INS[cr Polr-tN.ut<tr r\ Cittt,L\uot]sES
WHY ARE THERE SO MANY BUMBLEBEES SPECIES IN EYNE (FRANCE, PYRÉNÉES-ORIENTALES,
CERDAGNN)T
P. Rasmon4 E.-4. Durieux, S. IserbVt & M. Barscetti
SUMMARY
With 32 species of burnblebees found in an area of 2000ha, the commune
of Eyne may be the most diversified place in West-Palaearctic, perhaps
also in the whole world. The authors discuss shortly some hypotheses to
e x p l a i n t h i s h i g h s p e c i f i c r i c h n e s s . W i t h i n t h e s a m e c o m m u n e , t h e b i o g e o g r a p h i c a l a n d e c o l o g i c a i c o n d i t i o n s a r e e x t r e m e l y d i v e r s e .
However, the number of species is obviously too high to fit the rules of
resource sharing. The authors suggest that the well known Pyrenean climatic instability brings about a high, random non specific mortality.
This factor, linked with a very high floral diversity and abundance, could
reduce inter- and intra-specific competition, leading to the coexistence of a
large number of species.
INTRODUCTION
The origin of bumblebee diversity has been often discussed. The
most extreme view in this freld has been held by Inouye (1977a, 1977b,
1978). His tempting but misleading sirnplification has been challenged by
Ranta & Vepsâlâinen (1981) who noted that in the tundra and the taiga of
northern Fennoscandia, up to twelve species may coexist. Hanski (1982)
m o d u l a t e d t h e t h e o r y o f I n o u y e b y c o n s i d e r i n g t h a t i n t e r s p e c i f i c
competition for resources plays a major role in the occurrence of the most
abundant species only. Hanski calls these dominant species, the "core"
species, each of them accounting for more than 3 o of the total bumblebee
population. In his view, the "satellite" species - i. e. those accounting for
less than 3 % - would be a rnere "random combination of the resional
s p e c i e s p o o l " .
A c c o r d i n g t o R a s m o n t (1 9 8 9 ) . t h e s e c o n c e p t i o n s o f I n o u y e a n d
Hanski may not apply to the bumblebee fauna of some locations in France,
w'here the local species diversity, as well as the number of "core" species
largely exceeds their predictions. The most species-rich regions in France
8 3
IL L I I t I I I I t T
a
Ta
Z t t
v
ç
u.troLtI Ltll le ql e^
'f lopuxll
Jâ ru ru êlq cs '{
ss r0 H
Ole
^lâ c'D
or
^u J'E
la elg '^
{qrog
sslê^\ zâred 'î
âusâl sroeSrnog
>lc od sy 'X
ênDcâqs J
eq8uo1 aq 'y
Jeqqcoq) ueonol v slêplot0u
9 L L L 8 il VL 9t 0t
7ç
vt
09 a9 98 00 t
6L S
409 vT tl
uç 6t
slâqI-unu sueurceds
uo uâ J'J
uo ue l\
'H ztEl[)
ur llu 'l ê c'c
ssorc 1c ,{
nn '1
)ezseueg '1
ls ro uâ g'u
lâ lâ sê ou 'j' c'd
uoqqsrv Y JJâoqrâ^
ll or ls s'v
p.lecrd nEorohl 'J
uo ue
f H'
^\
or e^
'H 'H 'l
luBlcêl 'c
luotuseu 'd
se LU lê c'u
slâprocêà
sJâqunu suaurceds -saaua.t.i4 uto.tf ( 'sdluuat.to
.tof :^.tdp,to)d.t u!DLli 'l alqru
'(
Z'3 9)
uo s
rlu râ pr su oc
JIlsIunBJ pll€^ lluJêd ol tuercgfns
sr sêleluorJo-seeuerÂ4;o lueugedap eql
3o 8ur:e,,loc aldurus luaserd
aq1 'uorSar aqt paroldxa
,{lpeleedar osle
s€q âH SurleurSuo srqt tuo{ pue uorÊa: fpua-r1e suorlJâlloJ ur âlqelre^e '(I 'qet)
suaurcads êql IIUJo sproJêr
aqt palrduror seq roqlne
]srg âqJ SOOHJEN T lVIYTJVI\I
',(1rsra,r1p srqlJo urSr.to âql tnoq€
seseqlodÂq âuros rno^.uâpuâ
II€qs ê16 âJâq,{\ 'eu8up:e3 lsetuar8 âqt sââqâlqlunqgo Âtrsle,trp uââq suq 'pepJoJâJ
ur t '(
'3tg) au,(g 1o âunurluof,
ârll lnoq€
sr uorleJrunrrllroJ tuaseld aq1
.(a,g0"z
N,zt.zt punoJe)
E N i" ,0 (E ue l p âq a3 pr Su au,6 ç.
