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<ecenant?erescsearscceadesr-teceanee 189

HE ICR GN GLO DI SHIDTARIGS CALLS USP) WAL O Mav a etas couasevaveeettcanacpeOeeccate es epPaeeai aces na cates ectaneen cect eadacasnovatesdacsanteatotsnecutsedecsssesenyesteceatescnacenestescosstats 195

ea CV ENO IS IPO GIG SIIMIIO VOR PEW ss i coaci vacicecausetaagenh Rosebe teen can tetcaaaccadecdedon eentavavstdesastost ceases teaakaistaeaceascesacees laaeUacesccseecbeaceeotanes 200

PES KILO MILE CLG CLIMEML US Mere tect Re seaentcuasacesusee wees a: fdne MUeo- Crem eweeke saa wustabenve Pees reece sods tava cebu ce suevsvevectaelasitestestthed Meme eee aeegneere ee 200

FREE CT ENG CS 2a sea cas ctet- nan ancss-n ave ccsn snans¥ceast teaohaemune Weg Wacis dunadedersoaadgagere ded aduasbasuctsetunsksudcbuceeeekesuaskedsstuavacteeius seve seaduvaeSetmes 204

Synopsis. A partial revision of the genus Paracyclops is made based on type material and on collections from numerous

localities around the world. The following Paracyclops species are redescribed: P. affinis (G.O.Sars, 1863), P. poppei (Rehberg,

1880), PR oligarthrus (G.O.Sars, 1909), P. canadensis (Willey, 1934), P. dilatatus Lindberg, 1952, P. hardingi nomen novum, P.

baicalensis Mazepova, 1961, P. yeatmani Daggett & Davis, 1974, P. waiariki Lewis,1974, P. pilosus Dussart, 1984, P. carectum

Reid, 1987, PR novenarius Reid, 1987 and P.smileyi Strayer, 1988. Four species are described as new to science: P. reidae sp. nov.,

P. rochai sp. noy., P. punctatus sp. nov., and P. bromeliacola sp. nov.

Detailed descriptions of these species are given including several previously overlooked microcharacters, such as the

ornamentation of the coxobasis of antenna, the cuticular ornamentation of urosomal somites and the posterior spinular

ornamentation of the swimming legs, that are shown to have significant taxonomic value at species level. The detailed description

of males is revealed to be important in differentiating between closely related species.

The geographical distributions of the species are re-evaluated on the basis of examined material and verifiable published

records. It is revealed that P. affinis does not occur in North America and all previous records of P. affinis in North America refer

to the newly discovered P. canadensis.

26 November 1998

HISTORY MUSEUM

1S NOV 1998

PRESENTED

RARY

SSA

INTRODUCTION

The genus Paracyclops Claus, 1893 is one of nine genera currently

recognised as constituting the sub-family Eucyclopinae (Dussart &

Defaye, 1985; Pospisil & Stoch, 1997). All species are known to be

benthic although they can sometimes occur in the water column in

the littoral zone. Paracyclops species are distributed worldwide and

have been recorded in all types of freshwater habitats (Karaytug,

1998): P. dilatatus Lindberg, 1952 was found in the Dniester estuary

(Ukraine) on the Black Sea (Monchenko, 1977), P. baicalensis

Mazepova, 1961 was collected from great depths in Lake Baikal

(Mazepova, 1978), and P. bromeliacola sp. nov. and P. reidae sp.

noy. inhabit pools in the leaf axils of terrestrial Bromeliads. P.

chiltoni (Thomson, 1882) was recently collected from freshwater

bodies on Easter Island and is the only freshwater copepod on this

S. KARAYTUG AND G.A. BOXSHALL

112

_— ee

= aa <a ——

Z Sere = See

ad, ——————

SES SS

( =

\ —

Z ; c ae ree x dt

ZL i

SK = . \ ‘

1 : par 1 Fm

= as

Sy cS

7 ee

ES an

bee

\ SS

j =

Uy, >

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.

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

\ See

= =

SSS ee

Any

IN = S

ne ; ee t c= ay

i MA SS >

\ aie “ ay,

\) N

Coy.

MW |

gj oil y rw

4 aha yy Wy

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

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

Z aa

ntee aa

ppp tL ee aa ee

LE LEE pare c= =

————— Zs

aan

WS 3)

NES

VAG

gts

. &

a. ISS mT

S wa aaaadadalll ;

=S>

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

eee Se

}

. Oe a ae ASS 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

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

A 7

AN Vif

owe

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 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

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

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

\ i J Fan

by I

. ar (-

a a

‘ Va \\

j y)

gp ROOT

ow \

SSS

eee Loe

SSS R Se :

LLLP Ie ye ee,

ABF|5

SSS SSS

ues

| Wi

iA hunny t

: i iN

IN j\

| NN

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

128 S. KARAYTUG AND G.A. BOXSHALL

rap

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)

¥ : 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

UU,

\ Sr A

NO WH Vis

\ by

hy,

S. KARAYTUG AND G.A. BOXSHALL

132

—— SS SSS SN

SS oO SS

Var ST

amit Wai G

Fig. 16 P. oligarthrus. Adult female. A, antennule; B, antenna, frontal; C, antenna, coxobasis, caudal. Scale bars in um.

133

PARACYCLOPS REVISION

yg,

A 8

=) ey

BSS

> |

Abhay

‘4

= WS

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

Naw

NL an abbd Dy

INS

——s

S = & fe °

WGK ¥ oS Senes

Rs ea \\ S a 50

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.

eat

i / MW , Vi

ro)

x es SS

& a

\ SE Wes eS Me NV

S Se SE His

\\ = Q\

Na = \

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

—

“

( f

4 A

ane

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

wd wf ‘/ HED

Pal id Maelpinp ee h

i TE i

; 4 yy rt ene

a OP anges

ee >

Vi Ons

WL jm ¢

144, <i 4 i Yi),

i i

wr FPN PbO

a =

pr ee

= Zac.

