New species of epacteriscids (Copepoda, Calanoida) from ...A new plesiomorphic genus with one species is established in this paper, plus one new species in Azygonectes and two in Bofuriella,

ORIGINAL ARTICLE

New species of epacteriscids (Copepoda, Calanoida) from anchialinecaves in the Caicos Islands and the Bahamas

AUDUN FOSSHAGEN1 & THOMAS M. ILIFFE2

1Department of Biology, University of Bergen, P.O. Box 7800, NO-5020, Bergen, Norway, and 2Department of Marine

Biology, Texas A&M University at Galveston, Galveston, Texas, USA

AbstractOne new monotypic genus, Caiconectes , and one new species in each of the genera Azygonectes and Bofuriella are describedfrom the Caicos Islands. In the Bahamas, a third new species of Bofuriella is described from Cat Island and the hithertounknown female of Minnonectes melodactylus Fosshagen & Iliffe is recorded from Great Exuma Island. Caiconectes showsmany plesiomorphic features, with some slight reductions in mouthparts, probably related to raptorial feeding. Swimminglegs show maximum segmentation and setation; leg 1 has long and thin outer setae on the exopod, and the third segment ofthe endopod bears seven setae. The species is considered pelagic. The first male specimen of Azygonectes is described fromA. plumosus sp. nov.; in both sexes, caudal seta V on the left side is approximately 2.5 times body length. The single femalespecimen of Bofuriella spinosa sp. nov. from Cat Island differs from other congeners in having a lateral genital aperture onthe left side, and more setae on the mandibular endopod and on the coxal and basal endites of the maxillule. All cavecalanoids hitherto described from the Caicos Islands are not recorded from any other parts of the Caribbean; thus pointingto an isolated position of the islands.

Key words: Anchialine caves, Bahamas, Calanoida, Epacteriscidae, Taxonomy

Introduction

In Fosshagen et al. (2001), the family Epacterisci-

dae, with a total of 20 species distributed in 12

genera, was reviewed. The family was divided into

two subfamilies and a key to the genera provided.

Since then, five new genera have been described,

Bunderia from Western Australia (Jaume & Hum-

phreys 2001), Iboyella from Cuba (Boxshall & Jaume

2003), and Azygonectes , Cryptonectes and Minno-

nectes from the Bahamas (Fosshagen & Iliffe 2004a),

and a recent record of a second species of Balinella

from the Yucatan Peninsula (Suarez-Morales et al.

2006), bringing the number of described genera to

17 with 26 species. The recent records lead to a new

consideration of the family division and to a new key

to the genera.

Nearly all species of the family are cave-living. The

taxa are extremely disjunct and typically have a

Tethyan distribution and often co-occur with remi-

pedes. The family is one of the most primitive of the

Calanoida and there are few synapomorphies. Im-

portant characteristics include the stout body,

usually with a broad bilobed rostrum with filaments,

the dominance of the exopod and reduction of the

endopod of the mandible, raptorial mouthparts with

a strong gnathobase of the mandible with long sharp

teeth, modified spines of the maxilla and maxilliped,

and swimming legs with three-segmented rami.

Between the genera there are great differences in

the mouthparts, armature of the swimming legs and

in the structure of male leg 5, more than between

many calanoid families.

Most epacteriscids are obtained from net hauls in

anchialine caves in tropical and subtropical waters,

mainly from the Caribbean area, others from Ber-

muda, the Canary Islands, Western Australia, the

Philippines, Palau, Fiji, and the Galapagos Islands.It

is considered that most species are predators or

scavengers. The majority of the genera are mono-

typic, but Enantiosis Barr and Epacteriscus Fosshagen

have seven and three species, respectively. Species of

Correspondence: A. Fosshagen, Department of Biology, University of Bergen, P.O. Box 7800, NO-5020, Bergen, Norway. E-mail:

[email protected]

Published in collaboration with the University of Bergen and the Institute of Marine Research, Norway, and the Marine Biological Laboratory,

University of Copenhagen, Denmark

Marine Biology Research, 2007; 3: 73�92

(Accepted 6 February 2007; Printed 20 April 2007)

ISSN 1745-1000 print/ISSN 1745-1019 online # 2007 Taylor & Francis

DOI: 10.1080/17451000701274571

the last two genera have also been found in cryptic

environments outside caves (Fosshagen 1973; Fos-

shagen et al. 2001) and from emergence traps in

shallow water among corals in the Philippines

(Walter et al. 1982).

One species, Enantiosis belizensis Fosshagen, Box-

shall & Iliffe, has been obtained from baited traps,

from Giant Cave in Belize and also from plankton

hauls in the same cave (Fosshagen et al. 2001).

Other copepods might be possible prey. Jaume &

Humphreys (2001) observed the remains of a

misophrioid in the gut of Bunderia misophaga Jaume

& Humphreys from Western Australia. Few speci-

mens of epacteriscids have usually been obtained, as

may be expected of a predator, and they were often

present when there were great numbers of other

copepods, like cyclopoids, stephids and ridge-

wayiids, in the sample (Fosshagen & Iliffe 1991,

1998, 2003).

A new plesiomorphic genus with one species is

established in this paper, plus one new species in

Azygonectes and two in Bofuriella , and the hitherto

unknown female of Minnonectes melodactylus is

described.

This brings the total of the family to 18 genera and

29 species.

Material and methods

All material was collected from anchialine caves in

the Caicos Islands, except for Bofuriella from Cat

Island and the female of Minnonectes melodactylus

from Great Exuma Island in the Bahamas.

The copepods were mostly obtained using ad-

vanced diving techniques, when dragging a fine-

mesh hand net (ca. 100 mm) through the water. The

terminology used in the descriptions follows Huys &

Boxshall (1991). All type material is kept in the

Natural History Museum, London.

Systematics

Genus Caiconectes gen. nov.

Diagnosis

Female. Prosome in dorsal view widest posterior to

middle, five pedigerous somites well defined. Uro-

some four-segmented, genital double somite elon-

gate with genital aperture located ventrally in

middle of anterior half, anal somite slightly shorter

than preceding somite. Caudal rami asymmetrical

with plumose setae, seta V on left ramus extremely

elongate. Rostrum small with two closely set

filaments. Antennule 27-segmented, elongate with

apical segment incorporating XXVII and XXVIII.

Antenna with exopod and endopod of about the

same length, endopod with well-developed inner

lobe. Mandible with relatively well-developed two-

segmented endopod situated distally on basis;

gnathobase with strong pointed teeth, ventralmost

tooth strongest. Maxillule with five setae on coxal

endite, four on proximal basal endite, and presence

of a seta on basal exite; reduction in number of

setae on distal basal endite and endopod with one,

one, one, and five setae, respectively. Maxilla and

maxilliped well developed, both with long whip-like

setae distally; basal endite of maxilla strong with

five elements and process distally.

