Odonata diversity in a tropical dry forest of Mexico, 1. Sierra de Huautla, Morelos

2008·2.1 MB·English

by  González-Soriano, E

Odonatologica37(4):305-315 December 1,2008
Odonatadiversity ina tropical dry forest of
Mexico, 1. Sierra deHuautla, Morelos
E.González-Soriano¹,F.A.Noguera ²S,.Zaragoza-Caballero¹,
M.Á.Morales-Barrera¹,R.Ayala-Barajas²,A. Rodríguez-Palafox²†
andE.Ramírez-García²
1DepartamentodeZoologla,InstitutodeBiologla,Universidad Nacional Autonoma deMexico,
Avenida Universidad 3000,MX-04510 Ciudad Universitaria,D.F,Mexico
2EstaciondeBiologiaChamela,InstitutodeBiologia,Universidad Nacional Autonoma deMexico,
Km. 59CarreteraBarra deNavidad-Puerto Vallarta,MX-48980 San Patricio,Jalisco Mexico
t deceased
Received October19,2007/Reviewed andAcceptedJanuary,21, 2008
Astudy ofthefaunaofOdonata ofatropicaldeciduous forestispresented.Col-
lectionsweremademonthlyduringa1-yrperiod(Nov. 1995-Oct.1996)during5days
eachmonth. Atotalof2595adult specimenswerecollected,belongingto57species,
33generaand8families. Estimated richnessvalueusingthenon-parametricestima-
torICEwas76.28.
INTRODUCTION
This paperispartofalong-termproject to studythe diversityandpatternsof
distributionofthefaunaofinsectsassociatedwiththetropical dryforest(TDF)
inMexico (NOGUERA etal., 2002; ZARAGOZAetal„ 2003).
TheTDFisoneofthemostdiverseecosystems inAmericaandalso oneofthe
mostendangered (JANZEN, 1988). In Mexicoitcovers almost8%ofthecoun-
try’s surfaceandharborsa large numberofendemic species (TOLEDO & OR-
DONEZ, 1993;FLORES& GEREZ, 1994;CEBALLOS& GARCIA, 1995).
Thisplant community extendsinamoreorless continuousfashionfromSonora
to Chiapas stateswithotherintrusionsintheYucatanPeninsulaandinsomear-
easoftheGulfofMexico(PENNINGTON& SARUKHAn,2005). Since 1990
only 27%ofitssurfacehasremainedintact(TREJO &DIRZO,2000).
Odonataare predacious insects whichdo not depend directly upon plants, at
leastinthetrophic level, as mostoftheotherinsectsdo. However, they useveg-
etationin severalotherways: e. g.as breeding sites forlarvae(e. g.phytotelmata
306 E.Gonzalez-Soriano et.al
andground litter)(PAULSON, 2006) orforshelter, oviposition, roostingsites or
habitatselectionfor adults(BUCHWALD, 1992;CORBET. 1999; PAULSON.
2006). Vegetation isalso usedforconcealing adultsfrompredators and as mat-
ing areas and feedingperches (BUCHWALD, 1992).Thestudy ofthe diversity
anddistributionofOdonataassociatedto the TDFisalmost non-existentand
only a few published works dealwith this topic for Mexico(DUNKLE, 1976;
GONZALEZ-SORIANO etal., 2004).
MATERIALANDMETHODS
STUDY AREA - Thestudyareaislocatedin aportionoftheSierra ofHuautla BiosphereRe-
serve(SHBR), locatedin southcentralMorelosState,Mexico(l8"20T0”to18°34'20”Nand98o51'20”
to99°08T5”W)(DORADO, 1997).Theclimateis warm subhumid,typeAwo"(w) (i') gaccording
theKoeppenclassification modifiedbyGARCIA(1981).Averageannualprecipitationfortheperiod
1981-1997 was 824mm,with80%oftherain fallingfromJune toSeptember.Mean temperaturefor
thesameperiodwas24.7°C,withanaveragemaximumtemperature of34.3°Cand anaveragemini-
mumtemperatureofI5,0°C (CNA.2000).Thesiteisincluded within theAtoyacand theAmacuzac
riverbasins. The areaiscovered mostlybyTDFalthough otherplantassociations arealso found
here(e.g.galleryforest,secondaryvegetationandagriculturelands).