= ere

ae ke eee

: =

)

| Fig. 24 P. canadensis. Adult female. A, leg 3, anterior; B, intercoxal sclerite of leg 3, posterior; C, intercoxal sclerite and coxa of leg 4, posterior; D, leg 4,

| anterior. Scale bar in um.

142 S. KARAYTUG AND G.A. BOXSHALL

: \ ney |

Wy mi We

Tir inl |

Fig. 25 P. canadensis. Adult male. A, urosome, ventral; B, urosome, dorsal; C, antennule showing proximal segmentation, ventral; D, antennule, detail of

segments 11 to 14, ventral. Scale bars in um.

PARACYCLOPS REVISION

Antennule 8-segmented (Figure 26A,H); third segment with par-

tial suture line. Coxobasis of antenna with spinular row near base of

inner setae on caudal surface (arrowed in Figure 26C); second

endopodal segment with 9 setae, one seta transformed into massive

recurved claw (arrowed in Figure 26C); third endopodal segment

armed with 7 setae around apex; 2 of which modified into claw-like

setae (arrowed in Figure 26C).

Spine and seta formula of swimming legs (Figure 27A—D) as

follows:

Coxa Basis Exopod Endopod

Leg | 0-1 1-I I-1;1-1;101,5 0-1;0-1;1,1,4

Leg 2 0-1 2-0 I-1;1-1;10,1,5 0-1;0-2;1,1,4

Leg 3 0-I 1-0 I-1;1-1;11,1,5 0-1;0—2;1,1,4

Leg 4 0-I 1-0 I-1;I-1;1L,15 0-1;0—2;1,11,2

Leg 5 (Figure 26L) comprising single free segment, armed with |

long (almost twice as long as inner spine) outer spinulose seta, |

inner spine surrounded with spinules at base, and 1 plumose seta

centrally.

ADULT MALE. Unknown.

DIFFERENTIAL DIAGNOSIS. P. dilatatus is remarkable because of

the highly transformed seta which forms the massive claw on the

second endopodal segment of the antenna (arrowed in Figure 26C).

No other Paracyclops species possesses such a modified seta on this

segment.

REMARKS AND COMPARISONS

Sars (1927a) initially published Platycyclops dilatatus as a nomen

nudum but his death prevented him from describing the new species.

In 1952 Lindberg, on the basis of Sars’s specimens, described this

taxon under the name Paracyclops dilatatus. Its incomplete and

inaccurate description as P. dilatatus (Lindberg, 1952) led

Monchenko (1977) to describe a new subspecies from the Black Sea

without examining the type specimens. Lindberg’s (1952) interpre-

tation of the antennule as 10-segmented is wrong since the

examination of type material left no doubt that P. dilatatus has an 8-

segmented antennule. Lindberg (1952) admitted that that it was

difficult to examine and measure the specimens in his original

publication on P. dilatatus. However, Monchenko (1977) estab-

lished his new subspecies of P. dilatatus on the basis of having an

8-segmented antennule. The other main morphological characters of

the caudal rami and its setal elements are so similar to the type that

the description of anew subspecies P. dilatatus ivanegai Monchenko,

1977 is unjustified. Monchenko’s description should be considered

to be the first good redescription of P. dilatatus Lindberg, 1952.

DISTRIBUTION. P. dilatatus is known only from the Caspian Sea

and the Black Sea basin in Ukraine.

Paracyclops hardingi nom. nov.

(Figures 28-32)

Paracyclops fimbriatus andinus Lindberg, 1957

non Paracyclops andinus Kiefer, 1957

ORIGINAL DESCRIPTION. Paracyclops fimbriatus andinus Lindberg,

1957: Folia Biol. Andina, 1: 39-52.

TYPELOCALITY. Lindberg’s material (Lindberg, 1957) came from

two different sites in Peru, one from Lake Huampucocha and the

other from Lake Conococha. Since Lindberg did not specify on

143

which material his description was based, therefore both lakes are

type localities.

MATERIAL EXAMINED

Since Lindberg’s material of P fimbriatus andinus has not been

obtained, some material collected (originally identified under the

name P. finitimus) during The Percy Sladen Trust Expedition to Lake

Titicaca in 1937 under the leadership of Mr H. Cary Gilson was used

to describe P hardingi. A series of collections taken during the

expedition is stored in The Natural History Museum, London.

According to Harding (1955), the localities for the P hardingi

examined in this study are as follows: the shores of Taman Bay,

Laguna Arapa, Laguna Umayo and the Lagunillas, from springs by

the Lagunillas, from the River Ramis and from a ditch by the River

Urubamba. These localities are mostly in the Altiplano surrounding

Lake Titicaca. The examined material is 2 9 9 (PFH 227/2), 1 9 (G/

G 1/93/5), 39 9(PFH 245), 32 Qand 1c°(PFH 139); BMNH

1946. 11. 26. 216-225.

REDESCRIPTION OF ADULT FEMALE

Body length (um) not including caudal setae, 894-1129, mean =

975, n = 10. Genital double-somite with surface ridge extending

either side of copulatory pore on ventral surface (Figure 28A).

Urosomal somites without surface pits (Figure 28A,B). Seminal

receptacle divided into small conical anterior and broad posterior

lobes as figured (Figure 28A). Anal operculum broad and smooth.

Caudal rami parallel and short, about 2.1 times longer than broad;

anterolateral seta (II) long; terminal accessory seta plumose (VI) and

1.5 times longer than posterolateral seta (III); outer terminal seta

(IV) and inner terminal seta (V) well developed, spinulose and

homogeneously ornamented (Figure 28B).

