Note (February 2012): My webguide is
a work in progress, with daily edits and new family pages
and photographs added continuously. My focus at present
is on a few large families that account for about a third
of all reef fishes in the region: the gobies with about
150 species, the serranids with about 100 species, the
labrisomids with about 50 species, and the chaenopsids
with another 50 species. I am concentrating on the early
life history stages of labrisomids
and chaenopsids
right now- in more detail than before, with more and better
photos as the technique improves, and as a basic model
for my future larval pages. My long-range plan is to first
photograph my Caribbean larval collections, then go on
to my eastern Pacific and Indo-Pacific collections.
Why keep my guide on the web? The main
reason is that books and papers require that information
be frozen in time: essentially a snapshot of knowledge at
a particular moment. We used to accept this as inevitable
and many of us probably don't even think about it. But it
is clear that scientific information is continuously being
updated and remodeled and it is somewhat discordant, and
certainly inefficient, to produce and package the results
in an occasional bolus. Unfortunately, most of us accept
that authors will have their additions, changes, and errata
somewhere on their desk and inaccessible (we hope only until
the next edition). Even worse, when the subject is a compendium
of information, such as a guide, authors often wait until
it is in some form "complete" before submitting it to publication.
Woe be unto the perfectionist in this case, for it may take
a lifetime for some people to decide they are ready. That
is all 20th-century thinking- there is no reason in the
age of the internet not to have a work in progress made
generally accessible.
INTRODUCTION
Virtually all of the thousands of species
of tropical reef fishes have a larval stage that spends
weeks to months in the open ocean before returning to the
reef to settle. This transition from pelagic larvae to settled
juveniles is a profoundly important time for reef fishes.
During this settlement transition, which usually occurs
on moonless nights, the larvae have critical decisions to
make: they need to select an appropriate habitat, avoid
the ubiquitous predators, and change their shape and color
for life on the reef. Understanding this early-life-history
process is clearly important to understanding the population
dynamics and, more broadly, the biogeography, ecology, and
evolution of coral reef fishes.
Despite
its undoubted importance, this transition
has not been comprehensively studied and one
of the reasons this may be true is the lack
of information on late-stage larvae. I have
been collecting these larvae, mostly by netting
at a nightlight directly over the reef, but
also with crest-nets and other techniques
for many years. In addition, I have focused
my daytime collecting on what some of us call
"new recruits", the recently-transformed juvenile
fishes that suddenly appear in the morning
on the reef.
For some reason, almost all of the reef-fish
larvae I collect at a nightlight are late-stage larvae ready
to settle onto the reef. This is not an artifact of their
attraction to the light: I have towed plankton nets in the
area and I get a similar size complement of larvae. In addition,
other techniques to collect incoming larval fish, such as
crest nets, yield the same size categories of larvae.
The
simplest explanation for this phenomenon is
that larval fish that are ready to settle somehow
maneuver themselves into on-reef currents, perhaps
just by rising to the surface water layer. There
has been recent research indicating that reef-fish
larvae are good swimmers and can actively orient
towards reefs. Whichever way they manage it,
almost all of the larvae I capture are around
the particular settlement size range for their
species. This can be very helpful in identifying
larvae.
REFERENCES
There are a number of excellent books
on larval identification and the early life history stages
of fishes. Most include copious drawings of reef-fish larvae
and also focus on the earlier stages of fish larvae: Jeff
Leis at the Australian Museum has published his opus
(or opuses) with comprehensive coverage of the Indo-Pacific
species of reef and shore fishes.
For
the Atlantic, there is the latest larval fish
book edited by Bill Richards:
Richards, W.J. (2005) Early Stages Of Atlantic
Fishes: An Identification Guide For The Western
Central North Atlantic, Vols. 1 and 2
It covers pelagic fishes and deepwater fishes
as well as shore and reef species and captures
the state of the art from a couple of years
ago. Portions of this otherwise prohibitively
expensive and hard-to-find book can be viewed
on Google, but it is stark testimony to the
limitations of spreading information with
paper. Lastly, the classic book by Michael
Fahay (1983), the Guide to the Early Stages
of Marine Fishes Occurring in the Western
North Atlantic Ocean, deals mostly with
the temperate ichthyoplankton of the Atlantic
coast of the US, but with some overlap of
coverage.
