Bm (Oss VY) Zoology Series Sy THE NATURAL HISTORY MUSEUM VOLUME 64 NUMBER2 26 NOVEMBER 1998 7 The Bulletin of The Natural History Museum (formerly: Bulletin of the British Museum (Natural History) ), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology. The Zoology Series is edited in the Museum’s Department of Zoology Keeper of Zoology Prof P.S. Rainbow Editor of Bulletin: Dr N.R. Merrett Assistant Editor: Dr B.T. Clarke Papers in the Bulletin are primarily the results of research carried out on the unique and ever- growing collections of the Museum, both by the scientific staff and by specialists from elsewhere who make use of the Museum’s resources. Many of the papers are works of reference that will remain indispensable for years to come. All papers submitted for publication are subjected to external peer review for acceptance. 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(Zool.) © The Natural History Museum, 1998 Zoology Series ISSN 0968-0470 Vol. 64, No. 2, pp. 111-211 The Natural History Museum Cromwell Road London SW7 5BD Issued 26 November 1998 Typeset by Ann Buchan (Typesetters), Middlesex Printed in Great Britain by Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset Bull. nat. Hist. Mus. Lond. (Zool.) 64(2): 111-205 Partial revision of Paracyclops Claus, 189 (Copepoda, Cyclopoida, Cyclopidae) with descriptions of four new species Aid BS EEL SUPHAN KARAYTUG , Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 SBD, UK & School of Biological Sciences, Queen Mary and Westfield College, Mile End Road, London El 4NS, UK GEOFFREY A. BOXSHALL Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK CONTENTS IaH OTOH TAG nim tea men acy Renee nore eBCONe, cet etcetera SHORE ee AUR 3.2 Me AA ae ae ne nO eR Rene OPE mcr AeP oar 111 WTAE RTAISTANCINI GIRO S oeziczesepeanaersenenneneset sensation ex tepesac cuae ac vas conigM xis tonS ances conc xncis dpoxuatch nan vawsdes dae ta dias San aig ancane icavsnesvecasunsuoeane 111 NOMA GLAS {CI | ELOS (Gra) SATS UO S)) act taste co ae eco tae sa cscs es Uncan sccm eed cosas ata - nag abe see aca naen en veb sc ceSae tun densavos cers sans 113 EONACVCLO IS PORDET INCI DEL OMS O UL) o stecatccnuxsccsresseraaunvyneuaay rt aarsaar ecu cesvanesecetuanstesscastenadtetectssracdecpotsuaceseuas «sus cuescetsensresisavertGdase 120 EATAGYGIGDS OUSAKINTUS (GOs Sats IO) acccessseucstte-aovasvacnscanacieicnn tt ncoenh« Deas tetes Meee sades Matas Ayan Astecse nk cans sade tyeuneseoseaveasssearencees 128 PARAGY CLO DCAM AGETISTS CW MLLEYs UGS) ast voces ceeccse etenezet schaossdavccasocy atte ates Na eae eateries dah ate Nowsatigs Meszcextuteeesee tous teeeeds 136 EMAC CLO TISIOU CICTUS eM CL Ee ODA | wereqacveissvanerasccteiinxacestslsdeva cost suscdvenasssensaunade ot cd aves Mucvat ane debenetsodshunstaaee st Mase suansvadtt ssditvaae ss 142 Lexaipgarea Coop aby LA Tog @ Lay Neg 75810) 0 tO)! Hacer Cer eoce eo Bo RR SORES Peer Pech 7 CSREES EEA Per A PR PEER ORE oe Coe aero RR) eee 145 Paracyclops baicalensis Mazepova, 1961 ......ccceccccceeees Ney Paracyclops yeatmani Daggett & Davies, 1974 151 Paracyclops waiariki Lewis, 1974 162 Paracyclops pilosus Dussart, 1984 169 Paracyclops carectum Reid, 1987 ye: Paracyclops novenarius Reid, 1987 176 Paracyclops smileyi Strayer, 1988 176 MGCERELCM GL) ISU CLLLEAS PION tara cavecerwscecstewcxtacencxivtevavsssitetsusevevetndcacsGarccscurcapstesedhstevacenns suateerceteseTeceesesnsseatesasess tuacsvasveeeestuserdasvers 184 PaCOP ACN CCI ISU PO TIE LECAG ODL EL STO I) OM cane eons weasel /an ate rate ua pees temic «cadens taettevee as fesdbenacaneaat idenatstetecevenvoue qT Fig.1 P affinis. Adult female. A, maxillule; B, maxilliped; C, body, dorsal; D, maxilla; E, labrum; FG, mandible; H, detail of caudal seta. Scale bars in um. PARACYCLOPS REVISION remote island (Dumont & Martens, 1996). P. oligarthrus (G. O. Sars, 1909) occurs only in Lake Tanganyika. The lack of sufficient detail in the original description of the type-species P. fimbriatus (Fischer, 1853) and the publication of various incompletely described species or forms that are closely related to the type-species has created considerable taxonomic confusion. This has been exacerbated by the use of a limited set of traditional characters for differentiating between species within the genus, such as the morphology of the caudal rami and leg 5. The P. fimbriatus complex is a particular problem and has been addressed in a separate paper in which a neotype is designated for P. fimbriatus and P. fimbriatus, P. chiltoni and P. imminutus Kiefer, 1929 are all redescribed (Karaytug & Boxshall, in press a). Most early records of Paracyclops species are unreliable (Karaytug, 1998). The genus now contains 26 species and 2 subspecies. P. fimbriatus is the type species of the genus. The redescription of P. fimbriatus (Karaytug & Boxshall, in press a) from a neotype collected from one of the type localities has stabilised the taxonomy of P. fimbriatus and its closely related species P. chiltoni (Thomson, 1882) and P imminutus Kiefer 1929. Two new species, PR. longispina and P. altissimus, from Africa are described elsewhere (Karaytug et al., in press). No material of P. aioiensis It6, 1957, P. uenoi It6, 1962, P. timmsi Kiefer, 1969, P. fimbriatus paropamisi Lindberg, 1960, P. eucyclopoides Kiefer, 1929, P. fimbriatus euchaetus Kiefer, 1939 could be obtained. The remaining species of Paracyclops are exam- ined in this paper in detail including numerous previously overlooked microcharacters that have significant taxonomic value at the species level. Only partial redescriptions of P smileyi Strayer, 1988, P. dilatatus Lindberg, 1952 and P. pilosus Dussart, 1984 were possible due to the poor condition of the original slides. Four new-species are recognized; P. reidae sp. nov., P. rochai sp. nov., P. punctatus sp. noy., and P. bromeliacola sp. nov. MATERIALS AND METHODS Specimens were dissected and mounted in lactophenol. Broken glass-fibres were added to prevent the appendages from being compressed by the coverslip and to facilitate rotation and manipula- tion which allowed viewing from all sides. All drawings were made with the aid of a camera lucida using an Olympus BH-2 microscope with Nomarski differential interference contrast and all measure- ments made with an ocular micrometer. Body lengths were measured from the base of the rostrum to the posterior edge of the caudal rami. Body width is given as the widest part of the cephalothorax. In the spine and seta formula of the swimming legs Roman numerals and Arabic numerals are used for spines and setae, respectively. The terminology proposed by Huys & Boxshall (1991) is adopted. The new nomenclature system for the setation elements of caudal rami was established by Huys (1988) who identified 7 setae (Figure 2B): anterolateral accessory seta (I) is usually missing in mem- bers of the family Cyclopidae but is present in some, for example Metacyclops pseudoanceps (Boxshall & Braide, 1991), II — the anterolateral seta, III — the posterolateral seta, IV — the outer terminal seta, V — the inner terminal seta, VI — the terminal accessory seta, VII — the dorsal seta. The terminology proposed by Karaytug & Boxshall (in press b) to identify the individual setae on the first segment of male antennule is used. The terms ‘frontal’ and ‘caudal’ introduced by Van de Velde (1984) to denote the anterior and posterior surfaces of the antennary coxobasis are adopted here. LS SPECIES DESCRIPTIONS Paracyclops affinis (G. O. Sars, 1863) (Figures 1-7) Cyclops affinis Sars, 1863: Brady (1878), Vosseler (1886), Schmeil (1892), Brady (1892), Van Douwe (1909), Lilljeborg (1901). Cyclops pygmaeus Rehberg, 1880 Cyclops (Heterocyclops) affinis Sars, 1863: Claus (1893a) Platycyclops affinis (Sars, 1913-18): Lowndes (1930, 1932) Paracyclops sitiseiensis Harada, 1931: Kiefer (1938) Cyclops (Paracyclops) affinis Sars, 1863: Gurney (1933) ORIGINAL DESCRIPTION. Cyclops affinis Sars, 1863: Forh. Vidensk.- Selskab. Christiana (Jahr 1862); p. 256. TYPE LOCALITY. Norway TYPE MATERIAL. Three specimens of P. affinis collected by Sars including | slide (1 female, Reg. No: F 7380 Zool. Mus. Oslo); one tube with 1c’and | cop. V 2( Reg. No: F 20480) examined. Since the locality data of Sars’ material are not known precisely, the redescription of P. affinis is based on all material examined. OTHER MATERIAL EXAMINED — TheNatural History Museum, London:22 2 9 ,1c'fromRingmere, England, Reg. No: 1950. 9. 20. 194. Coll: R. Gurney; Calthorpe, England, 32 9, 10°, BMNH 1950. 9. 20. 193; Norfolk, England, 162 2, 2¢0°'o°, BMNH 1937. 11. 16. 619; Devon, England, 22 2, Norman coll., BMNH 1911. 11. 8. 40555-556. — Germany, Karlsruhe, | 9 dissected on 2 slides, coll: Kiefer in 1935. — The Natural History Museum, London: 192, 1o’from Upsala, Sweden, Norman coll., BMNH 1911. 11. 8. 40550-554. — The Natural History Museum, London: 1c’ from Palestine, BMNH1938. 3. 9. 83-89 (1030). — Japan, 32 2, Hokkaido, coll: T. Ishida on 4 Nov 1987; 29 9, Ryuky; Lake Biwa, 5 9 9 dissected on 5 slides; Desaru Beach, Malaya (0°21'N, 104°4'E), 29 9 undissected and mounted on | slide, | 9 dissected on | slide; Abiro, Hokkaido, 1 9 dissected on 1 slide (42°48'N, 141°50'E); R. Hichi, 22 9, 1 O’dissected on 3 slides. — Ethiopia, | slide (1 &), Lac Haik. Coll: C. H. Fernando on 11 Aug. 1984. Dissected on | slide: Urosome (dorsally), leg 4 (anteriorly) and antennule could be examined but all other ap- pendages were in poor condition. REDESCRIPTION OF ADULT FEMALE Body length and width not including caudal setae given in Table 1. Genital double-somite, second and third abdominal somites with dorsal surface ridges extending round sides to ventral surface as figured (Figure 2A,B). Seminal receptacle divided into broad butterfly-shaped anterior and posterior lobes (Figure 2A). Anal cleft with irregularly arranged spinules (Figure 2B,D). Caudal rami short, about twice as long as broad (Figure 2A,B); outer terminal seta (IV) and inner terminal seta (V) with complex spinular ornamentation (Figure 1C); spinular row at base of ante- rolateral seta (II) extending proximally near inner margin, almost halfway along ramus; terminal accessory seta (VI) shorter than posterolateral seta (III). Antennule | 1-segmented (Figure 3C). Segment 6 with spiniform seta. Segment 9 with aesthetasc (Figure 3C). Setal formula 8, 4, 2, 6, 4,2,2,3,4+aesthetasc, 2 + aesthetasc, 7 + aesthetasc. Coxobasis of antenna with complex ornamentation on caudal and frontal surfaces as figured (Figure 3A,B); with spinular row near inner setae (arrowed 114 in Figure 3B). First endopodal segment with spinular row near base of inner distal seta caudally (arrowed in Figure 3B). 4 Labrum with 3 spinules at either side of free posterior margin (arrowed in Figure 1E). Mandible with spinular row near base of gnathobasic blades (arrowed in Figure 1F). Maxillule with proximalmost spine ornamented with spinules (arrowed in Figure 1A). Maxilla (Figure 1D) with praecoxa bearing spinular row dorsally and with spinular row on outer margin. Coxa with scattered spinules along outer edge. Syncoxa of maxilliped without spinules near base of 3 setae (arrowed in Figure 1B); basis with spinular row on anterior surface and 2 diffuse groups of spinules on posterior surface. First endopodal segment with 2 tiny spinules on anterior surface. Strong seta fused to second endopodal segment, claw-like and ornamented with spinules (arrowed in Figure 1B). Legs | to 3 without mid-distal spinular row on posterior surface of coxa (arrowed in Figures 4B,C; 5C). Coxae of legs 24 with spinular row on anterior surface and with inner spine bearing large postero- lateral spinule (arrowed in Figures 4A; 5A,B); basis with spinular row on anterior surface near inner margin (arrowed in Figures 4A; 5A,B). Inner coxal seta of leg 1 semispinulose (arrowed in Figure 4D). Terminal endopodal segment of leg 3 with spine about half as long as segment (Figure 5B). Coxa of leg 4 with complex ornamen- tation on posterior surface; intercoxal sclerite with spinular rows on anterior and posterior surfaces, and along distal margin (Figure 5A,D). Spine and seta formula as follows: Coxa Basis Exopod Endopod Leg | 0-1 1-I I-1;I-1;H1,5 0-1;0-1;1,1,4 Leg 2 0-I 1-0 13 ES WOES) 0-1;0-1;1,1,4 Leg 3 0-I 1-0 I-1;L-1;11,5 0-1;0-2;1,1,4 Leg 4 0-1 1-0 1-1;J-1;111,5 0-1;0-1;1,11,2 Leg 5 (Figure 2C) with long inner spine, about 4 times as long as segment; outer seta simple, just less than half as long as inner spine and with spinules at base (arrowed in Figure 2C). DESCRIPTION OF ADULT MALE Body length of specimen from England (Norfolk): 619 um and body width: 213 um. Differing from adult female as follows: Genital somite separate, ornamented with 3 complete, 1 incomplete dorsal surface ridges and 4 incomplete ventral surface ridges; first, second and third free abdominal somites each with 2 complete dorsal and ventral surface ridges (Figure 6A,B). Antennule digeniculate (Figure 7A,B), indistinctly 16-segmented. Segment | armed with 8 setae; setaA simple (arrowed in Figure 7B). Segment 10 (= ancestral segment XV) produced on one side into sheath enclosing segment 11 ventrally: armed with 2 setae, one of which pear-shaped and constricted apically, constricted part bent slightly inwards and with small terminal seta-like process, other seta long and naked. Segment 11 bearing curved seta ornamented with double row of strong denticles but not as strong as in P. fimbriatus group; plus | naked seta (Figure 7E,F). Segment 12 armed with curved seta similar to that of eleventh segment, plus short highly chitinized seta. Segment 13 armed with 2 short naked setae. Seg- ment 14 armed with | short spinulate setae proximally, 2 short naked S. KARAYTUG AND G.A. BOXSHALL setae, plus 1 modified element (Figure 7F), main part of element lying along surface of segment and ornamented with longitudinal ridges and small central pore. Segmental fusion pattern as follows I— V, VI-VIL, VIII, IX, X, XI, XII, XI, XIV, XV, XVI, XVII, XVII, XIX-XX, XXI-EXXII, XXIV-XXVII. Coxobasis of antenna with spinules near base of inner setae but spinules smaller than those of female (Figure 6E). Sixth leg (Figure 6C) armed with | inner spine surrounded by spinules at base; middle seta plumose and as long as inner spine; outer seta naked and about half as long as inner spine. VARIABILITY, FEMALES. Arrangements of spinules on anal cleft may vary (Figure 2D). Coxobasis of antenna sometimes with extra spinular row on caudal surface (Figure 3D). DIFFERENTIAL DIAGNOSIS, FEMALE. P. affinis is distinguished from other Paracyclops species by the combination of its 11-segmented antennule; the surface ridges on the urosomal somites, the spinular ornamentation of the anal cleft, and the presence of | seta on the second endopodal segment of leg 4. P. affinis and P. canadensis are very closely related, but P. affinis can easily be differentiated from P. canadensis by the possession of three spines on the terminal exopodal segment of leg 3 (Figure 5B), by the presence of spinules at the base of the outer seta of leg 5 (arrowed in Figure 2C); by having fewer surface ridges on the genital, second and third free abdominal somites (Figure 2A,B); by the spinular row not extending either side of