Odonatologica30(1): 67-86 March 1, 2001 An annotatedlist of Odonata collected in Ghana in 1997, a checklist of Ghana Odonata, and comments on West African odonatebiodiversity and biogeography G.O’Neill¹ and D.R. Paulson² 1 DepartmentofBiology, University ofPuget Sound, Tacoma, WA 98416,United States (present address: 14 LehighAve., Wilmington,DE 19805,United States) 2SlaterMuseum ofNatural History, University ofPuget Sound, Tacoma, WA98416, United States; — e-mail: [email protected] Received August 8, 2000/ RevisedandAccepted September 4, 2000 Collections were madeat8 localities in southern Ghana duringthe summerof 1997. Three regions were sampled: coastal savanna, wooded savanna, and rainforest. 71 spp. werecollected, 24 ofwhich are new for the country,bringing the Ghana list to 123 spp. A list of spp. known from the country is included. Trithemis dejouxiPinhey, 1978,is raised tospecificrank. Individual variation in Phaon iridipennis and Palpopleura lucia is quantified. West African Odonata biodiversity and biogeography are discussed. INTRODUCTION Todate, only fourstudieshavefocusedspecifically onthedragonflies ofGhana (KARSCH,1893;NEVILLE,1960;MARSHALL&GAMBLES,1977;D’ANDREA & CARFI, 1994). Littlehas been published about the biology ofthe species occurring there,andnolistof speciesknownfromthecountry hasbeencompiled. Fromthesepapersandothers, especially PINHEY(1962a),99species ofOdonata have beenrecorded inGhanato date. The landscape ofGhanavaries fromwet forest to drysavanna due to a sharp rainfall gradient. The southernportion ofthecountry is covered by wet, semi- moistandsemi-dry forests, whilefarthernorth,incentralGhana,forestgives way to thetallgrasses, short shrubs, andscatteredtrees ofthesavanna (SAYERetal., 1992).Terrestrialorganismsexhibitadiversitygradient along thishabitatgradient, butto datenoonehasdoneasystematic survey ofaquatic organismsalongit. The 68 G. O’Neill& D.R. Paulson research presented herebeganas anattempttodeterminewhetherasimilardiversity gradient occurred in Odonata. Theoriginal intentwas to testthehypothesis that two changes in biodiversity willoccur along the gradient as itchanges from wet forest to savanna woodland,thefirst a declineinspecies richness ofOdonataand the seconda shiftin species composition. Due to fieldconditions,only two sites along thegradientcouldbesampled verywell,buttwo previous studies(NEVILLE, 1960;MARSHALL&GAMBLES, 1977) fallatdifferentplaces along thegradient andallow furthercomparisons. Thegradient fromwet todry inGhanaoccurs fromsouthtonorth,withincreased distancefromthecoast,but italso occurs fromwest toeast. Ghanais oneofthe easternmost countriesin theWestAfrican forest belt, and, moving fromwest to east,theclimatebecomesdrierand drieruntiltheDahomey GapisreachedinTogo andBenin. TheDahomey Gap, where the dry savannaextendsall theway tothe coast andthereis no wet forest, represents awell-knownclimaticbarrierinWest Africa(SAYERetal., 1992).This gaphasbiogeographic significance, becausefor manyterrestrial wet-forest species it represents abarrier ofinhospitable habitat. Thiscausessomespecies tooccurononlyonesideofthegap.Thepresentcollections also contributeto an attemptto understandthe significance ofthat barrier for Odonata. METHODS Field work was conductedby thefirst authoratthreesites in Ghana from June toAugust 1997. The