p tt
un oJ u) JE T, JE a np ê ss ne J e ql âir l
S'.
-lS :lO HN qll ll) N l NO I- Lb Nl' l'l o.
l èlo .l S, l, lt lll tll ,\
Llt l .lO :^
.) lJ SlN ll\
(lN :l ÀJ V lS lli l^
IC T MH
INsecr Por-r-txATIoN IN GREENHoUsES
Eyne
Figure 1. Map of France and adjacent countries showing the location of Eyne (Pyrénées - Orientales, C erdagne)
Table II. Main recorders of bumblebees in the commune of Eyne (Pyrénées-Orientales, Cerdagne)
Recorder R. DELMAS F. LECLANT P. RASMONT R. BENOIST J. HAMON H. MARION unl*town
specimens numbers
rr82
316 291 2
1 I 75
8 5
'pesfpuu Sureq p1s eru
ulup SurlpseJ eql 'serceds
Je1(oIJ perpunq uuqt eJoru
uo '666I Jeruuns eql
Suunp pe^Jesqo ueeq e^er{
seeqêIqumq LZ1E'euîtJJruoc eql;o uo4eroldxe
cqsrunuJ qEnoroql go lceford
u peu?ls sJoqlne eql
'Jeê srqJ
'Oeil'ç66I 'tuoruseg æ .rerqeg)
erolg
euned oUeJ qU^\ epuru
eæ sduru eq; 'GOet
1e rerq.reg) ''p
erolg Buned
BluC eru^uJos êql qtr,rr
(tuoursug 'd æ redseg 'J
sJord) suory te xnolqtuâC
ep senbruned seguuoq ep
enbuug eqt ,(q pe8euuru
ere etep eql
'(g g0 t ) luoruseg pw (9t6
I ) suutleq'(gçO t ) rrcuesn-D{
ur,{pmur'ern1ere1r1
ur êIqBIrB^e ere Etep
arêd '(II 'qe1) roqlnu
lsrrJ êql pue luBIce.I
sroSuug lsoru) surulêC Surluq sqnser petuêserd ueeq 'seruleq ur 'lotd'(gLil
uâqou 'Jord etel âql ,(q
epsru uêâq enuq eu,(g
uI suorls^resqo lsow
-savuvil7 ut a,utqwog oaptody.to{n,roc
Butqdwos Tuasatd aqr Eutmoqs ,";?';#i;tot
suenrrceds 6TTs arrmqurog Daplodv
oj o
*ffi'
dl ,t ,
C
/_-/
\ --*-^-- '
((i @
\r,ir"éffi
SIISnOHNEEàD Nr NOIJVNI-rIOd UOd
SiIgg[ITgINng dO SJTJSTNOVd ONV
ÀJISàSAICI sHI
INsncr PoLLrN,A.troN rt'r GRETNHoUSES
Lisl species
ol E,"ne up to 1998
Bombus nderarirs (Mùller) B o mbus s i c h e I i i Radoszkowski Bombus hortonm (L.) Bombus pvrenaeus Pérez Bombus st lvestris (Lepeletier) B om bus soroeensis (Fabricius) B om bus me so ntelas Gerstaecker Bonbus lucorum (L.)
Bonbus bohemicus Seidl Bom bus wurfl e nii Radoszkowski Bombus pratonm (L.)
