01992IBRA
Journalof ApiculturalResearch30(2):54-59 (1991)
lt
The limitationsof
biometriccontrol
on purerace
breedingin Apis
mellifera
RoerNF A Montrz'
Department
of Genetics,Universityof Natal,PO
Box 375, Pietermaritzburg,
3200SouthAfrica
.l
(Received24 April 991, acceptedfor
p u b l i c a t i o2n8 A u g u s t1 9 9 1 )
S UM M A R Y
Germanbee breedershavetried to reolacethe
autochthonoushoneybee populationApis
mellifera mellifera with A. m. carnica, a SE
Europeanraceof honeybee,for morethan 40
years.The latter race is believedto be more
In a biometrical
adaptedto bee management.
study of the honey bee populationin lower
Bavaria,samplesof at least20 workersper
colonyweretakenfrom9'1apiariesin autumn
1990.Thewing venationpatternsof the bees
discriminant
wereanalysedusingmultivariate
analysis,and the resultswerecomparedwith
those for 6 samplesof A. m. carnicaused for
commercialbreedingfrom Kirchhainand 7
samples of preservedA. m. melliferalrom
Erlangenand Kulmbachdatingbackto 1911.
Thedata indicatethat,in spiteof tremendous
breedingefforls,the bees of this areaform a
hybridtype betweenA. m. carnica andA. m.
mellifera.Althoughland-basedmatingstations
havethus failedto producepure racialtypes
in this area,it is consideredthat such stations
are of valueto practicalhoneybee breedingif
in open populationselecusedappropriately
tion schemes.
Keywords: Apis melliferamellifera, Apis
melIifera carnica, selectionprogramme,
breeding,biometry,matingstations,Bavaria
'Presentaddress:Institut für Biologie,TechnischeUniversität
Berlin,Franklinstr.
28/29, 1OOO
Berlin10, Germany.
55
Moritz
INTRODUCTION
I n b e e b r e e d i n gc e r t a i n r a c i a lt y p e s a r e o f t e n
undercerbelievedto showsuperiorperformance
tain conditionsto otherraces.This is clearlyso in
honeybee probthe Americaswith the Africanized
lem.Thereare severalprogrammesattemptingto
replacethe Africanizedtype with more desirable
stock.This is nothingnew in bee breeding.In the
after1950Germanbeekeep1940sand particularly
, 9 4 9 ,1 9 6 4 ) ,
e r s ,m a i n l yg u i d e db y G o e t z e( 1 9 4 0 1
'black'honeybee,
decidedthatthe locallyabundant
Apismelliferamellifera,was less suitablefor beekeeping than A. m. carnica,which has an
distributionin Austria,Yugoslavia
autochthonous
and Hungary(Ruttner,19BB).lt was believedthat A.
m. carnica bees were less aggressiveand better
adaptedto earlyspringhoneyflows. Nationwide,
supportedverystrictbreedingregulabeekeepers
tions in order to replaceA. m. melliferawith A. m.
carnicahoney bees and the breedingeffortswere
by statefunds.
supplemented
substantially
the problemof
Sincebothracesreadilyinterbreed,
was large.Forthis reason,severalbiohybridization
metricalcharacteristicsthat separated melIiferaw ell
'1
from carnica(Ruttner, 983) were used to try and
identifythosehybridsthat did occur.In particular,
the lengthof the abdominalbody hairs,the width
of the tomentum,andthe so-calledcubitalindexof
the cubitalcell in the wingvenationarestrikinglydif1969).
ferentbetweenthe two races(Ruttner,
to try to
lsolatedmatingapiarieswereestablished
betweenLhecarnicastock and
reducehybridization
population.