anal cleft (Figure 2B,D); by the structure of the inner coxal spines of legs 2 to 4; and by the presence of a spinular row on the anterior surface of the basis of legs 2 to 4 near to the inner margin (Figures 4A; 5A,B). REMARKS AND COMPARISONS Historically there has been some disagreement about the taxonomic position of P. affinis. This species was originally published by Sars (1863) under the name Cyclops affinis and this name was used by several subsequent authors (Brady, 1878, 1892; Vosseler, 1886; Schmeil, 1892; Van Douwe, 1909) even though Sars (1863) did not mention the ornamentation of the caudal rami and did not include any drawings in the original publication. Rehberg (1880) described Cyclops pygmaeus as a new species on the basis of the length of the caudal setae and the ornamentation of the caudal rami which he used to distinguish it from C. affinis. C. pygmaeus was regarded by Sars (1913-18) as a synonym of C. affinis and is here also considered to be a synonym of P. affinis. Claus (1893a) placed C. affinis in a new subgenus, Heterocyclops on the basis of the pattern of development of the antennule. Later Sars (1913-18) included C. affinis in a new genus, Platycyclops, but ignored or overlooked earlier work by Claus (1893). Platycyclops is a synonym of Paracyclops Claus, 1893.The inadequacy of Sars’s description of C. affinis (Sars, 1913— 18) prompted Lowndes (1932) to redescribe C. affinis, correcting some errors in Sars’s descriptions. Harada (1931) distinguished P sitiseiensis from P. affinis on the basis of the proportional length of the spines of leg 4 and the stronger inner spine of leg 5, however, these characters are not significantly different from P. affinis des- cribed herein. Therefore P. sitiseiensis is regarded as a synonym of P. affinis, as already indicated by Monchenko (1974). The length of the inner spines of fifth and sixth legs of male P. affinis from Lake Table 1 Body length (BL) and width (BW) measurements (in pm) of P. affinis from various localities (N = number of specimens measured) Locality Sex BL (mean + SD) England (Ringmere) Q 709 + 12 England (Norfolk) Q 692 + 20.2 Sweden (Upsala) fe) 827 + 60 Range BW(mean + SD) Range N 684-723 267 + 5.5 254-272 10 657-731 261 + 11.7 244-281 13 753-877 269 + 7.8 262-277 4 115 PARACYCLOPS REVISION ome, dorsal; C, leg 5, ventral; D, anal somite, dorsal. Scale bars in Lm. tral; B, uros me, ven Fig. 2 P. affinis. Adult female. A, uroso 116 S. KARAYTUG AND G.A. BOXSHALL Fig. 3 P. affinis. Adult female. A, antenna, coxobasis, frontal; B, antenna, caudal showing typical spinulation; C, antennule; D, antenna, coxobasis, caudal showing variant pattern of spinulation. Scale bars in um. PARACYCLOPS REVISION 117 Uusday a ee UU uh ey “sR AM Oe, SNS — Ge ( wid Jays 62" bndati\y, Fig.4 P. affinis. Adult female. A, leg 2, anterior; B, intercoxal sclerite and coxa of leg 2, posterior; C, intercoxal sclerite and coxa of leg 1, posterior; D, leg 1, anterior. Scale bar in um. 118 S. KARAYTUG AND G.A. BOXSHALL Cz \ See = = SSS ee EK Any N IN = S ne ; ee t c= ay i MA SS > \ aie “ ay, \) N Coy. MW | gj oil y rw 4 aha yy Wy 50 Fig.5 P. affinis. Adult female. A, leg 4, anterior; B, leg 3, anterior; C, intercoxal sclerite and coxa of leg 3, posterior; D, intercoxal sclerite and coxa of leg 4, posterior. Scale bar in um. PARACYCLOPS REVISION aaa \ \ TL i Fig.6 P. affinis. Adult male. A, urosome, ventral; B, urosome, dorsal; C, detail of leg 6, ventral; D, detail of leg 5, ventral; E, antenna, coxobasis, Scale bars in um. 119 caudal. S. KARAYTUG AND G.A. BOXSHALL 120 a Z aa ntee aa ppp tL ee aa ee LE LEE pare c= = ————— Zs LZ SN aan ae WS 3) NES VAG gts ? . & a. ISS mT S wa aaaadadalll ; =S> 7 aS ) eo 4 p z va pL = z > ip 5 ne (E Fay \ Fig.7 P. affinis. Adult male. A, antennule, ventral showing segmentation; B, dorsal showing segmentation; C, body, dorsal; D, detail of setation elements of caudal rami; E, anteroventral showing setation; F, detail of segments 12 to 15. Scale bars in um. PARACYCLOPS REVISION Tanganyika given by Lindberg (1951) is significantly shorter than in the material upon which this description is based. It is possible that Lindberg (1951) was dealing with a new species, but Lindberg’s (1951) description is based on the male only and lacks sufficient detail to make further comparisons. P. yeatmani is another species of Paracyclops possessing an | 1- segmented antennule, however, analysis of segmental homologies between P. canadensis, P. affinis and P. yeatmani has revealed that the antennulary segments in P yeatmani are not all homologous with those of P. affinis and P. canadensis (Karaytug & Boxshall, 1998). DISTRIBUTION: This revision has indicated that P. affinis does not occur in North America. All existing records of P. affinis from North America refer to the newly re-discovered P canadensis, however the presence of P. affinis in Africa as well as in Europe, Japan and Malaya has been confirmed. This species is probably widely distributed throughout the Palaearctic region but is not found in the Nearctic. Australian records (Sars, 1913—1918; Timms & Morton, 1988) of this species must be confirmed. For detailed references concerning the distribution of P affinis, see Karaytug (1998). Paracyclops poppei (Rehberg, 1880) (Figures 8-13) Cyclops poppei Rehberg, 1880 Cyclops crassicornis O. F. Miiller, 1785 sensu Herrick (1882) Cyclops fimbriatus Fischer, 1853 sensu Herrick (1884), Schmeil (1891), Marsh (1892, 1910), Byrnes (1909) Cyclops fimbriatus var Poppei Rehberg, 1880: Schmeil (1892), Van Douwe (1909) Cyclops (Paracyclops) fimbriatus poppei Rehberg, 1880: Gurney (1933) Cyclops fimbriatus poppei Rehberg, 1880: Harding & Smith (1960). Paracyclops fimbriatus poppei (Rehberg, 1880): Yeatman (1959) ORIGINAL DESCRIPTION. Cyclops poppei Rehberg, 1880: Abh. Natur. Ver. Bremen, 6, p. 550, Taf. VI, figs. 9-11. TYPE LOCALITY. Bremen, Germany TYPE MATERIAL. Lost MATERIAL EXAMINED — Germany, Oldenburg, 49 9, 1c’, collected by T. Ishida on 2 Aug. 1996. — The Natural History Museum, London: Derby, Mauchline, Catrine: 20°o",8 2 2 , collected by M.A. Learner. BMNH 1968. 8. 19. 3-6. — National Museum of Natural History, Smithsonian Institution, Washington: Louisiana, New Orleans, E. New Orleans, in tyre at Grant Street near Old Gentilly Road and Almonaster Avenue (21 June 1988), USNM cat: 252018, Acc. No: 373882, 792 9, 50’ CO", 6 copepodids were examined and one of each sex was dissected; Louisiana, New Orleans, in tree hole at Louisiana Science and Nature Center, 89 9, 50°O", 2 copepodids (1 female dissected), USNM cat: 252019; Virginia, Giles C., Hill- side seep near mountain lake, tiny pool on path around lake (37°21'33"N, 080°32'11"W), 100 + 9 ,c’ collected by J. W. Reid on 15 June 1990 (1 Q and 1C’dissected), USNM cat: 250443, Acc. No: 359834; New York, Pond at town landfill, town of Northeast, Dutchess ca., NY, 1C’and 1 2 mounted on | slide, collected by D. Strayer on 8 Oct 1985, USNM cat: 235366; New 121 Mexico, Guadalupe River, Jemez National Forest about 40 km NE of San Ysidro, 1 9 (35°45'N 106°50'W), 26 May 1991, USNM cat: 251151; Mexico, Aguascalientes, Calvillo, Presa Penuelas (23 March 1987), 2 2 9, 3 copepodids, USNM cat: 234218; Japan, Lake Biwa, Shiga Prefecture, 32 9 and 1 mounted on | slide collected by T. Ishida on 17 March 1986, USNM cat: 250682. — Russia, R. Ravan 100 km East of St.Petersburg district, 72 2, 30°C’, collected by V. Alekseev (22 July 96). — Canadian Museum of Nature: Ontario, Frontenac Cty, nearArden on Hwy. 7, pond, collected by Brenda J. Hann (2 June 1972), 29 Qand lo’, CMNC: 1984-0348, Acq: IZ 1984-064; Ontario, New Islands, collected by L. Kerr (29 Aug 1969), 2 9 2, CMNC: 1984-0370, Acq: 1969-227. — Japan, Lake Biwa, Honshu, T. Ishida collection (17 March 1986), 22 P dissected on 2 slides; 20" CO'dissected on 2 slides. REDESCRIPTION OF ADULT FEMALE Body length and width given in Table 2. Urosome (Figure 8A,B) with genital double-somite and second and third abdominal somites ornamented with fine pits on dorsal surface as figured (Figure 8B). Anal operculum smooth; spinular rows present on either side of anal cleft as figured (Figure 8B). Caudal rami length and width given in Table 3. Caudal rami parallel, with fine cuticular depressions on ventral surface. Dorsal row of spinules on rami extending proximally, nearly reaching base of rami (Fig- ure 8B). Antennule 8-segmented. Setal formula 8, 12, 6,5, 2 + aesthetasc, 2, 2 + aesthetasc, 7 + aesthetasc; coxobasis of antenna without spinular row on caudal surface near base of inner spinulose setae (arrowed in Figure 9H). Terminal endopodal segment of leg 2 (Figure 10D) with stout spine, as long as segment; intercoxal sclerite of leg 3 with spinular row on anterior surface (Figure 11A) and with 3 spinular rows posteriorly (Figure 11C); intercoxal sclerite of leg 4 with patch of spinules on anterior surface and with 3 spinular rows on posterior surface (Figure 11D,E). Spine and seta formula as follows: Coxa Basis Exopod Endopod Leg | 0-1 1-I J-1;I-1;11,5 0-1;0-1;1,1,4 Leg 2 0-I 1-0 J—-1;]-1;I11,1,5 0-1;0-2;1,1,4 Leg 3 O-I 1-0 I-1;1-1;111,1,5 0-1;0-2;1,1,4 Leg 4 0-1 1-0 I-1;I-1;IL1L5 O-1;0-2;1, 11,2 Leg 5 (Figure 8C) comprising single free segment, armed with 1 inner spine, well developed outer spinulose seta as long as inner spine, | plumose seta in middle. DESCRIPTION OF ADULT MALE Body length and width given in Table 2. Caudal rami short (Figure 12C,D), about 2.5 times longer than broad; coxobasis of antenna with spinular row on caudal surface at base of inner spinulose setae (arrowed in Figure 12F), first endopodal segment with 2 spinular rows on frontal surface. Antennule digeniculate (Figure 13A—F), indistinctly 15-seg- mented. Segment | armed with 8 setae; setae (A) and (C) are large and modified (arrowed in Figure 13E) by ornamentation of strong spinules in proximal and mid sections, tapering to fine point distally (Figure 13A,E,F). Segmental fusion pattern as follows: I-V, VI-VIL, VIll, IX, X, XI, XII, XII, XIV, XV, XVI, XVII, XVII-XX, XXI- XXII, XXIV—XXVII. S. KARAYTUG AND G.A. BOXSHALL 122 Hi / | A, LEE == z LL == —— —— \ es es aad AGL) = WSANSSSSSSISSEKSSRESSESES ASS ie) UJ SIIITS = © yqmae (EES eee Se } . Oe a ae ASS SS oO = — G SS al. Scale bars in Lm. ome, dorsal; C, leg 5, ventr: tral; B, uros me, ven Fig. 8 P. poppei. Adult female. A, uroso PARACYCLOPS REVISION AN Y ) vy M SN hi ee AN 3 Ka 2 ju 4 Ma “Dy “fh a Li Fig.9 P. poppei. Adult female. A (Virginia, U.S.A.), caudal ramus, dorsal; B (New Orleans, U.S.A.), caudal ramus, dorsal; C (Lake Biwa, Japan), leg 2, intercoxal sclerite, anterior; D (Lake Biwa, Japan), leg 4, intercoxal sclerite, anterior: E (Lake Biwa, Japan), antennule, segments 2 to 4, showing incomplete suture; F (Virginia, U.S.A.), leg 4, intercoxal sclerite, posterior; G, antenna, coxobasis, frontal; H, antenna, caudal. Scale bars in lum. 123 124 S. KARAYTUG AND G.A. BOXSHALL irr sy Alley iy |p Z Wy PH A 7 AN Vif owe bb AD AdbAAdy yppdd Fig. 10 P. poppei. Adult female. A, intercoxal sclerite and coxa of leg 1, posterior; B, leg 1, anterior; C, intercoxal sclerite and coxa of leg 2, posterior; D, leg 2, anterior; E, adult male, third endopodal segment of leg 1. Scale bar in um. PARACYCLOPS REVISION A A sng i y 3 oe GZ 22 Lif A [RE SE 2 NING Y) it or xk . iy) | | Wes Sy, : EF — ~*~ \ C D Lf Sass hed cal a ys wo : a ayssJO be AMM (AIBe SS eS LLU WW SSN Fig. 11 P poppei. Adult female. A, leg 3, anterior; B, coxa and basis of leg 3, posterior; C, intercoxal sclerite of leg 3, posterior; D, intercoxal sclerite and coxa of leg 4, posterior; E, leg 4, anterior. Scale bar in tum. 126 a S. KARAYTUG AND G.A. BOXSHALL Table 2 Body length (BL) and width (BW) measurements (in tm) of Paracyclops poppei in various localities. (N = number of specimens measured) Locality Sex BL (mean + SD) Germany °) 756 + 34.7 Gi 736 England Q 758 + 16.7 Russia (St-Petersburg) Q 828 + 43.6 of 655 + 21.4 United States (Virginia) Q 725 + 66 (of 601 + 33.8 United States (New Orleans) Q 741 +50 Gi 613 +8.5 Canada 2 819 + 109.6 Table 3. Caudal rami length (CL) and width (CW) measurements (in um) of Paracyclops poppei in various localities. L:W, ratio of length to width. (N = number of specimens measured) Locality Sex CL (mean + SD ) Germany 2 95 +6.7 England Q 102 +5.7 Russia (St-Petersburg) ie) 99 + 9.2 Mexico Q 95 +7 U.S.A. (New Orleans) fe) 60+7.9 U.S.A. (Virginia) Q 84+ 8.3 Japan (Lake Biwa) e) 104 + 14.7 VARIABILITY, FEMALES. The length and width of the caudal rami varied considerably especially inAmerican specimens (Figure 9A,B), and measurements are given in Table 3. One female from England and one from Lake Biwa (Figure 9E) had antennules with an incomplete suture line on the posterior margin subdividing segment 3. In some specimens from Virginia, U.S.A., the spinular rows on the intercoxal sclerite of leg 4 were unusually small (Figure 9F). Finally, in specimens from Lake Biwa (Japan), the dorsal spinular row of the caudal rami reached almost midway along the ramus in some specimens and the inner coxal spines of legs 2 and 4 were ornamented with longer setules (Figure 9C,D) than in material from elsewhere. However no other consistent variation was observed and these few slight differences do not justify the creation of a new taxon. DIFFERENTIAL DIAGNOSIS. P. poppei can easily be distinguished from its congeners by the continuous line of spinules on the dorsal surface of the caudal rami (Figure 8B) and by the possession of 2 large modified setae (setae A and C arrowed in Figure 13E) on the first segment of male antennule (Figure 13E,F).P. poppei also differs from P. fimbriatus, P. chiltoni and P. imminutus in the form of the spinular rows on either side of the anal operculum in the female (Figure 8B). REMARKS AND COMPARISONS P. poppei was originally described by Rehberg (1880) but subse- quently some authors considered that the differences between C. fimbriatus and P. poppei were not significant and therefore assigned this taxon firstly to Cyclops crassicornis O. F. Miiller, 1785 (Herrick, 1882) and then to C. fimbriatus Fischer, 1853 (Herrick, 1884; Schmeil, 1891; Byrnes, 1909; Marsh, 1892, 1910). Schmeil (1892) claimed that the differences could allow P. poppei to be recognized as a variety of P. fimbriatus and this opinion was shared by Van Douwe (1909). Gurney (1933) had doubts as to its status as subspe- cies and species, but its rank as subspecies was accepted by Lindberg (1958), Yeatman (1959) and Harding & Smith (1960). It was Kiefer (1929b) who first re-established P poppei as a valid species and in subsequent publications P. poppei gradually became accepted (Rylov, 1963; Dussart, 1969; Einsle, 1993; Ishida, 1993). The material identified by Sars (1927) as Platycyclops poppei Range BL (mean + SD) Range N 728-806 282 +8 276-283 4 282 1 736-778 288 + 11.7 274-309 4 786-913 PSST] a2 AS) 278-317 6 641-680 236 + 3 233-239 3) 640-849 261 + 19.4 230-291 10 538-615 PN EENIES 198-235 5 691-822 Pps ae il) 217-272 5 605-622 207 + 15.5 198-225 3 741-896 286 + 2.8 284-288 2 Range CW (mean + SD) Range N L:W 86-102 28+ 0.8 27-29 + 3.4 94-112 30+0.8 28-31 8 3.4 80-111 PSya= M9 27-31 6 3.4 90-100 29 + 0.7 28-29 2 3.2 48-72 2EIES 23-28 12 2.4 70-97 26i= 2: 24-28 11 322. 