoschevnikovi samples are found in a re-

mote, compact aggregation (Fig. 5).One sample (No. 991; 20 bees) from

Sumatra, collected in 1978 by one of theauthors (Koeniger) in Muaro near Solok, isseparated from the A, c. indica cluster by agreat distance, but very close to the A.koschevnikovi samples (Fig. 3). These

honey bees show the same typical rufouscolor of legs and abdomen as A. koschev-nikovi; their size is somewhat smaller thanthat of A. koschevnikovi from Borneo, butthey are much larger than indica bees fromSumatra, and the cubital index and the in-dex of st6 show the typical extreme values

of A. koschevnikovi. The following data arearranged in the sequence A. koschevniko-vi Borneo — sample 991 (Sumatra) - A.c. indica Sumatra (6 samples; in mm) :length of fore wing, 8.541-8.264-7.769;length of hind leg, 7.612-7.558-6.839; ter-gites 3 + 4, 3.987-3.886-3.535; index ofsternite 6, 89.56-90.24-86.50; cubital in-

dex, 7.59-7.86-4.34. There is no doubt,therefore, that the sample from Sumatrabelongs to the same taxonomic unit as thesamples of A. koschevnikovi from Borneo.

DISCUSSION

Apis koschevnikovi can be separated fromall the other Apis species by quantitativecharacters (Figs. 1, 3, 5), but the level ofconfidence is low for discrimination from A.

cerana if very different, remote species areincluded such as the Miocenic A. armbrus-

teri, a fossil honeybee which lived 12 mil-lion years ago (Fig. 2). But even in this

case, almost no overlapping occurs withthe ellipse of confidence (95%) of sympa-tric A. c. indica from Borneo. A good dis-crimination is obtained, however, if onlythe closely related cavity-nesting speciesA. cerana and A. mellifera are included inthe analysis (Figs. 3, 4 and 5).

In A. koschevnikovi, only a few of thecharacters generally used in honey beemorphometrics are outside or at an ex-

treme end of the known range of A. cerana

(see Ruttner, 1988). A. koschevnikovi is

very long-tongued (5.870 mm), slender

(st6-1 89.6), with narrow tomenta (tom-10.389) and high CI (7.59). All the other

characters, especially those of size, are

well within the range of cerana populationsof the temperate zone. The most conspicu-

ous character by which &dquo;this species canvery easily be distinguished from any otherhoney bee species is the uniform rufous

pattern&dquo; (Maa, 1953).Evidently, the overlapping of the A. ko-

schevnikovi and the cerana clusters is

mainly due to characters of size. Axis 3 ofa DA, which contains no characters of size(length of vertical bars in Fig. 3), clearly in-creases the distance A. koschevnikovi -

A. cerana. Size in honey bees, however, ishighly correlated to geographic latitudewithin the range of a species. Therefore it

seems appropriate to introduce an ad-ditional parameter into the taxonomic anal-ysis : geographic latitude. All the Apis spe-cies investigated so far (mellifera, cerana,florea; Ruttner, 1988) clearly follow Berg-mann’s rule. A correlation coefficient of0.7-0.9 was found for most characters ofsize and geographic latitude in A. mellifera(Ruttner, 1985). Therefore, a taxonomicdistortion will result if equatorial popula-tions are compared with those of higher

geographic latitudes. The samples of the&dquo;Red Bee&dquo; were collected in N. Borneo

(Sabah) at 2° N latitude. The sympatric A.cerana subsp. indica is much smaller (Rin-derer et al., 1989), very much like the pop-ulations of Java, Sumatra, Sri Lanka andS. India. A. koschevnikovi compares in

size with the cerana bees of the mountainsof Yunnan (25°) and from Honan (near To-kyo, 36°, Table I). A similar gradation isfound in A. mellifera (Ruttner, 1986); onthis scale, A. koschevnikovi compares in

size with the bees from Cameroon (4 °S)and Lamu (coast of Kenya, 2°N). As-

suming the same scale of gradation, an A.koschevnikovi of the temperate zone

would be of the size of A. mellifera (TableI). This, however, is a completely differentscale of size variation than in A. cerana,thus furnishing another factor to distinguishthis species.

