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| Serranids are ubiquitous predators on Caribbean
coral reefs and come in all shapes and sizes. Groupers are, or more accurately
were, the basis of important fisheries throughout the region but have become uncommon
on fished reefs. This very large and diverse family is composed of several clearly-defined
subfamilies. Those subfamilies with Caribbean reef-associated representatives
include the very numerous small seabasses and hamlets (Serraninae),
the groupers (Epinephelinae),
the basslets (Liopropominae),
the soapfishes (Grammistinae),
and the small reef bass (Pseudogramma
gregoryi), which may be considered either a Grammistinine or a Pseudogrammatinine).
There are many deep-water serranid genera and species that are not reef-associated,
but are included here for completeness. | | |
| Larval serranids can be recognized
by their realtively wide body, large terminal mouth, large round eye, and stout
spines on the head and in the medial fins (often serrated), with three spines
in the anal fin. Unfortunately, most of these characters are shared with many
other percoids, including the abundant snappers (Lutjanidae)
and grunts (Haemulidae).
These families broadly overlap in fin ray counts with serranids and would be a
problem for identification of late-stages if it were not for the very different
sizes of their larvae at settlement stage. Snapper larvae mature at a large size,
about 15 mm SL or more, and have a typical appearance that is shared by all of
the reef-associated species in the Caribbean (a few deep-water forms can be problematic).
Grunt larvae mature at a very small size, about 5 to 7 mm SL, and are thus easily
separated. Soapfishes differ from other percoids in having only a few small dorsal
spines and long soft dorsal and anal fins in an otherwise classic serranid body
form. Other percoids that can be difficult to separate from serranids include
hawkfishes (Cirrhitidae), with only six anal fin soft rays (seven or more for
Caribbean serranids). | | | |
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| | This
large group of small serranid fishes can be distinguished from other regional
serranids by their anal fin ray modal count of III,7 and having non-serrated fin
ray spines; the remaining subfamilies of Serranidae have eight or more anal fin
soft rays and those larvae often have markedly-serrated fin ray spines. Some snappers
(Lutjanidae) may have similar
fin ray counts and general appearances, but have eight or more anal fin soft rays
(for more detailed differences, see the lutjanid
web page). | |
| The deep-water anthiine serranines
are sometimes raised to their own subfamily Anthiinae, and they are the only serranines
to have some species with a mode of eight anal fin soft rays. | | |
| There are several reef-associated
Caribbean serranine genera, generally separating into two groups by dorsal fin
ray counts: the numerous hamlets (Hypoplectrus
spp.) with D-X,15 and the large genus of basslets (Serranus
spp. +), typically with D-X,12. | | |
| Shallow-water Serranus
spp. separate out somewhat by modal pectoral fin ray counts: a group with
14 (S. baldwini with 14
(13-15), S. tigrinus with
14, and S. tortugarum
with 14 (13-15)), then S. tabacarius
with 15 (sometimes 14); S. subligarius with
16 (14-17), and S. flaviventris with 16 (or
17). There are a number of deeper-water species as well: S.
annularis with 13 (or 14), S. chionaraia
with 14 (or 13), S. luciopercanus with 14,
S. maytagi with 15-16, S.
notospilus and S. phoebe with 15-16
(14-17) and S. atrobranchus with 16 (15-17). |
| | | The
remaining shallow-water serranine genera have sometimes overlapping fin ray counts
with the basslets: three Diplectrum spp.;
D. bivittatum (X,12 Pect-15-16, occ. 14),
D. formosum (X,12 Pect-16-17, up to 18), and
D. radiale (X,12 Pect-16-18, mode 17, from the S. Caribbean);
Paralabrax dewegeri (X,13-14 Pect-17);
Serraniculus pumilio (X,10-11, Pect-14-15); and Schultzea
beta (X,11-12 Pect-16). | | |
| Two deep-water serranines have
only ten dorsal fin soft rays: Parasphyraenops atrimanus
(X,10 III,6 Pect-17) and P. incisus
(X,10 III,7 Pect-17). The Centropristis spp.,
from the Gulf of Mexico north, have 11 (one with 12) dorsal soft rays and A-III,7
and comprise C. fuscula (with D-X,12), C.
