11 Bull. not. Hist. Mus. Lond. (Zool.)64(2): 111-205 —Issued26November1998 1 TIICNATLffifrt Partial revision of Paracyclops Claus, 1893 (Copepoda, Cyclopoida, Cyclopidae) with PRESented descriptions of four new species xxTjqqs^ SUPHAN KARAYTUG_ / DepartmentofZoology. The"NaturalHistoryMuseum, CromwellRoad. London SW75BD, UK& Schoolof BiologicalSciences, Queen Maryand WestfieldCollege, MileEndRoad, London El 4NS, UK GEOFFREY BOXSHALL A. DepartmentofZoology, TheNaturalHistoryMuseum, CromwellRoad, London SW75BD, UK CONTENTS Introduction 11 MaterialsandMethods 11 Paracyclopsaffinis(G.O.Sars, 1863) 113 Paracyclopspoppei(Rehberg, 1880) 120 Paracyclopsoligarthrus(G.O. Sars, 1909) 128 Paracyclopscanadensis(Willey. 1934) 136 ParacyclopsdilatatusLindberg. 1952 142 Paracyclopshardinginom. nov 145 ParacyclopsbaicalensisMazepova, 1961 151 ParacyclopsyeatmaniDaggett& Davies. 1974 151 ParacyclopswaiarikiLewis, 1974 162 ParacyclopspilosusDussart, 1984 169 Paracyclopscareclum Reid. 1987 172 ParacyclopsnovenariusReid, 1987 176 ParacyclopssmileyiStrayer, 1988 176 Paracyclopsreidaesp.nov 184 Paracyclopsbromeliacolasp.nov 189 Paracyclopspunctatussp.nov 195 Paracyclopsrothatsp.nov 200 Acknowledgements 200 References 204 SYNOPSIS. A partial revision ofthe genus Paracyclops is made based on type material and on collections from numerous localitiesaroundtheworld.ThefollowingParacyclopsspeciesareredescribed: P.affinis(G.O.Sars, 1863),P.poppei(Rehberg, 1880),P.oligarthrus(G.O.Sars, 1909),P.canadensis(Willey, 1934),P.dilatatusLindberg, 1952,P.hardinginomennovum,P. baicalensisMazepova. 1961,P.yeatmaniDaggett&Davis, 1974,P.waiarikiLewis,1974,P.pilosusDussart, 1984,P.carectum Reid, 1987.P.novenariusReid, 1987andP.smileyiStrayer. 1988.Fourspeciesaredescribedasnewtoscience:P.reidaesp.nov., P. rochaisp. nov.,P.punctatussp. nov.,andP. bromeliacolasp.nov. Detailed descriptions of these species are given including several previously overlooked microcharacters, such as the ornamentation of the coxobasis of antenna, the cuticular ornamentation of urosomal somites and the posterior spinular ornamentationoftheswimminglegs,thatareshowntohavesignificanttaxonomicvalueatspecieslevel.Thedetaileddescription ofmalesisrevealedtobe importantindifferentiatingbetweencloselyrelatedspecies. The geographical distributions ofthe species are re-evaluated on the basis ofexamined material and verifiable published records.ItisrevealedthatP.affinisdoesnotoccurinNorthAmericaandallpreviousrecordsofP.affinisinNorthAmericarefer tothenewlydiscoveredP. canadensis. INTRODUCTION have been recorded in all types offreshwater habitats (Karaytug, 1998):RdilatatusLindberg, 1952wasfoundintheDniesterestuary (Ukraine) on the Black Sea (Monchenko, 1977), P. baicalensis Thegenus ParacyclopsClaus, 1893 isoneofnine generacurrently Mazepova, 1961 was collected from great depths in Lake Baikal recognisedasconstitutingthe sub-family Eucyclopinae (Dussart& (Mazepova, 1978), and P. bromeliacola sp. nov. and P. reidae sp. Defaye, 1985; Pospisil & Stoch, 1997).All speciesareknowntobe nov. inhabit pools in the leaf axils of terrestrial Bromeliads. P. benthic although they can sometimes occurin the watercolumn in chiltoni (Thomson, 1882) was recently collected from freshwater the littoralzone. Paracyclopsspeciesaredistributedworldwideand bodies on EasterIsland and is theonly freshwatercopepodon this ©TheNaturalHistoryMuseum, 1998 112 S. KARAYTUG ANDG.A. BOXSHALL Fig.1 P.affinis.Adultfemale.A.maxillule;B,maxilliped;C,body,dorsal;D,maxilla;E,labrum;EG,mandible;H.detailofcaudalseta.Scalebarsin|im. 