The phylogeny of the genus Leucorrhinia and the evolution of larval spines (Anisoptera: Libellulidae)

8 Pages·2000·0.97 MB·English

by  Westman, A

Odonatologica29(2): 129-136 June 1, 2000
Thephylogenyof the genusLeucorrhinia
and theevolutionof larval spines
(Anisoptera: Libellulidae)
A.Westman,F. JohanssonandA.N.Nilsson
DepartmentofEcology and Environmental Science, Umeå University,SE-90187 Umeå,Sweden
Received March22, 1999/Revised andAcceptedSeptember1,1999
A cladistic analysis ofthe genus Leucorrhinia, based on adult morphological
characters, found onemostparsimonoustree with aconsistencyindex of0.35, The
evolution oflargedorsallarval spineswasmappedontheresultingtree.Thismapping
suggeststhat thepresence ofspinesistheprimitivestatewithinLeucorrhinia andthat
theyhave disappearedonfive differentoccasions,orhave disappearedtwice onlower
branches andreappearedthree timeshigherupin the tree.
INTRODUCTION
Odonatelarvae are common prey ofmany predator groups:birds (KENNEDY,
1950) fish(CROWDER&COOPER, 1982;RASK, 1986),andotherinvertebrates
(PRITCHARD, 1964).As prey they showavarietyofbehavioural(PIERCE, 1988;
McPEEK, 1990),and morphological adaptations (JOHANSSON & SAMUELS-
SON, 1994;McPEEKetal., 1996) against predation.
Thepresence of spines on the body has been shown to provide an effective
protection against predators in many animal groups (EDMUNDS, 1974;HAVEL
&DODSON, 1984;ABRAHAMS, 1995). Manyodonategenera,especially among
the anisopterans, have larvae with prominent lateraland dorsal spines on the
abdomen(WALKER, 1958;WALKER & CORBET, 1975;ASKEW, 1988).In
odonates, the larval spines mayfunction as an aidin climbing and sprawling in
vegetation andon the substrate (DA SILVA AGUIAR, 1989)and as protection
againstpredators (JOHANSSON&SAMUELSSON,1994).Currently thereissome
supportfor thepredator defencehypothesis in Leucorrhiniadubialarvae. Larvae
ofL. dubia grow longer spines in habitats with fish, fish have longer handling
times when eating long-spined larvae,andfish inducedifferencesin spine shape
underlaboratory conditions(JOHANSSON&SAMUELSSON, 1994;ARNQVIST
130 A.Westman, F. Johansson &A.N. Nilsson
& JOHANSSON, 1998).Considering thepotential useofthesespines itwouldbe
interesting to know how they have evolved in a genus. Being potentially highly
adaptive andconnectedtohabitatshifts, convergentevolutionofspinesis expected.
Inthisstudy we focus ontheevolutionoftheabdominalspines in larvaeofthe
genusLeucorrhinia(Anisoptera,Libellulidae,Sympetrinae), inwhichsomespecies
have prominent spines while others have very small spines on the abdomen
(WALKER&CORBET, 1975;ASKEW, 1988). The genusconsistsof 14species
andseveralsubspecies (DAVIES & TOBIN, 1985). Thepurposeofour study was
to provide aphylogeny ofthe genusandmaptheevolutionoflarval spines on the
resulting tree.