Odonatecollections weremadeatthree sites. The first ofthese islocated alongthe Quilamulariv-
er(18°27’671”N,99°02’475”W)nearthe facilities ofthe Center ofEnvironmental Education and
Research Sierrade Huautla (CEAMISH),the second alongacanyon made by theAjuchitlanriver
(18°27’065”N,98°59’546”W)nearthetownofAjuchitlan andthe lastone(ArroyoLos Idolos)ata
secondarystreamconfluentwiththeAjuchitlanriverneartheentranceofthetown.The firstsite(Qui-
lamulariver)haswaterthroughouttheyearspeciallydownriverduetothepresence oftheLorenzo
Vazquezdam. Thelasttwo streamslackwater duringthe dryseason.Collections weremadeduring
fivedays ofeverymonth(betweenNovember 1995 and October1996)andtheywerecarried outby
twopersons; theseniorauthor and onestudent(Ma.delos AngelesMorales),exceptin September
whencollections weremadeonlybythestudent.
ANALYTICAL METHODS—Thevaluesofrichnessand abundance correspondtothenumber
ofspeciesandindividuals recorded. Diversity andevennesswereanalyzedwithShannon Index,using
thenaturallogarithm,andvalueswereobtainedwiththeprogramBioDiversity Pro(McALEECEet
al., 1999).
Consideringthat theobserved numberofanysampleofindividuals fromaspeciesrich community
underestimates the truenumber ofspeciespresent,anon-parametricestimator ofspeciesrichness
wasperformedusingdata obtained duringthe yearofthestudyto determinehowclosetherichness
value recorded was representativeofthe truelocal richness. The estimatorused wasICE,aninci-
dence-basedestimator,becauseit bestsatisfiedtherequirementsforanideal species-richnessestima-
tor (CHAZDON etal., 1998).ICEestimator isbased onspecies found in < 10samplingunits and
itsformulais(COLWELL,2006):
S.ice=Sfreq +Sq.m,fr+£0. yl,2ce
ice ice
where Sf isthenumber ofspeciesfoundin> 10samples,Sinfristhenumber ofspeciesfoundin 10or
fewersamples; isthesampleincidencecoverageestimator.Q,isthenumberofspeciesthatoccurin
only onesampleand-y2iceistheestimated coefficient ofvariation oftheQ,’sforinfrequentspecies.
Theestimates werecalculated using Estimate S6.0bl (COLWELL,2006).The speciescollected
within eachdaywereconsidered onesampleunit(53intotal).
Odonatadiversityintropicaldryforest 307
Forphenologyanalysisofthedata,weconsidered therainyseasontolastfromMaytoNovember
and thedryseasontolast fromDecember toApril. Thiswasbased ontheoccurrenceofindividual
storm events totalinggreaterthen 15mm,because the canopyintercepts smalleramounts almost
completely(CERVANTES, 1988).
RESULTS
RICHNESS
Atotalof2595adultspecimens werecollected, belonging to57species, 33gen-
eraandeightfamilies. Thesevaluesrepresent61%ofthespecies, 82%ofthegen-
eraand89%ofthefamiliespreviouslyreported forthestateof Morelos, Mexico
(GARCIA, 1987;G0NZALEZ-S0RIAN0& NOVELO-GUTIERREZ, 1996,
2007; NOVELO, 1997).
Thefamilywiththegreatestnumberofspecies wasLibellulidaewith27species
(47% ofthetotal),followedby Coenagrionidae with 16(28%), Aeshnidaewith 7
(12%), Gomphidae with3(5%)andCalopterygidae, Lestidae, Platystictidaeand
Protoneuridaewith oneeach(2% each).
Libellulidaewas also thefamily withthe greatest numberof generawith 16
(48%)followedbyCoenagrionidae andAeshnidaewith5each(15%each), Gom-
phidae with3(9%); Calopterygidae, Lestidae,Platystictidae andProtoneuridae
withoneeach(the remaining 13%).
Argiawas themostdiversegenuswith9 species(15.8%) followedby Enallagma,
Erythrodiplax, Dythemisand Micrathyriawiththreespecieseach(21%), Ischnu-
ra, Rhionaeschna, Gynacantha, Libellula, Macrothemis, Orthemis,Perithemisand
Trameawithtwo species each(28.1%). Therest (21) ofthegeneraconsisted of
one species each(37%). Thus, 65% ofthegenerarecordedwere represented by
two orfewerspecies.