Larval descriptions have been illustrated
with line drawings for many years and even
now virtually all books and scientific papers
use line drawings almost exclusively. There
are a few benefits to the use of drawings
for early stage fishes; for example, otherwise-transparent
details such as head spines and pectoral fin
rays can be highlighted. Nevertheless, it
is most likely that the main reason for line
drawings is the cost of printing photographs
and the necessity to limit the number of pages
and illustrations in the publishing process.
But, in the near future, and however resistant
we may feel to change, libraries certainly
will be merged into central databases and
printing presses and paper will go the way
of the neighborhood bookstore. Line drawings
may seem as quaint as the watercolors used
for the illustrations in 19th century species
descriptions.
There are many limitations to line drawings.
Perhaps the most problematic is that melanophores (the black
spots usually prominent on fish larvae), especially small
or delicate ones, disappear against a black line drawing
and these melanophores are often the most critical element
in larval identifications. The idealization of the drawing
can also distort the real appearance of the larvae, emphasizing
outlines and de-emphasizing shading, form, and especially
color. Furthermore, there is usually marked variation in
the appearance of fish larvae, both functional (such as
melanophores expanded or pinpoint) and inherent (incomplete
melanophore complements are common). This variation among
individuals certainly argues against presenting an ideal.
In addition, the development of metamorphic markings and
morphological changes is a continuous process and it is,
of course, impossible to pick one or two images to capture
the information.
Now that digital photography and web publishing has streamlined
the use of photography, it is possible to present many
illustrations of a single larval type and get around the
limitations of choosing a single or only a few diagnostic
images. I hope to emulate the actual process of identification,
which is to use a variety of views and backgrounds to
create a gestalt that quickly identifies a larval type
without following a key or character list to arrive at
a diagnosis.
Along a similar vein, most of the literature to date
bravely (or is it quixotically) tries to verbally describe
larval shapes and melanophore patterns. More text is usually
favored over more illustrations, probably because of the
decreased costs of publication. This obviously limits
the amount of information captured since it is technically
impossible to render three-dimensional shapes and patterns
into words. Furthermore, the location of markings and
structures can only be given relative to other
landmarks and is always limited in precision by
the amount of text that can be printed, or even
tolerated, by a reader. I therefore will try to
avoid the temptation to add reams of description
to the photographs, but I will insert a hopefully
pithy verbal description when it is necessary to
highlight relevant features
I present the photographs of larvae usually
with a dark background, sometimes along with a matched photo taken
against a light background so that the black melanophore pattern,
and especially internal melanophores, can be seen. Photographs
against dark backgrounds, like line drawings, can hide the small
or edge melanophores that are often diagnostic for larval types.
I also include detail photographs on pertinent features of the
larvae that may be critical to identification or of specific interest.
If there are transitional forms of interest, photographs of those
are included. I also hope to add images of new recruits, especially
for those groups with many similar species and distinctive juvenile
markings.
THE
DEFINITION OF LARVAE
Please
note that most reef-fish biologists use the loose
definition of the word "larvae", i.e. the stages
of development before settlement to the reef, if
they show a morphological difference from settled
juveniles.
This definition of early stage fishes
excludes the pelagic juveniles of some species who are typically
indistinguishable from reef-based juveniles and often float
with drift objects: these fishes have essentially settled
onto a drifting platform. As one can imagine, the few species
that do this regularly are annoying to us mainly because
they defy the usual interpretations of what defines "larvae",
"settlement", and "pelagic larval duration". Reef fishes
in this category include the Sargassum
algal drift-associated juveniles, to some degree or another:
the damselfishes of Abudefduf and Chromis
(Pomacentridae), triggerfishes (Balistidae), filefishes
(Monacanthidae), barracudas (Sphyraenidae), tripletails
(Lobotidae), pipefishes and seahorses (Syngnathidae), and
needlefishes (Belonidae).
