A synopsis of the Argia fissa group, with descriptions of two new species, A. anceps sp.n. and A. westfalli sp.n. (Zygoptera: Coenagrionidae)

17 Pages·1996·1.9 MB·English

by  Garrison, R W

Odonatologica25(1): 31-47 March I.1996
A synopsis oftheArgia fissa group,
withdescriptions of twonewspecies,
A.ancepssp.n.andA. westfalli sp.n.
(Zygoptera: Coenagrionidae)
R.W.Garrison
Los Angeles CountyNatural History Museum,
900ExpositionBoulevard,Los Angeles,CA 90007, UnitedStates
Received June 12,1995/ReviewedandAcceptedAugust2, 1995
The Argiafissagroup, ofthe northern neotropicalregion,isanalyzed. A. anceps
sp.n.(holotypeS,allotype 9;CostaRica, SanJoseprov., San Pedro, 9-VIII-I979;in
USNM)andA.westfallisp.n.(holotype3,allotype5:Mexico,Nuevo Leon,Municipio
deSantiago, 2.5 km NW ofRancho Los Atascosos, 2-VII-I987;in ESCA) arede-
scribed,keyed, illustrated,and comparedwiththe northern SouthAmerican A.fissa.
The types aredepositedinFSCA,Gainesville,FL,USA.
INTRODUCTION
Argiafissawas originallydescribedby SELYS(1865) fromanunspecified number
ofmalesandfemalesfrom“Bogota, NouvelleGrenada[= Colombia]”. CALVERT
(1902) featuredthespecies in theBiologiacentraliamericana, extending the spe-
ciesnorthwardto25°38’NinNuevoLeon(nr. ElDiente, ca6 km SW Monterey),
Mexico.RIS(1918)extended itsrange southwardto 12°05’SinPeru(LimaProv.,
Callao).Malesofthisspecies are unique inpossessing slightlyamberwings, largely
pale blueabdominalcolorationlacking postbasal streaks, and long cerci with a
single decumbenttooth.LeonoraK. Gloyd, whowas working on thegenusup to
herdeathin June, 1993,realizedmanyyearsagothat threespecies were included
underthenameA.fissa. Oneofthe undescribedspecies is perhaps themost com-
monly encounteredzygopteran in Mexicoand south through Costa Rica. Gloyd
examinedthousandsof specimens andapplied a manuscript nametothis species.
Thisandtheotherundescribedspecies are thenortherncomponentsof theA.fissa
' Mailingaddress: 1030Fondale Street,Azusa,CA91702-0821,United States
32 R.W.Garrison
complex along thenorthernmostborderstatesofMexico. I describe themso that
theirnames willbeavailable foranupcoming manualon thedamselfliesofNorth
America by M. May andM.J.Westfall, Jr.
Terminologyfollows that ofGARRISON(1994). The illustrations ofthe structures in thispaper
weremadeusingaWildM-8microscopeequippedwithacameralucida.Underspeciesaccounts, app
=appendages,mp=mesostigmallobe(s),thx =thorax, synthx=synthorax, FW =fore wing,HW=
hindwing,and abd =abdomen. Venational countsare from 20 specimens ofeach sex, where avail-
able,from as many differentlocalitiesas possible. Specimenslisted as RWG arein my collection;
other acronyms canbe found intheacknowledgements.
Acriticalexaminationofthecerci (Fig.2c) inmalesand mesostigmal lobes(Fig.
1la’, c’)in femalesis necessary toassure accurate identificationofthethree spe-
cies. They may bedifferentiatedby thefollowing key.
