Bull. nat. Hist. Mus. Lond. (Zool.) 66(1): 148 Issued 29 June 2000

Generic concepts in the Clytemnestridae

(Copepoda, Harpacticoida), revision and

revival

RONY HUYS AND SOPHIE CONROY-DALTON

Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD

CONTENTS

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RUSE INNA LT OG an Ss canna cnn clan cas xeaians denna iin annvis aad sesiganpunsiiceiatnenieawace Sunsuewobs coxa «cae ach duasus thnaadaeanposwenarscbeuntewssartarsesuanseesuiaes'ecnecdseeeas 4

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(GETS GIVICIITICSTN AU As LOA sx ccyceateccnats esp caeea at agatancave suuenndteataderensaian opaciencn ip sasenerupsacrexnesnesnarcnanenss titans (anastetcavansteeaxsts =

CLYte MAESTEG SCULELIGSA DIANA LOT ccesccscencsarczaanspasvossosssessevensattactanserstasnesenvnctuariectnsditaencsseneasccesietesthcsnsaeithseaeescaprsvsitessrone 5

GUSICTIMEST Gi ShAGIES (CLAUS OLS Oca) COMO: DOVE, <ne.acevnssnastectaestauene ne atteraeasanacsacssnoatttarcaumrasesoreaketubesencsuseetestseatnasteceiseoe 15

GIPIZTIMN ESTE TATTGNU SPO Ve con ccactaveseasseocsvevsansescahcoansuacdaciessestese anes teunasecerassncrconeena tac avesUesctave di saseutivedtashasi:suceeohease coasest 20

Clytemnestra longipes SP. NOV. .........:00000+-

GISFETHTE SUT GIGSELOSG:SPNTIOV.creresscssceccccrerncedss-sevesesccsecenesasss

Clytemnestra hendorffi var. quinquesetosa Poppe, 1891 .....

OTERIRECOLGS, Arorttee testeoree tac ttues sorecorcveet tonal eaagceveetsdeenssecvees

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Goniopsyllus rostratus Brady, 1883 .......cccccecceseeseeseeseeseeteees

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BLAXOUOMIG MaIpPeEaimMment, ANG MAINE PLATO «css nasencacexadeasscenssesesedvencnswes=asovkneacexes <eaaussicuessestewsudnssayuex nr de sucwenssceseser«cesaenteree 45

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NCL HEI CES aso eteseceacertccsestecass cu ccnceNs teresa iacsisssticaneaus resaasdresuacessveas suedinenmerashevasusearaa sot sutern siti ecsvastavacnot evedaceesasacseveticeessessartensrcesieee 46

SYNopSsIS. The family Clytemnestridae is one of the very few holoplanktonic harpacticoid lineages, typically occurring in the

epipelagic zone of all oceans. Its monogeneric status and the cosmopolitan distribution of the only two species, Clytemnestra

scutellata Dana, 1847 and C. rostrata (Brady, 1883), have been universally accepted since 1891. Re-examination of the major

expedition collections (Challenger 1873-76, Cambridge Suez Canal Expedition 1924, Great Barrier Reef Expedition 1928-29,

Discovery) in the Natural History Museum proved both perceptions to be false. The generic concepts introduced by Claus (18915)

but rejected by subsequent authors are revived, resulting in the recognition of two valid genera Clytemnestra Dana, 1847 (syn.

Goniopelte Claus, 1891a) and Goniopsyllus Brady, 1883 (syn. Sapphir Car, 1890). Genera are separated on the basis of

antennulary segmentation, caudal ramus sexual dimorphism and differences in the armature of the antenna, maxillule, maxilla,

Pi and P2. Fundamental discrepancies are found in the female genital field and the male gonopores.

Species discrimination prior to this revision was exclusively based on generic characters. Detailed examination of NHM

material has quadrupled the number of species in the family. Redescriptions are provided for both C. scutellata and G. rostratus,

and descriptions are given for five new species previously confounded with these type species: C. farrani sp. nov., C. longipes

sp. nov., C. asetosa sp. nov., G. clausi sp. nov. and G. brasiliensis sp. nov.

Goniopelte gracilis Claus, 1891a is redescribed and reinstated as a valid species in Clytemnestra. It is believed to represent the

Atlantic-Mediterranean sister-species of C. scutellata which presumably assumes only a restricted eastern Indo-Pacific

distribution. Neotypes are designated for C. scutellata and C. gracilis. Mediterranean and other European records of G. rostratus

in reality refer to G. clausi sp. nov.

R. HUYS AND S. CONROY-DALTON

C. hendorffi Poppe, 1890 is a junior subjective synonym of C. scutellata. The doubtful status of Sapphir rostratus Car, 1890,

Clytemnestra tenuis Lubbock, 1860 and C. hendorffi var. quinquesetosa Poppe, 1890 is discussed.

The intricate taxonomic history of the family is reviewed, including the nomenclatural confusion surrounding the priority of

the family name. The phylogenetic relationships of the Clytemnestridae as well the ontogenetic processes underlying the caudal

ramus sexual dimorphism in Clytemnestra are discussed. The taxonomic impediment in marine plankton research caused by the

failure to recognize pseudo-sibling or cryptic species is highlighted.

INTRODUCTION

The greatest habitat shift performed by copepods was undoubtedly

the colonization of the open pelagic environment, covering 71

percent of the Earth’s surface and providing a volume of 1347

million cubic kilometres. This habitat was most successfully ex-

ploited by the calanoids which can be regarded as the marine

planktonic copepods par excellence (Huys & Boxshall, 1991), and

to a lesser extent by the cyclopoids and poecilostomatoids which can

be particularly abundant in small mesh net samples. The evolution-

ary history of harpacticoid copepods in the marine plankton is less of

a success story and is to be viewed as the result of multiple

colonization. Only three families are currently considered as exclu-

sively holoplanktonic, the Miraciidae, Euterpinidae and

Clytemnestridae, and each of them can be regarded as an evolution-

ary cul de sac. The Miraciidae contains 4 monotypic genera which

are typically associated with marine filamentous Cyanobacteria

(Huys & Bottger-Schnack, 1994). The Euterpinidae is represented

by asingle species Euterpina acutifrons (Dana, 1847) which is often

abundant in shallow neritic waters. The Clytemnestridae currently

comprises two cosmopolitan species which are primarily found in

the epipelagic zone but frequently penetrate into deeper layers. The

Aegisthidae, commonly regarded as typical holoplanktonic forms

found in the mesopelagic and bathypelagic zones, has recently been

shown to be only a secondary offshoot from a hyperbenthic ancestral

stock (Conroy-Dalton & Huys, 1999; Lee & Huys, in press). Other

pelagic harpacticoids exhibit an essentially benthic biology by their

association with ‘planktonic’ substrata, such as Microsetella spp.

which attach themselves to discarded and occupied larvacean houses

(Appendicularia) (Ohtsuka et al., 1993), and Parathalestris croni

(Kr@yer, 1846) which is typically associated with floating macroalgal

clumps (Ingolfsson & Olafsson, 1997).

Clytemnestrids have been known since the advent of the pioneer-

ing oceanographic expeditions such as the U.S. Explorer Expedition

(Dana, 1854) and the Voyage of the H.M.S. Challenger (Brady,

1883). They were originally classified as poecilostomatoids until

Claus (1891a) demonstrated their harpacticoid identity. Virtually all

of the taxonomic literature on this family was published in the

second half of the 1800s and apart from cursory treatment by Lang

(1948), Wells (1970) and Boxshall (1979) no significant contribu-

tions have been added since.

MATERIAL AND METHODS

The descriptive terminology is adopted from Huys et al. (1996).

Abbreviations used in the text are: ae, aesthetasc; P1—P6, first to

sixth thoracopod; exp(enp)-1(2, 3) to denote the proximal (middle,

distal) segment of a ramus. Specimens were dissected in lactic acid

and the dissected parts were placed in lactophenol mounting me-

dium. Preparations were sealed with glyceel (Gurr®, BDH Chemicals

Ltd, Poole, England) or transparent nail varnish. All drawings have

been prepared using a camera lucida on a Leitz Dialux or Leitz DMR

microscope equipped with differential interference contrast.

Clytemnestra gracilis and Goniopsyllus clausi were examined

with a Philips XL30 scanning electron microscope. Specimens were

prepared by dehydration through graded acetone, critical point

dried, mounted on stubs and sputter-coated with palladium.

Citations of articles in the International Code of Zoological

Nomenclature (ICZN) refer to the fourth edition published in Aug-

ust 1999 and superseding previous editions with effect from 1

January 2000. Type series and other material is deposited in the

collections of the Natural History Museum, London (BMNH).

TAXONOMIC HISTORY

The proliferation of generic names in this family at the end of the

19th century marked one of the most virulent episodes in the history

of harpacticoid taxonomy. The key players in this debate were the

eminent and influential Carl Claus and a cohort of opponents

including Wilhelm Giesbrecht, S.A. Poppe and Lazar Car. It is clear

that much of the confusion arose from observational errors made by

both Dana (1854) and Brady (1883).

Clytemnestra Dana, 1847

Dana introduced the genus Clytemnestra in the first part of his

“Conspectus Crustaceorum’ which was published in 1847 (for dis-

cussion of publication dates see Huys & Bottger-Schnack, 1994)

and included the families Cyclopidae and Harpactidae. This paper,

completely lacking in illustrations, provided a Latin diagnosis for

the genus and its only species C. scutellata which was placed in the

‘Harpactidae’ together with Harpacticus Milne Edwards, 1840 and

Setella Dana, 1846. Although no type locality was designated, the

author did mention that the species was found near the Gilbert

Islands and east of Tuamotu in the Pacific Ocean and in the South

China Sea. In his second volume of the Crustacea of the United

States Exploring Expedition (Dana, 1854) a more extensive and

illustrated description of C. scutellata was given based on speci-

mens from the Tuamotu samples.

Lubbock (1856) added a second species C. atlantica which he

described on the basis of a single female from an unspecified locality

in the Atlantic. The brief original description included illustrations

of the habitus and antenna only. Various authors (Poppe, 1891;

Giesbrecht, 1892; Lang, 1948) have questioned this identification

and referred the species to the genus Pachos Stebbing in the

Poecilostomatoida. Pesta (1909) considered C. atlantica as a syno-

nym of Pachos punctatum (Claus). In a later report Lubbock (1860)

described C. tenuis, again from a single female, collected east of

Mauritius. Lubbock himself had some reservations about the sexual

maturity of the specimen, and Poppe (1891) considered the species

as unrecognizable. Giesbrecht (1892) listed C. tenuis as a possible

synonym of C. rostrata.

Claus (1863) rejected Clytemnestra as a valid genus by stating

that the illustrations were so inadequate that they were worthless for

identification purposes.

GENERIC CONCEPTS IN CLYTEMNESTRIDAE

Goniopsyllus Brady, 1883

Brady (1883) established this genus for a single specimen found in

a tow-net gathering taken off the Argentinean coast during the

voyage of the H.M.S. Challenger. He regarded Goniopsyllus rostratus

as most closely related to the harpacticoid genera Enhydrosoma

Boeck and Cletodes Brady despite the marked differences in the

mouthparts. In addition, Brady remarked on the similarity in swim-

ming leg morphology with Peltidium and recognized a certain

affinity with the Sapphirinidae because of the rudimentary structure

of the mouthparts. The description of G. rostratus is fragmentary

and partly inadequate. Brady (1883) failed to observe the mandible.

Sapphir Car, 1890

Car (1890) described both sexes of Sapphir rostratus from plankton

samples taken off Trieste in the Adriatic. He used and revised

Brady’s (1878) classification, dividing the free-living copepods in 6

families (Calanidae, Cyclopidae, Harpactidae, Peltididae,

Corycaeidae and Sapphirinidae), but was apparently unaware of

Brady’s (1883) later paper describing the closely related Goniopsyllus

rostratus. Car (1890) placed Sapphir in the Sapphirinidae merely by

way of elimination and excluded the genus from the two harpacticoid

families known at that time (Harpactidae, Peltididae) by virtue of the

absence of (1) geniculate setae on the antennae, (2) a palp on the

mandible and maxillule, (3) modifications of the Pl, and (4) a

foliaceous PS. Allocation to the Sapphirinidae was substantiated by

the dorsoventrally depressed body, the 6-segmented antennules

which are similar in both sexes (Car did not recognize the sexual

dimorphism and male geniculation), the antenna lacking a defined

exopod and geniculate setae on the endopod, the reduced mouthparts,

the sexually dimorphic maxillipeds and the small PS.

In a short note Dahl (1890) considered S. rostratus a junior

subjective synonym of G. rostratus but gave no justification for this

course of action.

Car (1891a) admitted that he had overlooked Brady’s (1883)

Challenger report describing G. rostratus but maintained the dis-

tinction between both genera. His conviction was based on three

doubtful observations made by Brady (1883): (1) his statement that

all four swimming legs were ‘nearly alike’ having 3-segmented

rami; Brady only figured the P2 which he labelled ‘One of the

swimming feet’, (2) the maxillipeds which were described and

figured as 3-segmented, and (3) the 3-segmented fifth legs. Car

pointed out that in Sapphir the P1 exopod was clearly 1-segmented,

and both the maxillipeds and the P5 2-segmented, but did not

consider the possibility that this incongruity could be based on

observational errors made by Brady. It was largely this failure that

initiated the subsequent dispute between Car and Claus.

Goniopelte Claus, 1891a

Both sexes of Goniopelte gracilis were described in remarkable

detail by Claus (1891a) on the basis of scanty material (1 2 and 1d)

collected from an unspecified locality in the Eastern Mediterranean.

He recognized the male geniculation (‘elastischen Cuticularapparat’ )

and the ‘accessory’ aesthetascs of the antennules, the sexual dimor-

phism of the caudal rami and the presence of the male P6. Claus also

revealed details of the internal anatomy such as the tripartite nauplius

eye, the asymmetry of the male genital system and the presence of

integumental glands around the rostrum and the pleural areas of the

cephalothorax, pedigerous somites and abdomen.

Claus (1891a) severely criticized the quality of both Brady’s

(1883) and Car’s (1890) descriptions and like Dahl (1890) professed

that G. rostratus and S. rostratus were not only congeneric but also

conspecific. The differentiating characters used by Car (1890, 1891a)

he regarded as irrelevant to the issue. He presented convincing

arguments showing that Brady’s holotype of G. rostratus could not

possibly have been a male. Claus was also the first author to

reconsider Dana’s Clytemnestra scutellata. He placed the species

with reservations in the Scutellidiinae (“Scutellidinen’), a subfamily

of the Peltidiidae (‘Peltididen’), despite similarities in general body

shape and maxilliped structure with his new genus and species

Goniopelte gracilis.

Claus (1891a) remarked that the moderate flattening of the body,

the reduction of the mandible and maxillule, and the 1-segmented P1

exopod in G. gracilis would probably warrant the erection of a third

subfamily within the Peltidiidae. An alternative option suggested by

Claus was to regard it as a transitionary group between the Peltidiidae

and Harpacticidae.

Car’s (18915) re-examination of S. rostratus did not disclose new

information apart from the confirmation of the 4-segmented con-

dition of the antenna. Although his rebuttal was mainly aimed at

showing disapproval of Claus’ (1891a) provocative paper, it con-

tained clear indications of the author’s ambivalence about both the

conspecificity and familial placement of S. rostratus. Car main-

tained the latter as a valid genus and species but did not exclude

potential synonymy with G. rostratus. He kept the genus in the

Sapphirinidae but pointed out the close relationship between Sapphir,

Goniopsyllus and Goniopelte and the possible option of proposing a

new family for these three genera. Finally, he disagreed with Claus

(1891a) on the sexual identity of the holotype of G. rostratus, using

the unconfirmed presence of an internal spermatophore in Brady’s

(1883) habitus drawings as the only counterargument.

A breakthrough in unravelling the intricate synonymy was realized

by Poppe who had already recognized the identity between

Clytemnestra and Goniopsyllus in 1884 but did not publish his results

until 1891. Poppe’s (1891) comprehensive paper, which downgraded

Goniopsyllus and Sapphir to junior synonyms of Clytemnestra, was

based on a wide range of specimens including the holotype of G. ros-

tratus and a male of S. rostratus from Car’s collection. He described

anew species, Clytemnestra hendorffi from material collected in the

Java Sea, the Indian Ocean (south of Madagascar, Western Australian

Basin) and the South Atlantic (off Brazil and Argentina). Poppe

(1891) also re-examined Thompson’s (1888) material of G. rostratus

from Malta and identified it as C. hendorffi. Among the material from

the Java Sea he discovered a variety quinquesetosa which differed

from the typical form in the longer P5 which carried only 5 setae on

the exopod, amore stocky abdomen in both sexes and the caudal rami

which were relatively wider proximally.

Poppe (1891) synonymised G. rostratus and S. rostratus and

considered the previous distinction between them to be based on

erroneous observations of the PS by both Brady and Car, and the fact

that Brady had misidentified the holotype of S. rostratus as a male

and overlooked the Pl exopod in this species. For some unknown

reason he suspected the latter to be 2-segmented in G. rostratus. He

considered only 3 species as valid, all of which he placed in

Clytemnestra: C. scutellata, C. hendorffi and C. rostrata (Brady).

Poppe further regarded the inadequately described C. tenuis as a

probable synonym of C. scutellata and excluded Lubbock’s second

species C. atlantica from the genus on account of the different body

shape and the structure of the antennules.

Poppe (1891) did not accept Car’s (1890, 1891a—b) placement in

the Sapphirinidae and created anew family Pseudo-Peltididae which

showed similarities with the Peltidiidae but differed in the morphol-

ogy of the P1 (exopod not prehensile and 2-segmented (!) according

to Poppe’s diagnosis), the absence of a well defined antennary

exopod and strongly reduced mouthparts.

With Giesbrecht’s (1891a) claim that Goniopelte had already

been described under three different generic names the synonymy

issue surrounding Clytemnestra appeared to have come to a close.

Claus (18915), however, continued to defend his genus Goniopelte

with extraordinary persistence. After re-examination of Poppe’s

(1891) material, confirming the presence of the male P6, and the

vestigial antennary exopod, he acknowledged the conspecificity of

G. gracilis and C. hendorffi. Nevertheless, he adhered to his earlier

decision (Claus, 1863) to dismiss Clytemnestra as a valid genus. He

based this course of action on the rules drawn up by Raphael

Blanchard and Maurice Chaper and adopted, in part, at the First

International Congress of Zoology (Paris, 1889). They stipulated in

§ 7 that the valid name should be the oldest one provided that *. . . ce

nom etc. aura été clairement et suffisament defini’. Claus (1891)

rejected Poppe’s (1891) arguments as insufficient for the proposal of

a new family and instead created a third subfamily Goniopeltidinae

in the Peltididiidae. In this subfamily he recognized two genera,

Goniopsyllus (syn. Sapphir) and Goniopelte, which were differenti-

ated on the basis of antennule segmentation, antennary exopod

setation and caudal ramus sexual dimorphism.

Claus’ (1891b) generic concepts were finally rejected by

Giesbrecht (1892) who reviewed the intricate synonymy and rein-

stated Clytemnestra as the only valid genus on the basis of the

Principle of Priority. Giesbrecht (1891b, 1892) recognized only two

species, C. scutellata and C. rostrata, and regarded all other species

as subjective synonyms with the possible exception of C. tenuis.

This course of action was adopted by most subsequent authors such

as Lang (1944, 1948) and Boxshall (1979). The rapid accumulation

of plankton data during the 20th century fed the conjecture that both

species assumed a cosmopolitan distribution. Unfortunately, this

presumption made people loose sight of the possible existence of

other undescribed species and of the true identitiy of C. scutellata

and G. rostratus.

PRIORITY OF THE FAMILY NAME

Although various authors (Car, 18915; Claus, 1891a) had expressed

the need to introduce a new family or subfamily for Goniopsyllus,

Goniopelte and Sapphir it was finally Poppe (1891) who coined the

family name Pseudo-Peltididae for the only included genus

Clytemnestra. Claus (1891b) rejected the family status of Pseudo-

Peltididae and established a new subfamily Goniopeltidinae for

Goniopelte and Goniopsyllus. Giesbrecht (1892) did not consider

familial assignment which probably misled A. Scott (1909) who did

not consult the earlier literature and consequently proposed the new

family name Clytemnestridae for the type and only genus

Clytemnestra. Mori (1929) placed this genus in the Harpacticidae

whereas Wilson (1932) referred it to the Tachidiidae for some

unknown reason, an inexplicable assignment followed also by

Carvalho (1952) and Krishnaswamy (1953).

Most workers (e.g. Sars, 1921; Monard, 1927; Sewell, 1940; Klie,

1943) adopted Clytemnestridae as the valid family name until Lang

(1944, 1948) pointed out that Poppe’s Pseudo-Peltididae took prior-

ity over the latter. Boxshall (1979) remarked that this course of

action contravened ICZN Art. 11.7.1.1 since a family-group name

must, when first published, be based on the name then valid for a

contained genus. Poppe’s (1891) family name with its alternative

spellings Pseudo-Peltididae (Poppe, 1891), Pseudo-Peltidiidae

(Lang, 1944) and Pseudopeltidiidae (Wells, 1976) is therefore una-

vailable. Boxshall (1979) reinstated Clytemnestridae as the valid

name, but unfortunately ignored Claus’ (1891b) older and validly

R. HUYS AND S. CONROY-DALTON

introduced family-group name Goniopeltidinae. Other authors con-

tinued using Pseudopeltidiidae (e.g. Bowman & Abele, 1982).

Were priority to be rigorously enforced, Goniopeltididae should

replace its junior synonym Clytemnestridae and hence leave Claus,

at best, a pyrrhic victory. However, since the senior synonym

Goniopeltidinae has remained unused as a valid name since 1899

(ICZN Art. 13.9.1.1) and the junior synonym Clytemnestridae has

been used as the presumed valid name in at least 25 works

(Krishnaswamy, 1957; Marques, 1957; Bruce et al., 1963;

Kasturirangan, 1963; Cheng ef al., 1965; Owre & Foyo, 1967;

Fagetti, 1962; Chen et al., 1974; Boxshall, 1979; De Decker, 1984;

Citarella, 1986; Hicks, 1988; Huys & Boxshall, 1991; Razouls &

Durand, 1991; Campos Hernandez & Suarez Morales, 1994; Huys

& Bottger-Schnack, 1994; Kazmi & Muniza, 1994; Hirota, 1995;

Huys et al., 1996; Razouls, 1996; Bodin, 1997; Chihara & Murano,

1997; Hure & KrSinié, 1998; Reid, 1998: Suarez Morales & Gasca,

1998) published by at least 10 authors in the immediately preceding

50 years (and encompassing a span of not less than 10 years) (ICZN

Art. 13.9.1.2.) it is to be considered a forgotten name (nomen

oblitum). In accordance with Art. 23.9.1. prevailing usage is main-

tained and the junior name Clytemnestridae is treated as a nomen

protectum.

SYSTEMATICS

Claus’ (1891b) generic concepts of Goniopelte and Goniopsyllus

were based on differences in antennule segmentation, antennary

exopod setation and caudal ramus sexual dimorphism. Re-exam-

ination of material attributed to C. scutellata and C. rostrata have

revealed additional differentiating characters in mouthpart struc-

ture, swimming leg setation and female genital field morphology,

substantiating Claus’ recognition of two distinct genera. Secondly,

there is accumulating evidence that both C. scutellata and C. rostrata

represent species complexes, each of which can be justifiably

assigned generic rank. It has not been our intention to verify every

published record of these species since in most cases the information

contained in the numerous marine plankton studies did not permit

unambiguous identification. This paper is based almost solely on

BMNH collections and serves as a baseline study for future species

discrimination in the Clytemnestridae. It is aimed primarily at

reviving and elaborating Claus’ (1891b) original generic concepts,

albeit partly under different taxonomic names.

Family CLYTEMNESTRIDAE A. Scott, 1909

DIAGNOSIS. Body distinctly tapering posteriorly. Prosome dors-

oventrally flattened, urosome slender and cylindrical. First

pedigerous somite incorporated in cephalosome forming bell-shaped

cephalothorax. Pedigerous somites bearing P2—P4 with posteriorly

directed alate projections. Genital and first abdominal somites of

completely fused forming genital double-somite; original segmen-

tation marked by small chitinized internal ribs ventrally or laterally.

Anal operculum obsolete; anus terminal.

Sexual dimorphism in antennule, maxilliped, P6, urosomal orna-

mentation and in genital segmentation; often in rostrum shape,

occasionally in caudal ramus. No distinct sexual dimorphism in P1—

PS.

Rostrum large, fused to cephalic shield. Antennules slender; 6- or

7-segmented in 9; haplocer and distinctly or indistinctly 7-seg-

mented in 6, with geniculation between segments 6 and 7; aesthetascs

present on 4th and apical segments in 2, on 3rd, Sth and apical

GENERIC CONCEPTS IN CLYTEMNESTRIDAE

segments in 6; transformed aesthetasc-like setae present on seg-

ments 3, 4 and 6(or 7) in 2, and segments 3, 5 and 7 in d. Antenna

with separate basis and 2-segmented endopod; basis and proximal

endopod segment unarmed; distal endopod segment with 1 lateral

and 4-5 apical elements; exopod a minute segment with 1—2 long

setae. Mandibles, maxillules and maxillae reduced. Mandible with

stylet-like gnathobase, palp represented by | short seta. Maxillule a

small segment with 1 or 3 elements. Maxilla with 1-2 endites on

syncoxa; allobasis with articulating claw and 2 accessory elements.

Maxillipeds very large with elongate syncoxa and basis; syncoxa

with | seta, basis with 1 short seta and | pad-like element on palmar

margin; endopod represented by sexually dimorphic claw and 5

accessory elements.

Pl with 1-segmented exopod and 3-segmented non-prehensile

endopod; basis without inner seta/spine. P2—P4 with transversely

elongated basis bearing short outer seta; rami 3-segmented with

endopods longer than exopods. Outer spines of exopod segments

typically setiform, often with flagellate tip. Armature formula as

follows:

exopod endopod

Pl [0O-1]21 1.1.220

BZ 1.1.22[2-3] 222i

PS 1.1.32[2-3] 127321

P4 1.1.32[2-3] V2 221

P5 uniramous, comprising basis and 1-segmented exopod; later-

ally displaced; exopod elongate, with 5-6 setae.

Female genital field positioned anteriorly; genital apertures paired

or fused to median slit; closed off by vestigial P6 bearing I element;

copulatory pore unpaired. P6 d with | or 3 elements; closing off

median or asymmetrically positioned (sinistral/dextral) genital ap-

erture.

Caudal rami conical or rectangular, short; rear margin between

setae III and IV produced into conical process bearing apical pore;

setae I-II spiniform and strongly developed (seta I longer than II);

setae IV—V fused at base, without fracture planes.

One median egg-sac; spermatophores elongate, with very long

recurved neck.

Holoplanktonic, marine.

TYPE GENUS. Clytemnestra Dana, 1847

OTHER GENUS. Goniopsyllus Brady, 1883

Genus Clytemnestra Dana, 1847

Goniopelte Claus, 1891a [type species: G. gracilis Claus, 1891 —by

monotypy]

DIAGNOsIS. Clytemnestridae. Body without dorsal pattern of

denticles or spinules on urosomites. Antennule distinctly 7-seg-

mented in both sexes; ¢ segmental homologies: 1—I, 2—-(II-VIII),

3-(IX-XIID, 4-(XIV-XVII), 5-(X VIII), 6-(XIX—XX), 7-(XXI-

XXVIII); segment 5 in d with large spine. Antenna with | lateral

and 5 apical elements on distal endopod segment; exopod repres-

ented by well defined segment bearing 2 long setae. Maxillule

represented by bilobed segment with | lateral seta and 2 apical

spines. Maxillary syncoxa with 1—2 endites; proximal endite repres-

ented by very long seta, sometimes absent; distal endite bearing 3

setae.

P1 with outer seta on basis; exopod with 4 setae. P2 without outer

spine on exp—1. P1—P4 armature formula:

exopod endopod

Pl 121 1220

P2 1.1.22[2-3] 1.2.221

P3 1.1.32[2—3] 1.2.321

P4 1.1.32[2-3] 1.2.221

P5 exopod with 5 or 6 setae in both sexes.

Genital apertures paired in 9; closed off by paired P6 bearing 1

vestigial element; copulatory pore small, located anteriorly between

genital apertures; copulatory duct probably very short and definitely

not strongly chitinized.

Male P6 almost symmetrical, fused medially forming membra-

nous operculum closing off single median genital aperture; produced

into cylindrical process bearing 3 small setae.

Caudal rami parallel, almost cylindrical; sexually dimorphic with

setae [V—V short and pinnate in 2, long and multiplumose in d;

additional sexual dimorphism also noted in setae III and VI.

TYPE SPECIES. Clytemnestra scutellata Dana, 1847 [by monotypy].

OTHER SPECIES. C. gracilis (Claus, 1891a) comb. nov., C. farrani

sp. nov., C. longipes sp. nov., C. asetosa sp. nov.

SPECIES INQUIRENDAE. Clytemnestra hendorffi var. quinquesetosa

Poppe, 1891

REMARKS. Various authors, including Giesbrecht (1892), Sars

(1921), Mori (1937) and Boxshall (1979), have erroneously described

the 2antennule as 8-segmented. From the illustrations of Giesbrecht,

Sars and Mori it appears that the basal pedestal has been repeatedly

misinterpreted as an additional segment. Although his description

contradicts the accompanying illustration, the proportional segment

lengths given by Boxshall (1979) for the C. scutellata antennule

suggest a similar observational error.

Clytemnestra scutellata Dana, 1847

Clytemnestra Hendorffi Poppe, 1891: 132-136, Taf. I.

The form of the maxilliped and the 6-segmented urosome clearly

identify Dana’s (1854) illustrated specimen as a male. The append-

age labelled ‘extremity of a maxilliped’ (his Fig. 12d) is almost

certainly the PS exopod. We concur with Claus (1863, 1891a—b) that

the original description of C. scutellata does not provide the bare

minimum for unequivocal identification. In fact, the synonymy of

Clytemnestra with Goniopelte advocated by Giesbrecht (1891a,

1892) is justified solely by the long terminal setae of the caudal rami

figured in Dana’s (1854) habitus drawing. This sexually dimorphic

feature is the only character in Dana’s description which both

positively identifies his species as a Clytemnestra and excludes it

from the genus Goniopsyllus. If Dana had figured a female specimen

even this generic determination would not have been possible.

Since both Clytemnestra and C. scutellata have now been widely

accepted for almost a century, we have retained both names in the

interest of stability of nomenclature even though they are virtually

unidentifiable on the basis of Dana’s description. The original type

material no longer exists and the male specimen figured in Dana

(1854) is so badly illustrated that we have refrained from designat-

ing it as the lectotype. In order to settle the issue a neotype has been

designated from BMNH material collected from the Great Barrier

Reef by Farran (1936) which forms the basis of the description

below.

TYPE LOCALITY. The determination of the type locality presents

some difficulty. In his original diagnosis Dana (1847) listed three

6 R. HUYS AND S. CONROY-DALTON

t. & y ‘sta v

f) 44 2 Mi ’ a i

SS ee ae ee

oe ee ie eet

I. ce == =

om 7 < i

“ “ e

Fig. 1 Clytemnestra scutellata Dana, 1847. A, Habitus 9, dorsal; B, habitus ¢, dorsal; C, P5 9, anterior. [A, C based on neotype].

GENERIC CONCEPTS IN CLYTEMNESTRIDAE

localities, i.e. the South China Sea (300 miles NE of Singapore),

near Pitt’s Island (Kingsmill Group, Kiribati) and the eastern Pacific

Ocean at 18°S 124°W, but he did not designate a type locality. In his

illustrated description (Dana, 1854) he mentioned that the descrip-

tion and figures were based on specimens from the eastern Pacific

which could arguably be considered as the type locality.

Farran (1936) recorded a total of 11 specimens of C. scutellata

from 6 different stations sampled during the Great Barrier Reef

Expedition in 1928-29. Five specimens were found in serial

townettings inside the reef and another six specimens were discoy-

ered in deeper waters outside the reef. Examination of Farran’s spirit

preserved material in the Natural History Museum (BMNH

1948.4.28.121) revealed 3 29,5 dd and 1 damaged @ prosome,

representing at least 3 different species. According to Farran (1936)

the specimens from the reef flat were significantly smaller (0.80.9

instead of 1.05—1.20 mm) except for one male which measured 1.15

mm. The small specimens (2 99, 2 dd) are present amongst the

NHM material and represent a new species. The larger male could

also be identified and is described below as C. longipes sp. nov.

Among the remaining material, which must therefore have been

collected outside the reef, 1 female and 1 male agreed with (or at

least did not contradict) Dana’s (1854) description and are here

identified as C. scutellata primarily on the basis of cephalothorax

shape. Moreover, the close size correlation between Dana’s male of

C. scutellata (‘1—24th of an inch’ = 1058 pm) and the male from the

Great Barrier Reef (1064 pm) is striking. The single female speci-

men is designated here as the neotype, defining Farran’s (1936)

stations 19, 20 and 28 collectively as the new type locality (ICZN

Art. 76.3.) despite previously published statements of the place of

origin of Dana’s material. All three stations are situated outside the

Trinity opening to the reef off Port Douglas at 16°19-20'S, 146°3-

7E (Queensland). The depth ranges from 225 (stn 19) to >600 m

(stns 20, 28)

TYPE MATERIAL. Neotype 2 dissected on 11 slides (BMNH

1999.996); designated from material labelled Clytemnestra scutellata

(BMNH 1948.4.28.121); collected either on 20 October 1928 (stns

19, 20) or 23 November 1928 (stn 28) during the Great Barrier Reef

Expedition 1928-29 (Farran, 1936).

OTHER MATERIAL EXAMINED. One <4 dissected on 10 slides

(BMNH 1948.4.28.121); sampling data as for neotype.

REDESCRIPTION.

FEMALE. Total body length from tip of rostrum to posterior margin

of caudal rami: 1121 um. Maximum width (355 tm) measured at

posterior margin of cephalic shield. Posterolateral angles of

cephalothorax laterally expanded (Fig. 1A). Somites bearing P2—P4

successively decreasing in width posteriorly and bearing back-

wardly produced alate processes.

Genital double-somite (Fig. 5A) slightly constricted bilaterally;

original segmentation marked by paired transverse chitinous ribs

lateroventrally and laterally. Copulatory pore slit-like, located medi-

ally between genital apertures; leading to short posteriorly directed,

membranous duct connected to bilobate seminal receptacle. Genital

apertures located far anteriorly; closed off by small opercula derived

from vestigial P6; each with 1 vestigial seta at inner distal corner and

anterior tube-pore near base.

Urosomites without dorsal ornamentation (Figs 1A, 4E); penulti-

mate and anal somites with multiple rows of spinules around ventral

hind margin (Fig. 5A).

Caudal rami (Fig. 4E) about twice as long as wide, parallel;

slightly tapering towards rear margin, with stepped outer margin

marking insertion sites of setae I, II and III; produced into conical

process bearing terminal pore; posterior third with ventral spinular

patch (Fig. 5A). Setae -II minutely bipinnate, spiniform and strongly

developed. Seta III bipinnate. Setae IV—V basally fused; about

equally long and only slightly longer than caudal ramus; without

fracture planes, multipinnate and spiniform. Seta VI minute, bare;

seta VII small, biarticulate at base, bare.

Rostrum (Fig. 1A) triangular with rounded anterior margin, com-

pletely fused to cephalothorax; with numerous dorsal surface pores

as figured, none on ventral surface; with minute lateral sensillae near

apex.

Antennule (Fig. 2A) slender, 7-segmented; segment 7 longest.

Plumose setae present on segments 14. Segment 1 with small pore

near seta and few short spinules along anterior margin. Armature

formula: 1-[1 plumose], 2-[9 + 3 plumose], 3-[4 + 3 plumose + 1

transformed], 4-[1 + 1 plumose + (1 transformed + ae)], 5-[1], 6-[3],

7-[8 + acrothek]. Apical acrothek consisting of aesthetasc, long

transformed seta and short bare seta. Transformed setae on segments

3, 4 and 7 long and aesthetasc-like, with rounded tip; those on

segments 4 and 7 basally fused to aesthetasc. Rudimentary element

present at base of acrothek.

Antenna (Fig.3A) 4-segmented, comprising coxa, basis and 2-

segmented endopod. Coxa well developed, bare. Basis and proximal

endopod segment without ornamentation; unarmed. Exopod inserted

in membranous area between basis and endopod; represented by

small, well defined segment bearing 2 strong recurved setae apically;

exopodal setae multipinnate with long setules in proximal third.

Distal endopod segment (Fig. 3A, B) with several surface frills and

minute spinules on outer surface and patch of long setules on medial

surface; lateral armature consisting of | naked seta; distal armature

consisting of 5 apical, non-geniculate, bipinnate or multipinnate

elements, 2 of which spiniform, recurved and bearing long spinules

proximally.

Labrum (Fig. 3C) large, with 6 secretory pores on anterior sur-

face; distal margin spinulose medially and with spinular patch on

either lateral lobe.

Mandible (Fig. 3D) reduced. Palp represented by single naked

seta. Gnathobase long and narrow, stylet-like; produced into number

of cuspidate processes apically and subapically; without dorsal

seta(e).

Paragnaths (Fig. 3C) well developed hirsute lobes.

Maxillule (Fig.3E) reduced; represented by small bilobed seg-

ment bearing 2 naked apical spines and raised seta along outer

margin; posterior surface with distinct pore.

Maxilla (Fig. 3F) 2-segmented, comprising elongate syncoxa and

allobasis. Syncoxa with expanded basal portion and 2 endites; exit

of maxillary gland large (arrowed in Fig. 3F), partly concealed under

lobate extension; proximal endite represented by small cylindrical

process bearing very long plumose seta, distal endite cylindrical,

with | naked and 2 pinnate spines apically. Allobasis with large

articulating claw distally, smaller inner pinnate spine and naked seta

along outer margin.

Maxilliped (Fig. 4A, B) very large, articulating with well devel-

oped pedestal; 3-segmented, comprising syncoxa, basis and endopod.

Syncoxa extremely elongate, longer than basis; without ornamenta-

tion but with | anterior, plumose seta near membranous articulation

with basis. Basis elongate; distal third of palmar margin with double

spinule row (anterior spinules coarser than posterior ones) and 2

elements located closely to articulation with endopod; proximal

element spiniform and bare (arrowed in Fig. 4B), distal element pad-

like and spinulose. Endopod represented by short segment bearing

short naked claw; accessory armature consisting of 3 anterior and 2

posterior elements.

Swimming legs with wide, narrow intercoxal sclerites and well

R. HUYS AND S. CONROY-DALTON

Libr ayygi iii ‘

“as

| = < j LEE,

(

Fig. 2 Clytemnestra scutellata Dana, 1847. A, Antennule @, dorsal; B, antennule 4, ventral; C, antennulary segment 3 6, anterior; D, antennulary

segments 4~7 d, anterior [distal portion of segment 7 and proximal portion of segment 4 omitted]; E, antennulary segments 5S—6 d, ventral; F,

antennulary segment 7 2, distal portion, dorsal [arrow indicating rudimentary element]. [A based on neotype].

GENERIC CONCEPTS IN CLYTEMNESTRIDAE

Fig. 3 Clytemnestra scutellata Dana, 1847 (). A, Antenna, outer; B, distal antennary endopod segment, inner; C, oral area showing position of labrum,

paragnaths, mandibles, maxillules and right maxilla [position of maxilliped (Mxp.) indicated], ventral; D, mandible, posterior; E, maxillule, posterior; F,

maxilla [exit of maxillary gland arrowed], posterior. [all based on neotype].

R. HUYS AND S. CONROY-DALTON

—S—

—

——~

LSS)

LNT wi

— Ny.

“ LS

Ape

SSS

Fig. 4 Clytemnestra scutellata Dana, 1847. A, Maxilliped , posterior; B, maxilliped 9 distal half of basis and endopod, anterior [proximal palmar

element arrowed]; C, maxilliped d, anterior; D, maxilliped d, distal portion of basis and endopod [proximal palmar element arrowed], posterior; E, right

caudal ramus @, dorsal; F, right caudal ramus 4, dorsal. [A, B, E based on neotype].

iil

GENERIC CONCEPTS IN CLYTEMNESTRIDAE

anhth

a Va

iy,

100

ZZ We)

SS BEE Za

Ze = go aa

[inset showing setae IV—V at full length]; C, P6 3, ventral. [A

Fig.5 Clytemnestra scutellata Dana, 1847. A, Urosome 2 ventral; B, urosome 6, ventral

based on neotype]

developed praecoxa; both without ornamentation. Rami 3-segmented

except for P1 exopod.

P1 (Fig. 6A) separated from maxillipeds by large membranous

area. Coxa and basis prolonged along dorsoventral axis; without

surface ornamentation. Basis with plumose outer spine. Exopod 1-

segmented, represented by elongate segment bearing long setules

along outer margin; with subapical pore and 1 outer, 2 apical and 1

inner setae. Endopod 3-segmented; segments decreasing in size

distally, each with anterior pore; enp-1 and -2 with few setules along

outer margin, enp-2 and -3 with posterior spinules; enp-1 with very

long inner seta; ornamentation of inner elements typically

(multi)pinnate, distal elements of enp-3 plumose.

P2—P4 (Figs 6B; 7A, B) with transversely prolonged basis bearing

short outer seta. Endopods distinctly longer than exopods. Exopodal

outer spines setiform with flagellate tip. Exopod segments typically

with pore near outer distal corner; without ornamentation; exp-2

outer distal corner linguiform. Endopods with long proximal seg-

ment, particularly in P2—P3; segments with anterior pore, setules

along outer margin and spinules on posterior surface; setal ornamen-

tation typically combination of setular and spinular rows; inner seta

of P2—P3 enp-1! short. P1 exp-2 without outer spine. Spine and setal

formula of swimming legs as follows:

Exopod Endopod

Pl 121 1.1.220

P2 R228 2-220

P3 EES 28 e237

P4 1.1.323 P2221

P5 (Fig. 1C) uniramous, laterally displaced; 2-segmented; not

extending beyond posterior margin of genital double-somite (Fig.

5A). Basis with short outer seta and anterior pore. Exopod about

twice as long as basis, slightly curved inwards; outer margin with 4

pinnate setae; inner margin with long plumose seta; apex and inner

margin each with 1 long pinnate seta; anterior surface with 3 pores

and spinules near apex and in proximal third.

MALE. Total body length from tip of rostrum to posterior margin of

caudal rami: 1064 um. Maximum width (337 um) measured at

posterior margin of cephalic shield. Body (Fig. 1B) with similar

projections as in 2; urosome more slender with genital and first

abdominal somites separate (Fig. 5B).

Rostrum (Fig. 1B) more obtuse than in &.

Antennule (Fig. 2B) slender, distinctly 7-segmented with ances-

tral segment XIII completely incorporated into segment 4 (Fig. 2C);

haplocer, with geniculation located between segment 6 and 7.

Plumose setae present on segments 14. Segment | with small pore

near seta and few tiny spinules along anterior margin. Armature

formula: 1-[1 plumose], 2-[8 + 3 plumose], 3-[5 + 3 plumose + 1

pinnate + | transformed + ae], 4-[2 + 3 plumose + (1 transformed +

ae)], 5- [1 + 1 spine], 6-[2], 7-[9 + 2 modified elements + acrothek].

Apical acrothek consisting of aesthetasc, long transformed seta and

short bare seta. Transformed setae on segments 3, 4 and 7 long and

aesthetasc-like, with rounded tip; those on segments 4 and 7 basally

fused to aesthetasc. Rudimentary element present at base of acrothek

(arrowed in Fig. 2F). Segment 6 with 2 patches of spinules on

anterior surface (Fig. 2D—E). Segment 7 with 2 fused elements near

geniculation (Fig. 2D).