Antennule 8-segmented (Figure 29A); segment 3 with partial

suture line reaching nearly to outer margin of segment, and with

spiniform seta. Setal formula 8, 12, 6, 5, 2 + aesthetasc, 2, 2 +

aesthetasc, 7 + aesthetasc. Coxobasis of antenna with complex

ornamentation on caudal and frontal surfaces as figured (Fig. 29B,C),

and with well-developed spinular row near base of two inner setae

(arrowed in Figure 29B).

Intercoxal sclerite of leg | ornamented with spinular row on

anterior surface (Figure 30B), lacking spinules on posterior surface

(Figure 30A). Intercoxal sclerite of leg 2 ornamented with spinular

row on both anterior and posterior surfaces (Figure 30D,E).

Intercoxal sclerite of leg 3 without spinular row on anterior

surface (Figure 31D) and with 2 spinular rows on posterior surface

(Figure 31C). Intercoxal sclerite of leg 4 (Figure 31B) with groups

of spinules on posterior surface; first and second exopodal segments

without spinular row on posterior surface.

Spine and seta formula as follows:

Coxa Basis Exopod Endopod

Leg | 0-1 1-I I-1;1-1;111,5 0-1;0-1;1,L4

Leg 2 0-1 1-0 I-1;J-1;01,1,5 0-1;0—2:1,1,4

Leg 3 0-1 1-0 I-1;1-1;1,1,5 0-1;0-2:;1,1,4

Leg 4 0-I 1-0 I-1;I-1;IL1,5 0-1;0-2;1, 11,2

Leg 5 (Figure 28C) comprising single free segment, armed with

1 long (almost 1.5 times longer than inner spine) well-developed

outer spinulose seta, | inner spine, | plumose seta in centre. Leg 6

(Figure 28B) represented by | plumose seta and 2 tiny spinules

dorsolaterally

DESCRIPTION OF ADULT MALE

Urosomal somites (Figure 32A,B) without ornamentation of surface

144 S. KARAYTUG AND G.A. BOXSHALL

Pramnn? OVO

E %

—

= KENUS XS&\

Fig. 26 P. dilatatus. Adult female. A, antennule; B, labrum; C, antenna, caudal; D, maxillulary palp; E, maxillule praecoxal armature, distal; F, maxilla,

distal part, apical. G, anal somite and caudal rami, lateral; H, body, ventral; K, anal somite and caudal rami, dorsal. L, leg 5, lateral; Scale bars in um.

PARACYCLOPS REVISION

145

ma,

* Git: BZ

‘\

és ss |

a y ' NGS

1 ae ras \ :

Fig. 27 P dilatatus. Adult female. A, leg 1 with inset showing exopod, anterior; B, leg 2, anterior; C; leg 4, anterior; D; leg 3, anterior. Scale bar in um

146

pits. Caudal rami shorter than female. First antennulary segment

armed with 8 setae plus an aesthetasc, | seta large (seta A) and

modified.

Coxobasis of antenna with spinular row on caudal surface near

base of two inner setae (arrowed in Figure 32D); spinules more

elongate than those of female. One seta on terminal exopodal

segment of leg 1 spinulose (Figure 30C). Outer seta of fifth leg

(Figure 32C) plumose and less developed than that of female. Sixth

leg (Figure 32B,C) armed with 1 inner spine, longer than second

urosomal somite, and 2 outer plumose setae.

DIFFERENTIAL DIAGNOSIS. P. hardingi differs from other Para-

cyclops species by the combination of the following characters; the

presence of the spinular row on the caudal surface near the base of

the 2 inner setae (arrowed in Figures 29B; 32D) of the coxobasis of

the antenna in both sexes; by the structure of the seminal receptacle

(Figure 28A); by the absence of the spinular rows on the posterior

surface of the first and second exopodal segments of leg 4 (Figure

31A); by the wide anal operculum (Figure 28B); by the length of the

anterolateral seta (II) on the caudal ramus (Figure 28B); and by the

absence of cuticular pits from the urosomal somites in both sexes

(Figures 28A,B; 32A,B).

REMARKS

P. hardingi, P. altissimus Karaytug, Boxshall & Defaye (in press),

P. longispina Karaytug, Boxshall & Defaye (in press) and P.

imminutus Kiefer 1929 are closely related. All four species pos-

sess a well-developed spinular row near the base of the two inner

setae on the coxobasis of the antenna in both sexes (arrowed in

Figures 29B; 32D). P. hardingi can easily be differentiated from P.

altissimus by the length and spinulation of the outer seta of leg 5,

the structure of the seminal receptacle (Figure 28A), the presence

of the mid-distal spinular rows on the posterior surface of the coxa

of legs 1-3, and the relative length of the anterolateral seta (II) on

the caudal ramus (Figure 28B). It differs from P imminutus by

having a shorter outer seta of leg 5 (Figure 28C), in the structure

of seminal receptacle, the position of the mid-distal spinular row

on the posterior surface of the coxa of leg 1, the absence of the

surface pits on the genital somite and urosomal somites in the

male, and the length of the anterolateral seta (II) on the caudal

ramus. P. hardingi can be differentiated from P. longispina by the

shorter outer seta of leg 5, the absence of the surface pits on the

genital somite and urosomal somites in the male, the presence of

the aesthetasc on the first segment of the male antennule, the

structure of the seminal receptacle, and the length of the anterola-

teral seta (II) on the caudal ramus.

P. hardingi was first described by Lindberg (1957) under the name

P. fimbriatus andinus, but Kiefer used the name P. andinus (Kiefer,

1957) earlier in the same year. Therefore, Lindberg’s P. fimbriatus

andinus and Kiefer’s P. andinus are primary homonyms. According

to the priority principle P andinus Kiefer, 1957, published 1 March

1957, takes precedence over P. fimbriatus andinus Lindberg, 1957

published on 10 July 1957. The name P. fimbriatus andinus is a

junior homonym and 1s invalid (Article 52 (b)). P fimbriatus andinus

Lindberg, 1957 must be replaced by a new name (Article 60).

