Psammogammarus lucayensis - Instituto Mediterráneo de

Zootaxa 3700 (1): 048–064
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Copyright © 2013 Magnolia Press
Article
ISSN 1175-5326 (print edition)
ZOOTAXA
ISSN 1175-5334 (online edition)
http://dx.doi.org/10.11646/zootaxa.3700.1.2
http://zoobank.org/urn:lsid:zoobank.org:pub:B13ECE90-D744-40DD-B754-B42FEF9BAA70
A new Psammogammarus (Amphipoda: Eriopisidae) from anchialine pools on
the Exuma Cays, Bahamas
DAMIÀ JAUME1, THOMAS M. ILIFFE2 & JORIS L. VAN DER HAM3
1
IMEDEA (CSIC-UIB), Instituto Mediterráneo de Estudios Avanzados, C/ Miquel Marquès 21, 07190 Esporles, Balearic Islands,
Spain. E-mail: [email protected]
2
Texas A&M University at Galveston, Department of Marine Biology, 200 Seawolf Parkway, OCSB #251, Galveston, Texas 77553,
USA. E-mail: [email protected]
3
George Mason University, Department of Environmental Science and Policy, 4400 University Drive, 3018 David King Hall, MSN
5F2 Fairfax, Virginia 22030-4444, USA. E-mail: [email protected]
Abstract
Psammogammarus lucayensis sp. nov. is described from anchialine pools on Little Iguana Cay (Exuma Cays, Great Bahama Bank). It can be easily distinguished from the other 14 members of the genus by the combination of: 1) carpus of
G2 longer than broad; 2) male G2 palm margin non-excavated, evenly convex and devoid of strong mid-palmar robust
setae; 3) basis of P7 with subparallel margins; 4) armature arrangement of ventral margin of epimeral plates as 0–2–3; 5)
posteroventral angle of epimeral plate III strongly produced; 6) protopod of U2 with distomedial angle armed with comb
of 3–4 robust setae; 7) U3 endopod as long as exp1; and 8) telson with robust setae on tip. The generic diagnosis is amended in order to allow the precise characterization of members of Psammogammarus compared to other eriopisids.
Key words: Amphipoda, Senticaudata, Eriopisidae, Psammogammarus, anchialine, Bahamas
Introduction
The Bahama banks in the western Atlantic Ocean harbour the richest assemblage of stygobitic anchialine
crustaceans currently known. No less than 96 species occur in this region, including phyllocarids (1), decapods (8),
isopods (12), amphipods (5), mysids (2), thermosbaenaceans (2), bochusaceans (1), remipedes (17), and
representatives of several orders of copepods (25) and ostracods (23). Most of these species are endemic, many
represent phylogenetic and/or biogeographic relicts, and many represent exclusively anchialine genera or even
families (Daenekas et al. 2009; see also http://www.cavebiology.com).
The shallow water environment of the Bahamas represents a rare case of a habitat that persisted over a long
geological time scale. The constitutive limestone—of marine shallow-water origin— has been accumulating in situ
since the Cretaceous, up to its current thickness of at least 4448 m (Mullins & Lynts 1977; Sealey 1994). Extensive
karstification combined with tectonic fracturation (Mylroie & Carew 1995), have led to the development of a vast
network of voids within this huge mass of limestone, offering a suitable habitat for anchialine fauna. The
persistence of this habitat over the last 120 Ma and the buffered environmental condition of the subterranean
environment, may partly explain the unusual accumulation of subterranean taxa in the Bahama banks.
Contrary to other taxonomic groups, the biodiversity of subterranean Amphipoda in the Bahamas is rather low.
Only five truly stygobiontic species have been reported thus far, four in the hadziid genus Bahadzia Holsinger, and
one in the pardaliscid genus Spelaeonicippe Stock & Vermeulen. Only Bahadzia obliqua Stock, B. setimana Stock
and Speleonicippe provo Stock & Vermeulen occur in the Great Bahama Bank, the greatest platform of the
archipelago (Stock 1986; van der Ham 2002).
Here we describe a new species of stygobiontic eriopisid amphipod belonging to the genus Psammogammarus,
from a small cay of the Exumas (Great Bahama Bank; Fig. 1). This genus is currently composed of only 14 species,
and shows a broad but punctuated circum-global distribution in shallow interstitial and anchialine habitats of
48 Accepted by J. Lowry: 14 Jul. 2013; published: 12 Aug. 2013
tropical/subtropical seas. Although, one of the littoral species has recently been reported to occur also at bathyal
depths in the Mediterranean (see Vonk et al. 2011, and references therein).