ESTIMATEDRICHNESS. — Therichness estimatedwas larger thantherichness
observed:76.28(SD±
0.02) against 57 spe-
cies. This may mean
thatweonly recorded
75%of the true local
richness. On theother
hand, thespecies accu-
mulation curve com-
puted had began to
reach the asymptote
(Fig.l),which proba-
bly indicates that the
estimatedvaluecould
Fig. 1.Observed(circle)andestimated (square)species accumulation
beatrue reflectionof curvesofOdonatafaunaofHuautla,Morelos.
308 E.Gonzalez-Soriano et.at
theestimatedrichnessforthislo-
cality.
ABUNDANCE
Thedistributionperspecies of
2595adultspecimens washetero-
geneous;afewspecies werevery
abundantbut most others were
represented by oneorafewindi-
viduals(Fig.2). Themostabun-
dantspecies was Argiatezpiwith
443 individuals, followed by A.
pulla with 362, A. oenea with
194, Hetaerinaamericana with
190, Telebasis salva with 151,
Dythemis nigrescens with 119
and Protoneura cara with 109.
All abundantspecies (excepting
the libellulidD. nigrescens) are
zygopterans.Incontrast, 11 spe-
cies wererepresented by onein-
dividualand 17 by less than 10
individuals.
Fig. 2. Rank-abundance pattern of the Odonata of
Sierra deHuautla.
DIVERSITY
Thediversity valuecalculatedwiththeShannonIndexovertheentireyearwas
3.04andtheevenness indexwas 0.75.Thediversityvaluesbymonthvaried, with
thelowest valuerecordedinMay(2.2)andthehighestrecordedin July(3.1).The
lowestand highest values of theevenness indexwererecorded in April (0.755)
and July (0.91) respectively (SeeTable 1)
Table I
Diversityandevennessindexes obtained monthlyoftheOdonata faunafromHuautla,Morelos
Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sep Oct
Diversity 2.99 2.8 2.6 2.8 2.4 2.4 2.2 2.3 3.1 2.3 3.01 2.8
Evenness 0.84 0.85 0.82 0.87 0.79 0.75 0.78 0.76 0.91 0.88 0.85 0.89
Odonatadiversityintropical dryforest 309
Fig. 3.Patternofspeciesrichness(circles)andabundance (squares) ofOdonataofSierradeHuaut
laper month.
PHENOLOGY
Theannualactivitypatternshowedthatthe richnessreachedthehighest value
inNovember(36), attheend oftherainy seasonandthelowest in May(17), at
theend of the dryseason, witha gradual and nonconstant decrementbetween
thehighest andlowest value(Fig. 3).
The abundance showed a greater variationthan the richness, recording the
highestvaluesinNovember,MarchandAugust andthelowestin Decemberand
October(Fig. 3).
Fig.4.Number ofactivespeciesper month ofOdonataofHuautla.
310 E.Gonzalez-Soriano et. al
Despite thetemporal variationrecordedinrichness andabundance, therewas
not significant differencesofthesevariablesbetween the rainy and the drysea-
son(Mann-Whitney U, p=0.6andp =0-5 respectively).
On theotherhand,sixteen(28%) species wererecorded only inthe rainy sea-
son, seven(12%)during the dryseason, and34(60%) withinbothperiods.
Thenumberofmonthsadultswereonthewingwas variable.Eighteen species
(31.6%) wereactiveduringonlyonemonthand28(49%)wereactiveduring three
months or less. Contrastingly, 13 (22.7%) species wereactive at leastduring 11
months(Fig.4).
LISTOFODONATESPECIESRECORDED INTHESIERRADEHUAUTLA,MÉXICO
Thelistincludesnumbersofindividualscollected(in bold),informationwhen
theadultswere collectedand insome casesinformationaboutnaturalhistory.
ABBREVIATIONSUSED: YR=speciesthatflythroughoutthe year; — AYR= specieswhich
apparentlyflythroughoutthe year.
Zygoptera
CALOPTERYGIDAE
Hetaerina americana(Fabricius,1798), 190.November toOctober.YR.Lotic,
LESTIDAE
Archilestes grandis(Rambur, 1842).64. November toFebruary andApriltoOctober.
PLATYSTICTIDAE
Palaemnema dominaCalvert, 1903.3. July.Collected atthesunset, onepairin tandem.
PROTONEURIDAE
Protoneura caraCalvert, 1903. 109. NovembertoAugust,October.AYR.
COENAGRIONIDAE
Apanisagrionlais(BrauerinSelys, 1876).1.February.
Argiaanceps Garrison, 1996. 44.November, JanuarytoApril,June toSeptember.
ArgiacarlcookiDaigle, 1995. 18.November,February-March.
Argiaextranea (Hagen,1861).73. November toApril,June toAugust.