1 PARACYCLOPS REVISION 13 remote island (Dumont & Martens, 1996). P. oligarthrus (G. O. SPECIES DESCRIPTIONS Sars, 1909) occurs only in LakeTanganyika. The lack of sufficient detail in the original description of the Paracyclops affinis (G. O. Sars, 1863) type-species P.fimbriates (Fischer, 1853) and the publication of various incompletely described species or forms that are closely (Figures 1-7) related to the type-species has created considerable taxonomic CyclopsaffinisSars, 1863: Brady (1878),Vosseler(1886), Schmeil confusion.This has beenexacerbatedby the use ofalimited setof (1892). Brady (1892),Van Douwe (1909), Lilljeborg (1901). traditional characters for differentiating between species within Cyclopspygmaeus Rehberg. 880 the genus, such as the morphology of the caudal rami and leg 5. 1 Cyclops (Heterocyclops)affinis Sars, 1863: Claus (1893a) The P. fimbriatus complex is a particular problem and has been Platycyclops affinis (Sars, 1913-18): Lowndes (1930, 1932) addressed in aseparate paper in which a neotype is designated for Paracyclopssitiseiensis Harada. 1931: Kiefer (1938) P1.9f2i9mbarrieataullsraenddesPc.rfiibmebdri(aKtuasr,ayPt.ucghi&ltoBnoixasnhadlPl.,iimnmiprneustsusa).KiMefoesrt. Cyclops (Paracyclops) affinis Sars, 1863: Gurney (1933) early records of Paracyclops species are unreliable (Karaytug, OriginalDESCRIPTION. CyclopsaffinisSars, 1863:Fork.VidensL- 1998). Selskab. Christiana (Jahr 1862); p. 256. Thegenusnowcontains26speciesand2subspecies.P.fimbriatus Typelocality. Norway is the type species ofthe genus. The redescription of P.fimbriatus (Karaytug&Boxshall,inpressa)fromaneotypecollectedfromone TYPE MATERIAL. Three specimens of P. affinis collected by Sars ofthetypelocalitieshasstabilisedthetaxonomyofP.fimbriatusand including 1 slide (1 female. Reg. No: F7380Zool. Mus. Oslo);one its closely related species P. chiltoni (Thomson, 1882) and P. tube with 1cfand 1 cop. V9( Reg. No: F 20480) examined. Since imminutus Kiefer 1929. Two new species, P. longispina and P. the locality data of Sars' material are not known precisely, the altissimus, fromAfricaaredescribedelsewhere (Karaytugetal., in redescription ofP. affinis is basedon all material examined. press). No material of P. aioiensis Ito, 1957. P. itenoi Ito, 1962, P. Other material examined timmsi Kiefer, 1969, P.fimbriatuspawpamisi Lindberg, 1960, P. - TheNaturalHistoryMuseum.London:229 9 lcffromRingmere, eucyclopoides Kiefer, 1929, P.fimbriatus euchaetus Kiefer, 1939 . England, Reg. No: 1950. 9. 20. 194.Coll: R. Gurney:Calthorpe, cinoeudldinbtehiosbptaaipneerdi.nTdehteairleimnacilnuidnigngsnpeucmieersooufsPparreavciyoculsolypsovaererleoxoakme-d E1n6g9lan9d..23c9fc9f.,1BcfM,NBHMN1H9371.9501.1.9.1260..611993;;NDoervfoolnk,,EEnnggllaanndd,, microcharactersthathavesignificanttaxonomicvalueatthespecies 29 9. Normancoll., BMNH 1911. 11.8.40555-556. dleivlealt.atOusnlLyindpabretriga,l 1r9ed5e2scarnidptPi.opnisloosfusPD.usssmairlte,yi19S8tr4ayweerr.ep1o9s8s8i,blPe. - Germany, Karlsruhe, 1 9dissected on 2 slides, coll: Kiefer in 1935. duetothepoorconditionoftheoriginalslides. Fournewspeciesare nroevc.o,gnainzdedP;. bPr.ormeeildiaaecoslp.ansop.v.n,ovP.. rochai sp. nov., P. punctatus sp. -- TTShhweeedeNNanat,tuurNraaollrmHHiaisnsttocororylly.M.uMBsueMsueNmuH,m,L1o9L1nod1n.odn1o:1n.:81.9l.4c0f5lf5cr0fo-fm5r5o4mPa.lUepsstailnae,, BMNH1938. 3. 9. 83-89 (1030). - Japan. 