86-120 3144 25-33 4 3.4 from South Africa is not P. poppei nor can it be assigned to any other species of the genus. In fact, his material probably represents a new species. Sars assigned his specimens to P. poppei on the presence on the caudal rami of a single oblique row of small spinules across the dorsal surface, and he noted similarities in outward appearance to P affinis. Indeed, the dorsal spinular rows across the caudal rami are rather like P. affinis, however as Sars (1927) also stated, his species can easily be distinguished from P. affinis by its 8-segmented antennule. Sars’s species is also different from P. poppei as described herein in the structure of leg 5. DISTRIBUTION. P. poppei was considered to have a wide distribu- tion (Dussart & Defaye, 1985). Although its presence in Europe, North America and Japan has been confirmed, other records of P poppei, especially from Brazil and Paraguay (Lowndes, 1934), East Africa (Van Douwe, 1912), Tunisia (Dumontet al., 1979) and Hawaii (Sars, 1927) should be confirmed since there is insufficient descrip- tion provided for unequivocal identification. For detailed references concerning the distribution of P. poppei, see Karaytug (1998). Paracyclops oligarthrus (G. O. Sars, 1909) (Figures 14-20) Cyclops oligarthrus, Sars, 1909: Cunnington (1920) Platycyclops oligarthrus (Sars, 1909): Gurney (1928) ORIGINAL DESCRIPTION. Cyclops oligarthrus Sars, 1909: Proc. zool. Soc. Lond.: 31-77, pl. XXI. figs. 195-202. TYPE LOCALITY. Lake Tanganyika, Africa. MATERIAL EXAMINED. G. O. Sars, Lake Tanganyika 139 2, 5c co (Syntypes). BMNH 1909. 6. 24. 224-233. REDESCRIPTION OF ADULT FEMALE Body length (mean + SD) 555 + 32.6, range = 517-598, n=8. Body width 220 + 8, range = 206-233, n = 8. Prosome (Figure 14C) produced frontally, forming prominent rostral area. Fifth pedigerous somite with strong fringe of elongate setules at posterior margin PARACYCLOPS REVISION a \ i J Fan ° Yn by I . ar (- a a ‘ Va \\ j y) \ i LO = gp ROOT SS ow \ Se SSS eee Loe SSS R Se : LLLP Ie ye ee, Se ABF|5 SSS SSS ues | Wi iA hunny t : i iN IN j\ | | NN iN Fig. 12 FP. poppei. Adult male. A, detail of leg 6, anteroventral; B, leg 5, ventral; C, urosome, dorsal; D, urosome, ventral: E, body, dorsal; F, antenna, coxobasis and first endopodal segment, caudal. Scale bars in tm. 127 r 128 S. KARAYTUG AND G.A. BOXSHALL YR rap Ly WA) WL, LZ LI SK ts, S KL hag LL) Fig. 13 P. poppei. Adult male. Antennule. A, dorsal showing segmentation and with inset showing seta A; B, detail of terminal segments; C, ventral showing segmentation; D, anteroventral view of segments 14 and 15; E, anteroventral view of segments | to 12 with inset showing detail of seta C; F, anteroventral view of first segment with inset showing detail of seta B. Scale bars in um. PARACYCLOPS REVISION (Figure 15A). Genital double-somite, second and third abdominal somites lacking pits on dorsal and ventral surfaces (Figure 15A,B). Seminal receptacle divided into complex anterior and posterior lobes (Figure 15B). Caudal rami slightly divergent and 3.5 times longer than broad (Figure 15A,B). Posterolateral seta (IV) and inner terminal seta (V) with complex spinular ornamentation as figured (Figure 14C); terminal accessory seta (VI) strong and plumose. Antennule (Figure 16A) compact and 7-segmented; first (ances- tral segments I-V) and second (ancestral segments VI-XI) segments separated ventrally but incompletely separated dorsally. First and second segments here treated as distinct segments. Segment 3 with partial suture line (indicating boundary between ancestral segments XIII and XIV) and spiniform seta. Segment 5 with partial suture line (indicating boundary between ancestral segments XXI and XXII) and with characteristic short aesthetasc. Short aesthetasc located distally on anteroventral margin of segment 6. Setal formula 8, 12, 6, 5, 4 + aesthetasc, 2 + aesthetasc, 7 + aesthetasc. Coxobasis of antenna lacking spinular row near base of inner setae caudally (arrowed in Figure 16C); with complex ornamentation on caudal (Figure 16C) and frontal (Figure 16B) surfaces as figured. Terminal exopodal segments of legs 2—4 each with two semispinulose setae (arrowed in Figures. 17A; 18A,D). Praecoxa of leg 1 (Figure 17D) without spinular row at outer corner; basis with setiform spine on inner margin not extending beyond distal margin of second endopodal segment; intercoxal sclerite without spinular row on posterior surface (Figure 17E); terminal endopodal segment with 3 inner setae. Terminal endopodal spine of leg 2 (Figure 18A) strong, about as long as segment; coxa with complex ornamentation on posterior surface (Figure 18B). Intercoxal sclerite of leg 3 with spinules on anterior surface (Figure 18D) and with spinular row on posterior surface (Figure 18C); coxa with complex ornamentation on posterior surface (Figure 18C). Intercoxal sclerite of leg 4 with row of setules on anterior surface (Figure 17A) and with two spinular rows on posterior surface (Figure 17B); distal row well developed; inner coxal spine without cluster of setules posteriorly (Figure 17A); basis with long plumose outer angle seta; lacking setules along inner margin (Figure 17A); coxa with complex orna- mentation on posterior surface (Figure 17B); exopodal spines with dense spinules along margins (Figure 17A). Spine and seta formula as follows: Coxa Basis Exopod Endopod Leg 1 0-1 1-I 1-1;I-1;111,5 0-1;0-1;1,1,3 Leg 2 0-1 1-0 I-1;1-1;111,1,5 0-1;0-2;1,1,4 Leg 3 0-1 1-0 I-1;J-1;I01,1,5 0-1;0-2;1,1,4 Leg 4 0-1 1-0 I-1;1-1;1,5 0-1;0—2;1,11,2 Leg 5 (Figure 15C) comprising single free segment, armed with | short well developed outer spinulose seta, | strong inner spine and 1 strong plumose seta (slightly longer than inner spine) centrally. DESCRIPTION OF ADULT MALE Body length (mean + SD) 444 + 14.8, range = 433-454, n = 2. Body width 179 + 2.8, range = 177-181, n= 2. Urosomal somites without surface ornamentation (Figure 19A,B). Caudal rami short about 2 times longer than broad. Antennule digeniculate (Figure 20A,B), indistinctly 15-seg- mented. Segment | armed with 8 setae (Figure 20E,F); seta A large and modified by ornamentation of strong spinules in proximal and mid sections, tapering to fine point distally; aesthetasc absent. Segment 11 bearing strongly curved seta ornamented along convex surface with double row of strong denticles, plus 1 plumose seta (Figure 20E). Segmental fusion pattern as follows I-V, VI-VII, VIII, 129 IX, X, XI, XII, XIII, XIV, XV, XVI, XVIU, XVI-XX, XXI-XXII, XXIV-XXVIII. One seta on terminal exopodal segment of leg 1 semispinulose (Figure 17C). Intercoxal sclerite of leg 4 with stronger terminal spinular row than in female (Figure 19D). Fifth leg with strong inner spine and reduced outer and middle setae (Figure 19E). Sixth leg (Figure 19A,C) armed with | strongly developed inner spine sur- rounded by spinules at base, and 2 outer setae, outermost seta shorter than middle seta. VARIABILITY. Inner margin of basis of leg 4 lacks setules in most females examined and in one of the two males but was ornamented with setules in some females and the other male. DIFFERENTIAL DIAGNOSIS. P. oligarthrus can be distinguished from other Paracyclops species by the structure of the fifth leg in both sexes (Figures 15C; 19A,E), by the structure of the seminal receptacle (Figure 15B), by the 7-segmented antennule (Figure 16A and see remarks), by the absence of the proximal inner seta on the terminal endopodal segment of leg | (Figure 17D), by carrying 2 semispinulose setae on the terminal exopodal segment of legs 2-4 (Figures 18A,D; 17A) and by the sixth leg of the male being fully incorporated into the genital somite (Figure 19A). REMARKS AND COMPARISON P. oligarthrus is unique in the loss of the proximal inner seta on the terminal endopodal segment of leg 1 (Figure 17D). This segment carries 4 inner setae in all other species. P. oligarthrus also has 2 semispinulose setae on the terminal exopodal segment of legs 24 (arrowed in Figures 17A; 18A,D). The male sixth leg is unusual in the relatively large size of the inner spine and in being fully incorpo- rated into the genital somite (Figure 19A). Sars’s (1909) interpretation of the antennule as 6-segmented is incorrect. He appears to have overlooked the partial division of the proximal segments. DISTRIBUTION. Africa. P. oligarthrus is endemic to Lake Tanganyika, Paracyclops canadensis (Willey, 1934) (Figures 21—25) Cyclops affinis var. canadensis Willey, 1934 Paracyclops affinis (Sars, 1863) sensu Smith & Fernando (1977, 1978) ORIGINAL DESCRIPTION. Cyclops affinis var. canadensis Willey, 1934: Trans. R. Can. Inst. 20 (1): 77-98. TYPE LOCALITY. original paper) Canada, Quebec (no other detail is given in the TYPE MATERIAL. The type material of Willey (1934) could not be located. It is not deposited in the CMNC or the USNM. MATERIAL EXAMINED The redescription of P. canadensis is based on 29 2 which were obtained from Canadian Museum of Nature. Catalogue number: CMNC 1996-0019. Locality: Canada, Ontario, Parry Sound Dis- trict, 40 km N of Parry Sound on Hwy 69; collected by C. H. Fernando on 7 July 1972. — U.S.A, West Virginia; 82 2 collected on 23 May 1995 in Big Run Bog in the Monongahela National Forest by Robert Hamil- ton, Tucker County, 39°07° N, 79°35” W. USNM Acc. 417235. — CANADA, Jack Lake, Nova Scotia 3 9 2 dissected and mounted on | slide, 2 2 9 undissected and mounted on | slide, 1 9 dissected and mounted on 1 slide, 22 9 and 1 C'undissected and mounted S. KARAYTUG AND G.A. BOXSHALL 130 ALLE IP I PFPA PO Coco zz SSE Mee Gz 7h) yy Nf ¥ : SF Pe Z Fy ea : oie: Fig. 14 P. oligarthrus. Adult female. A, maxillulary palp; B, maxillule; C, body with inset showing the detail of setal elements IV and V of caudal rami, dorsal; D, maxilliped; E, maxilla; F, mandible; G, labrum. Scale bars in Um. | ronan | fii iN PSNNG Jip 38 : “a SS UU, \ Sr A NO WH Vis \ by po hy, S. KARAYTUG AND G.A. BOXSHALL 132 —— SS SSS SN SS oO SS a 4 Var ST amit Wai G 25 BC Fig. 16 P. oligarthrus. Adult female. A, antennule; B, antenna, frontal; C, antenna, coxobasis, caudal. Scale bars in um. 133 PARACYCLOPS REVISION yg, 25 \ A 8 f av s =) ey a Z 3 3 Z aS BSS > | Mi Abhay = g 4 ‘4 = WS ym Ye Le Vi fs lly fi Fig. 17 P. oligarthrus. Adult female. A, leg 4 with inset showing the inner coxal spine, anterior; B, intercoxal sclerite, coxa and basis of leg 4, posterior; C, adult male, terminal endopodal segment of leg 1, posterior; D, female, leg 1, anterior; E, intercoxal sclerite and coxa of leg 1, posterior; Scale bars in um. 134 S. KARAYTUG AND G.A. BOXSHALL LE 2 7 = f Z wa il A ‘nF Tp AV / ity! ays C AAKA 44h 6b : AN Naw NL N NL an abbd Dy f \ INS \ oe <> QE ——s S = & fe ° WGK ¥ oS Senes Rs ea \\ S a 50 SS TN wm NA Fig. 18 P. oligarthrus. Adult female. A, leg 2, anterior; B, intercoxal sclerite and coxa of leg 2, posterior; C, intercoxal sclerite and coxa of leg 3, posterior; D, leg 3, anterior. Scale bar in um. Sa , eat /; i / MW , Vi ai i : S < Oo z ro) E SS x es SS & a \ SE Wes eS Me NV S Se SE His \\ = Q\ Na = \ of sam IS Fig. 20 P. oligarthrus. Adult male. A, antennule, ventral showing segmentation; B, dorsal showing segmentation; C, body, dorsal; D, antennule, terminal segments showing setation; E, antennule showing setation, anteroventral; F, first segment showing setation, anteroventral;. Scale bars in um. 136 PARACYCLOPS REVISION on | slide, 1 h'dissected and mounted on | slide, 20” C'undissected and mounted on 2 slides. Dr H. Yeatman collection. These specimens are deposited in USNM. REDESCRIPTION OF ADULT FEMALE Body length and width not including caudal setae given in Table 4. Genital double-somite, second and third abdominal somites with more dorsal surface ridges extending round sides onto ventral surface than P. affinis (Figure 21A,B). Seminal receptacle divided into broad anterior and posterior lobes, anterior lobe slightly nar- rower than posterior (Figure 21B). Anal cleft with irregularly arranged spinules (Figure 21A,D) and with spinular row extending either side (arrowed in Figure 21D). Caudal rami short about 2 times longer than broad (Figure 21A,B); terminal accessory seta (VI) longer than posterolateral seta (III) (Figure 21A). Antennule 11-segmented. Segment 6 with spiniform seta. Setal formula 8, 4, 2, 6,4, 2,2, 3,4 + aesthetasc, 2 + aesthetasc, 7 + aesth- etasc. Coxobasis of antenna (Figure 22C,D) with complex ornamenta- tion on caudal and frontal surfaces as figured; lacking spinular row near base of inner setae on caudal surface (arrowed in Figure 22D). Basis of maxilliped with one spinular row on anterior surface and 2 groups of spinules on posterior surface (Figure 22B). First endopodal segment with | tiny spinule on anterior surface. Coxae of legs 2-4 with 2 spinular rows on anterior surface (arrowed in Figures 23D, 24A,D) and with inner spine bearing 2 or 3 large spinules posterolaterally (arrowed in Figure 24A); basis without spinular row on anterior surface near inner margin. Coxa of leg 3 with two mid-distal spinules on posterior surface. Intercoxal sclerite of leg 4 with well developed spinular row (stronger than that of P. affinis) along free margin and with spinular rows on anterior and posterior surfaces (Figure 24C,D) Spine and seta formula as follows: Coxa Basis Exopod Endopod Leg | 0-1 1-I I-1;I-1;01,5 O-1;0-1;1,1,4 Leg 2 0-1 1-0 I-1;1-1;01,1,5 O-1;0-1;1,1,4 Leg 3 0-I 1-0 I-1;J-1;01,1,5 O-1;0-2;1,1,4 Leg 4 0-I 1-0 L-1;I-1;01,5 0-1;0-1;1,01,2 Leg 5 with long inner spine about 4 times as long as segment; outer spinulose seta simple, about half as long as inner spine and without spinules at base (arrowed in Figure 21C). DESCRIPTION OF ADULT MALE Differing from adult female as follows: Genital somite with 5 or 6 irregular dorsal surface ridges and 5 incomplete ventral surface ridges; first to third free abdominal somites each with 3 complete dorsal and ventral surface ridges (best seen in Figure 25A,B). Antennule digeniculate (Figure 25C,D), indistinctly 16-seg- mented. Segment | armed with 8 setae plus an aesthetasc. Segment 10 (= ancestral segment XV) produced on one side into sheath enclosing segment 11 ventrally: armed with 2 setae, one of which pear-shaped and constricted at end in P. affinis but simple in P canadensis (arrowed in Figure 25D). VARIABILITY, FEMALES. Arrangements of spinules in the anal cleft 137 may vary (cf. Figures 21A and 21D). DIFFERENTIAL DIAGNOSIS. P canadensis is distinguished from other Paracyclops species by the combination of 11-segmented antennule, the surface ridges on the urosomal somites, the spinular ornamentation on the anal cleft and by the possession of only 1 seta on the second endopodal segment of leg 4. P. canadensis and P. affinis are very closely related but P. cana- densis can easily be differentiated from P. affinis by the possession of 4 spines on the terminal exopodal segment of leg 3 (Figure 24A), the lack of spinules at the base of the outer seta of leg 5 (Figure 21C); by the presence of more surface ridges on the genital and following 2 free somites (Figure 21A,B); by the extent of the spinular row either side of the anal cleft (Figure 21A,D); by the structure of inner coxal spines of legs 2 to 4 (Figures 23D; 24A,D) and by the absence of spinular rows near the inner margin of the