Psammogammarus lucayensis sp. nov., described herein, is the fifth member of the genus reported from the
Caribbean region, the other four species live in stygohabitats of the Dutch Antilles (Stock 1980; 1983; Vonk &
Stock 1987) and the coast of Nicaragua (Ortiz et al. 1993).
Material and methods
Specimens of the new Psammogammarus were collected from bottom sediments of an inland anchialine pool on
Little Iguana Cay in the Exuma Cays. Approximately 4 litres of fine muddy sediment were brought to the
laboratory and animals were collected from this sediment by hand. This tidal pool is one of several on the small
island and is about 6 m long by 4 m wide and 1 to 1.5 m deep. The pool is also inhabited by the anchialine shrimp
Parhippolyte sterreri (Hart & Manning), a species originally described from Bermuda caves. It is likely that
submerged cracks along the edge of the pool interconnect with deeper, but inaccessible anchialine voids.
Before study, specimens were treated with lactic acid to soften the cuticle and remove internal tissues to
facilitate observation. Drawings were prepared using a camera lucida on Olympus BH2 and Leica DM 2500
microscopes equipped with Nomarski differential interference contrast. Material preserved on slides was mounted
in lactophenol and the cover-slips sealed with nail varnish. Body measurements were derived from the sum of the
maximum dorsal dimensions (including telescoped portions) of head, pereionites, pleosomites and urosomites, and
exclude telson length. Material is deposited in the Crustacea collection of the Netherlands Centre for Biodiversity
Naturalis (RMNH) in Leiden, The Netherlands. Gnathopods 1 and 2, and pereiopods 3 to 7 appear abbreviated
elsewhere as G1–G2 and P3–P7, respectively; uropods 1–3, as U1–U3.
Taxonomy
Order AMPHIPODA Latreille, 1816
Suborder SENTICAUDATA Lowry & Myers, 2013
Family Eriopisidae Lowry & Myers, 2013
Genus Psammogammarus S. Karaman, 1955 amend
Diagnosis. Head lobe unnotched. A1 accessory flagellum ≥2-articulate. A2 flagellum with proximal articles not
fused to conform a composite article. Mandibular palp 3-segmented, third segment much shorter than second.
Maxillule [= maxilla 1] endopod [= palp] 2-segmented. Maxillipedal basal endite with up to four cuspidate robust
setae. G1 smaller than G2, with propodus longer than carpus; corresponding coxal plate not pointed forwardly. G2
coxal gill modified, sausage-like, elongated, and longer than basis of gnathopod. P7 basis not broadly expanded,
similar to P6. Epimeral plate 2 ventral margin lacking row of long setae. U3 dispariramous with reduced endopod;
exopod 2-segmented, not broadly expanded, both segments highly elongated and about same length.
Type species. Psammogammarus coecus S. Karaman, 1955, by original designation.
Composition. 14 taxa currently recognized, plus one described herein, distributed mainly in coastal anchialine
and marine shallow-water interstitial habitats of Japan, Indonesia, the Red Sea, Mediterranean, Cape Verde and
Canary archipelagos, the Dutch Antilles, the Caribbean coast of Nicaragua, and Baja California (Table 1; Vonk et
al. 2011, and references therein).
Remarks. Vonk et al. (2011) recently reviewed the main features of members of the so-called Eriopisa-group.
The foregoing generic re-diagnosis is based on data provided by these authors, complemented with features of the
new species described herein.
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-80°
-78°
-76°
Grand
Bahama
-74°
Abaco
26°
26°
Eleuthera
Nassau
Cat
Andros
San Salvador
24°
24°
Exumas
Little Iguana
Cay
Long
Acklins
Cuba
22°
22°
-80°
-78°
-76°
-74°
km
0
50
100
FIGURE 1. Map of the Bahamas showing position of Little Iguana Cay in the Exumas.
Psammogammarus lucayensis sp. nov.