39 9. Hokkaido, coll: T. Ishida on 4 Nov 1987; 299. MATERIALS AND METHODS Ryuky; Lake Biwa, 59 9dissected on 5 slides; Desaru Beach, Malaya(0o21'N. 104°4'E), 29 9undissectedandmountedon 1 Specimens were dissected and mounted in lactophenol. Broken slide, 1 9dissectedon 1 slide;Abiro,Hokkaido, 1 9dissectedon glass-fibres were added to prevent the appendages from being I slide(42°48'N, 141°50'E); R. Hichi, 29 9. lcfdissectedon 3 slides. ctioomnpwrheiscshedalblyotwheedcvoiveerwsilnigpfarnodmtaollfasciidleista.tAelrlotdartaiwoinnagnsdwmearneipmualdae- - Ethiopia. 1 slide (1 &), Lac Haik. Coll: C. H. Fernando on 1 withtheaidofacameraIucidausinganOlympus BH-2 microscope Aug. 1984. Dissected on 1 slide: Urosome (dorsally), leg 4 with Nomarski differential interference contrast and all measure- (anteriorly) and antennule could be examined but all other ap- mentsmadewithanocularmicrometer.Bodylengthsweremeasured pendages were in poorcondition. fromthebaseoftherostrumtotheposterioredgeofthecaudalrami. Redescription ofadultfemale Body width is given as the widestpart ofthe cephalothorax. In the Body length and width not including caudal setae given in Table spine and setaformulaofthe swimming legs Roman numerals and 1. Genital double-somite, second and third abdominal somites Arabic numerals are used for spines and setae, respectively. The withdorsal surfaceridgesextendinground sides to ventral surface terminology proposedby Huys & Boxshall (1991) is adopted. The as figured (Figure 2A.B). Seminal receptacle divided into broad new nomenclature systemforthe setation elementsofcaudal rami butterfly-shaped anterior and posterior lobes (Figure 2A). Anal was established by Huys (1988) who identified 7 setae (Figure cleft with irregularly arranged spinules (Figure 2B.D). Caudal 2B): anterolateral accessory seta (I) is usually missing in mem- rami short, about twice as long as broad (Figure 2A,B); outer bers ofthe family Cyclopidae but is present in some, forexample terminal seta (IV) and inner terminal seta (V) with complex Metacyclops pseudoanceps (Boxshall & Braide, 1991), II - the spinular ornamentation (Figure 1C); spinular row at base ofante- anterolateral seta, III - the posterolateral seta, IV - the outer rolateral seta (II) extending proximally near inner margin, almost terminal seta, V - the inner terminal seta, VI - the terminal halfway along ramus; terminal accessory seta (VI) shorter than accessory seta, VII - the dorsal seta. The terminology proposed posterolateral seta (III). by Karaytug & Boxshall (in press b) to identify the individual Antennule 11-segmented(Figure3C). Segment6withspiniform setae on the first segment of male antennule is used. The terms seta.Segment9withaesthetasc(Figure3C).Setalformula8,4,2,6, 'frontal' and 'caudal' introducedbyVan deVelde (1984) todenote 4,2,2,3,4+aesthetasc,2+aesthetasc,7+aesthetasc.Coxobasisof the anterior and posterior surfaces ofthe antennary coxobasis are antennawithcomplexornamentationoncaudalandfrontalsurfaces adopted here. asfigured(Figure3A.B);withspinularrownearinnersetae(arrowed 114 S. KARAYTUGANDG.A. BOXSHALL inFigure3B). Firstendopodal segmentwithspinularrownearbase setae, plus 1 modified element (Figure 7F), main part of element ofinnerdistal setacaudally (arrowedinFigure 3B). lying along surface of segment and ornamented with longitudinal Labrum with 3 spinules at either side of free posterior margin ridgesandsmallcentralpore.SegmentalfusionpatternasfollowsI- (arrowed in Figure IE). Mandible with spinular row near base of V,VI-VII,VIII, IX, X, XI, XII, XIII, XIV,XV, XVI, XVII, XVIII, gnathobasic blades (arrowed in Figure IF). Maxillule with