Maxilliped (Fig. 4C) much larger than in 2 articulating with well

developed pedestal; 3-segmented, comprising syncoxa, basis and

endopod. Syncoxa extremely elongate but not distinctly longer than

basis; without ornamentation but with 1 short anterior seta near

R. HUYS AND S. CONROY-DALTON

membranous articulation with basis. Basis elongate; more swollen

than in 9; middle and distal thirds of palmar margin forming

longitudinal furrow bordered by single row of spinules on both

anterior and posterior sides; with 2 elements located closely to

articulation with endopod; proximal element spiniform and bare

(arrowed in Fig. 4D), distal element pad-like and spinulose. Endopod

represented by short segment produced into very long naked claw

which in reflexed position typically fits in palmar furrow with the

apical part closely adpressed onto the anterior surface of the basis;

accessory armature consisting of 3 anterior and 2 posterior setae;

claw with spatulate apex.

P5 (Fig. 7C) very similar to that of 2, with identical proportions,

pore pattern and setation.

Sixth pair of legs (Fig. 5B) weakly asymmetrical, forming highly

membranous midventral area covering single, large median genital

aperture; each P6 produced into cylindrical process (Fig. SC) with 1

apical and 2 outer bare setae; few spinules along inner margin.

Urosomites 4—5 and anal somite with spinules around ventral hind

margin (Fig. 5B).

Caudal rami (Fig. 4F) somewhat shorter than in 9; seta II rela-

tively longer; seta III more slender and with longer pinnules; setae

IV-V long (60% of urosome length; Fig. 5B) and plumose; seta VI

much longer than in 2 and sparsely plumose.

Spermatophore with very long, recurved neck.

VARIABILITY. The right distal exopod segment of the male P2 has

only 2 outer spines (Fig. 6C).

REMARKS. There are very few published records of C. scutellata

that can be verified absolutely. There is little doubt that the species

described by Poppe (1891) under the name C. hendorffi is synony-

mous with C. scutellata. Poppe’s detailed description shows similar

posterolateral projections on the cephalothorax which are absent in

the other species from the Great Barrier Reef. C. hendorffi also

shows great consistency in body size (2: 1.09 mm; d: 1.07 mm),

relative proportions of the caudal rami and P5, and the ventral view

of the female urosome demonstrates the absence of spinular patches

on the second abdominal somite. The only significant discrepancy is

found in the armature of the P2 exopod which Poppe had figured

with an outer spine on the proximal segment. The absence of this

element is a generic character and we suspect that Poppe had

assumed its presence to be the rule in clytemnestrids and had altered

his figure accordingly. Poppe’s (1891) material came from two

localities in the Indian Ocean (West Australian Basin, south of

Madagascar), three localities in the southwest Atlantic off the coasts

of Brazil and Argentina, and the Karimata Strait in the Java Sea. He

also re-identified Thompson’s (1888) material of Goniopsyllus

rostratus from the Maltese Sea as C. hendorffi, confirming its

presence in the Mediterranean. From a zoogeographical point of

view (see below) it appears more conceivable that Thompson had

collected the species described by Claus (1891a) under the name

Goniopelte gracilis, the description of which was unknown to Poppe

(1891). We have been unable to confirm the presence of C. scutellata

in the Atlantic or the Mediterranean and therefore suspect that

Poppe’s records from the southwest Atlantic might have been based

on another species, possibly C. gracilis. Poppe based his illustra-

tions on specimens from the West Australian Basin, suggesting an

Indo-Pacific distribution pattern for C. scutellata.

The redescription by Giesbrecht (1892) has long been accepted as

the basis for identification of C. scutellata even though his material

was not from the type locality. However, from our revision it is clear

that Giesbrecht had redescribed Goniopelte gracilis (see below).

Both species are closely related, sharing the posterolateral projec-

tions on the cephalothorax and the presence of 3 outer spines on

GENERIC CONCEPTS IN CLYTEMNESTRIDAE

Ki \

W AN \\

AN \\

WR | + \ \

KALA

SANK

iN "

SSS

SSSA

Fig. 6 Clytemnestra scutellata Dana, 1847. A, P1 9, anterior; B, P2 Q anterior; C, right P2 exp-3 d, anterior, aberrant setation. [A, B based on neotype].

R. HUYS AND S. CONROY-DALTON

BRASS SY

Fig. 7 Clytemnestra scutellata Dana, 1847. A, P3 @, anterior; B, P4 9, anterior; C, P5 3, anterior. [A, B based on neotype].

GENERIC CONCEPTS IN CLYTEMNESTRIDAE

P2—P4 exp-3 and 6 elements on the P5 exopod in both sexes. They

can be separated by body size, length of caudal ramus setae IV—V,

length of the PS in both sexes and urosome ornamentation in the

female (Table I).

Clytemnestra gracilis (Claus, 1891a) comb. nov.

Goniopelte gracilis Claus, 1891a: 1-10; Taf. III.

Clytemnestra scutellata Dana, 1847 sensu Giesbrecht (1892): 568—

572; Taf. 1, fig. 9; Taf. 45, figs. 16-18, 21, 23-24, 27-30, 32,

34-38.

Clytemnestra rostrata (Brady, 1883) sensu T. Scott (1894): 106—

107; Pl. XII, figs. 47-57; Pl. XII, figs. 1-3.

Clytemnestra scutellata Dana, 1847 sensu Sars (1921): 100-101; PI.

LXVIII.

Clytemnestra scutellata Dana, 1847 sensu Vilela (1968): 44; Est.

XVII, fig. la—c.

Clytemnestra scutellata Dana, 1847 sensu Boxshall (1979): 232;

Fig. 1SA-K.

Clytemnestra scutellata Dana, 1847 sensu Huys et al. (1996): 301;

Fig. 120H.

TYPE LOCALITY. Claus (1891a) collected his material from an

unspecified locality in the eastern Mediterranean. The neotype

designation below redefines the type locality as follows: North-east

Atlantic, south-west of Azores, 35°N 33°W, 0-1 m.

TYPE MATERIAL. Claus’ (1891a) description was based on a single

specimen of either sex. Since the type material no longer exists a

neotype is designated here to secure stability of nomenclature: adult

2in alcohol (BMNH 1999.1024); collected during RRS Discovery

Cruise 121 (5-26 June 1981), station 10379; 13 June 1981, at night;

torpedonet; leg. Institute of Oceanographic Sciences.

OTHER MATERIAL EXAMINED.

(a) from type locality: 11 22 and 8 dd in alcohol (1 2 and 1 3

dissected in half, in separate vials), 1 2dissected on 6 slides (BMNH

1983.53); 2 2Pand 1 d on SEM stub; collection data as for neotype;

(b) Gulf of Guinea, Telegraph Steamer Buccaneer (BMNH

1999.1007—-1016): 9 22 (2 damaged) and 1 6 (damaged); misla-

belled as Clytemnestra rostrata, January—February 1886; leg. J.

Rattray, det. T. Scott. [body length of 7 29: 1381-1541 um, x= 1444

um];

F, Krsinié. [body length: 1309 pm].

DESCRIPTION. (based on Discovery material)

FEMALE. Total body length from tip of rostrum to posterior margin

of caudal rami: 1330-1562 um (xX = 1450 um; n = 10). Maximum

width (382 um) measured at posterior margin of cephalic shield.

| Posterolateral angles of cephalothorax slightly expanded (Fig. 8A).

| General body shape as in type species.

Genital double-somite (Fig. 8B) slightly constricted bilaterally;

original segmentation marked by paired transverse chitinous ribs

| lateroventrally and laterally, joining medially forming continuous

| but weakly defined rib. Copulatory pore slit-like, located medially

| between genital apertures (arrowed in Fig. 27B); leading to short

posteriorly directed, membranous duct connected to bilobate semi-

' nal receptacle. Genital apertures (Fig. 11D) separated by number of

| rounded swellings (also present in type species: Fig. SA); closed off

by small opercula derived from vestigial P6; each with 1 vestigial

seta (coarser than in C. scutellata) at inner distal corner and anterior

| tube-pore near base (arrowed in Fig. 11D).

Urosomites without dorsal ornamentation; penultimate and anal

| somites with multiple rows or patches of spinules around ventral

hind margin and lateroventral patches on second abdominal somite

(Fig. 8B).

Caudal rami (Fig. 8B) as in C. scutellata but setae IV distinctly

shorter than seta V.

Rostrum (Figs 8A; 10C) triangular with rounded anterior margin,

completely fused to cephalothorax; with numerous dorsal surface

pores; minute lateral sensillae flanking middorsal raised pore.

Antennule 7-segmented, with armature formula as in type species.

Antenna, mandible (Fig. 10A), maxillule and maxilla (proximal

endite on syncoxa present) as in type species. Palmar elements of

maxilliped as in Fig. 10B; proximal element fused to basis and with

apical pore; distal element pad-like, forming barbed, linguiform

extension posteriorly and bearing double spinule row and tube pore

anteriorly.

P2—P4 armature formula:

exopod endopod

Pp 1.0228 2221

P3 1.1.323 1.2.32]

P4 1323 1.2.221

P5 (Fig. 8B) elongate, extending clearly beyond posterior margin

of genital double-somite. Exopod about 2.4 times as long as basis,

with 6 setae.

MALE. Total body length from tip of rostrum to posterior margin of

caudal rami: 1420-1531 um (X= 1479 um; n=8). Body with similar

projections as in 2; urosome more slender with genital and first

abdominal somites separate (Fig. 9A).

Antennule with armature as in C. scutellata. Maxilliped much

larger than in 9; middle and distal thirds of palmar margin forming

longitudinal furrow bordered by single row of spinules on both

anterior and posterior sides (Fig. 10D).

P5 (Fig. 9A) very similar to that of 9, extending to distal margin

of first abdominal somite.

Sixth pair of legs (Fig. 9A) weakly asymmetrical, forming highly

membranous midventral area covering single, large median genital

aperture (Fig. 11A); each P6 produced into cylindrical process (Fig.

11B) with 1 apical and 2 lateral bare setae.

Urosomites 4—5 and anal somite with spinules around ventral hind

margin (Fig. 9A).

Caudal rami (Fig. 9A—B) longer and more slender than in 9; setae

III bare; setae IV—V long (68% of urosome length; Fig. 9A) and

plumose; seta VI longer than in 2 and sparsely plumose.

VARIABILITY. Some variability was noticed in the caudal ramus

length of the Buccaneer females, the majority having a slightly

longer ramus than in Fig. 8C. In the Adriatic 2 the spinular patches

on the first postgenital somite are wider medially forming an almost

continuous zone around the posterior margin.

REMARKS. Claus (1891b) himself surmised that Goniopelte graci-

lis was conspecific with Clytemnestra hendorffi which in turn

became relegated to a junior subjective synonym of C. scutellata by

Giesbrecht (1892). It is beyond any doubt that Giesbrecht’s excel-

lent redescription of C. scutellata was based on C. gracilis. His

illustrations were based on Naples material only, however, it is

likely that he included specimens of C. scutellata from the Pacific

(Giesbrecht, 18915) in his length measurements, possibly account-

ing for the lower end of his size range (2: 1.05-1.2 mm; ¢: 1.07-1.3

mm). C. gracilis is distinctly larger than C. scutellata and can be

distinguished from the latter by the slender caudal rami and the

longer P5 which extends clearly beyond the posterior margin of the

genital double-somite in the female and reaches to the rear margin of

16 R. HUYS AND S. CONROY-DALTON

t \

Arte 4

=N x.

Fig. 8 Clytemnestra gracilis (Claus, 1891a) comb. nov. (2) A, Habitus, dorsal; B, urosome, ventral; C, anal somite and right caudal ramus, dorsal.

GENERIC CONCEPTS IN CLYTEMNESTRIDAE

Fig.9 Clytemnestra gracilis (Claus, 1891a) comb. nov. (d) A, Urosome, ven

dorsal. Clytemnestra farrani sp. nov. C, P2 exp-3 9, anterior; D, PS 9, anterior; E, P5 6, anterior.

tral [inset showing setae [V—V]; B, anal somite and right caudal ramus,

R. HUYS AND S. CONROY-DALTON

Fig. 10 Clytemnestra gracilis (Claus, 1891a) comb. nov. SEM photographs. A, Mandibular gnathobase 9; B, maxilliped 9, palmar elements; C, rostrum

9, frontal; D, maxilliped ¢, palmar furrow.

GENERIC CONCEPTS IN CLYTEMNESTRIDAE

Fig. 11 Clytemnestra gracilis (Claus, 1891a) comb. nov. SEM photographs. A, Genital aperture and sixth legs 5; B, P6 d; D, genital field 9 [position of

copulatory pore arrowed]. Goniopsyllus clausi sp. nov. C, Genital aperture and sixth legs 6.

the first abdominal somite in the male. Females of both species can

be differentiated by the ventral ornamentation pattern of the urosome

(C. gracilis has lateral spinular patches on the first postgenital

somite) and the ventral transverse chitinous ridge (marking the

original segmentation of the genital double-somite) which is more

strongly developed in C. gracilis. Giesbrecht (1892) did not illus-

trate the second abdominal somite in the female, however, stated in

the text that spinules were present ventrally around the posterior

margin of all three postgenital somites. Caudal ramus seta IV is

distinctly shorter than seta V in females of C. gracilis (see also

Giesbrecht (1892): Taf. 45, Fig. 27; Sars (1921): Plate LX VII),

while both setae are equally long in the female of the type species.

Both sexes of C. gracilis have a propensity for developing asymme-

try in the caudal rami whereby one ramus is markedly narrower than

the other (see also Claus (1891a): Taf. I, Figs 1-2; Giesbrecht

(1892): Taf. 45, Fig. 27).

Despite his own arguments to the contrary, T. Scott (1894)

inexplicably identified his clytemnestrid material from the Gulf of

Guinea as C. rostrata. A. Scott (1909) re-identified the material as

C. scutellata. Re-examination of the Buccaneer material (BMNH

1893.4.22.268-275) has revealed it to be an amalgamate of two

species, containing 9 22 and 1 6 of C. gracilis and 7 99 of a

smaller Goniopsyllus sp. This might explain the discrepancy found

between the body length reported by T. Scott (1.25 mm) and our

measurements (X = 1.44 mm). Since males are usually larger than

females (Giesbrecht, 1892) it is doubtful whether Marques’ (1973)

male specimen (0.99 mm) of C. scutellata from S40 Tomé (Gulf of

Guinea) belongs to C. gracilis.

The only illustrated record of C. scutellata from northern Europe

is that by Sars (1921) who found a single female in Oslofjord and

described it in great detail. His specimen, 1.24 mm in length, agrees

in all aspects with C. gracilis and represents a significant range

extension for this species. Kasturirangan (1963) reproduced

Giesbrecht’s (1892) and Sars’ (1921) drawings of C. gracilis in his

identification key to the planktonic copepods of Indian coastal

waters, however its presence in the Indo-Pacific has yet to be

confirmed.

Vilela (1968) reported two females of C. scutellata, measuring

1.24-131 mm, from the Portuguese coast off Lisbon. Her illustra-

tions of the caudal rami and PS positively identify her material as C.

gracilis.

Clytemnestra farrani sp. nov.

TYPELOCALITY. Great Barrier Reef, Queensland, Australia. Farran

(1936) recorded a total of 5 specimens (4 belonging to C. farrani, 1

to C. longipes) from serial townettings (his stations 62, 65, 68) at 3

miles east of the laboratory on Low Island (off Port Douglas); depth

32 m.

ETYMOLOGY. This patronym commemorates the late G.P. Farran

for his comprehensive contributions to our knowledge of planktonic

copepods.

TYPE MATERIAL. Holotype 2 dissected on 6 slides (BMNH

1999.998); paratypes are 1 2 and 2 dd in alcohol (BMNH 1999.

999-1001). This material was originally registered as C. scutellata

under reg. no. 1948.4.28.121. Collected during Great Barrier Reef

Expedition 1928-29 on either 15 June (stn 62), 10 July (stn 65) or 18

July 1929 (stn 68).

OTHER MATERIAL EXAMINED. From R. Béttger-Schnack: 1 @ in

alcohol (BMNH 1999.1065); southern Red Sea, Meteor cruise 5/5,

stn 703 (15°34.8' N, 41°54.9' E); 03 August 1987; multiple opening-

R. HUYS AND S. CONROY-DALTON

closing net, 0.055 mm mesh, vertical hauling, 0-50 m (total water

depth 970 m).

DESCRIPTION.

FEMALE. Total body length from tip of rostrum to posterior margin

of caudal rami: 927—946 um (x = 937 um; n = 2). Maximum width

(252 um) measured halfway the cephalic shield length. Posterola-

teral angles of cephalothorax rounded, not expanded (Fig. 12A).

Backwardly produced alate processes of somites bearing P2—P4

distinctly shorter than in C. scutellata and C. gracilis.

Genital double-somite (Fig. 13A) not constricted bilaterally;

original segmentation marked by small, paired, chitinous patches

lateroventrally. Genital field as in type species.

Urosomites without dorsal ornamentation; penultimate and anal

somites with multiple rows or patches of minute spinules around

ventral hind margin and with lateroventral spinular patches on

second abdominal somite (Fig. 13A).

Caudal rami (Fig. 13A, C) shorter than in previous species; setae

IV slightly shorter than seta V but both setae distinctly shorter than

in C. scutellata (only slightly longer than ramus and as long as seta

III) and minutely pinnate.

Rostrum (Fig. 12A) rounded anteriorly, obtuse.

Antennule 7-segmented, with armature formula as in type species.

Antenna, mouthparts (proximal endite on maxillary syncoxa present)

and maxillipeds as in type species.

P2 exp-3 with only 2 outer spines (Fig. 9C). P2—P4 armature

formula:

exopod endopod

P2 He222, 220

P3 1323 APRS)

P4 E328 1.2.221

P5 (Fig. 9D) extending to posterior margin of genital double-

somite. Basis short, exopod about 3 times as long as basis, with 5

setae (3 outer, 1 apical, | inner).

MALE. Total body length from tip of rostrum to posterior margin of

caudal rami: 939-945 um (x = 942 um; n=2). Maximum width (257

lum) measured at posterior margin of cephalic shield. Body (Fig.

12B) with similar projections as in 2; urosome more slender with

genital and first abdominal somites separate (Fig. 13B).

Antennule, antenna, mouthparts and maxilliped with armature as

in C. scutellata.

P5 (Fig. 9E) distinctly shorter than in 2, not extending to distal

margin of first abdominal somite; exopod 1.9 times as long as basis,

apical and inner setae shorter than in @.

Sixth pair of legs (Fig. 13B) weakly asymmetrical; each P6

produced into short cylindrical process with 1 outer and 2 apical bare

setae.

Urosomites 4—5 and anal somite with spinules around ventral hind

margin (Fig. 13B).

Caudal rami (Fig. 13B) stubbier than in 2; setae I-II bare; setae

IV—-V very long (95% of urosome length) and plumose; seta VI

much longer than in 9.

REMARKS. C. farrani can be readily distinguished from its conge-

ners by the swimming leg setal formula, showing only 2 outer spines

on P2 exp-3 but 3 outer spines on P3—P4 exp-3. It is closely related

to C. asetosa which resembles it in the small size, the absence of

posterolateral processes on the cephalothorax and the presence of

only 5 setae on the P5 exopod. The number of endites on the

syncoxa, the spinulation pattern on the female urosome and the

GENERIC CONCEPTS IN CLYTEMNESTRIDAE

v. A, Habitus 2, dorsal; B, habitus 3, dorsal [inset showing setae IV—V at full length].

isp. no

Clytemnestra farran

Fig. 12

22 R. HUYS AND S. CONROY-DALTON

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=——

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0 MUI agts ager i

DRAK\ A} tH Hed Sinyr nae

HAV ghd g lta

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Sa Sea

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7 ae

4 eee Nyy 112 ‘3 All:

x ea

iN = ca

So \

: od A

o hae

i ; o

DUO ASE AULA Ae

MLN GEAR H YE AM 1

A ALIAY AL EP ACL eT

My, SHUNT Ferrin Tene arnt f one mit

Sees

100

PPA soy TD HY 1p 0. Wp

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WSS

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Fig. 13 Clytemnestra farrani sp. nov. A, Urosome &, ventral; B, urosome 6 (excluding P5-bearing somite), ventral [inset showing setae IV—V at full

length]; C, anal somite and right caudal ramus 9, dorsal.

GENERIC CONCEPTS IN CLYTEMNESTRIDAE

ZZZZZ

AAZAAVAAA

200

SDVZ te

—<$<$<——s

ZLDAZ-_ZZDZE

——— ae

SESS

SSS

—=S

Fig. 14 Clytemnestra longipes sp. nov. (3). A, Habitus, dorsal; B, urosome, ventral; C, P2 exp-3; D, P5, anterior; E, right P6.

relative length of the P5 exopod serve to distinguish both species. C.

farraniis currently known only from two widely separated localities

in the Indo-Pacific, suggesting that itis probably widespread through-

out this oceanic basin.

Clytemnestra longipes sp. nov.

TYPE LOCALITY. Great Barrier Reef — see C. farrani sp. nov.

ETYMOLOGY. The species name is derived from the Latin longus

(long) and pes (foot), and refers to the very long male P5 and P6.

TYPE MATERIAL. Holotype d in alcohol (BMNH 1999.997). This

material was originally registered as C. scutellata under BMNH

1948.4.28.121. Collected during Great Barrier Reef Expedition

1928-29 on either 15 June (stn 62), 10 July (stn 65) or 18 July 1929

(stn 68).

DESCRIPTION.

FEMALE. Unknown.

MALE. Total body length from tip of rostrum to posterior margin of

caudal rami: 1211 um. Maximum width (362 um) measured at

posterior margin of cephalic shield. Posterolateral angles of

cephalothorax angular, weakly produced (Fig. 14A). Backwardly

produced alate processes of somites bearing P2—P4 well developed.

Urosome with genital and first abdominal somites separate (Fig.

14B).

Urosomites without dorsal ornamentation; all postgenital somites

with multiple rows of minute spinules around ventral rear margin,

those on urosomites 3, 5 and 6 arranged in paired patches either side

of ventral midline (Fig. 14B).

Caudal rami (Fig. 14B) with bare seta II and minutely pinnate

setae I and III; setae IV—V long (54% of urosome length) and

plumose.

Rostrum (Fig. 14A) rounded anteriorly, protruding. Antennule,

antenna, mouthparts (proximal endite on maxillary syncoxa present)

and maxillipeds as in type species.

P2—P4 exp-3 with only 2 outer spines (Fig. 14C). P2—P4 armature

formula:

exopod endopod

P2 1.1.222 12221

P3 1322 232i

P4 S22 P2221

P5 (Fig. 14D) narrow and elongate, extending to distal margin

of first abdominal somite (Fig. 14B); exopod 2.7 times as long as

basis; with 3 outer seta and | long seta at apex and subdistal inner

corner.

Sixth pair of legs (Fig. 14E) forming very long cylindrical process

with 1 apical and 2 outer bare setae.

REMARKS. The male of this species differs from all known males

in (1) the ventral ornamentation pattern of the urosome, displaying

spinules on all postgenital somites, and (2) the extreme elongation

of the P5 and P6 (the distribution pattern of the 3 elements on the

latter indicate that allometric growth must have happened prima-

rily in the apical portion of the cylindrical process). C. longipes

has the same swimming leg setal formula as C. asetosa but, in

addition to the characters listed above, differs from the latter in

body size and the presence of the proximal endite on the maxillary

syncoxa.

R. HUYS AND S. CONROY-DALTON

Clytemnestra asetosa sp. nov.

TYPE LOCALITY. Suez Canal. Port Taufig, Bay of Suez (Egypt).

ETYMOLOGY. The species name alludes to the absence of the

proximal enditic seta on the maxillary syncoxa.

TYPE MATERIAL. Holotype d dissected on 10 slides (BMNH

1999.1025). Paratypes in alcohol are 3 22,2 dd (1 damaged) and 1

cop. V 6 (BMNH 1999.1026-1031); collected during the Cam-

bridge Expedition to the Suez Canal, 1924. This material was

originally identified as C. scutellata by Gurney (1927) and Boxshall

(i979):

OTHER MATERIAL EXAMINED. From R. Bottger-Schnack: 3

copepodid II stages in alcohol (BMNH 1999.1066—1068); central

Red Sea, Meteor cruise 5/5, stn 682 (21°13.9'N, 38°05.7' E); 25 July

1987; multiple opening-closing net, 0.055 mm mesh, vertical haul-

ing, 10-50 m (total water depth 1890 m).

DESCRIPTION.

FEMALE. Total body length from tip of rostrum to posterior margin

of caudal rami: 758-830 um (x = 801 um; n= 3). Maximum width

(226 um) measured halfway down the cephalic shield. Posterola-

teral angles of cephalothorax rounded, not produced. Backwardly

produced alate processes of somites bearing P2—P4 distinctly shorter

than in C. scutellata and C. gracilis. General body shape (Fig. 15A)

very similar to that of C. farrani (Fig. 12A).

Genital double-somite (Fig. 15B) weakly constricted bilaterally;

original segmentation marked by minute, paired chitinous patches

ventrally. Genital field as in type species.

Urosomites without dorsal ornamentation; penultimate and anal

somites with multiple patches of minute spinules around ventral

hind margin (Fig. 15B).

Caudal rami (Fig. 15C) with bare setae I and II; setae IV slightly

shorter than seta V, both plumose.

Rostrum (Fig. 15A) rounded anteriorly, not distinctly delimited

from cephalic shield.

Antennule (Fig. 16A) 7-segmented, with reduced armature on

segments 2 and 3. Armature formula: 1-[1 plumose], 2-[9 + 1

plumose], 3-[3 + 3 plumose + 1| transformed], 4-[1 + 1 plumose + (1

transformed + ae)], 5-[1], 6-[3], 7-[8 + acrothek].

Antenna with weakly defined exopod (Fig. 17G); one seta fused

basally to segment.

Mandible (Fig. 16B). Palp represented by minute seta; gnathobase

with large lateral tooth (arrowed in Fig. 16C).

Maxillule (Fig. 16D) produced into distal lash (derived from

armature element); with | lateral seta and 1 apical spine.

Maxilla (Fig. 16E) as in type species except for absence of

proximal endite on syncoxa (position in other species arrowed in

Fig. 16E). Maxilliped as in C. scutellata.

Pl (Fig. 17A) asin C. scutellata but setules along inner margin of

enp-1 absent. P2—P4 (Fig. 17B—D) with only 2 outer spines on exp-

3. P2—P4 armature formula:

exopod endopod

PD 1.1.222 1.2.221

128} TEES 22, 12321

P4 1.1.322 12221

P5 (Fig. 17E) nearly extending to posterior margin of genital

double-somite. Basis short, exopod about 2.5 times as long as basis,

with 5 setae (3 outer, 1 apical, 1 inner).

GENERIC CONCEPTS IN CLYTEMNESTRIDAE

—

ry YON aE A by

Santa Wy T9979 vp,

SOU — Wie oly,

apc

Ips

B 25

Fig. 15 Clytemnestra asetosa sp. nov. (9). A, Habitus, dorsal; B, urosome, ventral; C, anal somite and right caudal ramus, dorsal.

26 R. HUYS AND S. CONROY-DALTON

Fig. 16 Clytemnestra asetosa sp. nov. (). A, antennule, ventral [inset showing acrothek at full length]; B, mandible, posterior; C, mandibular gnathobase,

other view [secondary tooth arrowed]; D, maxillule; E, maxilla, posterior [small arrow: exit of maxillary gland; large arrow indicating position of

proximal endite in other Clytemnestra species].

GENERIC CONCEPTS IN CLYTEMNESTRIDAE

“a ZZ bs S 7

Fig. 17 Clytemnestra asetosa sp. nov. (9). A, P1, anterior; B

exopod, anterior; D, P4, distal portion of basis and exopod, anterior; E, PS, anterior; F, rostrum, dorsal; G, antennary exopod.

, P2, intercoxal sclerite, protopod and exopod, anterior; C, P3, distal portion of basis and

R. HUYS AND S. CONROY-DALTON

dal ramus, dorsal.

ntral; C, P5, anterior; D, P6; E, anal somite and right cau

ome, ve

v. (3). A, Habitus, dorsal; B, uros

sp. no

etosa

Fig. 18 Clytemnestra as

GENERIC CONCEPTS IN CLYTEMNESTRIDAE

MALE. Total body length from tip of rostrum to posterior margin of

caudal rami: 920 um (n= 1). Maximum width (232 um) measured at

posterior margin of cephalic shield. Body (Fig. 18A) with similar

projections as in 9; urosome more slender with genital and first

abdominal somites separate (Fig. 18B).

Antennule, antenna, mouthparts and maxilliped with armature as

in C. scutellata.

P5 (Fig. 18C) as in 2 not extending to distal margin of first

abdominal somite (Fig. 18B).

Sixth pair of legs (Fig. 18B, D) weakly asymmetrical; each P6

produced into short cylindrical process with 1 outer and 2 apical bare

setae.

Urosomites 4—5 and anal somite with spinules around ventral hind

margin (Fig. 18B).

Caudal rami (Fig. 18B, E) with bare setae I-II; setae [V—V very

long (75% of urosome length) and plumose; seta VI much longer

than in 9.

REMARKS. The early copepodid stages from the central Red Sea

were identified on the basis of the absence of the proximal endite of

the maxilla and the shape of the cephalothorax. C. asetosa, origi-

nally identified as C. scutellata by Gurney (1927), is the smallest

species in the genus. It is similar to C. farrani in many respects but

differs from it in the armature formula of the antennule, the loss of

the proximal endite of the maxilla, the presence of only 2 outer

spines on P3—P4 exp-3 and a different spinulation pattern on the

female urosome. The species is thus far known only from the Red

Sea and the Bay of Suez.

Clytemnestra hendorffi var. quinquesetosa Poppe, 1891

Poppe (1891) distinguished this variety on the basis of the following

characters: (1) female P5 exopod distinctly longer and bearing 5

setae; (2) urosome of both sexes less slender; (3) caudal rami

relatively wider proximally. This variety was collected from two

localities in the Java Sea. Most authors have followed Giesbrecht’s

(1892) decision to discard this variety and regarded it as a synonym

of C. scutellata. Our revision has revealed that only C. scutellata and

C. gracilis display 6 setae on the P5 exopod and that there are at least

three species in the Indo-Pacific which have only 5 setae. As far as

we could ascertain from the collections examined P5 setation is

never variable within populations and always identical between

sexes. Since Poppe (1891) did not provide any figures it is imposs-

ible to make any positive statement as to the identity of his material.

Other records

Chen et al. (1974) reported C. scutellata from the East China Sea

(one of the areas where Dana originally recorded the species from).

Unfortunately the few illustrations of the habitus and female PS are

_of no help in determining the specific identity of their material.

| Moreover, the extreme body size range (1.0-1.9 mm) strongly

| Suggests the co-occurrence of more than one species in their sam-

| ples. Cheng et al. (1965) also illustrated C. scutellata from the East

China Sea but their species has only 5 setae on the PS exopod, lacks

| posterolateral processes on the cephalothorax and has only 2 outer

| spines on at least P3 (which was mislabelled as the P2) and P4. Their

| reported size range ( 29: 0.86—-1.0 mm; 3d: 0.80-0.85 mm) strongly

| Suggests that they had identified C. asetosa or possibly a related

| species. Mori’s (1929) description of C. scutellata from the Sea of

Japan is equally brief. Posterolateral projections on the cephalothorax

| appear to be absent in his material (although they could be obscured

| by excessive squashing of the figured specimen), indicating that

| Mori was probably dealing with another species. Mori supple-

Kazmi & Muniza (1994) present sketchy figures of what they

believe to be C. scutellata in their samples from the Arabian Sea.

Nothing can be said about the real identity of their material other

than that were dealing with a Clytemnestra.

The Caribbean records of C. scutellata by Owre & Foyo (1967)

and Campos Hernandez & Suarez Morales (1994) require further

investigations. Both descriptions show the unique presence of lat-

eral protrusions halfway down the cephalothorax which may suggest

the occurrence of a distinct species in this region. It is impossible to

decide from Legaré’s (1964) inadequate illustrations whether this

modification also occurred in his Venezuelan material. Interestingly,

Morales & Vargas (1995) show similar protrusions in aclytemnestrid

from the Pacific coast of Costa Rica which they identified as C.

rostratus but has 7 segments in the antennule.

Genus Goniopsyllus Brady, 1883

Sapphir Car, 1890 [type species: S. rostratus Car, 1890 — by

monotypy]

DIAGNOSIS. Clytemnestridae. Body with dorsal pattern of denticles

and spinules on urosomites. Antennule 6-segmented in &, indis-

tinctly 7-segmented in d with segments 34 incompletely fused; 3

segmental homologies: 1—I, 2—(II—-VIII), 3-(IX—XI]), 4—XIII, 5-

(XIV-XVII), 6-(XVIII-XX), 7-(XXI-XXVIII). Antenna with |

lateral and 4 apical elements on distal endopod segment; exopod

represented by membranous segment bearing | long seta. Maxillule

represented by triangular segment with | apical spine. Maxillary

syncoxa with | endite bearing 2 setae.

P1 without outer seta on basis; exopod with 3 setae. P2 with outer

spine on exp-1. P1—P4 armature formula:

exopod endopod

Pl 021 1.1.220

P2 1.1.222 1.2.221

P3 1.1.323 Le PeBVeil

P4 L323 222

P5 exopod with 5 setae in both sexes.

Genital apertures fused in 2 forming common medial slit; closed

off by paired P6 bearing 1 well developed seta; copulatory pore

located medially in large circular depression halfway the length of

the genital double-somite; copulatory duct strongly chitinized.

Male P6 asymmetrical, forming membranous opercula closing

off single (sinistral or dextral) genital aperture; bearing | seta.

Caudal rami convergent, relatively short and conical; not sexually

dimorphic.

TYPE SPECIES. Goniopsyllus rostratus Brady, 1883 [by monotypy]

OTHER SPECIES. G. clausi sp. nov., G. brasiliensis sp. nov.

SPECIES INQUIRENDAE. Goniopsyllus tenuis (Lubbock, 1860)

comb. nov.; Sapphir rostratus Car, 1890

Since the type species is only known from the damaged female

holotype and no other material was available for study, G. clausi sp.

nov. is instead selected for the model description.

Goniopsyllus clausi sp. nov.

Clytemnestra rostrata (Brady, 1883) sensu Giesbrecht (1892): pp.

568-572; Taf. 45, Figs 22, 31.

Clytemnestra rostrata (Brady, 1883) sensu Vilela (1965): p. 21; Est.

IX, Fig. 2a—e; (1968): p. 44; Est. XVII, Fig. 2a—c.

Fig. 19 Goniopsyllus clausi sp. nov. A, Habitus °, dorsal; B, habitus of ovigerous 9, lateral.

R. HUYS AND S. CONROY-DALTON

GENERIC CONCEPTS IN CLYTEMNESTRIDAE Sil

Fig. 20 Goniopsyllus clausi sp. nov. A, Antennule 9, ventral; B, distal portion of antennulary segment 6 of 9, ventral [rudimentary element arrowed]; C,

antennule 3, ventral; D, antennulary segments 3-6 of 4, anterior; E, antennulary segment 7 of 3, ventral [rudimentary element arrowed].

32 R. HUYS AND S. CONROY-DALTON

~—_— Aa 3

SS ——

ae L-

—> el

Fig. 21 Goniopsyllus clausi sp. nov. A, antenna 9, outer; B, distal endopod segment of antenna @, inner [rudimentary elements arrowed]; C, mandible 9; D,

mandibular gnathobase 9; E, mandibular gnathobase of d specimen; F, maxillule 9, posterior; G, maxilla &, posterior [exit of maxillary gland arrowed]; H,

oral area 2 showing position of antenna (A,), labrum, paragnaths (P), mandible, maxillule, maxilla and maxilliped (Mxp.); I, rostrum Q, dorsal.

GENERIC CONCEPTS IN CLYTEMNESTRIDAE

WY quan rnannnsnnssssees

PL Stl ely

epee Mae

SSQOOn~sasyy

S WWMM

yyy

Sys

epee”

(o)

medial; D, same,

maxilliped 2 distal portion of basis and endopod, anterior; C, same

maxilliped 9, anterior;

posterior; E, maxilliped d, anterior; F, maxilliped d, distal portion and endopod, anterior; G, maxillipedal basis and endopod 6, medial; H

posterior.

Fig. 22 Goniopsyllus clausi sp. nov. A

same,

Clytemnestra rostrata (Brady, 1883) sensu Huys et al. (1996): pp.

300-303, Figs 120A—G, 121A—D.

Clytemnestra rostrata (Brady, 1883) sensu Boxshall & Huys (1998):

p. 782, Fig. 13(a)—(b).

Bay of Cadiz, 36°30'N 7°20'W (Spain).

ETYMOLOGY. The species is named in honour of Carl Claus, one of

the most prolific 19th century copepodologists, who first called

attention to the distinctiveness of the clytemnestrid genera.

TYPE MATERIAL. Holotype 2 dissected on 10 slides (BMNH

1999.1035). Paratypes are 2 dissected dd (on 2 and 5S slides,

respectively), 2 dissected 2(on 1 slide each), and9 2° (1 damaged),

1 3, 4 copepodids (2 Cop V, 1 Cop IV, 1 Cop II) in alcohol (BMNH

1999.1036-1055). In addition, 2 29 and 1 d were prepared for

SEM. Donated by J.M. Gee, collected by A. Lindley (Plymouth

Marine Laboratory), 1984.

TYPE LOCALITY.

OTHER MATERIAL EXAMINED. 4 99, 2d: Adriatic Sea, Station

CJ-008, Pelegrin, Hvar (Croatia), leg. F. KrSini¢é, ‘Bios’, 23 May

1998 (BMNH 1999.1072-1077).

DESCRIPTION.

FEMALE. Total body length from tip of rostrum to posterior margin

of caudal rami: 979-1067 um (x = 1017 um; n= 8). Maximum width

(306 um) measured at posterior margin of cephalic shield. Postero-

lateral angles of cephalothorax only weakly expanded laterally but

markedly produced posteriorly (Fig. 19A, B). Somites bearing P2—

P4 successively decreasing in width posteriorly and bearing

backwardly produced alate processes.

Genital double-somite (Figs 23A; 27C) slightly constricted bilat-

erally; original segmentation marked by two minute chitinous patches

ventrally. Copulatory pore (Figs 23C, D; 27A, C) located medially

in large circular depression, halfway the length of genital double-

somite; leading to anteriorly directed, strongly chitinized duct which

at level of P5-bearing somite enters median seminal receptacle.

Genital apertures located far anteriorly; closed off by small opercula

derived from vestigial P6; each with 1 well developed seta (Figs

23C; 27D).

Urosomites with zone of small denticles around dorsal hind

margin (not figured in Fig. 19A, but see Fig. 23B); penultimate and

anal somites also with larger spinules around ventral hind margin

(Fig. 23A).

Caudal rami short (Figs 23B; 26A), convergent; conical in shape

with stepped inner and outer margins marking insertion sites of setae

I, 11 and [V—V; produced into conical process bearing terminal pore;

with numerous ventral pores as illustrated in Fig. 26A. Setae LI

bipinnate, spiniform and strongly developed; seta I 1.85 times as

long as seta II, extending beyond apex of caudal ramus. Seta III

minutely bipinnate. Setae IV—V basally fused, without fracture

planes, multipinnate and spiniform; seta V about 2.1 times ramus

length. Seta VI minute, bare; seta VII biarticulate at base, bare.

Rostrum (Figs 19A; 211) triangular and well offset, completely

fused to cephalothorax; with numerous dorsal surface pores as fig-

ured, none on ventral surface; with minute lateral sensillae near apex.

Antennule (Fig. 20A) slender, 6-segmented; segment 6 very long.

Plumose setae present on segments 14. Segment | with small pore

near seta and few long setules along anterior margin. Armature

formula: 1-[1 plumose], 2-[6 + 1 plumose + 3 pinnate], 3-[5 + 2

plumose + | transformed], 4-[1 + 1 plumose + (1 transformed + ae)],

5-[1], 6-[11 + acrothek]. Apical acrothek consisting of aesthetasc,

long transformed seta and short bare seta. Transformed setae on

segments 3, 4 and 6 long and aesthetasc-like, with minutely spiniform

tip; those on segments 4 and 6 basally fused to aesthetasc. Rudimen-

R. HUYS AND S. CONROY-DALTON

tary element present at base of acrothek (arrowed in Fig. 20B).

Antenna (Fig. 21A, B) 4-segmented, comprising coxa, basis and

2-segmented endopod. Coxa well developed, bare. Basis and proxi-

mal endopod segment with few surface denticles; unarmed. Exopod

inserted in membranous area between basis and endopod; repres-

ented by small, weakly chitinized segment bearing strong recurved

seta apically; exopodal seta multipinnate, spinules in proximal third

distinctly longer. Distal endopod segment with 3 surface frills and

minute denticles on outer surface and patch of long setules on medial

surface; lateral armature consisting of 1 pinnate seta; distal armature

consisting of 1 subapical and 3 apical, non-geniculate, bipinnate or

multipinnate elements, 2 of which spiniform, recurved and bearing

long spinules proximally; distal margin with 2 rudimentary elements

on inner surface (arrowed in Fig. 21B).

Labrum (Fig. 21H) large, with 6 secretory pores on anterior

surface; distal margin smooth medially and with spinular patch on

either lateral lobe.

Mandible (Fig. 21C—E) reduced. Palp represented by single na-

ked seta. Gnathobase long and narrow, stylet-like; produced into

number of cuspidate processes apically and subapically; without

dorsal seta(e).

Paragnaths (Fig. 21H) well developed lobes without any con-

spicuous ornamentation.

Maxillule (Fig. 21F) reduced; represented by small triangular

segment bearing naked apical seta and raised pore along outer

margin.

Maxilla (Fig. 21G, H) 2-segmented, comprising elongate syncoxa

and allobasis. Syncoxa with expanded basal portion; exit of maxil-

lary gland large (arrowed in Fig. 21G), partly concealed under

lobate extension; coxal endite cylindrical, with 2 naked setae apically.

Allobasis with large articulating claw distally, smaller inner spine

and unipinnate seta along outer margin.

Maxilliped (Fig. 22A) very large, articulating with well devel-

oped pedestal; 3-segmented, comprising syncoxa, basis and endopod.

Syncoxa extremely elongate, longer than basis; without ornamenta-

tion but with 1 anterior, plumose seta near membranous articulation

with basis. Basis elongate; distal third of palmar margin with double

spinule row and 2 elements located closely to articulation with

endopod (Fig. 22B—D); proximal element spiniform and bare, distal

element stubby and spinulose. Endopod represented by short seg-

ment bearing naked claw; accessory armature consisting of 3 anterior

setae and 2 posterior setae (Fig. 22B—D).

Swimming legs with wide, narrow intercoxal sclerites and well

developed praecoxa; both without ornamentation. Rami 3-segmented

except for Pl exopod.

P1 (Fig. 23E) separated from maxillipeds by large membranous

area. Coxa and basis prolonged along dorsoventral axis; without

surface ornamentation. Basis without inner or outer seta (spine).

Exopod 1-segmented, represented by elongate segment bearing long

setules along outer margin; with subapical pore and 3 setiform

elements distally, outer one less than half the length of others.

Endopod 3-segmented; segments decreasing in size distally, each

with anterior pore and few spinules/setules along outer margin; enp-

1 with very long inner seta; ornamentation of inner elements typically

(multi)pinnate, distal elements plumose.

P2—P4 (Figs 24A, B; 25B) with transversely prolonged basis

bearing short outer seta. Endopods distinctly longer than exopods.

Exopodal outer spines setiform with distinct flagellate tip. Exopod

segments typically with pore near outer distal corner; without

ornamentation. Endopods with long proximal segment, particularly

in P2—P3; segments with anterior pore, setules along outer margin

and spinules (enp-2 and -3) or setular tuft (enp-1) on posterior

surface; setal ornamentation typically combination of setular and

: GENERIC CONCEPTS IN CLYTEMNESTRIDAE

x <>

ara INN

(iA aa aE “SS. > Sac SR

CR EN a.

o “0-6 V4 dif 25

Minas: 8

genital field, ventral; D, genital field,

Fig. 23 Goniopsyllus clausi sp. nov. (2). A, Urosome, ventral; B, anal somite and left caudal ramus, dorsal; C,

lateral; E, P1, anterior.

R. HUYS AND S. CONROY-DALTO:

ee,

Fig. 24 Goniopsyllus clausi sp. nov. (2). A, P2, anterior; B, P3, anterior; C, P5, anterior; D, aberrant P5, anterior.