Legs 1�5 with three-segmented rami and follow-

ing seta and spine formula:

Male. Urosome five-segmented, second urosomite

longest, first and third urosomites of equal length.

Right antennule weakly geniculated, 24-segmented,

segments II�IV fused. Leg 5 with slightly trans-

formed exopods, elongate on left side where third

segment tapers distally into thin curved process.

Type species

Caiconectes antiquus gen. et sp. nov.

Etymology. The generic name refers to the Caicos

Islands, where the animal was swimming in caves

(from the Greek nectos meaning swimming). Gender

masculine. The specific name refers to the suppo-

sedly ancient and primitive species (from the Latin

antiquus meaning ancient).

Caiconectes antiquus gen. et sp. nov.

Material examined. Cottage Pond, North Caicos

Island, 11 June 2003. One female and one male

collected with a plankton net from the water column

under a ledge in 25�30 m depths. Old Blue Hill

Cave, Providenciales, 12 June 2003. Four CV


at 1�2 m depths. Conch Bar Cave, Middle Caicos

Island, 15 June 2003. One female, one male, one

CV, and one CIV collected with a plankton net from

0.5�15 m depths.

Coxa Basis Exopod segments Endopod segments

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

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

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

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

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

74 A. Fosshagen & T. M. Iliffe

Type material. Holotype. Adult female, total length

2.82 mm from Cottage Pond, 11 June 2003.

Dissected and mounted on three slides. BMNH

2006.365.

Paratypes. Adult male, total length 2.70 mm from

the same locality as the holotype. Dissected and

mounted on three slides (leg 5 lost). BMNH

2006.366. Four CV from Old Blue Hill Cave, 12

June 2003 in one vial. BMNH 2006.367-370. One

female, one male, one CV, and one CIV from Conch

Bar Cave, 15 June 2003 in one vial. BMNH

2006.371-374.

Female. Body length of two specimens both 2.82

mm. All pedigerous somites separate. Body (Figure

1A) with the ratio of prosome length to urosome

length ca. 1.8:1. In dorsal view, body drop-shaped,

broadest across second pedigerous somite. Uro-

some (Figure 1C, D) four-segmented, genital

double somite elongate with genital aperture lo-

cated anteroventrally. Caudal rami asymmetrical,

longer than broad, longest on left side; seta I

minute; seta II long and plumose; seta V on left

side greatly elongate, approximately 1.7 times

length of body; setae IV and V with no fracture

planes. Rostrum (Figure 1B) relatively small as two

closely set lobes each with one filament.

Antennule (Figure 1E) 27-segmented, reaching

beyond caudal rami; segment I with three setae and

one aesthetasc; segment II with two setae; segments

III�XXI with trithek; apical segment incorporating

XXVII and XXVIII with seven elements, of which

tip bearing two long aesthetascs.

Antenna (Figure 1F) with endopod and exopod of

approximately the same length. First endopod seg-

ment with two inner setae located distally; second

segment with inner lobe bearing nine setae, terminal

lobe with seven setae. Exopod with small seta on first

and second segments, terminal segment approxi-

mately one-third length of ramus.

Mandible (Figure 1G) with gnathobase bearing

strong smooth ventralmost tooth separated by a gap

from the other three prominent teeth and seven less

strong teeth of a different shape (in the bottom of the

gap one gnathobase had a rounded stout tooth); all

other teeth pointed. Palp with one seta on basis;

endopod on extended basis, two-segmented with

one seta on first segment and four setae on second

segment.

Maxillule (Figure 2A) with praecoxal arthrite

bearing 15 elements, seven spinous setae, two long

and six short setae; coxal endite with five setae; coxal

epipodite with eight setae, five long and three short;

proximal basal endite with four setae, two long and

two short; distal basal endite represented by short

seta distally; minute seta on outer margin of basis.

Endopod three-segmented, first and second segment

with one inner seta, third segment with five terminal

setae. Exopod with nine setae.

Maxilla (Figure 2B) with relatively short setae on

praecoxal and coxal lobes; first praecoxal lobe

bearing four flexible setae, one short seta and one

spine-like seta; basal endite (Figure 2C) with distal

pointed setose process and five unequal elements,

one conspicuously strong spine, two setae of med-

ium length, two long setae, one of which is whip-like,

bearing modified setules in one row in distal half,

first cup-like then tooth-like setules towards the

distal part; first segment of endopod (Figure 2C)

with four unequal setae, distal segments compressed

bearing seven long whip-like setae all with modified

setules in the distal half.

Maxilliped (Figure 2D) well developed; second

and third lobes of coxa with one seta on each lobe

whip-like and modified like those of maxilla; en-

dopod six-segmented, formula 2, 4, 4, 3, 3�1, 3, of

which 10 setae whip-like and modified.

Leg 1 (Figure 2E): basis with outer seta; exopod

with slender setae along outer margin, terminal

spine with finely serrate outer margin; endopod

with seven setae (two outer ones) on third segment.

Terminal long spine of exopod legs 2�5 without

proximal transverse dividing line (breaking plane?)

commonly found in epacteriscids.

Leg 2 (Figure 2F): basis with outer spine; exopod

with three short spines along outer margin of third

segment; outer margin of first endopodal segment

with three rounded processes.

Leg 3 (Figure 2G): slightly longer than leg 2,

otherwise similar.

Leg 4 (Figure 3A): basis with small outer seta;

third segment of endopod with seven setae.

Leg 5 (Figure 3B): basis with small outer seta;

third segment of exopod attached to middle of inner

margin of preceding segment, outer spines of seg-

ment relatively long.

Male. Body length 2.70 mm. Urosome (Figure 3C)

five-segmented, genital somite approximately three-

quarters length of second urosomite, genital aper-

ture on the left side (spermatophore shown in

figure); caudal rami as in female.

Right antennule (Figure 3D, distal part) 24-

segmented, segments II�IV fused, free segments

17�19 bear strong spine along anterior margin;

distal to geniculation six free segments, of which

five distal ones are identical in both sexes.

Fifth legs (Figure 3E) with outer seta on basis and

three-segmented rami; endopods unmodified; exo-

pods slightly modified with same number of ele-

ments on both sides but with longer segments on left

side; first segment with outer distal spine; second

New epacteriscids 75

segment with outer distal seta, longest on left side, in

addition left side bears row of long setules along

distal inner margin; third segment on left side with

two unequal setae and terminal, long, thin, inward

curved tapering process; right side with two smaller

unequal setae and long terminal straight seta-like

process.

Ecological notes. The species was recorded from three

different caves in the Caicos Islands at Providen-

ciales, North Caicos and Middle Caicos. In Provi-

denciales at 1�2 m depth, only copepodids were

obtained; in North Caicos at 25�30 m, only adults;

in Middle Caicos at 0.5�15 m, both adults and

copepodids were found.

