Neoteny and the Plesiomorphic Condition of the Plesiosaur ...
New species of epacteriscids (Copepoda, Calanoida) from ...A new plesiomorphic genus with one...
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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:
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
collected with a plankton net from the water column
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.
76 A. Fosshagen & T. M. Iliffe
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.
New epacteriscids 77
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.
78 A. Fosshagen & T. M. Iliffe
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
New epacteriscids 79
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
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. One female
collected with a plankton net from the water column
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.
Dissected and mounted on three slides. BMNH
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
80 A. Fosshagen & T. M. Iliffe
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.
New epacteriscids 81
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.
82 A. Fosshagen & T. M. Iliffe
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.
New epacteriscids 83
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.
84 A. Fosshagen & T. M. Iliffe
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.
New epacteriscids 85
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.
Dissected and mounted on three slides. BMNH
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.
86 A. Fosshagen & T. M. Iliffe
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.
New epacteriscids 87
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
88 A. Fosshagen & T. M. Iliffe
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).
New epacteriscids 89
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.
References
Botosaneanu L, Iliffe TM. 2003. A new species of the stygobitic
cirolanid genus Bahalana from the Caicos Islands in the
Caribbean (Isopoda: Cirolanidae). Travaux du Museum d’His-
toire Naturelle ‘‘Grigore Antipa’’ 45:83�93.
Boxshall GA, Halsey SH. 2004. An Introduction to Copepod
Diversity. London: The Ray Society.
Boxshall GA, Huys R. 1998. The ontogeny and phylogeny of
copepod antennules. Philosophical Transactions. The Royal
Society of London B 353:765�786.
Boxshall GA, Jaume D. 2003. Iboyella a new genus of epacteriscid
copepod (Copepoda: Calanoida: Epacteriscidae) from Cuba.
Organisms, Diversity and Evolution 3:85�92.
Fosshagen A. 1970. Marine biological investigations in the
Bahamas. 15. Ridgewayia (Copepoda, Calanoida) and two
new genera from the Bahamas. Sarsia 44:25�58.
Fosshagen A. 1973. A new genus and species of bottom-living
calanoid (Copepoda) from Florida and Colombia. Sarsia
52:145�54.
Fosshagen A, Boxshall GA, Iliffe TM. 2001. The Epacteriscidae,
a cave-living family of calanoid copepods. Sarsia 86:245�318.
Fosshagen A, Iliffe TM. 1991. A new genus of calanoid copepod
from an anchialine cave in Belize. Bulletin of the Plankton
Society of Japan Special Volume (1991):339�46.
Fosshagen A, Iliffe TM. 1994. A new species of Erebonectes
(Copepoda, Calanoida) from marine caves on Caicos Islands,
West Indies. Hydrobiologia 292/293:17�22.
Fosshagen A, Iliffe TM. 1998. A new genus of the Ridgewayiidae
(Copepoda, Calanoida) from an anchialine cave in the Baha-
mas. Journal of Marine Systems 15:373�80.
Fosshagen A, Iliffe TM. 2003. Three new genera of the
Ridgewayiidae (Copepoda, Calanoida) from anchialine caves
in the Bahamas. Sarsia 88:16�35.
Fosshagen A, Iliffe TM. 2004a. New epacteriscids (Copepoda,
Calanoida) from anchialine caves in the Bahamas. Sarsia
89:117�36.
Fosshagen A, Iliffe TM. 2004b. A new species of cave-living
calanoid copepod from Grand Bahama. Sarsia 89:346�54.
Gregor VA. 1981. Karst and caves in Turks and Caicos Islands.
In: Proceedings of the Eighth International Congress of
Speleology. Bowling Green, Kentucky. p 805�7.
Hart CW, Manning RB. 1986. Two new shrimps (Procaridae and
Agostocarididae, new family) from marine caves of the western
North Atlantic. Journal of Crustacean Biology 6:408�16.
Huys R, Boxshall GA. 1991. Copepod Evolution. London: The
Ray Society.
Jaume D, Boxshall GA. 1996. A new genus and two new species of
cave-dwelling misophrioid copepods from the Balearic Islands
(Mediterranean). Journal of Natural History 30:989�1006.
90 A. Fosshagen & T. M. Iliffe
Jaume D, Humphreys WF. 2001. A new genus of epacteriscid
calanoid copepod from an anchialine sinkhole on northwestern
Australia. Journal of Crustacean Biology 21:157�69.
Koenemann S, Iliffe TM, van der Ham J. 2003. Three new
sympatric species of Remipedia (Crustacea) from Great Exuma
Island, Bahamas Islands. Contributions to Zoology 72:227�52.
Koenemann S, Iliffe TM, Yager J. 2004. Kaloketos pilosus , a new
genus and species of Remipedia (Crustacea) from Turks and
Caicos Islands. Zootaxa 618:1�12.
Ohtsuka S, Boxshall GA. 2004. A new species of the deep-sea
copepod genus Scutogerulus (Calanoida: Arietellidae) from
hyperbenthic waters of Okinawa, Japan. Systematics and
Biodiversity 22:49�55.
Ohtsuka S, Kase T, Boxshall GA. 2000. A new species of
Ridgewayia (Copepoda: Calanoida) from a submarine cave in
Palau, western Pacific. Species Diversity 5:201�13.
Schram F, Yager J, Emerson M. 1986. Remipedia. Part I.
Systematics. Memoirs of the San Diego Society of Natural
History 15:1�60.
Suarez-Morales E, Ferrari FD, Iliffe TM. 2006. A new epacter-
iscid copepod (Calanoida: Epacteriscidae) from the Yucatan
Peninsula, Mexico, with comments on the biogeography of the
family. Proceedings of the Biological Society of Washington
119:222�38.
Suarez-Morales E, Iliffe TM. 1996. New superfamily of Cala-
noida (Copepoda) from an anchialine cave in the Bahamas.
Journal of Crustacean Biology 16:754�62.
Walter C, Pasamonte JN, Talaue L. 1982. A preliminary
quantitative study on emergence of reef associated zooplankton
from a Philippine coral reef. Proceedings of the Fourth
International Coral Reef Symposium, Manila 1:443�51.
Yager J, Schram F. 1986. Lasionectes entrichoma , new genus, new
species (Crustacea: Remipedia) from anchialine caves in the
Turks and Caicos, British West Indies. Proceedings of the
Biological Society of Washington 99:65�70.
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
These characters not combined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
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
These characters not combined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8 Antennule with a digitiform process on segment IX; mandibular palp lacking an endopod,
the exopod with one extremely long seta distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oinella
These characters not combined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
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
New epacteriscids 91
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
These characters not combined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
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
92 A. Fosshagen & T. M. Iliffe