ETYMOLOGY. The new name has been given in honour of the late

Dr. J. P. Harding.

DISTRIBUTION. P. hardingi was recorded only once, by Loffler

(1963), under the name P. fimbriatus andinus Lindberg, 1957 from

Ecuador, since its original description from Peru.

Paracyclops baicalensis Mazepova, 1961

(Figures 33-37)

S. KARAYTUG AND G.A. BOXSHALL

ORIGINALDESCRIPTION. Paracyclopsfimbriatus baicalensis Maze-

pova, 1961: Trud. limnol. Inst. Moscou, 2,22: 172-195 (p.177, fig. 2).

TYPE LOCALITY. Russia, Lake Baikal

TYPE MATERIAL.

material.

Mazepova (1961) did not designate any type

MATERIAL EXAMINED. The redescription was based on two

topotypic females, collected on 19-20 August 1990 from a depth of

200 and 300 m in Lake Baikal.

REDESCRIPTION OF ADULT FEMALE

Body length excluding caudal setae 788—983 um, mean = 886 Um,

n = 2. Fifth pedigerous somite without fringe of elongate setules at

posterior margin (arrowed in Figure 34A). Genital double-somite,

second and third abdominal somites without surface pits on dorsal

and ventral surfaces (Figure 34A,B). Seminal receptacle difficult to

observe, anterior lobe apparently small and narrow (Figure 34B).

Anal operculum broad and smooth (Figure 34A,D). Caudal rami

(Figure 34A,B) very short, about 1.3 times longer than broad.

Anterolateral seta (II) long and plumose with spinules at base;

posterolateral seta (III) spinulose with spinular row laterally at base

extending dorsally; terminal accessory seta spinulose (VI); outer

terminal seta (IV) and inner terminal seta (V) well developed and

heterogeneously ornamented (Figure 34B).

Antennule short, compact, 8-segmented (Figure 35A,B): segment

3 with partial suture line and spiniform seta. Segment 5 with

characteristic short aesthetasc. Another short aesthetasc located

distally on anteroventral margin on segment 7. Apical segment with

aesthetasc fused to adjacent seta at base. Most of setal elements

highly spinulated. Setal formula 8, 12, 6, 5, 2 + aesthetasc, 2, 2 +

aesthetasc, 7 + aesthetasc. Coxobasis of antenna (Figure 35C,D)

with complex ornamentation on caudal and frontal surfaces; with

spinular row near base of inner setae on caudal surface (arrowed in

Figure 35D). Setal elements highly spinulated.

Labrum (Figure 33C) broad posteriorly; posterior margin form-

ing strong teeth and ornamented with 2 patches of spinules on frontal

surface; ornamented with paired clusters of long spinules near

posterior margin. Mandible (Figure 33D) consisting of well devel-

oped coxal gnathobase with 3 lateral spinules distally. Palp

represented by 3 spinulose setae, 1 of which very long.

Intercoxal sclerites of legs 1-3 (Figures 36A—D; 37A,B) without

spinular rows on anterior and posterior surfaces. Leg 1 with 3 setae on

terminal segment of exopod semispinulose; 2 setae on terminal

segment of endopod spinulose (arrowed in Figure 36C). Terminal

endopodal spine of leg 2 strong, naked and shorter than segment; three

setae on terminal segment of exopod and | seta on terminal segment of

endopod semispinulose (arrowed in Figure 36A). Leg 3 with 3 setae

on terminal segment of exopod and | seta on terminal segment of

endopod semispinulose (arrowed in Figure 37A). Intercoxal sclerite

of leg 4 without spinules on anterior surface (Figure 37C) and with

spinular row on posterior surface (Figure 37D); second endopodal

segment with 3 spinules on posterior surface; 3 setae on terminal

segment of exopod (one arrowed in Figure 37C) and | setaon terminal

segment of endopod semispinulose and 1 seta on terminal segment of

endopod spinulose (arrowed in Figure 37C).

Spine and seta formula as follows:

Coxa Basis Exopod Endopod

Leg 1 0-1 1-I I-1;1-1;101,5 0-1;0-1;1,1,4

Leg 2 0-I 1-0 I-1;/-1;10,1,5 0-1;0-2;1,1.4

Leg 3 O-I 1-0 -1;-1;0L1,5 0-1;0-2;1,1,4

Leg 4 0-I 1-0 I-1;I-1;111,5 0-1;0—2;1,11,2

147

PARACYCLOPS REVISION

SSSSAANSSSSSK SSSSSSSSASSS

YW plo

SSS

= =

—

Fig. 28 P. hardingi. Adult female. A, urosome, ventral; B, urosome, dorsal; C, leg 5, ventral. Scale bars in um.

148 S. KARAYTUG AND G.A. BOXSHALL

—e a

Ses

WSsy,

= Se

Fig. 29 P. hardingi. Adult female. A, antennule; B, antenna, caudal; C, antenna, coxobasis, frontal. Scale bars in um.

PARACYCLOPS REVISION

;

net

a CE

WHY

pers

’

;

}

{ ‘

\yast Awl?

(ld 54d \ ia

(

149

Fig. 30 P hardingi. Adult female. A, intercoxal sclerite and coxa of leg 1, posterior; B, leg 1, anterior; C, adult male, terminal endopodal segment of leg 1;

D, leg 2, anterior; E, intercoxal sclerite and coxa of leg 2, posterior. Scale bars in lum.