Argiaharknessi Calvert, 1899. 59.ApriltoAugust.
ArgiaoculataHageninSelys, 1865. 7.November toFebruary, July.
ArgiaoeneaHageninSelys, 1865. 194.November toOctober.YR,
ArgiapallensCalvert, 1902.73. MaytoAugust,October.
ArgiapullaHageninSelys, 1865.362.November toOctober.YR.
ArgiatezpiCalvert, 1902.443.November toOctober.YR.
Enallagmacivile (Hagen, 1861).I.March.
EnallagmanovaehispaniaeCalvert,1907.68.November,JanuarytoOctober.AYR.
Enallagmasemicirculare Selys, 1876, 60.November, JanuarytoMay,July-August,October.
Ischnurahastata(Say, 1839).6.December,February-March.
Ischnuraramburi (Selys, 1850).1.August.
Telebasis salva(Hagen,1861). 151.November toJune, AugusttoOctober.AYR.
AESHNIDAE
AnaxwalsinghamiMacLachlan, 1883. 1.July.
Coryphaeschnaadnexa(Hagen. 1861).1.November.
GynacanthahelengaWilliamson &Williamson, 1930. 1.September.
Odonatadiversityin tropicaldryforest 311
GynacanthanervosaRambur,1842. 1.January.
Remartinialuteipennisflorida (Hagen,1861).1.November.
Rhionaeschna multicolor (Hagen,1861).1.August.
Rhionaeschna psilus(Calvert, 1947).6.NovembertoJanuary,October.
GOMPHIDAE
Aphyllaprotracta (HageninSelys, 1859).14.July-August.
Phyllogomphoidespacificus (Selys, 1873).79.November,JunetoAugust.
Progomphusclendoni Calvert, 1905.4.July.
LIBELLULIDAE
Brachymesiafurcata(Gundlach,1889).4.April.
BrechmorhogapraecoxpostlobataCalvert, 1898.56.December, MaytoAugust.
DythemismayaCalvert, 1906. 27.November-December,February,July August.
DythemisnigrescensCalvert. 1899. 119.November toOctober,YR.
Dythemissterilis Hagen.6.November.
Erythemisplebeja{Burmeister,1839). 11.November-December,February-March.
Erythrodiplaxbasifusca(Calvert, 1895).19.November-December,February,April,October.
ErythrodiplaxfunereaHagen.2.April.
ErythrodiplaxfuscaRambur.5.November, January,April.
Libellula croceipenisSelys, 1868. 22.November-December,April,JunetoAugust,October.
Libellula saturataUhler,1857. 1.October.
Macrothemis inacutaCalvert, 1898. 6.November-December,April,JunetoAugust,October.
MacrothemispseudimitansCalvert,1898.37.November-December.FebruarytoAugust,October.
Miathyriamarcella (Selysin Sagra,1857).32.November toAugust,October.AYR.
Micrathyria aequalis(Hagen,1861).6. November,April,
Micrathyriadidyma(SelysinSagra, 1857).7.November.
Micrathyriasp.nov.1.July.
Orthemis discolor(Burmeister,1839). 12.November toFebruary,August.
Orthemisferruginea(Fabricius, 1775).12.November-December,April,JulytoOctober.
PaltothemislineatipesKarsch, 1890. 15.JunetoAugust.
Pantalaflavescens(Fabricius, 1798).5.December-January,August.
Perithemis domitia(Drury, 1773).2.August,October.
Perithemis intensaKirby, 1889.94.November toOctober.YR.
Pseudoleon superbus(Hagen, 1861).49.November toAugust,October.AYR.
Sympetrumillotum(Hagen,1861).1.November.
Trameaabdominalis (Rambur,1842).1.December.
TrameaonustaHagen, 1861.6.November,January,August.
DISCUSSION
Thenumberofspecies recorded inHuautlais lowerthanthatrecordedin the
regionofChamela, Jaliscostate(57 vs78spprespectively) (GONZALEZ-SORI-
ANO etah, 2004). Chamelaissituated athigher latitudethanHuautla(19°30’ vs
18°20’respectively) butataloweraltitude(<150vs900-940mrespectively) (BUL-
LOCK, 1988)and is also covered by TDF.Thedifferencesinrichness between
HuautlaandChamelaapparently are determinedby the sampling of a greater
diversityofhabitats inChamelathanin Huautla.In Huautlamostof thesam-
pled habitatswere lotic whereas inChamelaboth loticand lentichabitatswere
sampled.