ocyurus (Pect-17), C. philadelphica
(Pect-18) and C. striata (Pect 16-19). |
| | | The
rest of the deep-water taxa tend to have 13 or more dorsal fin soft rays and include
Bullsichthys caribbaeus (X,13-14 III,7 Pect-14-15)
and the anthiines. Anthiines comprise four Anthias
spp. all with D-X,14-15 Pect-18-21, i.e. Anthias
tenuis (III,8), A. nicholsi (III,7),
A. woodsi (X,14, III,7), and A.
asperilinguis (X,15, III,7); Hemanthias vivanus
(X,14, III,8 Pect-18-19), H. aureorubens (X,13-16,usually
15, III,8 Pect-16-17), and H. leptus (X,14
III,8); Pronotogrammus martinicensis (X,15
(13-16) III,7-8, Pect-16-18), and, finally, Plectranthias
garrupellus with the unusual fin ray count of X,16 III,7 and Pect-13. (Note:
some of these latter species with eight anal fin rays have identical fin ray counts
to some lutjanids.) | | | |
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| | Hypoplectrus spp. |
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| Hypoplectrus sp.
larva | | 7.2 mm SL | | San
Blas, Panama, SB86-1008 | | |
| Hypoplectrus sp.
larva | | 6.6 mm SL | | Slightly
vertically narrowed eye | | San Blas, Panama,
SB87-222 | | | |
| Hypoplectrus sp.
early transitional larva | | 7.3 mm SL |
| losing pectoral fin melanophores | | San
Blas, Panama, SB86-1004 | | | |
| Hypoplectrus sp.
larva | | 6.2 mm SL | Variant
pattern, row of melanophores on all anal fin ray membranes and an upper
caudal peduncle spot | | San Blas, Panama,
SB83-179 | | |
| Hypoplectrus sp.
larva | | 7.0 mm SL | Variant
pattern, with dorsal caudal peduncle spot | | San
Blas, Panama, SB86-425 | | |
| Hypoplectrus sp.
early transitional larva | | 7.0 mm SL |
| Variant pattern, with two caudal fin spots |
| San Blas, Panama, SB86-623 | |
|
| Hypoplectrus sp.
transitional larva | | 6.7 mm SL |
Variant pattern, with two ventral caudal peduncle
fin spots | | San Blas, Panama, SB83-156 |
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| Hypoplectrus sp.
transitional larva | | 6.7 mm SL |
| San Blas, Panama, SB83-156 | |
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Diagnosis: Modal fin ray counts of D-X,15 A-III,7
Pect-14 indicate the hamlets, Hypoplectrus spp..
There is a "species flock" of numerous color variations of these fishes in the
Caribbean, many of which can hybridize and the small juveniles are presumably
indistinguishable. Variant larvae with slightly different melanophore patterns
may either represent these different morphospecies or individual variation (or
both). DNA analysis may not be diagnostic, since it has been difficult to find
consistent sequence divergence between morphospecies, at least in Panama (McCartney
et al. 2003 and Ramon et al. 2003). The Caribbean morphospecies comprise
H. aberrans, H. chlorurus, H. gemma, H. gummigata, H. indigo, H. guttavarius,
H. nigricans, H. providencia, H. puella, and
H. unicolor. The medial fin ray count can be shared with some deep-water
Caribbean serranids (Anthias nicholsi and
A. asperilinguis, some
Hemanthias aureorubens, and Pronotogrammus
martinicensis), but the latter have more pectoral fin rays (16 or more).