basis of legs 2 to 4 (Figures 23D; 24A,D) REMARKS P. canadensis has been reported from North America under the name P. affinis by Yeatman (1959), Pennak (1963) and Smith & Fernando (1977, 1978). However, comparison between European and North American specimens of P. affinis led to the recognition here of P. canadensis as valid species in North America. P. canadensis was first recorded from North America in 1934 by Willey as a variety of P. affinis (Willey, 1934).According to the rules of zoological nomen- clature this taxon when raised to species rank must take Willey’s original variety name, becoming P. canadensis (Willey, 1934). DISTRIBUTION. P. canadensis occurs in the Eastern parts of Canada and United States. Paracyclops dilatatus Lindberg, 1952 (Figures 26-27) Platycyclops dilatatus Sars, 1927a [nomen nudum] Paracyclops dilatatus tvanegai Monchenko, 1977 ORIGINAL DESCRIPTION. Paracyclops dilatatus Lindberg, 1952: Bull. Soc. zool. France, 77, 1: p.80, fig. 1 TYPE LOCALITY. Caspian Sea. MATERIALEXAMINED. Syntypes: Zoologisk Museum, Oslo; 3 slides F6236, F6237a and F6237b contain parts of | dissected &. One tube F6237c contains female fragments. F6237a, 6237b and 6237c were separated from one original slide by Dr. P. Frenzel in 1979. The type specimens of P. dilatatus Lindberg, 1952 were obtained on loan from the Zoologisk Museum, Oslo. Unfortunately the 3 slides are not very informative and the available fragments of just one female were insufficient to redescribe P. dilatatus in detail. REDESCRIPTION OF ADULT FEMALE Body length (Figure 26H) not including caudal setae is 840 um (given by Lindberg, 1952 as approximately 770um to 810ym). Genital double-somite, second and third abdominal somites without surface pits on dorsal and ventral surfaces (Figure 26H). Anal somite with spinular row ventrally extending dorsally. Caudal rami (Figure 26K) short, about 2 times longer than broad. Table 4 Body length (BL) and width (BW) measurements (in um) of P. canadensis (N = number of specimens measured). Locality Sex BL(mean + SD) Canada, Ontario ie) 684 + 45.2 U.S.A. (West Virginia) Q 713 + 47.6 Range BL(mean + SD) Range N 652-716 A3\/ =) 3)-9) 254-259 2, 642-783 Pa) get.) 242-264 8 S. KARAYTUG AND G.A. BOXSHALL 138 ——— ntral; C, leg 5, ventral; D, anal somite, dorsal. Scale bars in um. is. Adult female. A, urosome, dorsal; B, urosome, ve adens: Fig. 21 P can 139 PARACYCLOPS REVISION dal. Scale bar in um. frontal; D, antenna, cau obasis, is. Adult female. A, maxilla; B, maxilliped; C, antenna, cox adens Fig. 22 P can S. KARAYTUG AND G.A. BOXSHALL 140 \ WV. | ‘ad — = ~ 4 v7 AS “ i ( f 4 A ane Z ats “ Fig. 23 P. canadensis. Adult female. A, intercoxal sclerite and coxa of leg 1, posterior; B, leg 1, anterior; C, intercoxal sclerite and coxa leg 2, posterior; D, leg 2, anterior. Scale bar in um. 141 _ PARACYCLOPS REVISION 9 wd wf ‘/ HED Pal id Maelpinp ee h i TE i ; 4 yy rt ene a OP anges ee > A Vi Ons WL jm ¢ 144, @® ] / pena Z it ZAI ee 3 aay e SIZ ! Z fe (yy! ZA / NY es econo er zl ST aS SS as 7 NY : = NS Se 5 100 — > NY AQ \ J p45), > x \. lf, fr “thin, jini \\ < i N/A IA } My | AY Xt << Ad a \ \ Fig. 46 P. waiariki. Adult female. A, leg 1, anterior; B, leg 2, anterior; C, intercoxal sclerite and coxa of leg 4, posterior; D, leg 4, anterior; E, leg 3, anterior; F, intercoxal sclerite of leg 3, posterior. Scale bar in um. 167 S. KARAYTUG AND G.A. BOXSHALL 50 AC C mn SF oy ® | ; | R s) " vyV Sen. cog ‘ d outer terminal set A (a) ntral; D, caudal rami, detail of the inner and outer terminal setae, i =z = ome, ve ome, dorsal; B, body, dorsal; C, uros rsal; E, legs 5 and 6, lateral. Scale bars in lum. Fig. 47 P. waiariki. Adult male. A, uros do 168 PARACYCLOPS REVISION Fig. 48 P. waiariki. Adult male. Antennule. A, dorsal showing segmentation; B, detail of segm anteroventral showing setation; D, detail of terminal segments showing setation; E, ventral s ents 12 to 15 showing setation, anterov howing segmentation. Scale bar in um. entral; C, 169 170 Outer setae of fifth leg plumose and about as long as inner spine (Figures 47C,E; 45G); sixth leg (Figures 47C,E; 45F) armed with 1 inner spine (much shorter than second urosomal somite) and 2 outer plumose setae. DIFFERENTIAL DIAGNOSIS. P. waiariki can be differentiated from other Paracyclops species by the combination of the following characters; its 12-segmented antennule, the absence of an inner seta on the first exopodal segment of leg 4, the structure of the seminal receptacle, the produced base for the outer seta of leg 5 in the female, the absence of the proximal spine on outer margin of terminal exopodal segment of leg 3, the spinular ornamentation on the frontal surface of coxobasis of the antenna, its wide anal operculum and the ornamentation of the fine spinules along the dorsal and ventral surfaces of caudal rami. P. waiariki, P. smileyi and P. eucyclopoides are closely related: however, P. wairaiki differs from P. smileyi and P. eucyclopoides by having 3 spines on the terminal segment of leg 3 rather than 4. In addition, P. waiariki can also be separated from P. eucyclopoides by the structure of seminal receptacle. P. waiariki also differs from P. smileyi in the length of outer seta of leg 5, in having a produced base for the outer seta of leg 5 and by the much shorter caudal rami. REMARKS Kiefer (1969) originally described Paracyclops timmsi from Aus- tralia. This species resembles P. waiariki in most respects, including the structure of leg 5, the number of segments on the female antennule and in spine and seta formula of swimming legs. As far as Kiefer’s description is concerned, the differences between the spe- cies are the structure of the seminal receptacle and the body shape. Lewis (1974) does not mention P. timmsi in her original description of P. waiariki which suggests that she was unaware of Kiefer’s work on P. timmsi. It is possible that P. waiariki may be a synonym of P. timmsi but P. timmsi needs to be redescribed to modern standards as it is clear that minor details of spinulation can represent significant differences at species level. DISTRIBUTION. Only known from its type locality in New Zealand. Paracyclops pilosus Dussart, 1984 (Figures 49-50) ORIGINAL DESCRIPTION. Paracyclops pilosus Dussart, 1984: Hydrobiologia, 113: p. 56., fig. 15. TYPE LOCALITY. Venezuela; Orinoco River. MATERIAL EXAMINED. This species was originally described from single male and female. Holotype (dissected on | slide, MNHN Cp 659) and Allotype (dissected between prosome and urosome on | slide, MNHN Cp 669) were obtained on loan from Museum Na- tional d’Histoire Naturelle in Paris. Due to the positioning of the female appendages on the slide and to the earlier partial dissection of the male it was not possible to describe every detail of this species but several characters could be clarified. REDESCRIPTION OF ADULT FEMALE Genital double-somite, second and third abdominal somites (Figure 49B) ornamented with fine pits on ventral surface (the dorsal surface could not be observed). Seminal receptacle divided into broad anterior and posterior lobes. Caudal rami given as 2.9 times longer than wide in original description (Dussart, 1984). Terminal acces- sory seta (VI) as long as posterolateral seta (III); posterolateral seta (IIL) unilaterally plumose, with spinules along dorsal surface; outer terminal seta (I[V) and inner terminal seta (V) well developed and heterogeneously ornamented (Figure 49B). S. KARAYTUG AND G.A. BOXSHALL Antennule 8-segmented (Figure 49D); first and second segments incompletely separated; third segment with two partial suture lines on dorsal surface and with spiniform seta; fifth segment with charac- teristic short aesthetasc; another aesthetasc located distally on anteroventral margin on segment 7 about 2.5 times longer than terminal segment. Setal formula 8, 12, 6, 5, 2 + aesthetasc, 2, 2 + aesthetasc, 7 + aesthetasc. Spinular ornamentation on coxobasis of antenna impossible to observe. Spine and seta formula of swimming legs as follows: Coxa Basis Exopod Endopod Leg 1 0-1 1-I J—-1;I-1;10,5 0-1;0-2:1,1,4 Leg 2 0-1 1-0 151-1500 15 0-1;0-2;1,1,4 Leg 3 0-I 1-0 I-11: 005 0-1;0—2;1,1,3 Leg 4 0-I 1-0 I-1;/-1;11,1,4 O—1;0—2;1,11,2 Leg 5 (Figure 49C) comprising single free segment, armed with | long (almost 1.5 times longer than inner spine) well developed outer spinulose seta, 1 serrate strong inner spine, plumose seta in centre slightly longer than inner spine. DESCRIPTION OF ADULT MALE Genital and 3 free abdominal somites (Figure 49A) without surface pits on ventral surface. Caudal rami short, about 2.5 times longer than broad. First segment of antennule with modified seta. Outer seta of fifth leg plumose, as long as inner spine (Figure 49A); sixth leg (Figure 49A) armed with | well-developed inner spine, slightly longer than second urosomal somite. DIFFERENTIAL DIAGNOSIS. P. pilosus is easily distinguishable by the incomplete separation of the first and second segments of the female antennule (Figure 49D), by the presence of 3 setae on the terminal endopodal segment of leg 3 (Figure 5OC), by the presence of 4 setae on the terminal exopodal segment of leg 4 (Figure SOD). It can also be separated from closely related species by the combina- tion of the presence of 2 setae on the second endopodal segment of leg | (Figure 50A), and the presence of cuticular depressions on the ventral surface of the caudal rami. REMARKS The presence of four inner setae on the terminal exopodal segment of leg 4, and of three inner setae on the terminal endopodal segment of leg 3 is remarkable. DISTRIBUTION. Venezuela: Orinoco River (type locality) (Dussart, 1984). Dussart (1984) also mentioned that this species was found in the littoral zone of flowing waters of the Orinoco at Barrancas at Ciudad Bolivar. Paracyclops carectum Reid, 1987 (Figures 51-53) ORIGINAL DESCRIPTION. Paracyclops carectum Reid, 1987: Hydrobiologia, 153, p. 124. (Figs. 1-12). TYPE LOCALITY. Vereda Grande Pond, Aguas Emendadas Bio- logical Reserve, Federal District, Brazil. MATERIAL EXAMINED. 4 Paratype 9 9(USNM Cat. No: 232176) from Brazil, shore of Vereda Grande pond 15 °32'30"S; 047 °34'S7"W: collected by Dr. J. W. Reid (May 1982-1986), 1 Qdissected; 1 Paratypeo (USNM Cat. No: 232175) from wet campo marsh, Fazenda Agua Limpa, Distrito Federal, Brazil; collected by Dr. J. W. Reid (Apr 1982).20° 0 (USNM 242425) from Brazil; Goias; marsh of Corrego Pocoes; collected by Dr. J. W. Reid (December 1983), 171 PARACYCLOPS REVISION var udal setae (IV) and (V), ventral; C, leg 5, ventral; ome with inset showing the ca ome, ventral; B, adult female, uros s. A, adult male, uros ntennule. Scale bars in um. Fig. 49 P pilosu D, a 172 S. KARAYTUG AND G.A. BOXSHALL 7 100 Ys ) GW rg K a if c = YS ZF 7. Y | h be r ‘¢ a Wf ND on if XG NS VA \ = Na\\y NS a i haa “ite sael inne AN WF ZS ee Ji = iN 4. & JN | A Poa SS / ; a Yj | A al At 7 it V, 4 ah Fig. 50 P. pilosus. Adult female. A, leg 1, anterior; B, leg 2, posterior; C, leg 3, anterior; D, leg 4, anterior. Scale bar in um. PARACYCLOPS REVISION l1O'dissected; 1 9(USNM 242423) from Brazil, Federal District; Brasilia Lagoada Peninsula Norte, collected by Dr. J. W. Reid; 1 9(USNM 242424) from Brazil; Federal District; Brasilia, Lagoa Jaburu, collected by Dr. J. W. Reid (Aug 1982). REDESCRIPTION OF ADULT FEMALE Body length measured within same range as given in original description (Reid, 1987) as 600-800 um, mean = 650 um. Genital double-somite ornamented with fine pits on dorsal surface as figured (Figure 51A,B). Seminal receptacle divided into broad anterior and posterior lobes. Caudal rami (Figure 51A,B) parallel, 3.2 times longer than broad; with groups of spinules and hairs along inner margin (Figure 51A,D); outer terminal seta (IV) and inner terminal seta (V) well developed and heterogeneously ornamented (Figure S1A). Antennule 8-segmented; Setal formula 8, 12, 6,5, 2 + aesthetasc, 2, 2 + aesthetasc, 7 + aesthetasc. Coxobasis of antenna (Figure 52G,H) with complex ornamentation on caudal and frontal sides as figured; without spinular row on caudal surface near base of two inner setae (arrowed in Figure 52G). Basis of leg 1 (Figure 52C) with setiform inner spine reaching midway along terminal endopodal segment; intercoxal sclerite ornamented with 2 spinular rows on posterior surface (Figure 52D). Intercoxal sclerite of leg 2 ornamented with spinular rows on anterior and posterior surfaces (Figure 52J,K). Intercoxal sclerite of leg 3 without spinular row on anterior surface and with 2 spinular rows on posterior surface (Figure 52E,F); coxa with complex orna- mentation on posterior surface as figured (Figure 52E). Intercoxal sclerite of leg 4 (Figure 52A,B) with 2 spinular rows on posterior surface; inner coxal spine with proximal group of setules mainly originating posteriorly (Figure 52B). Spine and seta formula as follows: Coxa Basis Exopod Endopod Leg | 0-1 1-I I-1;1-1;111,5 O-1;0-1;1,1,4 Leg 2 0-I 1-0 151-1500 15 0-1;0—2;1,1,4 Leg 3 0-I 1-0 I-1;I-1;11,1,5 0-1;0-2;1,1,4 Leg 4 0-I 1-0 I-1;I-1;IL,1L,5 0-1;0—2;1,11,2 Leg 5 (Figure 51C) comprising single free segment, armed with 1 long, well developed outer spinulose seta, | serrate, strong inner spine, and | plumose seta centrally. DESCRIPTION OF ADULT MALE Body length measured within same range as original description (Reid, 1987), length of allotype 580 um, lengths of paratypes 550 and 600 um. Genital, third, fourth and fifth urosomal somites (Figure 53A,B) ornamented with cuticular pits on dorsal surface. Caudal rami shorter than female, about 2.1 times longer than broad; with complex ornamentation along inner margin as figured (Figure 53A,B,D). First segment of antennule armed with 8 setae plus aesthetasc; 1 seta (A) large and modified by ornamentation of strong spinules in proximal and mid sections, tapering to fine point distally. All other appendages as in female except; outer setae of fifth leg plumose (Figure 53C); sixth leg (Figure 53A,C) armed with | inner spine as long as second urosomal somite, surrounded by spinules at base; 2 outer setae plumose. VARIABILITY. The outer margin of inner spine of the female leg 5 may be ornamented with a variable number of spinules (Figure 51C). DIFFERENTIAL DIAGNOSIS. P. carectum is unique within the genus in carrying ornamentation of spinules along the inner margin of the caudal rami (Figure 51A,D). 173 DISTRIBUTION. Brazil: Vereda Grande Pond, Aguas Emendadas Biological Reserve, Federal District, 15°32'30"S, 47°34'57"W; Wet campo Marsh, Fazenda Agua Limpa, Distrito Federal; Goias, marsh of Corrego Pocoes; Federal district, Brasilia, Lagoada peninsula norte; Federal district, Brasilia, Lagoa Jaburu (Reid, 1987). Paracyclops novenarius Reid, 1987 (Figures 54-57) ORIGINAL DESCRIPTION. Paracyclops novenarius Reid, 1987: Proc. Biol. Soc. Wash. 100(2), p. 262, figs. 1-20. TYPE LOCALITY. Colombia, Valle, Buenaventura. MATERIAL EXAMINED. Holotype: 1 9(USNM 231096) collected by Dr. Marco F. Suarez (5 Sept. 1985). Paratypes: 122 Q and 9S O(USNM 231099). 