Marine Biology Research, 2007; 3: 73 92
New species of epacteriscids (Copepoda, Calanoida) from anchialine caves in the Caicos Islands and the Bahamas
AUDUN FOSSHAGEN1 & THOMAS M. ILIFFE2 1
Department 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
Abstract One new monotypic genus, Caiconectes , and one new species in each of the genera Azygonectes and Bofuriella are described from the Caicos Islands. In the Bahamas, a third new species of Bofuriella is described from Cat Island and the hitherto unknown female of Minnonectes melodactylus Fosshagen & Iliffe is recorded from Great Exuma Island. Caiconectes shows many plesiomorphic features, with some slight reductions in mouthparts, probably related to raptorial feeding. Swimming legs show maximum segmentation and setation; leg 1 has long and thin outer setae on the exopod, and the third segment of the endopod bears seven setae. The species is considered pelagic. The first male specimen of Azygonectes is described from A. plumosus sp. nov.; in both sexes, caudal seta V on the left side is approximately 2.5 times body length. The single female specimen of Bofuriella spinosa sp. nov. from Cat Island differs from other congeners in having a lateral genital aperture on the left side, and more setae on the mandibular endopod and on the coxal and basal endites of the maxillule. All cave calanoids hitherto described from the Caicos Islands are not recorded from any other parts of the Caribbean; thus pointing to an isolated position of the islands.
Key words: Anchialine caves, Bahamas, Calanoida, Epacteriscidae, Taxonomy
Introduction In Fosshagen et al. (2001), the family Epacteriscidae, 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 & Humphreys 2001), Iboyella from Cuba (Boxshall & Jaume 2003), and Azygonectes , Cryptonectes and Minnonectes from the Bahamas (Fosshagen & Iliffe 2004a), and a recent record of a second species of Balinella from the Yucatan Peninsula (Sua´rez-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 remipedes. The family is one of the most primitive of the
Calanoida and there are few synapomorphies. Important 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 Bermuda, 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 monotypic, 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 (Accepted 6 February 2007; Printed 20 April 2007) ISSN 1745-1000 print/ISSN 1745-1019 online # 2007 Taylor & Francis DOI: 10.1080/17451000701274571
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the last two genera have also been found in cryptic environments outside caves (Fosshagen 1973; Fosshagen et al. 2001) and from emergence traps in shallow water among corals in the Philippines (Walter et al. 1982). One species, Enantiosis belizensis Fosshagen, Boxshall & 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 specimens 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 ridgewayiids, 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 advanced diving techniques, when dragging a finemesh 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.
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 twosegmented 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 15 with three-segmented rami and following seta and spine formula:
Leg Leg Leg Leg Leg
1 2 3 4 5
Coxa 0-1 0-1 0-1 0-1 0-0
Basis 1-1 I-0 I-0 1-0 1-0
Exopod segments Endopod segments 1-1; 1-1; 2,I,4 0-1; 0-2; 2,2,3 I-1; I-1; III,I,5 0-1; 0-2; 2,2,4 I-1; I-1; III,I,5 0-1; 0-2; 2,2,4 I-1; I-1; III,I,5 0-1; 0-2; 2,2,3 I-1; I-1; III,I,4 0-1; 0-1; 2,2,2
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 transformed 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 supposedly ancient and primitive species (from the Latin antiquus meaning ancient). Caiconectes antiquus gen. et sp. nov.
Diagnosis Female. Prosome in dorsal view widest posterior to middle, five pedigerous somites well defined. Urosome four-segmented, genital double somite elongate 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
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 12 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.
New epacteriscids 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. Urosome (Figure 1C, D) four-segmented, genital double somite elongate with genital aperture located 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 IIIXXI 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 segment 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 approximately 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 medium 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; endopod six-segmented, formula 2, 4, 4, 3, 31, 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 25 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 segment relatively long. Male. Body length 2.70 mm. Urosome (Figure 3C) five-segmented, genital somite approximately threequarters length of second urosomite, genital aperture on the left side (spermatophore shown in figure); caudal rami as in female. Right antennule (Figure 3D, distal part) 24segmented, segments IIIV 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; exopods slightly modified with same number of elements on both sides but with longer segments on left side; first segment with outer distal spine; second
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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.
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 Providenciales, North Caicos and Middle Caicos. In Providenciales 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 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.
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.
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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.
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 Kaloketos pilosus Koenemann et al., and the shrimp Agostocaris williamsi Hart & Manning (Hart & Manning 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 semicircular 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
New epacteriscids depths greater than 3 m. Previously described stygobites from this cave include Lasionectes entrichoma. 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 platform. Passages range from large, dry caverns previously 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, Erebonectoides macrochaetus (Fosshagen) and Fosshagenia ferrarii Suarez-Morales & Iliffe have been described from this locality (Fosshagen & Iliffe 1994; SuarezMorales & 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 extension 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 (XXVIIXXVIII) 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 appendages 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 Speleophriopsis 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 knowledge, this character of the endopod is novel to calanoids, where the maximum number is six with
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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 segments XXVI and distal compound segment. Antenna 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 segment offset near middle of inner margin of preceding segment. Male. Urosome five-segmented with first four somites of equal length, anal somite approximately half length of preceding somites. Right antennule geniculate, 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 12 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; urosomites 1 4 of subequal length, anal somite approximately 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 IIIV separate, segments 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
Figure 4. Azygonectes plumosus sp. nov., male. (A) Habitus, lateral; (B) rostrum; (C) urosome, dorsal; (D) right antennule; (E) distal part of left antennule.