Another intriguing discovery is the occasional
pelagic adult reef fish. This is new information and may explain
the sometimes contradictory findings of the degree of larval dispersal
derived from study of larval pelagic durations versus the observed
degree of gene flow in phylogenetic studies. The elusive pelagic
adult is quite intriguing, and I hope to pursue this curious creature
in future expeditions. Several sources of information indicate
that this is happening, although to what degree is completely
unknown. The original observation was by Ross
Robertson who saw adults of the slingjaw wrasse, Epibulus
insidiator, rising off of reefs and drifting away offshore
(in Palau, a part of Micronesia, in the Western Pacific). Then
there have been records of the rare adult reef fish caught in
light traps far offshore (reports from the Great Barrier Reef)
and observations of adult reef parrotfishes in schools in pelagic
waters far from reefs (Enric Sala reports from diving in Cuba).
The significance of this phenomenon
is unknown. Perhaps it is a way of relocating among reef
systems for more advantageous feeding or reproduction, although
this would require that the likelihood of returning to a
reef outcrop be greater than the likelihood of being eaten
by a tuna... somewhat hard to imagine. Alternatively, some
pelagic adults may not be reef emigrants, but could be lost
pelagic juveniles approaching maturity, perhaps associated
with drift structure for protection, and haplessly waiting
for a chance encounter with a reef. Larval mortality schedules
must have a tail to the distribution and thus permit some
small but real fraction of pelagic larvae to persist through
transformation (if obligatory at some point) and even grow
to adulthood offshore.
HOW DO
I IDENTIFY REEF FISH LARVAE?
This is perhaps the most common
question I am asked, since there is a bewildering variety of fish
larvae in almost any collection. In addition, the possible candidates
for any particular larva in the Caribbean or the eastern Pacific
include a thousand reef-fish species, and perhaps another thousand
other shorefish species and deepwater, midwater, and pelagic fishes
as well. That total rises to many thousands as one approaches
the center of diversity near Indonesia in the central Indo-Pacific.
Fortunately, larval fishes, especially at later stages, are not
uniform or even broadly similar; in fact they can often have even
more divergent characters than do the adults. For example, virtually
all reef fishes have round eyes after settlement, while their
larvae can have widely-varying eye morphologies, often narrowed
or tilted and sometimes exhibiting bizarre shapes.
In addition, some taxa have larvae with greatly-extended
fin spines or rays, sometimes many times longer than the larva
itself. There is also a great diversity in spine development,
with many larvae having complex spine ornamentation on the head
or fins, often lost or inconspicuous on adults. Finally, an important
character at higher taxonomic levels for fishes is the number
of myomeres, the muscle segments making up the body. These are
hidden under pigment and skin in juvenile and adult fishes, but
are clearly obvious and countable in most larval fishes.
Some
taxa fortunately develop the typical juvenile appearance as late-stage larvae
and are thus easy to identify from known-juvenile markings and coloration. Typically
these larvae still do show some subtle adaptations to pelagic life, often a silvery
sheen and shades of black, gray and brown or red instead of the usual bright colors.
Red does not penetrate seawater well and would appear gray, especially in low-light
conditions. Examples of fishes using this strategy include the angelfishes (Pomacanthidae),
the butterflyfishes (Chaetodontidae),
and the squirrelfishes (Holocentridae). A larval Pomacanthus arcuatus,
only 8.2 mm SL, is pictured at right.
Others live among drift algae and are camouflaged
by markings similar, but not identical, to those of settled juveniles.
The camouflage is an adaptation to avoid being eaten by the ubiquitous
pelagic predators which target floating objects in the open ocean,
such as tunas and mahi-mahi (dolphinfish). Larvae following this
strategy include some pipefishes and seahorses (Syngnathidae),
triggerfishes (Balistidae), and filefishes (Monacanthidae).
Identification to genus and species
in larvae with a more generic appearance can be much more difficult.
Most generic types of reef fish larvae are transparent and very
small and, while they are alive, look just like tiny slivers of
glass with eyes. When there are just a few genera or species in
a group, the fin ray counts are usually diagnostic to species.
In some of the more speciose groups with large sets of closely
related species, such as the gobies (Gobiidae)
and the gobioids (Gobioidei),
the scaled blennies (Labrisomidae),
the snappers (Lutjanidae),
the seabasses and groupers (Serranidae),
as well as parrotfishes (Scaridae),
wrasses (Labridae),
and some damselfishes (Pomacentridae),
the number of possible species can be overwhelming and I have
to resort to some other method of identification. There are a
variety of ways to do this, some simple and some using the latest
molecular techniques.
FIRST,
USE THE PROCESS OF ELIMINATION...