150 S. KARAYTUG AND G.A. BOXSHALL

“404 Mh

Fig. 31 P. hardingi. Adult female. A, leg 4, anterior; B, intercoxal sclerite and coxa of leg 4, posterior; C, intercoxal sclerite and coxa of leg 3, posterior;

D, leg 3, anterior. Scale bar in um.

PARACYCLOPS REVISION 151

\ aml

Afi

4) Nw Ny

yi | in

ha Nj Xa

} Al\

of y |

obi

Fig. 32 P hardingi. Adult male. A, urosome, dorsal; B, urosome, ventral; C, detail of leg 5, ventral; D, antenna, coxobasis, caudal. Scale bars in [um.

152

Leg 5 (Figure 34C) comprising single free segment, armed with 1

long (almost twice as long as inner spine) well developed outer

spinulose seta, | serrate-like strong inner spine, one spinulose seta in

middle (slightly shorter than outer seta). Leg 6 (Figure 34A) repre-

sented by | plumose seta and 2 tiny spinules dorsolaterally

ADULT MALE. Unavailable for redescription.

DIFFERENTIAL DIAGNOSIS. This endemic Baikalian species can

easily be distinguished from other Paracyclops species by the

absence of the fringe of elongate setules (arrowed in Figure 34A)

usually present around the posterior margin of the fifth pedigerous

somite in the female; by its very short caudal rami (Figure 34A,B),

by the structure of leg 5 (Figure 34C) and by the highly ornamented

setal elements on the female antennule (Figure 35A,B).

REMARKS

This species is remarkable by virtue of the highly ornamented setal

elements on nearly all the appendages. Mazepova (1961) described

this Baikalian endemic as a subspecies but later (Mazepova, 1978)

treated it as a distinct species.

DISTRIBUTION. P. baicalensis is endemic to Lake Baikal.

Paracyclops yeatmani Daggett & Davis, 1974

(Figures 38-42)

non Paracyclops yeatmani: Mahoon & Zia, 1985.

ORIGINAL DESCRIPTION. Paracyclops yeatmani Daggett & Davis,

1974: Can. J. Zool., 52, (2) : 301-304.

TYPE LOCALITY. Canada, Newfoundland, Highway 5 between

Bay Bulls and Witless Bay.

TYPE MATERIAL. ‘Type material of P yeatmani was obtained from

the Canadian Museum of Nature. CMNC1984-1121, paratypes, 2

microscope slides of 20’C’dissected between prosome-urosome.

CMNC 1984-1122, paratypes, | vial, 42 9. 1 9 dissected.

OTHER MATERIAL. 1Qundissected and mounted on one slide;

1 2 dissected and mounted on one slide from the type locality. Dr. H.

Yeatman collection, 11 September 1972.

REDESCRIPTION OF ADULT FEMALE

Body length (um), not including caudal setae, 778-798 (given by

Daggett & Davis as 750-860), mean = 785, n = 3. Body width 301—

331, mean = 319, n = 3. Prosome (Figure 38C) with cephalothorax

longer than 3 free pedigerous somites. Genital double-somite, sec-

ond and third abdominal somites without surface ornamentation

and posterior margins of abdominal somites more conspicuously

serrated ventrally than dorsally (Figure 38A,B). Seminal receptacle

divided into broad anterior and posterior lobes as figured (Figure

38B). Anal somite with spinular row ventrally extending dorsally

midway along either side of anal operculum (Figure 38A,B). Anal

operculum smooth; row of spinules present in anal cleft, either side

of midline. Caudal rami (Figure 38A,B) about 3.1 times longer than

broad; anterolateral seta (II) plumose with spinules originating at

base, extending midway along dorsal surface (Figure 38A); terminal

accessory seta naked (VI) and about 3 times longer than posterola-

teral seta (III). Outer terminal seta (IV) and inner terminal seta (V)

well developed and heterogeneously ornamented (Figure 38C).

Antennule 11-segmented (Figure 40A). Segment 3 with partial

suture line. Segment 5 with spiniform seta. Segment 8 with short

aesthetasc (see inset Figure 40A). Apical segment with aesthetasc

fused to adjacent seta at base, and another aesthetasc located distally

on anteroventral margin on segment 10. Setal formula 8, 4, 8, 4, 2, 2,

S. KARAYTUG AND G.A. BOXSHALL

3, 2 + aesthetasc, 2, 2 + aesthetasc, 7 + aesthetasc. Coxobasis of

antenna (Figure 40D,E) with complex ornamentation on caudal and

frontal surfaces as figured.

Proximal segment of maxillulary palp ornamented with 3 minute

spinules (arrowed in Figure 39F). Syncoxa of maxilliped (Figure

39A,B) without spinules near setal bases (arrowed in Figure 39B).

Basis ornamented with 2 transverse spinular rows near outer distal

angle and with 2 irregular spinular rows near bases of medial setae

(arrowed in Figure 39A). First endopodal segment with 4 spinules.

Coxa of leg 1 (Figure 41C) with spinular row near outer margin on

posterior surface; intercoxal sclerite without spinular rows on ante-

rior and posterior surfaces. Intercoxal sclerite of leg 2 (Figure 42A)

ornamented with spinular rows on anterior and posterior surfaces;

coxa with spinular row near outer margin on posterior surface; first

endopodal segment with spinular row on posterior surface. Intercoxal

sclerite of leg 3 (Figure 42B,C) with spinular row on anterior surface

and with 3 spinular rows on posterior surface; coxa with spinular

row near outer margin on posterior surface; first endopodal segment

with spinular row on posterior surface. Intercoxal sclerite of leg 4

with few spinules on anterior surface (Figure 41B) and with 3 long

spinular rows on posterior surface (Figure 41A); inner coxal seta

with group of setules mainly originating posteriorly; coxa with

complex ornamentation on posterior surface as figured (Figure

41A); basis with setules along inner margin.