MATERIALANDMETHODS
Weused 14taxainthe genusfor ouranalysis (Tab.I). L. ussuriensis Bartenef wasexcluded from
the study since nospecimens wereavailable. In addition,recent results suggest this species tobe a
synonym ofL. orientalis Selys (MALIKOVA, 1995). Since L. dubia orientalis Belyshev has been
TableI
The speciesusedin thisstudy,theirgeographicdistributionandinformationabout the specimens'sex
andwherethey werecollected.- [Vb=Vasterbotten;- Vrm=Varmland;- Upl=Uppland;- Dlr=Dalarna;
- Got=Gotland; - Vstm=Vastmanland; - Vg=Vastergotland],- Celithemis and Sympetrumwereusedas
outgroups
SSppeecciieess DDiissttrriibbuuttiioonn NNuummbbeerraannddoorriiggiinnooffssppeecciimmeennss
LL. aallbbiiffrroonnss((BBuurrnmt..)) EEuurrooppee I1d3,,VVbb;;l1dd,V,Vrrmm;;I1dd,,UUppppll;;22$9,,GGoott,,SSwweeddeenn
LL..bboorreeaalliissHHaagg.. CCaannaaddaa--NN.. UUSSAA 22cdJ,,YYuukkoonn TTeerr;; 1192,,SSaasskkaattcchheewwaann;; 1129,,
MMaanniittoobbaa,,CCaannaaddaa
LL.. ccaauuddaalliiss ((CChhaarrpp..)) EEuurrooppee 22<dJ,, 2229,,NNoorrwwaayy
LL.. dduubbiiaadduubbiiaa((VVaann ddeerrLL..)) EEuurraassiiaa 22dd,, 1129,,VVbb,, SSwweeddeenn
LL. dduubbiiaaoorriieennttaalliiss SSeell.. SSiibbeerriiaa--JJaappaann 22d3,,2229,,HHookkkkaaiiddoo,, JJaappaann
LL.ffrriiggiiddaaHHaagg,. NN.. AAmmeerriiccaa I1d6,, 1129,, OOnnttaarriioo,,CCaannaaddaa
LL..ggllaacciiaalliiss HHaagg.. NN,,AAmmeerriiccaa I1d3,,2229,,OOnnttaarriioo,,CCaannaaddaa;; IIdd,, 1129,,CCaalliiffoorrnniaia,,
UUSSAA
LL.. hhuuddssoonniiccaa ((SSeell..)) NN..AAmmeerriiccaa 333d,,3329,,QQuueebbeecc,,CCaannaaddaa;; 22d3,,CCaalliiffoorrnniiaa,,UUSSAA
LL. iInnttaaccttaa HHaagg.. NN..AAmmeerriiccaa 1192,,OOnnttaarriioo;;22d3,,2229,,QQuueebbeecc,,CCaannaaddaa
LL. iinntteerrmmeeddiiaa BBaarrtt,. AAssiiaa 22d3,, 1129,,HHookkkkaaiiddoo,, JJaappaann;;1129,,RRuussssiiaannffaarrEEaasstt
((HHeellssiinnkkii MMuusseeuumm))
LL..ppaattrriicciiaa WWaallkkeerr CCaannaaddaa 22d3,,2229,,OOnnttaarriioo,,CCaannaaddaa
LL..ppeeccttoorraalliissCChhaarrpp.. EEuurrooppee IIdd,, DDiirr;; 1129,,UUppppll;;IIdd,,1192,,VVssttmm;;IIdd,, 1129,,
VVgg,,SSwweeddeenn
LL.pprrooxxiimmoo CCaallvv.. CCaannaaddaa--NN.. UUSSAA 22dd,, 1129,,OOnnttaarriioo,,CCaannaaddaa
LL. rruubbiiccuunnddaa ((LL..)) EEuurraassiiaa 22dd,,2229,,VVbb,, SSwweeddeenn
CCeelliitlhheemmiisseeppoonniinnaa((DDmm..)) NN., AAmmeerriiccaa IIcdJ,, 1199,,FFlloorriiddaa,,UUSSAA
CC.. oormnaattaa((RRaammbb..)) UUSSAA IIdd,, 1129,,FHloonriddaa,, UUSSAA
SSyymmppeettrruummddaannaaee ((SSuullzz..)) HHoollaarrccttiicc 33dd,,1192,,VVbb,, SSwweeddeenn
SS.. oobbttrruussuumm ((HHaagg..)) NN..AAmmeerriiccaa 443d,,2229,,OOnnttaarriioo,,CCaannaaddaa
Phylogeny ofLeucorrhinia 131
treated asavalidspeciesby someauthors(BELYSHEV, 1973; ISHIDA, 1996),weincluded this taxon
inthe analysis. To generateahypothesis aboutthe phylogeneticrelationship among the specieswe
performedacladisticanalysis using 24adultcharacters(Tab.II).Ofthesecharacters, 7wereexamined
inbothsexes, 14inmalesonly and3 infemales only.Statesformostcharacters wereobtainedfrom
multiplespecimens(Tab.I).We included fourspeciesin the outgroup (TabI):the nearcticCelithemis
eponinaand C. ornata(Leucorrhiniini),and fromSympetrinithe twospeciesSympetrumdanae and