10°, 12, 4 copepodids (USNM 231100); all paratypes collected from the type locality. REDESCRIPTION OF ADULT FEMALE Body length measured within same range as in original description (given by Reid (1987) as 570-880 um, mean = 630). Genital double- somite, second and third abdominal somites ornamented with very fine pits on dorsal and ventral surfaces as figured (Figure 54A,B). Seminal receptacle divided into narrow anterior and broad posterior lobes as figured (Figure 54B). Caudal rami (Figure 54A,B) with fine cuticular depressions on ventral surface. Terminal accessory seta plumose (VI) and 1.5 times longer than posterolateral seta (IID; posterolateral seta (III) strong and unilaterally plumose, with spinules along dorsal surface; setae 1V and V well developed and heterogene- ously ornamented (Figure 54B). Antennule 8-segmented (Figure 55A). Segment 2 and 3 with complex partial suture lines. Segment 2 may be incompletely sepa- rated or with complete separation, best seen in Figure 55A,C,E,F. Segment 3 with partial suture line and spiniform seta. Setal formula 8, 12, 6, 5, 2 + aesthetasc, 2, 2 + aesthetasc, 7 + aesthetasc. Coxobasis of antenna (Figure 55B,D) without spinular row on caudal surface near bases of inner setae (arrowed in Figure 55D). Coxa of leg | lacking spinular row on posterior surface near intercoxal sclerite (Figure 56G). Intercoxal sclerite of leg 2 ornamented with spinular row on anterior surface (Figure 56B); without spinular row on posterior surface (Figure 56A). First and second exopodal segments lacking spinular row on posterior surface (Figure 56B). Intercoxal sclerite of leg 3 with spinular row on anterior surface (Figure 56E) and 2 spinular rows on posterior surface (Figure 56F). Intercoxal sclerite of leg 4 without spinular row on anterior surface (Figure 56D) and with 2 spinular rows on posterior surface (Figure 56C); coxa without mid-distal spinular row on posterior surface (arrowed in Figure 56C). Spine and seta formula as follows: Coxa Basis Exopod Endopod Leg 1 0-1 1-I I—-1;I-1;111,5 0-1;0-1;1,L4 Leg 2 0-1 1-0 I-1;1-1;11,1,5 0-1;0-2;1,1,4 Leg 3 0-I 1-0 -1;-1;00,15 0-1;0—2;1,1,4 Leg 4 0-I 1-0 I-1;I-1;IL1L5 O=1;0—251, 11,2 Leg 5 (Figure 54C) comprising single free segment, armed with | strong outer spinulose seta slightly longer than inner spine, | serrate- like strong inner spine with 3 spinules at base, | plumose seta in middle. DESCRIPTION OF ADULT MALE Body length measured within same range as in original description 174 S. KARAYTUG AND G.A. BOXSHALL =, MS /: A ma: WF SAV) : SSS 4 SS =<) rea 2 aye SS 2s 22 YY LL SSS SS = Ze ee ee ) 7) P. carectum. Adult female. A, urosome, dorsal; B, uros ;D,a ite and caudal rami, dorsal. Scale bars in um. Fig. 51 ome, ventral; C, leg 5 with inset showing variant pattern of spinulation on inner spine, ventral; D, anal som PARACYCLOPS REVISION VUPTPPHFE CE UF" ees eves emcee 0! ‘Wd . ) [ na A (Roos nlf my Myson IK 15 ABCDEFJK | Fig.52 P carectum. Adult female. A, leg 4, anterior; B, intercoxal sclerite and coxa of leg 4, posterior; C, leg 1, anterior; D, intercoxal sclerite and coxa ! of leg 1, posterior; E, intercoxal sclerite and coxa of leg 3, posterior; F, intercoxal sclerite of leg 3, anterior; G, antenna, coxobasis, caudal; H, antenna, coxobasis, frontal; J, intercoxal sclerite of leg 2, anterior; K, intercoxal sclerite and coxa of leg 2, posterior. Scale bars in um. 175 : S 4 ) Q E 1o) e 176 as vin" | [ al; D, anal somite and caudal rami, ome, dorsal; C, detail of leg 5 and leg 6, anteroventr: tral; B, uros ome, ven Fig. 53 P. carect dors: um. Adult male. A, uros' al. Scale bars in um. PARACYCLOPS REVISION given by Reid (1987) as 540-640 um, mean = 600). Genital, third, fourth and fifth urosomal somites each ornamented with cuticular pits on dorsal surface extending to ventral surface on third, fourth and fifth somites (Figure 57A,B). First segment of antennule armed with 8 setae plus an aesthetasc; one seta (A) large and modified by ornamentation of strong spinules in proximal and mid sections, tapering to fine point distally. All other appendages as in female except; one seta on terminal endopodal segment of leg 1 spinulose (Figure 56J). Outer seta of fifth leg plumose and less developed (Figure 57C). Sixth leg (Figure 57D) armed with | inner spine, shorter than second urosomal somite, bearing spinules at base; middle seta spiniform, short and stout; outer seta plumose. DIFFERENTIAL DIAGNOSIS. P. novenarius can be differentiated from other Paracyclops species by the combination of the following characters: — the dorsal subdivision of second segment of female antennule, observed in holotype and in one paratype (Figure 55C). Remain- ing specimens with second segment divided into 2 segments as in Figure 55E, F (partial suture line on second segment indicating boundary between ancestral segments VI—X and XI). — the structure of the seminal receptacle (Figure 54B), the absence of the mid-distal spinular row on the posterior surface of the coxa of leg 4 (arrowed in Figure 56C), the spinular pattern on the caudal surface of the coxobasis of antenna (Figure 55D), and the absence of spinular rows on the posterior surfaces of the first and second endopodal segments of leg 2 (Figure 56B). Distribution: Brazil: Vereda Grande Pond, Aguas Emendadas Bio- logical Reserve, Federal District, 15°32'30"S, 47°34'S7"W; Wet campo Marsh, Fazenda Agua Limpa, Distrito Federal; Goias, marsh of Corrego Pocoes; Federal district, Brasilia, Lagoada peninsula norte; Federal district, Brasilia, Lagoa Jaburu (Reid, 1987). Paracyclops smileyi Strayer, 1988 (Figures 58-60) ORIGINAL DESCRIPTION. logia 4 (3): 279-291. Paracyclops smileyi Strayer, 1988: Stygo- TYPELOCALITY. Type specimens were collected from the hyporheic zone of Coxing Kill, Town of Gardiner, Ulster County, New York, U.S.A, 17 December 1985 (Strayer, 1988). MATERIAL EXAMINED. Holotype (USNM Cat. No: 235368, one slide) and Paratype (USNM Cat. No: 235369, one slide) females were obtained on loan from United States National Museum of Natural History. REDESCRIPTION OF ADULT FEMALE Due to the positioning of the dissected appendages on the slides and to the poor condition of the slides it was not possible for this species | to be redescribed in detail. | Genital double-somite, second and third abdominal somites (Fig- | ure 58C) without ornamentation of surface pits dorsally; posterior margins of abdominal somites inconspicuously serrated dorsally. Caudal rami short (Figure 58C,E), length given as 2.5 times longer than broad by Strayer (1988). _ Antennule 12-segmented (Figure 59A); segment 6 with spiniform | seta (arrowed in Figure 59A); segment 9 with short aesthetasc | (arrowed in Figure 59A); apical segment with aesthetasc fused to | adjacent seta at base, and another aesthetasc located distally on | anteroventral margin of segment 11. Setal formula 8, 4, 2, 6, 4, 2, 2, 3, 2 + aesthetasc, 2, 2 + aesthetasc, 7 + aesthetasc. Coxobasis of Wa antenna (Figure 59D) with complex ornamentation on caudal and frontal surfaces as figured and without spinular row near base of 2 inner setae on caudal surface. Spine and seta formula of swimming legs (Figures 60A—E) as follows: Coxa Basis Exopod Endopod Leg 1 0-1 2-1 I-1;1-1;11,5 0-1;0-2;1,1,4 Leg 2 0-I 1-0 I-1;I-1;1,1,5 0-1;0-1(?);1,1.4 Leg 3 0-1 1-0 I-1;1-1;01,1,5 0-1;0-2;1,1,4 Leg 4 0-1 1-0 I-1;I-1;IL1,5 O-1;0—2;1, 11,2 Leg 5 (Figure 58B) comprising single free segment, armed with 2 outer setae about 3—4 times longer than inner spine, | strong inner spine. ADULT MALE. Unknown DIFFERENTIAL DIAGNOSIS. P. smileyi differs from other Paracyclops species except P. waiariki Lewis, 1974 and P. eucyclopoides Kiefer, 1929 in having a 12-segmented antennule in the female. P. smileyi differs from P. waiariki and P. eucyclopoides in the structure of leg 5 and in having shorter caudal rami (2.5 times longer than broad). P. smileyi can also be differentiated from P. waiariki by the presence of 4 spines on the terminal exopodal segment of leg 3. REMARKS There appears to be only a single inner seta on the second endopodal segment of leg 2 in the available type material. It is probable that the proximal seta (Figure 60C) is broken off in the types but it was difficult to observe any scar indicating the position of such a missing seta because of the poor condition of the slide. The presence or absence of this seta should be confirmed by examination of new material in the future. Similarly setal elements are missing from mouthparts such as the maxillule and maxilla. DISTRIBUTION. Known only from its type locality. Paracyclops reidae sp. nov. (Figures 61-64) TYPE LOCALITY. Pools in the leaf axils of a terrestrial bromeliad, El Tucuche, Trinidad, W.I.; 24 July 1994, coll. R. Martinez and M. Morton. TYPE MATERIAL. The type material (11 specimens) is stored in the collection of Smithsonian Institution, Washington D.C., USA, Reg. No: USNM 264163. | female and | male paratypes are stored in the collection of The Natural History Museum, London, Paratype fe- male, BMNH 1995.1668; paratype male, BMNH 1995.1669. DESCRIPTION OF ADULT FEMALE Body length not including caudal setae 778 um, body width 341 um. Urosomal somites (Figure 61A,B) without ornamentation on ventral and dorsal surfaces. Seminal receptacle divided into broad anterior and posterior lobes (Figure 61A). Fifth pedigerous somite with fringe of 34 elongate setules at posterior margin. Anal somite with spinular row on ventral surface (Figure 61A). Caudal rami (Figure 61A,B) with convex inner margin; about 2.5 times longer than broad. Anterolateral seta (II) longer than rami with 2 spinules near base (Figure 61B); posterolateral seta (III) with spinular row at base on ventral surface; terminal accessory seta (VI) plumose and about 2 times longer than caudal rami; outer terminal seta (IV) and inner terminal seta (V) well developed and plumose; dorsal seta (VII) about 1.5 times longer than ramus (Figure 61B). S. KARAYTUG AND G.A. BOX 2 f= ==e5 SS SS : = a ee a = ~ Se a M\ Z PARACYCLOPS REVISION SS ew => SS MW, Fig. 55 P. novenarius. Adult female. A, antennule ventral; B, antenna, frontal: C, antennule, second and third segments showing pattern of segmentation, dorsal; D, antenna, coxobasis, caudal; E, F, antennule second and third segments showing variant pattern of segmentation. Scale bars in [tm. 179 S. KARAYTUG AND G.A. BOXSHALL (aes 180 cae | Mili s Denes N SS SS A ‘ AN Vs AF ( BZ 1 Z Z q Nf SSS 90 yyperee ve iN ae MK B= Qn sss === eee \ \\ = ; = ST, \ \\ \S = WSK. KEK i i ———— “ity < Fig.56 P. novenarius. Adult female. A, intercoxal sclerite and coxa of leg 2, posterior; B, leg 2, anterior; C, intercoxal sclerite and coxa of leg 4, posterior; D, leg 4, anterior; E, leg 3, anterior; F, intercoxal sclerite and coxa of leg 3, posterior; G, intercoxal sclerite and coxa of leg 1, posterior; H, leg 1, anterior; J, adult male, terminal endopodal segment of leg 1 showing the sexually dimorphic seta, anterior. Scale bar in um. _ PARACYCLOPS REVISION SSS ~ = as = ss Ni IN | Fig.57 P. novenarius. Adult male. A, urosome, dorsal; B, urosome, ventral; C, leg 5, ventral; D, leg 6, anteroventral. Scale bars in lum. | | 182 S. KARAYTUG AND G.A. BOXSHALL SSSSSSE|TE|AE|AYY = \\ ON ay Bf re Fig. 58 P smileyi. Adult female. A, maxilla; B, leg 5, dorsal; C, urosome, lateral; D, maxillule with inset showing maxillulary palp; E, anal somite and caudal rami, lateral. Scale bars in um. // Wh | \\ S. KARAYTUG AND G.A. BOXSHALL 184 —— = v4 tm J j =a = SN Tl [SN Wd iy VK - SS Fig. 60 P. smileyi. Adult female. A, leg 4, anterior; B, leg 3 with inset showing endopod, posterior; C, leg 2, posterior; D, leg 1, anterior. Scale bar in im. PARACYCLOPS REVISION Antennule 8-segmented (Figure 62A). Segment 3 with partial suture line and spiniform seta. Segment 5 with characteristic short aesthetasc. Setal formula 8, 12, 6,5, 2 + aesthetasc, 2, 2 + aesthetasc, 7 + aesthetasc. Lengths of antennulary segments measured along posterior margin: 32, 42, 27, 59, 30, 20, 32, 35 respectively (length in ttm). Coxobasis of antenna with complex ornamentation on caudal (Figure 61D) and frontal surfaces (Figure 61E) and with spinular row on caudal surface, near base of inner setae as figured. Palp of mandible (Figure 62C) represented by 3 naked setae, 2 of which very long, third seta short. Legs | to 3 each without mid-distal spinular on posterior surface of coxa; without spinular row on anterior surface of intercoxal sclerite (Figure 63A,B,C). Inner coxal spine of leg 4 with group of setules mainly originating posteriorly. Exopodal segments | and 2 without spinular row on posterior surface (Figure 63D). Spine and seta formula as follows: Coxa Basis Exopod Endopod Leg 1 0-1 1-I I-1;I-1;111,5 O0-1;0-1;1,1,4 Leg 2 0-I 1-0 I-1;1-1;111,1,5 0-1;0-2;1,1,4 Leg 3 0-I 1-0 I-1;1-1;111,1,5 0-1;0—2:1,1,4 Leg 4 0-1 1-0 1-1;I-1;1L,1,5 O-1;0—2;1, 11,2 Leg 5 (Figure 63E) comprising single free segment, armed with | strong inner spine with spinules at base and 2 plumose setae about same length; bases of setae produced. Leg 6 (Figure 61B) repre- sented by | naked seta and 2 tiny spinules dorsolaterally. DESCRIPTION OF ADULT MALE Mean body length 552 um (n=2), mean body width 250 um (n=2). Outer lateral seta (III) of caudal rami naked (Figure 3.93B,C). First segment of antennule armed with 8 setae plus aesthetasc; 1 seta (A) large and modified by ornamentation of strong spinules in mid section. All other appendages as in female except for fifth and sixth legs (Figure 64C). Outer plumose seta of leg 5 shorter than in female. Sixth legs armed with | inner spine, | outer naked seta and | well developed spinulose seta in centre. The inner spine of the left leg 5 of a paratype male was abnormal (Figure 64D). ETYMOLOGY. ‘The new species is named in honour of Dr. Janet | Reid who made the specimens available for study, in recognition of | her contributions to cyclopoid systematics. | DIFFERENTIAL DIAGNOSIS. The new species can easily be differen- tiated from all species in the genus by the structure of leg 5 in both _ sexes (Figures 63E; 64E), by the absence of a dorsal spinular row either side of anal somite (Figure 61B), by the spinular ornamenta- | tion of the coxobasis of the antenna on caudal and frontal surfaces | (Figure 61D,E), by the shape of the caudal rami and the structure of its setae in both sexes (Figure 61A,B) and by the lengths of the distal | antennulary segments measured along the posterior margin in the | female (Figure 62A). REMARKS | P. reidae, P. altissimus (Karaytug et al., in press) and P. hardingi _ nom. nov. are closely related. The three species lack a spinular row on the posterior surface of the first and second exopodal segments of | legs 1-3 and possess a spinular row near the base of the two inner setae on the antennary coxobasis in both sexes. However P. reidae _ can easily be differentiated from P. hardingi and P. altissimus by the | relative length of the antennulary segments, the length and spinulation of the outer seta of leg 5, the structure of seminal receptacle and in 185 having the terminal accessory seta (VI) about 2 times longer than the caudal rami. Paracyclops bromeliacola sp. nov. (Figures 65-69) TYPELOCALITY. Brazil, State of Sao Paulo, Miracatu. In bromeliads from a farm at Iterei. March 1995. Collected by Léa P. Corréa. TYPE MATERIAL. Holotype, female dissected on 5 slides (Museu de Zoologia, Sao Paulo, Brazil; MZUSP 12788). Paratypes: 19, 1¢(BMNH 1997. 