312 E.Gonzalez-Soriano et. al
GONZALEZ-SORI ANOetal. (2004) commentedthatthespecies richnessof
Chamelais magnified by the presenceof species belonging to Libellulidaeand
someCoenagrionidae both inhabitantsofstanding waters.Species foundinsuch
habitatsare usually not foundflying oversmall streams or in smallopenings in
primary forest. LOUTON etal.(1996) consideredLibellulidae, Coenagrionidae
(minus Argia)and also Aeshnidaeas “weed”species by theirwide distribution
andgreatcapacity to colonizenewsites fortheirdevelopment. On thecontrary,
species not includedin thosegroups likely have narrow distributionsand they
characterizebetterthefaunaof oneregion. Ifweeliminatedthe weedspecies in
our comparison, theresulting numberof species betweenboth regions isalmost
identical: 17spp.in Chamelavs 16spp in Huautla.On the otherhand, acom-
parison betweenLibellulidaeandCoenagrionidae inbothareas show thatthere
were 42spp. of Libellulidaeand 14 of Coenagrionidae in Chamelaversus 27
and 16, respectively forHuautla.Inthisway, the bigger differencewas observed
in Libellulidaeandthis familyisresponsible for71%ofthe richnessdifferences
recorded between bothregions (see Fig 5). Species collectedin Huautla which
have not been collectedin Chamelaare: Palaemnemadomina, Argia carlcooki,
A. extranea, A. harknessi, A. pallens, Aphylla protracta and Phyllogomphoides
pacificus.
Theabsenceofaseasonalpatterninbothrichnessandabundancecontrastswith
thestrongly seasonalnature oftheareaandwiththoseobservedforothergroups
Fig. 5.Speciesandfamilies ofOdonataatChamela (greybars) and Huautla (solidbars).
Odonatadiversity in tropicaldryforest 313
of insects within thesame study area (NOGUERA etal., 2002; ZARAGOZA
etal.,2003). Although this patternhasbeenreported inothergroupsofinsects
(LEVINGS & WINDSOR, 1985;PESCADOR et al., 2002), the absenceof a
seasonal patterninbothvariablescould havebeeninfluencedin oneofthecol-
lecting sitesby theLorenzoVazquez damlocatedintheQuilamula river. During
the dry season the odonateswere collectedafterthe dammingofthe reservoir,
wherethewatersupplement wasconstant allyearlong, thuscreatingcontinuous
conditionsformany species along theentireyear.Thiseffectwas alsoobserved
intwo speciesofLampyridae, that during thedry seasonwere collectedonlyaf-
terthe dammingofthereservoirbutnotup-stream(ZARAGOZA etal.,2003) a
factthatseems tosupportour previous statement. Additionally, inChamela (an
areawithtemporalwaterbodies) theproportionofspecies recordedbetweensea-
sonsis inverse, with65%ofthespecies recorded onlyintherainy season(versus
28%inHuautla)and 32%recordedin bothseasons (versus 60%inHuautla).
CONCLUSIONS
Thespecies richness ofOdonataofthe SierradeHuautla Biosphere Reserve
(SHBR)is outstanding. TheSHBR withanextensionof59031hectaresconsti-
tutesalmost 12%oftheMorelosstatesurface. Inthissmallareaitisrepresented
almostthe56%ofthetotalMorelosodonatediversity(GONZALEZ-SORIANO
& NOVELO-GUTIERREZ,1996,2007). Although othersiteswith tropical dry
forestinMexicohaveahigher speciesrichness thantheSHBR,(forexample Cha-
melaregion), acloseanalysis showedthattheincreasedrichness ofthelastsiteis
dueto thehigher numberofspecies belonging to groupswithwidedistribution
as Libellulidae, AeshnidaeandsomeCoenagrionidae.
Finally, the following 7 species are recorded for the first timein the state of
Morelos: Argia anceps, A. carlcooki, Anax walsinghami, Gynacantha nervosa,
Dythemis sterilis, Macrothemispseudimitans and Trameaabdominalis.Argiaan-
cepswas formerly recorded as A. fissa(GARRISON, 1996; GONZALEZ-SO-
RIANO & NOVELO-GUTIERREZ,1996).
ACKNOWLEDGEMENTS
Wewishto thankDr ROSSERGARRISON (Sacramento,CA, USA)for criticallyreadingthe
manuscriptand byhis helpfulcommentaries. ToDrOSCAR DORADO (CEAMISH)forlogistic
support duringfield work in Huautla
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