The morphometric characters aside,various others can be listed to show theclose relation of A. koschevnikovi to A. ce-

rana : tomentum on the last tergite; &dquo;indicavein&dquo; of the hind wing; shape of endophal-lus; and pore in the capping of drone cells.It cannot be overlooked, however, thatthere are characters which indicate a cer-tain affinity to A. dorsata which is frequent-ly considered as an ancestral type in rela-tion to the cavity-nesting species: size of ahoneybee in S. Asia; slender abdomen;high cubital index; smokey tinge of wings.Methods of molecular biology might pro-vide further evidence of the phylogeneticrelationship among the honey bees of S.Asia.

As noted above, A. koschevnikovi is notrestricted to Borneo. A sample in the

Oberursel data bank, collected 10 yearsago near Solok in Sumatra and listed asan isolated, &dquo;aberrant&dquo; type, clearly be-

longs to this species. The true area of dis-tribution of the species has yet to be inves-tigated. A certain conclusion can be madeabout the age of the species. The Indone-sian Islands were separated from the con-tinent only during the sea level rise in thepost-pleistocenic period. The island popu-lation of A. c. indica, therefore, has noteven reached the status of a subspecies(Ruttner, 1988). The separation of the twospecies A. koschevnikovi and A. ceranacertainly occurred in a much earlier period.

ACKNOWLEDGMENTS

We wish to thank Dr Frank Koch, Museum furNaturkunde der Humboldt-UniversitAt Berlin, Dr

Christopher O’Toole, The University MuseumOxford, and Professor Charles Michener, SnowEntomological Museum, Lawrence, Kansas,USA; for their kind and helpful cooperation in

the process of clarifying the confusing nomen-clature of the &dquo;Red Bee&dquo;. Agnes Mohr, Oberur-sel, was as reliable as ever in taking all themeasurements and adapting the figures. E.

Huttinger and Dr. R. Keller assisted in comput-ing the graphs.

Résumé — La position de l’abeille

rouge, Apis koschevnikovi (Buttel-Reepen 1906) au sein du genre Apis.Depuis qu’il existe une science de l’abeille,des centaines d’espèces ont été décritesmais seules 4 espèces ont acquis une va-leur durable. Aussi cela fit-il du bruit lors-

qu’en 1988 Koeniger et al. et Tingek et al.montrèrent dans deux courtes notes qu’ilexistait à Bornéo une autre espèce nidi-

fiant dans les cavités, remarquable par sacouleur rouge cuivre et identifiée comme

espèce propre en raison de la forme de

l’endophallus et d’un décalage dans la pé-riode de vol des mâles empêchant le croi-sement : Apis vechti (Maa, 1953). Mais,entretemps des recherches ont montré

que la même abeille avait vraisemblable-ment déjà été décrite en 1906 par l’ento-mologiste berlinois H. von Buttel-Reepensous le nom d’A. indica koschevnikovi,d’après des exemplaires de musée dontl’origine était marquée parfois &dquo;Cameroun&dquo;

parfois «Bornéo&dquo;. Le préfixe «indica» (au-jourd’hui synonyme de cerana) devait indi-quer la ressemblance avec l’abeille orien-tale. Puisqu’aujourd’hui on sait

précisément qu’A. cerana est absente

d’Afrique, il faut admettre une erreur dans

l’étiquetage, dans le cas où les abeilles

présentent réellement des caractéristiquescerana. Grâce à l’amicale collaboration duDr Frank Koch, du Museum d’Histoire Na-turelle de l’Université Humboldt à Berlin,on a pu apporter la preuve sur des exem-plaires conservés intégralement, qu’ils neprovenaient assurément pas d’Afriquemaid du sud-est asiatique. Selon la règlede priorité leur nom d’origine A. koschevni-kovi doit être conservé.

Pour l’analyse morphologique de la nou-velle abeille, on disposait de 5 échantillons

de 20 abeilles chacun récoltés dans leNord de Bornéo (Sabah). Un autre échan-tillon de Sumatra, déjà présent dans la col-lection d’Oberursel, a pu être déterminé

plus tard comme &dquo;Abeille rouge&dquo;. Leséchantillons de comparaison provenaientde la même collection.