3i)/

GENERIC CONCEPTS IN CLYTEMNESTRIDAE

—THFALWY

TREE

(aa)

caarsl| 1 BIOS

\ | il \ f ey pe

VALS LL eg

Jee" — VAFL IT PSL

Ba SZ

. SOS

< SI

}

SKK LIES

== EES

SSS SSS

7 S Le

{

Oe (\ <— Es ae = SOS : WY

~~ x 2 = NN Se SSS SSS LAA: g Y \

= < — > “SS SSS ‘

Ry, ~ NEN Y ,

Y y/

Aan

}

ie}

vey

‘S)

ie)

B=)

-Q

ie}

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(e}

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ire)

—_

R. HUYS AND S. CONROY-DALTON

Ss:

sp I al

ventral. Goniopsyllus rostratus Brady, 1883 (holotype @). C, Antennule

, maxilliped, distal portion of basis and endopod, anterior; F, P5, posterior; G, anal somite and left caudal

®, lateral; B, urosome 6,

Fig. 26 Goniopsyllus clausi sp. nov. A, Caudal ramus

13)

(armature omitted); D, maxilliped, anterior;

ramus, dorsal.

GENERIC CONCEPTS IN CLYTEMNESTRIDAE

Fig. 27 Goniopsyllus clausi sp. nov. (2). SEM photographs. A, Circular depression surrounding copulatory pore (position obscured by remnant of

spermatophore neck); C, genital double-somite; D, genital aperture. Clytemnestra gracilis (Claus, 1891a) comb. nov. (?). B, Genital apertures and

copulatory pore [arrowed].

spinular rows; inner seta of P2—P3 enp-1 short. Spine and setal

formula of swimming legs as for genus.

P5 (Fig. 24C) uniramous, laterally displaced; 2-segmented, com-

prising basis and 1-segmented exopod; not extending to distal

margin of genital double-somite (Fig. 23A). Basis with short outer

seta and pore near outer distal corner. Exopod about twice as long as

basis, slightly curved inwards; outer margin with 2 pinnate setae and

3 pores; inner margin with long plumose seta; apex with 1 pinnate

and 1 plumose seta.

MALE. Total body length from tip of rostrum to posterior margin of

caudal rami: 1021 um (n= 1). Maximum width (304 um) measured

at posterior margin of cephalic shield.

Body (Fig. 25A) with similar projections as in 9; genital and first

abdominal somites separate.

Rostrum (Fig. 25A) more pointed than in 2.

Antennule (Fig. 20C) slender, indistinctly 7-segmented with seg-

ment 4 only demarcated dorsally (Fig. 20D); haplocer, with

geniculation located between segment 6 and 7. Plumose setae

present on segments 1—5. Segment 1 with small pore near seta and

few long setules along anterior margin. Armature formula: 1-[1

plumose], 2-[5 + 5 plumose], 3-[5 + 1 plumose + | pinnate + 1

transformed + ae], 4-[2 plumose], 5-[4 plumose + 1 pinnate+ (1

transformed + ae)], 6-[1 + 2 pinnate spines + 1 smooth spine], 7-[10

+ 2 vestigial elements + acrothek]. Apical acrothek consisting of

aesthetasc, long transformed seta and short bare seta. Transformed

setae on segments 3, 5 and 7 long and aesthetasc-like, with minutely

spiniform tip; those on segments 5 and 7 basally fused to aesthetasc.

Rudimentary element present at base of acrothek (arrowed in Fig.

20E). Segment 6 with continuous patch of spinules on anterior

surface (Fig. 20D). Segment 7 with 2 vestigial elements near

geniculation.

Maxilliped (Fig. 22E) very large, articulating with well devel-

oped pedestal; 3-segmented, comprising syncoxa, basis and endopod.

Syncoxa extremely elongate but not longer than basis; without

ornamentation but with | anterior, plumose seta near membranous

articulation with basis. Basis elongate; more swollen than in &,;

middle and distal thirds of palmar margin forming longitudinal

furrow bordered by multiple rows of spinules on both anterior and

posterior sides; with 2 elements located closely to articulation with

endopod; proximal element spiniform and bare, distal element

stubby and spinulose. Endopod represented by short segment pro-

duced into very long naked claw which in reflexed position typically

fits in palmar furrow with the apical part closely adpressed onto the

anterior surface of the basis (Fig. 22E, G); accessory armature

consisting of 3 anterior setae and 2 posterior setae (Fig. 22F—H).

P5 (Fig. 25C) very similar to that of 9, with identical proportions

and setation but lateral setae of exopod slightly shorter.

Sixth pair of legs (Figs 11C; 26B) asymmetrical, represented by

highly membranous non-articulating flaps covering single, large

genital aperture (Fig.11C); each lobe with 1 bare seta at outer distal

corner.

Urosomites 4—S and anal somite with spinules around ventral hind

margin (Fig. 26B).

Caudal rami (Fig. 26B) slightly more slender than in 9; conical

projection wider and setae I-II relatively shorter.

Spermatophore with very long, recurved neck (Fig. 26B).

VARIABILITY. The left P5 of the holotype @ shows slightly

different segmental proportions and pore pattern (Figs 23A; 24D).

REMARKS. This species was illustrated by Huys et al. (1996) as

‘Clytemnestra rostrata’. Their brief description which was based on

material from the Gulf of Cadiz contains some observational errors.

R. HUYS AND S. CONROY-DALTON

The most significant is the setation of the maxillule which was

actually based on C. gracilis. The armature on the genital field was

omitted in their Fig. 120B. The female P5 (their Fig. 121C) also

appears shorter but this is to be regarded as the result of excessive

squashing during mounting.

The distribution of G. clausi is thus far restricted to the Portu-

guese coast (Vilela, 1965, 1968) and the Mediterranean with

confirmed records from the Bay of Cadiz, Naples and the Adriatic.

Sapphir rostratus has also been recorded from the Adriatic but is

probably not synonymous with G. clausi (see below). The Naples

record refers to Giesbrecht (1892) who found 1 6 of ‘C. rostrata’ in

this area but also attributed Pacific specimens (3 29, 2 53) to this

species.

Goniopsyllus rostratus Brady, 1883

Clytemnestra rostrata (Brady, 1883) Poppe (1891)

TYPE LOCALITY. South Atlantic, off Argentinean coast; 42°32'S

56°29' W; net at 54 m depth.

MATERIAL EXAMINED. Holotype 2 dissected on slide (reg. no.

C.C.46); collected during Voyage of H.M.S. Challenger during the

years 1873-1876 (station 318); 11 February 1876. The dissection is

imperfect and incomplete (e.g. antenna and P1 are lacking), and the

specimen is partly aberrant in the swimming leg setal formula.

REDESCRIPTION.

FEMALE. Genital double-somite (Fig. 28A) relatively short in

comparison with other species, not constricted bilaterally; original

segmentation marked by two minute chitinous patches ventrally.

Copulatory pore (Fig. 28A) located medially in large circular de-

pression, halfway the length of genital double-somite; leading to

anteriorly directed, strongly chitinized duct which at level of P5-

bearing somite enters median seminal receptacle. Genital apertures

located far anteriorly; closed off by small opercula derived from

vestigial P6; each with 1 well developed seta.

Urosomites with zone of small denticles around dorsal hind

margin; penultimate and anal somites also with larger spinules

around ventral hind margin (Fig. 28A).

Caudal rami short (Figs 26G; 28A), convergent; similar in shape

to G. clausi but proportionally smaller. Setae II bipinnate, spiniform

and strongly developed; seta I 1.7 times as long as seta II, extending

beyond apex of caudal ramus. Seta III minutely bipinnate. Setae [V—

V basally fused, without fracture planes, multipinnate and more

setiform and distinctly longer than in G. clausi (compare Fig. 23B);

seta V about 3 times ramus length. Seta VI minute, bare; seta VII

biarticulate at base, bare.

Antennule (Fig. 26A) slender, 6-segmented; segment 6 longer

than in G. clausi (length ratio segment 6 : segment 5 being 6.0 in G.

rostratus, 5.0 in G. clausi). Armature pattern as in G. clausi.

Maxilliped (Fig. 26D) with similar armature as in G. clausi but

with different spinular ornamentation on palmar margin (Fig. 26E).

P2—P4 spine and setal formula of swimming legs as follows (left

P3 exp-3 and right P4 exp-3 with aberrant outer spine number):

Exopod Endopod

Right Left

P2 1.1.222 222 1.2.221

P3 Likey3) LES 22 L221

P4 1.1.322 1S 23 1.2.221

P5 (Fig. 26F) 2-segmented, comprising basis and 1-segmented

exopod;relative lengths as in G. clausi. Exopod outer margin with 2

GENERIC CONCEPTS IN CLYTEMNESTRIDAE

AVY 100 ns

TL Nt i

Ls Cie et i)

a <=

I Ay atadrnheS

eS ——

21778,

Roney i

Fig. 28 Goniopsyllus rostratus Brady, 1883 (holotype ¢). A, Urosome (excluding P5-bearing somite), ventral [distorted due to excessive squashing].

Goniopsyllus brasiliensis sp. nov. (2). B, Urosome, ventral; C, genital field, ventral; D, antennule (armature omitted).

pinnate setae and 3 pores; inner margin with long plumose seta; apex

with | pinnate and 1| plumose seta.

MALE. Unknown.

REMARKS. Upon re-examination Boxshall (1979) concluded that

the holotype, identified by Brady (1883) as a male, was in reality

female. The true sexual identity however, had already been noted by

both Poppe (1891) and Claus (1891a—b) who based their conclusion

on the 5-segmented urosome and the female facies of the antennule

and maxilliped. This opinion was also confirmed by Giesbrecht

(1892) but not by Car (18915) who continued regarding it as a male

on the basis of the internal spermatophore drawn by Brady. The most

plausible explanation is that Brady (1883) had misinterpreted the

strongly chitinized copulatory duct, a suspicion reinforced by in-

spection of the holotype.

Giesbrecht (1892: 573) pointed out the discrepancy between the

size mentioned in Brady’s text and that inferred from his habitus

figure reproduced at x80 magnification. According to Brady the

holotype is only 0.65 mm long (*1-40th of an inch’) but Giesbrecht

considered 1.16 mm a more realistic figure. Re-examination of the

slides strongly suggests that Brady must have made a morphometric

error of at least a factor 2. The urosome (excl. P5-bearing somite)

which is mounted intact measures 0.43 mm. Extrapolation by using

the urosome/body length ratio found in its congeners G. clausi and

G. brasiliensis (about 0.3) gives an estimated total body length of

1.43 mm. This large size rules out possible conspecificity with G.

brasiliensis (x = 0.96 mm).

Brady (1883) assumed all four swimming legs to be similar,

having 3-segmented rami and resembling the leg illustrated in his

Fig. 15 (i.e. the P2). His lateral habitus view suggests that the P1

possesses 3-segmented exopods and endopods, however Poppe

(1891) suspected that Brady had overlooked the exopod and instead

had superimposed both left and right endopods. For some unknown

reason he assumed the P1 exopod to be 2-segmented but failed to

confirm this against the holotype due to the absence of the P1 on

Brady’s slide.

G. rostratus can be readily identified from the other South-

American species G. brasiliensis by the large body size (compare

urosomes in Fig. 28A—B drawn at the same scale), the elongate

caudal ramus setae [V—V, the long seta I clearly extending beyond

the distal margin of the ramus, and additional differences in the

ornamentation of the maxilliped (spinule pattern on palmar margin).

Brady (1883) also illustrated well developed posterolateral exten-

sions on the cephalothorax which are completely absent in G.

brasiliensis.

Goniopsyllus brasiliensis sp. nov.

? Clytemnestra rostrata (Brady, 1883) sensu Ramirez (1966): 291;

Lam. II, figs 12-15.

TYPE LOCALITY. Rio Grande do Sul (Brazil); outside opening of

Lagoa dos Patos to ocean; 32°11'S 52°7'W.

ETYMOLOGY. The species name refers to the type locality.

TYPE MATERIAL. Holotype @ dissected on 8 slides (BMNH

1999.1056). Paratypes are 8 2° in alcohol (BMNH 1999.1057—

1064). Collected by G.A. Boxshall, February 1996, plankton haul.

DESCRIPTION.

FEMALE. Total body length from tip of rostrum to posterior margin

of caudal rami: 892-1057 um (x = 958 um; n= 8). Maximum width

(265 zm) measured at posterior margin of cephalic shield. Postero-

R. HUYS AND S. CONROY-DALTON

lateral angles of cephalothorax rounded, virtually not expanded

laterally (Fig. 29A). Rostrum (Fig. 29A) rounded and less pro-

nounced than in G. clausi. Backwardly produced alate processes of

somites bearing P2—P4 distinctly shorter and less pointed than in C.

clausi. Integument generally less chitinized than in G. clausi.

Genital double-somite (Fig. 28B) not constricted bilaterally and

relatively wider than in G. clausi; original segmentation marked by

minute, paired, chitinous patches ventrally. Genital field as in G.

clausi but with additional pores flanking copulatory pore (Fig. 28C).

Urosomites with zone of small denticles around dorsal hind

margin (Fig. 29B); penultimate and anal somites also with larger

spinules around ventral hind margin (Fig. 28C).

Caudal rami (Figs 28B; 29A—C) short, convergent. Setae III

bipinnate, spiniform and strongly developed; seta I 1.2 times as long

as seta II, not extending beyond apex of caudal ramus. Seta III

minutely bipinnate. Setae [V—V basally fused, multipinnate and

about as long as in G. clausi but seta IV more resilient (compare Fig.

23B); seta V about 1.5 times ramus length. Seta VI extremely small;

seta VII biarticulate at base, bare.

Antennule (Fig. 28D) slender, 6-segmented; segment 2 shorter

than in G. clausi but armature pattern identical.

Mandible and maxillule (Fig. 29D) somewhat more slender than

in G. clausi.

Maxilliped (Fig. 29E—-F) with similar armature as in G. clausi but

with different spinular ornamentation on palmar margin (Fig. 29F).

P1—P4 with setal formula as for genus.

P5 (Fig. 28B) markedly longer than in G. clausi, extending

beyond distal margin of genital double-somite.

MALE. Unknown.

REMARKS. Although many South-American authors have recorded

specimens that they attribute to C. rostrata, there is good reason to

believe that in fact often they have mistaken G. brasiliensis for this

species. In general, with the discovery of G. brasiliensis many of the

Brazilian records of G. rostratus are rendered doubtful (Bjérnberg,

1963; Bjornberg et al., 1981; Campaner, 1985; Carvalho, 1944;

Gaudy, 1963; Monti, 1980; Monti & Gloeden, 1986; Monti &

Cordeiro, 1988; Santos, 1973; Vega-Perez, 1993). The same applies

to Legaré’s (1961, 1964) records of C. rostratus from Venezuelan

coastal waters. The species illustrated by Ramirez (1966) as C.

rostrata from Mar del Plata in Argentina differs from the one figured

in his later paper (Ramirez, 1970) by the complete absence of

posterolateral projections on the cephalothorax and is almost cer-

tainly conspecific with G. brasiliensis. The author described the

female antennule as 7-segmented but this clearly contradicts his

illustration which shows only 6 segments as in other species of

Goniopsyllus. The only anomaly remaining is the body size which

according to Ramirez (1966) is 1.8 mm for the female and 1.5 mm

for the male. Based on his illustrations and the accompanying scale

bars the female only measures 0.74 mm and the male 0.77 mm.

It is not clear whether Carvalho’s (1952) material of C. rostrata,

consisting of 5 males from the Bay of Santés (Sao Paulo State), also

belongs to C. brasiliensis. His size range (0.50—0.85 mm) precludes

possible identity with C. rostratus but the illustrations accompany-

ing the brief description are completely worthless and erroneous.

The caudal rami are exceptionally long for this genus, the PS exopod

has only 4 elements, and the antennule is 8-segmented. The speci-

mens reported from Guaratuba (Parana State) in an earlier paper

(Carvalho, 1944) are also very small (0.5 mm) and their fragmentary

description is equally useless for identification purposes.

Finally, there is no possibility of identifying any specimens from

Campos-Hernandez & Suarez-Morales’ (1994) illustrations of C.

rostrata from the Gulf of Mexico.

GENERIC CONCEPTS IN CLYTEMNESTRIDAE

Vy ovvwo

o yan vy

WV Vi te y

yury yy WY, Yoye

¥ Wyvavyy vr vy

Fig. 29 Goniopsyllus brasiliensis sp. nov. (@). A, Habitus, dorsal; B, anal somite and caudal rami, dorsal; C, caudal ramus, lateral; D, mandible and

maxillule; E, maxilliped, posterior; F, maxilliped, distal portion of basis and endopod, anterior.

Goniopsyllus tenuis (Lubbock, 1860) comb. nov.

Clytemnestra tenuis Lubbock, 1860

Lubbock’s (1860) description is very incomplete and based on a

single specimen. The antennule was figured as 7-segmented but

comparison with other clytemnestrid descriptions indicates that the

author had erroneously shown the second segment as subdivided

into two distinct segments. The segmentation of the distal half of the

antennule conforms with the Goniopsyllus pattern, justifying its

placement in this genus. Giesbrecht (1892) regarded C. tenuis as a

likely synonym of G. rostratus but in the light of the discovery of

several closely related species we regard this course of action

premature. Conversely, Marques (1973) listed C. tenuis in the

synonymy of C. scutellata. Although Lubbock doubted the sexual

maturity of the holotype female this is contradicted by his state-

ments that the specimen was ovigerous and that the second and third

abdominal somites had almost completely coalesced (this being in

conflict with his illustration of a 6-segmented urosome lacking any

trace of a genital double-somite). With the scanty information

available it is extremely unlikely that C. tenuis will ever be recog-

nized; it is ranked here as species inquirenda.

Sapphir rostratus Car, 1890

Conspecificity between S. rostratus, described from Trieste (North

Adriatic), and G. clausi, recorded from the South Adriatic (this

paper), seems conceivable on zoogeographical grounds. The rela-

tive lengths of the distal antennulary segments in both sexes and the

length of caudal ramus seta II, however, do not agree with those of

G. clausi. It is questionable whether these discrepancies are real or

reflect observation bias since Car’s (1890) illustrations contain

other, more significant errors such as the P5 which is shown with

only 3 setae and the P4 which allegedly lacks an outer spine on the

distal endopod segment. A final obstacle to conspecificity is the

small size of S. rostratus which, based on the dorsal view of the

male, measures only 0.58 mm. Rather than proposing a new replace-

ment name in anticipation of potential secondary homonymy with

the type species, we maintain this species as species inquirenda

under its current name. If S. rostratus and G. clausi are conspecific

then the former becomes a invalid senior synonym of the latter.

Other records

Monard’s (1928) description of “C. rostrata’ from Banyuls-sur-Mer

contains several inconsistencies such as his illustration of the P5

exopod which shows only 4 setae and his statement that the P2—P4

enp-3 setal pattern is 6-5-5, indicating that he has confounded P2

and P3. The author also claims that the male P5 is modified and the

female antennule 7-segmented. The small size (0.65 mm) seems to

tule out conspecificity with G. clausi.

Chen et al.’s (1974) record of G. rostratus from the East China

Sea and Mori’s (1937) from Japanese waters are indeterminable on

the basis of the few illustrations provided. The short female P5

suggests a species different from G. rostratus. Similarly, Marques

(1958) did not give convincing evidence for her record from Angola

since only the habitus of the male and body length measurements (2

: 0.4- 0.94 mm; d: 1 mm) were provided.

DISCUSSION

Generic concepts and species discrimination

The generic concepts of Goniopsyllus and Clytemnestra (as

R. HUYS AND S. CONROY-DALTON

Goniopelte) introduced by Claus (1891b), but dismissed by subse-

quent authors, are reinstated here. Claus based the distinction on

differences in antennule segmentation and setation of the antennary

exopod, and on the presence or absence of sexual dimorphism in the

caudal rami. Goniopsyllus is clearly more advanced than

Clytemnestra, being illustrated by several reductions in the cephalic

appendages, P1 and male P6 which provide additional discrepancies

between both genera. In Goniopsyllus the number of distal setae on

the antennary endopod is reduced (the missing elements being

marked by rudiments; arrowed in Fig. 21B), the armature of the

maxillule is represented by a single apical element, the distal

syncoxal endite of the maxilla bears only 2 elements and the long

syncoxal seta representing the proximal endite is lost. The latter

character should be used with caution in generic discrimination

since convergent loss of the proximal endite has happened in at least

one representative of Clytemnestra (Fig. 16E). All species of

Goniopsyllus lack the outer basal seta of P1 and have lost the inner

seta of its exopod. The male sixth legs are weakly developed bearing

only 1 seta in Goniopsyllus (Fig. 11C) but are produced into con-

spicuous, elongate, trisetose processes in Clytemnestra (Fig. 11A—B),

resembling the condition found in the Aegisthidae and Cerviniidae.

Although Clytemnestra is the more primitive genus, it can be

readily identified by the absence of the outer spine on P2 exp-1. As

far as we could ascertain this is a unique character in harpacticoids

with a 3-segmented P2 exopod. The caudal ramus sexual dimor-

phism displayed only by Clytemnestra requires further ontogenetic

study before it can be considered a potential autapomorphy for the

genus. The typical caudal ramus condition found in the majority of

the Harpacticoida shows normally developed terminal setae IV and

V. In the Clytemnestridae this condition is exhibited only by the

males of Clytemnestra (e.g. Fig. 5B), the atypical female state (Fig.

5A) showing reduced setae. In contrast to swimming leg sexual

dimorphism which is nearly always the result of deviations in male

ontogeny, secondary sexual characters in the caudal rami are exclu-

sively expressed by the female, and as a rule are not expressed until

the final moult. This timing of expression has been demonstrated in

various families displaying caudal ramus sexual dimorphism, in-

cluding the Canuellidae, Cylindropsyllidae and Canthocamptidae.

In these families it is intrinsically linked with precopulatory mate

guarding where female caudal ramus modification shows substan-

tial congruence with male antennule morphology. Since the atypical

condition in female Clytemnestra is also found in both sexes of

Goniopsyllus — and thus unlikely to be the result of transformation at

the final moult — a different ontogenetic explanation must apply.

This is further corroborated by examination of early copepodids

(including Cop V 3d) of C. asetosa and G. clausi which revealed

similarly reduced caudal setae in both species. The male caudal

setae in Clytemnestra must therefore undergo transformation at the

final moult. Hence, it is assumed here that reduction of setae [V—V

represents the ancestral state in the family and that elongation

evolved only secondarily in male Clytemnestra, not being linked to

mate guarding but possibly enhancing its capacity during mate

location.

Examination of the genital field has revealed significant differ-

ences between both genera. In Goniopsyllus the copulatory pore is

located halfway down the genital double-somite in a large circular

depression (Fig. 27A) and connects via a strongly chitinized duct

with the anteriorly positioned seminal receptacles (Fig. 23C—D). In

Clytemnestra the copulatory pore is represented by a posteriorly

directed minute slit (arrowed in Fig. 27B), located between the

genital apertures far anteriorly on the genital double-somite, and a

copulatory duct is hardly differentiated (Fig. SA). The polarity of

copulatory pore displacementis difficult to assess, however, outgroup

GENERIC CONCEPTS IN CLYTEMNESTRIDAE 45

Table 1 Diagnostic characters of Clytemnestra species [Al = antennule; GDS = genital double-somite; AS = first adominal somite]. Length measure-

ments are based on material examined in this paper.

scutellata gracilis farrani longipes asetosa

size ? (in um) 1121 1309-1562 927-947 2 758-830

size 3 (in pm) 1064 1420-1531 939-945 1211 920

cephalothoracic processes present present absent obsolete absent

setal number segment 2 Al 12 12 12 u 10

proximal endite maxilla present present present present absent

P2 exp-3 formula 223 223 222 222 222

P3 exp-3 formula 323 323 323 322 322

P4 exp-3 formula 323 323 323 322 322

setal number P5 exopod $/d 6 6 5 5 5

P5 apex 2 vs GDS posterior margin coinciding distad coinciding ? proximad

P5 apex d vs AS posterior margin proximad coinciding proximad coinciding proximad

- spinules 2nd abdominal somite 2 absent present present ? absent

_ spinules 1st abdominal somite 3 absent absent absent present absent

comparison with the Tegastidae, Peltidiidae and Tisbidae suggests

that migration happened anteriorly and the condition in Clytemnestra

is apomorphic.

Species discrimination in Clytemnestra is most easily achieved

by comparing primarily cephalothorax shape, swimming leg spine

pattern, urosomal ornamentation and setation of the maxillae and

antennules (Table I). Conversely, identification of Goniopsyllus

species is strenuous and largely based on size, maxillipedal orna-

mentation and proportional lengths of caudal ramus setae. The

reported variability in body size and/or PS setation for both C.

scutellata and G. rostratus (e.g. Boxshall, 1979; Huys et al., 1996)

is based on erroneous identifications and observational errors.

Relationships

_ Prior to Claus’ (1891a) study the relationships of the Clytemnestridae

| were believed to lie with the planktonic poecilostomatoid families,

in particular the Sapphirinidae (Car, 1890). This concept was partly

_ based on the superficial similarity in dorsoventrally depressed body

shape, laterally displaced fifth legs and the failure to recognise the

geniculate antennules in the male (Car, 1890). More significantly,

this assignment was based also on the strongly reduced mouthparts

and the sexual dimorphism displayed by the maxillipeds, two

characters regarded as highly diagnostic for the Poecilostomatoida

(Huys & Boxshall, 1991).

Sexual dimorphism in the maxillipeds is uncommon in the

Harpacticoida. Huys (1988) reviewed the topic, showing that there

is clear dimorphism only in the Aegisthidae (as a result of male

atrophy), some Tisbidae (e.g. Boxshall, 1979) and deepwater

Huntemanniidae (Metahuntemannia, Talpina). Dahms & Schminke

(1993) demonstrated that in Tisbe bulbisetosa the male maxilliped is

involved in precopulatory mate guarding by holding the female’s

caudal setae IV and V prior to spermatophore transfer, the antennules

playing only an auxiliary role during this process. We speculate that

the modified male maxillipeds in clytemnestrids perform a similar

function, the elongate endopodal claw probably being involved in

holding the female’s caudal rami or swimming legs.

Boxshall & Huys (1998) pointed out that the antennulary chemo-

sensory system of C. rostratus (= G. clausi sp. nov.) is secondarily

enhanced in both sexes by transformation of three setae into

aesthetasc-like elements. The middle and distal of these elements

are fused basally to an aesthetasc. This study has revealed this

pattern to be diagnostic for all Clytemnestridae and can be consid-

ered an apomorphy for the family. Examination of copepodid stages

showed these transformed setae to be present from at least copepodid

III onwards. Modification of antennulary elements into putative

chemosensors is rare in harpacticoid copepods and has thus far only

been recorded in some deep-sea species. Gee & Huys (1991)

described a densely opaque, bulbous element on the distal antennulary

segment in both sexes of the paranannopid Leptotachidia iberica

Becker, 1974. The only report of a similar structure is that by Por

(1969) who figured a modified bulbiform element on the antennule

of Cerviniopsis obtusirostris Brotskaya, 1963 (Cerviniidae) which

he called the ‘Brodskaya organ’.

The complete lack of swimming leg sexual dimorphism impedes

an assessment of the relationships of the Clytemnestridae. The 1-

segmented P1 exopod is found in several interstitial Paramesochridae,

Leptastacidae and Laophontidae, yet it is diagnostic at the family

level only in the Rotundiclipeidae and Tegastidae. Lang (1948)

recognised a close relationship between the latter, the Peltididiidae

and the Clytemnestridae. He based this affinity solely on P1 mor-

phology, including the non-prehensile nature of the endopod and the

presence of maximum 5 elements on the distal exopod segment.

Within this group of tisbidimorph families he placed the Peltidiidae

as the sistergroup of the Clytemnestridae on account of the dorso-

ventrally flattened body and the reduction of the PS baseoendopod

in the female. The usefulness of Lang’s (1948) characters is limited

due to their homoplastic nature, however, there are at least two other

features which appear to substantiate a close relationship between

these three families. First, the aesthetasc pattern on the male antennule

(with an additional aesthetasc on ancestral segment XI) is displayed

by all three families. Secondly, the modification of the distal palmar

element on the maxillipedal basis into a pad-like sensory element

(Fig. 10B) is a unique synapomorphy (see Huys et al. (1996) for

examples in Peltidiidae and Tegastidae). A detailed phylogenetic

analysis of the Peltidiidae is nevertheless required before its

sistergroup relationship with the Clytemnestridae can be substanti-

ated. Indeed, an alternative evolutionary scenario could be that the

latter represent only a specialized terminal branch of the former.

Most species of the peltidiid genus Alteutha Baird are common

members of the coastal plankton, performing pronounced diurnal

vertical migrations in the water column. This may well be viewed,

either ecologically or evolutionary, as a transitionary step towards

the holoplanktonic lifestyle exhibited by the Clytemnestridae.

‘Taxonomic Impediment’ and Marine Plankton

The present revision has quadrupled the number of species in the

family solely by examination of the relatively limited material

deposited in the NHM. There is no doubt that this number would

have been significantly higher had the geographic coverage been

wider. Indicative of this is the discovery of three species of

Clytemnestra in a small sample from the Great Barrier Reef. Pre-

liminary examination of material from Brazilian waters (Rio Grande

do Sul) revealed a similar sympatry for both Clytemnestra and

Goniopsyllus. Although the discovery of several closely related

species in both genera is noteworthy, it is not unexpected nor

exceptional for a marine planktonic taxon. For example, recent

taxonomic studies have uncovered several important species com-

plexes in the Oncaeidae (Heron, 1977; Heron & Bradford-Grieve,

1995; Bottger-Schnack, 1999). Although this family is morphologi-

cally distinctive and arguably the most speciose in the marine

plankton, the continuing discovery of pseudo-sibling species and

frequent confusion about the validity of rank of its species and

morphs tarnish its literature, both taxonomic and ecological. Current

research on another planktonic poecilostomatoid genus, Pachos

Stebbing, resulted in the recognition of several new but previously

misidentified species (Huys & KrSinié, in prep.).

The taxonomy of pelagic harpacticoids is plagued by consider-

able conservatism and inadequate study of morphological features.

With the exception of the mesopelagic tisbid genera (Boxshall,

1979) all planktonic harpacticoids were known well before the turn

of the century (Kr@yer, 1846; Dana, 1847, 1849; Boeck, 1865;

Brady, 1883; Giesbrecht, 1891; T. Scott, 1894), yet, their morpho-

logical definition and supposedly cosmopolitan breadth of their

distribution have hitherto remained unchallenged. The genus

Microsetella Brady & Robertson currently encompasses only two

species, however, one can expect its number of species to increase

by an order of magnitude if the many undescribed sibling species are

considered (unpubl. data). Similarly, Euterpina acutifrons (Dana,

1847) is commonly regarded as a cosmopolitan species but compari-

son of distant ‘populations’ suggests that there is no factual

justification for this universally accepted view.

In Fleminger & Hulsemann’s (1977) scholarly study demonstrat-

ing the taxonomic divergence in three sympatrically occurring

sibling species of Calanus in the North Atlantic, one sentence

deserves wide currency: *. . ., the quality of knowledge about

circulating oceanic habitats and their entrained ecosystems rests

upon the reliability of three interrelated sets of information: system-

atics of the biota, routine identifications of species, and assessments

of their ranges, horizontally and vertically’. Unfortunately, routine

identifications in ecological investigations are generally not condu-

cive to the recognition of sibling species and all too often wide

geographical distributions have been uncritically accepted as the

natural consequence of potentially broad oceanic dispersal. The

latter perception is often coloured by underlying assumptions of the

lack of isolating physical barriers and global uniformity in the open

pelagic environment. Pseudo-sibling species can only be readily

distinguished once the appropriate characters are considered. Our

study demonstrated that for the last 110 years species discrimination

in the Clytemnestridae was based exclusively on generic characters,

the current recognition of cryptic species being only an artifact of

previous ignorance. Hence, there is considerable doubt involved in

collating records of the occurrence of these species from the litera-

ture to produce distribution maps. Though C. scutellata and G.

rostratus have universally been regarded as cosmopolitan, this

distributional concept is now no longer tenable and the compilation

of distribution records must start from scratch. It would be best to

consider earlier records primarily as evidence of the occurrence of

the respective genera, a useful attribute considering their virtual

absence at latitudes above 60° N and 45° S.

Although the geographic location of the collection and/or body

size can occasionally be used as indicators of species identity, these

R. HUYS AND S. CONROY-DALTON

approaches are limited in areas of sympatry where often more

sophisticated techniques are required. Like Clytemnestra in the

harpacticoids, Calanus is an unusual calanoid genus in that the

morphology of the female P5 does not discriminate all of the species

(Frost, 1971, 1974). Bucklin et al. (1995) showed however, that

despite their exceptional morphological similarity, species of Calanus

are quite distinct genetically. They obtained similar results for the

genus Metridia, confirming the distinctiveness of M. lucens (Boeck,

1865) and M. pacifica (Brodsky, 1948). Frost (1989) concluded,

based on morphological characters other than size, that there are

seven species within Pseudocalanus. For some, no absolute mor-

phological criterion could be found to distinguish females, however,

their validity was inferred from trends in several morphological

characters. Sévigny ef al. (1989) used patterns of allozyme variation

at the GPI (glucose phosphate isomerase) locus to show that Frost’s

(1989) sibling species were genetically isolated from each other.

Their results agreed with McLaren etal.’s (1989a—c) studies demon-

strating differences in genome size and life cycle characteristics

among Pseudocalanus species. Bucklin et al. (1998) showed by

DNA sequencing of two mitochondrial genes that the sibling species

P. moultoni and P. newmani can be reliably discriminated. Bucklin et

al.’s (1996) genetic analysis of DNA sequence variation separated

the widespread Nannocalanus minor into two genetically distinct

types that may represent the previously described N. m. forma major

and N. m. forma minor which differ primarily in size range and

geographic distribution. Finally, McKinnon ef al. (1992) demon-

strated the presence of three sympatric sibling species of Acartia

using allozyme electrophoresis.

Molecular analysis of marine planktonic copepods is likely to

continue to reveal taxonomically-significant genetic partitioning of

species populations, including cryptic species. The application of

molecular techniques should not however, be an end in itself.

Methods used to discriminate sibling species such as protein electro-

phoresis or discriminant function analysis profit significantly from

or even require a priori morphological recognition of groups or

morphotypes whose distinctiveness can be subsequently tested. In

fact, how can one demonstrate the accuracy and resolving power of

morphological analysis better than to refer to the thorough revisions

by Fleminger (1973) and Fleminger & Hulsemann (1974) who

presented most compelling evidence for sibling speciation in marine

calanoid copepods long before the deluge of molecular data. Failure

to recognize the numerous sibling species inevitably results in bad

science and has obvious implications for a large field like marine

plankton ecology, crippling our understanding of speciation and

resource partitioning in the ocean.

ACKNOWLEDGEMENTS. We thank A. Lindley and J.M. Gee (Plymouth

Marine Laboratory) for providing us with Goniopsyllus material from the

Gulf of Cadiz, and, M. Monti and W.J.A. Amaral (Fundagao Universidade

do Rio Grande) and Geoff Boxshall for putting Brazilian clytemnestrid

collections at our disposal. Prof. F. Krsini¢ is gratefully acknowledged for his

help in collecting material in the Adriatic. This research was partly supported

by the ALIS Programme (Project 041) which is jointly funded by the Croatian

Ministry of Science and Technology (MOST) and the British Council Croatia.

REFERENCES

Bjornberg, T.K.S. 1963. On the marine free-living copepods off Brazil. Boletim do

Instituto Oceanografico, Sao Paulo 13(1): 3-142.

, Campaner, F. & Jankilevich, S.S. 1981. Clasificacion de las especies presentes

en el Atlantico Sudoccidental. pp. 603-679. In: Boltovskoy, D. (ed.), Atlas del

zooplankton del Atlantico Sudoccidental y Métodos de Trabajo con el Zooplancton

GENERIC CONCEPTS IN CLYTEMNESTRIDAE

Marino. Instituto Nacional de Investigacién y Desarollo Pesquero, Mar del Plata,

Argentina.

Bodin, P. 1997. Catalogue of the new marine harpacticoid copepods. Studiedocumenten

van het K.B.1.N. 89: 1-304.

Boeck, A. 1865. Oversigt over de ved Norges Kyster jagttagne Copepoder henhorende

til Calanidernes, Cyclopidernes og Harpactidernes Familier. Forhandlinger i

Videnskabsselskabet i Kristiania 1864: 226-282.

Béttger-Schnack, R. 1999. Taxonomy of Oncaeidae (Copepoda, Poecilostomatoida)

from the Red Sea. I. 11 species of Triconia gen. nov. and a redescription of T. similis

(Sars) comb. nov. from Norwegian waters. Mitteilungen aus den Hamburgischen

Zoologischen Museum und Institut, in press.

Bowman, T.E. & Abele, L.G. 1982. Classification of the recent Crustacea. pp. 1-27.

In: Abele, L.G. (ed.), The Biology of Crustacea (editor-in-chief D.E. Bliss), 1

(Systematics, the Fossil Record, and Biogeography). Academic Press, New York &

London.

Boxshall, G.A. 1979. The planktonic copepods of the northeastern Atlantic Ocean:

Harpacticoida, Siphonostomatoida and Mormonilloida. Bulletin of the British Mu-

seum (Natural History), Zoology 35: 201-264.

Brady, G.S. 1878. A monograph of the free and semi-parasitic Copepoda of the British

Islands, 1. 148p. Ray Society, London.

—. 1883. Report on the Copepoda collected by H.M.S. Challenger during the years

1873-76. Report of the Scientific Results of the Voyage of H.M.S. Challenger 1873—

76, Zoology 8(23): 1-142.

Bruce, J.R. Colman, J.S. & Jones, N.S. 1963. Marine fauna of the Isle of Man.

L.M.B.C. Memoirs on Typical British Marine Plants and Animals 36: i-ix, 1-307.

Bucklin, A., Frost, B.W. & Kocher, D.T. 1995. Molecular systematics of six Calanus

and three Metridia species (Calanoida: Copepoda). Marine Biology 121: 655-664.

, LaJeunesse, T.C., Curry, E., Wallinga, J. & Garrison, K. 1996. Molecular

diversity of the copepod, Nannocalanus minor: genetic evidence of species and

populations structure in the North Atlantic Ocean. Journal of Marine Research 54:

285-310.

——,, Bentley, A.M. & Franzen, S.P. 1998. Distribution and relative abundance of

Pseudocalanus moultoni and P. newmani (Copepoda: Calanoida) on Georges Bank

using molecular identification of sibling species. Marine Biology 132: 97-106.

Campaner, A.F. 1985. Occurrence and distribution of copepods (Crustacea) in the

epipelagial. Boletim do Instituto Oceanogrdfico, Sado Paulo 33(1): 5-27

Campos Hernandez, A. & Suarez Morales, E. 1994. Copépodos peldgicos del Golfo

de México y Mar Caribe. I. Biologia y Systematica. 353p. Centro de Investigaciones

de Quintana Roo, Quintana Roo.

Car, L. 1890. Ein neues Copepoden-Genus (Sapphir) aus Triest. Archiv fiir

Naturgeschichte 56(1): 263-271.

——. 189 1a. Die Aufrechthaltung des Genus ‘Sapphir’ . Zoologischer Anzeiger 14(357):

72-73.

- 1891b. Erwiederung an Herrn Prof. C. Claus auf seine Arbeit *Goniopelte

gracilis’. Zoologischer Anzeiger 14(370): 271-275.

Carvalho, J. de Paiva. 1944. Copépodes de Caioba e Baia de Guaratuba. Arquivos do

Museu Paranaense 4(3): 83-116.

. 1952. S6bre uma colegao de Copépodos, nao parasiticos, da Baia de Santos e suas

adjacéncias. Boletim do Instituto Oceanogrdfico, Sao Paulo 3(\—2): 131-188.

Chen, Q.-c., Zhang, S.-z. & Zhu, C.-s. 1974. On planktonic copepods of the Yellow

Sea and the East China Sea. II. Cyclopoida and Harpacticoida. Studia marina sinica

9: 27-100.

Cheng, C., Zhang, S.-z., Li, S. & Li, S.-c. 1965. Marine planktonic copepods from

China. 210 p. Shanghai Scientific and Technical Publishing House, Shanghai.

_Chihara, M. & Murano, M. 1997. An illustrated guide to marine plankton in Japan.

1574 p. Tokai University Press, Tokyo.

Citarella, G. 1986. Les Copépodes des eaux portuaires de Marseille. Syllogeus 58:

276-282.

Claus, C. 1863. Die freilebenden Copepoden mit besonderer Beriicksichtigung der

Fauna Deutschlands, der Nordsee und des Mittelmeeres. 230p. W. Engelmann,

Leipzig.

—. 189la. Ueber Goniopelte gracilis, eine neue Peltide. Arbeiten aus dem

Zoologischen Instituten der Universitat Wien und der Zoologischen Station in Triest

9(2): 151-162.

—. 1891b. Die Beziehungen von Goniopelte gracilis Cls. = Clytemnestra Hendorffi

Poppe zu Goniopsyllus rostratus Brady = Sapphir rostratus L. Car, sowie deren

Stellung im System. Zoologischer Anzeiger 14(378): 424-432.

Conroy-Dalton, S. & Huys, R. 1999. A new genus of Aegisthidae (Copepoda,

Harpacticoida) from hydrothermal vents on the Galapagos Rift. Journal of Crusta-

cean Biology 19(2): 408-431.

Dahl, F. 1890. Berichtigung. Zoologischer Anzeiger 13(349): 633-634.

Dahms, H.-U. & Schminke, H.K. 1993. Mate guarding in Tisbe bulbisetosa (Cope-

poda, Harpacticoida). Crustaceana 65(1): 8-12.

Dana, J.D. 1847. Conspectus Crustaceorum, in orbis terrarum circumnavigatione, C.

Wilkes e classe Reipublice Foederate duce, collectorum auctore J.D. Dana. Proceed-

ings of the American Academy of Arts and Sciences 1: 149-155.

. 1849. Conspectus Crustaceorum que in Orbis Terrarum circumnavigatione,

Carolo Wilkes e Classe Reipublice Foederate Duce, lexit et descripsit Jacobus D.

Dana. Pars II. Proceedings of the American Academy of Arts and Sciences 2: 9-61.

. 1854. Crustacea. Part II. United States Exploring Expedition. During the years

1838, 1839, 1840, 1841, 1842, under the command of Charles Wilkes, U.S.N. 14:

691-1618.

De Decker, A.H.B. 1984. Near-surface copepod distribution in the south-western

Indian and South-eastern Atlantic Ocean. Annals of the South African Museum

93(5): 303-370.

Fagetti, E.G. 1962. Catalogo de los Copépodos plancténicos chilenos. Gayana,

Zoologia 4: 3-59.

Farran, G.P. 1936. Copepoda. Scientific Reports. Great Barrier Reef Expedition

1928-29 5(3): 73-142.

Fleminger, A. 1973. Pattern, number, variability, and taxonomic significance of

integumental organs (sensilla and glandular pores) in the genus Eucalanus (Cope-

poda, Calanoida). Fishery Bulletin NOAA 71(4): 956-1010.

— & Hulsemann, K. 1974. Systematics and distribution of the four sibling species

comprising the genus Pontellina Dana (Copepoda, Calanoida). Fishery Bulletin

NOAA 72(1): 63-120.

& . 1977. Geographical range and taxonomic divergence in North Atlantic

Calanus (C. helgolandicus, C. finmarchicus and C. glacialis). Marine Biology 40(3):

233-248.

Frost, B.W. 1971. Taxonomic status of Calanus finmarchicus and C. glacialis

(Copepoda), with special reference to adult males. Journal of the Fishery Research

Board of Canada 28(1): 23-30.

. 1974. Calanus marshallae, a new species of calanoid copepod closely allied to

the sibling species C. finmarchicus and C. glacialis. Marine Biology 26(1): 77-99.

. 1989. A taxonomy of the marine calanoid copepod genus Pseudocalanus.

Canadian Journal of Zoology 67(3): 525-551.

Gaudy, R. 1963. Campagne du navire océanographique ‘Calypso’ dans les eaux

cdtiéres du Brésil (Janvier-Février, 1962). Copépodes pélagiques. Recueil des Travaux

de la Station Marine d’Endoume 45 (Bulletin 30): 15-42.