Bombus lapidarius (L.) B ombus rupes tris (Fabricius) B o m bus ge rs ta e c ke r i Mor aw itz B ombus quadricolor (Lepeletier) B o mbus pas cuo runr (Scopoli) B o mbus sub ter raneus (L.) Bombus sylvarum (L.) Bombus flavidus Eversmann Bombus humilis llliger B o mbus nt e ndax Gerstaecker Bombus mucidus Gerstaecker Bombus monticola Smith Bombus hypnorunt (L.) Bombus cont'ilsus Schenck Bombus terrestris (L.) B o m b us ca m pe stris (Panzer) Bombus (A llo ps ith.vrus) s p.
Bombus cullumanus (Kirby) Bombus ruderatas (Fabricius) Bombus magnus Yog!
Bombus muscorum (L.)
Bombus norvegicas (Spane Schneider) Bombus barbutellus (Kirby)
Bombus jonellus (Kirby) Bombus mocsaryi Kriechbaumer
i n P Pyrénées- Orientales
l 1 9 l 4 1 3 5 8 6 256
J I J
534 3 7 8 550 262 464 407 477 t66 l 5 l 87 5 1 3 1 5 5 t 7 6 1 4 3
z t J
87 162 1 6 3 68 5 1 5
62 24 5 56 49 l 9 l 6
'ientules and in
Et,rre 337
t - ) /
1 3 0 126 1 2 0 l l 9 1 1 9 1 0 8 85 76 .70
) t
50 46
À a
J J
) z
30
z o z )
t 6 l 3 7 o
o
4 3 3 2
Nombretotal de spécimens : 8761 1859
Nombre total d'espèces : 36 30 (32-)
* New observations made bv the authors in the summer 1999.
RESULTS AND DISCUSSION
As much as 32 different bumblebees species have been observed in the Eyne commune (Tab. III).
Pekkarinen & Terâs (1993), reviewing the places of maximal
diversity in the West-Palaearctic region. mentions no other place where 32
species can be found on an area as small as 2000 ha.
8l
'lcrloJ IercelS-erd ereur
e se peureldxe
oq louuec ,{lrsre,trp I€Jol
âqJ 'serceds Jrrxepuo s18urs
€ lnoq}r.^a
qSnoqt oql;o ereueSqns 'uorSer crlcJeeel€d-lsolû uec 61 punoJ eq 'eu,(g ur
ZZ
âIIl Jo luqt passeJls aq ,{eur
'mârÀ go 1I
lurod crleueSolfqd eqi ruorg
'sâcuângurJo Â1euen e o1
uedo flrsrenrp eq .(eru ol pâqrrcs€ lecrqder8oe8 eqt Jo uorleool 'su8epre3 eql
Iecol eql go lcedse
su41- 'Dlorrluoa 'g eurdye-oeroq
êql pu€
(snpnnru d'g .t 1n to n. ru '( \ se 1v Iâ e I u s or elu s ou ed rc se ) w'g sa uo op 's 'g
'r un
"t oc
n1 i 'snuSou :seuo u€êueJJêlrpêJ 'sr4saila7 'g) ueedorng-pux '(snuorunnug 'g) :seuo seuo uerJâqrs-oJng'(snuoptdol'g 'g) : -oJnA '(snsntuoc 'g) serceds crtuellv pug
a^\ 'rvrsr^r,;o lurod yecrqder8oeSorq
e {uoJd 'slsryerceds
lcu1s
em yualpnm 'g pue
ua4canlsta8 'g se 'sruos 1nq (wn"tolotd
'wntoncsod)
cr1ce1,(1od ,(13uor1s ere
serceds '(ueyr1 'uor8er âql
ur sedfl uorlela8e,,
go ,ftrsranrp eql ,{q
peureldxe '(psee sr srql
'ure8e arc11 'snsnfuorg
se 'sâuo plag uado
to'untnqLs 'g se 'serceds eurlJâpJog
Jo <sJâllâ,ry\p ]seJoJ
en tuntoud[q 'g se'sercâds
euog 'uJ 006'7, o]
00t'l rrro{ : }crJlsrp êql
go a8uer opnlrllu êprÀ\
oql qlr.ry\
luolsrsuoc ,{lsnornqo sr snll
: serceds IIIq Jo
urelunou eJB sJâqlo leurdleqns
ere sercads etuos 'socueregerd
lecrqderSoe8 Suotuy satceds eql 'eur(g ruorJ pug â1r\ e3ru1 e Jo '{lrsrelrp -ocâ
'âJoq^\âslo uoturuocun lsotu oJ€
tlclq^\ 'snsntuot fqUo,troloN eql sr 1o ecueserd 'uatlcao\sa7 'g snuownl1n) 'g 'g pue
'ûU'qeD Mnro)nl 'g
pue s'Dpaosaut 'unlouoq 'snaDua"td 'g 'g 'g 'sulsaq{s 'g 'slsuaao)os
'g 'Ilaq)IS
'g 'snuDJapnJ snqluog :eru
eu,{E ur serceds luBpunq€
lsou or{J
'flrsrenrp sâoqâlqunq lseq8q eqt
Sutploq s€
poroplsuoc
eq '{uur eu,(g ;o
uor8er eql 'eEpeyrrou) Jno
Jo ot€ls luesard oql
uI ol seredutoc âloq,&\ eql 3o uor8er 'eu8epro3 9€ âJâqÀr uec serceds 'punoJ oq