Most
reef fish larvae do resemble juveniles and adults in basic form, and, especially,
in the number of fin spines and rays and myomeres. This is the primary method
of identification for early life history stages and works almost all the time
for family, most of the time for genus, and often for species.
Of
course, this is essentially a process of elimination and it is much easier in
the Caribbean, with only a few hundred possible candidates, versus the Indo-Pacific
where there can be thousands of reef fish species occurring at one particular
site. The eastern Pacific region is generally similar to the Caribbean in numbers
of reef fish species. The majority of my specimens are Caribbean, most from the
San Blas Islands of Panama, so I will start developing the guide for these fishes.
Other Caribbean sites from which I have collected include Belize, Puerto Rico,
and St. Thomas. I have also collected fishes at the Brazilian island of Fernando
de Noronha, where there is an unusual complement of reef fishes, some endemic
and some widespread Caribbean forms.
A particularly useful variant of
the process of elimination is the examination of fishes from isolated
regions where only a subset of the group is present (i.e. depauperate
locations). This is a very useful method in some groups where
the larvae of similar species can be very difficult to separate.
For example, there are numerous very similar gobies of Coryphopterus in the Caribbean, but only one species
occurs on the island of Noronha far off the coast of Brazil.
Another useful variant is
using the similarity with sibling species in the eastern
Pacific Ocean to refine the process of elimination. When
identifications of larval types becomes difficult because
there are several Caribbean species that all share fin ray
counts, the situation may be different on the Pacific side
of Central America. The isthmus of Panama rose about three
million years ago, and there are numerous reef fish genera
with species on both sides of the isthmus, often as very
similar species pairs or "sibling species".
On the
other hand, some genera occur on only one side, and the
presence (or even absence) of a particular larval type onone
side or the other can contribute to the process of elimination
and confirm an identification. For example, the sleeper
Erotelis
smaragdus shares its fin ray counts with other Caribbean
sleepers, making it a problem for identification. But, in
the eastern Pacific, the sibling species Erotelis armiger
has a unique fin ray count among the eastern Pacific sleepers.
I now have collected a series of sleeper larvae with that
fin ray count from near the Galapagos Islands, and they
are almost identical to the putative Erotelis
smaragdus larval type... thus confirming the identification
for the Caribbean larval type. Another example is an unusual
Caribbean goby larval type with a fin ray count matching
several Ctenogobius,
but also the monotypic Evorthodus
lyricus. Again, I have collected an identical larval
type from the eastern Pacific Ocean, where, fortuitously,
there are only two Ctenogobius species (neither of
which match the fin ray count), but there exists the sibling
of the Caribbean Evorthodus species, Evorthodus
minutus, with the appropriate fin ray count. That nicely
confirms the identification of the Caribbean larval type
as Evorthodus
lyricus.
THEN,
LOOK FOR MISSING LINKS (TRANSITIONAL LARVAE AND TRANSITIONAL RECRUITS)
On
close examination, many larval types have
very characteristic and remarkably consistent
patterns of melanophores. Often, they also
have specific patterns of other chromatophores.
Pigment cells in fishes include the classic
melanophores which comprise black and brown
spots, iridophores which are mirror-like spots
that simply reflect light and often appear
silvery, leukophores which are white, and
xanthophores which range from yellow to orange
to red. Unfortunately xanthophore pigments
dissolve rapidly in solvents and are not usually
preserved.
Now
why would these markings be so prominent and so specific? I think pelagic larvae
use these for species identification. Perhaps they prefer to travel together while
in the open ocean and settle together. I have noticed that sometimes several new
recruits of a particularly uncommon species will show up on the same rock on the
same morning, with no other recruits for miles around. Interesting....
Fortunately,
in most reef fishes the larval markings are quite different from the juvenile
markings. Larval melanophores are typically large, often stellate, individual
black spots that extend below the skin surface. These markings are replaced during
the settlement transition by metamorphic melanophores in the patterns shown by
juveniles on the reef. These arrays of additional melanophores (and sometimes
leukophores and iridophores), are usually much smaller and right at the skin surface.