Pour déterminer la place d’A. koschevni-kovi au sein du genre Apis, on a d’abordfait une analyse discriminante (limitée auxcaractéristiques de l’aile) sur toutes les es-pèces connues d’Apis, y compris A. arm-brusteri vieille de 12 millions d’années (Fig.1 Les exemplaires de l’Abeille rouge setrouvent dans un groupe individualisé,étroitement appuyé contre A. cerana maistrès éloigné d’A. mellifica. Si l’on calculeles ellipses de confiance à 95%, celles d’A.koschevnikovi et d’A. cerana se recouvrentfortement (Fig. 2).

Si l’on analyse seules les 3 espècesd’Apis qui nidifient dans des cavités, leséchantillons d’A. koschevnikovi sont situésentre les formes grandes (A. c. cerana) etles formes petites (A. c. indica) d’A. cera-na, bien loin des races tropicales d’A. met-lifica (Fig. 3). Les ellipses de confiance à95% ne montrent aucun recouvrement

(Fig. 4). Si finalement on teste seulel’Abeille rouge contre A. cerana avec

toutes ses races, une délimitation trèsnette apparaît (Fig. 5).

A. koschevnikovi est nettement plusgrande que la population locale d’A. c. ce-rana; sa taille correspond à peu près àcelle des races d’A. mellifica de la mêmelatitude (Tableau 1). Puisque chez toutesles espèces d’abeilles la taille augmentedu sud au nord, on peut admettre qu’A.koschevnikovi montrerait la même ampleurde variation de la taille qu’A. mellifica sielle avait réussi à pénétrer dans la zone

tempérée.

Zusammenfassung — Die Stellung derRoten Biene, Apis koschevnikovi (But-tel-Reepen 1906) in der Gattung Apis.Seit es eine Wissenschaft von der Honig-biene gibt, wurden hunderte von Bienen-arten beschrieben, aber nur vier Arten hat-ten bisher dauernde Gültigkeit. Darum er-regte es umso größeres Aufsehen, als

1988 Koeniger u.Mitarb. und Tingek u. Mi-tarb. in zwei kurzen Arbeiten zeigen konn-ten, daß es in Borneo eine weitere,höhlenbrütende Art gibt, auffällig durch

ihre kupferrote Körperfarbe und als eigeneSpezies ausgewiesen durch eine andereForm des männlichen Begattungs-schlauches sowie durch eine Verschie-

bung der Drohnenflugzeit als Kreuzungs-hemnis: Apis vechti (Maa, 1953). Inzwi-

schen angestellte Nachforschungen habenaber ergeben, daß vermutlich dieselbeBienenart schon 1906 von dem Berliner

Entomologen H. v. Buttel-Reepen an Handvon Museumsexemplaren, deren Herkunftz. T. als &dquo;Kamerun&dquo;, z.T. als &dquo;Borneo&dquo; an-

gegeben war, unter dem Namen A. indica-koschevnikovi beschrieben worden war.

Der Zuname &dquo;indica&dquo; (heute gleichbedeu-tend mit A. cerana) sollte auf die

Ähnlichkeit mit der Östlichen Honigbienehinweisen. Da man heute genau weiß, daßin Afrika A. cerana nicht vorkommt, ist einIrrtum in der Etikettierung anzunehmen,falls diese Bienen tatsächlich cerana-

Merkmale aufweisen. Dank der freundli-chen Mitarbeit von Dr. Frank Koch, Natur-kundliches Museum der Humboldt-Universität Berlin, konnte dieser Nachweisan den vollständig erhaltenen Exemplarengeführt werden, die somit sicherlich nichtaus Afrika, sondern aus SO Asien stam-men und nach der Prioritätsregel ihren

ursprünglichen Namen A. koschevnikovibehalten müssen.