(Figs 2–10)
Material examined. Anchialine pool on Little Iguana Cay (Exuma Cays, Great Bahama Bank). Holotype: Adult
female (oöstegites developed, setose) 3.77 mm, completely dissected and mounted on single slide
[RMNH.CRUS.P.10523]. Paratypes: Male 3.10 mm, completely dissected and mounted on single slide
[RMNH.CRUS.P.10524]; 75 specimens, all probably females, of which largest 3.73 mm, in single ethanol vial
[RMNH.CRUS.A.5041]. Collected by J. L. van der Ham & T. M. Iliffe, January 2003.
Diagnosis. Carpus of G2 longer than broad. Male G2 palm margin non-excavated, evenly convex and devoid of
strong mid-palmar robust setae. Basis of P7 with subparallel margins. Armature arrangement of ventral margin of
epimeral plates as 0–2–3. Posteroventral angle of epimeral plate 3 strongly produced. Protopod of U2 with
distomedial angle armed with comb of 3–4 robust setae. U3 endopod as long as exp1. Telson with robust setae on tip.
Etymology. The specific epithet is derived from the Lucayans, the presumed first inhabitants of the Bahamas.
Description of female. Eyeless. Body (Fig. 2A) slender, unpigmented, somites devoid of relevant armature or
sculpturing except for robust seta present on posteroventral angle of urosomite 3 (Fig. 9E). Head (Fig. 2B) lacking
rostrum; lateral lobes evenly rounded; antennal sinus hardly indicated. Epimeral plates (Fig. 9A) with acute
posteroventral angles, that of plate 3 more produced than rest; armature of ventral margin of plates (flagellate
robust setae) as 0–2–3.
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Antennule short, about half as long as body length (Fig. 2A, B). Peduncle segments relative length as 100: 92:
43; proximal segment provided with stout flagellate robust seta subdistally on ventromedial margin. Main
flagellum much longer than peduncle, articles each provided with single simple aesthetasc except most proximal
and terminal. Accessory flagellum 2-articulate, about as long as two proximal articles of main flagellum combined.
Antenna (Fig. 2A, B) attaining about three-quarters length of antennule. Gland cone slender, straight, pointing
anteriorly; relative length of three distal segments of peduncle as 45: 100: 89. Flagellum short, about as long as
distal segment of peduncle.
Labrum ordinary, globose, not figured. Paragnaths (Fig. 3C) with well-developed inner lobes; outer lobes each
with four multicuspidate terminal robust setae.
Left mandible (Fig. 3A) ordinary, with 5-denticulate incisor, 4-denticulate lacinia and columnar molar;
grinding surface of latter provided with pores as figured; molar seta pappose. Spine row comprising seven leaf-like
denticulated elements plus seven slender interspersed pappose setae. Mandibular palp distal segment shorter than
second, armed with two distal setae plus six setae along anterior margin; second segment with three setae along
anterior margin. Right mandible (Fig. 3B) different from left counterpart in multidenticulate lacinia and shorter
molar seta.
Maxillule (Fig. 4A) coxal endite [= inner plate] with 8+2 marginal setae. Basal endite [= outer plate] with nine
distal stout robust setae disposed in two rows (5+4), two of which tricuspidate, one 4-cuspidate, rest denticulated.
Endopod [= palp] 2-segmented, distal segment slightly expanded distally, with two unequal stout triangular
processes on distal margin; distal armature comprising two short, broad and stout denticulated robust setae, plus
four more slender denticulated setae placed subdistally on outer surface of segment as figured.
Maxilla (Fig. 4B) with oblique (“facial”) row on inner lobe composed of up to eight plumose setae; rest of limb
as figured.
Maxilliped (Fig. 4C) basal endite subrectangular with truncate distal margin, latter with four broad cuspidate
robust setae; remaining armature comprising oblique row of six plumose setae on posterior surface of endite, and
three simple setae on anterior surface as figured. Ischial endite (Fig. 4D) spoon-shaped with six robust setae on
distal margin, of which distomedial shortest and cuspidate, rest denticulated and progressively more slender
towards lateral; submarginal cluster of ca. 10 simple setae with blunt tip placed close to medial margin of endite on
its anterior surface. Merus-dactylus (= palp) slender, with nail (= dactylus + unguis) as long as broadly expanded
propodus (Fig. 4E).