KEY TO SPECIMENS OFTHEARGIA FISSA GROUP
MALES
I Cercusin dorsal viewwithmedialmarginconvexlycurved, followed by asmall concavemar-
ginneartip(Fig.3c),samestructureinmediodorsal view(Fig.3a)withmedialmarginnonlinear,
thisareacurvingventrallytomeetdecumbent tooth,then curvingdorsallyaftertooth;cercus
in mediodorsal view with decumbent tooth located at distal 0.70 sothat medial margin of
cercus appears slightlyangulate atregion oftooth; humeral stripeunbranched (Fig. 24) in
morenortherly Mexican states,orforked (Fig.25)inpopulationsfrom Chiapas.... westfalli
1' Cercus indorsal viewwith medial marginlinearorslightlyconcave(Figs 1c,2c), samestruc-
tureinmediodorsalview (Figs la, 2a)withmedialmarginlinear(exceptat regionoftooth);
cercusinmediodorsal viewwithdecumbenttoothlocated atca.0.30-0.60ofappendage,cercus
notappearingangulateat regionoftooth (Figs la, 2a);humeral stripeofvaryingcondition
(Figs 17, 18,20,21),but neverforked; northernborder statesofMexicosouththroughColom-
bia, Ecuador, andPeru 2
2(1’) Medial marginsofcerci in dorsal view contiguousornearlyso,transverse torusstronglyover-
lappingepiproct,torialmostcontiguous,eachtorus about lOx widerthangapseparatingthem
(Fig. 2c); decumbenttooth on medial marginofcercus inposteriorviewat basal 0.30-0.40
(Figs2d,5a-c);extreme easternCosta Rica south throughcordilleran regionofColombia and
northern Peru fissa
2’ Medialmarginsofcerciin dorsalviewseparatedby agapaboutaswideasbasalwidth ofeach
cercus,torionlyslightlyoverlappingepiproct,each torus about 3-4xwider thangap separating
them(Fig. 1c);decumbent tooth on medial marginofcercusinposterior viewatbasal 0.50-
0.60(Figs Id,4a-d);northernborderstates ofMexico south toeastern Costa Rica... anceps
FEMALES
1 Mesostigmallobes stronglyelevated,withlobe distinctlyerect (Figs 11b’,c’), anterior margin
ofmesepistemumdirectlyunderneath mesostigmallobewithnotubercle, thisareaforminga
concavejunctionwithbase ofmesostigmallobe (Fig. 16)[Note: Stronglightis necessary to
seethisstructure], humeralstripeunbranchedtoforked (Fig.26) westfalli
1’ Mesostigmallobes notasstronglyelevated(Figs 9b’, c’, 10b, c)comparedwith A. westfalli;
anterior marginofmesepistemumdirectlyunderneath mesepistemal lobewith a transverse
Argiafissagroup 33
tubercle ofvaryinglength(Figs 14-15),this structure sometimes visiblein lateral view(Fig.
9c’); humeral stripeneverforked (Figs22, 23) 2
2(1’) Distance between mesostigmallobes almostaswide(70-93%,N=8) aseach lobe; usuallywith
asmallgapbetweenmedial edgeofmesostigmallobe and anterior fork ofmiddorsal thoracic
carina(Figs 9a, a’, 12a-f) anceps
2’ Distance between mesostigmallobesapproximatelyhalf(50-63%,N=8) aswideaseach lobe;
medial marginofmesostigmallobe adjacentto anteriorfork ofmiddorsal thoracic carina(Figs
10a,13a-f) fissa
ARGIA ANCEPS SP. NOV.
Figures 1,4-app,7-penis,9, 12-mp, 17-bodyofholotype, 18-thx,abd segments 7-10 ofparatype,
19-abdsegments 7-10ofparatype,20-21-humeral stripeofmales,22-bodyofallotype
Argiafissa CALVERT 1899: 381 (descr. i from Tepic,Mex.); CALVERT, 1902:
89(allspecimensfromMiddleAmericamisidentifiedasA.fissa)
Material. - Holotype3, allotype 2:COSTARICA,San Joseprov.,San Pedro,dumpbeside
Holland House(Hotel),9-VIII-1979,R.W.& J.A.Garrison leg.Bothin USNM. - Paratypes(1233,
34 2):MEXICO;Sinaloa: stream 15.7miNEofConcordia, 1900ft,26-VIII-1965,D.R.Paulsonleg.,
1 3 (FSCA);- Hidalgo,Calnali,9-IV-1994,R. Novelo,leg. 1 3, 1 2 (IE);- Molango,Lagunade
Alezca, 1450m,20-XI-1983,J. Pena, leg, 3 3(IE);- LagunaAzteca[sic, should beAtezca], 2kmN
Molango,24-VI1-1992,G. Harp, leg., 1 3 (RWG); - Naopa, 1000 m, 25-IV-1984, R. Novelo, V.