1782-1785) from Brazil, State of Sao Paulo, Miracatu. In bromeliads from a farm at Iteref. March 1995. Col- lected by Léa P. Corréa; 29 9 in Museu de Zoologia, Sao Paulo, Brazil (MZUSP 12789). Paratypes: 62 9, 30° o'(BMNH 1997. 1786-1802) from Brazil, State of Sao Paulo, Salesopolis, Boracéia Biological Reserve, 7 August 1986. Carlos E. F. da Rocha col. from pools in soil bromeliads, from the Atlantic rain forest. 52 2, 20° Cin Museu de Zoologia, SAo Paulo, Brazil (MZUSP 12790). Paratypes: 39 9, SO O'(BMNH 1997. 1803-1822) from Brazil, State of Sao Paulo, Juréia Ecological Reserve (24°25'10"S, 47°13'50"W), 2 February 1987, Rubens M. Lopes col. In culture made from leaf litter. 6 9 9,30’ C'in Museu de Zoologia, Sao Paulo, Brazil (MZUSP 12791). DESCRIPTION OF ADULT FEMALE Body length and width measurements given in Table 5. Urosomal somites without surface pits on dorsal and ventral surfaces except genital double-somite with very fine surface pits on dorsal surface as figured (Figure 65A). Seminal receptacle (Figure 65B) with poste- rior lobe wider than anterior as figured. Caudal rami (Figure 65A,B) about 2.5 times longer than broad, with inner margin slightly convex distally. Terminal accessory seta (VI) plumose and 1.3 times longer than posterolateral seta (III); outer terminal seta (IV) and inner terminal seta (V) well developed and heterogeneously ornamented (Figure 65B). Antennule 8-segmented (Figure 66A). Segment 3 with partial suture line and spiniform seta. Setal formula 8, 12, 6, 5, 2 + aesthetasc, 2, 2 + aesthetasc, 7 + aesthetasc. Coxobasis of antenna with complex ornamentation on caudal (Figure 66C) and frontal (Figure 66B) surfaces as figured, and without spinular row on caudal surface near base of two inner setae (arrowed in Figure 66C). Basis of leg 1 (Figure 67D) with setiform spine on inner margin reaching halfway along terminal endopodal segment; intercoxal sclerite ornamented with spinular rows on anterior and posterior surfaces (Figure 67D). Intercoxal sclerite of leg 2 ornamented with spinular rows on anterior (Figure 67A) and posterior (Figure 67B) surfaces; coxa without mid-distal spinular row on posterior surface (arrowed in Figure 67B). Intercoxal sclerite of leg 3 without spinular row on anterior (Figure 68A) surface and with 2 spinular rows on posterior (Figure 68B) surface; coxa without mid-distal spinular row on posterior surface (arrowed in Figure 68B). Intercoxal sclerite of leg 4 with 2 spinular rows on posterior (Figure 68D) surface, without spinular row anteriorly (Figure 68C). Spine and seta formula as follows: Coxa Basis Exopod Endopod Leg 1 0-1 1-I I-1;1-1;I1,5 0-1;0-1;1,1,4 Leg 2 0-I 1-0 I-1J-1; 001.5 0-1;0-2;1,1,4 Leg 3 0-1 1-0 I-1;I-1;01,1,5 0-1;0-2;1,1,4 Leg 4 0-I 1-0 -1;I-1;IL1,5 O-1;0-2;1,11,2 Leg 5 (Figure 65C) comprising single free segment, armed with 1 S. KARAYTUG AND G.A. BOXSHALL 186 << Lf cand a Z - a ———— EE may SMe > >P>pVpes—— Sloe Fig. 61 P. reidae sp. nov. Adult female. A, urosome, ventral; B, urosome, dorsal; C, body, dorsal; D, antenna, caudal; E, antenna, coxobasis, frontal. Scale bars in um. _ PARACYCLOPS REVISION Ya ie VI f Mf =A Ee CEES Ss 50 ie Za ae Fig. 62 P. reidae sp. nov. Adult female. A, antennule; B, labrum: C, mandible; D, maxillulary palp; E, maxillule; F, maxilla: G, maxilliped. Scale bar in uum. 187 S. KARAYTUG AND G.A. BOXSHALL 188 ( WOW cone EAA = EX\N\) (Ae Militttdéz es, we ‘ wy of Lo sete f A MF way Ps KKK RXHOIOOOO ae i yf Y == Se FZ Fig. 63 P. reidae sp. noy. Adult female. A, leg 1, anterior; B, leg 2, anterior; C, leg 3, anterior; D, leg 4, anterior; leg 5, ventral. Scale bars in um. PARACYCLOPS REVISION (G SS EEE —— 189 PINAR rae SS SS 2 SSssSESTII953 250 A 100 BC \ 25 DE Fig. 64 P. reidae sp. noy. Adult male. A, body, dorsal; B, urosome, dorsal; C, urosome, ventral; D, abnormal leg 5 (right), ventral; E, normal leg 5 (left), ventral. Scale bars in um. long (almost twice as long as inner spine) well developed outer spinulose seta, | inner spine, 1 plumose seta in centre. Leg 6 (Figure 65A) represented by 1 plumose seta and 2 tiny spinules dorsolaterally. DESCRIPTION OF ADULT MALE Body length and width measurements given in Table 5. Genital somite with fine surface pits on dorsal surface, other somites without surface pits (Figure 69A,D). Caudal rami, short, only twice as long as broad, with inner margin convex distally (Figure 69A,D). First segment of antennule armed with 8 setae plus aesthetasc; | seta (A) large and modified by ornamentation of strong spinules in proximal and mid sections, tapering to fine point distally. 190 One seta on terminal endopodal segment of leg | spinulose (Figure 67C). Outer seta of fifth leg plumose and less well developed (Figure 69B) than in female; sixth leg (Figure 69C,D) armed with | inner spine, about as long as second urosomal somite, bearing spinular row at base, and 2 outer plumose setae, middle seta short and stout. VARIABILITY. Specimens from Juréia Ecological Reserve and Salesopolis were blackish in colour whilst specimens from Miracatu were pale brown. ETYMOLOGY. The species name is derived from the Bromeliaceae, name of the family of plants providing a microhabitat for this species, and from colere meaning to inhabit. It refers to the preferred cryptic habitat of the species. DIFFERENTIAL DIAGNOSIS. The new species can be differentiated from other Paracyclops species by the combination of the following characters: by the long (almost twice as long as inner spine) and well developed outer spinulose seta of leg 5 (Figure 65C), by the structure of the seminal receptacle (Figure 65B), by the absence of a mid- distal spinular row on the posterior surface of leg 2 (arrowed in Figure 67B) and leg 3 (arrowed in Figure 68B) and by the absence of surface pits from the dorsal and ventral surfaces of all urosomal somites except the genital double-somite, which has very fine surface pits on the dorsal surface as figured (Figure 65A,B). REMARKS P. bromeliacola is closely related to the other new Brazilian species, P. rochai sp. nov. and P. punctatus sp. noy. They share the absence of a mid-distal spinular row from the posterior surface of the coxa of leg 2 and leg 3, and they all lack the spinular row near the base of the two inner setae on the coxobasis of the antenna in both sexes (arrowed in Figure 66C). However P. bromeliacola differs from P. rochai and P. punctatus in the long (almost twice as long as inner spine) and well developed outer spinulose seta of leg 5 (Figure 65C) and in the structure of the seminal receptacle (Figure 65B). It also differs from P punctatus in the absence of surface pits from the dorsal and ventral surfaces of the urosomal somites except the genital double-somite of the female and genital somite of the male, both of which have very fine surface pits on the dorsal surface as figured (Figures 65A,B; 69A,D). Paracyclops punctatus sp. nov. (FIGUREs 70-73) TYPE LOCALITY. Brazil, State of Sergipe, Riachao do Dantas (11°02'S, 37°45'W), 24 July 1986, Carlos E. F. Rocha col. In leaf pools in soil bromeliads in an Atlantic rain forest remnant. TYPE MATERIAL. Holotype: (Museu de Zoologia, Sao Paulo, Bra- zil. MZUSP 12792) female dissected on 5 slides. Paratypes 29 9, S. KARAYTUG AND G.A. BOXSHALL 30 C(BMNH 1997. 1824-1834). 49 9, 40° Cin Museu de Zoologia, Sao Paulo, Brazil (MZUSP 12793). DESCRIPTION OF ADULT FEMALE Body length (um) not including caudal setae, 630-711, mean = 681, n = 5; body width 274-309, mean = 292, n = 5. Genital double-somite, second and third abdominal somites ornamented with conspicuous surface pits on dorsal (Figure 70A) and ventral (Figure 70B) surfaces as figured. Genital double-somite widest anteriorly, narrowing posteriorly. Seminal receptacle as figured (Figure 70B). Third and fourth urosomal somites with well-devel- oped hyaline frill dorsally on posterior margin (Figure 70A). Caudal rami 2.8 times longer than broad; with cuticular depres- sions on ventral surface (Figure 70B); with inner margin convex distally. Terminal accessory seta (VI) plumose and longer than posterolateral seta (III); outer terminal seta (IV) and inner termi- nal seta (V) well developed and heterogeneously ornamented (Figures 70B; 72B). Antennule 8-segmented (Figure 71A,B). Segment 3 with two partial suture lines on ventral and dorsal surfaces; with spiniform seta distally. Segment 4 with partial suture line ventrally, extending dorsally (Figure 71A,B). Segment 5 with characteristic short aesthetasc. Setal formula 8, 12, 6,5, 2 + aesthetasc, 2, 2 + aesthetasc, 7 + aesthetasc. Coxobasis of antenna with complex ornamentation on caudal (Figure 71D) and frontal (Figure 71C) surfaces; without spinular row on caudal surface near base of two inner setae (arrowed in Figure 71D). Coxa of leg 1 without mid-distal spinular rows on posterior surface (arrowed in Figure 72F); intercoxal sclerite without spinular row on posterior surface; basis with spinular row on anterior surface (arrowed in Figure 72H). Intercoxal sclerite of leg 2 ornamented with spinular row on anterior (Figure 72G) and posterior (Figure 72E) surfaces; coxa without mid-distal spinular row on posterior surface (arrowed in Figure 72E). Intercoxal sclerite of leg 3 with 2 spinular rows on posterior surface (Figure 72D); coxa without mid- distal spinular row on posterior surface (arrowed in Figure 72D). Intercoxal sclerite of leg 4 (Figure 72C) with 2 spinular rows on posterior surface. Spine and seta formula as follows: Coxa Basis Exopod Endopod Leg 1 0-1 1-I I-1;J-1;111,5 0-1;0-1;1,1,4 Leg 2 0-I 1-0 I-1;-1;0L,15 0-1;0-2:1,1,4 Leg 3 0-1 1-0 I-1;1-1;10,1,5 0-1;0-2;1,1,4 Leg 4 0-1 1-0 I-1;I-1;11,1,5 O-1;0—2;1, 11,2 Leg 5 (Figure 70C) comprising single free segment, armed with | long (1.6 times longer than inner spine) outer spinulose seta, 1 inner spine, | plumose seta in centre. DESCRIPTION OF ADULT MALE Body length (um) not including caudal setae, 657—701, mean = 688, Table 5 Body length (BL) and width (BW) measurements (in um) of Paracyclops bromeliacola in various localities. (N = number of specimens meas- ured) Locality Sex BL (mean + SD) Brazil (State of Sao Paulo) Q 654 + 62.1 Miracatu of 531 Brazil (State of Sado Paulo) Q T71 + 38.5 Juréia Ecological Reserve of 636+ 8.1 Brazil (State of Sao Paulo) Q 705 + 28.5 Salesdpolis (of 636 + 24.2 Range BL (mean + SD) Range N 595-741 265 + 32.8 240-313 4 210 1 716-847 330+ 9.3 314-343 9 617-647 245 + 11.1 227-262 10 657-741 274 + 14.3 254-296 10 605-662 238 + 5.4 230-242 4 192 S. KARAYTUG AND G.A. BOXSHALL Fig. 66 P. bromeliacola sp. nov. Adult female. A, antennule; B, antenna, coxobasis, frontal; C, antenna, caudal. Scale bars in um. 193 and basis of leg 2, posterior; C, adult male, terminal as cae SS Y Fw — Miao My v. Adult female. A, leg 2, anterior; B, intercoxal sclerite, coxa > vat, \ endopodal segment of leg 1, posterior; D, adult female, leg 1, anterior, Scale bar in [um. PARACYCLOPS REVISION | Fig. 67 P. bromeliacola sp. no 194 S. KARAYTUG AND G.A. BOXSHALL i> OI =, b Loe 7 i ‘ Se SF QZ — Py” = D (a nei Pees : WENN plassecan y Wa : 44449, tg IMU Steytgh sddayy 7 ol Fig. 68 P. bromeliacola sp. nov. Adult female. A, leg 3, anterior; B, intercoxal sclerite, coxa and basis of leg 3, posterior; C, leg 4, anterior; D, intercoxal sclerite, coxa and basis of leg 4. Scale bar in um. 195 PARACYCLOPS REVISION Se ae, SS 2s Pe Sn = ——— Se Sa ISS ral. Scale bars in um. me, vent ntral; D, uroso al; C, leg 6, anterove Fig. 69 P. bromeliacola sp. nov. Adult male. A, urosome, dorsal; B, leg 5, ventr; 196 S. KARAYTUG AND G.A. BOXSHALL Fig. 70 P. punctatus sp. noy. Adult female. A, urosome, dorsal; B, urosome, ventral; C, leg 5, ventral; D, anal somite, dorsal. Scale bars in um. PARACYCLOPS REVISION LESS ia Ds SS ee (CP 5 DSB ate \ ae \ 2 ‘ ‘ ' 197 | Fig. 71 P punctatus sp. nov. Adult female. A, antennule: B, antennule showing variant pattern of segmentation; C, antenna, coxobasis and first endopodal segment, frontal; D, antenna, caudal. Scale bars in im. 198 S. KARAYTUG AND G.A. BOXSHALL . \ ee Y LZ % shd44 SOAAMAMMA MANS dyavend® padded 4 N Fig. 72 P. punctatus sp. nov. Adult female. A, body, dorsal; B, detail of setal elements of caudal rami, dorsal; C, intercoxal sclerite, coxa and basis of leg 4, posterior; D, intercoxal sclerite, coxa and basis of leg 3, posterior; E, intercoxal sclerite and coxa of leg 2, posterior; F, intercoxal sclerite and coxa of leg 1, posterior; G, intercoxal sclerite of leg 2, anterior; H, basis of leg 1, anterior. Scale bars in um. 199 PARACYCLOPS REVISION | Fig. 73 P. punctatus sp. nov. Adult male. A, leg 5, ventral; B, urosome, dorsal; C, urosome, ventral; D, leg 6, ventral. Scale bars in Lum. 200 n=5; body width 274-291, mean = 279, n=5. Genital, third, fourth and fifth urosomal somites ornamented with cuticular pits on dorsal surfaces (Figure 73B,C). First segment of antennule armed with 8 setae plus an aesthetasc; | seta (A) large and modified by ornamen- tation of strong spinules in proximal and mid sections, tapering to fine point distally. Spinular ornamentation of the coxobasis of the antenna as in the female. Outer seta of fifth leg plumose and less well developed than in female (Figure 73A). Sixth leg (Figure 73D) armed with | inner spine, shorter than second urosomal somite, and 2 outer plumose setae; middle seta ornamented with long setules. VARIATION, FEMALE. The extent of the partial suture line on the fourth segment of the antennule varies (Figure 71A,B); it may be shorter than in the figured specimens. The dorsal hyaline frill on the posterior margin of the third urosomal somite may be more or less well developed (cf. Figure 70A and Figure 70D). ETYMOLOGY. ‘The name of the new species is derived from the Latin punctum meaning point. It refers to the ornamentation of cuticular pits on the surface of the urosomal somites. DIFFERENTIAL DIAGNOSIS. ‘The new species differs from other Paracyclops species by the combination of the following characters: the presence of conspicuous surface pits on the dorsal (Figure 70A) and ventral (Figure 70B) surfaces of the genital double-somite, and the second and third abdominal somites of the female, the structure of leg 5, the absence of a mid-distal spinular rows on the posterior surfaces of leg 1 (arrowed in Figure 72F), leg 2 (arrowed in Figure 72E) and leg 3 (arrowed in Figure 72D), the presence of integumental pits on the ventral surface of the caudal rami (Figure 70B), the subdivision of the fourth segment of the female antennule (Figure 71A,B), and the absence of a spinular row on the caudal surface of the antennal coxobasis near the base of the two inner setae in both sexes (arrowed in Figure 71D). REMARKS P. punctatus 1s closely related to the other new Brazilian species, P. rochai and P. bromeliacola. They share the absence of a mid- distal spinular row from the posterior surface of the coxa of leg 2 and leg 3, and they all lack the spinular row near the base of the two inner setae on the coxobasis of the antenna in both sexes (arrowed in Figure 71D). However P. punctatus differs from P. rochai and P. bromeliacola in the presence of conspicuous surface pits on the dorsal (Figure 70A) surface of the genital double- somite, and the second and third abdominal somites of the female, in the structure of the seminal receptacle (Figure 70A), in the subdivision of the fourth segment of the female antennule (Figure 71A,B) and in the weakly developed outer seta of leg 5 of the female (Figure 70C). Paracyclops rochai sp. nov. (Figures 74-76) TYPE LOCALITY. Brazil, State of Sergipe, Itabaina Mountains, at gruta, near to Areia Branca, 6/11/1993, Carlos E. F. da Rocha collection. TYPE MATERIAL. Holotype: (Museu de Zoologia, Sao Paulo, Bra- zil. MZUSP 12794) female dissected on 4 slides. Paratypes: 89 9, and 80°'o (BMNH 1997. 