(R) | |
| Analogues:
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| Description:
Body thin, wide, and short with a somewhat-narrowed to fully-round eye and very
large terminal mouth. Pectoral fins long, reaching to the vent. Pelvic fins long,
reaching about three-quarters of the way to the vent. Dorsal fin base long and
anal fin base short, caudal peduncle wide and short. The typical complement of
melanophores on the head consists of one at the angle of the jaw and a sparse
scattering on the top of the head. On the body there is one at the dorsal midline
just forward of the first dorsal spine and then two large melanophores on the
ventral midline of the caudal peduncle; one just behind the last anal fin ray
and one just before the first procurrent caudal fin ray (the latter are often
persistent through transition in serranines). Melanophores on the fins are prominent:
the full complement consists of a patch on the membranes of the third to fifth
dorsal fin spines, several near the base of the first three anal fin soft rays,
extensively lining the membranes of the pectoral and pelvic fins and finally one
at the base of the lower central caudal fin rays. Internal melanophores are present
around the sacculus and along the dorsal surface of the swim bladdder and the
peritoneum extending to the gut near the vent. Varying patterns are common: earlier-stage
larvae can be missing the melanophores on top of the head and/or the entire anal
fin, caudal peduncle, and caudal fin set of melanophores. Larvae approaching transition
progressively lose the markings on their pectoral and pelvic fin rays and some
larvae also develop a melanophore on the dorsal midline of the caudal peduncle
(the saddle characteristic of juvenile hamlets). One or both of the ventral caudal
peduncle spots occasionally are missing and sometimes the anal fin has additional
melanophores (sometimes a full row) just distal to the base of the rays. A variety
of additional melanophores occur in some individuals: just forward of the nasal
bones, along the ventral aspect of the lower jaw, an additional melanophore on
the caudal peduncle after the last anal fin ray or a second spot on the base of
the caudal fin rays (usually on the upper central caudal fin rays), or a few scattered
on the caudal fin rays. Pre-transitional larvae have a somewhat-narrowed vertical
oval eye becoming fully round as transition approaches. Transitional larvae develop
a fine scattering of discrete small surface melanophores, dense towards the anterior
and fading towards the tail and the larval melanophores progressively disappear
(usually starting with those on the pectoral fin rays). | | |
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| | There
are numerous species in this group and DNA sequence analyses indicate that it
is likely that the genus is polyphyletic (M. Craig pers. comm.). Several species
are very similar in general appearance and likely represent a clade of the wider-bodied
species. Based on the shared morphology and markings of juveniles, this clade
comprises S. annularis,
S. baldwini, S.
tabacarius, and S. tortugarum,
along with some, if not all, of the deep-water Serranus
spp. S. tigrinus
is longer-bodied and may group with some other deep-water serranines. The conclusion
awaits detailed phylogenetic studies. Nevertheless, the combination of modal pectoral
fin ray counts and the patterning of black spots along the upper body of transitional
larvae and juveniles should serve to separate most of the species. |
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| | Diagnosis:
Modal fin ray counts of D-X,12 A-III,7 Pect-14 (13-14) indicate a subset of Serranus
spp. Shallow-water species with this modal fin ray count comprise S.
baldwini, S. tortugarum,
and S. tigrinus (deep-water
species comprise S. chionaraia and S.
luciopercanus). S. annularis,
which is found in deeper water and is very similar in appearance to S.
baldwini, has a mode of 13 pectoral fin rays but cannot be excluded from
the larval type. Transitional larvae and recruits with four black patches along
the base of the spinous dorsal fin indicates S. baldwini.(ML) |
| | Analogues:
New recruit S. tortugarum
have only two black patches at the base of the spinous dorsal fin. New recruit
S. tigrinus have a longer
body and a pointed head and a black patch on the distal portion of the third through
fifth dorsal spines. | |
| Description:
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| Serannus tortugarum |
| | | | | |
| Serranus sp.