Garcia& J.Pena, leg. I 3 (IE);- Tecozautla,Rio Tecozautla, I4-IX-1992,R.Noveloleg.,2 3 (IE);
- Guerrero, MX-55betweenTaxcoJetand Huajintlan,22-VII-1993,W.F.Mauffray,leg., 1 3 (IORI);
- Jalisco,Sierrad*Autlan, 16-1X-1991,nocollector,1 3(RWG);- Morelos,Rt 115EofCuernavaca
nearTijalpa,23-VI-1963,R.E.Woodruff,leg.,1 3 (FSCA);- Las Fuentes, MunicipioofJiutepec, 10
km WofCuernavaca, 1300 m, 17-1X1983,R.W.Garrison, leg.,3 3 (RWG); - samedatabut 1-X-
-1983,6 3.2 2(RWG); - Itzamatitlan, 5km SWofYautepec,2-X-1983, R.W.Garrison leg.,23.1
2 (RWG);- Canon deLobos, 5km SW ofYautepec, l-X-1983,R.W.Garrison, leg.,1 3 (RWG);-
CanondeLobos, 19km EofCuernavaca,15-X-1986,E.Fisher, leg.,13(RWG);- Xochitepec,MX-
-95libreatBalnearioCampestreRioSabinos,21-VII-1992,W.F.Mauffray,leg.,173 (1 <Jincopula
with 2 ofAj. extranea[Hag.]),9 9(IORI,FSCA);- Cocoyotla,approx. 5kmN ofMXrt421,23-VII-
1992, W.F.Mauffray,leg.,2 <S(IORI,FSCA);- BalnearioonriverS ofTemixco onLibre95,21-VII-
1992, S.W. Dunkle, leg., 1 c3, 1 9 (SWD); - Ixcatepec, CampamentoCammohmila, 16-X-
-1993, R. Novelo,leg., 2<3 (IE);- Rio Sabinos,km 18.3 carr.95, 5km S Acatlipa, 14-X-1994, R.
Novelo, leg.,2 i (IE); - Michoacan, stream 3.9mi NofTuxpan, ca6000ft, 18-VIII-I965, D.R.
Paulson, leg,, 1 (3,(FSCA);-Veracruz, streamathighway,“Fincalas Animas”,ImiEofJalapa,4680
ft, 11-VIII-1957,G.H.BeattyIII,leg.,1 <3, I 9 (RWG);- stream6.9kmSofCordoba by Mex Hwy
150,700 m, 1-VIII-1976,R.W. &J.A.Garrison, leg., 1 S (RWG);- samedata but 12-VIII-1976,17
<3 (RWG);- samedata but 13-VIII-I976,1 o’ (RWG);-km 17carr.Cordoba-Veracruz,22-IX-1983,
R. Novelo, leg., 1 <J, 1 9 (IE);- Fortin delas Flores, 27-VI-1963,R.E. Woodruff, leg., 1 <3, 1 9
(FSCA);- stream4.2miNFortin,4100 ft,Il-VIII-1965, D.R.& ML. Paulson, leg.,2 i (FSCA);-
RioMetlac about3.5kmWNWFortindelasFlores, 900m, 10-V1II-1976,R.W.&J.A.Garrison,leg.,
4 d,2 9 (RWG); - 5 miNofHuatusco, 4000-4500 ft, 13-VIII-1963,H.V.Weems, Jr,leg., 1 9
(FSCA);- pond28km NEHuatusco by Mex Hwy 66,900m, 11-VIII-1976,R.W.& J.A.Garrison,
leg.,23,3 9 (RWG); - ParqueJavier Clavijero,Jalapa, 1300 m. 17-VIII-1982,R.W. & J.A. Garri-
son,leg.,2<3,I 9(RWG);- samelocality,8-V-1991,R. Novelo,leg,, 1 i (IE);- LaFratemidad, 17
kmEofCordoba,22-IX-1983,R.W.Garrison,leg.,1 <3 (RWG);- Puebla, Nuevo Necaxa(Presa), 18-
-VI1-1990, R. Novelo, leg., I c5 (IE);- Puente Necaxa, Hwy 130, 5 mi NEHuauchinango,B.C.
34 R.W. Garrison
Figs 1-3.Argiaancepssp.n. (holotype:Figs la-c),A.fissa (Figs2a-c),andA. westfallispn.(holotype:
Figs 3a-c):(a) caudal appendages,mediodorsa) view;- (b)same,lateral view;- (c)same,dorsalview.