In addition, they are typically numerous and in dense patches that often begin
on the head and develop posteriorly following the pattern of the juvenile markings
of the species. This transitional larva of a scaled blenny (Labrisomidae) shows the
prominent three larval melanophores surrounded by the patches of fine metamorphic
melanophores. In many species, this transition can take days and often the larval
markings coexist with the developing juvenile patterns. These metamorphic melanophores
are invaluable for identifications.
So
an excellent method of identifying larvae is to find the occasional pelagic larva
that has started to transform and is developing the characteristic juvenile markings
while still in the water column. Alternatively, I often have collected new recruits
on the reef that still have some identifiable larval melanophore patterns, i.e.
transitional recruits. Both of these forms
of missing links can indicate what species that type of larvae represents.
THE GOLD STANDARD:
DNA SEQUENCE MATCHING
The most recent advance is the use of DNA
sequencing to identify larvae. This technique
is rapidly becoming the gold standard for
larval identification. Once a library of sequences
is known for a group of species, then individual
larvae can be sequenced and matched to the
known species. This method can be expensive
and requires a full set of species-specific
sequences for the group, which is not always
easy to assemble. Nevertheless, I have started
a comprehensive larval-adult matching project
using mitochondrial DNA sequences and have
started to resolve some of the more tricky
ID situations, such as the damselfishes of
Stegastes,
the labrids of Halichoeres,
the parrotfishes of Sparisoma,
the snappers of Lutjanus,
and the gobies of Bathygobius,
Lythrypnus,
and Coryphopterus.
In addition to confirmatory IDs, sequence
analyses are wonderful tools for discovering
new and/or cryptic species: indeed, recent
analyses I have done have discovered cryptic
species among the Caribbean Coryphopterus
and that two cryptic species of bonefishes,
Albula, coexist in equal numbers in
my Panama collections. Another benefit of
sequencing reef fishes is discovering surprising
and unexpected patterns of relatedness (although
a long list of caveats should follow: "phylogenetic
relationships require multiple loci and appropriate
statistical analyses to ensure any robustness!").
My sequences for the labrids confirmed (in
agreement with other published trees) that
Halichoeres
maculipinna is only distantly related
to the other Halichoeres
species and may be closer to the
Thalassoma. In
addition, with collections over wide geographic
ranges, one can assess the degree of genetic
divergence of isolated island populations.
My sequences reveal that this divergence sometimes
seems to show little relation to the duration
of the pelagic larval phase. I cannot really
explain this counter-intuitive finding and
it surely indicates that the relationship
between dispersal and speciation is not a
simple one (to say the least). Clearly, a
plethora of profound insights are about to
arrive with the advent of widescale DNA sequencing
of reef fishes.
FINALLY, AQUARIUM
RAISING OF LARVAE
Another method is to isolate
the wild-caught larva and raise it in an aquarium tank until
it develops identifiable juvenile markings. This method can
be labor intensive and requires the larva not to be so delicate
that it dies from handling. In addition, there is a piscine
Heisenberg Uncertainty Principle, which is that one cannot keep
the larva in preservative and also raise it alive. If it is
possible to examine the living larva close enough (and even
photograph it) and then raise it, an identification can be made.
An
alternative, although much more intensive
and expensive, method to document the early
life history of reef fishes is to raise the
fish from the egg through the larval period
in the laboratory. This is being achieved
more recently as techniques for feeding notoriously
delicate reef fish larvae are perfected (see
for example the Red Snapper, Lutjanus
campechanus).
There are some indications, however, that
laboratory-raised larvae may look somewhat
different from "real" larvae since they are
certainly living in unnatural conditions and
may grow at different rates. In addition,
they would be undoubtedly confused as to when
and whether they should enter transition.
TERMINOLOGY
AND LAYOUT OF LARVAL IDENTIFICATIONS IN THE GUIDE
Identifications will be
presented to the lowest taxonomic level of which I am reasonably
sure. A question mark after the genus or species name indicates
that I am not reasonably sure of the identification. This could
mean that the identification is simply of the most likely species
without any other good reason to choose it, or, less commonly,
that the identification fits by some criterion but I am suspicious
because it doesn't "look right". Some larval types may, of course,
include other closely-related species that have identical larval
features. If this is the case, it will be indicated by a plus
sign after the species name. If the features of the larval type
are unique, or all of the related species sharing the fin ray
counts are accounted for, then there will be no plus sign.