S. obtrusum.
Character state distribution was entered in MacClade 3.03 (MADDISON & MADDISON, 1992),
andthereafterthematrixwasfurtheranalysedinboth MacCladeand PAUP 3.1(SWOFFORD, 1993).
Allmultistate characters weretreated asunordered. Ageneralheuristic searchwas doneinPAUPin
order tofind the mostparsimonious tree(s).Branch support was tested with bootstrappingin PAUP
(FOREY etal„ 1992).
Theoccurrenceofspineswastaken fromthe literature(WALKER&CORBET, 1975;CANNINGS
&STUART, 1977;ASKEW, 1988;NORLING&SAHLEN, 1997),andsomeunpublishedobservations
wereprovidedby S.Cannings.
Table II
Character and characterstates used inthecladisticanalysis
Characters present inboth sexes
(1) Labium,colour: (0)atleast outer marginlight; (1) all black
(2) Fore wing,number ofcellrows between IR3 andRspl: (0) 1;(1)2
(3) Fore wing, MAandCuP:(0)parallel;(1) divergent
(4) Base ofhind wing: (0)without distinctblack spot; (1)with distinct blackspot
(5) Metatibia,denserow ofspinesbelowtheknee; (0)absent;(1)present
(6) Abdominalsegments 2-4: (0)normal; (1)whitepowdered
(7) Abdominal superiorappendagescolour: (0) dark;(1) white
Characters present inmales
(8) Thorax, colourofsetation: (0)light;(1)mixture oflightand dark
(9) Thorax, colour: (0)notmetallic;(1)metallic
(10) Mesepistemumadjacenttomesepimeron,colour:(0)brown; (1)black
(11) Mesepimeron,shapeofbrownfield: (0)notwiderthantall;(1)wider than tall
(12) Metepimeron,colour:(0)predominantlyblack; (1)predominantlybrown
(13) Anterior lamina,posterioredge: (0)straight; (1)splitor excavated
(14) Hamulus,lateral outgrowth:(0)absent;(1)present; (2)present withanindentation
(15) Hamulus:(0)withoutspines; (1)withspines
(16) Hamulus,anterior tips:(0)bentoutwards;(1)notbentoutwards
(17) Hamulus, posterioredge: (0)bent upwards; (1)notbent upwards
(18) Genital lobe: (0)without spines;(1)with spines
(19) Abdominalsegment7, colourofdorsalspot: (0)red;(1)yellow;(2)absent
(20) Abdominal segment9,ventral placementofgonopore: (0) displacedforward;(1)in themiddle
ofthesegment
(21) Inferiorappendage,form:(0)sidesconvergingposteriorly;(1)sidesparallel;(2)sides diverging
posteriorly
Characters present infemales
(22) Vulvar lamina,form:(0)longerthan wide;(1)aslongaswide;(2)widerthan long
(23) Vulvar lamina,direction ofthe median gonapophyses:(0)backwards; (1)sidewards
(24) Vulvarlamina, surfacebehindthemediangonaphyses:(0)nohairs; (I)withhairs
132 A.Westman, F. Johansson & A.N. Nilsson
RESULTS
Thecharactermatrixis showninTableIII. Thecladogram presented inFigure 1,
isthesingle shortesttree foundby PAUP(TL79,Cl0.354,RI0.545).Thebootstrap
testprovided statisticalsupportonly for theingroup (79%), andtheL. albifrons +
L. caudalis clade (51%).
Thepresenceoflargelarval spines is not restrictedto asingle branchonthetree
(Fig. 1). Applying Farris optimization (FARRIS, 1970) to the tree gave several
equallyparsimonious solutions,inallofwhichthespinesarepartoftheLeucorrhinia
groundplan. Eitherthe spines have disappeared onfive differentoccasions within
thegenus, orthey have disappeared twiceonlowerbranchesandreappeared three
timeshigher upinthetree.Asingle disappearanceofthespines wouldincreasethe
tree length byat leastseven steps.