Argiafissa group 35
Figs 1-3(continued),4-8. A. ancepssp.n. (Figs Id,4,7), A.fissa (Figs2d, 5), and A.westfalli sp.n.
(Figs3d,6,8): (ld-3d)caudal appendages,posteriorview;- (4-6)right cercus, medioposteriorview;
- (7-8)penis,ventral(ectal)view, left,lateral view,right.
36 R.W.Garrison
Villegas,leg., 1 9(RWG);- Piedras Negrasarroyo,25-X-I987,A.Gomez, leg., 3S (IE);- Xicotepec
deJuarez, 20-V-I987,R. Novelo&A.Gomez,leg., 1 S, 1 9(IE);- Oaxaca, 43miWofTehuantepec,
2500-3000ft, 22-VIII-1963, H.V.Weems,Jr.,leg., 1 S (FSCA);- Chiapas,FincaSanJoseLaVicto-
ria, km 20carreteraTapachula-Cacahoatan,9-III-1982, R. Novelo,leg., 1 6 (IE);- FincaLaFlor, 3
km NEdeOcuilapan, Mun. deBerriozabal, I3TI1-1982,R. Novelo,leg., 1 S (IE);- La Boquilla,25
km Nde Tapachula,21-XI-1982, R. Novelo,leg., 1 <? (IE);- 13miSWofCintalapa, 1500-2000 ft,
19-VIII-I963,H.V.Weems,Jr.,leg.,1 S (FSCA);-Elchorreadero atkm25onTuxtla-Gutierrez-San
Cristobal Rd,24km S ofTuxtlaGutierrez,R.W.Garrison, leg.,13,19(RWG); - ElChorreadero,
carreteraTuxtla-Gutierrez-San Cristobal de las Casas,27-XI-1983,R. Novelo,leg., I <3 (IE);- BE-
LIZE,PrivassionCreek, 10-XII-1992,T. Boomsma, leg.,1 6(RWG); - GUATEMALA,DeptGuate-
mala,caldera, lakenrSanFrancisco deSales, 1800m, 22-VIII-1964, T.W.Donnelly,leg.;46 (FSCA,
RWG); - HONDURAS, Danli, 10-V-1966,J. Malta, leg., 13,19(FSCA);- Francisco Morazan
Dept,ElRancho,between ZamoranoandTegucigalpa,ll-XII-1987,S.W. Dunkle, leg.,2 3 (RWG);
- ElHatillo, 10km NofTegucigalpa, 1300 m,T.W.&A. Donnelly,leg.,28-VI-1974, 1 3 (FSCA);-
COSTARICA,CartagoProv,RioReventazon,SEofTurrialbaby Hwy 10,10-VIII-1979,R.W.&J.A.
Garrison,63,29 (RWG);- vicinityJuanVinas,RioReventazon,800m, 11-VI-1962,T.W.Donnelly,
leg., 1 3 (RWG); - Guanacaste Prov, smallstreamonRt 145justEofTilaron,27-11-1987, S.W.
Dunkle,leg., 1 J (SWD);-Puntarenas Prov, MacaconadeEsparta,I/II-1905,P.Biolley,Jr,leg., 1 3
(FSCA); - San Jose Prov,no locality, 7-V-1960, R.B.Gumming, leg.,2 3 (FSCA);- samedata as
holotype, 48,29(RWG); - Res. Biol. ElRodeo,7kmWofVillaColon, 9°54’N,84°16’W,800m,
10/13-VII-1990,T.W.Donelly.leg,23,I 9(FSCA,RWG);- SanPedrodeMontesdeOca,Quebrada
los Negritos, 1200m, 5-VIII-1983,E.A.Rojas,M., leg., I d,I 9 (RWG).
Etymology.- Latinfor double, in reference toits siblingrelative A.fissa.
MALE(holotype). -Head(Fig. 17).Labiumpaleblue,remainderofheadblue
with smallblackmedialspotatbaseof labrum;black onepicranium reducedwith
narrow longitudinal lineseparatinglarge postocular spot fromcross barofblueon
occiput; pale spotanterolateraltolateralocellus confluentwithremainderofblue;
anteriormargin ofblack withanterolateralmargin not reaching baseofantennae;
antennaeblack rearofheadchiefly pale, with smallareaaroundoccipital foramen
black.