manydifHowever,
the autochthonous
ficultiesarosebecauseof the largematingrangeof
1972).Strict
queensand drones(Ruttner& Ruttner,
matingcontrolwas achievedon NorthSeaislands,
but matingstationson the mainlandhad substantial
problems.Althoughferalhoneybees rarelyinter'pure'matings,unconferedwith efforlsto obtain
managedbee
with non-selected
trolledhybridization
coloniesfrequentlyoccurred.One attemptto solve
this problem,was the creationof purebredcarnica
areasaroundthe actualmatingapiary(Böttcher,
1989).For Bavarianmatingstationsthis protective
belt had a diameterof 15 km.A. m. carnicaqueens
weredistributedfreeof chargeamongthe beekeepers in thisarea.Thesequeenswereto producethe
'father'
. dditional
d r o n e sf o r t h e m a t i n ga p i a r y A
colonieswith a largenumberof carnicadroneswere
placedin the centreof the matingapiary,but, in
contrastto islandmatingstations,wereexpected
to haveonlya minorimpacton the matingpopulacarnicaqueensin largenumtion of drones.Vtrgin
berswere placedin the matingapiaryduringthe
matingseason.Severalthousandsof queenswere
matedat thesestationsyearly,which made many
believethat in fact the currentlymost
beekeepers
abundantracewould be carnicainsteadof mellifera.
In orderto justifythe extrawork-loadin honeybee
b r e e d i n gd u e t o b i o m e t r i c aal n a l y s i si t s e e m e d
to studywhetherthe objectiveof race
appropriate
was actuallybeingachieved.
replacement
M A T E R I A L SA N D M E T H O D S
Samples
apiariesweresampledin autumn1990
Ninety-one
in lowerBavaria.Eachsamplewas comprisedof at
least20 workersper colony.Tensampleswerecoilectedwithinthe protectivebelt of eachof two malTwo
and Bramandlberg).
ingapiaries(Schellenberg
o t h e r m a t i n g a p i a r i e s( R a c h e l d i e n s t h ü t taen d
K ö n i g s w a l d p) r o v i d e dn i n e s a m p l e se a c h . T e n
colonieseachweresampledin areasadjacentto the
protectivearea,and 13 samplescamefrom colonies
not in the vicinityof any matingapiary(morethan
20 km distance).
In orderto evaluateto whichdegreethe racialstatus
of the beeshad changedduringthe last40 years,
it seemedappropriateto comparethem with carnica
to replacethe origstockthat had beenintroduced
i n a l h o n e y b e e p o p u l a t i o nr a t h e r t h a n w i t h
a u t o c h t h o n o u sc a r n i c a p o p u l a t i o n so f c e n t r a l
Europe.Six samples(eachof a differentbreeding
family) were obtained from the Hessische
für Tierzucht,Departmentof Bee
Landesanstalt
Breedingin Kirchhain,FederalRepublicof Germany
(FRG),which maintainsseveralcarnicalinesfrom
insemAustriaand Sloveniathroughinstrumental
ination.Thismaterialincludesmoststrainsthat have
been and are currentlyin use in Germancarnica
breedingschemes.
Sevensamplesweretakenfrom preservedspeclmens in the collection of the Bayerische
L a n d e s a n s t a l tf ü r B i e n e n z u c h t ,F R G . T h e y
comprised20 workerbeesfor eachof the colonies
sampledin Erlangen,FRG(n = 6) anOKulmbach,
F R G( n = 1 ) i n 1 9 11 a n d 1 9 1 2r e s p e c t i v e lTyh. i si s
well before any carnicabreedingprogrammewas
in Bavariaand I willreferto thesebees
imolemented
as mellifera although they may vary from
foundtoday in otherlocals 'mellifera'
autochthonou
ities.
Biometry
The rightforewingof each workerwas placedon a
slide (35 x 35 mm) and projectedon a graphical
Thewingvenatabletusinga 1O{oldmagnification.
tion patternwas digitizedbasedon 18 intersection
pointsaccordingto DuPraw(1965a,1965b)and the
datawas processeddirectlyonline.Thirleenangles,
length,width,and the cubitalindexweremeasured'
The lengthand width were proportionallydeterminedfrom the venationpattern(DuPraw,1965b)
and do not representthe actualwing dimensionsas
56
Moritz
d i s c r i m i n a n tf u n c t i o n2
-
3
2
1
0
1
d i s c r i m i n a n ft u n c t i o n1
FlG. 1. Gentroids of the 75% confidence ellipses (method of Cornuet, 1982)using the two
significant discriminant functions 1 (x-axis)and 2 (y-axis).Mating apiary = samples from mating
apiary ; carnica = bees from contemporary commercial carnica stock (Kirchhainl; mellifera =
preserved specimens from northern Bavaria of 1911 and 1912 before the introduction of carnica
bees; non-apiary= p96-6ating apiary samples.
usedby Alpatov(1928)or Ruttner(1988).Thecubital
indexwas determinedby dividingdistanceBC by
AC (seeDuPrawfor pointnomenclature).