XIX-XX, XXI-XXIII, XXIV-XXVIII. proximalmost spine ornamented with spinules (arrowed in Figure Coxobasis ofantenna with spinules nearbase ofinner setae but 1A).Maxilla(Figure 1D)withpraecoxabearingspinularrowdorsally spinulessmallerthanthoseoffemale(Figure6E). Sixthleg(Figure andwithspinularrowonoutermargin.Coxawithscatteredspinules 6C)armedwith 1 innerspinesurroundedbyspinulesatbase;middle alongouteredge. Syncoxaofmaxillipedwithoutspinulesnearbase setaplumoseandaslongasinnerspine;outersetanakedandabout of3setae(arrowedinFigure IB);basiswithspinularrowonanterior halfas longas innerspine. surface and 2 diffuse groups ofspinules on posterior surface. First endopodal segmentwith2tiny spinules on anteriorsurface. Strong Variability, females. Arrangements of spinules on anal cleft may vary (Figure 2D). Coxobasis ofantenna sometimes withextra setafusedtosecondendopodal segment,claw-likeandornamented spinularrow on caudal surface (Figure 3D). with spinules (arrowed inFigure IB). Legs 1 to3withoutmid-distalspinularrowonposteriorsurfaceof Differentialdiagnosis,female. P.affinisisdistinguishedfrom coxa(arrowedinFigures4B,C;5C).Coxaeoflegs2-4withspinular other Paracyclops species by the combination ofits 11-segmented row on anteriorsurface andwith innerspine bearing largepostero- antennule; the surface ridges on the urosomal somites, the spinular lateral spinule (arrowed in Figures 4A; 5A,B); basis with spinular ornamentation ofthe anal cleft, and the presence of 1 seta on the row on anterior surface nearinnermargin (arrowed in Figures4A; secondendopodal segmentofleg4. 5A,B). Innercoxal seta ofleg 1 semispinulose (arrowed in Figure P. affinisandP. canadensisareverycloselyrelated,butP. affinis 4D).Terminalendopodal segmentofleg 3 with spine abouthalfas caneasilybedifferentiatedfromP. canadensisbythepossessionof longassegment(Figure5B).Coxaofleg4withcomplexornamen- threespinesontheterminalexopodal segmentofleg3 (Figure5B), tationonposteriorsurface;intercoxalscleritewithspinularrowson by the presence of spinules at the base of the outer seta of leg 5 anterior and posterior surfaces, and along distal margin (Figure (arrowed in Figure 2C); by having fewer surface ridges on the 5A,D). genital, secondandthirdfreeabdominal somites (Figure2A,B);by Spine and setaformulaas follows: the spinular row not extending either side of anal cleft (Figure 2B,D); bythe structureofthe innercoxal spinesoflegs 2to4; and Coxa Basis Exopod Endopod bythepresenceofaspinularrowontheanteriorsurfaceofthebasis oflegs 2 to4neartothe innermargin (Figures4A; 5A,B). Legl 0-1 1-1 I-1;I-1;III,5 0-1;0-l;1,1,4 Leg2 0-1 1-0 I-1;I-1;III,I,5 0-1:0-1;1,1,4 Remarks and comparisons Leg3 0-1 1-0 I—1;I—1;III,5 0-1:0-2:1,1,4 Historicallytherehasbeensomedisagreementaboutthetaxonomic Leg4 0-1 1-0 I-1;I-1;III,5 0-l;0-l;1,11,2 position ofP. affinis.This species wasoriginally publishedby Sars (1863) under the name Cyclops affinis and this name was used by Leg5 (Figure2C)withlonginnerspine,about4timesaslongas several subsequent authors (Brady, 1878, 1892; Vosseler, 1886; segment;outersetasimple,justlessthanhalfaslongasinnerspine Schmeil, 1892;VanDouwe, 1909)eventhoughSars(1863)didnot and with spinules atbase (arrowed in Figure 2C). mention the ornamentation ofthe caudal rami and did not include Description ofadultmale anydrawings intheoriginalpublication. Rehberg (1880)described BodylengthofspecimenfromEngland(Norfolk):619urnandbody Cyclopspygmaeusasanewspeciesonthebasisofthelengthofthe width: 213 urn. Differing from adult female as follows: Genital