Pereiopodal coxae 1–4 (Figs 5A, C; 6A, C) subrectangular, broader than long, each slightly overlapping one in
front (Fig. 2A); coxal plate 5 with broadly expanded anteroventral lobe (Figs 2A; 5A); coxal plate 6 anteroventral
lobe reduced (Fig. 7C), lobe wanting on plate 7 (Fig. 7E). Coxal gills on G2 and P3–P6, that on G2 sausage-like
(Fig. 5C), elongated, longer than corresponding basis; rest of gills (Figs 6A, C; 7A, C) ovoid and progressively
smaller towards posterior. Oöstegites on G2 and P3–P5 (Figs 5C; 6A, C; 7A).
Gnathopod 1 (Fig. 5A) medial surface of merus with dense patch of spinules; propodus (Fig. 5B) slightly
longer than carpus, 1.7 times as long as broad, broadest at palm angle; latter placed at 51 % of maximum [= dorsal]
length of segment, with armature comprising large flagellate robust seta and two shorter flagellate bifid robust
setae submarginally on medial surface, and two more slender, serrate flagellate robust setae on lateral surface of
segment; palm margin evenly rounded, lined with microserrate hyaline frill plus submarginal row of ca. 11 small
flagellate robust setae along medial surface of segment.
Gnathopod 2 (Fig. 5C) propodus 1.6 times as long as carpus, 1.8 times as long as broad, with parallel anterior
and posterior margins and with palm angle placed about midway of maximum [= dorsal] length of segment; palm
margin evenly convex. Armature of palm margin and palm angle similar to G1 counterpart except the two shorter
robust setae on medial margin are here unicuspid instead of bifid (Fig. 5D).
Pereiopods 3–4 (Fig. 6A–D) similar, P4 comparatively longer due to more elongated basis; nails similar, both
long and slender (seven times as long as broad, and about 44 % length of corresponding propodus), each provided
with two short blunt simple setae subdistally.
Pereiopods 5–7 (Fig. 7A, C, E) progressively longer towards posterior, with distal segments bearing numerous
flagellate robust setae distributed as figured. Basis of pereiopods moderately expanded, progressively broader
towards posterior, that of P7 with posterodistal angle strongly overhanging. Pereiopod 5 nail (Fig. 7B) shortest,
broadly overshoot by one of flagellate robust setae placed on distolateral angle of corresponding propodus. Nail of
P6 longer and more slender than P7 counterpart (compare Fig. 7D and F). All nails with single short blunt simple
seta subdistally.
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FIGURE 2. Psammogammarus lucayensis sp. nov. A, general aspect of female paratype 3.42 mm, lateral (notice distal portion
of P7 is missing); B, head of female holotype with right antennule and antenna attached, lateral. [Scale bar: 0.4 mm (A); 0.2
mm (B)]
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FIGURE 3. Psammogammarus lucayensis sp. nov., female holotype. A, left mandible; B, right mandible (palp omitted); C,
paragnaths; D, inset of right gnathopod 2 of male paratype, medial.
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FIGURE 4. Psammogammarus lucayensis sp. nov., female holotype. A, maxillule; B, maxilla; C, left maxilliped, anterior; D,
inset of maxillipedal ischial endite [= outer plate], anterior; E, inset of distal portion of maxillipedal endopod [= palp], lateral.
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FIGURE 5. Psammogammarus lucayensis sp. nov., female holotype. A, right gnathopod 1, medial; B, inset of palm; C, right
gnathopod 2, medial; D, inset of palm.
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FIGURE 6. Psammogammarus lucayensis sp. nov., female holotype. A, right pereiopod 4, lateral; B, inset of nail; C, right
pereiopod 3, lateral; D, inset of nail; E, left uropod 3 of female paratype 3.73 mm, dorsal. [Scale bar: 0.1 mm (B, D, E); 0.25
mm (A, C)]
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FIGURE 7. Psammogammarus lucayensis sp. nov., female holotype. A, left pereiopod 5, lateral; B, inset of nail; C, right
pereiopod 6, lateral; D, inset of nail; E, right pereiopod 7, lateral; F, inset of nail. [Scale bar: 0.25 mm (A, C, E); 0.1 mm (B, D,
F)]
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FIGURE 8. Psammogammarus lucayensis sp. nov., female holotype. A, right pleopod 1, posterior; B, right pleopod 2,
posterior; C, left pleopod 3, posterior.