Thorax.-Prothoraxblue, black above withfollowing areas pale: transverse
bandon frontallobe,finely dividednarrow dorsalspotandlarge dorsolateralspot
on middlelobe,narrow medialspotandlateralof0.30ofposterior lobe.Paleareas
ofsynthorax (Fig. 17)blue,withbroadblackmiddorsalstripe0.30aswideas pale
antehumeral stripe; black humeralstripe not forked, extending from base of
mesinfraepisternum andforming abroadlobe atbaseofmesepimeron, thenupper
halfofthisconstricting to0.25 widthofpale humeralstripe; black humeralstripe
abruptly narrowedto0.10widthofpale humeralstripeandwithslight enlargement
at dorsal0.10ofthorax,thirdlateralsuturewithnarrow black stripe.Wingsslightly
amber with venationblack; pterostigma dark brown,surmounting 1.5 cells in all
wings (one cell inlefthindwing); postnodals: 16/13;postquadrangular cells:FW
5/5; HW5/4, M, originating atFW 7/7; HW6/7. Coxae andtrochanterspale ex-
cept for black on dorsum; femorablack dorsally, pale ventrally; tibiaeblack on
innersurface, black onexteriorsurfaceofpro- andmesothoracictibiae,remainder
pale; tarsiand armature black.
Argiafissa group 37
Figs9-11.A.ancepssp.n. (Fig.9),A.fissa (Fig. 10),andA.westfallisp.n.(Fig. 11):(a,a’)mesostigmal
plates, dorsal view (note: dark areas behind each lobe areshadows for each lobe); - (b.b’) same,
obliquedorsolateral view;- (c,c’) same,lateral view.
38 R.W. Garrison
Abdomen(Fig. 17) blue, segment 1 pale with dorsalbrown spot atbasal
0.50;segment2 withblackdorsolateralstripe extending to0.80ofsegment,itsend
forming aninverted triangle extending to but not connecting above, asmall iso-
lated spot below, annulus black; segment3 with an incomplete bandofblack at
posterior 0.80of segment,isolatedspotofblueatlateral0.90ofsegment,annulus
black; segments 3-5 with apical 0.20ofeach segment, including annulus,black;
thisblack not meetingdorsallyon segment4,separated by anarrow middorsailine
onsegment5,andconnecting dorsally onsegment6;segment7similarto segment
6 butwith lateralprolongation ofblack extending to basal 0.15ofsegment; seg-
ments8-10pale with atouchofblack atlowerposterior margin ofsegment8.
Torifer (Fig. la, c) triangular, torus prominent, transversely twice as wide as
long, occupying lower0.50 oftorifer, each torus transversely about 1.5times as
wideas the distancebetweenthem; epiproct not prominent, extending to levelof
torus, largely blackexcept for pale apical margin, bilobed witha recessed small
medianlobe. Cercus (Fig. la) pale, abouttwice as long as wide, distally slightly
falcate, indorsal view with medialmargin slightly concave, cercus with asingle
smallblackdecumbenttoothvisibleatmiddlealong ventralmargin indorsomedial
view;medialmargin nearly linearindorsomedialview;inlateralview, cercus nar-
rowly triangular, withinnertoothnot visible;paraproct alittleshorterthancercus,
barely divided,withdorsallobeforming abroadly basedtriangle; ventralbrancha
small bluntly pointed tubercle(Fig, lb).
Penis(paratype.Fig. 7) withfourthsegmentforming anarrow, bluntly rounded
lobe.
Measurements (inmm).- HW 24.5,abdomen 31.0.
FEMALE (allotype). - Tan with headand thoracic patterns similar to malebut
withblack areas onepicraniumlessextensive(Fig. 22)andwithdiffuseblack spot
extending frombelowbaseofantennatoeyemargin,blackmiddorsalstripeamere
hairlinealongmiddorsalthoraciccarina, black humeralstripe similarto malebut
narrower. Wingsslightly amberwithvenationblack, pterostigmabrown,surmount-
ing onecell orslightly less inall wings, postnodal crossveins:FW 15/15,HW 13/
12; postquadrangular cells:FW 5/5, HW 4/4;M originating nearFW 8/8, atHW
2
6/6,
Abdomen(Fig. 22) tanwithdiffusebrown markingson dorsumandwithdarker
markings atdistal0.80of segments3-7, thesedarkmarkingscomposed ofalarger
dorsolateralspot andanarrow lateralstreak; annuli darkbrown on segments 2-6,
paler on 7; segment8 pale withawashofbrown at distalmargin, segments9-10
and cercipale.