Figure 1. Caiconectes antiquus gen. et sp. nov., female. (A) Habitus, lateral; (B) rostrum; (C) urosome, dorsal; (D) genital double somite,

ventral; (E) antennule; (F) antenna; (G) mandible.


Cottage Pond on North Caicos consists of a

water-filled sinkhole in the interior of the island. A

large crack in 10�15 m depths at the bottom of a

circular open-air pond gives way to an enormous

dome room with water depths to 70 m.

A profile view of Cottage Pond is shown in

Koenemann et al. (2004).

On 9 June 2003, salinity increased from 3 ppt at

the surface through a halocline between 8 and 24 m

to 31 ppt at depth, and the temperature decreased

from 30.88C at the surface through at abrupt

thermocline at about 10 m and then steadily

dropped from 25.08C at 13 m to 22.38C at 65 m.

Dissolved oxygen levels were below 0.5 mg l�1 at

depths greater than 21 m.

Other calanoids apart from epacteriscids observed

in Cottage Pond were a diaptomid, probably a

Mastigodiaptomus species, and a tiny ridgewayiid.

They were collected below the halocline at depths

between 25 and 41 m.

Figure 2. Caiconectes antiquus gen. et sp. nov., female. (A) Maxillule; (B) maxilla; (C) basal endite and first segment of maxilla; (D)

maxilliped; (E) leg 1; (F) leg 2; (G) leg 3.


Stygobitic species inhabiting this cave include the

cirolanid isopod Bahalana caicosana Botosaneanu &

Iliffe, the remipedes Lasionectes entrichoma Yager &

Schram, Godzillius robustus Schram et al. and Kalo-

ketos pilosus Koenemann et al., and the shrimp

Agostocaris williamsi Hart & Manning (Hart & Man-

ning 1986; Schram et al. 1986; Yager & Schram 1986;

Botosaneanu & Iliffe 2003; Koenemann et al. 2004).

Old Blue Hill Cave on Providenciales is a collapse

sinkhole with a 5 m high cliff bordering a semicir-

cular pond. Water depths in the cave reach 25 m. On

12 June 2003, salinity increased steadily from 17 ppt

at the surface to 33 ppt at 22 m, and the temperature

decreased from 29.38C at the surface through an

abrupt thermocline at 4 m to 25.28C at 22 m.

Dissolved oxygen levels were below 0.5 mg l�1 at

Figure 3. Caiconectes antiquus gen. et sp. nov., female (A, B), male (C�E). (A) Leg 4; (B) leg 5; (C) urosome, ventral; (D) distal part of right

antennule; (E) leg 5, posterior.


depths greater than 3 m. Previously described

stygobites from this cave include Lasionectes entri-

choma.

Conch Bar Cave on Middle Caicos is the longest

above sea level cave in the Bahamas archipelago,

with more than 3 km of mapped passageways. The

cave is situated below Village Hill, part of a

Pleistocene dune ridge approximately 500,000 years

old.

Flowstone in the cave has been dated at 200,000

years, establishing a minimum age of the cave.

Conch Bar is described as a flank-margin cave

because it was formed in the mixing zone below

the water table at the margin of a carbonate plat-

form. Passages range from large, dry caverns pre-

viously mined for guano to long partially or mostly

flooded galleries in multilevel sections. This cave is

inhabited by the mysid Stygiomysis clarkei Bowman,

Iliffe & Yager, the polychaete Pelagomacellicephala

iliffei Pettibone, the shrimp Barbouria cubensis

(Von Martens), and the isopod Bahalana caicosana

Botosaneanu & Iliffe. Among the calanoids, Erebo-

nectoides macrochaetus (Fosshagen) and Fosshagenia

ferrarii Suarez-Morales & Iliffe have been described

from this locality (Fosshagen & Iliffe 1994; Suarez-

Morales & Iliffe 1996). Erebonectoides macrochaetus is

also known from Providenciales, but Fosshagenia

ferrarii is only known from Conch Bar Cave.

Caiconectes has the appearance of a typical pelagic

calanoid, with its long antennule and relatively weak

outer spines of the exopods of swimming legs and

the extremely long caudal seta V on the left ramus.

These characters are in contrast to most other

epacteriscids where the antennule is short and the

outer spines of the swimming legs are long and

strong, thus pointing to hyperbenthic habits. Its way

of feeding seems to be raptorial, as indicated by the

pointed teeth of the mandibular gnathobase and its

long specialized setae on the endopods of the maxilla

and maxilliped.

Remarks

The most characteristic feature of the family is the

reduction of the endopod of the mandibular palp

and the dominance of the exopod forming the main

axis of the palp. This character is less expressed in

Caiconectes as the endopod is situated on an exten-

sion on the basis. However, the exopod is the larger

ramus. The genus is not closely related to any of the

17 described genera (Fosshagen et al. 2001; Jaume

& Humphreys 2001; Boxshall & Jaume 2003;

Fosshagen & Iliffe 2004a), nor does it fit into any

of the two subfamilies due to the plesiomorphic

conditions of legs 1 and 2 (see below).

Characters connecting Caiconectes to the family

are: the bilobed rostrum, raptorial mouthparts,

three-segmented swimming legs and fifth legs of

both sexes of a shape similar to that of most species

of the family.

Several plesiomorphic characters are present,

some of them unique to epacteriscids and even to

calanoids. There is no proximal fracture plane on

caudal setae IV and V. This state has not been

reported by Huys & Boxshall (1991). This lack of a

fracture plane may indicate the primitive state of a

plumose caudal seta in a copepod at the base of the

calanoid lineage.

The antennule is distinctly 27-segmented with all

segments separated, bearing three setae plus one

aesthetasc on segment I, with tritheks on segments

XIX and XX, and an elongate apical segment

(XXVII�XXVIII) with seven elements including

two aesthetascs at the tip. According to Huys &

Boxshall (1991) and Boxshall & Huys (1998), the

ancestral apical segment should only possess one

aesthetasc.

The maxillule shows a mixture of plesiomorphic

and apomorphic characters. In the first category, the

coxal endite and the first basal endite bear the

maximum number of setae in epacteriscids, five

and four, respectively; a basal exite with one seta,

only present in Erebonectes ; and a three-segmented

endopod only present in Azygonectes plumosus sp.

nov. (see below).

Apomorphies are reductions in the number of

setae in the praecoxa, second basal endite, exopod

and endopod.

Most epacteriscids have strong spinous setae on

the maxilla and maxilliped, whereas Caiconectes has

long whip-like setae with modified setules. These

setae are reminiscent of setae in the same appen-

dages from the Arietellidae (Ohtsuka & Boxshall

2004). Such modified setae are present on the basal

endite of the maxilla, which bears five elements, one

more than mentioned for the ancestral copepod

(Huys & Boxshall 1991); and on the coxa of the

maxilliped. In misophrioids, Jaume & Boxshall

(1996) described the primitive condition on the

allobasis of the maxilla in Speleophria and Speleo-

phriopsis with setal formula 5,3. Now it may be

considered that the ancestral copepod has the setal

formula 5,4 on the basal endite and proximal

endopodal segment.