Pleopods (Fig. 8A–C) ordinary, with spatial pattern and number of robust and simple setae on protopod, and of
simple setae on anterior and posterior surfaces of proximal articles of endopod apparently not fixed and submitted
to variability. Relative size of pleopods not corresponding to a progressive pattern of reduction towards posterior, i.
e. pleopod 2 shortest whereas pleopods 1 & 3 subequal in length.
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FIGURE 9. Psammogammarus lucayensis sp. nov., female holotype. A, left epimeral plates, lateral; B, right uropod 1; C, left
uropod 2; D, telson, dorsal; E, urosome of female paratype 3.73 mm, lateral. [Scale bar: 0.25 mm (A); 0.125 mm (B-D); 0.2
mm (E)]
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FIGURE 10. Psammogammarus lucayensis sp. nov., male paratype. A, left pleopod 1, anterior; B, right pleopod 2, anterior; C,
left pleopod 3, anterior; D, urosome, lateral; E, left uropod 1, posterior; F, right uropod 2, posterior; G, telson, dorsal.
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Uropod 1 (Fig. 9B) protopod distinctly longer than rami, latter unequal in length, with endopod distinctly
longest. Protopod provided with strong basofacial robust seta; rest of armature of segment comprising 5–6 robust
setae along posterolateral margin and 3–4 robust setae along posteromedial margin; distolateral and distomedial
angles of segment provided with one and two stout robust setae, respectively. Exopod with 1–2 robust setae along
lateral margin and five terminal robust setae; medial margin of segment unarmed. Endopod with one robust
marginal seta on lateral margin and 1–2 on medial margin; five robust setae plus simple seta placed terminally on
segment; in addition, simple seta implanted anteromedially on proximal portion of segment as figured.
Uropod 2 (Fig. 9C) protopod shorter than endopod; rami unequal in length, endopod distinctly longer than
exopod. Protopod with 1–2 flagellate robust setae on outer margin and 0–1 on inner margin; transverse row of 3–4
robust setae on distomedial angle, and single robust seta on distolateral angle. Exopod with two robust setae on
outer margin and single robust seta on inner margin; four robust setae, unequal in length, on tip; distolateral angle
of segment produced into short pointed process with bicuspidate tip. Endopod with 3–4 robust setae along inner
margin, 0–1 robust seta on medial margin, and five robust setae plus simple seta on tip.
Uropod 3 (Fig. 6E) elongated, about twice as long as urosome (Fig. 9E), strongly armed with robust setae
distributed as figured. Protopod short, slightly longer than telson, about 1.6 times as long as broad, expanded
distally; tiny simple seta implanted proximo-medially on dorsal surface of segment. Exopod 2-segmented, twice as
long as endopod; latter about as long as proximal segment of exopod.
Telson (Fig. 9D) cleft almost to base, much longer than broad. Lobes truncate with sinuose distal margin
provided with 2–3 robust setae; lateral margin of lobes with two short robust setae plus penicillate seta; two long
penicillate setae implanted subdistally on dorsal surface of each lobe as figured.
Description of male. The only male specimen collected, almost complete – only both U3 were missing–,
showed a remarkable sexual dimorphism on G2, protopod of U1 and U2, posteroventral robust seta of third
urosomite, and armature of telson. Thus the propodus of G2 (Fig. 3D) is a bit broader than in the female (twice as
long as broad vs. 1.8) and has a shorter posterior margin (palm angle placed at 44 % of maximum length of
segment vs. at 50 % length in the female); in addition, whereas in the female this segment attains its maximum
width between the anterior and the posterior margin, in the male it is reached between the anterior margin and the
proximal portion of the palm margin. Furthermore, the armature of the palm angle is reduced to two unicuspid
flagellate robust setae on medial margin (vs. three robust setae present in the female).
The U1 protopod is comparatively shorter than in the female, being as long as the corresponding exopod and
much shorter than the endopod, whereas in the female the protopod is longer than both rami (compare Figs 10D, E
and 9B, E). In addition, the robust seta present on the distomedial angle of protopod is hypertrophied in the male.
The U2 protopod is also shorter than in the female, being 1.6 times as long as broad vs. 2.7 times in the female
(compare Figs 9C and 10F).
The robust seta present on the posteroventral angle of the third urosomite is much longer than in the female,
being distinctly longer than half the U2 protopod length (compare Figs 10D and 9E).
The telson is more expanded proximally compared to the female, whereas its flagellate robust setae are much
larger (compare Figs 10G and 9D).