Gee, J.M. & Huys, R. 1991. A review of Paranannopidae (Copepoda: Harpacticoida)

with claviform aesthetascs on oral appendages. Journal of natural History 25: 1135-

1169.

Giesbrecht, W. 189 1a. Ueber secundire Sexualcharactere bei Copepoden. Zoologischer

Anzeiger 14(371): 308-312.

. 1891b. Elenco dei Copepodi pelagici raccolti dal Tenente di vascello Gaetano

Chierchia durante il viaggio della R. Corvetta *Vettor Pisani’ negli anni 1882-1885

e dal Tenente di vascello Francesco Orsini nel Mar Rosso, nel 1884. Arti della

Accademia nazionale dei Lincei. Rendiconti, Classe di Scienze fisiche, matematiche

e naturali (4)7(sem. 1): 474-481.

. 1892. Systematik und Faunistik der pelagischen Copepoden des Golfes von

Neapel und der angrenzenden Meeres-Abschnitte. Fauna und Flora des Golfes von

Neapel 19: 1-831.

Gurney, R. 1927. Report on the Crustacea: Copepoda and Cladocera of the plankton.

Zoological results of the Cambridge expedition to the Suez Canal, 1924. Trans-

actions of the Zoological Society of London 22: 139-172.

Heron, G.A. 1977. Twenty-six species of Oncaeidae (Copepoda: Cyclopoida) from the

southwest Pacific-Antarctic area. In: Biology of the Antarctic Seas, 6. Antarctic

Research Series Washington 26: 37-96.

& Bradford-Grieve, J.M. 1995. The Marine Fauna of New Zealand: Pelagic

Copepoda: Poecilostomatoida: Oncaeidae. New Zealand Oceanographic Institute

Memoirs 104: 1-57.

Hicks, G.R.F. 1988. Evolutionary implications of swimming behaviour in meiobenthic

copepods. Hydrobiologia 167-168: 497-504.

Hirota, Y. 1995. The Kuroshio. Part III. Zooplankton. Oceanography and Marine

Biology: an Annual Review 33: 151-220.

Hure, J. & Krsinic, F. 1998. Fauna Croatica X/I. Planktonic copepods of the Adriatic

Sea. Spatial and temporal distribution. Natura Croatica 7 (Suppl. 2): 1-135.

Huys, R. 1988. Sexual dimorphism in aegisthid cephalosomic appendages (Copepoda,

Harpacticoida): a reappraisal. Bijdragen tot de Dierkunde 58: 114-136.

— & Bottger-Schnack, R. 1994. Taxonomy, biology and phylogeny of Miraciidae

(Copepoda: Harpacticoida). Sarsia 79: 207-283.

— & Boxshall, G.A. 1991. Copepod Evolution. 486p. Ray Society, London, No.

159.

——,, Gee, J.M., Moore, C.G. & Hamond, R. 1996. Synopses of the British Fauna

(New Series): Marine and Brackish Water Harpacticoid Copepods Part 1. viii +

352p. Field Studies Council, Shrewsbury, United Kingdom.

Ingélfsson, A. & Olafsson, E. 1997. Vital role of drift algae in the life history of the

pelagic harpacticoid Parathalestris croni in the northern North Atlantic. Journal of

Plankton Research 19: 15-27.

Kasturirangan, L.R. 1963. A key for the identification of the more common plank-

tonic Copepoda of Indian coastal waters. Publications of the Council of Scientific

and Industrial Research, New Delhi 2:\—87.

Kazmi, Q.B. & Muniza, F. 1994. Notes on two harpacticoid copepods (Crustacea)

gathered from plankton collection during Naseer cruise I, January 1992 in the

Arabian Sea. Proceedings of the Pakistan Congress of Zoology 14: 151-156.

Klie, W. 1943. Copepoda-I. Sub-order: Harpacticoida. Fiches d’Identification du

Zooplancton 4: \-4.

Krishnaswamy, S. 1953. Pelagic Copepoda from Madras coast. Part II. Harpacticoida.

Journal of the zoological Society of India 5(1): 64-75.

. 1957. Studies on the Copepoda of Madras. 168 p. Thesis, University of Madras,

Madras.

Kreyer, H. 1846. Crustacés. In: Voyage de la Commission scientifique du Nord en

Scandinavie, en Laponie, au Spitzberg et aux Feroé, pendant les années 1838, 1839 et

1840, sur la corvette Le Recherche commandeée par M. Fabure, lieutenant de Vaisseau,

publiés par ordre du Roi sous la direction de M. Paul Gaimard. Atlas, pls. 41-43.

Lang, K. 1944. Monographie der Harpacticiden (Vorldufige Mitteilung). 39p. Almqvist

& Wiksells Boktryckeri Ab, Uppsala.

—. 1948. Monographie der Harpacticiden, volume I. pp. 1-896, volume II. pp. 897—

1683. Hakan Ohlssons Boktryckeri, Lund, Sweden.

Lee, W. & Huys, R. (in press). New Aegisthidae (Copepoda: Harpacticoida) from

western Pacific cold seeps and hydrothermal vents. Zoological Journal of the

Linnean Society, in press.

Legaré, J.E.H. 1961. Estudios preliminares del zooplancton en la region de Cariaco.

Boletin del Instituto Oceanografico de la Universidad de Oriente, Cumana 1(1): 3-30.

. 1964. The pelagic Copepoda of eastern Venezuela, 1. The Cariaco trench. Boletin

del Instituto Oceanografico de la Universidad de Oriente, Cumana 3(1-2): 15-81

Lubbock, J. 1856. On some Entomostraca collected by Dr. Sutherland, in the Atlantic

Ocean. Transactions of the Entomological Society of London (2)4: 8-39.

. 1860. On some oceanic Entomostraca collected by Captain Toynbee. Trans-

actions of the Linnean Society of London 23: 173-193.

McKinnon, A.D., Kimmerer, W.J. & Benzie, J.A.H. 1992. Sympatric sibling species

within the genus Acartia (Copepoda: Calanoida): a case study from Westernport and

Port Phillip Bays, Australia. Journal of Crustacean Biology 12: 239-259.

McLaren, I.A., Laberge, E., Corkett, C.J. & Sévigny, J.-M. 1989a. Life cycles of

four species of Pseudocalanus in Nova Scotia. Canadian Journal of Zoology 67(3):

552-558.

, Sévigny, J.-M. & Corkett, C.J. 1989b. Temperature-dependent development in

Pseudocalanus species. Canadian Journal of Zoology 67(3): 559-564.

, Sévigny, J.-M. & Frost, B.W. 1989c. Evolutionary and ecological significance

of genome sizes in the copepod genus Pseudocalanus. Canadian Journal of Zoology

67(3): 565-569.

Marques, E. 1957. Copépodes da Guiné portuguesa (coligidos pela Missao

geo-hydrografica da Guiné). Anais da Junta de Investigagées do Ultramar 10(4)(1):

1-25.

——. 1958. Copépodes dos mares de Angola. II. Ciclopoida e Harpacticoida. Trabalhos

da Missao de Biologia maritima. Anais da Junta de Investigacées do Ultramar 12(2):

1-20.

. 1973. Copépodes marinhos das aguas de S. Tomé e do Principe. Livro de

homenagem ao Prof. F-F. Viegas da Costa, 70.° aniversdrio — 27 de Abril de 1968:

231-260.

Monard, A. 1927. Synopsis universalis generum Harpacticoidarum. Zoologische

Jahrbiicher, Abteilung fiir Systematik 54(1—2): 139-176.

. 1928. Les Harpacticoides marins de Banyuls. Archives de Zoologie expérimentale

et générale 67: 259-443.

Monti, M. 1980. Zooplancton do estuario da Lagoa dos Patos. I. Estrutura e variogdes

temporais e espaciais da comunidade. Atlantica 4; 53-72.

& Cordeiro T.A.. 1988. Zooplancton del complejo estuarial de la Bahia de

Paranagua. 1. Composicion, dinamica de las especies, ritmos reproductivos y accion

de los factores ambientales sobre la comunidad. Neritica 3(1): 61-83.

& Gloeden, I.M. 1986. Atlas dos Cladocera e Copepoda (Crustacea) do estuario

da Lagoa dos Patos (Rio Grande; Brasil). Neritica 1(2): 1-134.

Morales (R.), A. & Vargas (Z.), J.A. 1995. Especies comunes de copépodos (Crusta-

cea: Copepoda) pelagicos del Golfo de Nicoya, Costa Rica. Revista de Biologia

Tropical 43: 207-218.

Mori, T. 1929. Chosen-kaikyo fukin yori sai skuseski Fuyu-To Kyakurni ni touhite,

oyobi 2 skinsku no kisai. An annotated list of the pelagic Copepoda from the S.W.

part of the Japan-Sea, with descriptions of two new species. Zoological Magazine,

Tokyo 41(486-487): 161-177, 199-212.

R. HUYS AND S. CONROY-DALTON

——. 1937. The pelagic Copepoda from the neighbouring waters of Japan. 15Op.

Tokyo (Reprinted 1964; 2nd edition edited by S. Shirai).

Ohtsuka, S., Kubo, N., Okada, M. & Gushina, K. 1993. Attachment and feeding of

pelagic copepods on larvacean houses. Journal of oceanography 49: 115-120.

Owre, H.B. & Foyo, M. 1967. Copepods of the Florida Current. Fauna Caribaea 1.

Crustacea, Part 1, Copepoda: 1-137.

Pesta, O. 1909. Zoologische Ergebnisse XV. Copepoden (I. Artenliste 1890). In:

Berichte der Kommission fiir Erforschung des dstlichen Mittelmeeres. Denkschriften

der Akademie der Wissenschaften. Wien, mathematisch-naturwissenschaftliche

Klasse 84: 19-31.

Poppe, S.A. 1891. Beitrag zur Kenntnis der Gattung Clytemnestra, Dana. Abhandlungen

herausgegeben vom Naturwissenschaftlichen Verein zu Bremen 12: 131-142.

Por, F.D. 1969. Deep-sea Cervinidae (Copepoda: Harpacticoida) from the western

Indian Ocean, collected with R/V Anton Bruun in 1964. Smithsonian Contributions

to Zoology 29: 1-60.

Ramirez, F.C. 1966. Copépodos ciclopoidos y harpacticoidos del plancton de Mar del

Plata. Physis 26(72): 285-292.

—. 1970. Copépodos plancténicos del sector bonaerense del Atlantico surocciden-

tal. Datos y resultados de las campanas Pesqueria. Contribuciénes del Instituto de

Biologia marina, Mar del Plata 98: \|-116.

Razouls, C. 1996. Diversité et répartition géographique chez les Copépods pélagiques.

2. Platycopioida, Misophrioida, Mormonilloida, Cyclopoida, Poecilostomatoida,

Siphonostomatoida, Harpacticoida, Monstrilloida. Annales de l’ Institut

océanographique, Paris 72(1): 1-149.

& Durand, J. 1991. Inventaire des Copépodes planktoniques Méditerranéens.

Vie et Milieu 41(1): 73-77.

Reid, J.W. 1998. Maxillopoda — Copepoda. Harpacticoida. pp. 75-127. In: Young, P.S.

(ed.), Catalogue of Crustacea of Brazil. Rio de Janeiro, Museu Nacional.

Santos, J.J. 1973. Estudo preliminar, principalmente do plancton, das aguas da bafa de

Todos os Santos. Boletim da Faculdade de Filosofia e Ciéncias, Universidade de

Sao Paulo, Zoologia e Biologia Marinha, N.S. 30: 419-447.

Sars, G.O. 1921. Copepoda Supplement. Parts [IX & X. Harpacticoida (concluded),

Cyclopoida. An Account of the Crustacea of Norway, with short descriptions and

figures of all the species 7: title-page, preface, 93-121, pls. 65-76.

Scott, A. 1909. The Copepoda of the Siboga Expedition. Part I. Free-swimming, littoral

and semi-parasitic Copepoda. Siboga Expedition Monographs 29a: \-323.

Scott, T. 1894. Report on Entomostraca from the Gulf of Guinea, collected by John

Rattray, B.Sc. Transactions of the Linnean Society of London, Zoology (2)6: 1-161.

Sévigny, J.-M., McLaren, I.A. & Frost, B.W. 1989. Discrimination among and

variation within species of Pseudocalanus based on the GPI locus. Marine Biology

102(3): 321-327.

Sewell, R.B.S. 1940. Copepoda, Harpacticoida. Scientific Reports of the John Murray

Expedition 7(2): 117-382.

Suarez Morales, E. & Gasca, R. 1998. Updated checklist of the free-living marine

Copepoda (Crustacea) of Mexico. Anales del Instituto de Biologia. Universidad

Nacional de México, Serie Zoologia 69(1): 101-119.

Thompson, I.C. 1888. Report on the Copepoda collected in Maltese seas by David

Bruce, M.B., during 1886—7-8. Proceedings of the Liverpool Biological Society 2:

137-151.

Vega-Perez, L.A. 1993. Estudo do zooplancton da regiao de Ubatuba, Estado de Sao

Paulo. Publicagao Especial do Instituto Oceanografico 10: 65-84.

Vilela, M.H. 1965. Copépodes da Ria de Faro-Olhao. Notas e estudos do Instituto de

Biologia Maritima, Lisboa 31: 1-38.

. 1968. Copépodes da campanha do N.R.P. «Faial», 1958-1959. Notas e estudos

do Instituto de Biologia Maritima, Lisboa 35: 1-55.

Wells, J.B.J. 1970. Copepoda. I. Sub-order Harpacticoida. Fiches d’Identification du

Zooplancton 133: 1-7.

. 1976. Keys to aid in the identification of marine harpacticoid copepods. 215p.

Department of Zoology, University of Aberdeen, Aberdeen.

Wilson, C.B. 1932. The copepods of the Woods Hole region, Massachusetts.

Smithsonian Institution United States National Museum Bulletin 158: 1-635.

Bull. nat. Hist. Mus. Lond. (Zool.) 66(1):49—107 Issued 29 June 2000

Basal resolution of laophontid phylogeny and

the paraphyly of Esola Edwards

RONY HUYS AND WONCHOEL LEE

Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD

CONTENTS

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FRE LET OMG ES pecaccs cra seee seater ts cue seecoenea cara estasce vo cats nag cuasve For casatvs hava euaeceeudede Seetscaeeoe actee vera ccasactrv evan ser nabetsns fukee<oeVipnesdediewseeceSes 105

SYNOPSIS. The phylogeny of the Laophontidae, currently the second most speciose family of harpacticoid copepods in the

marine environment, is poorly understood. Despite its well established monophyletic status interrelationships within the family

have not been re-assessed since Lang’s (1948) deceptive phylogenetic hypothesis based on 19 genera (6 being of non-laophontid

affinity). Quadrupling of the number of recognized genera in the last 50 years and the persistent failure to recognize the

paraphyletic or polyphyletic nature of many of them have severely compromised objective analysis of relationships.

Parsimony analysis employing all informative morphological characters supports a basal dichotomy dividing the family in two

clades which are attributed subfamilial status. The Laophontinae, containing 95% of the species, differs from the Esolinae sub

fam. nov. in 2 PS morphology, the loss of the outer spine on the distal endopod segment of P2 and additional losses of armature

elements on the maxillipedal syncoxa and P1 endopod which were primitively retained in the Esolinae. Based on P3 endopod

50 R. HUYS AND W. LEE j

sexual dimorphism Onychocamptus Daday and the Laophonte cornuta-group invariably form a clade in opposition to all other

Laophontinae, implying polyphyly of the type genus.

The Esolinae is a relict group, cosmopolitan in distribution and displaying a complex ecological radiation. Analysis at species

level identified Archilaophonte Willen as the basal node and Mourephonte Jakobi as the terminal branch, and provided strong

support for the paraphyly of Esola Edwards. Relationships within the Esolinae are largely determined by patterns of transformed

integumental pores, sexual dimorphism of P2—P3 and caudal rami, segmentation of 2 antennule and P1 exopod, and 2 P5

armature.

The genus Esola is redefined to include a crown-group of 8 species, the distribution of which primarily coincides with the

circumglobal Tethyan belt. The universally accepted cosmopolitan distribution of the type species E. Jongicauda Edwards is

rejected on morphological grounds, resulting in the resurrection of E. bulbifera (Norman), the upgrading of E. longicauda

galapagoensis Mielke and the recognition of four species previously confounded with the type (E. vervoorti sp. nov., E. lobata

sp. nov., E. canalis sp. nov.) or based on new collections (E. profunda sp. nov.). Laophonte rhodiaca Brian is regarded as a likely

synonym of E. bulligera.

Both E. hirsuta (Thompson & A. Scott) and E. bulligera (Farran) are allocated to monotypic genera, Applanola gen. nov. and

Corbulaseta gen. noy., respectively. The mediterranean E. rosei (Monard) is considered a junior subjective synonym of the

northwestern European C. bulligera. E. spelaea (Chappuis), representing an isolated freshwater incursion in Apulian caves, is

transferred to Troglophonte gen. noy. and various ambiguities contained in its original description are reviewed. Bathyesola

compacta gen. et sp. nov. was discovered at 2765 m depth on the North Fiji Ridge, representing the deepest record for the family

thus far. E. typhlops (Sars) forms an exclusively Atlantic boreo-arctic clade with E. longiremis (T. Scott) and Esola sp. sensu

Chislenko (1967). A fourth species, A. hamondi from Norfolk, is added to this group which is accorded generic rank (Archesola

gen. nov.) on the basis of neotenic development of the male P3 endopod.

A generic key to the Esolinae and a review of their ecological radiation are presented.

INTRODUCTION

Laophontids comprise one of the six extant families of the

Laophontoidea (Huys & Lee, 1999). They represent by far the most

speciose group in this superfamily, currently accommodating 269

valid species and subspecies in 57 genera (Lee & Huys, 1999).

Laophontidae are essentially marine, free-living, benthic andrestricted

to phytal or shallow subtidal and intertidal habitats. Their success in

the deep sea is modest and only very few lineages have radiated into

freshwater or have entered into associations with invertebrate hosts.

The current rate of new species descriptions indicates that only a

moderate fraction of their true diversity is known.

Lang’s (1948) phylogenetic scheme of the Laophontidae included

only 19 genera, six of which being placed in other, existing or new,

families since (Hicks, 1988a; Huys, 1990a,b; Huys & Lee, 1999;

Huys & Willems, 1989). Although this re-allocation has signifi-

cantly refined the taxonomic concept of the family and hence its

monophyletic status is no longer a matter of dispute (Huys & Lee,

1999), the relationships between genera are usually not well under-

stood. The justification for creating new genera has traditionally

been based on a purely comparative approach, usually by consider-

ing a particular combination of characters as unique, rather than on

phylogenetic grounds. Some authors (e.g. Noodt, 1958) attempted

to unravel the relationships within particular lineages but their kind

of analysis was not cladistic and considered only a limited number of

characters. Others considered a thorough revision of the type genus

Laophonte Philippi as a conditio sine qua non for a phylogenetic

analysis incorporating all genera (Hicks, 19885; Willen, 1996).

The recent discovery of the primitive genus Archilaophonte in the

Antarctic Weddell Sea (Willen, 1995) has shed some light on the

early evolution of the family. Willen (1995) proposed an evolution-

ary scenario placing Archilaophonte and Esola as sistertaxa at the

base of the laophontid tree. Her analysis did not include the genus

Mourephonte Jakobi, left the potential paraphyly of Esola unchal-

lenged and was based on few characters. In this paper we have first

concentrated on the relationships within the genus Esola and its

affinity to Mourephonte and Archilaophonte. In order to resolve the

basal dichotomy in laophontid evolution we found it necessary to

run the analysis at the species level. Re-examination of the majority

of these species revealed important new taxonomic information

which reinforces the early split of two major lineages in the

Laophontidae. In this paper we propose a new hypothesis of basal

evolutionary relationships in the Laophontidae which will hopefully

provide a solid baseline for future studies addressing the phylogeny

of the more advanced crown-group taxa.

MATERIAL AND METHODS

Specimens were dissected in lactic acid and the dissected parts were

mounted on slides in lactophenol mounting medium. Preparations

were sealed with Glyceel or transparent nail varnish. All drawings

have been prepared using a camera lucida on a Zeiss Axioskop,

Leitz Dialux or Leitz DMR microscope equipped with differential

interference contrast.

Esola bulbifera, Applanola hirsuta and Archesola typhlops were

examined with a Hitachi S-800 or Philips XL30 scanning electron

microscope. Specimens were prepared by dehydration through

graded acetone, critical point dried, mounted on stubs and sputter-

coated with gold or palladium.

The descriptive terminology is adopted from Huys et al. (1996).

Abbreviations used in the text are: Al, antennule; A2, antenna; ae,

aesthetasc; exp, exopod; enp, endopod; P1—P6, first to sixth

thoracopod; exp(enp)-1(2, 3) to denote the proximal (middle, distal)

segment of a ramus. Type series are deposited in the collections of

The Natural History Museum, London (BMNH), the Muséum

National d’Histoire Naturelle, Paris (MNHNP) and the National

Museum of Natural History, Smithsonian Institution, Washington,

D.C. (NMNH). Scale bars in figures are indicated in um.

GENERIC DIAGNOSES AND SPECIES

DESCRIPTIONS

Family LAOPHONTIDAE T. Scott, 1905

Genus Esola Edwards, 1891

Edwards (1891) described Esola longicauda from an unknown,

shallow coastal locality in the Bahamas. Although the author found

the species embedded in mucus inside the body cavity of the

BASAL LAOPHONTID EVOLUTION

holothurian Actinopyga agassizii (Selenka) [as Miilleria Agassizii],

he considered it to be essentially free-living. He noted the distinctly

hirsute appearance and recognized a similarity between Esola and

Cleta Claus, placing the genus in the ‘Harpactiden’. Monard (1927)

placed the genus in the Laophontidae but erroneously stated in the

generic key that the antennule is 5-segmented. Later he professed

that Esola was really a ‘hirsute Laophonte’, differing from its

congeners only by the 1-segmented P1 exopod and its commensal

lifestyle with holothurians (Monard, 1935). Nicholls (19415) also

regarded the genus as a ‘derivative’ of Laophonte, however main-

tained the generic name pending a redescription of the type species.

The genus remained monotypic until Lang’s (1944, 1948) revi-

sion of the Laophontidae which added 8 Laophonte species to the

genus: L. hirsuta Thompson & A. Scott, L. longiremis T. Scott, L.

typhlops Sars, L. bulligera Farran, L. rosei Monard, L. spelaea

Chappuis, L. bulbifera Norman, and L. rhodiaca Brian. Lang (1948)

regarded the latter two species as synonyms of E. longicauda. He

maintained E. longicauda, E. bulligera and E. rosei as distinct

species for convenience rather than conviction, believing that future

examination might well show all three to be mere forms of the same

species. Lang (1944) divided the genus into two groups, the spelaea-

group, including only E. spelaea, and the longicauda-group,

accommodating all other species.

Nicholls (19416) had adopted a more artificial approach in his

revision of the Laophontidae, subdividing the genus Laophonte

Philippi into five subgenera on the basis of the endopodal setation of

the P3, and to a lesser extent also that of P2 and P4. He referred L.

rosei, L. bulligera, L. bulbifera, L. typhlops and L. longiremis to the

nominate subgenus Laophonte, more specifically to the typhlops-

group which also included L. elongata Boeck, L. thoracica Boeck

and L. barbata Lang. In the subgenus Mesolaophonte Nicholls he

placed L. spelaea which he believed to occupy an isolated position

due to the presence of 5 setae on the distal endopod segment of P4.

Finally, he regarded both L. hirsuta and L. rhodiaca as species

inquirendae, the former because it was inadequately described, the

latter because it was only known from the male. This system was

heavily criticized by Lang (1948: 1620-1621) in a postscript to his

monograph. A similar unnatural division of the genus Laophonte

had also been proposed by Sewell (1940), using P1 exopod segmen-

tation as the primary divisive character.

With the exception of Vervoort (1964) most authors have

uncritically accepted Lang’s (1948) decision to consider E.

longicauda as a variable and cosmopolitan species. Wells & Rao

(1987) regard the species as ‘highly distinctive pan-temperate/

tropical’ and express severe doubts about Mielke’s (1981) justifica-

tion for establishing E. longicauda galapagoensis. Mielke (1997)

hinted at the possibility of E. Jongicauda being a complex of several

closely related species and our examination appears to substantiate

his conjecture. In this revision we have restricted the genus Esola to

E. longicauda and to those species which have mistakenly been

synonymized with the type or were incorrectly described under that

name. The major diagnostic characters of these species are tabulated

in Table 1. Only E. bulbifera will be described in detail below; the

descriptions of the other species will be largely confined to the

differences with this species.

DIAGNOSIS. Laophontidae. Body cylindrical; posterolateral cor-

ners of @ genital double-somite and second abdominal somite

laterally and backwardly produced. Integument of cephalothorax

and body somites with dense pattern of spinules and setules. Ros-

trum large, partly delimited at base. Four pairs of integumental

cup-shaped pores present: anterodorsally on cephalothorax, near

ventrolateral margins of cephalic shield, laterally on genital (¢) or

genital double-somite ( @) and ventrally on caudal rami. Anal oper-

culum spinulose. Caudal rami modified in 2, often forming bulbous

expansions dorsally, ventrally and medially; rectangular and longer

than wide in d.

Sexual dimorphism in body shape, antennule, P3 endopod, P5,

P6, genital segmentation and caudal rami.

Antennules slender; 6- or incompletely 7-segmented in Q,

subchirocer and 7-segmented in d; segment 1 with 2—3 spinous

processes along posterior margin; with aesthetasc on segment 4 ( 9)

or 5 (3) and as part of apical acrothek on distal segment; segment 5

3 swollen, bearing modified spine on anterior outgrowth; proximal

aesthetasc fused to 2 setae. Antenna with 4 setae on exopod;

allobasis with abexopodal seta. Labrum with overlapping scales

distally and dense pattern of spinules proximally. Mandible with

short 1- or 2-segmented palp; endopod free or incorporated, repres-

ented by 2—3 setae; exopod usually absent, sometimes represented

by single seta; basis represented by 1-2 setae. Maxillule with

minute, defined exopod. Maxilla with 3 endites on syncoxa; endopod

represented by 4 setae. Maxilliped slender; syncoxa with 2 setae;

entire palmar margin with spinules; endopodal claw elongate.

Pl with 2-segmented exopod bearing 4-5 setae on exp-2 and

elongate endopod; enp-1 without inner seta, enp-2 with minute seta

and long, slender claw. P2—P4 with 3-segmented exopods and 2-

segmented endopods. P2 basis with very long outer spine. Outer

spine of P2—P4 enp-2 very long and setiform. P3 endopod 6 3-

segmented; enp-2 with inner seta and outer, dentate or smooth,

spinous apophysis. Armature formula as follows:

Exopod Endopod

Be 0.1.123 (O-1].221

P3 0.1.223 {0O-1].321 [d: [0-1].1.220]

P4 0.1.223 (0-1].221

PS 2 with separate rami; exopod elongate, with 6 setae/spines;

baseoendopod slightly developed, with 4 setae/spines. PS d without

endopodal lobe; exopod short, with 1 inner, 2 apical and 2 outer

elements.

P6 2 forming opercula closing off paired genital apertures; with

one seta and 2 small processes at outer corner. P6 ¢ asymmetrical;

membranous flaps with 2 setae.

TYPE SPECIES. Esola longicauda Edwards, 1891 [by monotypy].

OTHER SPECIES. Esola bulbifera (Norman, 1911); E. galapagoensis

Mielke, 1981 grad. nov.; E. profunda sp. nov.; E. canalis sp. nov.; E.

lobata sp. nov., E. vervoorti sp. nov.

SPECIES INQUIRENDAE. E. longicauda Edwards, 1891 sensu Noodt

(1955); E. longicauda Edwards, 1891 var. sensu Vervoort (1964); E.

longicauda Edwards, 1891 sensu Wells & Rao (1986); Esola spec.

sensu Mielke (1997).

Esola longicauda Edwards, 1891

TYPE LOCALITY. Unspecified shallow water locality in Bahamas.

TYPE MATERIAL. Edwards (1891) found both sexes but the material

is presumably lost.

Lang (1948) pointed out Edwards’ observational errors in his de-

scription of the P1 such as the presence of 4 setae on the inner margin

of the proximal endopod segment and the 1-segmented exopod.

Using the insertion site of the endopod as a reference point Lang

inferred that Edwards had incorporated the proximal exopod seg-

ment into the basis and that the outer basal seta is in reality exopodal.

Although Nicholls (19415) had also questioned the presence of 4

inner setae on the Pl endopod, assuming that they were only long

ornamentation elements, he nevertheless used this feature in his

generic key. The wide acceptance of Lang’s re-interpretation of the

P1 exopod, removing the one remaining obstacle to synonymy with

L. bulbifera and L. rhodiaca, made most authors overlook another

P1 character, i.e. the presence of only 4 elements on the distal

exopod segment. This pattern is also recorded in the subspecies E.

longicauda galapagoensis described by Mielke (1981) from two

islands in the Galapagos and in Esola spec., known from a single

female collected in North Sulawesi (Mielke, 1997), however, in all

other descriptions a consistent number of 5 setae is found.

There has been substantial debate over the supposed variability of

the P4 endopod in the various ‘populations’ of E. longicauda. Most

authors have dismissed the significance of the absence or presence

and relative size of the inner seta on the proximal segment (Table 1).

The inner seta is completely absent in Willey’s (1935) material of L.

bulbifera from Bermuda, Sewell’s (1940) specimens of L. bulbifera

from the Nicobar Islands and the Addu Atoll (Maldive Archi-

pelago), Vervoort’s (1964) specimens of E. longicauda from the

Ifaluk Atoll, E. longicauda galapagoensis from the Galapagos

(Mielke, 1981), Wells & Rao’s (1987) single female from Havelock

Island (South Andaman), and Mielke’s (1997) typical form of E.

longicauda from Bunaken Island (North Sulawesi). It is represented

by a vestigial element in Vervoort’s (1964) single male of E.

longicauda var. from Ifaluk Atoll and Mielke’s (1997) single female

of Esola spec. from North Sulawesi. Finally, it is very well devel-

oped in the male of L. rhodiaca described from the Aegean Sea

(Brian, 1928a) and Noodt’s (1955) ovigerous female of E. longicauda

from the Sea of Marmara. The very long seta recorded in this

position in L. bulbifera by Norman (1911) proved upon re-exam-

ination of the holotype to be based on an observational error (see

below). Hamond (1969) illustrated a scar which he interpreted as a

socket where a seta had probably broken off. It is our contention that

these setal differences do not reflect real variability but (in conjunc-

tion with other characters) demonstrate that several closely related

and frequently sympatric species have been described under the

name E. longicauda. Unfortunately the condition of the P4 in

Edwards’ (1891) material is somewhat dubious. On the basis of the

[0.1.223] setation pattern of the exopod his Taf. Il-Fig. 21 must

either be the P3 or the P4 and not the P2 as labelled (BpII!). Edwards

is less specific in the accompanying legend which states “Fuss eines

der drei folgenden Segmente’. The presence of only 2 inner setae on

the distal endopod segment may indicate that he had figured the P4

in which case the inner seta on the proximal segment is very well

developed. Edwards’ material differs also in the extremely long and

slender claw of the endopod (its length being 83% of that of enp-1)

and the elongate caudal rami which are slightly swollen in the

female, about 1.7 times as long as wide and have ventrally posi-

tioned pores. From the lateral habitus view they appear to be even

more slender and elongate in the male. These characters in conjunc-

tion with the presence of only 4 setae on Pl exp-2 and the well

developed inner seta of P4 enp-1 readily differentiate E. lJongicauda

from its congeners. The male is 550 um long (inferred from the

habitus drawing reproduced at x97 magnification).

Fiers’ (1986) single damaged female from Crooked Island (Baha-

mas) is likely to be the only reliable record of this species. Willey’s

(1935) record of Laophonte bulbifera from Harrington Sound (Ber-

muda) is zoogeographically closest but his claims that the caudal

rami are shortly barrel shaped, being only slightly longer than wide,

and that the P4 enp-1 lacks an inner seta cast doubt on his identifica-

tion. The conspecificity of his smaller female displaying a significant

disproportion in size (0.42 mm instead of 0.6 mm) and an atypical

R. HUYS AND W. LEE y

0.022 pattern on the P2 endopod is also highly questionable. Willey

(1935) regarded L. bulbifera to be close to L. depressa T. Scott but

gave no justification for this relationship. Alheit & Scheibel (1982)

also recorded E. longicauda from Harrington Sound but it is un-

known whether their identification was based on Willey’s or Edwards’

description. Finally, Rouch (1962) recorded the species from

Pernambuco State in Brazil but gave no evidence to substantiate his

identification.

Esola bulbifera (Norman, 1911)

Laophonte bulbifera Norman, 1911

? Laophonte rhodiaca Brian 1928a

Esola longicauda Edwards, 1891 sensu Hamond (1969)

Esola longicauda var. bulbifera Norman, 1911 sensu Holmes &

O’Connor (1990)

TYPE LOCALITY. Lamlash Bay in Firth of Clyde (Scotland).

MATERIAL EXAMINED.

(a) Holotype ¢ dissected on slide (BMNH #396.5); leg. J. Murray &

A.M. Norman, July 1888; dredging;

(b) 2 2Gand 1 6 collected from West Runton, Norfolk (England), at

extreme low water, around and under rocks; leg. R. Hamond, 20

August 1993; 1 2dissected on 13 slides (BMNH 1999.984), 1 2 and

1 6 preserved in alcohol (BMNH 1999.985—986);

Clifden, Co. Galway, Ireland; leg. B. O’Connor, July 1980, on

Serpula reef; det. JMM.C. Holmes; 1 ¢d dissected on 11 slides

(BMNH 1999.987), 3 2° preserved in alcohol (BMNH 1999.988—

990).

OTHER MATERIAL. National Museum of Ireland, Dublin: (a) sey-

eral specimens: Salt Lake, Clifden, Co. Galway; leg. B. O’Connor,

July 1980, from Serpula reef (in alcohol); (b) 1 2: Lough Hyne, Co.

Cork; leg. J.M.C. Holmes, 23 September 1987, light trap, 5 m (in

alcohol); (c) 1 6: Lough Hyne, Co. Cork; leg. J.M.C. Holmes, 08

Augustus 1992 (on slide).

DESCRIPTION.

FEMALE. Body length from anterior margin of rostrum to posterior

margin of caudal rami 681 um (n=5; range: 643-714 um). Maxi-

mum width (181 um) measured at posterior margin of cephalothorax.

Body (Fig. 1A—B) cylindrical, not dorsoventrally depressed,

covered with dense pattern of minute spinules dorsally and laterally.

Cephalothorax slightly wider than free somites, posterolateral angles

backwardly produced forming lobate extension (Fig. 1B); with

paired cup-shaped pores both anterodorsally and anteroventrally on

either side of rostrum (arrowed in Fig. 1B), anterodorsal set partly

closed off by fringe of setular extensions; with distinct transverse

spinule row dorsally about halfway down the cephalothorax length

(Fig. 1A). Posterior margin of cephalothorax and all body somites

with row of long setules dorsally and laterally. Posterior margin of

body somites with minute spinules laterally and ventrally; ventrola-

teral areas of cephalic shield and pleurotergites of pedigerous somites

with longer spinules. Pleurotergite of P5-bearing somite narrowest.

Genital double-somite wide and dorsoventrally flattened; original

segmentation marked by bilateral constriction and spinule row

arising from transverse surface ridge dorsally and laterally; anterior

(= genital) half with large cup-shaped pores laterally, each partly

closed off by fringe of setular extensions (Fig. 1C); posterior half

with backwardly directed lobate extensions bearing spinular tuft;

ventral surface without spinular ornamentation; genital field located

near anterior margin (Fig. 1C). Sixth legs forming well developed

opercula closing off paired genital apertures; each with outer naked

BASAL LAOPHONTID EVOLUTION

PANS Wy Dy

ALT IIAR NSS enn fe

HINA

NSA

aren

Adare

1 i .

yD! vf,

ASR

EZ,

ere

Bee ag -

fe od,

=p t

Lita

‘tp,

bite NN

Hit it

Fe ray

Yigg

1 \

eda Th

haiti

PUNT PS HAMERSAS DSSS

VMOU NSS

Way x

LAU AN HWS

CER AERRNT

ht URN

ARN)

Lr LAAN Wy \&

AL AY AN

Handy

HD \

/ Hi hy

stilt

Lien

TARR

FN My ee RA

Br rind wera! AN

UNNI i

i ein!

PRED

WN WEEN

TRAHAN

VATA

I AMENDS

iin

A TD

HAVO

HGS

SANNA

SAN

Melty yt ZONSSSONT

COA ut

CHM hy

PMN

PWN tly

WAN,

iY]

ny)

Fig.1 sola bulbifera (Norman, 1911) (9). A, Habitus, dorsal; B, habitus, lateral [anteroventral cup-shaped pore arrowed]; C, urosome (excluding P5-

bearing somite), lateral; D, anal somite and caudal rami, dorsal.

54 R. HUYS AND W. LEE

Wy Hi

Bhi Q

nN Ht

By iy, i ty i

May AHH it

TSAI

oe i

Fig. 2 Esola bulbifera (Norman, 1911) (9). A, Rostrum, dorsal; B, labrum, anterior; C, antennule, dorsal; D, antenna; E, mandible; F, maxillule; G,

maxilla; H, genital field; I, right caudal ramus, ventral.

BASAL LAOPHONTID EVOLUTION

Fig. 3 Esola bulbifera (Norman, 1911) (9). A, P1, anterior; B, P1, distal endopod segment, anterior; C, P2, anterior; D, P5, anterior: E, maxilliped; F,

paragnath.

R. HUYS AND W. LEE

fea)

Win We a RY

WIRES 5 \..

\ NGS

<q My

ae = A

4 Bran BN x

VASA:

<2 Ss =

* 4 a law =

CEG , \

—~———"

Vall)

lai

= i Temas HRS

Lue [dae

Fy a

= a

pggZ i ap Wo y 4.

L4Eé AZ MLE

\ <= f-7 =

J Cs <2

HIS

an)

= = Se

LLL we

IS a

—<—s

SSS

La.

Wy oe oF bo SS ————— :

ig IG A Ml ges OSs SS

= Z Zz =< LEE ————

rior; D, P3 endopod (¢), anterior.

rior; C, P4 enp-1 (9), ante

rior; B, P4 (9), ante

Fig. 4 sola bulbifera (Norman, 1911). A, P3 (9), ante

BASAL LAOPHONTID EVOLUTION Sy)

SSS

IPE

tan Se

Wy Li 0\\\

Way LL. MULAN

\a\5

HAN AAR

LOS)

41S

“4

4M, 4

4 NZ

Onis

BONES

Ve All

We: \

ATR

a Wes A

WN it Ny

WM YA hn ih wl

THAR AUT a ey,

WN Sidi Lhe pe ea ity

Asif! wale

yy RNS

4 i] \

Wiig PW

LUN.

EDN,

7] AYE IE

Win,

AIMS

TINK

DWN SQN

Fig.5 sola bulbifera (Norman, 1911) (3). A, Habitus, dorsal; B, antennule, dorsal (armature of segments 3-6 omitted); C, antennulary segments 3-4,

ventral; D, antennulary segment 5, ventral; E, antennulary segments 6-7, ventral; F, left caudal ramus, ventral; G, right PS, anterior; H, left P6, anterior.

seta and 2 small processes; with 4 medially directed, spinous

processes (Fig. 2H).

First postgenital somite with backwardly produced lateral angles,

bearing spinular tuft (Fig. 1C); without ventral ornamentation.

Penultimate and anal somites distinctly narrower; ventral posterior

border with spinules. Anal somite (Figs. 1D; 31A) with spinulose

anal operculum.

Caudal rami (Figs. 1D; 31A) widely separated; slightly longer

than widest portion; proximal half distinctly bulbous with major

swelling medially, dorsally and ventrally (Fig. 1C); ventral surface

with very large semi-circular concavity (Fig. 21) leading to small

tube-pore; dorsolateral surface with minute spinules; seta I small,

setae II-III well developed, naked and closely set; setae [V and V

pinnate and with fracture planes, seta V 2.5 times as long as seta IV;

setae VI-VII naked.

Rostrum (Figs 1A; 2A) large, rounded anteriorly; delimited at

base by transverse surface suture; with paired sensillae anteriorly

and median tube-pore dorsally.

Antennule (Fig. 2C) slender, incompletely 7-segmented, with 1

minute (obscured by large distal one) and 2 well developed spinous

processes on posterior margin of segment 1, no processes on long

segment 2. Segment 1 with short spinules posteriorly between

processes and large spinular patch around anterior margin. Armature

formula: 1-[1], 2-[4 + 4 pinnate], 3-[6], 4-[(2 + ae)], 5-[1], 6-[2], 7-

[6 + 1 pinnate + acrothek]. Aesthetasc on segment 4 fused basally to

2 setae. Acrothek consisting of aesthetasc and 2 naked setae; set on

apical pedestal. Boundary between segments 6 and 7 only expressed

posteriorly.

Antenna (Fig. 2D) with elongate exopod bearing 2 lateral and 2

apical pinnate elements, and a longitudinal row of fine spinules.

Allobasis with pinnate abexopodal seta and spinular patch opposite

exopod. Endopod with lateral armature consisting of | pinnate spine

and 2 setae; distal armature consisting of 2 unipinnate spines and 3

geniculate setae (outermost fused basally to small tube-seta).

Labrum (Fig. 2B) with spinules around distal margin; anterior

face with dense pattern of fine spinules and distal patch of overlap-

ping scales.

Mandible (Fig. 2E) with short gnathobase and small 1-segmented

palp probably representing fused basis and endopod; with 2 lateral

(basal) pinnate setae and 3 distal (endopodal) setae (1 pinnate, 2

bare).

Paragnaths highly ornate lobes as in Fig. 3F.

Maxillule (Fig. 2F) with well developed praecoxal arthrite bear-

ing 1 seta on anterior surface and 9 elements around distal margin.

Coxal endite with 1 spine and 1 seta, basal endite with 1 spine and 2

setae. Exopod a short segment with 2 distal setae; endopod incorpo-

rated into basis, represented by 2 setae.

Maxilla (Fig. 2G). Syncoxa with very long spinules around outer

margin and dense surface spinulation as figured; with 3 endites;

praecoxal endite small, with 1 plumose seta; middle endite drawn

out into pinnate claw, with 2 tube-setae; distal endite with 3 ele-

ments. Allobasis produced into strong curved claw; accessory

armature consisting of 1 spine and | seta; with spinular patch

proximal to endopodal setae. Endopod incorporated into allobasis,

represented by 2 bare and 2 pinnate setae.

Maxilliped (Fig.3E) slender, with elongate basis and endopodal

claw. Syncoxa with 2 plumose setae. Basis with spinular ornamen-

tation as figured; spinules present along entire palmar margin.

Endopod represented by very long, minutely pinnate claw bearing 1

accessory seta and tube-pore at base.

P| (Fig. 3A) with dense ornamentation on praecoxa, coxa and

basis. Basis with pinnate seta on anterior surface and along outer

margin. Exopod 2-segmented, small compared to endopod; exp-1

R. HUYS AND W. LEE }

not extending to distal margin of basal pedestal, with pinnate outer

spine; exp-2 with 3 pinnate outer setae and 2 geniculate setae

apically. Endopod slender; enp-1 about 2.5 times as long as basis,

with long setules along inner margin and fine spinules along outer

margin; enp-2 about 3 times as long as wide, with slender minutely

pinnate claw and small accessory seta (Fig. 3B).

P2—P4 (Figs 3C; 4A—B) with 3-segmented exopods and 2-seg-

mented endopods. P2 basis with long, bipinnate outer spine; P3—P4

bases with bare outer seta. P2—P3 enp-1 with multipinnate inner

seta; P4-enp-1 (Fig. 4C) with basally swollen, minute seta. Outer

spine of P2—P4 enp-2 very long and setiform. Tube-pore present

near distal outer corner of P3—P4 enp-2. Armature formula as

follows:

Exopod Endopod

P2 0.1.123 1.221

iP) 0.1.223 iL2y2Il [d: 1.1.220]

P4 0.1.223 1.221

P5 (Fig. 3D). Endopodal lobe small, extending just beyond

insertion sites of proximal outer setae of exopod; with 1 short and 1

long pinnate seta apically, and 2 long widely separated setae along

inner margin; tube-pores present near articulation with exopod,

between apical setae and proximal to innermost seta. Exopod elon-

gate, produced apically into tubular extension bearing 1 bare seta;

inner margin with 1, outer margin with 4 pinnate setae; inner seta

distinctly longer than apical one. Both baseoendopod and exopod

with elaborate ornamentation pattern as figured.