first site was near Pantan, asmall villageinthe southern coastal savanna, approximately 10 miles north ofAccra. Specimens werecollected 2-6 June at a small stream and several small pondsin thearea(see below for habitatdescriptions). The second site was Bui National Park, a Guinea savanna/semi-dry forest mosaicinthewestern part ofthe central region.Specimens were collected 19June-24 July from the Djapolx,a medium-sized stream, and theBlack VoltaRiver, a large river. The third site was Kakum National Park, a dense rainforest site in the southwestern section ofGhana. Specimenswere collected 26-27 August at two small streams in denseforest. Identifications were made using published literature, the unpublished keys ofVICK (1997), and comparisons with material in the second author’s collection. All specimens have been depositedin that collection. Notes oneye color in life, ecology, and behavior are includedfor some species. COLLECTION LOCALITIES GREATER ACCRA REGION, PANTAN, 5°42’ N, 0°12’ W — (Oya) Oyarefacoastal savannapond,0.5-1 m deep30 m wide pond,1-2 m mud surrounding, then 1 m tallgrass, sparse trees and shrubs. — (Dak) Dakobi stream, 1-2 m wide coastal savannastream nearroad,marshy in places, 1 m tall grass bordering, sparse trees and shrubs. — (Pan)Pantan coastal savannapond, 30 m wide pond, surrounded by 1-2 m tall grass, 1 m deep water,then sparse trees and shrubs. (a)Dawa, an open coastal savannaarea, 1-2m tall grass, nearroad. Odonata of Ghana 69 CENTRAL REGION, SHAI HILLS GAMEPRODUCTION RESERVE, 5°52’ N,0°4’ E — (Sha) open coastal savannaarea with 0.5-1 m tall grass, more trees on nearby steeprocky hills,short grass nearsmall stream. BRONG AHAFO REGION, BUI NATIONAL PARK, 8°20’ N,2°19’ W — (Dja) Djapoli stream, 3-4 m wide stream with rocky/sandy bed, alternatingpools and trickles, 0.05-0.30 m deepwater, bordered by 1-2m tallgrass and largetrees. (a)Bui Camp,small farmlands surroundingvillage,0.5-1 m tallgrass with exposedareasof soil and rock, no waternearby. (Vol) Black Volta River, 50-70 m wide river, water 13-15 m deep,bordered by very dense shrubs aryl trees. CENTRAL REGION, KAKUM NATIONAL PARK, 5°22’ N, 1°22’ W — (Kakl) small rainforest stream,meandering0.5-1 m wide trickle with sandy soil/small rock bed through dense rainforest, sun shining throughonly in limited areas ofthinned canopy. — (Kak2) rainforest stream, slow flowing 1 m wide stream, sandy soil/pebblerock bed with areas of solid rock causing waterfalls/small pools, through dense rainforest, sun shining throughonly inlimited areas ofthinned canopy. ANNOTATEDLISTOFSPECIESCOLLECTED Afterthespecies namesare thelocalitiesandthenumberofeach sex collected. Thecodeforlocality referstothelist ofcollectinglocalitiesabove.Theasterisked species arenewforGhana.Thecollectionsare summarizedinTableI, whichisalso acomplete listoftheOdonataknownto occur inGhana. CHLOROCYPHIDAE (1) Chlorocypha curta(Hagen, 1853). Dja(8(3,89). — Thisspecies apparently favorsmoreopenstreamsthanthefollowing two species, whichwerefoundtogether atKakum. (2) Chlorocypha glauca (Selys, 1879). Kak2(2<3). (3) Chlorocypha selysi (Karsch, 1899). Kakl (6<3), Kak2 (1<3). CALOPTERYGIDAE (4) Phaon iridipennis (Burmeister, 1839). Oya (2d), Dak(3c?), Dja(26<3, 10 9),Vol(2c3),Kak1 (2c?),Kak2(1c3). — Thisspecies, widespread acrossAfrica, was collected atnearlyevery