'luelxâ ur ll
I eunuuroc eu'(g êql
ueql re8rel qcntu sI
'te,te,troq '1ctr1stp
srql'sarceds 0[ stsrl (OOOt)
Ieqzo'feryn13o uor8er turunzrg êq]
uI 'lcr4srp
eu,{g eql Jo seqcrr sarceds âqt
.^Aolâq ,(ets (e86I 'p
drcq7) eturogtle3 p
pus (1661 'sluerllr^(\.; se,{uleurrH
urâlsâ1y\ :eunEJ seeqelqulnq rIâgt
3o ,{lrsrerrrp âql JoJ
u,e\ouI 1ê,^
âJe 'rêrtantoq 'suot8ât Joqlo
o,4rrl 'pa:o1dxe
,{lluercrilnsur ururrrâJ seceld ,(ueut
'ppo^\
âql Jo lsâJ âq}
lod
SA S0 OH NA II N UD I NO IJ VN II IO d UO C S S3 tlE lg t^
ln g :I O S JI JS IN nV C CI NV ^ IlS tl:
{^
l(
l lr HJ
I NSrrcr PoLlrNetroN rN GnEENuousES
The particular situation of Eyne, with its large ecological and b i o g e o g r a p h i c a l d i v e r s i t y m a y p a r t l y e x p l a i n t h e d i v e r s i t y of its
bumblebees fauna. However, the same factors may be present in other
places through Europe, without resulting in the sarne effects. Other factors should therefore be looked for.
The species diversity in Eyne is in clear contradiction with Inouye's theory : there are well over four species in the commune, as well as in any of its biotopes.
The Hanski theory does not fare better: there is no clear-cut
distinction between core and satellite species, and well over four species
account, each, for more than 3 % of the total population. The diversity cannot be simply explained by a sharing of ecological and nutritional resources.
Ranta & Vepsâlâinen (1981) explain the great diversity of the
northern bumblebees faunas by the frequent occurrence of "catastrophic
environmental vicissitudes". These may randomly restrict the populations to levels below the competition threshold, allowing a great diversity of
small populations - and species - to coexist. This explanation is
appealing in the case of Eyne. Indeed, the Pyreneans are well known for their clirnatic instability. The weather may tum from torrid to ice-cold within the same day, with temperature differences of 30' C. and almost daily thunderstorms. Such harsh conditions can be expected to impose a heavy toll on bumblebees populations and make for the kind of situation
described by Ranta & Vepsâlâinen.
There is one difference, however, between the taiga and tundra biotopes described by these authors and the Eyne region. The very abundant and varied vegetation of the Pyreneans may have allowed the coexistence of many species, with a very low level of interspecific competition.
89
nuo"ti ut zlttutD,tow LtaT)aolsnS snqLuogto
uo17nq!,4s!cJ '€ arnar[
sueuloedssTT ynlîADtSnE SnqUIOfl
(,\= JÆ
SESnOHNAIIdD NI NOITVNI-I-IOd àOd
SgîIfllI-IgNng dO SJIJSINnVd CNV
ÀIISUSAIC AHI
INsrcr Polr-rNATroN tN GREENHoUSES
The case of Bombus gerstaeckerl is a special one. This species is a
strict specialist : it is found almost exclusively on Aconitum vulparia. It
lives in the subalpine stages and its distribution is very narrow. In W
Europe, close to 90 o of the recent observations concentrate in three
regions : the High Pyrénées, the eastem Pyrénées and the Alpes-Maritimes
(Fig. 3). More than half of the observations have been made in two
localities of the Pyrénées Orientales: the Eyne valley and the nearby
Madres massif. This species may be one of the more nalrowly localised in
Europe.