Diagnosis:
If necessary, I explain how the identification is
made in a diagnosis paragraph. The critical features
for the ID are highlighted. If it is needed, the
methods I have used for larval identification, such
as unique and specific morphology or fin ray counts
(U), the process of elimination (PE), DNA sequence
matching (DNA), or raising in captivity (R) are
indicated at the end of the diagnosis paragraph.
Analogues: For most larval descriptions,
the diagnosis paragraph is sufficient to separate
larval types, but, for some groups of species with
similar-appearing larvae, such as the gobies, I
often include an additional paragraph on how that
larval type can be separated from other closely-allied
larvae.
Terminology: In the text, larvae are immature
fishes caught in waters above or off the reef (usually
at night), typically with adaptations to open ocean
dwelling, such as transparency or silvery coloration,
extended spines and often a different set of markings
from juveniles of the species. Transitional larvae
are immature fishes caught in waters above or off
the reef (also usually at night) who have started
to develop juvenile markings and morphology. Transitional
recruits are fishes associated with the reef substrate
who have retained remnants of larval markings along
with their juvenile markings and morphology. They
typically are out during the day and behave as normal
juveniles. Recruits are newly-settled fishes associated
with the reef substrate (let's say less than a couple
of weeks on the reef). Juveniles are young fishes
on the reef who are no longer newly-settled.
Each
photograph is associated with a caption including
the identification to the lowest taxonomic level
possible, the standard length (SL) in millimeters
of the specimen, as well as the collection location
and collection code number. Lastly, important features
that need to be highlighted in the illustration
are often included in parentheses following the
collection information. The data associated with
the collection numbers (specific location, date,
etc.) can be found in my inventory
files, which are progressively being updated and
posted to the web.
PERMISSIONS
My photographs are freely available
for reproduction and use in non-commercial applications.
Please e-mail me for permissions, if warranted (e-mail
ben followed by at sign, then coralreeffish.com).
The web versions posted here are mostly about 500
pixels wide and run from 20 to 120 kb. I have larger
resolution versions, usually about 1.5 mb jpegs,
available for uses that require better resolution.
THE CARIBBEAN REEF FISH SPECIES LIST
(including the Gulf of Mexico)
Fish families following Randall's book,
but with updated, expanded, and revised species lists
excluding elasmobranchs
including numerous species found in non-reef
habitats (if the family has reef-associated members)
but sometimes excluding sets of deep-water
species (over ~30 m depth)
FAMILIES
(click on
the family name to jump down to species list)
Elops saurus Ladyfish
(US coastline to N Florida and Gulf of Mexico) Elops smithi Caribbean Ladyfish (new cryptic species
found in the Caribbean and S Florida)
FAMILY MEGALOPIDAE
Megalops atlanticus
Tarpon
FAMILY ALBULIDAE
Albula garcia (new
cryptic species, widespread Florida and Caribbean) Albula nemoptera Threadfin bonefish (estuarine) Albula vulpes Bonefish
FAMILY CLUPEIDAE
Alosa alabamae Alabama shad (anadromous, Gulf
of Mexico) Alosa chrysochlorus Skipjack shad (anadromous,
Gulf of Mexico)
Brevoortia gunteri Finescale menhaden (Gulf of
Mexico) Brevoortia patronus Gulf menhaden (Gulf of Mexico) Brevoortia smithi Yellowfin menhaden (Gulf of Mexico)
Pellona harroweri Coastal pellona (Panama to Brazil,
Pristigasteridae)
Sardinella aurita Round sardinella, Spanish sardine
(Atlantic Ocean) Sardinella janeiro (unresolved how it is different
from S. aurita, widespread; S. brasiliensis
is synonym)
FAMILY ENGRAULIDAE
Anchoa belizensis Belize anchovy (freshwater,
Belize) Anchoa cayorum Key anchovy Anchoa colonensis Narrow-striped anchovy (Caribbean
only, replaces A. hepsetus) Anchoa cubana (widespread) Anchoa filifera Longfinger anchovy Anchoa hepsetus Broad-striped anchovy (Gulf of
Mexico northwards, Venezuela southwards) Anchoa lamprotaenia Big-eye anchovy Anchoa lyolepis Shortfinger anchovy Anchoa mitchilli Bay anchovy (Gulf of Mexico northwards) Anchoa parva Little anchovy (Caribbean only, replaces
A. mitchilli) Anchoa spinifer Spicule anchovy (Panama to Brazil) Anchoa trinitatus Trinidad anchovy (Gulf of Venezuela
to Trinidad)
Anchovia clupeoides Zabaleta anchovy
Anchoviella blackburni (Gulf of Venezuela only) Anchoviella elongata Elongate anchovy (Belize to
Panama and Colombia) Anchoviella perfasciata (widespread)
Cetengraulis edentulus Atlantic anchoveta
Engraulis eurystole Silver anchovy (Florida northwards
and Venezuela southwards)
Lycengraulis grossidens Atlantic sabretooth anchovy
(Gulf of Venezuela eastward plus Belize)
(excluding numerous deep-water
taxa from 20-500 m depth)
Ahlia egmontis Key worm eel
Aplatophis chauliodus Tusky eel
Aprognathodon platyventris Stripe eel
Apterichtus ansp Academy eel
Bascanichthys bascanium Sooty eel (Gulf of Mexico
and Florida north) Bascanichthys inopinatus (Puerto Rico +?)