The legs show the most plesiomorphic condition

within the family. This is most pronounced in leg 1

bearing thin outer setae on the exopod, and in the

third endopodal segment where there are seven

setae, of which two are outer ones. To our knowl-

edge, this character of the endopod is novel to

calanoids, where the maximum number is six with


one outer seta (Huys & Boxshall 1991). The

presence on leg 2 of three outer spines instead of

two on the third exopodal segment is new to the

family.

In female leg 5, the attachment of the third

exopodal segment is in the middle of the inner

margin of the preceding segment, a character present

in Azygonectes and a key character in the family

Ridgewayiidae. Slightly modified fifth legs of males,

mainly in the third segment of the exopods, like

those of Caiconectes , are present in Enantronoides ,

Gloinella , Minnonectes and Oinella. Generally, this

segment has three elements; one of them may form a

curved process. However, similarities in the fifth legs

of males do not seem to reflect any closer similarities

between the genera in the morphology of other

appendages.

Genus Azygonectes Fosshagen & Iliffe, 2004

Diagnosis (emended)

Female. Prosome with five well-defined pedigerous

somites. Urosome four-segmented with second and

third urosomites of subequal length. Caudal rami

slightly asymmetrical, caudal seta II spinous or

plumose, seta V on left side extremely elongate.

Rostrum elongate with two closely set filaments at

tip. Antennule reaching beyond caudal rami, 26- or

27-segmented depending on fusion or not of seg-

ments XXVI and distal compound segment. An-

tenna with endopod slightly longer than exopod,

second endopodal segment with nine setae on inner

lobe. Gnathobase of mandible with one strong

smooth tooth or two multicuspid teeth ventrally;

palp with two-segmented endopod, bearing one seta

on first segment and four setae on second segment,

and with two to three setae on basis. Maxillule well

developed but with slight reductions in number and

length of setae along inner margin of endopod.

Maxilla and maxilliped rather unmodified, mostly

with long and flexible setae. Leg 1 with long and

slender outer spines on exopod, each with long

filament at tip.

Legs 2 and 3 with two outer spines on third

exopodal segment; leg 3 with stout outer spine on

basis. Legs 4 and 5 with three outer spines on third

exopodal segment; leg 5 with third exopodal seg-

ment offset near middle of inner margin of preceding

segment.

Male. Urosome five-segmented with first four so-

mites of equal length, anal somite approximately half

length of preceding somites. Right antennule geni-

culate, 23-segmented with long pointed process

distally on penultimate segment (XXIV�XXV).

Leg 5 with transformed exopods and unmodified

endopods. Left leg with curved process on inner

margin of basis. Left exopod two-segmented, each

segment bearing outer long curved spine. Right

exopod three-segmented, outer margin of second

segment with two irregular elements, third segment

tapers into long flexible tip.

Azygonectes plumosus sp. nov.

Etymology. The specific name alludes to the long and

feathery setae on the caudal rami (from the Latin

plumosus meaning feathery).

Material examined. Cottage Pond, North Caicos

Island, 9 June 2003. One female collected with a

plankton net from the water column at 25�46 m

depths. Cottage Pond, 11 June 2003. One male


under a ledge in 25�30 m depths. Old Blue Hill

Cave, Providenciales, 12 June 2003. One female


at 1�2 m depths.

Type material. Holotype. Adult male, total length

1.57 mm from Cottage Pond, 11 June 2003.

Dissected and mounted on four slides. BMNH

2006.375.

Paratypes. Adult female, total length 1.57 mm

from Cottage Pond, 9 June 2003. Slightly damaged.


2006.376. Adult female, 1.65 mm, from Old Blue

Hill Cave, 12 June 2003 in one vial. BMNH

2006.377.

Male. Body length of one male 1.57 mm. Body

(Figure 4A) with a ratio of prosome to urosome ca.

2.8:1. Urosome (Figure 4C) five-segmented; uroso-

mites 1�4 of subequal length, anal somite approxi-

mately half length of preceding somite. Caudal rami

asymmetrical, longest on the left side with seta V

extremely elongate to approximately 2.5 times body

length; seta II plumose.

Rostrum (Figure 4B) pointed with two closely set

filaments at the tip.

Antennule reaching slightly beyond caudal rami,

geniculate and 23-segmented on the right side and

26-segmented on the left side. Right antennule

(Figure 4D) with segments II�IV separate, seg-

ments 20�22 (XX�XXV), in connection with

geniculation, relatively elongate; segment 22

(XXIV�XXV) with anterodistal corner bearing a

slender pointed process reaching beyond the apex

of the limb, and posterodistal corner bearing seta

with conspicuously long setules. Left antennule

(Figure 4E, distal part) like those of females in


Balinella , Bofuriella , and Bomburiella , all 26-seg-

mented with different degrees of incomplete fusion

between segment XXVI and the compound term-

inal segment.

Antenna (Figure 5A) with endopod slightly longer

than exopod, otherwise as in A. intermedius .

Mandible (Figure 5B) differs from A. intermedius

in bearing one strong smooth ventral tooth and a

Figure 4. Azygonectes plumosus sp. nov., male. (A) Habitus, lateral; (B) rostrum; (C) urosome, dorsal; (D) right antennule; (E) distal part of

left antennule.


relatively long dorsal seta on the gnathobase. Palp

with two setae on the basis.

Maxillule (Figure 5C) with more setae than A.

intermedius. Coxal endite and basal endites with

three, four and five setae, respectively; endopod

three-segmented, partly fused distal segments, four

setae on first and second segments, seven setae on

terminal segment; exopod with 11 setae.

Maxilla (Figure 5D) and maxilliped (Figure 5E)

generally as in A. intermedius.

Leg 1 (Figure 6A) with a minute seta on the outer

margin of the basis; outer spines of the exopod with

a long flagellum and of equal length.

Legs 2�4 (Figure 6B�D) with the same seta and

spine formula as in A. intermedius. The outer distal

margin of the second endopodal segment ending in

two pointed processes in the new species.

Leg 5 (Figure 6E, F) with unmodified three-

segmented endopods and modified exopods, three-

segmented on the right side and two-segmented on

Figure 5. Azygonectes plumosus sp. nov., male. (A) Antenna; (B) mandible; (C) maxillule; (D) maxilla; (E) maxilliped.


the left side. The first segment of the right exopod

with a strong outer spine; the second segment with

two strong, curved, irregular outer elements; the

third segment with a spine on the proximal anterior

side, an outer pointed process, and a distal element

gradually tapering into a thin flexible tip. Left leg

with a curved process on the inner margin of the

basis; the first segment of the exopod bearing a

long curved outer spine reaching the end of the

endopod; the second segment broad proximally,

tapering irregularly into a pointed tip, bearing an

outer strong process, with a notch proximally on

Figure 6. Azygonectes plumosus sp. nov., male. (A) Leg 1; (B) leg 2; (C) leg 3; (D) leg 4; (E) left leg 5, posterior; (F) right leg 5, posterior.


the outer margin, the shape and length as in the

outer spine on the first segment; further distally on

the outer margin of the second segment seta

present.