Spine and seta formula as follows:

Coxa Basis Exopod Endopod

Leg | 0-1 1-I I-1;1-1;111,5 0-1;0-2;1,1.4

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;I-1;0,1,5 0-1;0-2;1,1,4

Leg 4 O-I 1-0 I-1;1-1;11,1,5 0-10-21, 11-2

Leg 5 (Figure 38D) comprising single free segment, armed with

short outer plumose seta (shorter than inner spine), 1 serrate-like

strong inner spine, and | plumose seta in middle about twice as long

as inner spine. Base of middle seta produced. Leg 6 (Figure 38A)

represented by | long plumose seta and 2 tiny spinules dorsolaterally.

DESCRIPTION OF ADULT MALE

Antennule (Figure 40C) 16-segmented. The poor condition of the

slides and the orientation of the antennule made it impossible to

confirm all details of the setation pattern.

DIFFERENTIAL DIAGNOSIS. P. yeatmani can easily be differentiated

from other Paracyclops species by the combination of the following

characters: the produced base of the middle seta of leg 5 (Figure

38D), the spinules originating at the base of the anterolateral seta

(II), extending midway along the dorsal surface of caudal rami

(Figure 38A), the very long, naked terminal accessory seta which is

about 3 times longer than the posterolateral seta (Figure 38A), the

three rows of long spinules on the posterior surface of intercoxal

sclerite of leg 4 (Figure 41A) and its 11-segmented antennule.

REMARKS

Paracyclops yeatmani Mahoon & Zia, 1985 is a junior primary

homonym of P. yeatmani Dagget & Davis, 1974 and therefore an

invalid name (ICZN Article 57 (b)). This species was based on

juvenile stages and belongs to a species not related to P. yeatmani

Dagget & Davis, 1974. It is regarded here as species incertae sedis

in the Cyclopidae.

There are only three species with 1 1-segmented antennules in the

genus Paracyclops; the other two being P. affinis and P. canadensis.

However, the 1 1-segmented state is not homologous in P. yeatmani

and in P. affinis-canadensis group. Segments 3 and 4 (ancestral

PARACYCLOPS REVISION

4. a

\\ eI

= \ m App

Sih 7

Woai( (=

Tt S

Fig. 33 P. baicalensis. Adult female. A, maxilla; B, body, dorsal; C, labrum; D, mandible: E, maxillule; F, maxilliped. Scale bars in tm.

S. KARAYTUG AND G.A. B

154

fs}

—S E

thn :

BBE :

Zz t Sen RELZZ =

(a)

Eo CoS 4 =

Le { (EN

x wax

MG |

—— We <

~ SSS QS SSS

—————

1595)

PARACYCLOPS REVISION

[\-

Z t

§ oS

gment 2 showing setation; C, antenna,

Fig. 35 P. baicalensis. Adult female. A, antennule, with setation omitted from segment 2; B, antennule, se

coxobasis, frontal; D, antenna, caudal. Scale bar in um.

S. KARAYTUG AND G.A. BOXSHALL

156

)

CD50

\ a

' ~~

Wes

Hh} LM GY S N\

IE : : 4 tite

A Ky Z <

= 4

Gi; Za

‘

G \\Y

7 7.) . ¥

i *\\s DNS \

Wk, NN ANS

y IN ON

i\e

a (TE met

AYU

al sclerite and coxa

tercox,

ior; D, in

of leg 1, posterior; C, leg 1, anter

xal sclerite and coxa

P. baicalens

Fig. 36

is. Adult female. A, leg 2, anterior; B, interco

rior. Sca

um.

bars in

of leg 2, poste

PARACYCLOPS REVISION

A Veal f

Z 73 Yip = = ~ Sa —— :

=e WZ) a

a SSS

y lp eo.

, | or en

i i} ny

y ii

WA Ay) N

% TAM!' Gl) WI, \\

_@®© ZI4R An! \

/ Sra fi! Zi! _Zay \\tIN X

| of | \te \

AVN |

yy ig

—

AY \

ARY\

WM \ 4

dd \X)

yey 3 aa Va

aM y

SQ v1

NaN COC Lm

Fig. 37 P. baicalensis. Adult female. A, leg 3, anterior; B, intercoxal sclerite and coxa of leg 3, posterior; C, leg 4, anterior; D, intercoxal sclerite and coxa

of leg 4. Scale bars in um.

158 S. KARAYTUG AND G.A. BOXSHALL

Fig. 38 P yeatmani. Adult female. A, urosome, dorsal; B, urosome, ventral; C, body, dorsal; D, leg 5, ventral. Scale bars in um.

159

PARACYCLOPS REVISION

)

én

ABCF

Fig. 39 P. yeatmani. Adult female. A, B, maxilliped; C, maxilla; D, labrum; E, maxillule: F, maxillulary palp; G, mandible. Scale bars in um.

S. KARAYTUG AND G.A. BOXSHALL

160

tion, dorsal; D, adult female,

le showing segmenta

le; B, adult male

ntenna, coxobasis, frontal; E, adult female, antenna, cau

Fig. 40 P. yeatmani. A, adult female, antennu

, urosome, dorsal; C, adult male, antennu

dal. Scale bars in um.

PARACYCLOPS REVISION

tel ices Wig SS

pin viv wns, Noell! 1X

Gs? A foun

Nae \

wm i Will

m\\ m

2. 7a

“= GZ \ &

Fig. 41 P. yeatmani. Adult female. A, intercoxal sclerite, coxa and basis of leg 4, posterior; B, leg 4, anterior; C, leg 1, anterior. Scale bar in um.

Sil

ZL 1}

—

- sal FR j

went \

jr Ne

a NATE

ai i

—

= c —w

aa \ < =

Dodd dg dy) SOND Wy

\ aS WH.