The means
of eachsamplewere analysedwith a multivariate
packdiscriminant
analysisusingthe Statgraphics
a g e ( t h i sp r o c e d u r ea s s u m e se q u a lc o v a r i a n c e
matrices).
The biometricalanalysiswas restrictedto
the wingvenationpatternto facilitatebothsimplicity
and speedin dataacquisition,
specimenpreparation
and storage.
RESULTS
Figure1 showsthe graphicalpresentatton
of the discriminantanalysisusingall samples.A. m. carnica
as well as A. m. melliferaform clearnon-overlapping
clusters.The Bavarianbees sampledfrom mating
apiariesand those sampledin other areasoverlap
substantially.
Only one of the non-matingapiary
sampleswas misclassifiedinto the melliferagroup
(table1). However,it shouldbe notedthat the data
might be biasedtowardscorrectclassificationdue
to the smallnumberof coloniesin someof the samotes.
The meansof all measuredcharactersusingthe
nomenclatureof DuPraw(1965b)are givenin table
2. The cubitalindex of the Bavarianhoney bees
(bothmatingapiaryand non-matingapiary)is similar
to the index of oure-bredA. m. carnica bees and
significantlylargerthan that of A. m. mellifera.The
TABLE 1. Glassificationresults of discriminant analysis using three discriminant functions
(the first two functions are highly significant; P < 0.00005)
Actual group
Matingapiary
Non-matingapiary
mellifera
carnica
Predicted group
Mating-apiary
Non-matingapiary mellifera
carnica
26 (68.4%)
15 (28.3%)
0
0
12(31.60/o)
37(69.8%)
0
0
0
0
0
6(100%\
0
1 (1.9%)
7(100%)
0
Purerace breedingin Aprs mellifera
57
TABLE2. Group means of the 16 characterstested.
Character
Angle41
Angle44
Angle84
AngleG7
AngleE9
A n g l eJ 1 0
A n g l eH 1 2
A n g l eJ 1 6
A n g l eM 1 7
A n g l eG 1 8
AngleQ21
AngleN23
Angle026
DistanceLG
DistanceAO
Cubitalindex
Mating apiary
Non-mating apiary
27+3'
30+2'
109+4"
23+1"
24+2
50+3'
20+2'
87+3'
38+3.
92r3'
36+2'
8713"
40+3'
2..15t0.05mm
4.37t0.03mm
z.t zlu.zv
26+4"
30+2'
109+4"
23+1"
2 3 1 1"
49r3"
19+2'
86+3'
38+3"
92+3"
37+2
86+3'
40+3'
1.99t0.04mm
4.36t0.05mm
2.5710.39
intracolonial
distribution
of the cubitalindexwas in
m o s t c a s e su n i m o d a al n d o n l y 1 4 c o l o n i e s( 3 i n
matingapiaries,11 in otherareas)revealedbimodal
distributions.However,in no case was therea stat i s t i c a l l ys i g n i f i c a ndt e v i a t i o nf r o m u n i m o d a l i t y .
Bimodality
is usuallyinterpreted
as a mixedmating
ol carnicaqueenswith both melliferaand carnica
1983).In spiteof this similarityto
drones(Ruttner,
carnica,the wings of the Bavariansampleswere
smallin overallsize(lengthand width)and similar
to those of the melliferasamples.Most characters
are intermediatebetweenthe two oure-bredraces.
However,for two charactersof the Bavarianhoney
bees(anglesJ16 and N23)the distanceto boththe
carnicaand the melliferasamplesis largerthan
betweencarnica and mellifera.Also angle M.17 is
clearlynot intermediate.
AnglesH12,Q21,and 026
show only littlevariationamongthe four groups.
DISCUSSION
In contrastto the beliefof manybee breeders,the
commonlykept honeybee in lowerBavariais not
pureA. m. carnicaas it is currentlymaintainedin
commercialbreedingstock,but actuallya synthesis
of introducedand autochthonousraces.Similar
resultswerealsofound in LowerSaxonyby Reinsch
e t a l . ( 1 9 9 1 )a l t h o u g ht h e i r ' L a n d r a s s es' a m p l e s
more closely resembled the carnlca type.