Mesostigmal lobe(Fig. 9a) well developed, moderately erect, and forming a
broadsemicircle;itsdistalmargin welldifferentiatedfromremainderofmesostigmal
plate, itsmedialmargin separatedfrom lateralarmofmiddorsalthoraciccarinaby
asmallgap; medialhalfoflobe black dorsally, entirely black ventrally; a distinct
transverse tubercle atjunction of mesostigmal lobe andanteriormarginofmese-
Argiafissa group 39
Figs 12-13. A.anceps sp.n. (Fig. 12)andA.fissa (Fig. 13):(a-f) mesostigmallobes, outline, dorsal
view.
40 R.W. Garrison
pisternum (Fig. 14, paratype); no mesepisternal tubercles.
Measurements (inmm).- HW26.0, abdomen 31.0.
REMARKS.- Much inter-and intrapopulational variability in body markings is
expressed in this widely distributedspecies. Overall black markings maybe re-
duced(Fig. 18),andsomespecimens ofbothsexes lack anyblack markings onthe
epicranium (Figs 18, 23). A corresponding reductioninthe widthofthemiddorsal
andhumeralstripesexists withinpopulations. Somemaleshavethemiddorsaltho-
racic stripe as wideas thepale antehumeralarea on themesepistemum(similar to
thatillustratedfor the holotype, Fig. 17), whileothers (Fig. 18) have the stripe
reduced almost to a hairline, as for females(Figs 22, 23), although the stripe is
usually widerin malesthanin femalesin thesamepopulaton. Theblack humeral
stripe can be greatly reduced (Figs 20a, b) or may form an almost continuous,
nearlyparallel stripe whichisslightly narroweddorsally(Fig. 21).Thelateralblack
streakon abdominalsegment7 may beisolated(Fig. 19)or reducedto thedistal
0.20 (Fig. 18). Ihavenoticedno correlationofthoracicmarkings withgeography:
intraspecific variability seems as greatas interpopulation variability among the
specimens Iexamined.
Paratype measurements: M a 1 e s (N = 20): HW 22-26.5mm, abdomen29-33
mm. Postnodalcrossveins (FW/HW) 14-18/11-15;origin ofM 6-8.5/6-8;postquad-
2
rangular cells4-6Z3-5.- Females(N=20): HW22.5-27mm,abdomen27.5-32
mm;postnodal crossveins 14-18/12-15; origin ofM 6-8Z6-7.5; postquadrangular
2
cells4-5Z3-4.
CALVERT(1899) first misidentifiedthis species as A.fissa afterhe compared
two malesfromTepic, Mexico,with“specimens fromGuatemalainBarondeSelys’
collection...”.Becausenobonafidespecimens ofA.fissaareknownnorthof south-
eastern CostaRica, Calvertalmostcertainly hadcompared hismaterialwithGua-
temalanspecimens ofA. anceps. I doubtwhetherCalverthadever seenspecimens
ofthetrue A.fissa. Allprevious records ofA.fissafromMiddleAmericaprobably
refertoA.anceps.
BIOLOGY. - A.anceps is anabundantstream-dwelling species. Itis likely to be
encountered atany smallstreamor trailbordering largerstreamsalmostanywhere
in MiddleAmerica. Itapparently flies year roundas collectiondatesrange from
January-February to 11 December.
distribution. - According to Gloyd’s unpublished distributionrecords and
GONZALEZ& NOVELO (1996), thisspecies has beenrecorded from every state
inMexicoexcept thefarnorthwesterly states(BajaCalifornia, Sonora,Chihuahua,
Coahuila, Durango), or poorly collected states (Zacatecas, Aguas-Calientes,
Guanajuato, Queretaro, Tlaxcala, DistritoFederal, and Mexico). I have seen no
records form the Yucatan peninsula (Tabasco, Campeche, Yucatan, and Quintana
Roo), though itwillprobably befoundinwesternTabasco.Its distributionextends
to approximately 9°5’ N inSan Jose, Cartago andthe northernpartof Puntarenas
provinces, Costa Rica. Itis apparently parapatric to A.fissa, the most northerly