site visited. Commonperching places includedrocks in the streambed, vineson thewalls ofthestreambed,andin 1-2m grassbordering thestream.Oviposition tookplace inthestemsofa20 cm tallplant next toaslow- flowing 1.5 mwidestream atDjapoli. Many individualswereobserved feeding by making short(5-20 cm) flights, capturing smallflying insects, andlanding onthe original perch for consumption ofthe prey. The intensity ofthe greenthoracic markings variedgreatly inbothmales andfemales,not apparently correlatedwith age,and some specimens hada lightblue pruinosity ontheventral surfaceofthe thorax. PINHEY(1961a, 1961b)discussedbriefly thevariationinthe relativesize 70 G.O'Neill & D.R. Paulson and colorofthethoraxandthesizeandpresenceofthepterostigmainthisspecies. Thusweanalyzed thesubstantialseriesfromGhanacollectedforthisstudy.Variation insizewasnot especially noteworthy (Fig. 1),exceptforasinglemalefromDjapoli streamthatfellwell belowtherangeoftheothers, withahindwing length of32.3 mm.Thismaleis dullerthanmostoftheotherspecimens, andits cerciandepiproct arerelatively short,butotherwiseitappearsidenticaltotherest oftheseries.With theexception oftheespecially smallmale, therangeinhindwing length of 34.4- -37.2 mm in males (N = 35) and37.2-39.6mm in females(N = 10) seems not especially large. PINHEY(1951, 1961a) gives therangeformalesas 37-40mm, implying distinctlylargeranimalsin southern and eastern Africathanthosefrom Ghana,andindeed5 males inthePaulson collectionfromKenya andZimbabwe ranged from37.9-39.0mm. Figure 1 doesshow that pterostigma length is quite variablein males.Thepolymorphism apparent(pterostigma vs nopterostigma) is surely not arealpolymorphismbutinsteadrepresents continuousvariationinsize, because thesmallest pterostigmapossible corresponds tothelength ofthecellsat thatpoint onthe wing.Twoofthemales withthesmallestpterostigmas alsohave relatively littlepigmentation inthecellthatwouldbe consideredthepterostigma, andthatcell is not very welldifferentiatedfromadjacent cells. Males thatlack a pterostigma merely lack pigment inany cell. (5) Sapho bicolor Selys, 1853.Kakl (1c?), Kak2 (seen). (6) Sapho ciliata (Fabricius, 1781). Kak2(56, 1$). — Thisspecies wasmore Fig. 1. Hindwing and pterostigma lengthofPhaon iridipennisfrom Ghana. Odonata of Ghana 71 common thanS. bicolorandwas foundatthelargerofthetwo streams inthewet forestatKakum.All specimens collectedwere perched 1-1.5mover thestream. (7) Umma cincta (Hagen inSelys, 1853). Kakl (19),Kak2(26). - Observed both under theshadeofthecanopyand insun undergapsin thecanopy. PROTONEURIDAE (8) Chlorocnemiselongata Hagen in Selys, 1853. Kakl (66, 19). — Males have wings suffusedwithyellow, andtheonly femalecollectedhas clearwings. (9) * Elattoneuranigra Kimmins, 1938. Dja (8c?, 79). - Nearest previous records Coted’IvoiretotheWestandNigeria totheEast.Fivepairs werecollected in tandem,oneover a5 cm deep trickleandoneovera2 mwidestream. (10) Isomecocnemis subnodalis (Selys, 1886). Kakl (19). — The specimen collectedwas ateneral,andthespecific identificationisnot certain,butsubnodalis is theonly speciesknown fromGhanato date. (11) Prodasineuravilliersi Fraser, 1948. Kakl (9c?,29), Kak2 (3c?). - This species isknown fromonly GhanaandtheCote d’Ivoire. PLATYCNEMIDIDAE (12) * Mesocnemisrobusta (Selys, 1886).Vol(5c?). — Thisrecordextendsthe