Discriminant
function2 indicatesthat the Bavarian
bee clustersare not intermediateto melliferaand
carnica.Therefore,the input of other yet unknown
geneticmaterialintothe Bavariangenepool cannot
be excluded.However,geneticdrift,linkage,or simply the small samplesize of the referencecarnica
and melliferasamples may also account for this
onenomenon.
mellifera
carntca
20+1'
32+1"
105+3'
22+1'
20+1"
48+2'
20+0.5"
81!2
35+2'
97t2"
3610.5"
83+2'
40+3'
1.94t0.03mm
4.37i0.05mm
1. 8 6 1 0 . 1 6
29+2"
29+2'
112+2'
24+2
25!2
5312"
19+0.5'
80r2"
32+2'
92!2"
37+2'
82+2'
40!2"
2.06t0.03mm
4.46+0.04mm
2"94t0.34
This evidenceis certainlydisappointingfrom the
viewof the originalbreedingconcept(replacement
of melliferawith carnica).lt also has far reaching
consequences
for bee breedingin general.In spite
of a hugebreedingeffortwith extensiverequeening
operationswilh carnicaqueenscombinedwith a
strictmorphometriccontrolof all selectedcolonies,
it was obviouslynot possibleto replaceone race
with anotherone.Thereare severalootentialreasonsfor this failureof the breedinoconcept.
Mating apiary management and mating
range
The controlledmatingapiariesapparentlyallowed
The densityof bee colonies
too manymismatings.
and beekeepersis high in the study areaand the
numberof coloniesrangedfrom 400 to morethan
1 000 withinthe protectivebelt of the matingapia r i e s .I n t h e l i g h t o f t h e s e f i g u r e sa n d i r r e g u l a r
r e q u e e n i n gi n t e r v a l so f m o r e t h a n t h r e e y e a r s
(Kauhausen,
1991),it becomesobviousthat mating
is not sufficientlycontrolledto guaranteemating
that the mating
withinthe samerace.Considering
distanceof honeybees can exceed16 km under
experimental
conditions(Peer,1975),and the vast
matingrangeof drones(about78 km', Ruttner&
for
Ruttner,1975)the bio-geographical
requirements
matingcontrolbecomeclear.The maintenance
of
a pure-bredprotectivebeltgrowsintoa herculean
task,whichmaysurmountthe limitsof logisticsand
m a n p o w e ra v a i l a b l et o t h e f e w b e e k e e p e r si n
chargeof eachmatingstation.lt is clearlyunrealistic
to use lowlandmatingstationsto maintainpurebred honeybee stock underthe conditionsfound
in the studyarea.
Moritz
58
Biometrical analysis
Bee breederswere well aware of the flaws of the
land-basedmatingapiaries.To overcomethe problemsof hybridization
they usedbiometrical
toolsin
theirselectionwork basedon the work of Ruttner
(1969).The ideawas to identifythe racialtypes by
threebiometriccharactersthat clearlyseparatedthe
autochthonou
s carnica and melIifera populations.
Yet,apparentlythis was not sufficientto eliminate
hybridcoloniesin the breedingstock.This is not
necessarily
surprisingbecauseof the extremeselection intensityfor thesethreecharacters.
lf a colony
did not fulfillthe racialrequirements
it was eliminated, irrespective
of its honeyproductionor any other
behavioural
this leads
characteristics.
Unavoidably,
to a honeybee populationthat matchesthe three
charactersin question.Yet,althoughthe threecharactersfit the carnicastandard,the resultingphenotype may be very differentfrom the autochthonous
carnicaracein manyothercharacters.
Thisis exactl y i n a g r e e m e nw
t i t h t h e p r e s e n td a t a s e t . T h e
cubitalindex(oneof the lhreecarnicatypicalchara c t e r s )o f t h e B a v a r i a np o p u l a t i o nm a t c h e st h e
selectioncriteriareasonably
well.Othercharacters
t h a t w e r e n o t s e l e c t e df o r , i n d i c a t ee x t e n s i v e
hybridization
betweenthe racialtypes.
b e c a u s et h e y d o n o t f i t a c e r t a i nm o r p h o m e t r i c
ideal.