Female. Body length of two females 1.57 and

1.65 mm. Differs from male in urosome, right

antennule and leg 5.

Urosome (Figure 7A) four-segmented with a

genital aperture located posteroventrally on the

genital double somite. Seta IV on caudal rami

thickened, abruptly tapering distally; seta V on the

left side, as in male, extremely long.

Antennule 26-segmented, as on the left side of the

male.

Leg 5 (Figure 7B) as in A. intermedius, but lacking

the outer seta on the basis and the inner seta on the

first exopodal segment.

Ecological notes. One female and one male were

recorded at Cottage Pond, North Caicos on separate

dates in June 2003 at depths between 25 and 46 m in

clear seawater below the halocline, and one female at

Old Blue Hill Cave, Providenciales at 1�2 m depths

at salinity about 20 ppt (for cave descriptions

and water quality see Caiconectes). At Old Blue

Figure 7. Azygonectes plumosus sp. nov., female (A, B). (A) Urosome, dorsal; (B) leg 5. Bofuriella spinosa sp. nov., female (C�I). (C)

Urosome, dorsal; (D) antenna; (E) mandibular palp; (F) mandibular coxa with gnathobase; (G) maxillule; (H) leg 1; (I) leg 5.


Hill, the oxygen levels below 3 m depth were less

than 0.5 mg l�1. This might have been a limiting

factor for the vertical distribution of the animals.

In both caves, small cyclopoids dominated; a few

specimens of Caiconectes antiquus were also obtained.

In Cottage Pond, several specimens of a diaptomid,

probably a Mastigodiaptomus species, and a tiny

ridgewayiid were recorded.

Remarks

Only one species, A. intermedius Fosshagen & Iliffe

described from two female specimens, is known in

the genus. It was recorded from South Andros Island

and Great Exuma Island, Bahamas (Fosshagen &

Iliffe 2004a). The new species is known from both

sexes and compared with A. intermedius.

The two species of the genus are recorded on two

banks separated by deep water, A. intermedius on

Great Bahama Bank and A. plumosus sp. nov. on

Caicos Bank. In A. intermedius , only the female is

known.

The two species are most readily distinguished by

caudal seta II, in A. plumosus sp. nov. long and

plumose, and in A. intermedius short and spinous;

one strong and smooth ventral tooth is present on

the mandibular gnathobase in A. plumosus sp. nov.,

and two strong ventral teeth with distal projections

are present in A. intermedius ; furthermore, the

endopod of the maxillule is elongate and in a more

plesiomorphic state than in A. intermedius , three-

segmented and equipped with more setae; the distal

basal endite carries five setae and the exopod 11

setae, conditions that are among the most plesio-

morphic in calanoids.

A characteristic for both species is the extremely

elongate seta V on the left caudal ramus.

Genus Bofuriella Fosshagen, Boxshall & Iliffe, 2001

Diagnosis (emended)

Female. Prosome oval in dorsal view, widest anterior

to middle; separation of cephalosome and first

pedigerous somite not clearly visible. Urosome

four-segmented, with genital aperture ventral or

lateral on left side; third and fourth urosomites of

subequal length. Caudal rami with seta II spiniform,

seta V of subequal length on both rami; seta VI on

left ramus ornamented with tuft of setules proxi-

mally. Rostral lobes weakly developed, each with one

filament. Antennule 26-segmented with segment

XXVI incorporated into apical double segment

XXVII�XXVIII. Endopod of antenna approxi-

mately two-thirds length of exopod and with well-

developed inner lobe. Mandibular palp with small,

two-segmented endopod carrying two to five setae;

gnathobase with strong, bicuspid ventralmost tooth.

Maxillule with coxal and basal endites each armed

with two to three unequal setae; coxal epipodite with

five long setae. Maxilla with well-developed endites

on praecoxa and coxa; endopod compressed as

typical for family, armed with very strong spines

distally. Maxilliped strongly developed, endopod

short, carrying claw-like spines on endopod, one or

two of which are coarsely serrated. Legs 1�5

generally with same segmentation and setation as

in Bomburiella , sometimes with reduced length of

outer spines on second and third segments of exopod

leg 1.

Bofuriella spinosa sp. nov.

Etymology. The specific name alludes to the long

outer spines on leg 1 in contrast to B. vorata , which

has shorter spines (from the Latin spinosus meaning

thorny).

Material examined. Conch Bar Cave, Conch Bar,

Middle Caicos Island, 15 June 2003.

One female obtained from the water column in

0.5�15 m depth collected with a plankton net.

Type material. Holotype. Adult female, body length

1.45 mm from Conch Bar Cave. Dissected and

mounted on three slides. BMNH 2006.378.

Female. Cephalosome and first pedigerous somite

apparently separate.

Urosome (Figure 7C) four-segmented; genital

double somite slightly produced on the left side

with the genital aperture located laterally on the

same side. Seta VI on the left caudal ramus more

densely covered with setules proximally than the

corresponding seta on the right side.

Rostrum with small, rounded, closely set lobes at

the tip, each with a small filament.

Antennule 26-segmented and very similar to that

of B. vorata.

Antenna (Figure 7D) with one minute seta on the

first segment of the exopod and one medium-sized

seta on the second segment; the inner lobe of the

endopod with seven setae.

Mandible (Figure 7E, F) with a two-segmented

small endopod; one small seta on the first segment,

two long and two small setae on the second segment.

Maxillule (Figure 7G) with a praecoxal arthrite

bearing 13 spinous or flexible setae; the coxal endite

with one strong seta, one medium-sized seta and

one small seta; the proximal basal endite with two

long setae and one small seta.


Maxilliped with one small seta on the praecoxal

endite; weaker spines on the endopod than in B.

vorata ; only one spinulate claw present, this origi-

nates on the third free segment of the endopod;

other strong spines on the endopod with fine

spinules along the concave margin.

Leg 1 (Figure 7H) with a small outer seta on the

basis, the outer spines on the second and third

exopodal segments relatively slender, the outer distal

corner of the first endopodal segment bifid with a

rounded and pointed process terminally.

Legs 2�5 with the inner distal corner of the basis

produced into a pointed tip, contrary to the rounded

corner in B. vorata.

Leg 5 (Figure 7I) lacking the prominent process

between the second and third outer spines of the

third exopodal segment present in B. vorata ; the

terminal spine on the third exopodal segment is

twice the length of the third outer spine.