Bascanichthys scuticaris Whip eel (Gulf of Mexico
and Florida north)
Callechelys bilinearis Twostripe snake eel Callechelys guineensis Shorttail snake eel
Echiophus intercinctus Spotted spoon-nose eel Echiophus punctifer Stippled spoon-nose eel
Ichthyapus ophioneus Surf eel
Myrichthys breviceps Sharptail eel Myrichthys ocellatus Goldspotted eel
Myrophis anterodorsalis Longfin worm eel Myrophis platyrhynchus Broadnose worm eel Myrophis punctatus Speckled worm eel
Ophichthus cylindroideus Ophichthus gomesii Shrimp eel Ophichthus ophis Spotted snake eel Ophichthus puncticeps Palespotted snake eel Ophichthus rex (Gulf of Mexico)
Quassiremus ascensionis Black-spotted snake eel
FAMILY CONGRIDAE
(excluding numerous deep-water
taxa from 20-500 m depth)
FAMILY OPHIDIIDAE, SUBFAMILIES BROTULINAE
AND NEOBYTHITINAE
(excluding numerous deep-water
taxa from 20-8000 m depth)
Brotula barbata
Petrotyx sanguineus
FAMILY OPHIDIIDAE, SUBFAMILY OPHIDIINAE
(excluding numerous deep-water
taxa from 20-500 m depth)
Lepophidium brevivarbe
Ophidion beani (Florida and Gulf of Mexico) Ophidion grayi (Florida and Gulf of Mexico) Ophidion holbrookii (= Ophidion holbrooki) Ophidion josephi (Florida and Gulf of Mexico) Ophidion lagochila Ophidion marginatum Ophidion nocomis (Antilles) Ophidion selenops (Florida and Gulf of Mexico)
Ophidion welshi (Florida and Gulf of Mexico, incl.
Ophidion marginatum?)
Gramma dejongi Dejong's
basslet (new species 2010, S. Cuba only) Gramma linki Yellowlined basslet Gramma loreto Royal gramma Gramma melacara Blackcap basslet
Cynoscion acoupa (Panama to Argentina) Cynoscion arenarius (Florida and Gulf of Mexico) Cynoscion jamaicensis Cynoscion leiarchus (Nicaragua to Brazil) Cynoscion microlepidotus (Venezuela to Brazil) Cynoscion nebulosus Spotted weakfish (Florida and
Gulf of Mexico) Cynoscion nothus Silver seatrout (Florida and Gulf
of Mexico) Cynoscion similis (S Caribbean) Cynoscion virescens (Nicaragua to Brazil)
Larimus breviceps Shorthead drum Larimus fasciatus (Florida and Gulf of Mexico)
Leiostomus xanthurus Spot (Florida and Gulf of
Mexico)
Lonchurus elegans (S Caribbean) Lonchurus lanceolatus (S Caribbean)
Macrodon ancylodon King weakfish (SE Caribbean)
Menticirrhus americanus Southern kingfish Menticirrhus littoralis Menticirrhus saxatilis (Florida and Gulf of Mexico)
Micropogonias furnieri Micropogonias undulatus Atlantic croaker (Florida
and Gulf of Mexico)
Nebris microps (Colombia to Brazil)
Odontoscion dentex Reef croaker
Ophioscion costaricensis (= Stellifer microps?,
Costa Rica and Suriname) Ophioscion panamensis (valid?, Belize to Panama) Ophioscion punctatissimus (Panama to Brazil, and
Puerto Rico)
Eugerres brasilianus (Belize, Cuba south to Brazil?) Eugerres mexicanus (Mexico Guatemala FW, Rio Usumacinta) Eugerres plumieri Striped mojarra (Florida, Gulf
of Mexico to Cuba, Panama?)