DISCUSSION
It is clear from the phylogeny, that the presence of large larval spines is not
restrictedtoasingle clade.McPEEK(1995) suggested thatchanges inmorphology
ofdamselfly larvaewere associated withhabitatshifts.After such ahabitatshift,
selectionpressureexertedbytheenvironmentshouldfavourthemorphologies best
adaptedtothenewenvironmentalconditions.Undertheassumption thatlargespines
are beneficialin someenvironmentsbutnot inothers, themultiple disappearance
ofspines in the tree suggests several such habitatshifts. These shifts might have
triggered a reduction of the spine length. Currently we do not know what
environmentalconditionsthatcause the reductionoflarge spines. We know that
fish predators have difficultiesin handling L. dubia larvae with large spines
(SAMUELSSON & JOHANSSON, 1994),butwe do not know the disadvantage
ofhaving long spines, besidesa general cost inproducing them.
Thefact that L. dubialarvae show very low abundancein thepresence offish
compared to in the absence offish (HENRIKSON, 1988; SAMUELSSON &
JOHANSSON, 1994), suggests that spines are not very effective against fish
predators. However, arelevantcomparison inthiscontext wouldbebetween spine
and spineless species withregard toabundance with fish. Itis interesting to note
that the longitudinal darkbands present on the ventral side ofthe abdomenis
negatively correlatedwith spine length in the genus. Whilethe correlationis less
clear amongthe palaearctic species, a strong negative correlationexists forthe
NorthAmericanspecies (WALKER & CORBET, 1975;NORLING & SAHLEN,
1997).Crypticcolorationhasbeenshowntobeaneffective defenceagainstpredators
inmanyspecies (EDMUNDS, 1974),butas farasweknow, nostudieshavelooked
attheeventualfunctionofcryptic colorationinodonatelarvae.Moreworkis clearly
needed,especially withregard toabundancepatternsofspinedandspineless species,
larvalcolorationandhow thesearerelatedtoenvironmentalconditions.
PhylogenyofLeucorrhinia 133
mark 2244 ? 0 1 1 0 1 0 0 0 0 1 1 1 1 0 1 1 1
question 2233 0 0 7 0 1 1 0 1 1 1 1 0 1 1 1 1 1 1
2222 0 2 10 2 1 0 1 0 0 2 1 0 0 1 1 2 2
a 1010100001
by 2211 0 0? 0 0 1 0 1 1 0 1 2 2 1 1 1 1 1
denoted
2200 0 0 00 10 1 0 1 1 1 0 1 0 0 1 0 0
1000???100000210010
are 1199 0 2? i 2 0 2 0 0 2 2 0 1 0 0 1 0 0
certainty 1188 000000000000000000000000? 00000000001010100002002000000100?00000010010?0?1?10 00011111011101101002102100 1 01011100000101100000001111 1011111110011101012010000 1011100100100101000011010 1111100101100111000110110 10011100100011111111110010111020011110010002112101 1011100100100111010021011 010111001011002001000101111011100100111211110011010 101110010110020110111111 101110010011110011001211 101110010110100010001211
1177 0 0 1 1 00 0 0 0 0 0 0 0 1 1 1 1 1
with 1166 0 0 0 0 10 1 1 1 0 0 1 1 0 1 1 0 0
determined 1155 0 0 0 1 00 0 0 1 0 1 1 1 0 1 0 0 0
1144 0 1 1 1 1 1 1 1 1 0 0 1 1 2 2 2 1 0
be 1133 0 0 0 0 1 1 1 0 0 1 0 0 1 0 1 0 1 1
not
1122 0 1 0 1 0 0 1 0 0 ? 1 0 1 0 1 0 1 0
could
1111 0 0 0 0 1 1 0 1 1 7 1 1 0 1 1 1 1 1
that 01010000010101
HI
1100 0 1 0 1 1 00 0 1? 0 0 0 1 0 I 0 1
Table
Characters
99 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0
88 0 0 0 0 0 01 1 1 0 1 1 1 1 1 1 1 1
species.