Ecological notes. Conch Bar Cave at Middle Caicos is

the largest and most significant cave in the Turks and

Caicos group (Gregor 1981). Specimens were col-

lected during a plankton tow in shallow (0.5 m deep)

pools and entirely submerged galleries (to 15 m

depth). Both the previously described calanoids,

Erebonectoides macrochaetus and Fosshagenia ferrarii ,

were rediscovered in the June 2003 sample, together

with B. spinosa sp. nov. and the new genus Caico-

nectes described in this paper.

Remarks

Only one species, Bofuriella vorata Fosshagen, Box-

shall & Iliffe, from Oven Rock Cave, Great Guana

Cay (Fosshagen et al. 2001) and Sanctuary Blue

Hole, South Andros, Bahamas (Fosshagen & Iliffe

2004a) is known in the genus.

The new species is compared with the female of B.

vorata . The single specimen was broken dorsally

along the dividing line between the cephalosome and

the first pedigerous somite, and the genital double

somite was infested by several protozoans.

The female of the new species is most readily

distinguished from B. vorata by its lateral genital

aperture, its higher number of setae on the endopod

of the mandible, the larger outer spines on the

exopod of leg 1, and a pointed inner corner of the

basis of legs 2�5.

It is somewhat surprising to find closely related

species with different locations of genital aperture;

ventral in B. vorata and lateral on the left side in B.

spinosa sp. nov. Such a lateral position on the left

side is present in the monotypic epacteriscid genus

Bomburiella , on the right side in the ridgewayiid

genus Exumella (Fosshagen 1970; Fosshagen et al.

2001), and ventrally on the left side in Ridgewayia

stygica (Ohtsuka et al. 2000). The midventral

position is most common among calanoids and

considered ancestral (Huys & Boxshall 1991).

The consensus tree from the phylogenetic analysis

by Fosshagen et al. (2001) shows that Bofuriella and

Bomburiella are closely related. Females of the two

genera have nearly identical antennule and antenna,

and legs 1�5 with the same spine and seta formula.

Slight differences are present in the mandibular

teeth, and in the number of setae on the coxal

epipodite of the maxillule. However, the maxilla and

maxilliped show different structures. The endopod

of the maxilliped in Bofuriella has strong claw-like

spines, whereas Bomburiella has long flexible setae in

same position. This may indicate differences in

feeding habits and prey.

Bofuriella paravorata sp. nov.

Etymology. The specific name alludes to the similar-

ity to B. vorata (the Latin voratus meaning the

devouring one) and referring to the very strong

spines on the maxilliped.

Material examined. Big Fountain, Orange Creek, Cat

Island, Bahamas, 12 and 18 August 2004. Speci-

mens were obtained with a plankton net and

individual vials from the water column in 30�45 m

depths below a hydrogen sulphide layer. 12 August:

two males, one female, one copepodid; 18 August:

one male and two females.

Type material. Holotype. Adult male, body length

1.73 mm from Big Fountain, 12 August 2004.


2006.379.

Paratypes. One adult female, body length 1.53

mm, from Big Fountain, 18 August 2004. Dissected

and mounted on three slides. BMNH 2006.380-

382. Two males and two females in one vial from the

same locality. BMNH 2006.383-384.

Female. Very similar to B. vorata. Cephalosome and

first pedigerous somite with a distinct dividing line.

Genital double somite (Figure 8A) broadest in the

posterior half and apparently more produced ven-

trally and more posterior than in B. vorata. One

specimen was regarded as abnormal with one leg 1

bearing one inner seta on the second segment of the

endopod and one leg 2 with four inner setae on the

third segment of the exopod and seven setae on the

third segment of the endopod. All other appendages

without any noticeable differences from those of B.

vorata.


Male. Right antennule 21-segmented (Figure 8B),

with a pointed process on the anterodistal corner of

segment 20 shorter than in B. vorata and reaching

approximately one-third along the length of segment

21. At geniculation, segment 18 bears one long spine

on the anterior margin, serrated on the outer margin

(in B. vorata this serration is suggested to be along

the margin of the antennule, but this is probably

wrong; Fosshagen et al. 2001) and one distal seta;

segment 19 bears two shorter similarly serrated

Figure 8. Bofuriella paravorata sp. nov., female (A), male (B�D). (A) Urosome, dorsal; (B) right antennule; (C) left leg 5, anterior; (D)

right leg 5, anterior. Minnonectes melodactylus Fosshagen & Iliffe, 2004, female (E�H). (E) Habitus, lateral; (F) caudal rami, dorsal; (G)

antennule; (H) leg 5.


spines, one aesthetasc in the middle and one distal

seta. The left antennule is similar to the female

except that segment XIX bears two setae of unequal

length where in the female they are equal.

The fifth legs, in particular the left one, differ

slightly from B. vorata. Left leg (Figure 8C) with the

basis less pronounced at the inner corner in B.

paravorata than in B. vorata , and the third segment

of the exopod with a pointed process inserted distally

on the outer margin, and the third segment of the

endopod with more (ca. 12) unequal flattened

processes on its anterior surface than B. vorata ;

right leg (Figure 8D) apparently similar to that of B.

vorata.

Ecological notes. Big Fountain at Cat Island is an

open sinkhole pool extending down into a more than

60 m deep cave with a strong hydrogen sulphide

layer at 5�10 m. Most animals were collected

around 40 m depth where dissolved oxygen levels

slowly begin to recover. Of other copepods present

were large numbers of a misophriid and a few

specimens of Bomburiella gigas. It is not unlikely

that the epacteriscids were preying on the miso-

phriids, as observed in Bunderia misophaga by Jaume

& Humphreys (2001). At least four species of

remipedes have been collected from this cave.

Remarks

This is the third species in the genus. It is compared

with the closely related B. vorata from Oven Rock

Cave, Great Guana Cay, Bahamas (Fosshagen et al.

2001).

Genus Minnonectes Fosshagen & Iliffe, 2004

Diagnosis

Female. Urosome four-segmented with very short

anal somite. Caudal seta VI on left side with dense

unilateral tuft of setules near base. Antennule 27-

segmented with partial fusion of segments I�IV on

posterior side. Leg 5 with three-segmented rami,

short bulbous inner seta on first segment of exopod.

Minnonectes melodactylus Fosshagen & Iliffe, 2004

Material examined. Only one male specimen has

hitherto been obtained from Basil Minn’s Blue

Hole, Great Exuma Island, Exuma Cays, Bahamas,

18 March 2000 (Fosshagen & Iliffe 2004a). Now,

two female specimens have been collected from the

same cave on 13 January 2003 with a suction bottle

from 15�40 m depth of the dome room. One female

in vial BMNH 2006.385. One female 1.77 mm on

three slides BMNH 2006.386.

Female. Total body length 1.76 and 1.77 mm. Differs

from the male in the urosome, antennule and leg 5.