caudalsetaeandtheornamentationofthecaudalramiwhichheused somite separate, ornamented with 3 complete, 1 incomplete dorsal todistinguish itfrom C. affinis. Cpygmaeus wasregardedby Sars surfaceridgesand4incompleteventralsurfaceridges;first, second (1913-18) asasynonymofC. affinisandisherealsoconsideredto and third free abdominal somites each with 2 complete dorsal and beasynonymofP. affinis. Claus (1893a)placed C. affinisinanew ventral surface ridges (Figure 6A.B). subgenus,Heterocyclopsonthebasisofthepatternofdevelopment Antennuledigeniculate(Figure7A,B),indistinctly 16-segmented. oftheantennule. LaterSars (1913-18) included C. affinisin anew Segment 1 armedwith8setae;setaAsimple(arrowedinFigure7E). genus, Platycyclops, but ignored or overlooked earlier work by Segment 10 (= ancestral segment XV) produced on one side into Claus (1893). Platycyclops is a synonym of Paracyclops Claus, sheath enclosing segment 11 ventrally: armed with 2 setae, one of 1893.TheinadequacyofSars'sdescriptionofC.affinis(Sars, 1913- which pear-shaped and constricted apically, constricted part bent 18) prompted Lowndes (1932) to redescribe C. affinis, correcting slightlyinwardsandwithsmallterminalseta-likeprocess,otherseta some errors in Sars's descriptions. Harada (1931) distinguished P. long and naked. Segment 11 bearing curved seta ornamented with sitiseiensisfromP. affinisonthebasisoftheproportional lengthof doublerow ofstrong denticles but notas strong as in P.fimbhatus the spines ofleg 4 and the stronger inner spine ofleg 5, however, group; plus 1 naked seta (Figure 7E,F). Segment 12 armed with these characters are not significantly different from P. affinis des- curved seta similar to that ofeleventh segment, plus short highly cribedherein. Therefore P. sitiseiensisis regardedas asynonymof chitinized seta. Segment 13 armed with 2 short naked setae. Seg- P. affinis,asalready indicatedbyMonchenko(1974).Thelengthof ment 14armedwith 1 shortspinulatesetaeproximally,2shortnaked the innerspines offifth and sixth legsofmale P. affinis from Lake Table1 Bodylength(BL)andwidth(BW)measurements(inurn)ofP.affinisfromvariouslocalities(N=numberofspecimensmeasured) Locality Sex BL(mean±SD) Range BW(mean±SD) Range N England(Ringmere) 9 709± 12 684-723 267±5.5 254-272 10 England(Norfolk) 9 692±20.2 657-731 261 ±11.7 244-281 13 Sweden(Upsala) 9 827±60 753-877 269±7.8 262-277 4 115 PARACYCLOPS REVISION Fig.2 P. affinis.Adultfemale.A,urosome,ventral;B,urosome,dorsal;C,leg5,ventral;D,analsomite,dorsal.Scalebarsin\im. 116 S. KARAYTUGANDG.A. BOXSHALL Fig.3 P. affinis.Adultfemale.A.antenna,coxobasis,frontal;B,antenna,caudalshowingtypicalspinulation;C,antennule;D,antenna,coxobasis,caudal showingvariantpatternofspinulation.Scalebarsinu.m. PARACYCLOPS REVISION 117 50 Fig.4 P.affinis.Adultfemale.A,leg2,anterior;B,intercoxalscleriteandcoxaofleg2,posterior;C,intercoxalscleriteandcoxaofleg 1,posterior;D, leg 1,anterior.Scalebarin\xm. 118 S. KARAYTUGANDG.A. BOXSHALL Fig.5 P. affinis.Adultfemale.A,leg4.anterior;B,leg3,anterior;C,intercoxalscleriteandcoxaofleg3,posterior;D. intercoxalscleriteandcoxaofleg 4,posterior. Scalebarin|im. PARACYCLOPS REVISION 119 Fig.6 P. affinis.Adultmale.A,urosome,ventral;B,urosome,dorsal;C,detailofleg6,ventral;D,detailofleg5,ventral;E,antenna,coxobasis,caudal. Scalebarsinp.m. 120 S. KARAYTUGANDG.A. BOXSHALL Fig.7 P.ajfinis.Adultmale.A,antennule,ventralshowingsegmentation;B,dorsalshowingsegmentation;C,body,dorsal;D,detailofsetationelements ofcaudalrami;E,anteroventralshowingsetation;F,detailofsegments 12to 15.Scalebarsinurn.