X id & WMMMh) 4

Mill \ Nee

al g

It female. A, leg 2, anterior; B, intercoxal sclerite of leg 3, posterior; C, leg 3, anterior. Scale bar in im.

PARACYCLOPS REVISION

segments VIII and IX—XI) of P. affinis have failed to separate in P.

yeatmani whereas segments 8 and 9 (ancestral segments XXI—XXIII

and XXIV) of P yeatmani have failed to separate in P. affinis

(Karaytug & Boxshall, 1998). The 11-segmented states are, there-

fore, convergent.

Daggget & Davis (1974) mentioned that there is a seta swollen at

its base on the first segment of the antennule of male P. yeatmani. It

would be remarkable should this seta be homologous with modified

seta (A) of the Paracyclops fimbriatus-group since modified seta (A)

is an important synapomorphy of that group which is only distantly

related to P yeatmani.

In the original description only one inner margin seta was re-

ported from the second endopodal segment of leg 1, but 2 setae were

observed in all material examined, including the paratypes. It is

likely that the presence of this seta was overlooked in the original

description

DISTRIBUTION. Canada, Newfoundland, Highway 5 between Bay

Bulls and Witless Bay (Daggett & Davis, 1974; Daggett & Davis,

1975). U.S.A: Wisconsin, no locality specified (Torke, 1979).

Paracyclops waiariki Lewis, 1974

(Figures 43-48)

ORIGINAL DESCRIPTION. Paracyclops waiariki Lewis, 1974: New

Zealand J. Freshwat. Res., 8 (2) : 275-281.

TYPE LOCALITY. New Zealand. Details of the type locality were

given by Lewis (1974) as follows: The type specimens were netted

in shallow water (up to 50 cm depth) along the roadside edge of Lake

Rotowhero, beneath Rainbow Mountain, about 15 miles from

Rotorua, just beyond the junction of State Highways 30 and 38, Grid

reference: NZMS | Sheet N85; 835 817. The water temperature was

Pio.

TYPE MATERIAL. Not available.

MATERIAL EXAMINED. Topotypic specimens including 3 adult 2 9 ,

1 cop. IV and | adultc’of P. waiariki were obtained on loan from

Museum of New Zealand Te Papa Tongarewa. 1 adult male and

female dissected. Registration number: MNZ Cr 1928.

REDESCRIPTION OF ADULT FEMALE

Body length, excluding caudal setae 672-938 um (given by Lewis

(1974) as 700-800 um), mean = 805, n = 2. Body width 229-240,

mean = 235, n = 2. Prosome as in Lewis (1974); Rounded appear-

ance of cephalothorax (Figure 43B) due to state of preservation.

Genital double-somite, second and third abdominal somites (Figure

44A,B) without ornamentation of pits on dorsal and ventral surfaces.

Posterior margins of abdominal somites more conspicuously ser-

rated ventrally than dorsally. Seminal receptacle divided into

butterfly-shaped anterior and posterior lobes as figured (Figure

44B). Anal somite with spinular row ventrally, extending dorsally

and with 2 small spinules on midsection of ventral surface (Figure

44B). Anal operculum broad and smooth (Figure 44A). Caudal rami

(Figure 44A,B) parallel, about 4.2 times longer than broad;

ornamented with fine spinules along dorsal and ventral surfaces;

anterolateral seta (II) on dorsolateral surface with spinules at base;

posterolateral seta (III) unilaterally plumose, surrounded with

spinules along dorsal surface and with spinular row around base

ventrally, extending dorsally; terminal accessory seta (VI) plumose

and slightly longer than posterolateral seta; outer terminal seta (IV)

and inner terminal seta (V) well developed and heterogeneously

ornamented (Figure 43B,E).

Antennule 12-segmented (Figure 43A); segment 6 with spiniform

163

seta (arrowed in Figure 43A); segment 9 with short aesthetasc

(arrowed in Figure 43A). Setal formula 8, 4, 2, 6, 4, 2, 2, 3,2 +

aesthetasc, 2, 2 + aesthetasc, 7 + aesthetasc. Coxobasis of antenna

(Figure 43C,D) with complex ornamentation on caudal and frontal

surfaces as figured; with spinular row near 2 inner setae on caudal

surface (arrowed in Figure 43C); first endopodal segment with

transverse spinular row on caudal surface (arrowed in Figure 43C).

Labrum (Figure 45D) narrowing posteriorly; posterior margin

with strong teeth; ventral surface ornamented with paired groups of

long spinules. Gnathobasic blades of mandible (Figure 45E) mostly

simple with 3 spinules laterally; palp represented by 3 setae, 2 of

which very long, third short and naked. Proximalmost spine of

maxillule (arrowed in Figure 45C) with spinules. Praecoxa of max-

illa (Figure 45A) with complex spinular rows on dorsal surface; coxa

with group of spinules near base of proximal endite. Syncoxa of

maxilliped (Figure 45B) without long spinules near base of endites;

basis armed with 2 spinulose setae ornamented with 6 long spinules

near base of endites; first endopodal segment with group of long

spinules.

Legs | to 3 without mid-distal spinular row on posterior surface of

coxa. Basis of leg 1 (Figure 46A) with setiform spine on inner

margin reaching almost to end of terminal segment; intercoxal

sclerite without spinular row on posterior surface; seta next to

outermost spine of terminal exopodal segment semispinulose.

Intercoxal sclerite of leg 2 (Figure 46B) ornamented with spinular

rows on anterior and posterior surfaces. Intercoxal sclerite of leg 3

with spinular row on anterior surface (Figure 46E) and with 2

spinular rows on posterior surface (Figure 46F); first endopodal

segment with spinular row on posterior surface.