Zur morphometrischen Analyse derneuen Art standen 5 in Nordborneo (Sa-bah) gesammelte Proben zu je 15 Bienenzur Verfügung. Eine weitere Probe aus Su-matra, die schon in der Sammlung Oberur-sel vorhanden war, konnte später eben-falls als &dquo;Rote Biene&dquo; bestimmt werden.Die Vergleichsproben stammten ebenfallsaus dieser Sammlung.Um die Stellung von A. koschevnikovi

innerhalb der Gattung Apis zu bestimmen,wurde zunächst eine Diskriminanz-

Analyse (beschränkt auf Flügelmerkmale-mit sämtlichen bekannten Apis-Arten, alsomit Einschluß der 12 Millionen Jahre altenA. armbrusteri aus dem Randecker Mar

durchgeführt (Abb. 1 Die Exemplare derRoten Biene sind in einer geschlossenenGruppe zu finden, eng angelehnt an A. ce-rana, aber weit entfernt von A. mellifera.Berechnet man den Vertrauensbereich derstatistischen Abgrenzung bei 95%, so er-gibt sich eine starke Überlappung der kon-fidenzellipsen von A. koschevnikovi und A.cerana (Abb. 2).

Werden die drei höhlenbrütenden Apis-Arten für sich analysiert, so liegen die Por-ben von A. koschevnikovi zwischen den

großen (A. c. cerana) und den kleinen For-men (A. c. indica) von A. cerana, weit ent-fernt von tropischen Rassen von A. mellif-era (Abb. 3). Die Konfidenzellipsen von95% zeigen keine Überlappung (Abb. 4).Wird schließlich die Rote Biene gegenüberA. cerana mit all ihren Rassen allein getes-tet, so ergibt sich eine sehr klare Abgren-zung (Abb. 5).

A. koschevnikovi ist ganz wesentlich

größer als die lokale Population von A. ce-rana indica; sie entspricht in ihrer Größeetwa den Rassen von A. mellifera aus der-selben geographischen Breite (Tab. I). Dabei allen Bienenarten die Formen von

Süden nach Norden zu größer werden,kann man annehmen, daß A. koschevniko-vi dieselbe Größen-Variations breite wie A.mellifera hätte, falls es ihr gelungen wäre,in die gemäßigte Zone vorzudringen.

REFERENCES

Buttel-Reepen H. (1906) Apistica. Beitrage zurSystematik, Biologie, sowie zur geschichtlichenund geographischen Verbreitung der Honig-biene (Apis mellifica L.), ihrer VarietAten und deriibrigen Apis-Arten. Veroff. Zool. Museum Berlin117-201

Cornuet J.M. (1982) Representation graphiquede populations multinormales par des ellipsesde confiance. Apidologie 13, 15-20Koeniger N., Koeniger G., Tingek S., Mardan M.& Rinderer T.E. (1988) Reproductive isolation

by different time of drone flight between Apis ce-rana Fabricius 1793 and Apis vechti Maa 1953.Apidologie 19, 103-106Maa T.C. (1953) An inquiry into the systematicsof the tribus Apidini or honeybees (Hymenopte-ra). Treubia 21, 525-640

Rinderer T.E., Koeniger N., Tingek S., MardanM. & Koeniger G. (1989) A morphological com-parison of the cavity dwelling honey bees of Bor-neo, Apis koschevnikovi (Buttel-Reepen, 1906)and Apis cerana (Fabricius 1793). Apidologie20, 405-411 1

Ruttner F. (1985) Graded geographic variabilityin honeybees and environment. Pszcz. Zesz.Nauk29, 81-92

Ruttner F. (1988) Biogeography and Taxonomyof Honeybees. Springer Verlag, HeidelbergRuttner F., Tassencourt L. & Louveaux J. (1978)Biometrical-statistical analysis of the geographicvariability of Apis mellifera. Apidologie 9, 363-381

Ruttner F., Wilson R., Snelling G., Vorwohl G. &Kauhausen D. (1986) The evolution of the hon-eybee wing venation. Apidologie 17, 349Smith F. (1858) Catalogue of the Hymenopter-ous insects collected at Sarawak, Borneo,Mount Ophir, Malacca, and at Singapore, byA.R. Wallace. J. Proc. Linn. Soc. London Zool.

2, 42-130Smith F. (1865) On the species and varieties ofthe honey-bee belonging to the genus Apis.Ann. Mag. nat. Hist. London 15, 372-380

Tingek S., Mardan M., Rinderer T.E., KoenigerN. & Koeniger G. (1988) Rediscovery of Apisvechti (Maa: 1953): the Saban honey bee. Api-dologie 19, 97-102