Starksia
atlantica Bahamas smootheye blenny (now part of
a species complex; Bahamas) Starksia
culebrae Antillean checkered blenny (Antilles) Starksia elongata Elongate blenny (widespread) Starksia fasciata Blackbar blenny (Bahamas and
Greater Antilles) Starksia
greenfieldi Tobago chessboard blenny (new
species 2011, Testigos to Tobago) Starksia
guttata Southern checkered blenny (SE Caribbean) Starksia hassi Ringed blenny (widespread Caribbean) Starksia
langi Bay chessboard blenny (new
species 2011, Bay of Honduras and south) Starksia
lepicoelia Bahamas blackcheek blenny (part of
a species complex; Bahamas) Starksia leucovitta Whitesaddle blenny (new species
2003, Navassa) Starksia melasma Blackspot blenny (new species
2003, PR and St.Croix) Starksia
nanodes Dwarf blenny (species complex) Starksia
ocellata Florida checkered blenny (now part of
a species complex; Florida) Starksia
occidentalis Western checkered blenny (MAB to
Panama) Starksia rava (new species 2003, Tobago) Starksia
robertsoni Panama blackcheek blenny (new
species 2011, Panama) Starksia sella (new species 2003, Tobago) Starksia
sangreyae Bay smootheye blenny (new
species 2011, Belize) Starksia
sluiteri Southern chessboard blenny (now part
of a complex; S. Neth. Antilles) Starksia smithvanizi Brokenbar blenny (new species
2003, Antilles) Starksia
springeri Southern smootheye blenny (new
species 2011, S. Neth. Antilles) Starksia starcki Key blenny (Florida to Belize) Starksia
variabilis Colombian checkered blenny (Colombia) Starksia
weigti Bay blackcheek blenny (new
species 2011, Bay of Honduras) Starksia
williamsi Antillean blackcheek blenny (new
species 2011, N. Antilles) Starksia y-lineata Forkbar blenny (Cayman and Nicaraguan
Shelf)
FAMILY CHAENOPSIDAE
Acanthemblemaria aspera
Roughhead blenny (widespread, not SE Caribbean) Acanthemblemaria betinensis Speckled blenny (Costa
Rica, Panama, and Colombia) Acanthemblemaria chaplini Papillose blenny (FL, Bahamas,
Cuba, Yucatan) Acanthemblemaria cubana (Cuba, Panama) Acanthemblemaria greenfieldi False papillose blenny
(Belize & W Caribbean Islands) Acanthemblemaria harpeza (Navassa) Acanthemblemaria johnsoni (Tobago, NE Venezuela) Acanthemblemaria maria Secretary blenny (widespread,
not Colombia, Neth Ant, or VZ) Acanthemblemaria medusa (SE Caribbean) Acanthemblemaria paula Dwarf spinyhead blenny (Belize
and Panama) Acanthemblemaria rivasi Spotjaw blenny (Costa Rica,
Panama, Colombia, Venezuela) Acanthemblemaria spinosa Spinyhead blenny (widespread,
not mainland S. Amer?)
Microgobius
carri Seminole goby (widespread) Microgobius gulosus Clown goby (GOM and US east
coast) Microgobius
meeki Microgobius microlepis Banner goby (widespread) Microgobius
signatus Microgobius thalassinus Green goby (GOM and US
east coast)
n. sp.: saber goby (deep walls, Bahamas, Puerto Rico,
and Belize)
FAMILY PTERELEOTRIDAE
Ptereleotris
calliura blue dartfish (Florida and N Gulf of Mexico) Ptereleotris
helenae Hovering dartfish (widespread) Ptereleotris randalli bluestripe dartfish (SE Caribbean
to Brazil)