early transitional larva | | 8.1 mm SL |
| 13 pectoral fin rays | | San
Blas, Panama, SB86-101 | | |
| Serranus tortugarum
transitional larva | | 7.8 mm SL |
| San Blas, Panama, SB86-623 | |
|
| Serranus tortugarum
late transitional larva | | 8.0 mm SL |
| San Blas, Panama, SB86-1004 | |
|
| Serranus tortugarum
transitional recruit | | 9.9 mm SL |
| San Blas, Panama, SB82- | |
| |
| | Diagnosis:
Modal fin ray counts of D-X,12 A-III,7 Pect-14 (13-15) indicate a subset of Serranus
spp. Shallow-water species with this modal fin ray count comprise
S. baldwini, S. tortugarum, and S.
tigrinus (deep-water species comprise S.
chionaraia and S. luciopercanus). Transitional
larvae and recruits with two black patches at the base of the spinous dorsal fin
and the black markings on the mid-spinous dorsal fin on the proximal membranes
indicate S. tortugarum. (ML) |
| | Analogues:
S. tabacarius recruits
have the melanophores on the mid-spinous dorsal fin on the distal portion of the
membranes (vs. proximal). New recruit S.
annularis and S. baldwini
have four black patches at the base of the spinous dorsal fin (vs. two). New recruit
S. tigrinus have a longer
body and a pointed head and a black patch on the distal portion of the third through
fifth dorsal spines. | |
| Description:
Body thin, wide, and relatively short with a large round eye and very large terminal
mouth. Pectoral and pelvic fins relatively long, reaching much of the way to the
vent, dorsal fin base long and anal fin base short, caudal peduncle wide and short.
Melanophores on the head only at the angle of the jaw. On the body there is a
large melanophore at the ventral midline of the caudal peduncle just after the
last anal fin ray and a smaller one before the start of the procurrent caudal
fin rays. On the dorsal fin there are patches of melanophores on the membranes
between the third and sixth spines (mid-length on the third, proximal on the fourth
and fifth). Internal melanophores line the dorsal surface of the swim bladder
and peritoneum extending to the gut near the vent. Transitional larvae develop
melanophores in a large patch on top of the head and in a row along the dorsal
midline, starting with four patches: the first just forward of the dorsal fin
origin, the largest on and below the fin membranes at the fourth and fifth dorsal
fin spines, then on and below the last three dorsal fin spines, then on and below
the base of the third through fifth soft dorsal fin rays. At the same time the
larval melanophores are lost from the pectoral and pelvic fin membranes and then
from the body, with the last persisting larval melanophore the one just before
the lower procurrent caudal fin rays. A broad irregular stripe of fine melanophores
extends rearward from the upper edge of the operculum. Melanophores appear on
the iris at about 2, 3, 5, 8, and 11 o'clock. | | |
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| Serranus tabacarius |
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| Serranus tabacarius
larva | | 8.2 mm SL | | note
internal melanophores | | San Blas, Panama,
SB86-927 | | | |
|
|
Diagnosis: Modal fin ray counts of D-X,12 A-III,7
Pect-15 indicate several Serranus spp. and
Diplectrum bivittatum.
Serranus tabacarius is the shallow-water species
with this modal fin ray count (deep-water species comprise S.
maytagi, S. notospilus and S. phoebe).
Transitional larvae and recruits have two black patches at the base of the spinous
dorsal fin (vs. S. annularis and S. baldwini)
and the black markings at the distal portion of the spine membranes indicates
Serranus tabacarius (vs. S.
tortugarum). (ML) | |
| Analogues:
S. tortugarum recruits
have the melanophores on the mid-spinous dorsal fin on the proximal portion of
the membranes (vs. distal). New recruit S.