Consequences for bee breeding
in general
T h e c o n c l u s i o n so f t h e p r e s e n ts t u d y a r e n o t
restrictedto the populationin lowerBavaria.They
a r e v a l i d w h e n e v e ro n e r a c e o f h o n e y b e e s i s
plannedto be replacedwith anotherone via landbasedmatingstations.Currentlythis may be germaneto the Africanized
bee problem.sincedrone
controlhas been suggestedas a meansto slow
(Rinderer
et a/.,
down the progressof Africanization
1985).The experiencein Bavariashowsthat even
underradicalbreedingstrategies,in a population
with an extremelysmallnumberof feralcolonies,it
to replacean autochthonous
is virtuallyimpossible
populationwith anotherone by artificialselectionin
a time scaleof morethanfour decades.Both racial
typesin Bavariaarewelladaptedto the temperate
climateand naturalselectionis unlikelyto form a
majordifferential
selectiveforce betweenthe two
races.How difficultwill it be to maintaina population
where
of Europeanhoneybeesin an environment
naturalselectionfavoursa larqeferalAfricanized
oooulation?
Maintenance
of pureracialtypes requiresrigorously
insembee-controlled
areas(islands)
or instrumental
ination.Land-basedmatingstations,likethosestudTheconceptof land-basedmatingstationsseems ied in this paper,can only be usedfor breedingwork
t o b e i n a p p r o p r i a t et o c o n d u c t a n y b r e e d i n g not in connectionwith racialtypes.Neveftheless,
schemerequiring
controlledmatings.As morethan the matingstationscertainlyareof valueto practical
In fact this studyshowsthat breeding
40 years have not been sufficientto replacethe beekeeping.
(e.9.cubitalindex)is poscharacters
autochthonousmelliferaracewith carnicathis pro- for selectable
cedurehas clearlyfailed.In the long run,a limited sibleon landmatingstations.Cornuet& Chevalet
set of biometrical
toolscannotverifythe racialstatus (1987)and Cornuet& Moritz(1989)presentedvarIn spite
for theirusein bee breeding.
o f a h o n e y b e e c o l o n y i f h y b r i d i z a t i o no c c u r s . iousstrategies
Currentlyit is uselessfor the bee breedersin lower of the limitedmatingcontrol,substantialgenetic
Bavariato determine
the cubitalindexbecausethis progresscan be expected,althoughlessthan under
characterno longerdiscriminates
the hybridsfrom a strictcontrolledmatingregime.
ourecommercialcarnicastock.Realisticalternatives
are:(1)discardbiometrycompletelyand establish
a recurrentselectionscheme(for examolessee A C K N O W L E D G E M E N T S
P a g e& L a i d l a w1, 9 8 2 ;M o r i t z ,1 9 8 4 ;C h e v a l e &
t
T h i s s t u d y w a s c o n d u c t e da t t h e B a y e r i s c h e
1982;Cornuet& Chevalet,
Cornuet,
1987)withinthe Landesanstalt
für Bienenzucht
in Erlangen,FRG,
localavailablehoneybee populationor (2)useother am gratefulto E Härtlfor providingthe samplesfrom
biometriccharactersfor the discrimination
between Lower Bavariaand U Bosch for technicalassiscarnicaand the hybridpopulation.However,solution tance.
(2)againcan only be of temporaryvalue,because
of the highprobability
of repeating
the failureof the
lastfour decadesof carnicabreedingwork. In the
REFERENCES
courseof such an attemptit will be increasingly
difW W (1929)Biometricalstudieson variationand raceso{
ficultto identifythe hybridsand finallya resultsimilar ALpAroV
the honeybee Apis mellifera L. QuarterlyReview in
to the one presentedin this paperwillbe obtained.
Biology4.1-57.
