The phylogenetic significance of colour patterns in
marine teleost larvae
CAROLE C. BALDWIN*
Division of Fishes, National Museum of Natural History, Smithsonian Institution, Washington, DC
20560, USA
Received 5 April 2012; revised 2 March 2013; accepted for publication 4 March 2013
Ichthyologists, natural-history artists, and tropical-fish aquarists have described, illustrated, or photographed colour
patterns in adult marine fishes for centuries, but colour patterns in marine fish larvae have largely been neglected.
Yet the pelagic larval stages of many marine fishes exhibit subtle to striking, ephemeral patterns of chromatophores
that warrant investigation into their potential taxonomic and phylogenetic significance. Colour patterns in larvae of
over 200 species of marine teleosts, primarily from the western Caribbean, were examined from digital colour
photographs, and their potential utility in elucidating evolutionary relationships at various taxonomic levels was
assessed. Larvae of relatively few basal marine teleosts exhibit erythrophores, xanthophores, or iridophores (i.e.
nonmelanistic chromatophores), but one or more of those types of chromatophores are visible in larvae of many basal
marine neoteleosts and nearly all marine percomorphs. Whether or not the presence of nonmelanistic chromato-
phores in pelagic marine larvae diagnoses any major teleost taxonomic group cannot be determined based on the
preliminary survey conducted, but there is a trend toward increased colour from elopomorphs to percomorphs. Within
percomorphs, patterns of nonmelanistic chromatophores may help resolve or contribute evidence to existing
hypotheses of relationships at multiple levels of classification. Mugilid and some beloniform larvae share a unique
ontogenetic transformation of colour pattern that lends support to the hypothesis of a close relationship between
them. Larvae of some tetraodontiforms and lophiiforms are strikingly similar in having the trunk enclosed in an
inflated sac covered with xanthophores, a character that may help resolve the relationships of these enigmatic taxa.
Colour patterns in percomorph larvae also appear to diagnose certain groups at the interfamilial, familial,
intergeneric, and generic levels. Slight differences in generic colour patterns, including whether the pattern
comprises xanthophores or erythrophores, often distinguish species. The homology, ontogeny, and possible functional
significance of colour patterns in larvae are discussed. Considerably more investigation of larval colour patterns in
marine teleosts is needed to assess fully their value in phylogenetic reconstruction.
? 2013 The Linnean Society of London, Zoological Journal of the Linnean Society, 2013, 168, 496?563.
doi: 10.1111/zoj.12033
ADDITIONAL KEYWORDS: chromatophore ? erythrophore ? fish larvae ? iridophore ? western Caribbean
? xanthophore.
INTRODUCTION
The pelagic larval stages of most marine fishes
inhabit an evolutionary arena distinct from that of
adults, and morphological specializations that pre-
sumably enhance survival in the planktonic realm
have evolved in numerous teleost groups (Moser,
1981; Moser et al., 1984). In a few cases, marine fish
larvae and adults are so different morphologically
that they were initially classified as separate genera
or families (Cohen, 1984; Johnson et al., 2009). Dis-
tinctive pigment patterns are among the transient
features that characterize the pelagic larval phase
of marine fishes (Moser et al., 1984), and Kendall,
Ahlstrom & Moser (1984) included pigment patterns
in their list of characters of early life-history stages
commonly utilized in taxonomic and systematic
*E-mail: baldwinc@si.edu
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Zoological Journal of the Linnean Society, 2013, 168, 496?563. With 52 figures
? 2013 The Linnean Society of London, Zoological Journal of the Linnean Society, 2013, 168, 496?563496
studies of fishes. Although pigment patterns were
noted by Kendall et al. (1984) to be among the most
useful larval characters at specific and generic levels,
only melanophores, not other types of chromato-
phores, were discussed. Chromatophores of poikilo-
thermic vertebrates are dermal pigment units that
comprise light-absorbing erythrophores (red/orange
pigment), xanthophores (yellow pigment), and
melanophores (dark brown/black pigment), as well as
light-reflecting iridophores (structural colour, often
silver/blue but many colours possible) ? e.g. Bagnara
& Hadley (1973), Grether, Kolluru & Nersissian
(2004). Colour patterns in adult marine fishes are
well documented, particularly those of tropical reef
fishes, but pigment other than melanin in marine fish
larvae has received little attention. This is not sur-
prising because erythrophores and xanthophores fade
upon conventional preservation (melanophores gener-
ally do not), and plankton samples are typically pre-
served soon after capture. Only when larvae have
been observed, illustrated, or photographed prior to
preservation has colour in marine teleost larvae been
reported in the scientific literature (e.g. Brownell,
1979; Baldwin & Smith, 2003; Yasir & Qin, 2007;
Baldwin et al., 2009a; 2011; Miller, 2009; Wittenrich,
Baldwin & Turingan, 2010).