annularis and S. baldwini
have four black patches at the base of the spinous dorsal fin (vs. two). New recruit
S. tigrinus have a longer
body and a pointed head and a black patch on the distal portion of the third through
fifth dorsal spines. D. bivittatum
recruits have a longer body and often have 16 pectoral fin rays. |
| |
Description: Body thin, wide, and relatively short
with a large eye and very large terminal mouth. Pectoral and pelvic fins relatively
long, reaching much of the way to the vent, dorsal fin base long and anal fin
base short, caudal peduncle wide and short. Melanophores on the head only at the
angle of the jaw and at the isthmus. On the body there is a large spot below the
last dorsal spines, another at the ventral midline of the caudal peduncle just
after the last anal fin rays and a smaller one before the start of the procurrent
caudal fin rays. On the dorsal fin there are patches of melanophores on the distal
membranes between the third and eighth spines, and there are melanophores lining
the distal pectoral fin ray membranes and on most of the membranes of the pelvic
fins. There is often a small melanophore near the base of the first anal fin soft
ray and some melanophores along the tips of the anal fin ray membranes. Internal
melanophores line the dorsal surface of the swim bladder and peritoneum extending
to the gut near the vent. There is an additional deep melanophore above the spine
at the level of the last anal fin ray. | | |
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| Diplectrum bivittatum |
|
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| Diplectrum bivittatum
larva | | 12.7 mm SL | | San
Blas, Panama, SB84-627 | | |
| Diplectrum bivittatum
transitional larva | | 12.8 mm SL |
| San Blas, Panama, SB86-1001 | |
|
| Diplectrum bivittatum
late transitional larva | | 12.3 mm SL |
| San Blas, Panama, SB86-1002 | |
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| Diplectrum bivittatum
late transitional larva | | 12.9 mm SL |
| San Blas, Panama, SB86-1008 | |
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Diagnosis: Modal fin ray counts of D-X,12 A-III,7
Pect-15 and often 16, includes a wide group of serranines including Diplectrum
spp. in shallow waters and Serranus atrobranchus,
S. maytagi, S. notospilus, and S. phoebe
along with Schultzea beta, all in deeper waters.
Shallow-water Serranus spp. are less likely
candidates with S. flaviventrus reportedly
with 16 and sometimes 17 pectoral fin rays and S.
subligarius supposedly a northern Caribbean species. Recruits and juveniles
with prominent lateral stripes indicate Diplectrum
spp. The two species are separated only slightly by modal pectoral fin
ray counts (and scale counts): 15-16 in D. bivittatum
(54-75 lateral-line scales) and 16-17 in D. formosum
(46-55). This larval type could potentially include larvae of D.
formosum, but it may not occur in the Western Caribbean. The remaining
species in the genus, D. radiale, has a mode
of 17 pectoral fin rays. (ML) | |
| Analogues:
| |
| Description:
Body thick, somewhat wide, and long with a large eye and very large terminal mouth.
Pectoral and pelvic fins long, reaching more than half-way to the vent, dorsal
fin base long and anal fin base short, caudal peduncle wide and short. Melanophores
typically at the angle of the jaw, a cluster on the top of the head, and two stripes
of small melanophores, one along the upper body only below the spinous dorsal
fin and one full-length along the lateral midline. There is a row of melanophores
along the base of the anal fin soft rays, one per ray, and several along the ventral
midline of the caudal peduncle ending before the start of the procurrent caudal
fin rays. On the fins, melanophores concentrate on the distal membranes between
the dorsal spines and extensively speckled along the pectoral and pelvic fin ray
membranes. There are often a few melanophores between some of the anal fin rays.
On the caudal fin, there are melanophores at the base of several of the central
lower segmented fin rays and a larger patch at the base of the first two or three
upper segmented caudal fin rays that presents as a slight upward curve of the
main body mid-lateral stripe. Internal melanophores are present along the dorsal
surface of the swim bladder and peritoneum extending to the gut near the vent.
Transitional larvae intensify the two stripes and the mid lateral stripe extends
forward to the tip of the upper jaw and the upper stripe continues irregularly
backwards to the dorsal caudal peduncle. A third short stripe develops on top
of the head from the braincase towards the origin of the dorsal fin. Smaller melanophores
develop between the anterior portions of the main two stripes and just above the
anal fin, as well as in a series of fine patches just below the mid-lateral stripe.