Body (Figure 8E) with prosome length to urosome

length ca. 2:1.

Urosome four-segmented, genital double somite

slightly produced midventrally, anal somite much

reduced. Caudal rami (Figure 8F) slightly asymme-

trical with finely plumose setae, seta VI on the left

side with a unilateral tuft of fine setules near the

base.Antennule (Figure 8G) 27-segmented with

segments I�IV incompletely separated on the poster-

ior side; segments XIX and XX each possessing

two setae; segments XXVI and XXVII�XXVIII

separate.

Mouthparts as in the male; strong distal spinous

setae on the maxilla and maxilliped apparently not as

distinctly black pigmented at the tips as in the male.

Swimming legs 1�4 as in the male.

Leg 5 (Figure 8H) with the inner distal corner of

the basis extended into a pointed process, a short

bottle-shaped seta on the inner corner of the first

exopodal segment.

Ecological notes. Basil Minn’s Blue Hole is a wholly

submerged cave that crosses beneath Great Exuma

Island from northeast to southwest. Further descrip-

tions of the cave and associated animals are found in

Koenemann et al. (2003) and Fosshagen & Iliffe

(2004a). Three epacteriscids are present; in addition

to M. melodactylus only found here, Cryptonectes

brachyceratus is also recorded from Acklins Island,

Norman’s Pond Cay, Exumas and from Sweetings

Cay, Grand Bahama Island and finally Azygonectes

intermedius also from South Andros Island. Other

copepods include the ridgewayiids Stargatia palmeri

and Ridgewayia spp.; unidentified species of Para-

misophria and Pseudocyclops (Fosshagen & Iliffe

2004a). The co-occurrence in this cave of three

sympatric remipedes in the genus Speleonectes is

remarkable (Koenemann et al. 2003). As pointed

out by Fosshagen et al. (2001), epacteriscids and

remipedes do often seem to co-occur.

Remarks

When the single male of M. melodactylus was

described (Fosshagen & Iliffe 2004a), similarities

to the female of Enantronia canariensis , only known

from one sex, were suggested. Now that both

females are known, further similarities may be

pointed out.

In both species, left caudal seta VI bears a

proximal tuft of setules; the antennule has similar


modified setae in the proximal part and the two

distal free segments are of the same shape; the

ventralmost tooth of the mandibular gnathobase is

prominent, bifid and rounded. Differences are pre-

sent in the stronger and modified distal claws on the

maxilla and maxilliped of Minnonectes , and in a

strong outer spine on the basis of leg 3 instead of a

small seta in Enantronia. These apparently small

differences may indicate that the two species are

congeneric, but the answer has to await the discovery

of the male of Enantronia.

The modified tips of claws on the maxillipeds of

Minnonectes and Bunderia , although slightly differ-

ent, may indicate a relationship or a similar way of

feeding.

Leg 5 of the Minnonectes male is reminiscent of

several genera in the family, such as Enantronoides ,

Bunderia , Oinella and Gloinella , where the exopods

have slight modifications in the third segment.

Concluding remarks

The four species of cave-living epacteriscids de-

scribed from the Caicos Islands are unique to the

Bahamas archipelago; one belonging to each of the

genera Caiconectes gen. n., Erebonectoides , Azygo-

nectes and Bofuriella.

Caiconectes gen. nov. and Erebonectoides are mono-

typic, a second species of Azygonectes is previously

recorded from South Andros Island and the Exu-

mas, Bahamas, and two species of Bofuriella are

recorded from the Exumas and Cat Island, Bahamas

(Fosshagen & Iliffe 1994, 2004a; Fosshagen et al.

2001).

Another cave-living calanoid from the Caicos

Islands belonging to the family Fosshageniidae is

Fosshagenia ferrarii (Suarez-Morales & Iliffe 1996);

recently, a second species of the genus was recorded

from Grand Bahama Island (Fosshagen & Iliffe

2004b).

This may indicate that the Caicos Islands, sur-

rounded by deep water, have been isolated from

other islands in the Bahamas and an endemic cave

fauna has developed.

There are still unidentified cave calanoids from

the islands and new species in the Ridgewayiidae and

Diaptomidae will probably be recorded.

Previously, only two genera in the Epacteriscidae,

Epacteriscus and Enantiosis , were polytypic, with

three and seven species, respectively (Fosshagen

et al. 2001). Now with the addition of Azygonectes ,

Bofuriella and a second species of Balinella , five

genera are polytypic, but still 13 genera are mono-

typic.

The Epacteriscidae is a heterogeneous family of

cave-living calanoids with few synapomorphies, but

with several plesiomorphic characters. Its members

show several modifications of the mandible, which

speak for predatory habits. In most species, the

gnathobase is strongly developed with prominent

and sharp teeth. This is most clearly expressed in

Epacteriscus , with its enormous scythe-like gnatho-

base extending ventrally.

Further support for predatory habits is present

with the development of strong and modified spines

in the maxilla and maxilliped of Bofuriella , Oinella

and Minnonectes , and in the long whip-like setae

distally on the maxilliped of Bomburiella .

A phylogenetic analysis of the family (Fosshagen

et al. 2001) showed that a key synapomorphy was in

the mandible with the dominance of the exopod and

the reduction of the endopod. At that time, the

family was divided into two subfamilies, Erebonec-

tinae and Epacteriscinae; the first one containing the

two genera Erebonectes and Erebonectoides and the

second one all other remaining genera. Erebonecti-

nae has two outer spines on the third exopodal

segment of swimming legs 3 and 4, whereas Epac-

teriscinae has three spines at the same site.

A key to the genera provided by Boxshall & Halsey

(2004) was based on this subdivision of the family,

but since that time, the four genera Azygonectes ,

Caiconectes , Cryptonectes and Minnonectes have been

added. In the armature of the swimming legs, in

particular the number of outer spines of the third

exopodal segment, Caiconectes , Azygonectes and

Cryptonectes are distinctly different from previously

known genera. Now as Azygonectes bears two outer

spines on leg 3 and three outer spines on leg 4,

Caiconectes has three outer spines on legs 2�4 and

Cryptonectes two outer spines on the same legs; a

revision of the family as well as a new key to the

genera has to be provided.

A new key based mainly on the armature of the

swimming legs still seems to be workable, but a

revision of the family is pending a new phylogenetic

analysis. A division into new families may also be a

possibility.

A key based on the male leg 5 is problematic

because of its complex structure in many genera and

because homologous characters are not easily seen.

Generally, the rami are three-segmented, often with

highly modified exopods and relatively unmodified

endopods. Most complex exopods are present in

Enantiosis with its seven known species, and in

Erebonectes , Erebonectoides , Bomburiella , Bofuriella

and Azygonectes , whereas the other genera have a

simpler or more uniform structure. In Enantiosis , the

females are very similar, but the males show great

variations in their highly complex leg 5 (Fosshagen

et al. 2001).