Our current research program may shed some more light on the
questions raised here, and confirm invalidate - some of our
hypotheses. Whatever the prospect, we'll keep marvelling at the amazing
diversity of the bumblebees fauna in Eyne.
ACKNOTTLEDGMENTS
The authors acknowledge the OPlE-Languedoc-Roussillon (Prof. J-
P. Lumaret, G. Pinault), the Mairie and M. the Maire of Eyne for their funds and their help.
REFERENCES
BARBIER, Y. & P. RASMONT, 1995. Carto Fauna-Flora - Cartographie des données biologiques - Cartography of biological data. Université de Mons-Hainaut, 93+38 pp,4 disquettes.
BARBIER, Y., & P. RASMONT, 1996. Carto Fauna-Flora 1.2 - Cartographie des données biologiques - Cartography ofbiological data - Nouveautés de la
version 1.2 et utilitaires - Changes in version 1.2 and tools. Université de
Mons-Hainaut,62pp., I CD-rom ^
BARBIER, Y., P. RASMONT, M. DUFRENE & J.-M. SIBERT, 1999. Data Fauna
Flora. Guide d'utilisation. Université de Mons-Hainaut, Région Wallonne, Mons, Namur,94pp, I CDROM.
DELMAS, R., 1976. Contribution à l'étude de la faune française des Bombidae
(Hymenoptera, Apoidea, Bombidae). Annls Soc. ent. Fr. QV.S'), 12:247-290.
HANSKI, 1., 1982. Communities of bumblebees: testing the core-satellite species
hypothesis. Annales Entomologici F ennici, 19 : 65-73.
INOUYE, D., 1977A. Species Structure of Bumblebee Communities in North America
and Europe. pp. 35-49 ft.' Mattson, W.J., The role of Arthropods inforest
ecosystems. Springer, Berlin - Heidelberg - New york.
INOUYE, D., 19778. Resource partitioning in bumblebees. New York entomological
So c ie ty, 85 (4): 253 -25 4.
INOUYE, D., 1978. Resource partitioning in bumblebees: experimental shrdies of
foraging behavior. Ecologr, 59(4): 67 2-67 8.
9 l
r0Z- | :(l)09' ' (G o1
out otu g)
(ûo7sg17 7otn7o61)
wnasnyg qs14tg aryto uuailng
'(rurqruog'eeprdy
:ereldouaufg) e,{eleug rltur{se) eqlJo
soâq elqunq eqJ 't66
''H'd 'SWVI"I'II1y\ t
61 'dd
',{e1eryeg'sser4
erruoJrle3 Jo flsrantun'
@op1 dV : otaldouaw{17) o1utot11o3
Jo saa I oolzr1)
puD saag anwng'c86t 'DNtNNnc
'l"I
? uf DNtNuoH
's'c ''AÏu 'duoHJ
71 ec 'dd
'1961 erqruenou'sttu4 ',,anbltslunottgtVlul
purtÊ ap sandsg,p
aJuDilD^ins o7 n uolloc1fltuap11 tnod satq?ua^ultp
sasgntutoJu-r sarloua^ul
sap uolt0stlttn T,, uunBIAI
'H t?