knownrangeofthisspecies, previously known fromNigeriato Sudan,andis the firstrecordWestoftheDahomey Gap.Specimens collectedwere foundless than1 m over theBlackVoltaRiver, wherevegetation hung downfromthebank. (13) * Mesocnemis singularis (Karsch, 1891).Vol (6c?). — Nearest records from Cote d’Ivoireto theWestand Togo to theEast. Specimens were collected only fromthe Black VoltaRiver, where the species coexisted in apparentequal numberswithM. robusta.Bothspecies ofthegenus were coloredsimilarlyinlife, heavily pruinose. (14) * Platycnemis guttifera (Selys, 1886).Kakl(lc?). — Thisrecordextends theknownrangeofthisspecies, previously known fromGuineatoCoted’Ivoire, slightly to theEast. (15) Platycnemis sikassoensis (Martin, 1912).Dja(6c?, 69).During copulation the male opened its wings, then closed them when straightening the abdomen afterwards.Threepairs werecollected intandem. COENAGRIONIDAE (16) * Aciagrion hamoniFraser, 1955.Dja (2c?). — NearestrecordsfromCote d’IvoiretotheWestandtheRepublic ofCongototheSoutheast.Onemalecollected at duskin an openarea with 0.5 mtall grass, somebare soil, andtrees 10-15m apart.PINHEY (1972)describedtheonly maleofthisspecies heretoforereported. 72 G. O'Neill & D.R. Paulson and thetwo specimens collectedinGhana matchhis description quitewell. (17) Agriocnemiszerafica LeRoi, 1915.Oya (1d, 39), Dak (1d,2?). (18) * Ceriagrion bakeri Fraser, 1941.DJa (1d). — Nearestprevious records Coted’Ivoireto theWestandTogo to theEast. (19) Ceriagrion glabrum (Burmeister, 1839).Dak(8c?, 39), Pan(8c?, 7 9),Dja (4d, 19). — Five tandempairs were collected, oneofthemduringoviposition. (20) * Ischnurasenegalensis (Rambur, 1842).Oya (1d, 1 9). — We couldfind norecords closerthanGuineato theWestand Nigeriato theEast. Although this species is oneofthe more ubiquitous odonates ofAfrica (PINHEY, 1962a), it is clearly notwidespread andcommoninGhana,aswerecordeditonlyfromacoastal pond,andneitherNEVILLE(1960)norMARSHALL&GAMBLES(1977)found itattheirstudy sites. Itmaybe generally rare intheWestAfricanforestregion, as itwas not reported fromtherelatively well-sampled NimbaMountainsinGuinea and Cote d’Ivoire (LEGRAND, 1983; 1985),or at Korhogo (LINDLEY, 1974), Lamto(LEGRAND,1982),orTai(LEGRAND&COUTURIER, 1985)intheCote d’Ivoire. (21) * Pseudagrion epiphonematicum Karsch, 1899.Kakl(4d). — Firstrecord betweenGuineato theWest and Nigeria to theEast.This species is restrictedto WestAfrica. (22) Pseudagrion glaucescens Selys, 1876.Dak (3d). — Recordedonly from aslow-moving coastal stream. (23) Pseudagrion hamoniFraser, 1955.Dak (21d, 11 9),Dja (20d, 129). — The thorax of the specimens collectedwas green dorsally and pruinose blue ventrally. Theeyes ofmalesareblack dorsallyandred ventrally,thoseoffemales browndorsally andgreenventrally.Thespecies emerged abundantly fromDjapoli; eight teneralswerepreserved. Pairsoviposited intandeminthestream. (24) Pseudagrionkersteni (Gerstaecker, 1869).Dak (1 d),Dja (9d,2 9). — The eyes areblackdorsally and greenventrally inlife. (25) Pseudagrion melanicterum Selys, 1876.Dja(21 d,69),Kakl (6d),Kak2 (8d,29). (26) * Pseudagrion nubicumSelys, 1876.Oya (Id),Dak(Id). — Knownfrom Cote d’IvoiretotheWestand Nigeria totheEast. (27) Pseudagrion sjostedti Forster, 1906.Dja(6d, 19), Vol(4 d, 19). (28) Pseudagrion sublacteum (Karsch, 1893).Dak (2d),Dja (1d, 1 9). AESHNIDAE (29) * Anaxephippiger (Burmeister, 1839).Pan (Id). — Known from Cote d’Ivoireto theWestand Nigeria totheEast.Recorded only fromthecoastalarea. (30) * Gynacantha cylindrata Karsch, 1891.Dja (29). — KnownfromCote d’Ivoireto the West and Benin to the East. This species was collected at dusk flyingoveranareaofbaresoil40mfromthestream. Odonata of Ghana 73 (31) Gynacantha manderica Griinberg, 1902.Dja (Id, 19). — This species was collectedat dusk. GOMPHIDAE (32) * Crenigomphus reneiFraser, 1936.Dja(Id). — KnownfromCoted’Ivoire totheWestand Nigeria totheEast. (33) * Gomphidia madi Pinhey, 1961.Dja (1 d), Vol (2d, 19). — ThisWest Africanspecies waspreviously knownonlyfromCoted’Ivoire,Nigeria,andUganda. (34) Ictinogomphusferox (Rambur, 1842).Dak (1 d). (35) Lestinogomphus africanus (Fraser, 1926).Vol (Id). — Collected20 m fromalarge river in 1 mtallgrass. (36) * Paragomphus genei (Selys, 1841).Dja (2d),Vol(19). — Knownfrom Coted’IvoiretotheWestandNigeria totheEast,aswellas muchmoreextensively on thecontinent.Theeyes are greendorsally andlightblue ventrally. (37) * Paragomphus nigroviridis (Cammaerts, 1969).Kak2 (1d). — Known fromCoted’IvoiretotheWestandCameroonandZairetotheEast,itappearstobe aspecies ofwet forest, unlike theprevious species, whichis distributedwidely overAfrica. CORDULIIDAE (38) Phyllomacromia bifasciata Martin, 1912.Dja(1 d). (39) * Phyllomacromia pseudafricana Pinhey, 1961.Dja(1d). — Knownfrom Coted’IvoiretotheWestandNigeria totheEastandfairlywidespread throughout WestAfrica. (40) * Phyllomacromia sp. Dja (19). — Thisfemalehasnot beenidentified, butit is surely notany ofthe species knownfrom thecountry, including thetwo species listedaboveaswellas P.melania(Selys),andP.sophia (Selys). Itdoesnot appeartofitineitherthepicta orsophia groupsas definedby GAMBLES(1979). Itisasmall(abdomen38.6mm,hindwing 36.5mm)species withthreeconspicuous stripes oneach sideofthethorax. Thefaceis brown,with asmallyellow spot at eachendofthepostclypeus, thefronsmetallicblue-violetabove.Theabdomenis mostly black, withthefollowing yellow markings: large squarelateralmarksand pair oftransversely orientedmiddorsalspots on 1;fine yellowrings atbases and paired dorsal spotsatmiddlesofsegments3to6, becoming smallertorear; anda conspicuous yellow bandonthebasalthirdofsegment7.The vulvarlaminaeare small(0.7 mm), narrow, andpointed. LIBELLULIDAE (41) Acisomapanorpoides Rambur, 1842.Dak (2c?, 19),Pan (19). 74 G. O'Neill & D.R. Paulson (42) * Aethriamantarezia Kirby, 1889.Dak (1c?). — Recordedonly fromthe coastal area. KnownfromCoted’Ivoireto theWestandNigeria totheEast. (43) * Brachythemis lacustris (Kirby, 1889). Vol(5c?, 89). — Knownfrom Coted’IvoiretotheWestand Nigeria totheEast.Recordedonly fromalargeriver. This species was foundinhigh populations amongthe smallshrubsthatgrew on islands nearthebank oftheriver. (44) Brachythemis leucosticta (Burmeister, 1839).Oya(7<?,29),Dak(2c?), Vol (1c?). — Recordedfromthecoastalareaandonalarge river. (45) Bradinopyga strachani (Kirby, 1900).Sha(2c?, 1?).This genusis usually associatedwith rocky areas, andthe Shai Hillshave abundantrock outcrops. (46) Chalcostephiaflavifrons Kirby, 1889.Dak (4c?), Pan(3c?). (47) Crocothemisdivisa Baumann,1898. Dja-a(2), Sha (It?). — As withthe species of Bradinopyga, thisspecies is typically associatedwithrocky areas, and