The larval melanophores on the pectoral and pelvic fin ray membranes disappear
rapidly, those between the dorsal spines disappear last. Melanophores appear on
the iris at about 3, 7, 9, and 10 o'clock. | | |
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| Subfamily Grammistinae | | Rypticus
sp. A | | |
|
| Rypticus sp. A
larva | | 9.6 mm SL | | San
Blas, Panama, SB83-168 | | | |
| Rypticus sp. A
larva | | 10.5 mm SL | | San
Blas, Panama, SB83-169 | | |
| | Diagnosis:
Reduced spinous dorsal fin, only 2-4 dorsal spines, and no prominent anal fin
spines indicate the soapfishes, Rypticus spp.
There are five species in the southern Caribbean separated into two groups by
the number of dorsal spines: R. saponaceus
has 3 dorsal spines (and 23-25 dorsal rays) and R.
subbifrenatus has 3 or 4 dorsal spines (and 23 dorsal rays, 15 anal rays).
The remaining species have only two dorsal spines:
R. bistrispinus and R. bornoi (= R.
macrostigmus) (the latter with 24-27 dorsal rays, 14-17 anal rays). R.
maculatus is found in US waters only and R.
randalli occurs from Panama to Brazil. This larval type has a much shorter
second dorsal spine (which should exclude R. saponaceus),
along with juvenile markings of a broad dark stripe ending in a triangular point
on the caudal fin, characteristic of R. subbifrenatus
and R. bistrispinus). | |
| Description:
Body relatively thick, wide, and short with a large eye and large terminal mouth.
Pectoral fin rays very long, pelvic fins short, dorsal fin base relatively long
and anal fin base medium, caudal peduncle wide and short. First dorsal spine prominent
and covered in soft tissue, subsequent spines very short. Fins generally covered
in soft tissue membranes. Very few melanophores, typically only along the pectoral
rays. | | |
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| | Diagnosis:
Reduced spinous dorsal fin, only 2-4 dorsal spines, and no prominent anal fin
spines indicate the soapfishes, Rypticus spp.
There are five species in the southern Caribbean separated into two groups by
the number of dorsal spines: R. saponaceus
has 3 dorsal spines (and 23-25 dorsal rays) and R.
subbifrenatus has 3 or 4 dorsal spines (and 23 dorsal rays, 15 anal rays).
The remaining species have only two dorsal spines:
R. bistrispinus and R. bornoi (= R.
macrostigmus) (the latter with 24-27 dorsal rays, 14-17 anal rays). R.
maculatus is found in US waters only and R.
randalli occurs from Panama to Brazil. This larval type has three dorsal
spines with a large second dorsal spine and a dark juvenile with reticulations
and light-edged medial fins characteristic of R.
saponaceus. | |
| Description:
Body relatively thick, wide, and short with a large eye and large terminal mouth.
Pectoral fin rays very long, pelvic fins short, dorsal fin base relatively long
and anal fin base medium, caudal peduncle wide and short. Fins generally covered
in soft tissue membranes. Very few melanophores, typically only along the pectoral
rays. | | |
| Rypticus
sp. larva, 9.6 mm SL (San Blas, Panama, SB83-168) | |
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| Subfamily Pseudogrammatinae | | Pseudogramma gregoryi |
| | | | |
| | Diagnosis: Modal
fin ray counts of D-VII,18-19 A-III,15-16 indicate Pseudogramma
gregoryi. (U) | |
| Description: Body
relatively thin, somewhat long and narrow with a large eye and large terminal
mouth. Pectoral fin rays very long, pelvic fins short, dorsal and anal fin bases
relatively long, caudal peduncle wide and short. | |
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| All contents © copyright 2006 Benjamin Victor.
All rights reserved. | | | |