1840-1870). 79 9, 70’ Cin Museu de Zoologia, Sao Paulo, Brazil (MZUSP 12795). DESCRIPTION OF ADULT FEMALE Body length (um) not including caudal setae, 506-674, mean = 596, n= 10; body width 204—247, mean = 227, n= 10. Urosomal somites S. KARAYTUG AND G.A. BOXSHALL (Figure 74A,B) without surface pits on dorsal and ventral surfaces. Seminal receptacle as figured (Figure 74B). Caudal rami (Figure 74A,B) parallel and about 2.7 times longer than broad; terminal accessory seta (VI) plumose and about as long as posterolateral seta (II]); outer terminal seta (IV) and inner terminal seta (V) well developed and heterogeneously ornamented (Figure 74B). Antennule 8-segmented (Figure 75A); segment 3 with two partial suture lines and spiniform seta. setal formula 8, 12, 6, 5, 2 + aesthetasc, 2, 2 + aesthetasc, 7 + aesthetasc. Coxobasis of antenna with complex ornamentation on caudal (Figure 75B) and frontal (Figure 75C) surfaces as figured and without spinular row near base of two inner spinulose setae (arrowed in Figure 75B). Coxa of leg 1 without mid-distal spinular row on posterior sur- face. Intercoxal sclerite of leg 4 with 2 spinular rows on posterior surface (Figure 75G). Spine and seta formula as follows: Coxa Basis Exopod Endopod Leg | 0-1 1-I I-1;[-1;111,5 O-1;0-1;1,1,4 Leg 2 0-I 1-0 J—-1;I-1;10, 15 0-1;0-2:1,1,4 Leg 3 0-1 1-0 I-1;I-1;10,15 0-1;0-2;1,1,4 Leg 4 O-I 1-0 I-1;I-1;I1,1,5 0-1;0—2;1,11,2 Leg 5 (Figure 74C) comprising single free segment, armed with 1 long (almost twice as long as inner spine), well developed outer spinulose seta, | strong inner spine with 3 spinules at base. Other appendages as in P. bromeliacola. DESCRIPTION OF ADULT MALE Body length (um) not including caudal setae, 459-560, mean = 519, n= 10; body width 185—205, mean = 196, n= 10. Urosomal somites without surface pits on dorsal (Figure 76B) and ventral (Figure 76B) surfaces, except that some fine pits visible on genital somite on dorsal surface. First segment of male antennule armed with 8 setae plus an aesthetasc; | seta (A) large and modified by ornamentation of strong spinules in proximal and mid sections. Spinular rows on posterior surface of intercoxal sclerite of leg 4 well developed (Figure 75F). Outer seta of leg 5 plumose and about as long as inner spine (Figure 76C). Sixth leg (Figure 76C) armed with | inner spine, half as long as second urosomal somite. VARIABILITY, FEMALE. The length of the terminal endopodal seg- ment of leg 4 varied as indicated in Figure 75D,E. One female showed complete subdivision of the third segment of the antennule (Figure 75A). ETYMOLOGY. ‘The species is named after Prof. Carlos Eduardo Falavigna da Rocha (University of Sao Paulo), the collector of this material, in recognition of his many contributions to the study of cyclopoid copepods. DIFFERENTIAL DIAGNOSIS. P rochai can be differentiated from other Paracyclops species by the combination of the following characters: the structure of leg 5 in the female (Figure 74C), the absence of surface pits on the dorsal and ventral surfaces of the urosomal somites in the female (Figure 74A,B), and the absence of a spinular row near the base of the two inner spinulose setae in both sexes (arrowed in Figure 75B). REMARKS P. rochai is closely related to P. chiltoni and P, punctatus. However, it differs from P. chiltoni in the absence of a spinular row near the base of the two inner spinulose setae in the male (arrowed in Figure 201 PARACYCLOPS REVISION a ee aS Io) n iS - =) } us} iS) i=] oy n § S £ qu + 6 ob a Fig. 76 P. rochai sp. nov. Adult male. A, urosome, ventral: B, urosome, dorsal; C, detail of leg 5 and leg 6, ventral. Scale bars in um. 203 204 75B), by the length of the outer seta of leg 5 in the female (Figure 74C), by the absence of surface pits on the dorsal and ventral surfaces of the urosomal somites in the female (Figure 74A,B), and by the absence of integumental pits on the ventral surface of the caudal rami in the female (Figure 74B). P. rochai differs from P. punctatus by the absence of surface pits on the dorsal and ventral surfaces of the urosomal somites in the female (Figure 74A,B), by the absence of integumental pits on the ventral surface of the caudal rami in the female (Figure 74B), by the structure of leg 5 (Figure 74C) and by the ornamentation of the outer terminal seta (IV) and the inner terminal seta (V) of the caudal rami in both sexes (Figure 74B; 76B). ACKNOWLEDGEMENTS. We would like to thank Dr Rony Huys for the translation of much German literature and commenting on the manuscript. This study would not have been possible without the assistance of many people who generously made material and information available. The follow- ing people are therefore gratefully acknowledged: Dr. Janet Reid (Smithsonian Institution, Washington D.C., U.S.A); Dr. Danielle Defaye (Museum Na- tional d’Histoire Naturelle, Paris, France); Dr. Teruo Ishida (Japan); Prof. Carlos E. F. Da Rocha (Universidade de Sao Paulo, Brazil); Janice Clark Walker (Smithsonian Institution, Washington D.C., U.S.A); Prof. Dr. Henri Dumont (Universiteit Gent, Belgium); Dr. Y. Ranga Reddy (Nagarjuna University, India); Prof. Victor R. Alekseev (Zoological Institute, Russian Academy of Science, St. Petersburg, Russia); Dr. Jouko Sarvala (University of Turku, Finland); Prof. Dr. Marit E. Christiansen (Zoologisk Museum, Universitetet I Oslo, Norway); Judith C. Price (Canadian Museum of Na- ture); Dr Iskandar M. Mirabdullayev (Institute of Zoology, Tashkent, Uzbekistan); Dr. Harry Yeatman (The University of the South, Sewanee, U.S.A); Dr. Silvina Menu Marque (Universidad de Buenos Aires, Argentina). This research was funded by Balikesir University, Turkey. REFERENCES Boxshall, G. A. & Braide, E. I. 1991. The freshwater cyclopoid copepods of Nigeria, with an illustrated key to all species. Bulletin of the British Museum (Natural History), Zoology 57(2): 185-212. Brady, G. S. 1878. A monograph of the free and semi-parasitic Copepoda of the British Islands. Vol. 1: 1-148. The Ray Society, London. 1892. A revision of the British species of freshwater Cyclopidae and Calanidae. Transactions of the natural History Society of Northumberland, Durham and Newcastle-upon-Tyne, 11: 68-120. Byrnes, E. F. 1909.The fresh-water Cyclops of Long Island. Cold Spring Harbour Monographs, 7: 1\-43. Claus, C. 1893. 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Some laurentian copepods and their variations. Transactions of the Royal Canadian Institute 20(1): 77-98. Yeatman, H.C. 1959. Free-living Copepoda: Cyclopoida. In: Ward, H.B. & Whipple, G.C.; Freshwater Biology, 2nd ed., 1959 (ed. W.T. Edmondson) : 795-815. weve aS ee e ° : Herve! «~Si9 we, re ef - = : es TH ~ ; = OS, aah eee le é (2 fot «ood, ome fee he - m~ re *-4 an -) =——t aah ee + © ape ee — pa” mind . ++)? ~ oR . rn £ cS fap : ao bp ; 3 acs 7 SD a 2 a4 © tw intel : Pa) Y meat pw | ty ah tA AED Pd TS oe Se Ok ceo ~~ wr. | A ek ) ig PQS oS & chi |, Alleles > —— why ee Os ave a”, re wu ‘se i ~~ a ne Bull. nat. Hist. Mus. Lond. (Zool.) 64(2): 207-211 Issued 26 November 1998 First records and a new subspecies of Rhinolophus stheno (Chiroptera, Rhinolophidae) from Vietnam. GABOR CSORBA Department of Zoology, Hungarian Natural History Museum, Baross u. 13, H-1088 Budapest, Hungary PAULINA D. JENKINS *Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD Synopsis. The recently discovered populations of Rhinolophus stheno from North Vietnam, along with specimens previously collected in Thailand, are described as a new subspecies, Rhinolophus stheno microglobosus. The median anterior rostral swellings of the new subspecies are notably smaller than those of the nominate subspecies. A morphological and statistical comparison is given between the two subspecies of R. stheno, and the closely related R. malayanus. INTRODUCTION Rhinolophus stheno Andersen, 1905 was originally described from peninsular Malaysia. The known range of the species was later extended to Thailand (Lekagul & McNeely, 1977), Sumatra and Java (Corbet & Hill, 1992; Koopman, 1994) and Tioman Island, off the coast of Malaysia (Csorba et al., 1997). Recent expeditions to Vietnam led by British and Hungarian researchers have discovered the first specimens of R. stheno to be recorded from that country. Comparative examination of these specimens with other populations in the collections of The Natural History Museum, London revealed that specimens from Vietnam were most similar to those from Thailand, and that both were sufficiently different from material from Malaysia, Sumatra and Java to represent an undescribed subspecies. Andersen (1905) considered R. stheno to belong to the borneensis subgroup of the simplex-group of Rhinolophus, which Tate & Archbold, 1939 subsequently termed the ferrumequinum-group. Andersen distinguished R. stheno from other members of the borneensis sub-group by the much more projecting anterior nasal swellings of the rostral part of the skull. Lekagul & McNeely (1977) reported that R. stheno resembles R. malayanus Bonhote, 1903 but that the two are separable by a set of external features (body size, shape of lancet and relative proportions of the first and second phalanges of the third digit). Subsequently, McFarlane & Blood (1986) concluded that, although there are no reliable differences between R. stheno and R. malayanus in these features, they are instead distinguishable by supraorbital and rostral characters of the cranium. They suggested that the general similarity of the noseleaf and skull of R. stheno and R. malayanus implied a closer relationship than formerly supposed. This view was accepted by Corbet & Hill (1992), who continued to group both species in the ferrumequinum group, and keyed the two species on the basis of the shape and size of the anterior and posterior rostral compartments. Bogdanowicz (1992), ina phenetic analysis of the whole family, proposed different group-level classifications for the two species (R. malayanus in the megaphyllus group but R. stheno, with a question mark indicating uncertainty, in the ewryotis group). Specimens of R. malayanus and R. borneensis Peters, 1861 were *Address for correspondence. © The Natural History Museum, 1998 also collected during the recent expeditions, confirming the pres- ence of R. borneensis in Vietnam (see Hill & Thonglongya, 1972, Corbet & Hill, 1992 and discussion below). In view of the various theories outlined above concerning the relationship between R. stheno and R. malayanus, morphological comparisons and a Princi- pal ComponentsAnalysis are given below between the two subspecies of R. stheno and R. malayanus. MATERIALS AND METHODS All available specimens were included in the morphological com- parisons but for the multivariate analysis, which requires the use of complete sets of measurements, the reduced number of specimens is given in parentheses as follows: 12 (8) specimens of R. s. microglobosus described below, 21 (13) specimens of the nominate subspecies of R. stheno (from Sumatra, Java and Malaysia) and 14 (11) specimens of R. malayanus (from Thailand and Malaysia). External measurements, to the nearest 0.1 mm, were taken from dry and alcoholic museum specimens using digital calipers. Cranial measurements, to an accuracy of 0.01 mm, were collected using digital calipers and a binocular microscope. Characters for the multivariate analysis included one external and nine cranial meas- urements, as follows, with the abbreviation in parentheses: 1. forearm length (FA) 2. greatest skull length (GSL) — measured from the anterior of the canine to the posteriormost part of the occiput; 3. maxillary toothrow length (MTL) — the crown length from the anterior of the upper canine (C) to the posterior of the third upper molar (M3); 4. zygomatic width (ZW) — the greatest distance across the zygoma; 5. mastoid width (MW) — the greatest distance across the mastoid region of the braincase; 6. mandible length (ML) — the distance from the most posterior portion of the articular process to the anteriormost edge of the alveolus of the first lower incisor (il ); 7. lower toothrow length (LTL) — the crown length from the anterior of the lower canine (c) to the posterior of the third lower molar (m3); 8. interorbital width (IW) — the least width of the interorbital constriction; 9. rostral swelling width (RSW) — the greatest width of the nasal swellings; 10. median anterior rostral swelling width (MARW) — greatest width in dorsal view. To reveal the taxonomic differences and relations between the taxa included in this study the Mann-Whitney U Test and Principal Component Analysis (PCA) were used. For the univariate analysis a non-parametric method was applied since the raw data did not meet the criteria for a normal distribution based on the F-test Both statistical methods were performed by Statistica 5.1, 1984-1995 statistical programme of StatSoft Inc. run on a 486 PC. Abbreviations used for institutions are: BMNH — The Natural History Museum, London, formerly the British Museum (Natural History); HNHM — Hungarian Natural History Museum, Budapest; MNHN — Muséum National d’Histoire Naturelle, Paris; IEBR — Institute of Ecology and Biological Researches, Hanoi. RESULTS Rhinolophus stheno microglobosus ssp. nov. Figs 1-4, Table 1 HOLOTYPE. BMNH 1997.360 (field number 9601/B 11), adult male in alcohol, skull extracted, collected by members of ‘Frontier’ the Society for Environmental Exploration — Vietnam, between 17 January and 18 March 1996. Type LocALity. Na Hang Nature Reserve, Tuyen Quang Prov- ince, Vietnam, between 22°16' and 22°31'N, 105°22' and 105°29'E, altitude 100-1082 m. Highly diverse evergreen and semi-evergreen primary limestone rainforest (see Hill & Kemp, 1996). PARATYPES. The same collection data as the holotype: BMNH 1997.359 (field number 9601/B10) adult female in alcohol, skull extracted; BMNH 1997.361 (field number 9601/B25) adult male in alcohol, skull extracted; BMNH 1997.362 (field number 9601/B28) adult female in alcohol; 2 km SE of Pac Ban, Na Hang Nature Reserve, Tuyen Quang Province, Vietnam, 22°19'N, 105°25'E, alti- tude 300 m, 3 March 1997, collected by Gabor Csorba and Pham Duc Tien: HNHM 98.1.1. (field number CSOVI 30) adult female, skin, skull and skeleton; HNHM 98.1.2-3. (field number CSOVI 32, 33) adult males, in alcohol, skull extracted; IEBR (not catalogued, field number CSOVI 31) adult female, in alcohol, skull extracted. Referred material: Tham Tap Tao, Fang, Chiangmai, Thailand, 19°55'N 99°13'E BMNH 1978.974, adult female in alcohol, skull extracted, BMNH 1978.2301, adult skull only; Chanthaburi, Pong Nam Ron, Khao Soi Dao Tai, Thailand, 12°36'N 102°09'E 850 m: BMNH 1978.2298—2300, adult skulls only. COMPARATIVE MATERIAL. Rhinolophus stheno stheno —West Ma- laysia: Selangor (BMNH 1898.3.13.1 [holotype], 1898.3.13.2-3, 1973.606-607); Gunong Benom, Pahang (BMNH 1967.1492, 1967.1494, 1967.1497, 1967.1533—1534); Batu Pahat, Kangar, Perlis (BMNH 1968.817—818); Tioman Island (HNHM 95.55.2-4); Indo- nesia: Saekaranda, N. E. Sumatra (BMNH 1907.1.9.2, MNHN 1903.3); Kalipoetjang, Tji-Tandoei River, Java (BMNH 1909.1.5.179-182). Rhinolophus malayanus — Yhailand: Biserat, Jalor (BMNH 1903.2.6.83 [holotype], 1903.2.6.84, 1908.2.5.24—25); Phu Nam Tok, Saraburi (BMNH 1970.1462); Phu Nam Tok Tap Kuang, Khaeng Khoi, Saraburi (BMNH 1978.973); Satun, Muang, Wang Bla Chan (BMNH 1978.2295); Chiangmai, Fang, Tham Tap Tao G. CSORBA AND P. JENKINS (BMNH 1978. 