bothlocalities whereitwas takenhadexposed rocks. (48) Crocothemiserythraea (Brulle, 1832). Dak (19). — Recordedonly from thecoastal area.Thisspecies is very widespread and commonthroughout Africa, and it is surprising that it was nowhere commonat the sites visited duringthis study in Ghana. UnlikeIschnura senegalensis, ithas been foundata numberof WestAfricanforestsites. (49) * Eothemiszygoptera Ris, 1909.Kakl (2c?). — Recordedonly fromwet forest. KnownfromCoted’Ivoireto theWestand Nigeria totheEast. (50)Hemistigmaalbipuncta (Rambur, 1842). Dak (3c?), Pan(29). (51) * Nesciothemispujoli Pinhey, 1971.Dak (6c?), Pan(19). — This species wasdescribedfromtwo malesfromtheCentralAfricanRepublic(PINHEY, 1971) andhas notbeendiscussedsubsequently. Ourspecimens matchPinhey’s description quitewelland differfromaseriesofmaleN. farinosa (Forster, 1898)examined(2 fromKenya, 1 fromMalawi,4fromZimbabwe,2fromSouthAfrica).Thedifference inthe hamulesis readily apparentin theshape ofthe outerhamule, broaderand less deeply indentedinpujoli, narrower andwith a deep longitudinal groove on the outer surface infarinosa. Although Pinhey notedthatpujoli differedfrom farinosa inbeing darkly pruinose, we suspect this might havebeen an artifactof preservation, as the Ghana specimens as preserved show a shade oflight blue pruinosity identicalto thatoffarinosa. The extentofabdominalpruinosity may furnish anotherspecies difference:the six pujoli males are pruinose to thetip of segment3 orextremebaseof4;twoKenyafarinosaarepruinose tothetip orjust shortofthetipof4; andtwo ZimbabweandoneMalawifarinosa arepruinose to thetipof5. PINHEY (1951)stated thatthe basal4-6segmentsofmalefarinosa became pruinose, but there is no indication of the geographic source of the specimens from which he came to this conclusion. In addition, the malepujoli have entirely black labia, whilethe labiaofthe malefarinosa examinedare pale (thetwofromKenyahaveblackonthemedianedges ofthepalpi). Thelabrumand legs ofpujoliare entirely black,unlikethoseofN. minorGambles, 1966,asimilar Odonata of Ghana 75 butsmallerspecies.ThesinglefemaleNesciothemiswas takenatadifferentlocality fromthe malesbutisassociatedwiththembecauseofhersize, wing venation,and general appearance. She appears tobe quite mature, dark all over with light pruinosity on the thoraxand three basalabdominal segments.Unlike the males, herentirefaceandlabiumare lightbrown,asare theouter surfacesofherfemora andtibiae.Thefoliationson abdominalsegment8 are shaped as infarinosa and are pale with dark borders. As the two species are superficially very similar, it wouldbeofinteresttore-examineall specimens of ‘farinosa”fromWestAfrica, whereitisconsideredwidespread(PINHEY, 1962a). (52) * Olpogastra fuelleborni Ris, 1912.Vol(Id, 1$). — Thisrecordextends theknown rangeofthisspecies, previously known fromNigeria to SouthAfrica andKenya, andisthefirstrecordWestoftheDahomey Gap.Recordedonly froma large river.Theeyes weregreeninlife.Thefemalecollectedwas a teneral,perched 2 mabovethewater. (53) Olpogastra lugubris Karsch, 1895. Dak (Id, 19), Vol (7d, 19). — Recordedfromthecoastalareaandalargeriver.Amatingpair perched0.5mabove thewateron abranchapproximately 15mfromthebankof theriver. (54) Orthetrumbrachiale (P. deBeauvois, 1805).Pan-a(Id). (55) Orthetrumchrysostigma (Burmeister, 