Intercoxal sclerite of leg 4 (Figure 46D) without spinules on

anterior surface and with 2 irregular spinular rows on posterior

surface (Figure 46C); inner coxal spine with group of setules mainly

originating posteriorly; coxa with complex ornamentation on poste-

rior surface as figured (Figure 46C); basis with setules along inner

margin; first endopodal segment with spinular rows on anterior and

posterior surfaces (Figure 46D).

Spine and seta formula as follows:

Coxa Basis Exopod Endopod

Leg 1 0-1 1-I I-1;I-1;11,5 0-1;0-2;1,1,4

Leg 2 0-I 1-0 I-1;1-1;01,1,5 0-1;0-2;1,1,4

Leg 3 0-1 1-0 J—-1;I-1;01,1,5 0—1;0-2;1,1,4

Leg 4 0-I 1-0 I-0;J-1;11,1,5 O-1;0—2;1, 11,2

Leg 5 (Figure 44C) comprising single free segment, armed with 2

outer setae almost equal in length; strong inner spine shorter than

outer setae. Leg 6 (Figure 44A) represented by | plumose seta and 1

tiny dorsolateral spinule.

DESCRIPTION OF ADULT MALE

Body length, excluding caudal setae = 568 um (given by Lewis

(1974) as 600-700 um), body width = 173 (Figure 47B). Urosomal

somites without surface ornamentation (Figure 47A,C); genital

somite broader than abdominal somites; caudal rami about 3.5 times

as long as broad (Figure 47C,D).

Antennule digeniculate (Figure 48A—E), indistinctly 16-seg-

mented. Segment | armed with 8 setae plus aesthetasc (arrowed in

Figures 48C and E); seta A not modified. Segment 11 bearing small

curved seta ornamented with row of strong denticles, plus 1 plumose

seta (Figure 48B,C). Segmental fusion pattern as follows: I-V, VI-

VU, VII, IX, X, XI, XII, XI, XIV, XV, XVI, XVII, XVIII,

XIX-XX, XXI-XXIII, XXIV-XXVIIL.

164 S. KARAYTUG AND G.A. BOXSHALL

NMG, ~ ay

fone 2d el

Fig. 43. P. waiariki. Adult female. A, antennule; B, body, dorsal; C, antenna, coxobasis, caudal; D, antenna, frontal; E, detail of terminal setal elements of

caudal rami, dorsal. Scale bars in um.

=o ee yy,

on ee

O —————

4 Ze

f :

a € =

m 7

166

leg 6; G, adult male, leg 5. Scale bars in um.

S. KARAYTUG AND G.A. BOXSHALL

Fig. 45 P. waiariki. Adult female. A, maxilla with inset showing accessory armature; B, maxilliped; C, maxillule; D, labrum; E, mandible; F, adult male,

PARACYCLOPS REVISION

@®

] /

pena Z it

ZAI ee

3 aay e SIZ

! Z fe (yy!

ZA / NY es econo

er zl

SS as 7

NY : =

NS Se 5

100

—

NY AQ

J p45), > x \.

lf, fr “thin, jini \\ <

i N/A

} My

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

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,

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

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 )

rg K

a if c =

YS ZF

Y | h be r ‘¢

a Wf

if XG NS

VA \ =

Na\\y

NS a

haa

“ite sael

inne AN

WF ZS ee

Ji =

iN 4. &

| A

Poa SS

/ ; a

| A

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:

SAV) :

SSS

=<)

rea 2

aye

SSS

SS =

Ze ee ee

)

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

Myson IK

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

)

1o)

176

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

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

: = a ee

= ~ Se

a M\

PARACYCLOPS REVISION

=> 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

‘ AN Vs

(

1 Z

SSS

yyperee ve iN

MK B=

Qn sss ===

eee

\ \\ = ; = ST,

\ \\ \S = WSK. KEK<LEE ; LZ

\ See, }

& SS Sa en , f AS

SS GSS A eA Saree Mh

SS She: | TRIN Nee

S-> 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 =

| 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

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.

S. KARAYTUG AND G.A. BOXSHALL

184

——

tm J

Tl [SN

Wd iy

VK -

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

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

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,

‘ wy of Lo sete

f A MF

way Ps KKK

RXHOIOOOO

ae i

yf Y

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

EEE

——

189

PINAR rae

SSssSESTII953

250

100

BC \

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

—

Miao

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

OI =,

Loe 7

‘

QZ —

Py”

= D (a nei Pees

: WENN plassecan y

Wa :

44449,

tg IMU Steytgh sddayy 7

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

= ———

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

ee (CP

5 DSB ate \

\ 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

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

Io)

us}

iS)

i=]

S. KARAYTUG AND G.A. BOXSHALL

202

SOL ey

; fle lec ire =a = s

WL 5 PBL.

“44 astyyy NAAN

AE of 2 os

ule with inset showing variant pattern of segmentation; B, antenna, coxobasis, caudal; C, antenna,

—

UIE eae)

Fig. 75 P. rochai sp. nov. Adult female. A, antenn

coxobasis, frontal; D, terminal endopodal segment of leg 4, anterior; E, same, another specimen; F, adult male, intercoxal sclerite of leg 4, posterior; G,

and basis of leg 4, posterior. Scale bars in [um.

adult female, intercoxal sclerite, coxa

PARACYCLOPS REVISION

Aaron,

Lif

OT hee

SS SS SSS

ZPD SD SoS

ZZ,

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.

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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.

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

GSL

MTL

LIL

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

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

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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.

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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

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A review of the serranochromine cichlid fish genera

Pharyngochromis, Sargochromis, Serranochromis and Chetia

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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

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The status of the Persian Gulf sea snake Hydrophis lapemoides

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Taxonomic revision of some Recent agglutinated foraminifera

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Whittaker.

Foregut anatomy, feeding mechanisms, relationships and

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Association of epaxial musculature with dorsal-fin

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