2296-2297); West Malaysia: Batu Pahat, Kangar, Perlis (BMNH 1968.812); Kisap Forest Reserve, Pulau Langkawi (BMNH 1968.813-816). DIAGNOSIS. Anterior median rostral compartments abruptly el- evated but narrow and globular in outline; posterior median rostral compartments very small but slightly inflated dorso-laterally. Skull slender, rostral swelling width < 5.1, zygomatic and mastoid width <9) II. DESCRIPTION. A medium-sized horseshoe bat belonging to the ferrumequinum group (sensu Corbet & Hill, 1992), forearm length 43.8-47.2, mean 45.46, SD 1.21, n=8; head and body length 38.8— 45.2, mean 43.39, SD 2.73, n= 8; tail length 17.7—23.0, mean 20.0, SD 1.49, n=8; hindfoot length 7.8—8.4, mean 8.0, SD 0.18, n=8; ear length 16.9-18.7, mean 17.89, SD 0.64, n = 8; weight 9-9.5 grams, mean 9.33, SD 0.24, n = 3. Ear medium in length, just reaching the tip of nose when laid forward. Noseleaf with sella almost parallel- sided, only narrowing very slightly, rounded at tip; the connecting process rounded, typical for the ferrumequinum group; the lancet long, straight-sided, its tip cuneate; the supplementary noseleaf clearly visible; the lower lip has three groves (Fig. 1). The dorsal pelage is light yellowish-brown at the base of hairs, reddish cinna- mon-brown above and c. 8 mm long, that of the venter paler and shorter. The wing membranes are uniformly dark brown. The fifth metacarpal is subequal or slightly longer than the fourth, the third shorter than fourth. Ratio of first to second phalange of third digit 1.56-1.67, mean 1.62, SD 0.04, n= 8. Skull averaging smaller than in R. s. stheno; slender, rostral swelling width less than 5.1 mm, zygomatic and mastoid width subequal, not exceeding 9.1 mm (see Table 1). The anterior median rostral compartments are high and abruptly elevated but narrow and not forming the lateral walls of the rostrum, in profile they are posteriorly concave but less sharply so than in R. s. stheno; the posterior median rostral compartments are slightly inflated dorso- laterally so that the anterior region of the supraorbital depression is shallow and narrow, unlike the deep broad depression of R. s. stheno; lateral rostral compartments slightly inflated (see Fig. 2). The sagittal crest moderately developed. Palatal bridge less than one- third of the upper toothrow length. Anterior upper premolar well Fig. 1 Lateral (left) and frontal (right) views of noseleaves of R. s. microglobosus (HNHM 98.1.2. [paratype]). L = lancet; C = connecting process; S = sella; SN = supplementary noseleaf. Scale = 5 mm. NEW SUBSPECIES OF RHINOLOPHUS STHENO 209 Table 1 Selected external and craniodental measurements (in mm) of R. s. microglobosus, R. s. stheno and R. malayanus presented as range, mean + standard deviation and number of specimens in parentheses. Column |: character. Column 2-4: taxon. Column 5—7: Mann-Whitney U Test p-levels between groups. Character FA GSL MTL ZW MW ML LIL IW RSW MARW R. s. microglobosus 43.8-47.2 45.46+ 1.21 (8) 18.22-19.38 18.73+ 0.35 (10) 6.74-7.45 7.05+ 0.20 (12) 8.81—9.06 8.93+ 0.08 (11) 8.71-9.07 8.87+ 0.11 (12) 11.83-12.85 12.23+ 0.29 (12) 7.04—7.80 7.45+ 0.22 (12) 1.49-1.85 1.66+ 0.11 (12) 4.78-5.07 4.91+ 0.11 (12) 3.53-4.00 3.82+ 0.12 (12) Taxon R. s. stheno 43.6-47.2 45.55+ 1.06 (21) 18.84-19.92 19.37+ 0.31 (15) 7.25—-7.78 7.49+ 0.15 (16) 9.29-9.75 9.56+ 0.16 (15) 9.09-9.66 9.41+ 0.16 (15) 12.59-13.31 12.95+ 0.21 (16) 7.68-8.40 7.98+ 0.17 (16) 1.64 —2.00 1.82+ 0.13 (15) 5.01-5.38 5.13+ 0.10 (15) 4.13-4.36 4.23+ 0.07 (15) R. malayanus p-levels between groups R. s. stheno— R.malayanus R. s. stheno- R. s. microglobosus R. s. microglobosus— R. malayanus 38.3-42.4 40.45+ 1.18 (14) 17.25=17.79 17.56+ 0.22 (11) 6.54-7.09 6.75+ 0.16 (11) 8.47-8.94 8.77+ 0.13 (11) 8.14-8.45 8.23+ 0.11 (11) 11.18-12.08 11.79+ 0.25 (11) 6.86—7.36 7.1L7£O0R7GD) 2.13—2.67 2.44+ 0.18 (11) 4.94-5.37 5.16+ 0.12 (11) 3.99-4.4] 4.15+ 0.13 (11) Fig. 2 Dorsal view of rostral part of skulls of a.) R. malayanus (BMNH 3.2.6.83 [holotype]), b.) R. s. stheno (BMNH 98.3.13.1 [holotype]) and c.) R. s. microglobosus (HNHM 98.1.1. [paratype]), Scale = 5 mm. 0.828 0.000 0.000 0.001 0.000 0.000 0.001 0.000 0.001 0.000 0.000 0.019 0.000 0.000 0.000 0.000 0.000 0.008 0.000 0.000 0.001 0.033 0.000 0.000 0.001 0.885 0.002 0.000 0.016 0.000 developed with distinct cusp, included in the toothrow. Lower middle premolar (p3) small and fully extruded from the toothrow; first (p2) and last (p4) lower premolars in contact or nearly so; p2 moderately small and narrow, antero-posterior axis only slightly displaced rela- tive to main axis of toothrow, unlike R. stheno stheno in which p2 is slightly larger, overlaps more with the lower canine and p4, and in which the axis is more skewed. ETYMOLOGY. The Latin word microglobosus refers to the size and shape of the median anterior rostral swellings which are considerably smaller than those of the nominate subspecies. COMPARISONS WITH OTHER TAXA. Besides the classical morpho- logical comparisons of the new subspecies, R. s. microglobosus and the nominate subspecies, R. s. stheno, the Mann-Whitney U Test was also performed to reveal if statistically significant differences were present in morphological characters. In the course of the analysis highly significant differences (highest p < 0.01) were shown in greatest skull length, maxillary toothrow length, zygomatic width, mastoid width, mandible length, lower tooth-row length, rostral swelling width and median anterior rostral swelling width; in all cases the new subspecies was smaller (Table 1). The same method was used for pair-wise comparisons between R. malayanus and R. s. steno, and between R. malayanus and R. s. microglobosus (see Table | for p-levels between groups). Significant differences (at p < 0.01) were shown for the following variables: forearm length, greatest skull length, lower toothrow length, mastoid width, in which R. malayanus was smaller in each parameter, and interorbital width where R. stheno was smaller. To help elucidate the relationships of the three taxa, a Principal Component Analysis (PCA) was performed using the characters recorded in Table 1. The scatterplots of the specimens against the factor | (F1) and factor 2 (F2) axes showed aclear separation of three groups (Fig. 3) supporting the view that R. s. microglobosus repre- sents a distinct taxonomic unit. The first two factors represent more than 89% of the total variance where F2 was identified as the ‘rostral 210 : R. s. stheno ° (eo) 1 r r " r ° fe} fe F ° Oe oO oL ° ° os e ft ir [ a 1b x -2 -1.5 -1 -0.5 0 0.5 1 1.5 FACTOR1 G. CSORBA AND P. JENKINS R. malayanus raed Een oy eae Fig. 3. Principal components analysis of R. s. microglobosus, R. s. stheno and R. malayanus specimens based on 10 external and craniodental characters. swellings component’ in which the two measurements of the ros- trum (rostral swelling width and median anterior rostral swelling width) were the most important, and Fl pertained to the other characters (Table 2). Table 2 Factor loadings of external and craniodental characters obtained by Principal Component Analysis Character Factor | Factor 2 Forearm length —0.873 —0.344 Greatest skull length —0.985 —0.040 Maxillary toothrow length 0.954 0.075 Zygomatic width —0.887 0.283 Mastoid width —0.972 —0.003 Mandible length —0.943 0.126 Lower toothrow length —0.955 0.096 Interorbital width 0.740 0.569 Rostral swelling width —0.050 0.929 Median anterior rostral swelling width —0.206 0.894 Variance explained 67.67% 22.17% DISCUSSION The new records of R. stheno extend the known distribution of the species to North Vietnam, and represent a new subspecies which is characterised by its generally smaller, narrower skull and above all, by the small, globular anterior median rostral swellings. Specimens of R. stheno from Thailand in the collection of The Natural History Museum also proved to belong to the new subspe- cies. It seems possible also, that specimens recorded by Osgood (1932: 219) refer to the same subspecies as described here. His specimens, listed as ‘Rhinolophus sp.’, derived from Tonkin (North Vietnam) and Osgood stated that *. . . it is possible that the present [form] is a northern representative of the larger Malayan form stheno’. An alternative suggestion, that Osgood’s specimens might be referable to R. borneensis was, however, made by Hill & Thonglongya (1972). This supposition is equally probable, as af- firmed by specimens of R. borneensis which were also collected during the recent expeditions to Vietnam. It appears likely that the section on R. stheno in Lekagul & McNeely (1977) also refers to the new subspecies; unfortunately, however the accompanying photo- graph is of a specimen in which the diagnostic characters are not visible on the damaged rostrum. According to the literature, R. stheno and the closely related R. malayanus may be distinguished by the shape of the rostral swell- ings. On the basis of our data set, the width of the interorbital constriction also distinguishes the two species (Table 1). As regards the external characters, according to Koopman (1994) there is a definite gap between the two species in forearm length (45— 48 mm against 40-43 mm) but McFarlane & Blood (1986) concluded ‘that there is a probability of overlap between specimens of the two species’. Indeed, during the examination of larger series derived from different geographical regions only very slight differences may be observed between extreme values of forearm length of small R. stheno and large R. malayanus. Furthermore the ratio of first to second phalange of the third digit in R. s. microglobossus shows overlap in size between the smaller R. malayanus and the larger R. s. stheno, as figured by McFarlane & Blood (1986). ACKNOWLEDGEMENTS. We are very grateful the Society for Environ- mental Exploration — Vietnam, especially the co-ordinators Mike Hill and Neville Kemp assisted by Dung Ngoc Can of the Institute of Ecology and Biological Resources, Hanoi, but also the other members involved in the bat surveys, including Maysie Harrison, Daniel Fitton, Vicky Jenkins and Joe Walston, and to Pham Duc Tien for his generous help in the field. We are indebted to Prof. Cao Van Sung, Institute of Ecology and Biological Re- sources, Hanoi for organising and co-ordinating the research programs in Vietnam, to Dr. Gyérgy Topal for his helpful advice during the preparation of the manuscript, to Dr. Jacques Cuisin, Muséum National d’ Histoire Naturelle, Paris for access to the collection and to Péter Ujhelyi for the drawings. In particular we thank Dr. Wieslaw Bogdanowicz, Polish Academy of Sciences for his very helpful review comments and Clive Moncrieff, NHM for his constructive criticism of the statistical methodology. The work of GCS was supported by the Royal Society of London and the Hungarian National Scientific Fund (OTKA) grant no. F 17700. NEW SUBSPECIES OF RHINOLOPHUS STHENO REFERENCES Andersen, K. 1905. On some bats of the genus Rhinolophus, with remarks on their mutual affinities, and descriptions of twenty-six new forms. Proceedings of the Zoological Society of London, 2: 75-145. Bogdanowicz, W. 1992. Phenetic relationships among bats of the family Rhinolophidae. Acta Theriologica 37: 213-240. Bonhote, J. L. 1903. Report on the mammals. Fasciculi Malayenses. Zoology 1: |-45, 4 pls. Corbet, G. B. & Hill, J. E. 1992. The mammals of the Indomalayan region: a systematic review. Natural History Museum Publications, Oxford University Press, London, 488 pp. Csorba, G., Fuisz, T. & Kelen, B. 1997. New birds and bats from Pulau Tioman, Pahang, Malaysia. Malayan Nature Journal, 50: 197-200. Hill, J. E. & Thonglongya, K. 1972. Bats from Thailand and Cambodia. Bulletin of the British Museum (Natural History) Zoology 22: 173-196. Hill, M. & Kemp, N. 1996. Biological survey of Na Hang Nature Reserve, Tuyen 21 Quang Province, Vietnam. Part 1; Ban Bung sector. S.E.E. — Vietnam Scientific Report No. 1. Hanoi, 45 pp. Koopman, K. F. 1994. Chiroptera: systematics. Handbook of zoology. Mammalia, part 60. Walter de Gruyter, Berlin, 217 pp. Lekagul, B. & McNeely, J. A. 1977. Mammals of Thailand. Association Conservation Wildlife, Bangkok, 758 pp. McFarlane, D. A. & Blood, B. R. 1986. Taxonomic notes on a collection of Rhinolophidae (Chiroptera) from Northern Thailand, with a description of a new subspecies of Rhinolophus robinsoni. Zeitschrift fiir Sdugetierkunde 51: 218-223. Osgood, W. H. 1932. Mammals of the Kelley-Roosevelt and Delacour Asiatic expedi- tions. Publications Field Museum of Natural History. Zoology 18: 193-339. Peters, W. 1861. Uber die von Hrn. F. Jagor bisher auf Malacca, Borneo, Java und den Philippinen gesammelten Saugethiere aus den Ordnungen der Halbaffen, Pelzflatterer und Flederthiere. Monatsberichte der Koniglichen Preussischen Akademie Wissenschaften zu Berlin: 706-712 Tate, G. H. H. & Archbold, R. 1939. Results of the Archbold Expeditions. No. 24. Oriental Rhinolophus, with special reference to material from the Archbold Collec- tions. American Museum Novitates. (1036): 1-12. No. 2 Volume 59 No. 1 No. 2 Volume 60 No. | No. 2 Volume 61 No. | No. 2 Notes on the anatomy and classification of ophidiiform fishes with particular reference to the abyssal genus Acanthonus Giinther, 1878. G. J. Howes Morphology and morphogenesis of the soil ciliate Bakuella edaphoni noy. spec. and revision of the genus Bakuella Agamaliev & Alekperov, 1976 (Ciliophora, Hypotrichida). W. Song, N. Wilbert and H. Berger A new genus and species of freshwater crab from Cameroon, West Africa (Crustacea, Brachyura, Potamoidea, Potamonautidae). N. Cumberlidge and P. F. Clark On the discovery of the male of Mormonilla Giesbrecht, 1891 (Copepoda: Mormonilloida) R. Huys, G. A. Boxshall and R. Bottger-Schnack £40.30 A new snake from St Lucia, West Indies. G. Underwood Anatomy of the Melanonidae (Teleostei: Gadiformes), with comments on its phylogenetic relationships. G. J. Howes A review of the serranochromine cichlid fish genera Pharyngochromis, Sargochromis, Serranochromis and Chetia (Teleostei: Labroidei). P. H. Greenwood A revision of Danielssenia Boeck and Psammis Sars with the establishment of two new genera Archisenia and Bathypsammis (Harpacticoida: Paranannopidae). R. Huys and J. M. Gee. A new species of Syrticola Willems & Claeys, 1982 (Cope- poda: Harpacticoida) from Japan with notes on the type species. R. Huys and S. Ohtsuka Erratum £40.30 The status of the Persian Gulf sea snake Hydrophis lapemoides (Gray, 1849) (Serpentes, Hydrophiidae). A. Redsted Rasmussen. 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