Calanoids are predominantly pelagic in open

water and adapted for floating, often with a long

antennule and a transparent body. In some families,

such as the Aetididae and Arietellidae, several

members are more or less associated with the bottom

and live an epibenthic life (Boxshall & Halsey 2004).

Typical benthic families are the Ridgewayiidae and

Pseudocyclopidae.

Species associated with the bottom are often

robust, have short antennules and strong outer

spines on the exopods of swimming legs.

Epacteriscids seem to have a wide ecological

separation in caves, some are pelagic or benthic,

other species may show adaptations to both environ-

ments; the last feature indicated by species of

Enantiosis having various lengths of the antennules.

Species of Caiconectes and Azygonectes are con-

sidered pelagic; both with a delicate and transparent

body, a long antennule, one extremely long seta on

the left caudal ramus in both sexes and rather weakly

developed outer spines on the exopods of swimming

legs. The spine and seta formula of the legs differs

between the two genera; the most plesiomorphic

condition is present in Caiconectes , being also the

most primitive among calanoids.

A life close to the bottom is indicated by species of

Epacteriscus , Cryptonectes , Bomburiella , Bofuriella

and Bunderia , having compact bodies, relatively

short antennules, and strong outer spines of the

swimming legs. Also here, the spine and seta formula

of legs shows much variation between the genera.

According to the previous division into subfamilies

based on swimming legs, Epacteriscus and Crypto-

nectes should belong to different taxa (Fosshagen et

al. 2001). A classification based mainly on the

structure of mouthparts rather than on legs seems

at this moment to be a more suitable one for this

family of predators.

Acknowledgements

This research was funded by grants from NOAA’s

Caribbean Marine Research Center and NSF’s

Biodiversity Surveys and Inventories Program

(NFS # 0315903) to T. Iliffe. During the 2003

Caicos Islands expedition, Texas A&M University

graduate student Darcy Gibbons and Caicos Caves

Project divers John Garvin, James Hurley and Mark

Parrish provided assistance with cave collections.

Cave diver Gregg Stanton helped with diving and

sorting specimens from Cat Island caves in 2004.

Team members during the 2003 expedition to the

Exuma Cays included Fernando Alvarez (Universi-

dad Nacional Autonoma de Mexico); Todd Haney

and Leslie Harris (National History Museum of Los

Angeles County); Texas A&M graduate students

Rebecca Belcher, Darcy Gibbons, Lara Hinderstein

and Colin Kliewer; University of Louisiana at

Lafayette graduate student Joris van der Ham and

expert cave divers Brian Kakuk (Bahamas Caves

Foundation) and Curt Bowen (Advanced Diver

Magazine). Their assistance with this fieldwork is

greatly appreciated. The collection of biological

specimens from caves within the Bahamas was

carried out under a marine resource collecting

permit issued by the Bahamas Department of Fish-

eries. We would like to thank the reviewers for their

comments.

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Editorial responsibility: Matz Berggren

Appendix

Key to the genera of Epacteriscidae

1 Legs 2�4 with three outer spines on the third exopodal segment; leg 1 with seven

setae on the third endopodal segment. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caiconectes

These characters not combined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2 Leg 3 with two outer spines on the third exopodal segment and leg 4 with three

outer spines on the third exopodal segment. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Azygonectes


3 Legs 3 and 4 with two outer spines on the third exopodal segment . . . . . . . . . . . 4

Legs 3 and 4 with three outer spines on the third exopodal segment . . . . . . . . . . 6

4 Antenna with the endopod about twice the length of the exopod; leg 2 with a strong

outer spine on the basis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Erebonectes

Leg 2 with an unarmed basis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

5 Antennule reaching beyond the prosome; the endopod of the antenna markedly longer

than the exopod. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Erebonectoides

Antennule approximately half the length of the prosome; the endopod of the antenna

approximately half the length of the exopod . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cryptonectes

6 Gnathobase of the mandible projecting frontally, with the tip modified into a coarse

rake-like blade; labrum elongate, tapering to a single point . . . . . . . . . . . . . . . . . . . . . . . .Epacteriscus

Gnathobase of the mandible not projecting frontally, and not of this form; labrum not

as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

7 Antenna with the endopod just longer than the exopod; mandibular palp with two setae

on the basis, the endopod clearly two-segmented with setal formula 1,4. . . . . . . . . . . . . . . . . Balinella


8 Antennule with a digitiform process on segment IX; mandibular palp lacking an endopod,

the exopod with one extremely long seta distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oinella


9 Articulation in the non-geniculate antennule between segments II and III not expressed;

the exopod of the antenna slightly longer than the endopod . . . . . . . . . . . . . . . . 10

Antenna with the exopod approximately twice as long as the endopod . . . . . . . . . 11

10 Mandibular palp lacking an endopod; maxillule with five setae on the

exopod . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Enantronoides

Mandibular palp with a one-segmented endopod; maxillule with eight setae on the

exopod . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Enantronia


11 Rostrum weakly developed as a broad plate with short filaments; articulation between

antennular ancestral segments XXVI and double segment XXVII�XXVIII not

expressed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12


12 Maxilliped with reduced enditic setation on the syncoxa, basis elongate with a group

of three setae distally, endopod with long flexible setae. . . . . . . . . . . . . . . . . . . . . . . . . . .Bomburiella

Maxilliped of a different shape . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

13 Maxilliped strongly developed, basis compressed, endopod with strong spinulate

claws . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bofuriella

Maxilliped slender, not particularly modified. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Iboyella

14 Rostrum bifurcate with widely separated tips. . . . . . . . . . . . . . . . . . . . . . . . . . . 15

Rostrum elongate with two closely set lobes bearing a pair of thick

filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gloinella

15 Strongly modified legs 5 in the male, particularly on the left exopod and endopod;

first basal endite of the maxillule with one seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Enantiosis

Legs 5 in the male with a moderately modified third segment of the exopods,

endopods unmodified; the first basal endite of the maxillule with more

than one seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

16 Proximal segments (I�VII) of the antennule without modified setae; anal somite

concealed beneath the posterior rim of the preceding urosomal somite. . . . . . . . . . . . . . . . . Edaxiella

Proximal segments (I�VII) of the antennule with modified setae, flattened proximally

and with a filament at the distal part; anal somite not concealed by

the preceding somite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

17 Antennular segments II�IV fused; mandibular palp lacking an endopod. . . . . . . . . . . . . . . . .Bunderia

Antennular segments II�IV partially separated; mandibular palp with a

one-segmented endopod bearing two setae; strong setae on the maxilla and

maxilliped with black-pigmented tips . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Minnonectes


Documents

New species of epacteriscids (Copepoda, Calanoida) from ...A new plesiomorphic genus with one species is established in this paper, plus one new species in Azygonectes and two in Bofuriella,