MALE. Body length from anterior margin of rostrum to posterior

margin of caudal rami 512 um (n=2, range 500-524 um). Maximum

width (168 tm) measured at posterior margin of cephalothorax.

Body (Fig. 5A) more compact and abbreviated than in 2; covered

with similar dense pattern of minute spinules. Pattern of cup-shaped

pores as in @ except for paired lateral pores present on genital

somite. Cephalothorax wider than free somites; body constricted at

level of genital somite. None of urosomites with backwardly pro-

duced posterolateral corners.

Genital somite with large cup-shaped pores laterally, each partly

closed off by fringe of setular extensions (Fig. 5H). Sixth legs

represented by well developed opercula, one articulating and clos-

ing off left or right genital aperture; each produced into cylindrical

process bearing | lateral and 1 apical seta.

Antennule (Fig. 5B—E) 7-segmented, subchirocer, with

geniculation between segments 5 and 6. Segment 1 with spinules/

setules around anterior margin and 2 spinous processes along poste-

rior margin. Segment 2 longest; segment 4 minute, represented by

incomplete sclerite. Segment 5 with large proximal process anteriorly,

bearing modified bifid spine (Fig. 5D); forming cylindrical process

bearing long aesthetasc. Segment 6 with 3 spinous processes along

anterior margin. Distal portion of segment 7 elongated, displacing

acrothek to position isolated from other armature. Armature for-

mula: 1-[1], 2-[4 + 5 pinnate], 3-[6 + 1 pinnate], 4-[2], 5-[7 + 2

pinnate + | bifid spine + (2 + ae)], 6-[1 + 3 processes], 7-[7 +

acrothek]. Apical acrothek consisting of aesthetasc and 2 bare setae.

P3 endopod (Fig. 4D) 3-segmented; enp-1 as in 9, enp-2 with

inner seta and short outer apophysis; enp-3 small, with tube-pore, 2

lateral and 2 apical setae.

P5 (Fig. 5G) medially fused, positioned ventrolaterally.

Baseoendopod without endopodal lobe; medial margin with 2

spinules and 2 tube-pores; outer basal seta arising from short spinulose

pedestal. Exopod free; with 1 apical and 1 inner and 3 outer pinnate

setae.

BASAL LAOPHONTID EVOLUTION

Caudal ramus (Fig. 5F) rectangular, without bulbiform expan-

sions; about 1.6 times as long as wide; with medioventral cup-shaped

concavity as in 9; ventral ornamentation more elaborate than in 9.

REMARKS. Lang (1948) synonymized L. bulbifera with E.

longicauda, alluding to the congruence in the female P5

baseoendopod between Gurney’s (1927) description of L. bulbifera

and Edwards’ (1891) original description of E. longicauda (i.e. with

4 setae; Norman (1911) figured only 3), and in the number of

processes on the first antennulary segment between the males of L.

rhodiaca (cf. Brian, 1928a) and E. longicauda (cf. Edwards, 1891)

and the female of L. bulbifera (cf. Norman, 1911). He also referred

to Willey’s (1935) discovery of L. bulbifera in Bermuda as addi-

tional zoogeographical evidence for this course of action. It is clear

however that (1) the morphological grounds for this synonymy only

prove generic identity and not conspecificity, (2) Willey’s (1935)

record is both unreliable and unconfirmed, and (3) Gurney’s (1927)

records from the Suez Canal in reality refer to another species E.

canalis sp. nov. (see below).

Our redescription diverges from Norman’s illustrations in only

two aspects: (1) the presence of 4 setae on the baseoendopod of the

2 PS, the innermost being overlooked by Norman as already sus-

pected by Lang (1948), and (2) the inner seta of P4 enp-1 which is

minute (checked against the holotype) instead of very well devel-

oped as figured by Norman. E. bulbifera can be differentiated from

its congeners on the basis of the following combination of characters:

antennule indistinctly 7-segmented, P1 enp-1 2.5 times as long as

basis, P1 enp-2 3 times as long as wide, P2-P4 enp-1 with inner seta

(that of P4 minute), outermost seta of 2P5 baseoendopod extending

to distal margin of exopod, caudal rami @ distinctly bulbous.

E. bulbifera is widely distributed around the British Isles with

reliable records from Ireland (Farran, 1913, 1915; Holmes &

O’Connor, 1990), the west coast of Scotland (Norman, 1911) and

Norfolk (Hamond, 1969). Moore’s (1973) record of E. longicauda

from St. Abb’s probably also refers to this species. It has not been

reported anywhere else in northwest Europe, however, its syn-

onymy with L. rhodiaca Brian, first suspected by Nicholls (1941b)

and later confirmed by Lang (1948), has considerably extended its

distribution, including the Mediterranean, Gulf of Suez and Western

Australia. Nicholls based his conviction on similarities in the

antennule, antennary exopod, P1 and P4 and the modified caudal

rami although he admitted that the latter were not bulbous in L.

rhodiaca. Brian’s (1928a) original description, based on a single

male specimen from Rhodes in the Aegean Sea (Brian, 1928a—b),

shows very few discrepancies with our material from Ireland and

Norfolk. The caudal rami are somewhat longer in the Mediterranean

specimen, the inner seta on P4 enp-! is more developed, the antennule

shows an additional segment distal to the geniculation and small

proportional length differences can be noted in the antennulary

segments and P4 endopod. Lang (1948) had already pointed out that

Brian had overlooked one of the outer spines on the P2 exopod. We

regard these differences insufficient to warrant the reinstatement of

L. rhodiaca and tentatively regard it as a junior subjective synonym

of E. bulbifera. Nicholls’ (1945) few illustrations of a male from

Port Denison in Western Australia which he attributed to L. rhodiaca

do not contradict Brian’s description. In the absence of information

on the swimming legs (except P3 endopod) and the female this

geographically widely separated record cannot be verified abso-

lutely.

Monard’s (1928) brief description of L. bulbifera from the Banyuls

area does not contain the level of detail to either confirm or deny his

identification. The setae on the P5 baseoendopod were probably not

drawn at their full length even though the outermost one appears to

be exceptionally short, his spine formula would infer a 123 pattern

on P4 exp-3 and the size of his female specimens (0.8 mm) falls

outside our recorded range. Monard (1937) recorded the species a

second time from Algers but the specimens were apparently dis-

tinctly smaller (0.64 mm).

The Croatian records of L. bulbifera from Rovinj and Split in the

northern Adriatic (Douwe, 1929; Klie, 1941) could not be con-

firmed. It is conceivable that VriSer’s (1984, 1986) records of E.

longicauda from the Gulf of Trieste and Petkovski’s (1955) record

from Montenegro refer to the same species.

Esola galapagoensis Mielke, 1981 grad. nov.

Esola longicauda galapagoensis Mielke, 1981

TYPE LOCALITY. Cabo Douglas, Fernandina (Galapagos).

Wells & Rao (1987) expressed reluctance about the subspecific rank

attributed to the Galapagos population of E. longicauda. Although

Mielke (1981) acknowledged the reported variability and

cosmopolitanism of the latter to some extent, he considered the

differences exhibited by his material sufficient to warrant the recog-

nition of a distinct subspecies. Mielke diagnosed E. longicauda

galapagoensis on the basis of the following characters: (1) P1 exp-

2 with 4 setae/spines, (2) Pl enp-2 with remarkably short claw, (3)

P4 enp-1 without inner seta, and (4) P5 baseoendopod @ with

strongly reduced outer apical seta. Additional diagnostic features

not mentioned by the author include (1) inner seta of P6 d extremely

reduced, (2) outer setae of PS exopod ¢ naked, (3) outer spine of P2—

P4 enp-2 remarkably short, and (4) caudal rami very elongate with

conspicuous medial swelling in °. Based on this suite of characters

we feel it justified to upgrade Mielke’s form to full species rank as E.

galapagoensis. The species has thus far been recorded from two

localities in the Galapagos archipelago (Mielke, 1981).

Esola canalis sp. nov.

Laophonte bulbifera Norman, 1911 sensu Gurney (1927)

TYPE LOCALITY. Suez Canal, Port Taufiq (Egypt).

TYPE MATERIAL. Holotype @ dissected on 10 slides (BMNH

1999.993): paratype 2 in alcohol (BMNH 1999.994); from material

originally registered as Laophonte bulbifera (BMNH 1928.4.2.116)

collected during the Cambridge Expedition to the Suez Canal in

1924; det. R. Gurney.

ETYMOLOGY. The species name refers to the type locality.

DESCRIPTION.

FEMALE. Body length from anterior margin of rostrum to posterior

margin of caudal rami 621 um (n=2; range: 585-658 um). Maxi-

mum width (137 pm) measured at posterior margin of cephalothorax.

Body as in E. bulbifera; cephalothorax with paired cup-shaped

pores both anterodorsally and anteroventrally on either side of

rostrum, and with distinct transverse spinule row dorsally about

halfway down the cephalothorax length.

Genital double-somite (Fig. 6A) wide and dorsoventrally flat-

tened; original segmentation marked by bilateral constriction and

spinule row arising from transverse surface ridge dorsally and

laterally; anterior (= genital) half with large cup-shaped pores

laterally; ventral surface without spinular ornamentation. First

postgenital somite with backwardly produced lateral angles, bearing

spinular tuft (Fig. 6A); without ventral ornamentation. Penultimate

and anal somites distinctly narrower; ventral posterior border with

spinules (Fig. 6C). Anal somite (Fig. 6B) with spinulose anal

operculum; spinules coarser than in E. bulbifera.

60 R. HUYS AND W. LEE

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HF ayy ‘wt SNS

ANTIK

Mop Tree

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Wily

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Fig. 6 Esola canalis sp. nov. (2). A, Urosome (excluding P5-bearing somite), lateral; B, anal somite and left caudal ramus, dorsal; C, anal somite and

caudal rami, ventral; D, antennule, dorsal; E, mandibular palp; F, antennary exopod.

LLL L—L£L_S

BASAL LAOPHONTID EVOLUTION

NINN

~ = =e SS =

: SSS Z =

Sex

Ww = ——

Vv i

tAZAl

XZ ‘ :

<— = GA

Ss Wil le

Ss a. Zi

LSS =

SSE oe ter EY

SSS SST

WS Sa

rior; C, P3 endopod, anterior; D, P4 endopod, anterior; E, P5, anterior

v. (9). A, P1, anterior; B, P2 endopod, ante

sp. no

lis

Fig.7 Esola cana

maxilliped.

R. HUYS AND W. LEE

Table 1. Diagnostic characters of Esola species [species A = Esola spec. sensu Mielke (1997)]. CR = caudal rami, SD = sexual dimorphism.

longicauda _ bulbifera galapagoensis canalis profunda vervoorti lobata species A

2 size (um) ? 643-714 330-460 585-658 500-529 510 490-530 470

3 size (um) 550 500-524 300-360 2 2 389-415 380-450 ?

cephalothorax dorsal spinule row ? present u present present absent ? ?

mandible — endopod ? fused, 3 setae fused,2setae free,3setae free,3setae free,3setae free, 3 setae free, 2 setae?

— exopod ? absent absent 1 seta absent absent absent absent

— basis ? 2 setae 2 setae 1 seta 2 setae 1 seta 2 setae 2 setae?

P1 exp-2 setal number 4 3) 4 5) 5) 3) 3) 4

P1 exp-2 outer apical seta SD eo = = ? ? + ~ y

ratio Pl enp-1 : enp-2 claw 0.83 0.50 0.37 0.50 0.55 0.55 0.55 y

P2—P3 enp-1| inner seta present present present present present absent present present

P3 enp-2 apophysis d ? smooth smooth ? ? dentate smooth ?

P4 enp-1 inner seta normal vestigial absent vestigial absent absent absent vestigial

P5 benp 2 outer apical seta plumose plumose naked plumose plumose plumose plumose uy

— length* short long vestigial very short short short ~short ?

CR — pore position ventral medioventral _ ventral mediodorsal ventral medial ventral 2

— medial swelling weak strong strong moderate moderate moderate slight moderate

— @length : distal width 3.0 28} 3.4 3.0 DS Del 3.8 ?

— 6 length : distal width 2 PhS) 3.4 i ? i157) 37) ?

*: very short = not extending to insertion level of middle outer exopodal seta; short = extending to about insertion level of middle outer exopodal seta; long = extending to

about apex of exopod [Note that in E. lobata sp. nov. the endopodal lobe is secondarily elongated so that its outer apical seta extends beyond the insertion level of the middle

outer exopodal seta despite being short].

Caudal rami (Fig. 6A—C) widely separated; gradually tapering

posteriorly and about as long as anal somite; proximal half expanded

with major swelling medially and dorsally, and to a lesser extent

ventrally (Fig. 6A); large cup-shaped pore located mediodorsally

(Fig. 6A) leading to small tube-pore. Armature as in E. bulbifera.

Antennule (Fig. 6D) slender, 6-segmented, with 1 large (proxi-

mal) and 2 small spinous processes along posterior margin of

segment 1. Segment 1 with long spinules around anterior margin.

Segments 2 and 3 equally long. Armature formula: 1-[1 pinnate], 2-

[7 + 1 pinnate], 3-[6], 4-[(2 + ae)], 5-[1], 6-[9 + acrothek]. Aesthetasc

on segment 4 fused basally to 2 setae. Acrothek consisting of

aesthetasc and 2 naked setae; set on apical pedestal.

Antennary exopod (Fig. 6F) elongate exopod bearing 2 lateral and

2 apical pinnate elements, and a longitudinal row of coarse spinules

proximally.

Labrum with ornamentation as in E. bulbifera.

Mandible (Fig. 6E) with small 2-segmented palp; proximal seg-

ment with 1 inner (basal) seta and 1 small outer seta representing

exopod; endopod a free segment with 3 setae.

Maxilliped (Fig.7F). Basis more slender than in E. bulbifera and

spinules along outer margin coarser.

P1 (Fig. 7A) similar to that of E. bulbifera but basis forming

shorter pedestal for endopod, and both exopod (but exp-1 extending

to distal margin of basal pedestal) and enp-2 somewhat shorter; exp-

2 with 3 outer setae and 2 geniculate setae apically.

P2—P4 (Fig. 7B—D). P2—P3 enp-1 with multipinnate inner seta; P4

enp-1 with vestigial inner seta. Outer spine of P2—P4 enp-2 shorter

than in E. bulbifera. Armature formula as follows:

Exopod Endopod

P2 0.1.123 122

P3 0.1.223 1.321 [d: probably 1.1.220]

P4 0.1.223 122A

P5 (Fig. 7E). Endopodal lobe small, not extending beyond inser-

tion sites of proximal outer setae of exopod; with 2 apical and 2

widely separated inner setae; outer apical seta very short. Exopod

more slender than in E. bulbifera; with 1 apical, | inner and 4 outer

setae; length of inner (ratio to exopod length 1.15 vs 1.5 in

E. bulbifera) and apical seta (ratio to exopod length 1.25 vs 2.2 in E.

bulbifera) distinctly shorter.

Unknown.

REMARKS. Gurney (1927) collected this species from the plankton

at Port Taufiq and Le Cap, and in sediment samples from El Ferdane.

He attributed his material to L. bulbifera but remarked on some

differences with Norman’s (1911) holotype, such as the discrepancy

in body size (0.68 mm instead of 0.80 mm), the P1 endopod which is

more slender in the Scottish specimen and the presence of an

additional seta (the innermost) on the PS baseoendopod. We have

found these differences to be of no value in discriminating both

species. Norman (1911) clearly overlooked the innermost seta (as

indicated by the gap along the medial margin in his figure of the

baseoendopod). Also, based on a larger sample of E. bulbifera we

found this species on average to be significantly smaller than Nor-

man’s observed size of 0.8 mm, approximating the mean length of E.

canalis (681 xm vs 621 um; see also Table I). There is no significant

difference in the P1 endopod of both species although the proximal

segment appears to be longer in E. canalis and the distal segment to be

longer in E. bulbifera. Gurney (1927) illustrated the P5, caudal rami

and the female habitus in lateral view. His illustration of the caudal

rami gives a slightly distorted view in that the rami appear to be much

longer than in reality. Por & Marcus (1972) recorded E. longicauda

from four localities in the Suez Canal; itis likely that these records and

Por’s (1967) previous record from the Gulf of Elat refer to E. canalis.

E. canalis is most closely related to E. bulbifera. Females of the

former can be differentiated by the conical caudal rami, the medi-

odorsal position of the cup-shaped pores on these rami, and the P5

endopodal lobe which is significantly shorter and has a much

smaller outer apical seta. Additional differences can be found in the

proportional lengths of the proximal antennulary segments, the

slenderness of the maxilliped and the size of particular setae on the

P2—P4 endopods and PS exopod. E. canalis is the only species of the

genus which has retained a vestige of the mandibular exopod.

MALE.

Esola lobata sp. nov.

Esola longicauda (Edwards, 1891) sensu Mielke (1997)

TYPE LOCALITY. Bunaken Island near Manado, North Sulawesi

(Indonesia); sublittoral sand between seagrass and corals.

BASAL LAOPHONTID EVOLUTION

ETYMOLOGY. The species name refers to the well developed

endopodal lobe of the P5 in both sexes.

P2—P4 setal formula:

Exopod Endopod

P2 0.1.123 1.221

P3 0.1.223 1.321 [d: 1.1.220]

P4 0.1.223 0.221

Mielke (1997) provided an excellent description of Sulawesi

females and males which he attributed to E. Jongicauda. His illustra-

tions show sufficient differences to warrant separate species status.

E. lobata is similar to E. profunda from the Mediterranean and both

E. vervoorti and E. galapagoensis from the Pacific in the loss of the

inner seta on P4 enp-1. The species can, however, be readily

distinguished by the long endopodal lobe in the @ PS, a well

developed bulbous extension on the baseoendopod of the d PS, the

elongate caudal rami which are relatively little modified in the

female, and the short P1 endopod. Discrepancies are also noted in

the female antennule, particularly in the relative lengths of the

proximal segments, and the size and precise position of the spinous

processes on segment |. The species is thus far known only from the

type locality.

Esola profunda sp. nov.

TYPELOCALITY. Ligurian Sea (Western Mediterranean; 42°39’ 12"

N, 08°39°30" E), northwest of the Bay of Calvi (Corsica); depth 760

TYPE MATERIAL. 2 22from type locality. The bottom sample was

taken on 10 June 1986 with a small, modified Reineck box corer

(170 cm?) by K. Soetaert. The median grain size of the sediment is

4 um and the silt-clay amount averages 78.5%. The CPE value is

about 0.59 g/cm? of which 12.6% is represented by chl a. Holotype

dissected on 11 slides (BMNH 1999.991), paratype @ preserved in

alcohol (BMNH 1999.992).

ETYMOLOGY. The species name is derived from the Latin profun-

dus (meaning deep) and refers to the bathyal distribution of this

species.

DESCRIPTION.

FEMALE. Body length from anterior margin of rostrum to posterior

margin of caudal rami 515 um (n=2; range: 500-529 um). Maxi-

mum width (129 um) measured at posterior margin of cephalothorax.

Body (Fig. 8A) as in E. bulbifera but constrictions between

pedigerous somites less defined; cephalothorax with paired cup-

shaped pores both anterodorsally and anteroventrally on either side

of rostrum, and with distinct transverse spinule row dorsally about

halfway down the cephalothorax length.

Urosomites with dense spinulation and irregular pattern of sur-

face ridges laterally and dorsally (Fig. 8B). Genital double-somite

(Fig. 8A—B) with large cup-shaped pores laterally in anterior half;

ventral surface without spinular ornamentation; posterolateral angles

slightly produced. First postgenital somite with backwardly pro-

duced lateral angles, bearing spinular tuft; without ventral

Ormamentation. Penultimate and anal somites distinctly narrower

(Fig. 8A); ventral posterior border with spinules (Fig. 8D). Anal

somite (Fig. 8C) with spinulose anal operculum.

Caudal rami (Fig. 8C—D) widely separated; with slight swelling

medially and virtually no expansion ventrally (Fig. 8B); dorsal

surface with 2 chitinous processes in posterior half; large cup-

shaped pore located ventrally (Fig. 8D) leading to small tube-pore.

Armature as in E. bulbifera.

Antennule (Fig. 9A) slender, 6-segmented, with 1 large (proxi-

mal) and 2 small spinous processes along posterior margin of

segment 1.Segment | with long spinules around anterior margin.

Segment 2 distinctly longer than segment 3. Armature formula: 1-[1

pinnate], 2-[7 + 1 pinnate], 3-[6], 4-[(2 + ae)], 5-[1], 6-[9 + acrothek].

Aesthetasc on segment 4 fused basally to 2 setae (Fig. 9B). Acrothek

consisting of aesthetasc and 2 naked setae; set on apical pedestal.

Antennary exopod (Fig. 9D) elongate exopod bearing 2 lateral

and 2 apical pinnate elements; no ornamentation discernible.

Labrum with ornamentation as in E. bulbifera.

Mandible (Fig. 9E) with small 2-segmented palp; proximal seg-

ment with 2 inner, (basal) setae; endopod a free segment with 3 setae.

Maxillule (Fig.10D) as in E. bulbifera but outer apical seta of

exopod naked and shorter and distal spine on basis stouter.

P1 (Fig. 8E) similar to that of E. bulbifera but both endopodal

segments and terminal claw shorter; exp-1 extending to distal

margin of basal pedestal; exp-2 with 3 outer setae and 2 geniculate

setae apically.

P2—P4 (Figs 9C; 10A-—B). Outer basal spine of P2 distinctly

shorter and more setiform. P2—P3 enp-1 with multipinnate inner

seta; P4 enp-1 inner seta absent. Outer spine of P2—P4 enp-2 shorter

than in E. bulbifera. Armature formula:

Exopod Endopod

P2 0.1.123 1.221

P3 0.1.223 1.321 [d: probably 1.1.220]

P4 0.1.223 0.221

P5 (Fig. 10C). Endopodal lobe elongate, clearly extending bey-

ond insertion sites of proximal outer setae of exopod; with 2 apical

and 2 widely separated inner setae; outer apical seta distinctly

shorter. Exopod more slender than in E. bulbifera; with 1 apical, 1

inner and 4 outer setae; anterior proximal seta and distalmost outer

seta much shorter.

MALE. Unknown.

REMARKS. E. profunda is known only from the type locality and

represents the deepest record for the genus. It is similar to E. lobata

in the elongate endopodal lobe of the 2 P5, the mandibular palp

setation, the ventral position of the caudal ramus pores and the

absence of the inner seta on P4 enp-1. It differs from this species in

the elongate 2 PS exopod, caudal ramus shape (presence of dorsal

chitinous processes) and the longer P1 enp-2.

Esola vervoorti sp. nov.

Esola longicauda (Edwards, 1891) sensu Vervoort (1964)

TYPELOCALITY. Ifaluk Atoll, Caroline Islands, North Pacific; stn

592 (Vervoort, 1964).

TYPE MATERIAL. National Museum of Natural History, Washing-

ton, D.C.: holotype 2 dissected on 12 slides (NMNH 109702);

paratypes are 2 dd in alcohol (NMNH 288048). Originally labelled

E. longi-cauda; det. W. Vervoort; 16 October 1953. Two other vials

with identical labels contained different species: (a) NMNH 109789:

cope-podid V 2 of E. longicauda var. sensu Vervoort (1964), from

stn 591; (b) NMNH 109790: 1 2and 1 3 of Paralaophonte sp., from

stn 590.

ETYMOLOGY. The species is named in honour of Dr Willem

Vervoort (Rijksmuseum van Natuurlijke Historie, Leiden) who first

illustrated this species.

64 R. HUYS AND W. LEE

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Fig. 8 Esola profunda sp. nov. (¥). A, Habitus, dorsal; B, urosome (excluding P5-bearing somite), lateral; C, anal somite and caudal rami, dorsal; D, right

caudal ramus, ventral; E, P1, anterior.

BASAL LAOPHONTID EVOLUTION

rior; D, antennary exopod; E,

lary segment 4; C, P4, ante

Fig.9 sola profunda sp. nov. (9). A, Antennule, dorsal; B, cylindrical outgrowth on antennu

mandibular palp.

R. HUYS AND W. LEE

\ Sal

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v. (9). A, P2, anterior; B, P3, anterior; C, P5, anterior; D, maxillule, anterior.

Fig. 10 Esola profunda sp. no

BASAL LAOPHONTID EVOLUTION

DESCRIPTION.

FEMALE. Body length from anterior margin of rostrum to posterior

margin of caudal rami 510 um; maximum width 120 um (Vervoort,

1964).

MALE. Body length from anterior margin of rostrum to posterior

margin of caudal rami 401 um (n=3; range: 389-415 um). Maxi-

mum width (121 um) measured at posterior margin of cephalothorax.

Body (Fig. 11A) more compact and abbreviated than in 9; covered

with similar dense pattern of minute spinules. Cephalothorax with

small cup-shaped pores anterodorsally and anteroventrally on either

side of rostrum; wider than free somites; without transverse spinule

row dorsally. Urosome distinctly narrower than prosome; none of

urosomites with backwardly produced posterolateral corners.

Genital somite with ventrolateral cup-shaped pores (Fig. 11B-—C).

Sixth legs (Fig. 11B—C) represented by well developed opercula,

one articulating and closing off left or right genital aperture; each

produced into cylindrical process bearing | lateral and | apical seta.

Antennule (Fig. 12A—D) 7-segmented, subchirocer, with

geniculation between segments 5 and 6. Segment 1 with spinules/

setules around anterior margin and 2 spinous processes along poste-

rior margin. Segment 4 minute, represented by incomplete sclerite.

Segment 5 longest, with large proximal process anteriorly, bearing

modified spine; forming cylindrical process bearing long aesthetasc

fused basally to 2 setae (Fig. 12C). Segment 6 with 3 spinous

processes along anterior margin. Segment 7 triangular. Armature

formula: 1-[1 pinnate], 2-[7 + 2 pinnate], 3-[6], 4-[2], 5-[7 + 2

pinnate + | spine + (2 + ae)], 6-[1 + 3 processes], 7-[7 + acrothek].

Apical acrothek consisting of aesthetasc and 2 bare setae.

Mandibular palp (Fig. 11F) small, comprising elongate basis with

1 pinnate seta and free endopod bearing 3 apical setae.

Pl (Fig. 12E) with broader basal pedestal and more robust

endopod than in E. bulbifera; enp-1 stouter and enp-2 slightly

shorter. Exopod small; exp-1 not extending to distal margin of basal

pedestal, with stout outer spine; exp-2 with 3 outer setae and 2 apical

setae, outer apical seta much shorter than in 2 and not geniculate.

P2—P4 without inner seta on enp-1 (Fig. 12F—H). P3 endopod

(Fig. 12G) 3-segmented; enp-1 as in 2; enp-2 with inner seta and

dentate outer apophysis; enp-3 small, with tube-pore, 2 lateral and 2

apical setae. Armature formula:

Exopod Endopod

P2 0.1.123 0.221

P3 0.1.223 0.1.220 [ 2: 0.321]

P4 0.1.223 0.221

P5 (Fig. 11E) medially fused, positioned ventrolaterally.

Baseoendopod without endopodal lobe; medial margin with 2 tube-

pores; outer basal seta arising from short spinulose pedestal. Exopod

free; with 1 inner seta and 1 apical plus 3 outer pinnate spines; spines

markedly shorter than in E. bulbifera.

Caudal ramus (Fig. 11B—D) rectangular, without bulbiform ex-

pansions; about 1.7 times as long as wide; with medial cup-shaped

concavity as in 2.

REMARKS. Vervoort (1964) inclined to assign specific status to his

material from the Ifaluk Atoll, however, refrained from doing so due

_ to the uncertainty about the widely recorded variability for E.

longicauda. E. vervoorti occupies an isolated position in the genus

for a number of reasons: (1) the absence of the inner seta on P2—P4

enp-1, (2) the dentate type of apophysis on the male P3 endopod, (3)

absence of transverse spinular row on cephalothorax, (4) reduced

mandibular palp, (5) very short 2 caudal rami, and (6) the sexual

dimorphism of the outer apical seta on P1 exp-2. The latter character

is unique within the Laophontidae; Vervoort (1964) also illustrated

this sexual dimorphism but did not mention it as a feature of high

significance.

Esola longicauda Edwards, 1891 sensu Noodt (1955)

Noodt (1955) illustrated a single ovigerous female of E. Jongicauda

recorded from the Sea of Marmara. His specimen is much larger

(0.79 mm) than any other species in the genus (Table I) and like

Edwards’ (1891) types shows a strongly developed seta on P4 enp-

1. It resembles E. bulbifera in the bulbiform caudal rami and the

incompletely 7-segmented antennule which according to Noodt

(1955) displays a partly subdivided apical segment. His statement

that the endopodal lobe has only 3 setae is clearly based on an error.

Without further information the identity of this specimen cannot be

determined.

Esola longicauda Edwards, 1891 var. sensu Vervoort

(1964)

This variety, known from a single male, differs from Vervoort’s

(1964) typical specimens of E. longicauda (here designated as E.

vervoorti sp. nov.) in the slender and almost haplocer antennule, the

presence of an inner seta on P2—P4 enp-1 and the shorter P4 endopod

and P5 exopod. This combination of characters rules out

conspecificity with both E. vervoorti and E. lobata, the only estab-

lished species from the Western Pacific. It also differs from Mielke’s

(1997) Esola spec. from Sulawesi by the presence of 5 setae on the

distal exopod segment of P1. It is conceivable that this variety

represents yet another species, however, the discovery of the female

is crucial before it can be attributed such status.

Esola longicauda Edwards, 1891 sensu Wells & Rao

(1986)

Wells & Rao’s (1986) record of a single female from Havelock

Island (South Andaman) is virtually indeterminable. It is probably

conspecific with Sewell’s (1940) specimens of Laophonte bulbifera

recorded from Nankauri Harbour in the Nicobar Islands and Addu

Atoll in the Maldive Archipelago. Both share the absence of the

inner seta on P4 enp-1 and their antennulary segments have similar

proportional lengths.

Esola spec. sensu Mielke (1997)

Mielke (1997) provided figures and additional information of a

single female which is potentially sympatric with E. lobata in North

Sulawesi. This form differs from the latter in the size of the proc-

esses on the first antennule segment, the shape and setal length of the

antennary exopod, mandibular armature, P1 exp-2 setation, pres-

ence of a vestigial seta on P4 enp-1 and caudal ramus shape. The

presence of only 4 setae on the distal exopod segment of P1 relates

it to E. galapagoensis and E. longicauda, however, differences in

the antennules and P4 endopod make conspecificity unlikely.

Genus Mourephonte Jakobi, 1953

Moerephonte Jakobi, 1953: lapsus calami by Vervoort (1964).

Jakobi (1953) established this genus to accommodate a new species

M. catharinensis described from the coast of Santa Catarina, Brazil.

Vervoort (1964) expressed severe doubts as to the validity of this

genus, assuming that the completely reduced P2 endopod and the

68 R. HUYS AND W. LEE

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Fig. 11 Esola vervoorti sp. noy. (3). A, Habitus, dorsal; B, urosome (excluding P5-bearing somite), ventral; C, same, lateral; D, anal somite and caudal

rami, dorsal; E, left PS, anterior; F, mandibular palp.

BASAL LAOPHONTID EVOLUTION

Fig.12 Esola vervoorti sp. nov. (3). A, Antennule, ventral (armature of segments 2—5 and 7 omitted); B, antennulary segments 2-4; C, antennulary

segment 5, ventral; D, antennulary segment 7; E, P1, anterior; F, P2 endopod, anterior; G, P3, exopod, anterior; H, P4, endopod, anterior.

aberrant setal formula most likely resulted from imperfect dissec-

tion. In addition, he suspected that Mourephonte was a junior

subjective synonym of Esola and claimed that M. catharinensis was

probably nothing more than an inadequately illustrated specimen of

Esola longicauda. Lang (1965) pointed out that Jakobi’s species had

already been described as Laophonte longiseta by Nicholls (1941a)

and regarded the absence of the P2 endopod and the reduced

armature of P2—P4 as sufficient grounds to maintain Mourephonte as

a distinct genus.

Jakobi’s material, consisting of an unspecified number of males

collected from the tidal zone at Itapocoroy and Porto Belo, is no

longer extant, the only specimen available being Nicholls’ holotype

male of L. longiseta deposited in the South Australian Museum,

Adelaide. This specimen forms the basis of the redescription given

below. The female is as yet unknown.

DIAGNOSIS (based on 6 only). Laophontidae. Body cylindrical.

Integument of cephalothorax and body somites with dense pattern of

spinules and setules. Rostrum large, partly delimited at base. Cup-

shaped pores present both anterodorsally and anteroventrally on

cephalic shield, laterodorsally on caudal rami; absent on genital

somite. Anal operculum dentate. Caudal rami rectangular, short.

Sexual dimorphism in antennule, P3 endopod, P5, P6, genital

segmentation and caudal rami.

Antennules slender; haplocer and 7-segmented in d; segment 1

with 2 small processes along posterior margin; swollen segment 5

with very long aesthetasc (fused basally to 2 setae) but without

distinct anterior outgrowth. Antenna with 4 setae on exopod; allobasis

with abexopodal seta. Labrum with marginal spinules distally. Man-

dible with small 1-segmented palp bearing 1 lateral and 3 apical

setae. Maxillule without defined exopod, represented by | setae.

Maxilla with 3 endites on syncoxa; endopod represented by 4 setae.

Maxilliped slender; syncoxa with 2 setae; entire palmar margin with

spinules; endopodal claw elongate.

P1 very large compared to other legs; with 2-segmented exopod

bearing 4—5 setae on exp-2 and elongate endopod; enp-1 without

inner seta, enp-2 with minute seta and long, slender claw. P2—P4

with 3-segmented exopods; endopods entirely absent (P2) or 2-

segmented (P3—P4). Bases with plumose (P2) or naked (P3—P4)

short outer seta. P2—P4 without inner setae on exp-2 and -3. P4 enp-

2 with widely separated apical setae. P3 endopod ¢ indistinctly

3-segmented with incomplete surface suture between enp-2 and -3;

enp-2 with inner seta and short outer, spinous apophysis. Armature

formula as follows:

Exopod Endopod

P2 0.0.022 —

P3 0.0.022 1.1.110 [in 2 presumably 1.211]

P4 0.0.022 0.111

P5 6 without endopodal lobe; exopod short, with 1 inner, 2 apical

and 2 outer setae/spines.

P6 asymmetrical; membranous flaps with 2 setae arising from

cylindrical process.

TYPE AND ONLY SPECIES. Laophonte longiseta Nicholls, 1941a =

Mourephonte longiseta (Nicholls, 1941a)

Laophonte longiseta Nicholls, 1941a

Mourephonte catharinensis Jakobi, 1953

TYPE LOCALITY. ‘Tidal zone at Itapocoroy and Pérto Belo, Santa

R. HUYS AND W. LEE

Catarina State, Brazil; holdfasts of Endocladia and Codium.

MATERIAL EXAMINED. South Australian Museum, Adelaide:

Holotype 3 of Laophonte longiseta, dissected on slide Tc 13437

(SAM C5550); Sellick Beach, south of Port Willunga, South Aus-

tralia; coll. H.M. Hale, 31 January 1937, froma stone in 1.5 mat low

tide on south edge of reef. Jakobi’s (1953) type material of M.

catharinensis is lost.

REDESCRIPTION.

Unknown.

MALE. Body length 0.25 (Jakobi, 1953) to 0.30 mm (Nicholls,

1941a). Cephalic shield with paired cup-shaped pores both

anterodorsally and anteroventrally on either side of rostrum.

Antennule (Fig. 14A—F) 7-segmented, haplocer; geniculation

between segments 5 and 6; proximal segments without conspicuous

spinous processes but segment | with 2 small protuberances; seg-

ment | with spinular row distally and tiny spinules along anterior

margin; segment 2 longest; segment 5 with very long aesthetasc

(150 um). Armature formula: 1-[1], 2-[8 + 1 pinnate], 3-[6], 4-[2], 5-

[8 + 1 pinnate + 1 spine + (2 + ae)], 6-[1 + modified seta], 7-[7 +

acrothek]. Acrothek consisting of 2 basally fused setae.

Antennary exopod (Fig. 13D) with 2 pinnate setae laterally and 2

pinnate spines distally.

Labrum with marginal spinules distally; without overlapping

scales.

Mandibular palp (Fig. 13C)1-segmented, bilobate, rami com-

pletely incorporated; with 1 pinnate seta laterally (probably basal in

origin) and 3 bare setae distally (representing incorporated endopod).

Maxillule and maxilla as in the genus Esola.

Maxilliped (Fig. 13B) slender; syncoxa with 3 spinular rows and

2 pinnate setae; basis elongate, with long spinular row on palmar

margin and spinular patch on outer margin; endopod represented by

tiny setule and very long claw, exceeding length of basis.

Pl (Fig. 13A) large compared to P2—P4; protopodal segments

with rows and patches of fine spinules as illustrated; basis with outer

spine near joint with coxa and inner pinnate spine on anterior

surface. Exopod 2-segmented, exp-1 with pinnate outer spine; exp-2

with 2 spines and 3 geniculate setae. Endopod very long; enp-1

without inner seta; enp-2 with 3 spinular rows, | setule and long,

denticulate claw.

P2—P4 (Fig. 14G-I) with 3-segmented exopods; endopod 2-

segmented (P3—P4) or entirely absent (P2). P3 enp-2 partly

subdivided along anterior surface by short transverse suture; apo-

physis on outer margin short and slightly sigmoid, bare. P4 enp-2

with apical setae widely separated and flanking secretory tube-pore.

Armature formula of P2—P4 as for genus.

P5 (Fig. 14J) with baseoendopod fused to somite, endopodal lobe

not developed. Exopod rectangular; with 2 outer, 1 apical and 2

inner setae. P6 asymmetrical, produced into cylindrical process at

outer corner, bearing long apical and shorter inner seta.

Pleural areas of genital somite without modified pores. Posterior

margins of abdominal somites with row of long spinules (Fig. 13F).

Anal operculum dentate (Fig. 13E).

Caudal rami (Fig. 13E—F) short, about 1.3 times as long as wide;

with 6 setae (seta I absent), seta VII tri-articulate at base and

plumose, setae [TV and V well developed and fused at base. Inner

proximal margin with cup-shaped depression (specialized pore)

dorsally, marked by row of tiny spinules set on strongly chitinized

margin; cup filled with secretory substance.

REMARKS. Neither Jakobi (1953) nor Nicholls (1941a) illustrated

cup-shaped pores on the cephalic shield. Vervoort (1964) pointed

out that Jakobi had shown an anteriorly directed middorsal spinous

FEMALE.

BASAL LAOPHONTID EVOLUTION

| i , as

/} Ny

i Ip y

bin

h EN NANNY Hypa ram

r Dili ya

PPD Nyy p yqyy yyy

AAR

i i _"_

SLLSLA TTI

959 EE ~ | f,

oe <

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LLL |

anal somite and right caudal ramus, dorsal; F, posterior urosomites and right caudal ramus, ventral.

Hm me ey

Ge uy

| yy

—

oS =

\\\

ae ME,

——

Vai

oA ees

Fig. 13 Mourephonte longiseta (Nicholls, 1941a) (3). A, P1, anterior; B, maxilliped; C, mandibular palp; D, coxa, allobasis and exopod of antenna; E,

S555

= Se

Lkllezs

a eee

SS PVS=s SS SSS

LES ee 2

SoS Le

) SS SSS

Ss aS = | = Co

—7 largely omitted); B—F, antennulary segments 3-7;

Fig. 14 Mourephonte longiseta (Nicholls, 1941a) (3). A, Antennule, ventral (armature of segments 3

G, P2, anterior; H, P3, posterior; I, P4, anterior (apical tube-pore arrowed); J, right P5, anterior.

BASAL LAOPHONTID EVOLUTION

process which was not illustrated in the lateral habitus view, possi-

bly indicating that the author had indeed observed but incorrectly

figured the cephalic pores. We have re-examined Nicholls’ slide

material and found remnants of the cephalic shield, confirming the

presence of both anterodorsal and anteroventral pores as in Esola.

Scrutinous observation failed to reveal any such structures on the

genital somite.

With only two records known, M. longiseta appears to display a

remarkably disjunct distribution. Topotype material from Brazil is

required to confirm whether the morphometric discrepancies in

Jakobi’s description result from imperfect observation or reflect

species level differences between the Brazilian and Australian popu-

lations.

}

Genus Archilaophonte Willen, 1995

DIAGNOSIS. Laophontidae. Body elongate and slender; cephalo-

thorax slightly wider than rest of body; posterolateral corners of 2

genital double-somite and second abdominal somites not laterally or

backwardly produced. Integument of cephalothorax and body somites

with dense pattern of spinules and setules; cup-shaped pores on

cephalothorax, genital (double-)somite and caudal rami absent.

Rostrum very large, partly delimited at base. Integumental cup-

shaped pores absent. Anal operculum spinulose. Caudal rami very

long, cylindrical with posterior halves diverging.

Sexual dimorphism in body shape, antennule, P3 endopod, P5, P6

and in genital segmentation.

Antennules short; 6-segmented in 2, subchirocer and 7-seg-

mented in d;posterior margin of segment 1 with small blunt process,

that of segment 2 with distinct spinous process; with aesthetasc on

segment 4 () or 5 (6) and as part of apical acrothek on distal

segment; segment 6 of ¢ not particularly modified; proximal

aesthetasc fused to 1 seta. Antenna with 4 setae on exopod; allobasis

with abexopodal seta. Mandible with biramous palp bearing discrete

1-segmented rami; basis with | lateral seta, exopod with 1, endopod

with 3 apical setae. Maxillule with seta at base of exopod. Maxilla

with 3 endites on syncoxa; endopod represented by 4 setae.

Maxilliped moderately slender; with 3 setae on syncoxa; endopodal

claw long and slender.

Pl with 2-segmented exopod bearing 5 setae on exp-2 and

elongate endopod; enp-1! with inner seta, enp-2 with minute seta and

long, slender claw. P2—P4 with 3-segmented exopods and 2-seg-

mented endopods. P2 basis with normally developed outer spine.

Outer spine of P4 enp-2 not very long. P3 endopod d 3-segmented;

enp-2 with inner seta and very long, slender, sigmoid apophysis. P3

exopod d weakly modified with exp-3 being shorter than in 9

Armature formula as follows:

Exopod Endopod

P2 0.1.123 Pl

P3 0.1.223 1.321 [d: 1.1.220]

P4 0.1.223 1.221

P5 2with separate rami; exopod large and elongate, with 6 setae/

spines; baseoendopod well developed, with 5 setae/spines. PS 3

with trapezoid endopodal lobe bearing 2 long setae; exopod rectan-

gular, with 1 inner, 1 outer and 2 apical setae/spines.

P6 2forming opercula closing off paired genital apertures; with 2

‘long setae. P6 d asymmetrical; membranous flaps with | tiny seta.

TYPE AND ONLY SPECIES.

[by monotypy].

Archilaophonte maxima Willen, 1995

TYPE LOCALITY. 72°52.3'S, 19°34.7' W, Weddell Sea, Antarctic;

495 m depth.

REMARKS. Willen (1995) described A. maxima in great detail; the

slight sexual dimorphism illustrated for the P3 exopod was not

mentioned in the text. The species is known from two localities in

the Weddell Sea.

Genus Applanola gen. nov.

DIAGNOSIS. Laophontidae. Body strongly depressed and com-

paratively short; cephalothorax much wider than rest of body;

posterolateral corners of ¢ genital double-somite and second ab-

dominal somites laterally and backwardly produced. Integument of

cephalothorax and body somites with dense pattern of spinules and

setules. Rostrum very large, partly delimited at base. Four pairs of

integumental cup-shaped pores present: anterodorsally on

cephalothorax, near ventrolateral margins of cephalic shield, later-

ally on genital (3) or genital double-somite (9) and ventrally on

caudal rami. Anal operculum spinulose. Caudal rami short, squar-

ish.