uoJnsâg ep'1 :u!'uoutBlB(lse^ dt Blued
ep euogql erluoc e,(nou1,p
aFoggl '(aepldy'ereldoueru,(g) ecuerg 3p
suoprnog
sep euneJ e1 ap gla.rnned
op ser1uâc le ossaqcu
âp serlueJ '686
''d 'JNOI^trSYU I
Itxt 'dd
+ 0l
'xnolquleg 'xnolqurag €
ep senbruouorEe secuetcg sop
s?llnceC '1ero1cop
âp uorlepossrcl 'Qowqutog 'anpldy
'otaldouaut11) anb8lag ap ta
æuo.ti
ap suoprnog sap anbrt1dotSoaSooz
p anblSolocV arydo.tSouoyg
'8861 ''d 'JNOI
trS1trU lerodurel pue ,{ileueEoreleq t 'tgO {qlf âegelqunq tuâquou oreqt ârB ,{uetu os 'sorp6r 'serltunrutuoc -oged5 ;serceds 'Vt-gZ igt
'NENIyIVSdE^
4l ')
'g 'VINYU
'802-t8 t :gg'rctuuag 1ooz
'uuv'(eepqdy 'eteldoueru,(g) edorng uâ1se^\qlrou
u snt,(qils4 pue snqwoq
Jo ,{qderSoeSooz't661 'SYUAI 'l
'V 'N!INIUV))!Id ?
'Zgl-tsl'ununztg '066 t znluwal 67-91 ptBuoy
1!o1o,t1g losnln 'X'rs€uneJ
(eaprody :ereldoueruf g) ue
uelEep uuzedeBny â^ uê{opu€led'066
I''H'y.,Eg;ZO suottcâlloc :(72)9'nluotn a6t'sêsrepuelrê?u I'0ll-l9l 'ld
sâl suep s,xqluog erueS
np suegugr,{d suoprnoq sel
rns soloN '8ç6
''C 'NVI t
IAS1U)
sa sn oH Ng gu I N D N OT TV NI -I 'I Od UO C S AA SE 'I SN nB S CO JI JS IN nV C C NV ^J IS UE
^I C A HJ
INSE,CT POLLINATTON IN GREEI{HOUSES
Edited by Marinus J. Sommeijer and Aad de Ruijter
Proceedings of the specialists' meeting held in Soesterberg,
The Netherlands, 30 September to 2 October 1999
ureBs,r,tnar5l'serlcnpord âqcsrJelD 1h,r,u1ur1E-eppn€l
..lutrd Xqnceg :u8rseprerro3 '11 ',{3o1org;o droq ue,r;a1er4 .lotl1 ue^ Sursseco:da8eul 'u8rseq pue
:â^eoqsnrsorquy :oloqd .lo^\olJ Âueq,trerls
Burlrsr,t ooqelqung ..râ^oJ
.JSggOru'JTgddOX'Sggg NYWXNIV{ )NIJNN{,NYXNT'NOIJVDOSSY
4OVVJ O4IS HJJnq
:Suqaatu aLfl parosuods suounrusur
Buruollotaql 'AlrsJe^run
1{Je4n tcâsul JoJ roJ uorssruruoJ âC uorleurlyod ââg tueld suor}ulâU â^âoqsnrsorqurv q (Ugdll) âql pue sjcêsul lerJoS qlr.,rl uorleredooc otnlusq oql go lueugede6l
Isuorl€uretuJ âqlJo uorlcos uorl€urllod âqt
pus uorluurllod pu8 srol{
snorêJrTlâhJ
roJ uorssruuo3 Surpuelg VICINOhtrIdy
eqt,{q pezwe8rc se,r Buqeeru
srql
suorurdo âql JoJ
Jo uortcnpord aqt
qlt./l,r pecnporder
'sJoqlne âql qlr^\ slsor
sâlcru" aa,rlcedser eql ur
pesserdxe
pue luêtuoc orll JoJ
,{lpqrsuodser ls8urpeecord eseql
JoJ posn âJe,ry\
sroqlne eql ,{q pouruIqns s1xa1 1ur3e1ur
eq1 ueilrJ./y\ Jo uorssrrurad 'sJolrpg
aq ,(eu looq srqt
Jo sged 'pe^rêsor slq8rr
IIV 0002 O
Surdeeleeq'sesnoqueolS'seeqelqrunq'uorl€urllod :s8urpeeq lceiqng
8-t-668ç08-06 NgSr
re1[nA op pBV pue refreuuos .f
snurre6 :.spo
I 6661 0E o1 requreldes reqotco 7 ,{uprn1e5
'spuelrâqtâN eq1 ,8:eqretsâos uI plâq
Sutloeu .slsryerceds eqt.Sto
sSurpaacord :sosnoquââJ8
ur uorleurllod 1cesu1
tseeH uâc'Tââqlorlqrg êItrpluruo) vJVC-dI:)