In sucha scenario,biometrywouldactuallyhinder
geneticprogressfor characters,likehoneyproduc- BöficHER,F K (19891Haltungund Zucht der Biene. VerlagEugen
U l m e rS
; t u t t g a r tG, e r m a n y3; 5 2 p p .
tion or defensivebehaviour,
that areof interestto the
beekeeper,if productivecoloniesare discarded
Consequences for practical bee breeding
in Bavaria
Purerace breedingin Apis mellifera
CHevnrrr,
C; Conrurr,J I/ (1982)Etudetheoriquesur la selection
du caractere'productionde miel'chezi'abeille.I Modöle
gönetiqueet statistique.Apidologie13: 39-65.
graphiquede populations
CoRNUET,
J M (1982)Repräsentation
multinormalespar des ellipsesde confiance.Apidologie
13: 15-20.
Conruurr,
J M; CHEVALET
C (1987)Etudethöoriquesur la selectron
du caractöre'productionde miel' chez I'abeille.ll Plan
de s6lectioncombin6ede reinesen f6condationnaturelle.
Apidologie18: 253-266.
Conrurr,J M; Moarrz,R F A (1989)Selectiontheoryand selection
programmes./n Moritz, R F A (ed) The instrumental
insemination
of the queenbee. Apimondia;Bucharest,
Romania;pp 125-141
DuPnnw,E (1965a)Non-Linneantaxonomyand the systematics
Zoology 14:1-24.
of honeybees.Systematic
) herecognition
DUPRAW
E (, 1 9 6 5 b T
a n d h a n d l i n go f h o n e y b e e
s p e c i m e n s i n n o n - L i n n e a nt a x o n o m y .J o u r n a l o f
Apicultural Research4(2).7 1-84.
G o E r z EG, ( 1 9 4 0 )D i e b e s t e8 l e n e . L i e d l o f f L
, o t h& M l c h a e l i s :
Leipzig,Germany;200 pp.
GoErzE,G (1949) ImkerlicheZüchtungspraxis.Landbuchverlag;
H a n n o v eG
( e r m a n y1; 7 0 p p .
GoErzE,G (1964)Dle Honigbienein natürlicherund künstlicher
Zuchtauslese.Yerlag
PaulParey;Hamburg,Germany;210
pp.
KAUHAUSEN,
D. (1991)Belegstellenführung
in Bayern(unpublished).
MoRrz,R F A (1984)Selectionin smallpopulationsof the honeybee
(Apis mellifera l.). Zeitschrift für Tierzüchtungund
Züchtungsbiologle101(5):394-400.
PAGE,
R E; LATDLAW,
H H, JR.(1982)Closedpopulationhoneybee
breeding.
2. Comparative
methodsof stockmaintenance
and selectivebreeding.Journalof ApiculturalResearch
2 1 ( 1 \3: 8 - 4 4 .
PEER,
D F ('1957)Fudherstudieson the matingrangeof the honeybee, Apls mellifera. Canadian Entomologist 89:
108-110.
RE!NscH, N; ScHUSTER,H; BIENEFELDK
, i PRcHNEF, F (1991)
Morphologischer
Vergleichvon Völkernder'Landbiene'
in Niedersachsen
mit typischerApls melliferacarnicaund
Apis melliferamellifera.Apidologie 22(1):75-80.
R T N D E BTEER: ,H e l l v r c u R
, L , l l ; D n r u x eR, G ; C o r l r s , A M ( 1 9 8 5 )
Malereproductiveparasitism:a factor in the Africanization
o f E u r o p e a nh o n e y - b e ep o p u l a t i o n s .S c i e n c e2 2 8 :
.t
.l
1 19-1 2.t.
R u r r N E R ,F ( 1 9 6 9 ) B i o m e t r i s c h e C h a r a k t e r i s i e r u n gd e r
Österreichischen Carnica-Biene. Zeitschrift tur
Bienenforschung 9: 469-49 1.
RurrNER,
F (19831Zuchttechnikund Zuchtauslesebei der Biene.
EhrenwirthVerlag;Munich,Germany;141 pp (5thedition).
Rurrren, F (1988)Alogeographyand taxonomyof honeybees.
SpringerVerlag;Heidelberg,Germany;284 pp.
R u r r N E FH
, ; R u r r r e R , F ( 1 5 7 2 ) U n t e r s u c h u n g e nü b e r d i e
Flugaktivität
und das Paarungsverhalten
der Drohnen.5.
Drohnensammelplätze
und Paarungsdistanz.
Apidologie
3ß\:203-232.
59