Sexual dimorphism in body shape, antennule, P2—P4 exopods, P3

endopod, P5, P6 and in genital segmentation.

Antennules short; 6-segmented in 9, subchirocer and 7-seg-

mented in 3; segments 1—2 without distinct processes; with aesthetasc

on segment 4 () or 5 (d) and as part of apical acrothek on distal

segment; segment 6 of d with large bilobate outgrowth dorsally;

proximal aesthetasc fused basally to 2 setae. Antenna with 4 setae on

exopod; allobasis with abexopodal seta. Labrum with distal patch of

long spinules. Mandible with elongate 1-segmented palp with 1

lateral and 3 apical setae. Maxillule with elongate defined exopod.

Maxilla with 3 endites on syncoxa; endopod represented by 4 setae.

Maxilliped large and robust; with 2 setae on syncoxa; endopodal

claw relatively short.

P1 with 2-segmented exopod bearing 5 setae on exp-2 and robust

endopod; enp-1 without inner seta, enp-2 with minute seta and short,

strongly curved claw. P2—P4 with 3-segmented exopods and 2-

segmented endopods. P2 basis with very long outer spine. Outer

spine of P4 enp-2 very long. P3 endopod d 3-segmented; enp-2 with

inner seta and outer dentate apophysis. P3 exopod ¢ strongly

developed with modified outer and distal spines on exp-3; exopods

of P2 and P4 similar in size to 2 but with stronger ornamentation on

outer spines. Armature formula as follows:

Exopod Endopod

P2 0.1.123 1.220

PS 0.1.223 1.321 [d: 1.1.220]

P4 0.1.223 1.221

P5 ¢ with separate rami; exopod elongate, with 6 setae/spines;

baseoendopod slightly developed, with 4 setae/spines. P5 d without

endopodal lobe; exopod short, with 1 inner, 2 apical and 2 outer

setae/spines.

P6 ¢ forming opercula closing off paired genital apertures; with

one seta and 2 small processes at outer corner. P6 ¢ asymmetrical;

membranous flaps without armature.

TYPE AND ONLY SPECIES. Laophonte hirsuta Thompson & A.

Scott, 1903 = Applanola hirsuta (Thompson & A. Scott, 1903)

comb. nov.

ETYMOLOGY. The generic name is derived from the Latin ad (to)

and planatus (flattened), and alludes to the dorsoventrally depressed

body. Gender: feminine.

Applanola hirsuta (Thompson & A. Scott, 1903) comb.

nov.

Laophonte hirsuta Thompson & A. Scott, 1903

Esola hirsuta (Thompson & A. Scott, 1903): Lang (1948)

Thompson & A. Scott (1903) described Laophonte hirsuta from

washings of pearl oysters and other unidentified invertebrates dredged

in the Gulf of Manaar, Sri Lanka. A. Scott (1909) reported the species

from 1595 m in the Banda Sea (Indonesia) but this record is almost

certainly the result of contamination by ashallow water sample (Lang,

1948; Lee & Huys, 1999). The unknown male was described by

Gurney (1927) from Port Taufiq in the Suez Canal. Por’s (19645)

records from Haifa Bay and off the coast of Caesarea are likely the

result of Lessepsian migration. Both Krishnaswamy (1957) and

Krishna Murty (1983) reported the species from the Bay of Bengal.

Krishnaswamy collected adults and developmental stages from

sponges taken off the Krusadai Islands. Krishna Murty reported some

occasional specimens in algal washings from the Visakhapatnam

coast. The only other record outside the Indo-Pacific is that by Pesta

(1916) from Sao Tomé in the Gulf of Guinea. Lang (1944, 1948)

placed the species in the Jongicauda-group of Esola.

TYPE LOCALITY. Muttuvaratu, Sri Lanka; washings of pearl oys-

ters and other dredged invertebrates.

MATERIAL EXAMINED. Cambridge Suez Canal Expedition 1924;

Port Taufiq (Egypt): 1 2 dissected on 14 slides (BMNH 1999.982),

1 d dissected on 11 slides (BMNH 1999.983); 3 29(1 damaged), 2

36 and | copepodid V 6 in alcohol (BMNH 1928.4.2.111).

REDESCRIPTION.

FEMALE. Body length from anterior margin of rostrum to posterior

margin of caudal rami 664 um (n=3; range: 650-690 um). Maxi-

mum width (281 wm) measured at posterior margin of cephalothorax.

Body very dorsoventrally depressed, covered with dense pattern

of minute spinules dorsally (Fig. 30A). Cephalothorax much wider

than free somites, posterolateral angles backwardly produced; with

paired cup-shaped pores both anterodorsally and anteroventrally on

either side of rostrum (arrowed in Fig. 15A—B and 30A-B),

anterodorsal set partly closed off by fringe of setular extensions.

Posterior margin of cephalothorax and all body somites with row of

long spinules dorsally and laterally. Ventrolateral areas of cephalic

shield and pleurotergites of first two pedigerous somites with long

spinules and setules (Fig. 31B; ventral surface with distinct vent-

pore at level of mandibles (Fig. 31B). Pleurotergite of P5-bearing

somite wide.

Genital double-somite (Fig. 17A—B) only slightly narrower than

pedigerous somites (Fig. 15A); original segmentation marked by

bilateral constriction and dorsal transverse spinule row; anterior (=

genital) half with large cup-shaped pores laterally (Fig. 29A), each

partly closed off by fringe of setular extensions (Fig. 29B—C);

posterior half with backwardly directed lobate extensions bearing

spinular tuft (Fig. 29A); ventral surface without spinular ornamen-

tation except for spinule row around posterior margin; genital field

located near anterior margin. Sixth legs (Fig. 17C) forming well

developed opercula closing off paired genital apertures; each with

naked seta and 2 small processes at outer corner; inner corner

produced into paired, medially directed, spinous processes.

Postgenital somites with spinules around ventral hind margin;

second abdominal somite with posteriorly directed lateral angles,

bearing spinular tuft; penultimate and anal somites distinctly nar-

rower. Anal somite with paired oblique spinule rows on ventral

surface; anal operculum spinulose.

Caudal rami (Fig. 15A, C) widely separated; shorter than wide;

R. HUYS AND W. LEE

inner margin with medial protrusion; ventral surface with 2 spinule

rows and large slit-like pore (arrowed in Figs. 15C; 30C) connected

with spacious subsurface duct, extending into anal somite; entrance

to pore with fine setules (Fig. 30D); dorsal surface with minute

spinules; setae I-III all well developed, naked and closely set; setae

IV and V pinnate and with fracture planes, seta V twice as long as

seta IV; setae VI-VII naked.

Rostrum (Fig. 15A) large, rounded anteriorly; partly delimited at

base by transverse surface suture (Fig. 30A); with paired sensillae

anteriorly.

Antennule (Figs 15A; 16A) short, 6-segmented, without proc-

esses on segments 1—2. Segment 1 with dorsal spinular patch.

Armature formula: 1-[1 pinnate], 2-[4 + 4 pinnate], 3-[2 + 2 pin-

nate], 4-[(2 + ae)], 5-[1], 6-[6 + 3 pinnate + acrothek]. Acrothek

consisting of aesthetasc and 2 naked setae; set on apical pedestal.

Antenna (Fig. 16B) with well developed exopod bearing 2 lateral

and 2 apical pinnate elements. Allobasis with pinnate abexopodal

seta accompanied by setular patch. Endopod with lateral armature

consisting of 2 spines and | seta; distal armature consisting of 2

unipinnate spines and 3 geniculate setae (outermost shortest and

fused basally to setule).

Labrum with elaborate ornamentation around distal margin (Fig.

20E) but without spinules or scales on anterior face (Fig. 29D).

Mandible (Fig. 16C) with elongate gnathobase and long 1-seg-

mented palp (Fig. 31B) probably representing fused basis and

endopod; with | lateral and 3 distal pinnate setae.

Paragnaths densely hirsute lobes as in Fig. 20D.

Maxillule (Fig. 16D) with well developed praecoxa bearing | seta

on anterior surface and 8 elements around distal margin. Coxal

endite with 2 setae, basal endite with 1 spine and 2 setae. Exopod an

elongate segment with 2 distal setae; endopod incorporated into

basis, represented by 2 setae.

Maxilla (Fig. 20F). Syncoxa with long coarse spinules around

outer margin; with 3 endites; praecoxal endite small and unisetose;

middle endite drawn out into pinnate claw, with 2 setae; distal endite

with 3 elements. Allobasis produced into strong curved claw; acces-

sory armature consisting of | spine and 1 seta. Endopod a minute

segment with 4 setae of different lengths.

Maxilliped (Fig. 17D) compact, with relatively short basis and

endopodal claw. Syncoxa with 2 pinnate setae. Basis with spinular

ornamentation as figured. Endopod represented by unipinnate claw

bearing | accessory seta and tube-pore at base.

P1 (Fig. 19E) with narrow coxa and basis. Basis with pinnate seta

on anterior surface and along outer margin. Exopod 2-segmented,

small compared to endopod; exp-1 with pinnate outer seta; exp-2

with 3 distinctly pinnate outer setae and 2 geniculate setae apically.

Endopod robust; enp-1 with long setules along inner margin; enp-2

with short, hook-like, naked claw and small accessory seta.

P2-P4 (Figs 17F; 18A, C) with 3-segmented exopods and 2-

segmented endopods. P2 basis with very long, multipinnate outer

spine; P3—P4 bases with bare outer seta. P2—P4 exp-2 with well

developed inner seta. P2—P4 enp-1 small, with inner seta. P2 enp-2

without outer spine; outer spine of P3—P4 enp-2 very long. Tube-

pore present near distal outer corner of P3—P4 enp-2. Armature

formula as for genus.

P5 (Fig. 17E). Endopodal lobe reduced, not extending beyond

proximal outer setae of exopod; with 1 short and 1 long pinnate seta

apically, and 2 long widely separated setae along inner margin;

anterior face with 2 tube-pores. Exopod elongate, produced apically

into tubular extension bearing | bare seta; inner margin with 1, outer

margin with 4 pinnate setae; inner seta much shorter than apical one.

Both baseoendopod and exopod with elaborate ornamentation pat-

tern as figured.

BASAL LAOPHONTID EVOLUTION 15

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| Fig. 15 Applanola hirsuta (Thompson & A. Scott, 1903) comb. nov. (9). A, Habitus, dorsal; B, habitus, lateral; C, left caudal ramus, ventral. [Arrows

indicating anteroventral cup-shaped pores in A-B, ventral one in C].

R. HUYS AND W. LEE

Sa ae 7

v. (9). A, Antennule, dorsal; B, antenna; C, mandible; D, maxillule.

Fig. 16 Applanola hirsuta (Thompson & A. Scott, 1903) comb. no

BASAL LAOPHONTID EVOLUTION 77

Fig.17 Applanola hirsuta (Thompson & A. Scott, 1903) comb. nov. A, Urosome 2 (excluding P5-bearing somite), ventral; B, same, lateral; C, left genital

aperture 9, ventral; D, maxilliped; E, PS Q anterior; F P2 2 anterior; G, P2 exopod d, anterior.

R. HUYS AND W. LEE

5B,

rior; D, P4 exopod 4, anterior

; C, P4 9 ante

rior; B, P3 d, anterior

Fig. 18 Applanola hirsuta

(Thompson & A. Scott, 1903) comb. nov. A, P3 &, ante

rticulated enp-2 and -3, anterior.

P3 6, disa

BASAL LAOPHONTID EVOLUTION

7 it

7 141

Fig. 19 Applanola hirsuta

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TUSTIN

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(Thompson & A. Scott, 1903) comb. nov. A, Habitus 3, dorsal; B, urosome ¢, ventral; C, PS 6, anterior; D, genital apertures

6, ventral [arrows indicating absence of armature]; E, P1, anterior.

80 R. HUYS AND W. LEE ;

cece

Fig. 20 Applanola hirsuta (Thompson & A. Scott, 1903) comb. nov. A, Antennule 6, dorsal [armature of segments 4—5 omitted]; B, antennulary

segments 3-4 of d, dorsal; C, antennulary segment 5 of d, anterior; D, left paragnath; E, labrum, anterior; F, maxilla.

BASAL LAOPHONTID EVOLUTION

MALE. Body length from anterior margin of rostrum to posterior

margin of caudal rami 581 um (n=3, range 570-595 um). Maximum

width (248 um) measured at posterior margin of cephalothorax.

Length of d copepodid V: 571 um.

Body (Fig. 19A) very dorsoventrally depressed, covered with

dense pattern of minute spinules as in 2. Pattern of cup-shaped pores

as in 2 except for paired lateral pores present on genital somite.

Cephalothorax much wider than free somites, posterolateral angles

backwardly produced. Posterior margin of cephalothorax and all

body somites with row of long spinules dorsally and laterally.

Pedigerous somites decreasing in width posteriorly. Urosome (Fig.

19B) slender and narrow; pleurotergite of P5-bearing somite nar-

row; posterolateral corners of all urosomites with spinular tuft and

posterior margin with spinules all around.

Genital somite with large cup-shaped pores laterally, each partly

closed off by fringe of setular extensions (Fig. 19D); ventral surface

without spinular ornamentation except for spinule row around pos-

terior margin. Sixth legs represented by membranous flaps, one

articulating and closing off left or right genital aperture; without

armature at outer corner.

Antennule (Fig. 20A—C) 7-segmented, subchirocer, with

geniculation between segments 5 and 6. Segment | with spinules/

setules around anterior margin. Segment 2 with minute knob near

dorsal posterior margin. Segment 4 minute, represented by incom-

plete sclerite. Segment 5 with spinous outgrowth on anterior margin,

probably interlocking with similar processes on segment 6 (Fig.

20C); forming cylindrical process bearing long aesthetasc. Segment

6 with bilobed outgrowth on ventral surface near posterior margin.

Distal portion of segment 7 elongate, displacing acrothek to position

isolated from other armature. Armature formula: 1-[1 pinnate], 2-[4

+5 pinnate], 3-[7 + 1 pinnate], 4-[2], 5-[7 + 1 pinnate + 1 spine + (2

+ ae)], 6-[1 + 2 processes], 7-[7 + acrothek]. Apical acrothek

consisting of aesthetasc and 2 bare setae.

P2 exopod (Fig. 17G). Outer spines of all segments with much

longer pinnules than in &.

P3 (Fig. 18B, E). Exopod more robust than in &, slightly bent

medially; outer spine of exp-1 with longer pinnules than in 2; middle

and distal outer spines and apical spine of exp-3 enlarged, with

minute spinules; inner and inner apical setae reduced in length.

Endopod 3-segmented; enp-1 larger than in 9, densely setulose

along outer margin; enp-2 with inner seta and short outer apophysis

bearing small spinous processes along both inner and outer margin

(Fig. 18E); enp-3 small, with long tube-pore and 4 setae.

P4 exopod (Fig. 18D). Proximal segment slightly more robust

than in 2 Outer spines of exp-2 and -3 stubby and somewhat

enlarged; spinules typically longer than in &.

P5 (Fig. 19C) medially fused (Fig. 19B) positioned ventrolater-

ally. Baseoendopod without endopodal lobe; medial margin with

setules and tube-pore; outer basal seta arising from short spinulose

pedestal. Exopod free; with 3 multipinnate (1 apical, 2 outer) and 2

bipinnate (inner) setae, all well developed.

REMARKS. Thompson & A. Scott (1903) illustrated the female P5

with only 3 setae on the baseoendopod, a character included with

hesitation by Lang (1948) in the diagnosis of the species. Re-

examination revealed that the innermost seta on the endopodal lobe

was overlooked. This seta is implanted medially at considerable

distance from the others and was also missed by Norman (1911) in

his description of Laophonte bulbifera. According to Lang’s (1948)

table XXIV the swimming leg armature formula is constant within

the Jongicauda-group, including amongst other patterns the pres-

ence of the outer spine on P2 enp-2. One cannot but conclude that

Lang (1948) must have overlooked Gurney’s (1927) statement that

this segment has 2 inner and 2 apical setae. The present redescription

has revealed the sexual dimorphism of the exopods of P2 and P4, the

presence and pattern of integumental cup-shaped pores, and the

detailed morphology of the genital area in both sexes.

Krishnaswamy’s (1957) redescription is grossly inadequate and

potentially misleading. The ramus labelled *P2 end 2” is the male P3

endopod, his illustration of the female P2 is in fact based on the P3

and the real P2 is figured as the P7(!). In view of these inaccuracies

the tabulated setal formula and the author’s remarks on the generic

placement of the species are best ignored. Krishnaswamy’s descrip-

tion of the first copepodid is of similarly abominable quality.

Genus Archesola gen. nov.

This genus is proposed to include Esola typhlops and a number of

closely related species. It is difficult to understand why Lang (1965)

regarded Laophonte lamellipes Nicholls as most closely related to E.

typhlops. This doubtful statement was based on the similarity in the

long caudal rami and the erroneous fact that males of both species

show no modifications on the P3 endopod. Noodt (1955) suggested

arelationship with the Laophonte setosa-group but did not elaborate

on this view. Re-examination of Nicholls’ (1944) type material

(BMNH 1947.10.6.23—27) revealed the true nature of the modified

male P2 endopod, confirming close affinity with the genus

Paralaophonte Lang.

DIAGNOSIS. Laophontidae. Body cylindrical or dorsoventrally

depressed; posterolateral corners of 9 genital double-somite and

second abdominal somite laterally but not backwardly produced.

Integument of cephalothorax and body somites with irregular pat-

tern of minute surface lamellae. Rostrum large, partly delimited at

base by surface furrow. Integumental cup-shaped pores absent on

cephalothorax, genital (double-)somite and caudal rami. Anal oper-

culum smooth or bordered with spinules. Caudal rami cylindrical

and elongate; not sexually dimorphic.

Sexual dimorphism in antennule, P3 endopod, P5, P6 and in

genital segmentation.

Antennules slender; 7-segmented in 9, haplocer and 7-segmented

in 6;segments 1—2 without spinous processes along posterior margin;

with aesthetasc on segment 4 ( 9) or 5 (3) andas part of apical acrothek

on distal segment; segment 5 d not swollen, without anterior out-

growthbut with very long cylindrical pedestal for aesthetasc; proximal

aesthetasc fused to 2 setae. Antenna with 4 setae on exopod; allobasis

with abexopodal seta. Labrum with distal spinular ornamentation.

Mandible with discrete 1-segmented exopod bearing | seta; endopod

(3 setae) and basis (2 setae) incompletely fused. Maxillule with

minute, defined exopod. Maxilla with 3 endites on syncoxa; endopod

represented by 4 setae. Maxilliped slender; syncoxa with 2 setae;

palmar margin naked; endopodal claw elongate.

Pl with 3-segmented exopod bearing 4 setae on exp-3 and

elongate endopod; enp-1 with inner seta, enp-2 with minute seta and

strong claw. P2—P4 with 3-segmented exopods and 2-segmented

endopods. P2 basis with long outer spine. Outer spine of P2—P4 enp-

2 setiform and very long in P3—P4. P3 endopod ¢ 2-segmented;

enp-2 with 3 inner setae and short outer basally fused spine. Arma-

ture formula as follows:

Exopod Endopod

P2 0.1.123 1.221

P3 0.1.223 1.321 [2and 3]

P4 0.1.223 1.221

P5 ¢ with separate rami; exopod elongate, with 6 setae/spines;

baseoendopod slightly developed, with 5 setae/spines. P5 d without

endopodal lobe; exopod short, with 2 outer, 1 apical and 2 inner

elements (distal inner spiniform). Outer basal seta arising from long,

articulating, cylindrical setophore in both sexes.

P6 ¢forming opercula closing off paired genital apertures; with 2

small setae at outer corner. P6 ¢ asymmetrical; membranous flaps

with 1 apical and 1 lateral seta.

TYPESPECIES. Laophonte typhlops Sars, 1908 =Archesola typhlops

(Sars, 1908) comb. nov.

OTHER SPECIES. Laophonte longiremis TY. Scott, 1905 = A.

longiremis (T. Scott, 1905); A. hamondi sp. nov.

SPECIES INQUIRENDAE. Esola sp. sensu Chislenko (1967); Esola

typhlops pontoica Por, 1959 = A. typhlops pontoica (Por, 1959)

comb. nov.

ETYMOLOGY. The Greek prefix arche alludes to the primitive

position of the genus.

Archesola typhlops (Sars, 1908) comb. nov.

Laophonte typhlops Sars, 1908

Esola typhlops (Sars, 1908) Lang (1948)

TYPE LOCALITY. Flekker6, south coast of Norway, 36 m depth.

MATERIAL EXAMINED.

(1) West Runton, Norfolk, England: 1 6d dissected on 8 slides

(BMNH 1999.1079); collected among Polyclinum and Morchellium

under rocks; leg. R. Hamond, September 1971;

(2) Frierfjord/Langesundfjord, Norway: 4 damaged 2° (3 in alco-

hol: BMNH_ 1999.1081—1083; 1 dissected on 5 slides: BMNH

1999.1080); 99 m, mud, leg. R. Huys, 1985;

(3) Gullmar Fjord, Sweden: 1 2 in alcohol (NMNH 90955); 30 m,

sand; leg. K. Lang, 08 July 1942.

REDESCRIPTION.

FEMALE. Body length from anterior margin of rostrum to posterior

margin of caudal rami 585 um (n=4; range: 575-592 um).

Body cylindrical, not dorsoventrally depressed, covered with

dense pattern of minute surface ridges dorsally and laterally.

Cephalothorax with almost parallel lateral margins in posterior two-

thirds, without paired cup-shaped pores. Posterior margin of

cephalothorax and all body somites with row of long setules dorsally

and laterally. Posterior margin of urosomites with spinules all

around (Fig. 23B); ventrolateral areas of cephalic shield and

pleurotergites of pedigerous somites with longer setules. Pleurotergite

of P5-bearing somite narrowest.

Genital double-somite (Fig. 23A) wide and dorsoventrally flat-

tened; original segmentation marked by bilateral constriction and

transverse surface ridge dorsally; without cup-shaped pores in ante-

rior half; lateral lobes in both anterior and posterior halves with

backwardly directed strong spinules; ventral surface without orna-

mentation except for spinules around hind margin and 2 pairs of

medial tube-pores. Genital field located near anterior margin (Fig.

23A); copulatory pore minute. Sixth legs forming well developed

opercula closing off paired genital apertures; each with 2 naked

setae.

Anal somite (Fig. 23B) with coarse spinules on anal operculum.

Caudal rami (Figs. 23B; 24F; 31C—D) widely separated, cylindri-

cal and slightly tapering posteriorly; without cup-shaped pores;

about 4 times as long as wide; setae I-III closely set, I minute, I-III

very long and thin; setae IV and V pinnate and with fracture planes,

R. HUYS AND W. LEE

seta IV distinctly longer than caudal ramus; setae VI-VII naked.

Vent-pore and small tube-pore present ventrally near insertion sites

of setae I-III (Fig. 31C—D).

Rostrum as in d (Fig. 22B); large, trapezoid with straight anterior

margin; delimited at base by transverse surface suture; with paired

sensillae anteriorly and median tube-pore ventrally.

Antennule (Fig. 22A) slender, 7-segmented; segments 1—2 with-

out processes. Segment | with spinules around anterior margin;

segment 4 forming large cylindrical pedestal ventrally. Armature

formula: 1-[1], 2-[8 + 1 pinnate], 3-[6], 4-[(2 + ae)], 5-[1], 6-[2], 7-

[7 + acrothek]. Aesthetasc on segment 4 fused basally to 2 setae.

Acrothek consisting of aesthetasc and 2 naked setae; set on small

tubercle.

Antenna (Fig. 23C) with elongate exopod bearing 2 lateral and 2

apical pinnate setae, and a longitudinal row of coarse spinules. Coxa

with few large spinules, allobasis with pinnate abexopodal seta.

Endopod with lateral armature consisting of | seta, 1 large and 1

small spine; distal armature consisting of 2 unipinnate spines and 3

geniculate setae (outermost fused basally to small seta).

Labrum as in A. hirsuta.

Mandible (Fig. 25A) with short gnathobase and small bilobed

palp representing partially fused basis and endopod; with 2 lateral

(basal) pinnate setae and 3 distal (endopodal) setae; exopod repres-

ented by minute segment bearing 1 apical seta.

Maxillule (Fig. 25B) and maxilla as in E. bulligera.

Maxilliped (Fig.23D) slender, with elongate basis and endopodal

claw. Syncoxa with 2 pinnate setae. Basis with naked palmar margin

and setules around outer margin. Endopod represented by very long,

naked claw bearing 1 accessory seta at base.

P1 (Fig. 22F) with sparse ornamentation on coxa and basis. Basis

with pinnate seta on anterior surface and along outer margin. Exopod

3-segmented, well developed; exp-1 with long pinnate outer spine;

exp-2 with | naked outer spine; exp-3 with 2 unipinnate lateral setae

and 2 geniculate setae apically. Endopod long and slender; enp-1

with long setules along inner margin and shorter spinules along

outer margin, with thin inner seta in distal quarter (arrowed in Fig.

22F); enp-2 about twice as long as wide, with strong minutely

pinnate claw and small accessory seta.

P2—P4 as in Sars (1908). P3 enp-2 (Fig. 24B) with setiform outer

spine (arrowed). Armature formula typical for genus.

P5 (Fig. 23E). Endopodal lobe well developed, not extending

beyond insertion sites of proximal outer setae of exopod; with

distinctly stepped inner margin bearing 2 strong spines and | long

distal seta (extending beyond apex of exopod); apex with 2 setae,

outer one about twice length of inner one; tube-pores present near

apical setae and proximal to innermost spine; outer basal seta

inserting on cylindrical articulating setophore. Exopod narrow and

elongate, produced apically into long tubular extension bearing |

bare seta; inner margin with 1, outer margin with 1 naked and 3

pinnate setae. Both baseoendopod and exopod with elaborate or-

namentation pattern as figured.

MALE. Body length from anterior margin of rostrum to posterior

margin of caudal rami 475 um. Sexual dimorphism in antennule, P3

endopod, P5, P6 and genital segmentation.

Antennule (Fig. 22B—E) 7-segmented, haplocer, with geniculation

between segments 5 and 6. Segment 1 with spinules/setules around

anterior margin; segment 2 longest; segment 4 minute, represented

by incomplete sclerite (Fig. 22C). Segment 5 with large process

proximally but forming long cylindrical pedestal distally (Fig. 22D).

Segment 6 with 3 spinous processes along anterior margin (Fig.

22E). Distal portion of segment 7 elongated, displacing acrothek to

position isolated from other armature (Fig. 22E). Armature formula:

BASAL LAOPHONTID EVOLUTION

prey ya on

A eee

; $

UO RS

TAC vin aS AX

ESE

d ‘ =o

TS ah

rie A

Rat

Pee

Mach

pity

ytd

A gare

itp

i, \

Hig tae

7 rs!

t 1

Si AML ris | = ¥

BI AFRAIC RS ES ST

RE OSFK OEM KA, YY,

LES ae ST t\. ‘4

Bes

SOYA

A+100 yd 7 ea B} 100,

i ; 1

ton fe

Mi PEPPPPER TODD.

5 eo sp AN RU

| ca rr

“nm " 8 <

LI ~s

a4 eo :

R&S

‘a

Vy\y Vas

pep ayy \ wi

Tifity VASSA ASSES iN

' S

pa yaa AS

1 ! \ MK

1 ee

Say

\ ys

AMANITA

it ww

is Y/ Mh \ \

% an \ al

he .

Fig. 21 Archesola hamondi gen. et sp. nov. A, Habitus 9, dorsal. Corbulaseta bulligera (Farran, 1913) comb. nov. B, habitus 9, dorsal.

R. HUYS AND W. LEE

SEE ADR Es

pS ——

GRY,

mae Nttisres

Fig. 22 Archesola typhlops (Sars, 1908) comb. noy. A, Antennule °, ventral; B, rostrum and antennule 4, dorsal [armature of segments 3—7 omitted]; C

antennulary segments 3-4 6; D, antennulary segment 5 d; E, antennulary segments 6-7 6; F, P1 Q anterior [inner seta on enp-2 arrowed].

a; D,

(

tral; B, anal somite and right caudal ramus, dorsal; C, antenn

v. F, PS , anterior.

v. (2). A, Genital double-somite, ven

Fig. 23. Archesola typhlops (Sars, 1908) comb. no

maxilliped; E, P5, anterior. Archesola hamondi gen. et sp. no

BASAL LAOPHONTID EVOLUTION

R. HUYS AND W. LEE

SSS — ’

Sif iil =

sa! ;

udal ramus, dorsal. Archesola

v. B, P3 endopod 9 anterior [elongate outer spine arrowed]; C, P3 protopod and endopod 4, anterior [apophysis

arrowed]; D, right P5 d, anterior [endopodal tube-pore arrowed]; F, left ca

np-2 arrowed]; E, anal somite and left ca

udal ramus, lateral; G, P3 enp-2 d.

ta on e

v. (@). A, P1, anterior [inner se

Fig. 24 Archesola hamondi gen. et sp. no

typhlops (Sars, 1908) comb. no

BASAL LAOPHONTID EVOLUTION

1-[1], 2-[8 + 1 pinnate], 3-[7], 4-[2], 5-[8 + 1 pinnate spine + 1

spinous process + (2 + ae)], 6-[1 + 3 processes], 7-[8 + acrothek].

Apical acrothek consisting of aesthetasc and 2 bare setae.

P3 endopod (Fig. 24C, G) 2-segmented; enp-1 as in 2; enp-2 with

3 inner setae (proximal one being distinctly shorter than in 9), 2 long

apical setae and short pinnate outer spine (fused basally to segment);

tube-pore present near outer apical seta.

PS (Fig. 24D) medially fused, positioned ventrolaterally.

Baseoendopod without endopodal lobe or armature; medial margin

with few setules and 2 tube-pores (longest arrowed); outer basal seta

arising from long, articulating, cylindrical setophore. Exopod free,

rectangular; with | long pinnate seta apically; inner margin proximal

seta and distal bipinnate spine; outer margin with 2 bare setae.

Sixth legs represented by well developed opercula, one articulat-

ing and closing off left or right genital aperture; each produced into

cylindrical process bearing 1 lateral and 1 apical seta.

REMARKS. Drzycimski (1969) corrected two major errors in Sars’

(1908) description. First, he pointed out the presence of the thin

inner seta on the proximal endopod segment of P1. Within the

Laophontidae this element is further only found in Archilaophonte

maxima. Secondly, Drzycimski remarked that the inner seta on the

baseoendopod of the male P5 is not well developed as in Sars’

illustration but greatly reduced. In reality, Drzycimski referred to the

short hyaline tube-pore located closely to the exopod whereas in

Sars’ (1908) illustration it was the longer medial tube-pore (arrowed

in Fig. 24D) which was misinterpreted as a genuine seta. One

character that has traditionally been used to differentiate A. typhlops

from A. longiremis is the setation of the female P5 exopod. This

distinction is invalid since it is based on the erroneously reported

absence of the proximal surface seta in Sars’ description of A.

typhlops. The same error also served to distinguish E. typhlops

pontoica from the type population (Por, 1959, 1964a).

Reliable records of A. typhlops include Flekkeré (Sars, 1908),

Bergen ( Drzycimski, 1969) and Frierfjord/Langesundfjord (this

account) in Norway, Gullmar Fjord (Lang, 1948) and the Isle of

Bonden (Por, 1964a) in Sweden, and Norfolk in England (this

account). The Scottish records from the River Ythan (Aberdeen-

shire) by Hockin & Ollason (1981) and Hockin (1982a—b, 1984) and

that from Newbiggin (Northumberland) by Moore (1973) may be

based on A. longiremis.

Archesola longiremis (T. Scott, 1905) comb. nov.

Laophonte longiremis T. Scott, 1905

Esola longiremis (T. Scott, 1905) Lang (1948)

TYPE LOCALITY. Granton, Firth of Forth, Scotland; old quarry

opening to the sea (T. Scott, 1905, 1906).

TYPE MATERIAL. _T. Scott (1905) recorded an unspecified number

of females; this material has not been deposited in any of the British

museums (London, Newcastle-upon-Tyne, Edinburgh) and is there-

fore almost certainly lost.

REMARKS. This species is very close to A. typhlops and can be

differentiated primarily by the shorter caudal rami (only twice as

long as wide) and the smaller body size (0.6 mm). Lang (1948)

pointed out that T. Scott’s (1905) drawing of the P5 showed an

aberrant setation on the endopodal lobe (total of 7 setae: 3 inner, 3

apical, 1 outer). The short apical seta is almost certainly the equiva-

lent of the long tube-pore found in this position in A. typhlops,

however, the presence of the supernumerary outer seta is more

difficult to explain since no laophontoidean is known to display

more than 5 elements on the endopodal lobe of the female P5 (Huys,

1990a; Huys & Lee, 1999). We suspect that this seta is the result of

an observational error.

The species has never been figured again since T. Scott (1905) nor

has the male been discovered. Wells (1961) illustrated some features

of a male specimen from St. Martin’s (Isles of Scilly) which he

attributed to E. longiremis. The P5 shows only 3 setae on the exopod

and the endopodal armature is represented by 2 fine setae (one of

which likely to be a tube-pore). The P6 bears 2 strong setae but is not

drawn out into a cylindrical process as in other species of the genus.

These characters in conjunction with his statement that the male

antennule is subchirocerate and the endopod 3-segmented clearly

exclude the possibility that Wells was dealing with a species of

Archesola or any other esolinid genus. Wells (1963) also recorded

the species from Exmouth (Devon) but this record remains uncon-

firmed.

The genus Archesola consists of a complex of closely related

species which can be differentiated primarily by morphometric

characters, such as caudal ramus length and Pl exopod: endopod

ratio, and various setal length differences on the PS. Coull’s (1971)

identification of E. longiremis from North Carolina suggests an

amphi-Atlantic distribution for the genus Archesola, however, in

view of the relatively subtle differences between congeners, the

specific identity of his record remains to be confirmed.

Archesola hamondi sp. nov.

TYPE LOCALITY. 53°10.34'N 00°56.34'E; depth 12-13 m; fine

sand with high silt and shell gravel content.

TYPE MATERIAL. This species is only known from the holotype 2

(leg. R. Hamond; 06 May 1992) which unfortunately was acciden-

tally destroyed before the description could be completed. The brief

description below provides sufficient information to warrant the

proposal of a new species.

ETYMOLOGY. This patronym is dedicated to Dr Richard Hamond

who collected the holotype, in recognition of his significant contri-

butions to laophontid systematics.

DESCRIPTION.

FEMALE. Body length from anterior margin of rostrum to posterior

margin of caudal rami 600 pm.

Body (Fig. 21A) dorsoventrally depressed and much wider than

in A. typhlops; covered with dense pattern of minute surface lamel-

lae dorsally and laterally. Cephalothorax bell-shaped, distinctly

widening towards posterior margin; lateral and hind margins fringed

with long setules; without paired cup-shaped pores. Setular fringes

also present laterally on pedigerous somites and urosomites, some-

times forming tufts locally. Posterior margin of urosomites without

distinct ornamentation dorsally except for penultimate somite bear-

ing transverse row of fine spinules.

Genital double-somite (Fig. 21A) wide and dorsoventrally flat-

tened; original segmentation marked by bilateral constriction only;

without cup-shaped pores in anterior half; lateral lobes without

backwardly directed strong spinules. Genital field as in A. typhlops.

Anal somite (Fig. 24E) with distinct setular fringe around anal

opening; anal operculum completely bare; posterolateral margins

with fine spinules.

Caudal rami (Fig. 24E) cylindrical and slightly swollen in anterior

half; distinctly wider than inA. typhlops; about 3 times as long as wide.

Seta II shorter and seta II posteriorly displaced compared to A.

typhlops; setalV reduced, lacking fracture planes, shorter than caudal

ramus; seta V well developed, pinnate, without fracture planes; setae

VI-VII naked. Large vent-pore present at outer subdistal corner.

Rostrum (Fig. 21A) longer than in A. typhlops; trapezoid with

straight anterior and concave lateral margins; delimited at base by

transverse surface suture; with paired sensillae anteriorly and me-

dian tube-pore ventrally.

Antennules to maxillipeds as in A. typhlops.

P1 (Fig. 24A) as in A. typhlops except for (a) basal pedestal

bearing endopod wider, (b) inner seta on enp-1 inserting more

distally, and (c) enp-1 about twice the length of exopod (distinctly

shorter in type species). P2—P4 as in A. typhlops.

P5 (Fig. 23F). Endopodal lobe well developed, well extending

beyond insertion sites of proximal outer setae of exopod; with

distinctly stepped inner margin bearing 2 strong spines (more closely

set than in A. typhlops) and 1 bare distal seta (not extending beyond

apex of exopod); with 2 apical setae, outer one about 1.5 times

length of inner one; tube-pores present near apical setae and proxi-

mal to innermost spine; outer basal seta inserting on cylindrical

articulating setophore. Exopod elongate but distinctly shorter than

in A. typhlops, produced apically into short tubular extension bear-

ing 1 bare seta; inner margin with 1, outer margin with 4 pinnate

setae. Both baseoendopod and exopod with elaborate ornamentation

pattern as figured.

MALE. Unknown.

REMARKS. Differentiation of A. typhlops and A. hamondi is best

achieved by comparison of the general body shape, caudal ramus

outline and armature pattern, and 2 PS morphology and morpho-

metry

Esola sp. sensu Chislenko (1967)

Chislenko illustrated a male which he obtained in Laminaria

saccharina washings from the White Sea and identified as Esola sp.

His drawings of the caudal ramus, P3 endopod and P1 leave little

doubt that this species belongs to Archesola and is obviously close

to A. typhlops and A. longiremis. The caudal ramus L:W ratio

appears to be intermediate between the latter two species and the P1

endopod and exopod have slightly different proportions. Chislenko’s

male (0.35 mm) is smaller than those of A. typhlops recorded by

Drzycimski (1969) from the Bergen area (0.45 mm) and our single

male from West Runton (0.475 mm). The antennary exopod bearing

the atypical number of 5 setae is obviously based on an aberrant

specimen. His illustration of the P3 endopod lacks the proximal

inner seta on the distal segment, its location being indicated by the

distinct step in the inner margin. Finally, the small inner seta

illustrated on the P5 baseoendopod is probably a tube-pore. The

White Sea material identified by Brotskaya (1961) as E. longiremis

is likely to be conspecific with this species, the true identity of which

is as yet uncertain. Consequently, Chislenko’s species is tentatively

ranked specues inquirenda in Archesola.

Archesola typhlops pontoica (Por, 1959) comb. nov.

Esola typhlops pontoica Por, 1959

TYPE LOCALITY. Black Sea coast, Rumania.

TYPE MATERIAL. Dr Ileana Negoescu (Museum ‘Grigore Antipa’,

Bucharest) informed us that the syntypes no longer exist.

REMARKS. Por (1959, 19645) established this subspecies for 3 22

found at 61-69 m depth off the Rumanian coast, however it is

doubtful whether his material deserves such status. The author

discriminated the Black Sea population on the basis of the presence

of 6 setae on the 2P5 exopod (Sars (1908) erroneously figured only

5), the slightly shorter caudal rami and the incompletely 3-seg-

mented P1 exopod (a feature displayed in only 1 specimen!). The

R. HUYS AND W. LEE

most significant difference, not mentioned by Por, is found in the

proportional lengths of the distal antennulary segments (segments 6

and 7 being of equal length). Examination of new material is

necessary to resolve the identity of the Rumanian population; E.

typhlops pontoica is considered here as subspecies inquirenda.

Genus Corbulaseta gen. nov.

The diagnosis below is based on Vervoort’s (1964) redescription of

E. bulligera and personal observations of Wells’ (1970) material

from Great Britain Rock, Isles of Scilly, and additional specimens

collected from the Belgian North Sea coast by the senior author.

DIAGNOSIS. Laophontidae. Body cylindrical; posterolateral cor-

ners of 2 genital double-somite and second abdominal somite

laterally and backwardly produced. Integument of cephalothorax

and body somites with dense pattern of spinules and setules. Ros-

trum large, partly delimited at base by incomplete surface furrow.

Cephalothorax with one pair of large, anterodorsal cup-shaped

pores; such pores absent on genital (double-)somite and caudal rami.

Anal operculum spinulose. Caudal rami rectangular, short; not

sexually dimorphic.

Sexual dimorphism in antennule, P3 endopod, P5, P6 and in

genital segmentation.

Antennules slender; 6-segmented in 9, subchirocer and 7-seg-

mented in 6; segment 1 with 1—2 minute processes along posterior

margin; with aesthetasc on segment 4 () or 5 (d) and as part of

apical acrothek on distal segment; segment 5 ¢ swollen, without

anterior outgrowth; proximal aesthetasc fused basally to 2 setae.

Antenna with 4 setae on exopod; allobasis with abexopodal seta.

Labrum with distal spinular ornamentation. Mandible with 1-seg-

mented palp; exopod and endopod represented by small tubercles

bearing 1 and 3 setae, respectively; basis represented by 2 apical

setae. Maxillule with minute, defined exopod. Maxilla with 3 endites

on syncoxa; endopod represented by 3 setae. Maxilliped slender;

syncoxa with 2 setae; entire palmar margin with long setules;

endopodal claw elongate.

P1 with 2-segmented exopod bearing 5 setae on exp-2 and

elongate endopod; enp-1 without inner seta, enp-2 with minute seta

and long, slender claw. P2—P4 with 3-segmented exopods and 2-

segmented endopods. P2 basis with short outer spine. Outer spine of

P2—P4 enp-2 setiform and very long in P3-P4. P4 endopod modified

in both sexes; distal inner seta proximally dilated, bearing enlarged

spinules which enclose long secretory tube-pore arising from seg-

ment. P3 endopod d 3-segmented; enp-2 with inner seta and short

outer spinous apophysis. Armature formula as follows:

Exopod Endopod

P2 0.1.123 Ail

P3 0.1.223 e372 [d: 1.1.220]

P4 0.1.223 0.221

*: or 1.220 in Vervoort’s (1962) 2 specimen of E. bulligera from New Caledonia.

P5 2 with separate rami; exopod elongate, with 6 setae/spines;

baseoendopod slightly developed, with 4 setae/spines. PS d without

endopodal lobe; exopod short, with 1 inner, 2 apical and 2 outer

setae/spines.

P6 forming opercula closing off paired genital apertures; with 2

small setae at outer corner. P6 d asymmetrical; membranous flaps

with 1 long and 1 minute seta.

BASAL LAOPHONTID EVOLUTION

TYPE AND ONLY SPECIES. Laophonte bulligera Farran, 1913 =

Corbulaseta bulligera (Farran, 1913) comb. nov.

ETYMOLOGY. The generic name is derived from the Latin corbula

(little basket) and seta (bristle) and refers to the modified distal inner

seta of P4 enp-2, the proximal setules of which form a trapping

basket typically enclosing a secrete bolus.

Corbulaseta bulligera (Farran, 1913) comb. nov.

Laophonte bulligera Farran, 1913

Esola bulligera (Farran, 1913) Lang (1948)

Laophonte rosei Monard, 1926

Laophonte Rosei Monard, 1926: Monard (1928)

Esola rosei (Monard, 1928) Lang (1948)

TYPE LOCALITY. Blacksod Bay, Co. Mayo (Ireland); 1.8—5.4 m

depth

MATERIAL EXAMINED. Farran’s (1913) type material is lost (J.M.C.

Holmes, pers. comm).

(a) Isles of Scilly, Great Britain Rock: 1 @ in alcohol (BMNH

1967.10.31.76); coll. University of London Sub-Aqua Expedition

1966; det. J.B.J. Wells;

(b) Belgium, North Sea coast, 51°30"N 2°00"E: 19, 1 3; 08 April

1986, depth 14.1 m, sandy substrate; leg. R. Huys.

ADDITIONAL OBSERVATIONS.

FEMALE. Body length from anterior margin of rostrum to posterior

margin of caudal rami 570-590 um. Body (Fig. 21B) cylindrical,

slightly depressed; covered by irregular pattern of minute surface

spinules. Cephalothorax widest, subrectangular; with pair of large

cup-shaped pores anterodorsally (Fig. 25C); ventral pores absent;

posterior margin and anterior half of ventral margin with setular

fringe; posterior half of ventral margin bordered by tiny spinules;

posterolateral corner produced forming distinctive lobate extension

(Fig. 25C). Prosome gradually tapering posteriorly; all somites with

dorsal transverse spinular row and setular fringe around hind mar-

gin.