1839).Dak(3d),Pan(Id),Sha(6d), Dja(ld,39). (56) Orthetrumhintzi Schmidt, 1951.Pan (Id). (57) Orthetrumjulia Kirby, 1900. Dja (2d, 29), Kak2 (Id). — Oneofthe femalescollectedwas ovipositing, guarded by amale. (58) Orthetrumstemmale (Burmeister, 1839).Dak (1d). Pan(1d), Dja(12d, 129),Vol(Id). — Theeyesofthisspecies areblueinlife.Onepairwascollected intandem.Anotherpairwas collectedwhilematingin1mtallgrass.Sevenfemales werecollectedwhileovipositing atDjapoli. Allovipositionoccurredby thefemale dipping the tipofherabdomeninto smalltrickles andpools. Thisis the species listed previously in WestAfrican publications as O. brachiale kalai Longfield, 1936(PINHEY,1979). (59) Palpopleura deceptor (Calvert, 1899). Sha(Id). (60) Palpopleura lucia (Drury, 1773).Dak (lOd, 39), Pan(3d,69),Sha(6d, 49),Dja(13d,69),Vol(Id). — This commonand verywidespread species was collectedateverysiteexceptthewetforest.Twofemaleswereobservedovipositing atDjapoli whilethemalehoveredoverthefemale.Ofthe33malescollected,only two fromDjapoli wereof theformportia,withmuchreducedwing markings. The situation with these two forms remains one of the more puzzling cases of polymorphism in theOdonata. Bothtypes apparently occur throughout therange ofthe species, yet at some localities, only one ofthem isreported. Differences between the forms in extentof wing markings, although long known, have not been quantified, andwe attempted thatby examining 58 male specimens in the PaulsoncollectionfromSierraLeone(2),Liberia(1),Coted’Ivoire(1), Ghana(35), 76 G. O'Neill & D.R. Paulson Fig. 2. Size and wing markings ofPalpopleura lucia from Africa, Congo (2), Uganda (1), Kenya (3), Malawi(1), Zambia(1), Zimbabwe(10),and SouthAfrica(1). Theblackcolorationinthehindwing is typically widest distalto the nodusandnarrowest at orjustproximal to thenodus.Thewidthoftheblack was measured at both of these points, and the differencebetween them (the “indentation”) was taken as a measure oftheextent oftheblack. This was then expressed as aproportion ofthehindwinglength, totakethedragonfly’s size into account. Thesemeasurements are showninFigure2.Itcanbeseenthatthespecimens aredistributedbimodally, corresponding totheformslucia(indentation/hindwing <0.2) andportia (indentation/hindwing>0.2). Twopoints areapparenthere. (I) Thetwo formsoverlapinsize,althoughportiaaveragessmaller, asusuallydescribed (PINHEY, 1951).(2)WestAfricanspecimens ofluciahavewings moreextensively markedwithblack,ontheaverage,thanthosefromEastandSouthAfrica; PINHEY (1961a) noted that the black extended farther out the wing in West African specimens. Although the specimens examined here clearly fall intotwo distinct groupsbasedon wing markings, intermediatesareknown(PINHEY 1984),andthe erectionofportia to specific statusby GAMBLESetal.(1998) ispremature,with nonew informationaboutthebiology ofthetwo forms. (61) Pantalaflavescens (Fabricius, 1798).Dak (2c?, 29),Pan(1c?), Sha(3c?, 49),Dja(2c?, 49),Vol(1c?). — Thisvery widespreadspecies was observedflying in large numbers over an open area at Shai hills. One pair was collectedwhile copulating 15m fromthestream atDjapoli. (62) Tholymis tillarga (Fabricius, 1798).Dja (3c?, 29). — The eyes ofthis species are red dorsally and blue ventrally in life.Four ofthe specimens were collectedinlow flightat dusk nearthestream. (63) Trameabasilaris (P. deBeauvois, 1805).Sha (19).