77 0 0 7 0 1 01 0 0 00 0 0 0 0 0 0 0
66 0 0 0 0 1 01 0 0 10 0 0 0 0 0 0 0
examined
55 0 0 0 0 1 1 1 1 1 11 1 1 1 1 1 1 1 1
the 44 0 0 1 1 1 1 1 1 1 11 1 1 1 1 1 1 1 1
among 33 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1
101
states 22 0 0 0 1 0 00 0 1 0 1 0 0 0 0 0 0 0
11 0 0 0 0 0 1 1 1 1 1 1 I 0 1 1 1 1 1
character
the
nnoo..
of
distribution SSppeecciieess//CChhaarraacctt.. oorriieennttaalliiss
oobbttrruussuumm ddaannaaee eeppoonniinnaa oormnaattaa aallbbiiffrroonnssbboorreeaalilsis cacuaduadlaislis dduubbiiaa ffrriiggiiddaa ggllaacciiaalliisshhuuddssoonniiccaa Iinnttaaccttaa iinntteerrmmeeddiiaa ppaattrriicciiaappeeccttoorraalliiss pprrooxxiimmoorruubbiiccuunnddaa
dd.. dd..
The
SS.. SS.. CC.. CC.. LL. LL. LL. LL.. LL.. LL.. LL. LL. LL. LL.LL. LL. LL.LL..
134 A.Westman, F. Johansson &A.N.Nilsson
As seen in the low Cl and
lack of strong statistical
support, our phylogenetic
hypothesis is weak. A more
firmly based phylogenetic
hypothesis would probably be
possible tofindbyadding also
characters from larvae to the
analysis. However,thiswasnot
possible at this stage of our
investigation duetothelackof
larval material of several
species. Adding only the
abdominalspine character to
the analysis resulted in four
shortest trees, one of which
was the same as the tree in
Figure 1. Theotherthreetrees
differedonly intheposition of
L. borealis and L. rubicunda
,
showing thesame dynamicsin
the evolution of the larval
spines as our firsttree.
The best supported mono-
phyletic group within the
genus is the L. albifrons + L.
Fig. 1.The shortest tree found. The presence (*) oflarge caudalisclade, supported by
abdominalspinesinlarvaeis shownfor each terminaltaxon. characters no. 7 (abdominal
superior appendages), 9(thorax colour) and 16(hamulus anteriortips).This clade
isgroupedtogetherwithL. frigidaby characterno. 13(posterioredge oftheanterior
lamina)andno. 6(abdominalsegments2-4 white-powdered). As L. albifrons and
L. caudalis lookvery muchalikeintheiroverallappearancewithwhite-powdered
abdomens, itis interesting to note that they also have genital similarities. An
interestingthing tonoteinthetreeis thatL. dubiadubiaandL.dubiaorientalisdid
not grouptogether as suggested in some odonata lists e.g. DAVIES & TOBIN
(1985),HARITONOV &MALIKOVA(1998).InsteadL. dubiaorientalisappeared
asthesister-taxonofL.glacialis.Themonophylyofthisgroupdependson character
no.2, sayingthatthesetwo species are theonly onesthathavedoublerowsofcells
betweenIR3 andRspl., and characterno. 15 (spines on hamulus).
Phylogeny ofLeucorrhinia 135
ACKNOWLEDGEMENTS
Wewould liketothank HANSOLSVIK,GORANSAHLEN,MATTIHAMALAINEN andLARRY
HULDEN fromHelsinki museum,andTOHROYOKOYAMAforprovidinguswithadultspecimens.
SYDCANNINGS,GUNTHERPETERS,DENNISPAULSON, andHIDENORIUBUKATAprovided
valuable species information. Parts of this research were supported by aNational Sciences and
EngineeringResearch Council ofCanadatograntLockeRowe(University ofToronto),andaSwedish
Natural Science Research Council postdoctoralfellowshiptoFrankJohansson,
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