Genital double-somite dorsoventrally depressed; original seg-

mentation marked by bilateral constriction and dorsal transverse

spinular row set on surface ridge; ventral surface without conspicu-

ous ornamentation; cup-shaped pores absent. Genital aperture closed

off by sixth legs bearing | naked seta. Posterolateral corners of

second abdominal somite backwardly produced; remaining

urosomites distinctly narrower. Ventral posterior margin of penulti-

mate somite with medial fringe of fine setules flanked by strong

spinules (decreasing in length ventrolaterally). All urosomites with

spinules around dorsal posterior margin. Anal operculum spinulose.

Caudal rami short, slightly longer than wide; all setae arranged in

posterior quarter; setae IV and V well developed, pinnate, with

fracture planes; no conspicuous pores present.

Rostrum (Fig. 21B) trapezoid, delimited at base by incomplete

surface suture; with 2 long sensilla apically and tube-pore ventrally.

Antennule 6-segmented; posterior margin of segment | with

slight bulbous swelling but no real spinous processes; aesthetasc on

segment 4 fused basally to 2 long setae; armature formula: 1-[1], 2-

[7 + 1 pinnate], 3-[6], 4-[(2 + ae)], 5-[1], 6-[9 + acrothek]; acrothek

consisting of aesthetasc and 2 naked setae. Antennary exopod with

strong pinnate outer apical spine and 3 pinnate setae. Labrum with

Sparse ornamentation resembling condition in A. hirsuta. Mandibu-

lar palp 1-segmented, with ancestral setation, i.e. 2 basal, 1 exopodal

and 3 endopodal setae. Maxillule as in E. bulbifera, with endopod

represented by 2 setae. Maxilla as in E. bulbifera. Maxilliped with 2

setae on syncoxa; palmar margin with long fine spinules; endopodal

claw slender and longer than basis, with 1 accessory seta.

P1 as in Farran’s (1913) description except for outer spine of exp-

1 being longer and pinnate and proximal and middle outer spines of

exp-2 distinctly shorter. P2 basis with bipinnate outer spine, P3—P4

bases with smooth outer seta. Outer spine of P3—P4 enp-2 very long

and setiform (Fig. 25D).

P4 (Fig. 25D) with 2-segmented endopod; enp-1 short, without

inner seta; enp-2 (Fig. 25E-F) highly distinctive: distal inner seta

with dilated base bearing comb of long curved setules on both

anterior and posterior outer margins; this ornamentation forming

trapping basket enclosing large secrete bolus produced by long

anterior surface tube-pore located near distal margin of enp-2.

PS as in original description.

MALE. Body length from anterior margin of rostrum to posterior

margin of caudal rami 530 um. Body more slender than in °; none of

urosomites with backwardly produced posterolateral corners. Ven-

tral posterior margin of postgenital (except anal) somites with

median fringe of fine setules flanked by strong spinules.

Antennule subchirocerate; 7-segmented with geniculation between

segments 5 and 6. Segment | without distinct processes, segment 5

without anterior outgrowth.

P3 endopod 3-segmented; very similar to that of EF. bulbifera (Fig.

4D).

PS without endopodal lobe; medial margin frimged with long

spinules and | tube-pore; outer basal seta arising from short setophore.

Exopod elongate, about 3.5 times as long as wide; with | seta and 1

spine along inner margin, apex with | long bipinnate seta, outer

margin with 2 bipinnate spines.

P6 asymmetrical; each opercular flap with cylindrical extension

at outer corner bearing long outer seta and minute inner seta.

REMARKS. Nicholls (19416) pointed out that Laophonte rosei,

described from Banyuls (Monard, 1926) may well be a junior

synonym of L. bulligera since the difference between them appears

to be based on two doubtful characters. The ‘sensory organ’ illustrated

on the P4 endopod of L. bulligera by Farran (1913) was not

described for L. rosei by Monard (1926) although the latter did

illustrate the adjoining modified seta (see also Monard (1928)).

Secondly, the different number of setae on the P5 endopodal lobe is

based on Monard’s failure to observe the seta near the base of the

baseoendopod, a portion of which appears to have been lost in L.

rosei. Lang (1948) also expressed strong reservations about the

distinctiveness of L. rosei but like Nicholls (1941b) and Vervoort

(1967) nevertheless maintained it as a valid species. We can see no

justification for this distinction and formally relegate E. rosei to a

junior subjective synonym of E. bulligera. Pesta (1959) published

an incomplete description of the male (as E. rosei) but did not

mention the transformed P4 endopod. The discrepancy found in the

length of the P1 endopod casts some doubt on his identification.

Pending the re-examination of mediterranean material, the known

records suggest an almost continuous boreo-mediterranean distri-

bution pattern with records from Ireland (Farran, 1913, 1915), Isles

of Scilly (Wells, 1970), Belgian coast (unpubl.), Banyuls-sur-Mer

(Monard, 1926, 1928) and possibly Naples (Pesta, 1959). Por &

Marcus (1972) recorded the species also in the Great Bitter Lake and

off Port Taufig in the southern part of the Suez Canal and considered

the species an Atlantic (anti-Lessepsian) immigrant. There is no

morphological evidence supporting Alheit & Scheibel’s (1982)

record from Harrington Sound in Bermuda.

The isolated record from New Caledonia by Vervoort (1962) is

difficult to interpret, particularly because his single female speci-

men deviates from European E. bulligera in the absence of the outer

spine on P2 enp-2. Vervoort (1962) did not remark on this character

ee R. HUYS AND W.LEE —

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LES

Fig. 25 Archesola typhlops (Sars, 1908) comb. nov. A, Mandibular palp; B, maxillule, anterior. Corbulaseta bulligera (Farran, 1913) comb. nov. C,

cephalothorax, lateral; D, P4 2, anterior; E, P4 endopod 2, anterior; F, P4 enp-2 °, medial [contours of secrete bolus stippled in E-F]. Laophonte parvula

Sars, 1908. G, P5 2, anterior [anteriorly displaced outer seta arrowed].

BASAL LAOPHONTID EVOLUTION

presumably because of the lack of a base for comparison in Farran’s

(1913) description and illustrations which omitted P2 and P3. The

problem is exacerbated by the aberrant left-right asymmetry (1.220

vs 1.320) displayed on the P2 endopods. It is unclear whether the

reduced setal formula is real and therefore indicative for the pres-

ence of a second species in the western Pacific. There is very little

additional evidence pointing in this direction except for the different

cephalothorax shape (in lateral aspect: compare Fig. 25C) and some

morphometric discrepancies in the caudal rami, which appear to be

longer, and in the exopods of P2—P4, which are more abbreviated.

E. bulligera cannot be retained in the genus Esola because of the

absence of (1) distinct spinous processes on the first antennulary

segment, (2) cup-shaped integumental pores on the genital (double-)

somite and caudal rami, (3) characteristic labral ornamentation and

(4) caudal ramus sexual dimorphism. It is reminiscent of Bathyesola

compacta (see below) in the presence of only one pair of cup-shapes

pores on the cephalothorax but differs from it in the reduced

armature on the 2 P5 exopod and the transformed P4 endopod which

is the most significant autapomorphy of E. bulligera, justifying its

placement in a new genus Corbulaseta.

Genus Bathyesola gen. nov.

DIAGNOSIS (based on 2 only). Laophontidae. Body cylindrical;

posterolateral corners of 2 genital double-somite and second ab-

dominal somite laterally and backwardly produced. Integument of

cephalothorax and body somites with dense pattern of spinules and

setules. Rostrum large, partly delimited at base. Anterolateral pair of

small integumental cup-shaped pores present on cephalothorax.

Caudal rami not modified in 9, cylindrical and elongate.

Sexual dimorphism presumably in antennule, P3 endopod, P5,

P6, and genital segmentation.

Antennules slender; 7-segmented in 2; segment | without spinous

processes along posterior margin; with aesthetasc on segment 4

(fused basally to 2 setae) and as part of apical acrothek on segment

7. Antenna with 4 setae on exopod; allobasis with abexopodal seta.

Labrum without overlapping scales distally but with pattern of

spinules anteriorly. Mandible with 2-segmented palp; endopod free,

with 3 setae; exopod represented by single seta; basis represented by

2 setae. Maxillule with defined exopod. Maxilla with 3 endites on

syncoxa; endopod represented by 3 setae. Maxilliped robust; syncoxa

with 2 setae; entire palmar margin with spinules; endopodal claw

relatively stout.

P1 with large 3-segmented exopod bearing 4 setae on exp-3 and

relatively short endopod; enp-1 without inner seta, enp-2 with

minute seta and short, curved claw. P2—P4 with 3-segmented exopods

and 2-segmented endopods. P2 basis with moderately long outer

spine. Inner seta of P2—P4 exp-2 reduced. Outer spine of P2—P4 enp-

2 setiform, short in P2—P3, long in P4. Armature formula as follows:

Exopod Endopod

P2 0.1.123 e222

P3 0.1.123 0.321 [d presumably

0.1.220]

P4 0.1.123 0.221

P5 2 with separate rami; exopod relatively short, with 6 setae/

spines; baseoendopod well developed, with 5 setae/spines, apical

setae reduced; outer basal seta on short setophore.

P6 Yforming opercula closing off paired genital apertures; with 2

small setae.

TYPE AND ONLY SPECIES. Bathyesola compacta gen. et sp. nov.

ETYMOLOGY. The generic name refers to the bathyal distribution

of the type species.

Bathyesola compacta gen. et sp. nov.

TYPE LOCALITY. 18°50'S, 173°29'W, ‘White Lady’ site on North

Fiji Ridge, west of Fiji; 2765 m depth. Accompanying harpacticoid

fauna: several 92 and dod of Xylora bathyalis Hicks, 1988

(Thalestridae: Donsiellinae).

TYPE MATERIAL. Holotype 2 dissected on 6 slides, deposited in

Muséum National d’ Histoire Naturelle, Paris under MNHNP Cop-

1869; collected during STARMER II expedition, station 14 (Kaiyo

87), dive 19; 14 July 1989; leg. L. Laubier.

ETYMOLOGY. The species name alludes to the compact P1, dis-

playing a short and robust endopod.

DESCRIPTION.

FEMALE. Body length from anterior margin of rostrum to posterior

margin of caudal rami 360 um. Maximum width (105 um) measured

at posterior margin of cephalothorax.

Body (Fig. 26A—B) cylindrical, slightly dorsoventrally depressed,

covered with dense pattern of minute spinules dorsally and laterally.

Cephalothorax slightly wider than free somites, posterolateral angles

backwardly produced forming small lobate extension (Fig. 26B);

with pair of small lateral cup-shaped pores. Posterior margin of

cephalothorax and all body somites with row of long setules dorsally

and laterally. Pleurotergite of P5-bearing somite almost as wide as

anterior somites.

Genital double-somite wide and dorsoventrally flattened; with

lateral, backwardly produced extensions in posterior (=abdominal)

half; original segmentation marked by bilateral constriction and

spinule row arising from transverse surface ridge dorsally and

laterally; posterior half with backwardly directed lobate extensions

bearing spinular tuft; ventral surface without spinular ornamenta-

tion; genital field located near anterior margin. Sixth legs forming

well developed opercula closing off paired genital apertures; each

with 2 small setae.

First postgenital somite with backwardly produced lateral angles,

bearing spinular tuft; without ventral ornamentation. Penultimate

and anal somites distinctly narrower; ventral posterior border with

long spinules. Anal somite with spinulose anal operculum.

Caudal rami (Fig. 26A—B) widely separated; about 4 times as long

as average width; maximum width measured at base; dorsal surface

with minute spinules; seta I small, setae II-III well developed, naked

and closely set; setae IV (naked) and V (pinnate) with fracture

planes, seta V 2.8 times as long as seta IV; setae VI-VII naked.

Rostrum (Figs 26A) large, blunt anteriorly; delimited at base by

transverse surface suture; with paired sensillae anteriorly and me-

dian tube-pore dorsally.

Antennule (Fig. 26A—B) relatively short, distinctly 7-segmented,

without spinous processes on segments 1—2. Segment 1| with large

spinular patch around anterior margin. Armature formula: 1-[1], 2-

[4 + 4 pinnate], 3-[6], 4-[1 + (1 + ae)], 5-[1], 6-[2], 7-[5 + 1 pinnate

+ acrothek]. Acrothek consisting of aesthetasc and 2 naked setae; set

on apical pedestal.

Antenna (Fig. 27A). Coxa with spinules on both inner and outer

margins. Exopod short, bearing 2 lateral and 2 apical pinnate

elements, and a longitudinal row of fine spinules along outer margin.

Allobasis with pinnate abexopodal seta. Endopod with lateral arma-

ture consisting of 2 spines and 1 minute seta; distal armature

consisting of 2 naked spines and 3 geniculate setae (outermost fused

basally to minute seta).

Fig. 26

R. HUYS AND W. LEE

im im Tan Ne

i Tit 1 ni ty Tree fn \f stu eA

ail i nmin i mtn i Me

= Gee ecr

— oi

TTA ee

Hh)

AHN)

( a op Tan RTT aes

i rari om

Gh oe

ene itl ar ea Fj aN iN

ee uy, NN

pe =

— WAY, Lt

100 yu

Bathyesola compacta gen. et sp. nov. (@). A, Habitus, dorsal; B, habitus, lateral.

- BASAL LAOPHONTID EVOLUTION

ity,

_ Fig.27 Bathyesola compacta gen. et sp. nov. (2). A, Antenna; B, mandible; C, maxillule, anterior; D, maxilla; E, maxilliped; F, P1, anterior.

‘d

R. HUYS AND W. LEE

pee

phe hb =

heeds. pit WA wth pb eee —~

IF bf Y o li A Se Na

/ ac eX

es, Se

Se aes

e Yipee

UTE

oe Srna aaa

aS SS Se =

NG 2 :

Lh SES en ie 8

RNS

50u

Fig. 28 Bathyesola compacta gen. et sp. nov. (@). A, P2, anterior; B, P3, anterior; C, P4, anterior; D, P5, anterior. Paralaophonte pilosoma Vervoott,

1964. E, P3 endopod 4, anterior.

BASAL LAOPHONTID EVOLUTION

Labrum with spinular patches on anterior face but no overlapping

scales.

Mandible (Fig. 27B) with short gnathobase and small 2-seg-

mented palp representing free endopod and fused basis and exopod;

endopod a minute segment with 3 pinnate setae; basal armature

represented by 2 lateral pinnate setae, exopod represented by single

seta.

Paragnaths highly ornate lobes as in E. bulbifera.

Maxillule (Fig. 27C) with elongate arthrite bearing | seta on

anterior surface and 9 elements around distal margin. Coxal endite

with | spine and | seta, basal endite with 1 spine and 2 setae. Exopod

a short segment with 2 distal setae; endopod incorporated into basis,

represented by 2 setae.

Maxilla (Fig. 27D). Syncoxa with very long setules around outer

margin and few additional spinule rows as figured; with 3 endites;

praecoxal endite small, with | naked seta; middle endite drawn out

into spine, with 2 setae; distal endite with 3 elements. Allobasis

produced into strong curved claw; accessory armature consisting of

2 setae. Endopod incorporated into allobasis, represented by 3 bare

setae.

Maxilliped (Fig.27E) compact, basis and endopodal claw not

particularly elongate. Syncoxa with 2 pinnate setae. Basis with

spinular ornamentation as figured; spinules present along entire

palmar margin. Endopod represented by stout, minutely pinnate

claw bearing | accessory seta and tube-pore at base.

P1 (Fig. 27F) with dense ornamentation on praecoxa, coxa and

basis. Basis with pinnate seta on anterior surface and along outer

margin. Exopod large, 3-segmented; exp-1 with pinnate outer seta;

exp-3 with 2 unipinnate outer spines and 2 geniculate setae apically.

Endopod robust and relatively short; enp-1 about 2.2 times as long

as basis, with long setules along inner margin and fine spinules along

outer margin; enp-2 about as long as wide, with short unipinnate

claw and small accessory seta.

P2—P4 (Figs 28A—C) with 3-segmented exopods and 2-segmented

endopods. P2 basis with long, bipinnate outer spine; P3—P4 bases

with bare outer seta. P2 enp-1 with pinnate inner seta, P3—P4 enp-1

unarmed. Inner seta of P2 exp-2 reduced. Outer spine of P2—P4 enp-

2 setiform, very long in P4. Pore present near distal outer corner of

P3—P4 enp-2. Armature formula as for genus.

P5 (Fig. 28D). Endopodal lobe well developed, extending to

halfway down the exopod; with 2 reduced bare setae apically, and 2

long widely separated setae along inner margin; pores present near

articulation with exopod, at base of apical setae and proximal to

innermost seta. Exopod relatively short, produced apically into short

tubular extension bearing | bare seta; inner margin with 1, outer

margin with 4 pinnate setae; inner seta slightly longer than apical

one. Both baseoendopod and exopod with spinulation as figured.

MALE. Unknown.

REMARKS. The discovery of B. compacta at 2765 m depth at the

North Fiji Ridge represents the deepest record thus far for the family

Laophontidae (Lee & Huys, 1999). It displays a mozaic of primitive

(7-segmented 2 antennule; 3-segmented P1 exopod; 2 P5 endopodal

lobe with 5 setae/spines) and advanced characters (P3—P4 enp-1

without inner seta; P3—P4 exp-3 with | inner seta) which serves to

distinguish the species from other esolinids.

Status of Esola Spelaea (Chappuis, 1938)

Lang (1944, 1948) placed Laophonte spelaea in the genus Esola

without giving any explicit reasons. From his generic diagnosis and

the phylogenetic scheme presented on p. 1450 (Lang, 1948), one can

infer that his course of action was based solely on the presence of an

outer spine on the distal endopod segment of P2. Although this

character was diagnostic for Esola in Lang’s sense it is clearly a

symplesiomorphy shared by all genera in the Archilaophonte-Esola

lineage (with the exception of Mourephonte) and consequently of no

value in inferring relationships. Lang (1944, 1948) subdivided

Esola into two species groups, diagnosed by the number of setae on

the male P5 endopodal lobe and the armature of the P3 in both sexes.

His spelaea-group included only E. spelaea and has until now

remained monotypic. It differed from the Jongicauda-group in the

presence of 2 setae (rather than | or 0) on the d P5 endopodal lobe

and a reduced armature on the P3 exopod (exp-3 with only 2 outer

spines) and endopod (enp-2 @ with only 2 inner setae; endopod 3

without inner seta on enp-2 and with only 3 setae on enp-3).

Chappuis’ (1938) description is very brief and provides illustra-

tions of the male P2—P5 only. Unfortunately the author did not give

any information about the position of the setae on the female P5

which could have provided the justification for including L. spelaea

in the Archilaophonte-Esola lineage since in all of its members (1)

the proximal seta of the endopodal lobe is medially displaced and (2)

the insertion sites of the 2 proximal setae of the exopod are superim-

posed. Chappuis’ statement that there are 4 setae on the baseoendopod

and 5 or 6 setae on the exopod can be interpreted in the light of this

generalized pattern. His reservation about the correct number of

exopodal setae might indicate the close or overlapping position of

some of these elements. Secondly, due to its strong medial displace-

ment the proximal endopodal seta has frequently been overlooked or

lost during dissection (Thompson & A. Scott, 1903; Norman, 1911;

Monard, 1926, 1928; Noodt, 1955), leaving open the possibility of

a similar observational error made in Chappuis’ (1928) description.

The actual number of endopodal setae on the female P5 of L. spelaea

could therefore be five rather than four. Chappuis’(1928) armature

formula of P2 exp-3 tabulated as 222 (i.e. with 2 outer spines) is

unlikely to be correct when both P2 and P4 reportedly have 3 outer

spines on exp-3. No laophontid described thus far displays a [3-2-3]

outer spine pattern for P2—P4 and hence we suspect 123 (as in B.

compacta) to be the correct formula for P2 exp-3.

Chappuis (1938) described L. spelaea from three caves in Apulia,

southern Italy (Abisso and La Zinzulusa near Castro, Grotta dei

Diavoli near Badisco) and regarded it as a marine relict. The caves

exhibit a tidal regime but the salinity approaches that of freshwater

(‘. . . das Wasser schmeckt aber fast stiss’) which appears to be

confirmed by the presence of the stygobiont mysids Spelaeomysis

bottazzii Caroli and Stygiomysis hydruntina Caroli and the

palaemonid Typhlocaris salentina Caroli, all of which are endemic

to coastal caves and phreatic waters in the Apulia region. Both Pesce

(1985) and Rouch (1986) consider the species as a descendant from

a marine ancestral stock which successfully colonized subterranean

freshwater habitats via littoral karstic systems, possibly during

regression periods in the Tertiary (“Regression Model Evolution’ ).

Laophonte spelaea cannot be accommodated in any of the exist-

ing laophontid genera. It appears to be related to Bathyesola in

certain aspects (see above) but differs from it in the presence of an

inner seta on P3—P4 enp-1, only 2 inner setae on P3 enp-2 and more

primitive setal formula on the P4 exopod. In view of the strong

ecological divergence between B. compacta and L. spelaea we

prefer to establish a new genus for the latter. The male P3 endopod

in Troglophonte gen. nov. does not accord with the pattern found in

the other esolinid genera. The absence of an inner seta on the middle

segment could be related to the reduced 1.221 pattern in the female

but might also indicate a relationship with a large group of other

laophontid genera which typically lose the proximal inner seta

during male P3 ontogeny.

96 R. HUYS AND W. LEE

Fig. 29 SEM micrographs. Applanola hirsuta (Thompson & A. Scott, 1903) comb. nov. (2). A, Lateral margin of genital double-somite, ventrolateral; B,

lateral cup-shaped pore on genital double-somite [pore exit arrowed]; C, setular extensions bordering dorsal margin of cup-shaped pore; D, labrum and

mandibular gnathobases. [Scale bars: 2 um (C), 10 um (B, D), 20 um (A)].

BASAL LAOPHONTID EVOLUTION

Fig. 30 SEM micrographs. Applanola hirsuta (Thompson & A. Scott, 1903) comb. nov. (2). A, Cephalothorax and rostrum, frontal [anterodorsal pore

arrowed]; B, cephalothorax, ventral [anteroventral pore arrowed]; C, anal opening and caudal ramus, ventral [ventral pore arrowed]; D, right caudal

ramus, ventral, showing cup-shaped pore. [Scale bars: 6 um (D), 15 um (C), 20 um (B), 60 um (A)].

Genus Troglophonte gen. nov.

DIAGNOSIS. Laophontidae. Body shape unknown but somites not

well demarcated. Rostrum short, presumably fused at base.

Integumental cup-shaped pores unconfirmed. Anal operculum

spinulose. Caudal rami short, squarish.

Sexual dimorphism in antennule, P3 endopod, P5, P6 and in

genital segmentation.

Antennules 7-segmented in 9, segmentation unknown in d. An-

tenna with 4 setae on exopod; allobasis with abexopodal seta.

Mouthparts unknown. Maxilliped very slender.

P1 with 3-segmented exopod bearing 4 setae on exp-3 and slender

endopod; enp-1 without inner seta, enp-2 with minute seta and

slender, strong claw. P2-P4 with 3-segmented exopods and 2-seg

R. HUYS AND W. LEE

x009Z

dx3 emubey

4016

ubew

ole

dx3

Fig. 31 SEM micrographs. Esola bulbifera (Norman, 1911). A, Anal somite and caudal rami, dorsal. Applanola hirsuta (Thompson & A. Scott, 1903)

comb. noy. B, mandibular palp and inner face of cephalothorax showing vent-pore [arrowed]. Archesola typhlops (Sars, 1908) comb. nov. C, caudal

ramus, ventral; D, caudal ramus, area around setae I-III showing pores [arrowed]. [Scale bars: 5 um (D), 10 um (B), 20 um (A, C)].

BASAL LAOPHONTID EVOLUTION

mented endopods. P3 endopod d 3-segmented; enp-2 with outer

pinnate apophysis but without inner seta. Armature formula as

follows:

Exopod Endopod

Pp 0.1.123 1.221

PS 0.1.123 (or 0.1.2226) 1.221 [d: 1.0.120]

P4 0.1.223 22

P5 2 with separate rami; exopod elongate, with 5 or 6 setae/

spines; baseoendopod slightly developed, with 4 setae/spines. PS 3

with trapezoid endopodal lobe; exopod short, with 1 inner, 2 apical

and 2 outer setae/spines.

P6 unknown in both sexes.

TYPE AND ONLY SPECIES. Laophonte spelaea Chappuis, 1938 =

Troglophonte spelaea (Chappuis, 1938) comb. nov.

ETYMOLOGY. The generic name is derived from the Greek trogle,

meaning hole, and refers to the stygobiont life style of the type

species. Gender: feminine.

MATERIALEXAMINED. None. Chappuis’ (1938) material no longer

exists and the species has not been recorded again since its original

description.

PHYLOGENETIC ANALYSIS

Taxa and characters

The analysis was executed at species level in order to test the

monophyly of the genus Esola and its relationships to both

Mourephonte and Archilaophonte. Onychocamptus and the cornuta-

group of the genus Laophonte were also included as separate taxa in

the analysis on the basis of their ancestral P3 endopod sexual

dimorphism. The highly advanced genus Folioquinpes Fiers &

Rutledge, although having been positively identified as the

sistergroup of Onychocamptus (Lee & Huys, 1999), was excluded

from the analysis. The residual Laophontidae were replaced by their

hypothetical ancestor (Table 4: Other Laophontidae) which was

constructed by combining the most plesiomorphic state encountered

for each character.

Table 2. Laophontidae with 3 inner setae [3(1-2)(0-1) setation pattern] on P3 enp-2 9.

exp enp

Laophonte Group I OM23 1.220

Laophonte adduensis 0.1.122 1.220

Laophonte ciliata 0.1.122 1.220

Onychocamptus Group I 0.1.123 0.220

Onychocamptus besnardi O23 0.220

Onychocamptus anomalus 0.1.123 0.220

Onychocamptus taifensis 0.1.123 0.120

Onychocamptus krusensterni 0.1.123 0.220

Laophonte galapagoensis 0.1.123 0.220

Laophonte confusa 0.1.123 0.220

Laophonte Group II 0.1.123 0.220

Laophonte lignosa 0.1.123 0.220

Laophonte setosa 0.1.123 0.220

Laophonte elongata Oaln23 0.220

Laophonte Group III 0.1.123 0.220

Laophonte nordgaardi O23 0.120

Bathylaophonte spp. 0.1.123 0.220

Microlaophonte trisetosa 0.1.122 0.220

Pseudonychocamptus carthyi 0.1.123 0.220

Paralaophonte Group I 0.1.123 0.220

Paralaophonte panamensis 0.1.123 0.220

Paralaophonte Group II 0.1.123 0.220

Paralaophonte tenera 0.1.123 0.220

Paralaophonte innae 0.1.123 0.220

Paralaophonte aenigmaticum 0.1.123 0.220

Heterolaophonte campbelliensis 0.1.123 0.220

Heterolaophonte Group I 0.1.123 0.220

Heterolaophonte Group II 0.1.123 0.220

Heterolaophonte manifera 0.1.123 0.220

Heterolaophonte hamata ONE123 0.220

Heterolaophonte minuta 0.1.123 0.220

Paronychocamptus spp. 0.1.123 0.1—220

Asellopsis hispida 0.1.123 0.220

Asellopsis duboscqui 0.1.122 0.120

Folioquinpes chathamensis 0.1.123 0.220

P3 P4

exp enp 2 enp 3 exp enp

0.1.223 1.321 1.1.220 0.1.223 1.221

0.1.223 S21 1.1.220 0.1.223 1.221

0.1.222 1.321 1.1.220 0.1.222 1.221

0.1.123 0.321 0.1.220 0.1.123 0.111

0.1.123 0.321 0.1.220 0.1.022 0.111

0.1.123 0.321 0.1.220 0.1.122 0.111

0.1.123 0.321 0.1.220 0.1.123 0.111

0.1.123 0.321 0.1.220 0.1.122 0.111

0.1.223 0.321 0.0.220 0.1.223 1.121

0.1.223 0.321 0.0.220 0.1.223 1.120

0.1.223 0.321 0.0.220 0.1.223 0.221

0.1.223 0.321 0.0.220 0.1.223 0.121

0.1.223 0.321 0.0.220 0.1.223 0.111

0.1.223 0.321 0.0.220 0.1.123 0.111

0.1.123 0.321 0.0.220 0.1.123 0.111

0.1.123 0.311 0.0.210 0.0.023 0.111

0.1.223 0.321 0.0.220 0.1.223 0.221

0.1.222 0.321 0.220 0.1.222 0.221

0.1.223 1.321 0.220 0.1.223 ii

0.1.223 0.321 0.0.220 0.1.223 0.121

0.1.223 0.321 0.0.220 0.1.222 0.121

0.1.123 0.321 0.0.220 0.1.123 0.121

0.1.123 0.321 0.0.120 0.1.123 0.121

0.1.223 0.320 0.320 0.1.223 0.121

0.1.123 0.320 0.320 0.1.022 0.120

0.1.223 0.321 0.0.220 0.1.223 0.121

0.1.123 0.321 0.0.220 0.1.022 0.121

0.1.123 0.321 0.220 0.1.123 0.121

0.1.123 0.321 0.220 ONeI22 0.121

0.1.123 0.321 0.220 0.1.022 0.121

0.1.123 0.321 0.220 0.0.022 0.121

0.1.223 0.321 0.0.220 0.1.122 0.111

0.1.223 0.321 0.0.220 0.1.223 0.111

0.1.222 0.321 0.0.220 0.1.222 0.111

0.1.123 0.321 0.321 0.1.123 0.120

Laophonte: Group I = cornuta, expansa, plana; Group II = inornata, parvula, serrata; Group III = adamsiae, thoracica.

Onychocamptus: Group | = mohammed, bengalensis, vitiospinulosa

Paralaophonte: Group I = asellopsiformis, brevirostris, congenera, dieuzeidei, gurneyi, hyperborea, lacerdai, majae, meinerti, ormieresi, pacifica, pilosoma, royi; Group II =

karmensis, lunata, spitzbergensis, zimmeri.

Heterolaophonte: Group I = discophora, variabilis,; Group Il = murmanica, stromi, uncinata.

100

Characters used in the analysis are listed in Table 3. Apomorphic

character states are explained inside square brackets using the multistate

system. The scores for each character and taxon are compiled in

matrix format in Table 4. A question mark indicates missing data,

either because the appendage or structure is unknown in that species

(certain sexually dimorphic characters could not be scored because

only one sex is known) or because it was impossible to score the

character accurately due to incompleteness or the lack of detail in the

original descriptions. Esola typhlops pontoica, E. longicauda vat.

sensu Vervoort (1964) and the unnamed forms of E. longicauda

identified by Noodt (1955) and Wells & Rao (1987) were excluded

from the analysis because of their questionable status.

Huys & Boxshall’s (1991) study of ordinal copepod phylogeny

demonstrated that oligomerization was the dominant trend of evolu-

tionary transformation within the Copepoda. Armature counts used

in this analysis were scored according to this overall polarisation

mode. Most characters in Table 3 are self-explanatory but additional

notes are provided for the following:

Integumental pores (characters I-4)

The conspicuous cup-shaped integumental pores on the

cephalothorax and genital (double-)somite have remained unnoticed

Table 3. Characters used in phylogenetic analysis. Apomorphic character

states are referred to in square brackets.

1 Paired anterodorsal cup-shaped pores on cephalothorax absent

[present]

2 Paired anteroventral cup-shaped pores on cephalothorax absent

[present]

3 Paired cup-shaped pores on genital double-somite of 9 and genital

somite of ¢ absent [present]

4 Caudal rami without large pore medially or ventrally [present]

5 Cephalothorax without transverse spinular row dorsally [present]

6 Caudal rami not sexually dimorphic [modified in 9]

7 Antennule ?7-segmented [6-segmented; failure in separation of

segments 6 and 7]

8 Antennule d with 3 segments distal to geniculation [with 2

segments: segments 7 and 8 fused]

9 Aesthetasc of segment 4 in 2(and segment 5 in d) fused basally to

seta [fused to two setae forming trifid compound element]

10 Antennule segment 1 without processes in 9/d [with 3 spinous

processes along posterior margin]

11 Antennule segment 2 with large spinous process arising from

posterior margin in 9/d [absent]

12 Antennule segment 5 of 3d without anterior cylindrical process

(bearing large spine) [present]

13 Labrum without conspicuous ornamentation on anterior surface

[with overlapping scales distally and dense pattern of fine spinules

proximally]

14 Maxillulary endopod represented by 3 setae [2 setae, outermost seta

lost]

15 Maxillipedal syncoxa with 3 setae [state 1: 2 setae, proximal seta

lost; state 2: 1 seta]

16 Pl exopod 3-segmented [2-segmented; exp-2 and -3 fused]

17 Pl exopod 2-segmented, exp-2 with 3 outer spines and 2 apical

geniculate setae [exp-2 with 2 outer spines and 2 apical geniculate

setae]

18 P1 enp-1 with inner seta [absent]

19 P2 enp-2 with outer spine/seta [absent]

20 P3 endopod d 3-segmented [2-segmented; neotenic development]

il P3 enp-2 6 with inner seta [absent]

22 P5 baseoendopod @ with 5 setae [state 1: with 4 setae, middle inner

seta lost; state 2: with 3 setae]

23) P5 baseoendopod ¢ with 2 setae [setae absent]

24 P5 basoendopod 9/¢ without distinct setophore for outer basal seta

[basal seta positioned on long cylindrical setophore]

25 P5 exopod 2 with all outer setae arranged around margin [proximal

2 outer setae displaced with overlapping insertion sites]

R. HUYS AND W. LEE

in previous descriptions except for Vervoort (1962, 1964) who

briefly described the anterodorsal pores in C. bulligera and E.

vervoorti and suspected them to be eyes. Various authors (e.g.

Jakobi, 1953; Hamond, 1969; Mielke, 1981, 1997) have uninten-

tionally figured the modified pores on the caudal rami, however,

incorrect interpretation of the internal chitinized walls of the ducts as

external ridges (“Chitinleiste’) made them fail to recognize these

structures as true pores. Huys (1990b) pointed out that the trans-

formed cup-shaped pores in Esola are not serially homologous with

the pleural glands of the Adenopleurellidae and consequently can-

not serve as a basis for phylogenetic affinity. With the exception of

Archilaophonte and the typhlops-group of Esola all other esolinids

appear to exhibit a propensity for developing modified secretory

pores. The functional correlation between pores of different body-

regions is unknown and in view of their positional disparity and

structural differences it is unlikely that their expression is controlled

by a single gene. We postulate that the cup-shaped pore type evolved

from a surface precursor pore by major integumental invagination

and secondary development of setular extensions. These marginal

extensions either protect the depression or (more likely) maintain

the secrete bolus in close contact to the body wall. The degree of

invagination is obviously morphologically constrained and this is

particularly the case in swimming leg segments which are typically

depressed along the antero-posterior body axis. Although the ‘trap-

ping basket’ seta on the P4 endopod of C. bulligera represents a

radically divergent modification, it can be viewed as an external

analogue of the internal cup-shaped pore which developed in response

to this constraint. The tube-pore, which is also found in most other

esolinids, is enclosed by the long setules arising from the proximally

dilated distal inner seta (Fig. 25E-F) which hold the secrete bolus in

position. Since there are no differences in pore pattern between the

sexes a possible role in mate recognition is considered unlikely.

Huys (1992) demonstrated that in the interstitial Leptastacidae the

mucopolysaccharid strands produced by the caudal ramus glands

are intimately involved in mucus-trap feeding. We suggest that in

esolinids the secretory products discharged by the cup-shaped pores

perform a similar role in trophic gardening. It should be noted that

the caudal ramus pores located near the insertion sites of setae I-III

in E. typhlops (Fig. 31C—D) are not homologous to the large slit-like

pores found in Esola and Mourephonte.

Caudal ramus sexual dimorphism (character 6)

Females of Esola typically have bulbous caudal rami, displaying a

variety of swelling medially, ventrally and/or dorsally. Although the

secondary expansion appears to be correlated with the size of the

transformed pores, it is decoupled here from character 4 (presence of

caudal ramus pores) and scored separately. This is justified by the

absence of caudal ramus sexual dimorphism in A. hirsuta despite the

presence of modified pores in both sexes.

Setal fusion on antennules (character 9)

In most esolinids (except Archilaophonte) the proximal aesthetasc

(on segment 4 in 9, segment 5 in <) is fused at the base to 2 setae.

This trifid compound element is a unique character in the

Harpacticoida.

Antennulary processes (characters 10-12)

Within the esolinid grouping a spinous process along the posterior

margin of the second antennulary segment (character 11) is present

only in Archilaophonte. This is not an autapomorphy for the genus

but considered a retention of the ancestral state, based on outgroup

comparison with the remaining families of the Laophontoidea (Huys,

1990a; Huys & Lee, 1999). The presence of auxiliary processes

BASAL LAOPHONTID EVOLUTION

along the posterior margin of the first segment (character 10) is a

unique feature displayed by the species related to E. longicauda.

There are no equivalent structures known from other Laophontidae

and consequently this feature should be regarded an evolutionary

novelty for this species-group. In males of the same group the

enlarged fifth segment has produced an anterior sub-cylindrical

outgrowth bearing a stout modified spine (character 12). Minute

outgrowths are found on the first segment of C. bulligera but these

are not considered important enough to warrant a separate score.

Maxillulary endopod armature (character 14)

The maxillulary endopod typically bears 2 setae along the outer

margin of the basis, representing the incorporated endopod. This

condition is found in all esolinids while in several other laophontid

genera the endopod is represented by a cluster of 3 setae (e.g.

Langia, Quinquelaophonte: Mielke (1997)). A notable exception is

Archilaophonte in which the outermost third seta is secondarily

displaced to a more proximal position, i.e. at the base of the exopod.

Consequently, character 14 is scored 0 for A. maxima despite the

clearly derived positional pattern.

Male P3 endopod segmentation (character 20)

The P3 endopod in the males of A. typhlops and Esola sp. sensu

Chislenko (1967) is 2-segmented as in the female. The outer spine

forming the apophysis in the males of other esolinids has remained

largely unmodified except for reduction in size and basal fusion.

This virtual absence of sexual dimorphism is considered the

apomorphic state on the basis of ontogenetic evidence. Huys (1990a)

demonstrated that the typical 3-segmented condition is accom-

plished at the final moult by secondary subdivision of the distal

segment and allometric growth of the spinous apophysis. The

atypical pattern in A. typhlops, resembling the condition of a

copepodid V stage, is interpreted here as the result of neoteny, i.e.

the decrease in developmental rate has delayed the segmentation

beyond the final moult.

Male P3 endopod armature (character 21)

The modification of the male P3 endopod in esolinids has no effect on

the number of armature elements. In particular, the homologue of the

outer spine in the female is transformed into a spinous process or

apophysis arising from the middle segment in the male (but see

character 20), and the proximal inner seta on enp-2 of the 2-segmented

endopod in the female is retained on enp-2 of the 3-segmented

endopod in the male [typically 1.1.220 pattern]. The presence of the

latter seta in males is a particularly conservative character in primitive

laophontids, however, outside the esolinid grouping it is found only

in Onychocamptus and one species group of the genus Laophonte. The

fate of this seta during male development can only be traced in

Laophontidae displaying the full complement of 3 inner setae in the

female enp-2. In these species (Table 2) the endopodal armature

pattern is most commonly [0-1.321] but can also be [0.311] in

Laophonte nordgaardi Sars or [0.320] in some species of

Paralaophonte Lang. Except for 6 species of Laophonte and all

species of Onychocamptus, the proximal inner seta is consistently lost

in the male, resulting in a 0.0.220 pattern. The only exceptions with

3 inner setae in the male are those that have lost sexual dimorphism

altogether (Folioquinpes, Paralaophonte innae Chislenko, P.

aenigmaticum Wells, Hicks & Coull). Vervoort (1964) reported a very

long inner seta on the middle segment of Paralaophonte pilosoma but

re-examination of the holotype (USNM reg. no. 109763) has proven

this to be erroneous (Fig. 28E).

The loss of the proximal inner seta in the male is an apomorphy of

pivotal importance in laophontid evolution since it unifies nearly

101

95% of all species. Since many genera have only 0, 1 or 2 inner setae

in the female we have assumed that they are descendants from an

ancestral stock which displayed the 3-setae condition in the female

but lost the proximal one in the male.

Female P5 exopod armature (character 25)

Female esolinids can be readily identified by the setal arrangement

around the outer margin of the PS exopod. The two proximal setae

are displaced so that their respective insertion sites have become

superimposed on one another. Lang (1948) and Willen (1995)

pointed out that a similar displacement also occurs in Laophonte

parvula Sars (arrowed in Fig. 25G), however, we concur with the

latter author that this is the product of convergence.

Results

Analysis was performed with PAUP 3.1.1 (Swofford, 1993) using

the exact Branch and Bound algorithm (Hendy & Penny, 1982)

that is guaranteed to find all most parsimonious trees (MPTs),

with all characters set irreversible up and arbitrary solutions (zero-

length branches) suppressed. Analysis of the complete data (Table

4) produced 84 MPTs with tree length 40 and consistency index

0.675. The strict component consensus tree is illustrated in Fig.

32 and has a slightly longer length (42) and lower consistency

index (0.643). Relationships within the crown-group Esola are

poorly resolved, however construction of the majority-rule com-

ponent consensus tree revealed an additional group

(bulbifera-canalis-profunda). This boreo-mediterranean majority

component appears in 48 (57%) of the trees. A. longiremis, A.

hamondi and Esola sp. sensu Chislenko (1967) all have different

combinations of missing entries, however each is also a potential

taxonomic equivalent of A. typhlops (Table 4) and can therefore

be safely deleted (Wilkinson, 1995). Safe taxonomic reduction of

these taxa reduces the number of MPTs to 14 but does not alter

tree length or consistency index.

The strict component consensus (Fig. 32) reveals a strongly

supported basal dichotomy which divides the Laophontidae into two

major clades. In order to reflect the robustness of this dichotomy,

subfamilial rank is attributed to the two corresponding lineages. The

Esolinae subfam. nov. includes Archilaophonte, Mourephonte and

all species previously assigned to Esola. It is supported by male

antennulary segmentation (character 8) and the female P5 exopodal

setation pattern (character 25).

The primitive position of Archilaophonte conjectured by Willen

(1995) is confirmed. The genus represents the first offshoot in the

evolution of the Esolinae and is tentatively defined by the following

suite of autapomorphies: (a) 6-segmented @ antennule (segment 6

compound), (b) 2-segmented P1 exopod (fusion exp-2 and -3), (c)

Pl enp-2 secondarily elongated P2, (d) P2 enp-2 with only 1| inner

seta, (e) P3 enp-2 d with very long sigmoid apophysis, (f) P5 exopod

6 with 4 setae (loss of proximal inner seta), and (g) extremely

elongation of caudal rami. In addition, the maxillulary palp shows a

peculiar setal arrangement along the outer margin with 1 seta

positioned at the base of the bisetose exopod. Outgroup comparison

with the Normanellidae indicates that this seta is of endopodal origin

and must therefore have been secondarily displaced to a more

proximal position. The basal position of Archilaophonte is sup-

ported by the presence of (a) a spinous process on the posterior

margin of the 2nd antennulary segment, (b) maxillulary endopod

represented by 3 setae, (c) 3 setae on the maxillipedal syncoxa, and

(d) the well developed ¢ P5 endopodal lobe bearing 2 long setae.

The apomorphic alternatives of these characters (Table 3) in con-

junction with the formation of a trifid compound element on

102

ESOLINAE subfam. nov.