Balinella , Bofuriella , and Bomburiella , all 26-segmented with different degrees of incomplete fusion between segment XXVI and the compound terminal 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
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Figure 5. Azygonectes plumosus sp. nov., male. (A) Antenna; (B) mandible; (C) maxillule; (D) maxilla; (E) maxilliped.
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 threesegmented endopods and modified exopods, threesegmented on the right side and two-segmented 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 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
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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 12 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.
New epacteriscids Hill, the oxygen levels below 3 m depth were less than 0.5 mg l1. 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 , threesegmented and equipped with more setae; the distal basal endite carries five setae and the exopod 11 setae, conditions that are among the most plesiomorphic 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 proximally. Rostral lobes weakly developed, each with one filament. Antennule 26-segmented with segment XXVI incorporated into apical double segment XXVII XXVIII. Endopod of antenna approximately two-thirds length of exopod and with welldeveloped 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.
A. Fosshagen & T. M. Iliffe
Maxilliped with one small seta on the praecoxal endite; weaker spines on the endopod than in B. vorata; only one spinulate claw present, this originates 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 25 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 collected 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 Caiconectes described in this paper. Remarks Only one species, Bofuriella vorata Fosshagen, Boxshall & 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 15 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 similarity 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. Specimens 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 2006.380382. 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 ventrally 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.
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.
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
A. Fosshagen & T. M. Iliffe
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 misophriids, 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 27segmented 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 1540 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 asymmetrical 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 posterior 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 descriptions 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 Paramisophria 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
New epacteriscids 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 present 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 different, 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 described from the Caicos Islands are unique to the Bahamas archipelago; one belonging to each of the genera Caiconectes gen. n., Erebonectoides , Azygonectes and Bofuriella. Caiconectes gen. nov. and Erebonectoides are monotypic, a second species of Azygonectes is previously recorded from South Andros Island and the Exumas, 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, surrounded 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 monotypic. 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 gnathobase 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, Erebonectinae and Epacteriscinae; the first one containing the two genera Erebonectes and Erebonectoides and the second one all other remaining genera. Erebonectinae has two outer spines on the third exopodal segment of swimming legs 3 and 4, whereas Epacteriscinae 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).
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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 environments; the last feature indicated by species of Enantiosis having various lengths of the antennules. Species of Caiconectes and Azygonectes are considered 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 Cryptonectes 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 (Universidad 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 Fisheries. 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’Histoire 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 Bahamas. 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.
New epacteriscids 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.
Sua´rez-Morales E, Ferrari FD, Iliffe TM. 2006. A new epacteriscid 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. Sua´rez-Morales E, Iliffe TM. 1996. New superfamily of Calanoida (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 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 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 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 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 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 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 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 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 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
A. Fosshagen & T. M. Iliffe
Rostrum weakly developed as a broad plate with short filaments; articulation between antennular ancestral segments XXVI and double segment XXVII XXVIII not expressed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . These characters not combined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Maxilliped with reduced enditic setation on the syncoxa, basis elongate with a group of three setae distally, endopod with long flexible setae. . . . . . . . . . . . . . . . . . . . . . . Maxilliped of a different shape . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Maxilliped strongly developed, basis compressed, endopod with strong spinulate claws . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Maxilliped slender, not particularly modified. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rostrum bifurcate with widely separated tips. . . . . . . . . . . . . . . . . . . . . . . . . . . Rostrum elongate with two closely set lobes bearing a pair of thick filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Strongly modified legs 5 in the male, particularly on the left exopod and endopod; first basal endite of the maxillule with one seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . 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 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Proximal segments (IVII) of the antennule without modified setae; anal somite concealed beneath the posterior rim of the preceding urosomal somite. . . . . . . . . . . . Proximal segments (IVII) of the antennule with modified setae, flattened proximally and with a filament at the distal part; anal somite not concealed by the preceding somite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Antennular segments IIIV fused; mandibular palp lacking an endopod. . . . . . . . . . . Antennular segments IIIV partially separated; mandibular palp with a one-segmented endopod bearing two setae; strong setae on the maxilla and maxilliped with black-pigmented tips . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
12 14 . . . . Bomburiella 13 . . . . . Bofuriella . . . . . . .Iboyella 15 . . . . . . Gloinella . . . . . Enantiosis
16 . . . . . Edaxiella
17 . . . . . .Bunderia
. . . Minnonectes