Presumed differences in number of armature elements on U1, U2 and pleopods (Fig. 10A–C) compared to the
female are pending confirmation once additional material is eventually collected. Nevertheless, the male pleopod 1
displays a bifid seta proximally on the inner margin of the proximal article of the endopod, whereas this seta is
unicuspid in the female (compare Figs 10A and 8A).
Discussion
Three out of the 14 currently recognized species of Psammogammarus share an evenly convex, not excavated palm
margin on male G2 with the new Bahamian taxon. These species are also unique with the new taxon in the display
of a male G2 carpus longer than broad (see Table 1). Namely, P. burri Jaume & Garcia, 1992, from an anchialine
cave in the Balearics, P. longidactylus Vonk & Stock, 1987, from an anchialine cave on Bonaire (Dutch Antilles),
and P. longiramus (Stock & Nijssen, 1965), from an anchialine pool on Entedebir Island (Dahlak Archipelago;
South Red Sea).
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The new species differs from P. burri in the lack of strong mid-palmar robust setae on the palm margin of male
G2 (vs. three setae present in P. burri); the U3 endopod is about as long as the proximal segment of exopod (vs.
much shorter in P. burri); the telson bears several strong robust setae distally (vs. telson unarmed distally in P.
burri); the posteroventral angle of epimeral plate 3 is strongly produced (vs. angle straight in P. burri); the P7 basis
is more expanded (length: width ratio 1.79 vs. 2.13); the ventral margin armature arrangement on epimeral plates
also differs (0–2–3 vs. 2–4–4); and there is a comparatively reduced number of setae on endites of maxillule,
among other features (Table 1; Jaume & Garcia 1992).
The new species differs from P. longidactylus in the outline of basis of P7, which is more slender and with
subparallel anterior and posterior margins (vs. basis broadly expanded with convex margins in P. longidactylus); in
the armature of distal angles of U2 protopod (with one and comb of 3–4 robust setae on distolateral and distomedial
angle, respectively, vs. 0 and 2 in P. longidactylus); and in the armature of epimeral plates (0–2–3 vs. 0–1–2),
among other features (Table 1; Vonk & Stock 1987).
The new species differs from P. longiramus in the armature of distomedial angle of protopod of U2 (with comb
of 3–4 robust setae vs. only 2 setae in P. longiramus); in the number of marginal setae on the coxal endite of
maxillule (10 vs. 15); and in the lower number of setae comprising the oblique row on the inner lobe of maxilla (8
vs. 12), among other features (Table 1; Stock & Nijssen 1965).
Two species, P. bluefieldensis Ortiz, Lalana & Beltrán, 1993, and P. scopulorum Stock, 1983, are known only
from female specimens. Nevertheless, both differ markedly from the new species, P. bluefieldensis displaying a
much shorter endopod of U3 (attaining only 34 % length of exp1, vs. segments equal in length in P. lucayensis sp.
nov.); two basofacial robust setae on the protopod of U1 (vs. one in lucayensis); only two robust setae on
distomedial angle of protopod of U2 (vs. comb of 3–4 in lucayensis); no lateral armature on telson (vs. two robust
setae at each side in lucayensis); coxal endite of maxillule with a much lower number of marginal setae (only 4 vs.
10); and the oblique row on inner lobe of maxilla composed of a lower number of setae (only 3 vs. 8), among other
features (Table 1; Ortiz et al. 1993).
Psammogammarus scopulorum differs from the new species in the armature of the distomedial angle of the
protopod of U2 (consisting of only two robust setae vs. comb of 3–4 setae in P. lucayensis sp. nov.); the reduced
armature of ventral margin of epimeral plates (0–1–1 vs. 0–2–3); the telson devoid of distal armature (vs. 2–3 distal
robust setae in lucayensis); and the lower number of robust setae on the endites of maxilliped, among other features
(Table 1; Stock 1983).
Acknowledgements
Contribution to Spanish MCINN project CGL2012-33597, partially financed with FEDER funds. Biological
collections from caves in the Bahamas were carried out under the terms of a Marine Resource Collecting Permit
issued by the Bahamas Department of Fisheries to Thomas Iliffe. Cave investigations in the Bahamas were funded
by grants from NOAA’s Caribbean Marine Research Center and the National Science Foundation (NSF #9870219
and 0315903) to T. Iliffe.
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