Se a ee ee a eee

Xx 2 2

Sy Re: ie) —

x Sg

= S & = = se

U4y 46S Mimo Se RS

Sess g ss 8 wk

eon ee ee a

TOSS eS S Kei

Noy ral

e :

| Applanola gen. nov.

hirsuta

R. HUYS AND W. LEE

LAOPHONTINAE

ade

3 = =; :

es °: >

: 9 -#& (=)

Sw nc bake =

vo i—| °

ee :

Ss : 8D

go “ss Ss

A oe 6s S jaa)

= 8 8 > <x

S 3S 8 v (e

sees § B

So. 8 x i)

Oe & x S)2

SSS ee Sas Wy

e £#Aa {a

alesis ule

2 & zw lO

aA 0 <

3 Se ee ls Se

Siar Shoe Bucs he Beurthre Ruse

Ln a ee ee SS =

a ae Ses ie eek Sabina et =

= eee eee ee es ee =

SS) Sh 3) Se fh cS tai OQ ||O

laa)

= a

a a

co N

Cl re)

% “4

1,18

9,11, 14,

15, 23

Fig. 32 Strict component consensus tree of 84 MPTs produced by parsimony analysis. Numbers refer to apomorphic character states listed in Table 3 (15!

and 15? denote multistep states).

antennulary segment 4 (or 5 in d) provide overwhelming support for

the monophyletic status of its sistergroup comprising Mourephonte

and ‘Esola’ sensu lato.

The phylogenetic analysis unequivocally identifies the paraphyly

of the genus Esola (as originally and pre-cladistically conceived).

Three northwestern European species and the unidentifiable Esola

spec. sensu Chislenko (1967) form a basal monophyletic group

(Archesola gen. nov.) defined by the 2-segmented ¢ P3 endopod and

the presence of an articulating basal setophore on the fifth legs of

both sexes. The degree of resolution within this clade will undoubt-

edly increase upon the discovery of the males of A. hamondi and A.

longiremis.

Evolution in the outgroup of Archesola is marked by a stepwise

addition of modified integumental pores. Initially, only paired

anterodorsal (or -lateral) pores were present on the cephalothorax (in

compacta, bulligera and possibly spelaea). This condition was

BASAL LAOPHONTID EVOLUTION

103

Table 4. States for characters listed in Table 3 [0 = plesiomorphic; 1 = apomorphic; 2 = further derived state]. Characters 14 and 22 are multistep

characters.

Taxon

ARCHILAOPHONTE

MOUREPHONTE

bulbifera

bulligera

galapagoensis

hirsuta

longicauda

longiremis

spelaea

typhlops

canalis sp. nov.

compacta sp. nov.

hamondi sp. nov.

lobata sp. nov.

profunda sp. nov.

vervoorti sp. nov.

spec. sensu Chislenko (1967)

spec. sensu Mielke (1997)

Laophonte cornuta-group

Onychocamptus

OTHER LAOPHONTIDAE

SOOVVHEH HRB VOHHOVOH HWE HO

SOOM VRE RHR VOOHOVOR RP VORHO|N

SOOv eH RBDTOOMroaCOoOnre {,OMSoSG|

SOCK OHHH OOH OCOORHBPH ORF O/]aA

SOOVVOHKPVOOHOVVVOVORVO|!|N

COOK VE HH OOH COOF OF OR VO! a

OPE EP VE HH OOH OO OHP HP Bee Ve

oe en

—

par

—

lon

—

(ore)

nas

(=)

—

Ie a NN a a a at >)

oo 0 YY OeH yy Oa YO yt OF OF or S'S

SOO srs OO) Ot (1 SS) a SS

oot ee ee a a a a ia)

SBNN VW Ve Se Vee ee yee eee eR oO

oF Ee SH OF KS KP OOK Coo r YE HS EHP -

SS So Sore Ss OO SS SS SS SS SOOO

Py py) Pe uy fer) Pet ee) eed ey SS Sis)

Sem oQooc”*ooddqdcoocroooraqooqcond

cooo7yrRrOowovrvvVFROoOwooooocno°o

eoovoovonvrvvornrvnvyooocnoeoocoeo

ON OF NER KH OOr OF OF eRe Ere yo

COrOowr Re NR NIN NOR Re DOR RR Re RRO

qooocorococorocorcrdqocoqocooo°qc”oo

a oe el el

further elaborated in both Mourephonte and the residual species of

Esola by the development of an accessory pair of anteroventral

pores on the cephalothorax (character 2) and of ventral or medial

pores on the caudal rami (character 4). Finally, the lateral pores on

the genital (double-)somite (character 3) evolved not until after the

divergence of Mourephonte.

The genus Esola is redefined here to encompass the terminal

polychotomy containing the type species E. longicauda and 7 other

species (Fig. 32). This strongly supported, cosmopolitan crown-

group is characterized by distinctive labral ornamentation, caudal

ramus sexual dimorphism, formation of 3 spinous processes on the

first antennulary segment and modification of segment 5 in the male

antennule E. hirsuta is the only species that shares genital cup-

shapes pores with this clade, however it is excluded from Esola and

placed in a monotypic genus Applanola on account of the following

autapomorphies: (1) dorsoventrally depressed body morphology,

(2) elongation of mandibular palp, (3) modification of P1 endopod,

(4) exopodal sexual dimorphism of P2—P3, (5) loss of outer spine on

P2 enp-2, and (6) strong reduction of the male sixth legs. The sexual

dimorphism on the P2—P3 exopod is unique in the Esolinae. Al-

though this character is globally homoplastic within the Laophontidae

it can be informative locally (see Lee & Huys, 1999) and should not

therefore be routinely ignored in phylogenetic analyses.

The three remaining species, compacta, spelaea and bulligera,

are identified as independent lineages splitting off successively

between the basal Archesola clade and the terminal ((Mourephonte

+ Esola) + Applanola) clade. Corbulaseta gen. nov., accommodat-

ing E. bulligera, is most closely related to the latter clade because of

shared fusions in the female antennule (segments 6-7) and Pl

exopod (exp-2 and -3). The modified distal inner seta forming a

trapping-basket is a unique autapomorphy for this genus. The

position of Troglophonte is tentative pending the confirmation of

cup-shaped pores on the cephalothorax and of the armature patterns

of P2 exopod and P5 in both sexes. The basal position of the genus

Bathyesola is caused by its retention of the maximum number of

setae on the female PS baseoendopod.

The genus Mourephonte is radically divergent from other esolinids.

The extreme development of the P1, the complete absence of the P2

endopod, the loss of the inner seta on the P2—P4 exopods and the

wide separation of the apical setae on P4 enp-2 form a remarkable

combination of autapomorphies which places it on a distinct evolu-

tionary lineage, ruling out possible inclusion in the genus Esola

under a broader concept.

The residual laophontids, comprising 95% of the known species,

are grouped in the subfamily Laophontinae. All 54 genera have lost

the inner seta on P| enp-1 and the outer spine on P2 enp-2, and bear

a maximum of 2 setae on the maxillipedal syncoxa (absence of

proximal seta). With the exception of the genus Onychocamptus and

the Laophonte cornuta-group all Laophontinae are characterized by

the P3 endopod sexual dimorphism involving the loss of the proxi-

mal inner seta of enp-2 (character 21). The isolated position of the

cornuta-group (= Laophonte Group I + adduensis + ciliata: Table 2)

testifies to the widely accepted polyphyletic status of the genus

Laophonte and has major nomenclatural consequences because of

its inclusion of the type species L. cornuta Philippi. Restriction of

the generic concept to the cornuta-group will require the other 37

species of Laophonte to be re-allocated to other existing or new

genera. This is a major task which can only be accomplished by

sound phylogenetic analysis involving the remaining laophontinid

genera. The sistergroup relationship between the cornuta-group and

Onychocamptus depicted in Fig. 32 is not be taken as absolute since

other advanced but closely related genera such as Folioquinpes have

deliberately been omitted from the outgroup to the Esolinae. Al-

though inclusion of these genera in future analyses may introduce

additional basal nodes changing the relative position of Laophonte

and Onychocamptus, we envisage that the latter will consistently

show up as an early speciation event predating the evolution of the

other Laophontinae.

Subfamilial division

ESOLINAE subfam. nov.

Rostrum delimited at base by surface suture; antennule 2 6- or 7-

segmented, usually without spinous process on segment 2 but

frequently with processes on segment 1; 7-segmented and haplocerate

or subchirocerate in 6, with only 2 segments distal to geniculation;

104

proximal aesthetasc typically fused to 2 setae (except Archilao-

phonte). Antennary exopod with 4 well developed setae. Mandible

typically biramous (except Applanola and Mourephonte). Maxilla

with 3 endites on syncoxa. Maxilliped with 2—3 setae on syncoxa.

P1 with 2- or 3-segmented exopod, retaining full complement of

setae (0.0.022 or 0.023); enp-1 occasionally with inner seta. P2 enp-

2 with outer spine (except Applanola) or entire P2 endopod absent

(Mourephonte). P3 endopod ¢ retaining proximal inner seta of 2

enp-2 (except for Troglophonte where it is lost in both sexes).

Armature formula as follows:

Exopod Endopod

p2 0.1.123 0-1.(1—2)2(0-1) or absent

P3 0.1.(1—2)23 0-1.321

P4 0.1.(1=2)23 0-1.221

P5 @ with separate rami; exopod elongate, with 6 setae/spines;

proximal two setae along outer margin with superimposed insertion

sites; baseoendopod trapezoid, slightly developed, with 4—5 setae/

spines. P5 d without endopodal lobe (except for Archilaophonte,

bearing 2 long setae), no endopodal armature; exopod 5 setae/

spines.

Typically with cup-shaped transformed pores on cephalothorax,

genital (double-)somite, and/or caudal rami.

TYPE GENUS. Esola Edwards, 1891

OTHER GENERA. Mourephonte Jakobi, 1953; Archilaophonte

Willen, 1995; Applanola gen. nov.; Archesola gen. noy.; Bathyesola

gen. nov.; Corbulaseta gen. nov.; Troglophonte gen. nov.

Laophontinae T. Scott, 1905

Antennule d with up to 3 segments distal to geniculation; proximal

aesthetasc fused to 1 seta. Mandible typically uniramous. Maxilliped

with maximum 2 setae on syncoxa. P| enp-1 without inner seta. P2

enp-2 without outer spine. P3 endopod ¢ typically not retaining

proximal inner seta of 2enp-2 (except for Laophonte cornuta-group

and Onychocamptus).

Proximal outer setae of 2 P5 exopod with distinctly separated

insertion sites.

Cup-shaped transformed pores on cephalothorax, genital (double-)

somite, and/or caudal rami never present.

TYPE GENUS. Laophonte Philippi, 1840

OTHER GENERA. Fifty-five; see Lang (1948), Bodin (1997), George

(1997) and Lee & Huys (1999) for complete list.

KEY TO GENERA OF ESOLINAE

[PR 2endopodkabsemtgeceer secre ee Mourephonte Jakobi, 1953.

PAendopodspresenty2-Se SMe Med meecsee esas eee eee De

2. Antennulary segment 2 with large spinous process along anterior

margin; P2 enp-2 with 1 inner seta; P5 baseoendopod d with 2 long

ISVS Spo acen aa ee ses ssoe soe eSoeeeceeuassendebocoocececaee Archilaophonte Willen, 1995.

Antennulary segment 2 without spinous process along anterior margin;

P2 enp-2 with 2 inner setae; P5 baseoendopod ¢ without setae ....... 3:

3. Antennule 2 6-segmented; P1 exopod 2-segmented; caudal rami with

Mecialormven thal anno chiki dap OLesmeeewes ee see eee eee ene eee eee 4.

Antennule 2°7-segmented; P1 exopod 3-segmented; caudal rami with-

QULBISUCIY POLES acess vest es eo eonicees cae sms asctecepae cere conn aAcattucs seeeeeeaees heen 6.

4. Body short, dorsoventrally flattened; P2 enp-2 outer spine absent; P3

R. HUYS AND W. LEE

exopod ¢ strongly modified 00.0.0... Applanola gen. nov.

Body elongate, sub-cylindrical; P2 enp-2 outer spine present; P3 exopod

G MOt tmOdified ez. c, Be ececkeeacecee eo ose scene seed se a:

5. Antennulary segment | with 3 spinous processes along posterior mar-

gin; distal inner seta of P4 endopod not transformed; caudal rami 2

modified, with bulbous swelling dorsally, ventrally and medially .......

ss hevanstehs Seth ts «ons dane dR ences aicasessaaiitenessseneae ani Esola Edwards, 1891.

Antennulary segment | without distinct spinous processes; distal inner

seta of P4 endopod transformed; caudal rami not sexually dimorphic,

Gylim det Calls. 5 lake cs eae Mees seete casks een eee Corbulaseta gen. nov.

6. P3—P4 exp-3 with 1 inner seta; P3—P4 enp-1 without inner seta .........

SEE Er ee ee 2 ae eee Bathyesola gen. nov.

P3—P4 exp-3 with 2 inner setae *; P3—P4 enp-1 with inner seta....... 7.

7. P3 enp-2 with 3 inner setae; P3 endopod d 2-segmented; P5

baseoendopod with long articulating setophore in both sexes .............

seus nstnn jedan ato sugbhes bevdteocaeettneds voscunecentettte stot rence weer Archesola gen. nov.

P3 enp-2 with 2 inner setae; P3 endopod ¢ 3-segmented; P5

baseoendopod of both sexes without articulating setophore................

sdaacdibiaigia snasat Biaesov satel niveasees tosvOueciede eect Troglophonte gen. nov.

* Note that Chappuis’ (1938) setal formula of P3 exp-3 can also be

interpreted as 123, implying the presence of only 1 inner seta.

ECOLOGICAL RADIATION OF ESOLINAE

Although none of the 18 species can be considered as truly cosmo-

politan, the subfamily as a whole occurs in all oceanic basins,

including the Antarctic Ocean. Superimposing habitat utilization

upon the phylogeny presented in Fig. 32 reveals an interesting but

complex ecological radiation pattern. Esolinae are essentially shal-

low water inhabitants, however, the variety of additional habitats

exploited by this lineage is startling for its small number of known

species. Considered against the background of the overwhelming

evolutionary success of their sister-lineage Laophontinae, esolinids

can be viewed as relicts of a formerly diverse group.

Lee & Huys (1999) reviewed published deepwater records of

Laophontidae and regarded the colonization of the deep sea by this

family as remarkably unsuccessful. There is no single lineage

containing all deepwater forms, and the three exclusively bathyal

genera in the Laophontinae, Cornylaophonte Willen, Weddellao-

phonte Willen and Bathylaophonte Lee & Huys can be considered as

independent colonists of this habitat. Colonization of the deep sea by

the Esolinae follows a similarly erratic trend with early attempts by

the monotypic genera Archilaophonte in the Antarctic and Bathyesola

in the western Pacific. Within the genus Esola, E. profunda repres-

ents a third, secondary deepwater invasion derived from a shallow

water inhabiting ancestral stock (Fig. 32).

According to Pesce (1985) and Rouch (1986) the genus

Troglophonte is likely to be derived from a marine ancestor stranded

during the lowering of sea level during the Tertiary. It is highly

endemic to freshwater lenses in several Apulian caves in southern

Italy (Chappuis, 1938). These caves are separated from the littoral

zone by macroporous karstic rock and exhibit a detectable tidal

current which appears insufficient to ensure substantial mixing of

the water inside the caves. The strong stratification with freshwater

lenses overlying the poorly oxygenated deeper layers has clearly

prevented the establishment of a diverse marine benthic fauna.

Rather than considering Troglophonte a Tethyan relict, its present

restricted distribution can also be regarded as a relatively recent

landward habitat range extension from a primarily shallow-subtidally

residing ancestral stock. Although some Laophontidae are regularly

BASAL LAOPHONTID EVOLUTION

found in salt-marsh and mudflat habitats within river estuaries

(Noodt, 1957; Barnett, 1968; Bodin, 1976) or in brackish lagoons

(Heip, 1969; Hamond, 1972), tolerance to oligohalinity may have

appeared convergently only twice in the family. Both colonization

events presumably occurred early in the evolution of the family (Fig.

32), however their nature is fundamentally different. The evolution-

ary success of the Troglophonte lineage has clearly remained limited,

both in dispersal and speciation. It can be considered as a freshwater

incursion without further radiation or diversification. The second

invasion of low salinity environments is cosmopolitan in scope and

probably of Tethyan origin, containing the genera Onychocamptus

Daday and Folioquinpes Fiers & Rutledge (Lee & Huys, 1999).

Little is known about the possible dispersal of Laophontidae in

marine caves. Pesta (1959) reported E. rosei from a submarine cave

near Naples and several unidentified Laophontidae were recorded

by Huys (1996) from the anchialine Walsingham Cave on Bermuda.

Examination of samples from Caye Chapel Cave in Belize, the type

locality of the recently discovered family Novocriniidae (Huys &

Iliffe, 1998), resulted in the discovery of a single male belonging to

a new genus of Esolinae. The new genus has several characters in

common with Archilaophonte such as the presence of a spinous

process on the second antennulary segment, the displacement of the

outermost endopodal seta on the maxillule, the presence of 3 setae

on the maxillipedal syncoxa, Pl with 2-segmented exopod and

elongate enp-2, P2 enp-2 with only | inner seta and presence of a

very long apophysis on P3 endopod. Phylogenetic analysis identi-

fied the Belize genus unambiguously as the sistergroup of

Archilaophonte, suggesting the evolution of an independent

cavernicolous lineage in the western Atlantic.

The genus Archesola is exclusively boreo-arctic in distribution

and restricted to the Atlantic basin, with a single known outlier from

the Black Sea (Por, 1959). Its southernmost limit based on reliable

records is Norfolk (England), however, confirmation of the doubtful

records of A. longiremis from the south coast of England (Wells,

1961, 1963, 1970) and North Carolina (Coull, 1971) may extend this

limit further southward. The genus occurs primarily at higher lati-

tudes, showing limited dispersal in Arctic waters such as the White

Sea (Brotskaya, 1961; Chislenko, 1967). It is suggested that the

strongly discontinuous, bipolar distribution of the two basal clades,

with Archesola restricted to northern Europe and Archilaophonte to

the Antarctic, indicates a wider, perhaps continuous, horizontal

zonation of primitive stenothermal esolinids at greater depths. This

trend of ‘Equatorial Submergence’ appears to be supported by the

discovery of Bathyesola in the deep tropical western Pacific, the first

lineage to diverge after Archesola (Fig. 32).

A major event in the evolution of the Esolinae was the episode of

rapid speciation within the genus Esola. This event is revealed as a

polychotomy in the cladogram (Fig. 32) although it is clear that the

low resolution is partly attributable to the abundance of missing

entries for several taxa which are known from one sex only (Table

3). Many of these species are small-sized (Table 1) and adapted to a

mesopsammic life-style in shallow subtidal localities and sandy

beaches, while others are frequently found associated with algal

substrates. Results show that only a fraction of the species is known.

Although the genus assumes a cosmopolitan distribution it is pre-

dominantly restricted to the circum-tropical belt. This zone coincides

with the former Tethyan seaway separating the northern and south-

ern continents, which continued into Palaeogene times with free

marine continuity along its length not being interrupted until the

beginning of the Neogene. One Pacific-Caribbean subgroup, com-

prising E. longicauda, E. galapagoensis and Esola sp. (Fig. 32),

probably originated from an ancestral stock in the western Pacific.

From there, eastward dispersal was greatly influenced by tectonic

105

plate movement, particularly the formation of the Caribbean plate at

the beginning of the Oligocene. This was established by decoupling

of the eastward protruding tongue of the East Pacific plate, causing

the formation of a subduction zone along what is now the western

coast of southern Central America, and the subsequent westward

motion of North and South America past a nearly stationary Carib-

bean plate (Malfait & Dinkelman, 1972; Coney, 1982). The entry of

the ancestor of E. longicauda into the Caribbean must have preceded

the closing of the Panama land bridge approximately 3.1—3.5 Ma

(Keigwin, 1978).

Applanola displays amore disjunct distribution than its sistergroup

Esola, provided that Pesta’s (1916) record from the Gulf of Guinea

is correct. The dorsoventrally depressed body, robust maxillipeds

and powerful Pl endopod indicate that A. hirsuta may be loosely

associated with invertebrate hosts. Thompson & A. Scott (1903)

obtained the species from washings of pearl oysters and other

dredged invertebrates but did not present any firm evidence for a

clear association.

ACKNOWLEDGEMENTS. Dr Danielle Defaye (Muséum National d’ Histoire

Naturelle, Paris), Dr Chad T. Walter (National Museum of Natural History,

Washington D.C.), Karen Gowlett-Holmes (South Australian Museum) and Dr

Mark Holmes (National Museum of Ireland) kindly made type and other

material available. Dr Richard Hamond (Norfolk, U.K.) provided us with

specimens of E. bulbifera and A. typhlops. The material of Bathyesola

compacta was collected during the STARMER II expedition to the Fiji Basin

(chief scientist: L. Laubier). One of us (W.L.) acknowledges financial support

from the Korea Research Foundation provided for the programme year 1997.

REFERENCES

Alheit, J. & Scheibel, W. 1982. Benthic harpacticoids as a food source for fish. Marine

Biology, Berlin 70(2): 141-147.

Barnett, P.R.O. 1968. Distribution and ecology of harpacticoid copepods of an intertidal

mudflat. /nternationale Revue der gesammten Hydrobiologie 53: 177-209.

Bodin, P. 1976. Les Copépodes Harpacticoides (Crustacea) des cétes Charentaises

(Atlantique). Données écologiques et biologiques sur les espéces principales. Bulle-

tin du Muséum national d’ Histoire naturelle, Paris (3)353 (= Ecologie générale 29):

1-45.

. 1997. Catalogue of the new marine harpacticoid copepods. Studiedocumenten

van het K.B.I.N. 89: 1-304.

Brian, A. 1928a. Descrizione di specie nuove 0 poco conosciute di Copepodi bentonici

del mare Egeo. (Nota preliminare). Bollettino dei Musei di Zoologia e Anatomia

comparata della R. Universita di Genova (2)7(18): 1-37.

. 1928b. I Copepodi bentonici marini. Archivio zoologico italiano 12: 293-343.

Brotskaya, V.A. 1961. Materialy po faune Harpacticoida (Crustacea, Copepoda)

Velikoi salmy i prilezheshchikh uchastkov Belogo morya. Contributions to the fauna

of Harpacticoidea (Crustacea: Copepoda) of the Velikaya Salma Strait and of

adjacent region of the White Sea. Biologiya Belogo Morya (= Trudy belomorskoi

biologicheskoi Stantsit. M.G.U.) 1: 109-129.

Chappuis, P.A. 1938. Subterrane Harpacticoiden aus Siid-Italien. Bulletinul Societafii

de Stiinte din Cluj 9: 153-181.

Chislenko, L.L. 1967. Garpaktitsidy (Copepoda Harpacticoida) Karelskogo

poberezh’ ya Belogo morya. [Copepoda Harpacticoida of the Karelian coast of the

White Sea. /ssledovaniya Fauny Morei 7(15): 48-196.

Coney, P.J. 1982. Plate tectonic constraints on the biogeography of Middle America

and the Caribbean region. Annals of the Missouri Botanical Garden 69: 432-443.

Coull, B.C. 1971. Meiobenthic Harpacticoida (Crustacea, Copepoda) from the North

Carolina continental shelf. Cahiers de Biologie marine 12(2): 195-237.

Douwe, C. yan 1929. Marine Litoral-Copepoden: Zur Verbreitung des Genus Laophonte

Philippi im Mittelmeer. Zoologischer Anzeiger 83(11—12): 283-294.

Drzycimski, I. 1969. Harpacticoida (Copepoda) wé6d morskich okolic Bergen

(Zachodnie Wybrzeze Norwegii) i ich ekologia. Harpacticoida (Copepoda) of sea

waters in Bergen region (West Coast of Norway) and their ecology. (Polish with

English and Russian summaries). Wyzsza Szkola Rolnicza w Szczecinie 17: 1-72.

Edwards, C.L. 1891. Beschreibung einiger neuen Copepoden und eines neuen

copepodenahnlichen Krebses, Leuckartella paradoxa. Archiv fiir Naturgeschichte

57: 75-104.

106

Farran, G.P. 1913. Marine Entomostraca. In: A biological survey of Clare Island in the

county of Mayo, Ireland, and of the adjoining district. Proceedings of the Royal Irish

Academy (B)31(45): 1-20.

. 1915. Results of a biological survey of Blacksod Bay, Co. Mayo. Scientific

Investigations. Fisheries Branch, Department of Agriculture for Ireland, Dublin

1914(3): 1-72.

Fiers, F. 1986. Harpacticoid copepods from the West Indian Islands: Laophontidae

(Copepoda, Harpacticoda). Amsterdam Expedition to the West Indian Islands,

Report 48. Bijdragen tot de Dierkunde 56(1): 132-164.

George, K.H. 1997. Mielkiella spinulosa gen.n. sp.n., anew taxon of the Laophontidae

(Copepoda, Harpacticoida) from Porvenir (Tierra del Fuego, Chile). Microfauna

Marina 11: 71-86.

Gurney, R. 1927. Report on the Crustacea. — Copepoda (littoral and semi-parasitic).

Zoological results of the Cambridge expedition to the Suez Canal, 1924, no. 35.

Transactions of the zoological Society of London 22: 451-577.

Hamond, R. 1969. The Laophontidae (Copepoda, Harpacticoida) of the shore at West

Runton, Norfolk, England. Crustaceana 16(1): 1-14.

——. 1972. Some marine and brackish-water copepods from Wells-next-the-Sea,

Norfolk, England. Transactions of the Norfolk and Norwich Naturalists’ Society

22(4): 237-243.

Heip, C. 1969. Drie Copepoden nieuw voor de Belgische fauna. Biologisch Jaarboek

Dodonaea 37: 42-49.

Hendy, M.D. & Penny, D. 1982. Branch and bound algorithms to determine minimal

evolutionary trees. Mathematical Bioscience 59: 277-290.

Hicks, G.R.F. 1988a. Systematics of the Donsiellidae Lang (Copepoda, Harpac-

ticoida). Journal of natural History 22(3): 639-684.

—. 1988b. Harpacticoid copepods from biogenic substrata in offshore waters of New

Zealand. 1: New species of Paradactylopodia, Stenhelia (St.) and Laophonte.

Journal of the Royal Society of New Zealand 18(4): 437-452.

Hockin, D.C. 1982a. The harpacticoid copepod fauna of the River Ythan and its

estuary, Aberdeenshire, Scotland. Journal of the marine biological Association of

the United Kingdom 62(3): 729-736.

. 1982b. The effect of sediment particle diameter upon the meiobenthic copepod

community of an intertidal beach: a field and laboratory experiment. Journal of

animal Ecology 51(2): 555-572.

. 1984. Records of symbiotic Protozoa from harpacticoid copepods of a sandy

beach. Crustaceana 46(3): 319-320.

& Ollason, J.C. 1981. The colonization of artificially isolated volumes of

intertidal estuarine sand by harpacticoid copepods. Journal of experimental marine

Biology and Ecology 53(1): 9-29.

Holmes, J.M.C. & O’Connor, J.P. 1990. A provisional list of the Harpacticoida

(Crustacea: Copepoda) of Ireland. Bulletin of the Irish biogeographical Society 13:

44-130.

Huys, R. 1990a. A new family of harpacticoid copepods and an analysis of the

phylogenetic relationships within the Laophontoidea T. Scott. Bijdragen tot de

Dierkunde 60(2): 79-120.

—. 1990b. Adenopleurella, new genus, Proceropes, new genus, Sarsocletodes

Wilson (ex Laophontidae), and Miroslavia Apostolov (ex Cletodidae): representati-

ves of a new family (Copepoda: Harpacticoida). Journal of Crustacean Biology

10(2): 340-363.

—. 1992. The amphiatlantic distribution of Leptastacus macronyx (T. Scott, 1892)

(Copepoda: Harpacticoida): a paradigm of taxonomic confusion; and, a cladistic

approach to the classification of the Leptastacidae Lang, 1948. Mededelingen van de

Koninklijke Academie voor Wetenschappen, Letteren en schone Kunsten van Belgié

54(4): 21-196.

—. 1996 Superornatiremidae fam. nov. (Copepoda: Harpacticoida): an enigmatic

family from North Atlantic anchihaline caves. Scientia Marina 60: 497-542.

— & Gee, J.M., Moore, C.G. & Hamond. R. 1996. Synopses of the British Fauna

(New Series): Marine and Brackish Water Harpacticoid Copepods Part 1. viii +

352p. Field Studies Council, Shrewsbury, United Kingdom.

& lliffe, T.M. 1998. Novocriniidae, a new family of harpacticoid copepods from

anchihaline caves in Belize. Zoologica Scripta 27: 1-15.

& Lee, W. 1999. On the relationships of the Normanellidae and the recognition

of Cletopsyllidae grad. nov. (Copepoda, Harpacticoida). Zoologischer Anzeiger,

237: 267-290.

& Willems, K.A. 1989. Laophontopsis Sars and the taxonomic concept of the

Normanellinae (Copepoda: Harpacticoida): a revision. Bijdragen tot de Dierkunde

59(4): 203-227.

Jakobi, H. 1953. Novos Laophontidae (Copepoda-Crustacea) da costa Brasileira.

(Neue Laophontiden von der Brasilianischen Kiiste). Dusenia 4(1): 47-60.

Keigwin, L.D. 1978. Pliocene closing of the isthmus of Panama, based on

biostratigraphic evidence from nearby Pacific Ocean and Caribbean Sea cores.

Geology 6: 630-634.

Klie, W. 1941. Laophonte-Arten (Cop. Harp.) aus dem Mittelmeer mit verkimmertem

Nebenast der zweiten Antenna. Archiv fiir Naturgeschichte n. ser. 10: 259-277.

Krishna Murty, P.V.M. 1983. Distribution of phytal harpacticoid copepods along

Visakhapatnam coast. Mahasagar 16(1): 47-54.

R. HUYS AND W. LEE

Krishnaswamy, S. 1957. Studies on the Copepoda of Madras. 168p. Thesis, Univer-

sity of Madras.

Lang, K. 1944. Monographie der Harpacticiden (Vorldufige Mitteilung). 39p. Almqvist

& Wiksells Boktryckeri Ab, Uppsala.

——. 1948. Monographie der Harpacticiden, volume I. pp. 1-896, volume IJ. pp. 897—

1683. Hakan Ohlssons Boktryckeri, Lund, Sweden.

. 1965. Copepoda Harpacticoida from the Californian Pacific coast. Kungliga

Svenska VetenskapsAkademiens Handlingar (4)10(2): 1-560.

Lee, W. & Huys, R. 1999. Bathylaophonte gen. nov. from deep-sea hydrothermal vents

and the polyphyly of Paronychocamptus Lang (Copepoda: Harpacticoida). Cahiers

de Biologie marine 40: 293-328.

Malfait, B.T. & Dinkelman, M.G. 1972. Circum-Caribbean tectonic and igneous

activity and the evolution of the Caribbean plate. Bulletin of the Geological Society

of America 83: 251-272.

Mielke, W. 1981. Interstitielle Fauna von Galapagos. XXVIII. Laophontinae

(Laophontidae), Ancorabolidae (Harpacticoida). Mikrofauna des Meeresbodens 84:

1-106.

—. 1997. On a small collection of Laophontidae (Copepoda) from Sulawesi.

Microfauna Marina 11: 223-250.

Monard, A. 1926. Description de quelques espéces nouvelles d’Harpacticides marins

de la région de Banyuls. Revue suisse de Zoologie 33: 619-628.

. 1927. Synopsis universalis generum Harpacticoidarum. Zoologische Jahrbiicher,

Systematik 54(1—2): 139-176.

. 1928. Les Harpacticoides marins de Banyuls. Archives de Zoologie expérimentale

et générale 67: 259-443.

. 1935. Etude sur la faune des Harpacticoides marins de Roscoff. Travaux de la

Station biologique de Roscoff 13: 5-88.

. 1937. Les Harpacticoides marins de la région d’ Alger et de Castiglione. Bulletin

de la Station d’Aquiculture et de Péche de Castiglione 1935(2): 9-93.

Moore, P.G. 1973. The kelp fauna of northeast Britain Il. Multivariate classification:

turbidity as an ecological factor. Journal of experimental marine Biology and

Ecology 13: 127-164.

Nicholls, A.G. 1941a. Littoral Copepoda from South Australia. (1) Harpacticoida.

Records of the South Australian Museum 6(4): 381-427.

. 1941b. A revision of the families Diosaccidae Sars, 1906 and Laophontidae T.

Scott, 1905. (Copepoda, Harpacticoida). Records of the South Australian Museum

7(1): 65-110.

. 1944. Littoral Copepoda from the Red Sea. Annals and Magazine of natural

History (11)11: 487-503.

. 1945. Marine Copepoda from Western Australia. III. Littoral harpacticoids from

Port Denison. Journal of the Royal Society of Western Australia 29: 1-16.

Noodt, W. 1955. Marmara denizi Harpacticoid’leri (Crust. Cop.). Marine Harpacticoiden

(Crust. Cop.) aus dem Marmara Meer. Istanbul Universitesi Fen Fakiiltesi Mecmuas1

(B)20(1-2): 49-94.

. 1957. Zur Okologie der Harpacticoidea (Crust. Cop.) des Eulitorals der deutschen

Meereskiiste und der angrenzenden Brackgewasser. Zeitschrift fiir Morphologie und

Okologie der Tiere 46: 149-242.

—. 1958. Die Copepoda Harpacticoidea des Brandungsstrandes von Teneriffa

(Kanarische Inseln). Abhandlungen der mathematisch-naturwissenschaftlichen

Klasse. Akademie der Wissenschaften und der Literatur in Mainz 1958(2): 53-116.

Norman, A.M. 1911. Three species of harpacticoid Copepoda. Transactions of the

Linnean Society of London, Zoology (2)11: 137-143.

Pesce, G.L. 1985. The groundwater fauna of Italy: a synthesis. Stygologia 1(2): 129-

159.

Pesta, O. 1916. Crustacea I: Copepoda. In: W. Michaelsen, ed., Beitrdge zur Kenntnis

der Meeresfauna Westafrikas: 1-10.

. 1959. Harpacticoiden (Crust. Copepoda) aus submarinen Hohlen und den

benachbarten Litoralbezirken am Kap von Sorrent (Neapel). Ergebnisse der

Osterreichischen Tyrrhenia-Expedition 1952. Teil: VI. Pubblicazione della Stazione

zoologica di Napoli 30, suppl.: 94-177.

Petkovski, T.K. 1955. Zweiter Beitrag zur Kenntnis der Harpacticidenfauna unserer

Meereskiiste. Fragmenta balcanica 1(15): 125-139.

Por, F.D. 1959. Harpacticoide noi (Crustacea, Copepoda) din milurile Marii Negre.

(Harpacticoides nouveaux (Crustacés, Copépodes) des vases de la mer Noire). Studii

si Cercetari de Biologie, Seria Biologie animala 4(11): 347-368.

. 1964a. Les Harpacticoides (Crustacea, Copepoda) des fonds meubles du Skagerak.

Cahiers de Biologie marine 5(3): 233-270.

. 1964b. A study of Levantine and Pontic Harpacticoida (Crustacea, Copepoda).

Zoologische Verhandelingen, Leiden 64: 1-128.

. 1967. Level bottom Harpacticoida (Crustacea, Copepoda) from Elat (Red Sea),

part I. Israel Journal of Zoology 16: 101-165.

— & Marcus, A. 1973. Copepoda Harpacticoida of the Suez Canal. In: Contribu-

tions to the knowledge of Suez Canal migration. /srael Journal of Zoology 21(3-4):

249-274.

Rouch, R. 1962. Harpacticoides (Crustacés Copépodes) d’ Amérique du Sud. Biologie

de l’Amérique Australe 1: 237-280.

. 1986. Copepoda: Les Harpacticoides souterrains des eaux douces continentales.

BASAL LAOPHONTID EVOLUTION

pp. 321-355. In: Botosaneanu, L. (ed.) Stygofauna Mundi, a faunistic, distributional,

and ecological synthesis of the worldfauna inhabiting subterranean waters (includ-

ing the marine interstitial). E.J. Brill/Dr. W. Backhuys, Leiden.

Sars, G.O. 1908. Copepoda Harpacticoida. Parts XXI & XXII. Laophontidae (contin-

ued). An Account of the Crustacea of Norway, with short descriptions and figures of

all the species 5: 241-256.

Scott, A. 1909. The Copepoda of the Siboga Expedition. Part I. Free-swimming, littoral

and semi-parasitic Copepoda. Siboga Expedition, Monographs 29a: 1-323.

Scott, T. 1905. On some new and rare Crustacea from the Scottish seas. Report of the

Fishery Board for Scotland 23(3): 141-153.

—. 1906. Notes on new and rare Copepoda from the Scottish seas. Report of the

Fishery Board for Scotland 24(3): 275-280.

Sewell, R.B.S. 1940. Copepoda, Harpacticoida. Scientific Reports of the John Murray

Expedition 7(2): 117-382.

Swofford, D.L. PAUP Phylogenetic Analysis Using Parsimony, version 3.1.1. Docu-

mentation and Software. Illinois Natural History Survey, Champaign.

Thompson, I.C. & Scott, A. 1903. Report on the Copepoda collected by Professor

Herdman, at Ceylon, in 1902. In: W.A. Herdman, Report to the Government of

Ceylon on the Pearl Oyster Fisheries of the Gulf of Manaar 1, suppl. 7: 227-307.

Vervoort, W. 1962. Report on some Copepoda collected during the Melanesia Expedi-

tion of the Osaka Museum of Natural History. Publications of the Seto marine

biological Laboratory 10(2): 393-470.

——. 1964. Freeliving Copepoda from Ifaluk Atoll in the Caroline Islands with notes

on related species. Smithsonian Institution United States National Museum Bulletin

236: i-ix, 1-431.

107

VriSer, B. 1984. Strukturne in kvantitativne znatilnosti meiofavne v notranjosti

Piranskega, Strunjanskega in Koprskega zaliva. [The structure and abundance of

meiofauna in the inner parts of Piran, Strunjan and Koper Bays (Gulf of Trieste,

North Adriatic)]. Bioloski Vestnik 32: 121-136.

——. 1986. Vpliv organskega onesnazenja na meiogavno priobalnih glinastih muljev

Koprskega zaliva. [The effect of organic pollution on mud meiofauna communi-

ties (Bay of Koper, Gulf of Trieste, Northern Adriatic)]. Bioloski Vestnik 34:

93-104.

Wells, J.B.J. 1961. Interstitial copepods from the Isles of Scilly. Crustaceana 2(4):

262-274.

——. 1963. Copepoda from the littoral region of the estuary of the River Exe (Devon,

England). Crustaceana 5(1): 10-26.

——. 1970. The marine flora and fauna of the Isles of Scilly. Crustacea: Copepoda:

Harpacticoida. Journal of natural History 4(2): 255-268.

& Rao, G.C. 1987. Littoral Harpacticoida (Crustacea: Copepoda) from Andaman

and Nicobar Islands. Memoirs of the Zoological Survey of India 16(4): 1-385.

Wilkinson, M. 1995. Coping with abdundant missing entries in phylogenetic inference

using parsimony. Systematic Biology 44(4): 501-514.

Willen, E. 1995. Archilaophonte maxima gen. n., spec. n., a new taxon of the

Laophontidae (Copepoda, Harpacticoida) from the high Antarctic (Weddell Sea).

Hydrobiologia 302: 241-255.

——. 1996. Two new genera of Laophontidae (Copepoda: Hapacticoida) from the high

Antarctic Weddell Sea. Journal of natural History 30: 1297-1327.

Willey, A. 1935. Harpacticoid Copepoda from Bermuda. — Part II. Annals and Maga-

zine of natural History (10)15: 50-100.

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the word-processor programme used. Indicate whether IBM or

Apple Mac (IBM preferred).

3. Supply the file in the word-processor format; if there is a

facility to save in ASCII please submit the file in ASCII as well.

4. Specify any unusual non-keyboard characters on the front

page of the hard copy.

5. Do not right-justify the text.

6. Do not set a left-hand margin.

7. Make sure you distinguish numerals from letters, e.g. zero (0)

from O; one (1) from 1 (el) and I.

8. Distinguish hyphen, en rule (longer than a hyphen, used

without a space at each end to signify ‘and’ or ‘to’, e.g. the Harrison—

Nelson technique, 91-95%, and increasingly used with a space at

each end parenthetically), and em rule (longer than an en rule, used

with a space at each end parenthetically) by: hyphen, two hyphens

and three hyphens, respectively. Be consistent with rule used paren-

thetically.

9. Use two carriage returns to indicate beginnings of paragraphs.

10. Be consistent with the presentation of each grade of heading

(see Text below).

Title. The title page should be arranged with the full title; name(s)

of author(s) without academic titles; institutional address(es); sug-

gested running title; address for correspondence.

Synopsis. Each paper should have an abstract not exceeding 200

words. This should summarise the main results and conclusions of

the study, together with such other information to make it suitable

for publication in abstracting journals without change. References

must not be included in the abstract.

Text. All papers should have an Introduction, Acknowledge-

ments (where applicable) and References; Materials and Methods

should be included unless inappropriate. Other major headings are

left to the author’s discretion and the requirements of the paper,

subject to the Editors’ approval. Three levels of text headings and

sub-headings should be followed. All should be ranged left and be in

upper and lower case. Supra-generic systematic headings only

should be in capitals; generic and specific names are to be in italics,

underlined. Authorities for species names should be cited only in the

first instance. Footnotes should be avoided if at all possible.

References. References should be listed alphabetically. Authori-

ties for species names should not be included under References,

unless clarification is relevant. The author’s name, in bold and lower

case except for the initial letter, should immediately be followed by

the date after a single space. Where an author is listed more than

once, the second and subsequent entries should be denoted by a long

dash. These entries should be in date order. Joint authorship papers

follow the entries for the first author and an ‘&’ should be used

instead of ‘and’ to connect joint authors. Journal titles should be

entered in full. Examples: (i) Journals: England, K.W. 1987. Certain

Actinaria (Cnidaria, Anthozoa) from the Red Sea and tropical Indo-

Pacific Ocean. Bulletin of the British Museum (Natural History),

Zoology 53: 206-292. (ii) Books: Jeon, K.W. 1973. The Biology of

Amoeba. 628 p. Academic Press, New York & London. (iii) Articles

from books: Hartman, W.D. 1981. Form and distribution of silica in

sponges. pp. 453-493. In: Simpson, T.L. & Volcani, B.E. (eds)

Silicon and Siliceous Structures in Biological Systems. Springer-

Verlag, New York.

Tables. Each table should be typed on a separate sheet designed to

extend across a single or double column width of a Journal page. It

should have a brief specific title, be self-explanatory and be supple-

mentary to the text. Limited space in the Journal means that only

modest listing of primary data may be accepted. Lengthy material,

such as non-essential locality lists, tables of measurements or details

of mathematical derivations should be deposited in the Biological

Data Collection of the Department of Library Services, The Natural

History Museum, and reference should be made to them in the text.

Illustrations

DRAWINGS — Figures should be designed to go across single (84 mm

wide) or double (174 mm wide) column width of the Journal page,

type area 235 x 174 mm. Drawings should be in black on white stiff

card with a line weight and lettering suitable for the same reduction

throughout, ideally not more than 40%. After reduction the smallest

lettering should be not less than 10 pt (3 mm). Tracing paper should

ideally be avoided because of the possibility of shadows when

scanned. All artwork must have bulletin, author and figure number

included, outside of the image area, and must be free of pencil, glue

or tape marks.

PHOTOGRAPHS — All photographs should be prepared to the final size

of reproduction, mounted upon stiff card and labelled with press-on

lettering (eg Letraset). They can be mounted on white or black

background; a black background must be evenly black all over; any

background must be free of all pencil and glue marks within the

image area. All figures should be numbered consecutively as a

single series. Legends, brief and precise, must indicate scale and

explain symbols and letters. Photos, when components of figure-

plates should be abutted, trimmed as regular rectangles or close

trimmed up to edge of specimen. Joins etc. can be removed at the

scanning stage but at extra cost. Cropping instructions, if any, should

be indicated on an overlay or marked on a photocopy of the figure.

SIZE — Maximum size of artwork for use of flatbed scanners is A3.

Larger artwork has to be reduced photographically prior to scan-

ning, therefore adding to expense.

Symbols in text. Male and female symbols within the text should

be flagged within curly brackets to enable setter to do a swift global

search.

Reprints. 25 reprints will be provided free of charge per paper.

Orders for additional reprints can be submitted to the publisher on

the form provided with the proofs. Later orders cannot be accepted.