The transient colour patterns in marine fish larvae
generally bear little resemblance to those of adults
and may result from different ontogenetic (larval and
adult) populations of chromatophores (Nakamura
et al., 2010). The ontogeny of pigment patterns in
marine fishes is poorly understood relative to that
of many freshwater fishes, especially zebrafishes
(Danio spp.), which have been studied extensively (e.g.
Johnson et al., 1995; Parichy et al., 2000; Parichy,
2003, 2006; Kelsh, 2004; Budi, Patterson & Parichy,
2011). In Danio, the colour pattern of the recently
hatched fish transforms directly into the adult colour
pattern through incorporation of embryonic chromato-
phores and differentiation of new chromatophores
from stem cells at metamorphosis (Parichy, 2003,
2006). There is no pelagic larval stage in Danio and
most other freshwater fishes comparable to that in
most marine fishes, and there is no accompanying
distinctive pigment phase between the recently
hatched and adult stages (Bagenal & Nellen, 1980;
Kendall et al., 1984). Colour variation in this ?extra?
pigment phase in marine fish larvae was the focus of
this study. The presence of a specialized larval stage
in certain freshwater fishes that are evolutionarily
derived from marine fishes corroborates the distinc-
tion between early life stages of most marine and
freshwater fishes. For example, larvae of basal fresh-
water percoids typically lack the head spination char-
acteristic of pelagic larvae of basal marine percoids,
but young Lates from Lake Tanganyika retain head
spination that evolved in their marine, Indo-Pacific
ancestors (Kinoshita & Tshibangu, 1997).
Colour patterns in the young of some freshwater
fishes are highly conserved and thus of little potential
phylogenetic value. For example, Quigley et al. (2004)
noted that the young of several Danio species have
virtually indistinguishable pigment patterns, and
Kelsh (2004) noted the same for five Danio species
and Tanichthys albonubes. In contrast, Baldwin &
Smith (2003) and Baldwin et al. (2009a, 2011) high-
lighted the utility of patterns of erythrophores
and xanthophores in species identification of larval
Gobiidae and Apogonidae, and Baldwin et al. (2011)
suggested that chromatophore patterns may be of
value in resolving the generic classification of western
Atlantic Apogonidae. The potential utility of larval
colour patterns at higher taxonomic levels has not
been properly investigated. Kendall et al. (1984)
noted that pigment (melanophore) patterns in fish
larvae are of limited use in systematic studies in part
because convergence has resulted in the occurrence
of strikingly similar patterns in unrelated groups.
Convergent evolution can be detected if pigment char-
acters are examined in a phylogenetic context. A
phylogenetic tree that includes clades composed pri-
marily of marine teleosts was used herein to examine
the distribution of nonmelanistic chromatophores in
larvae among major groups of marine teleosts. The
exclusion of clades of freshwater fishes renders the
tree but a partial view of teleost phylogeny, but in this
first attempt to provide comparative information on
colour patterns among marine teleost larvae, it is
desirable to contemplate the results from both broad
and more focused phylogenetic perspectives. Numer-
ous recent hypotheses of phylogenetic relationships
among various groups of teleosts were utilized for
comparisons at lower taxonomic levels. The purposes
of this paper were to describe the distribution of
nonmelanistic chromatophores among a broad spec-
trum of marine teleost larvae and comment on the
potential of selected chromatophore patterns to
inform phylogeny.
MATERIAL AND METHODS
This study was based largely on colour photographs of
fish larvae collected off Belize, Central America.
Larvae were collected in a plankton net of 505 mm
mesh fitted onto a 0.5 ? 1 m rectangular frame and
deployed from a dock at Carrie Bow Cay (16?48.5?N,
88?05?W). Specimens were submerged in a photo
tank, photographed with a Nikon D1 or Fuji FinePix
S3 digital camera, and then tissue sampled for DNA
analysis prior to preservation. Species identification
of larvae was accomplished by matching cytochrome
oxidase-c subunit I (COI) sequences (DNA barcodes)
COLOUR IN MARINE TELEOST LARVAE 497
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of larvae to those of known adults (Weigt et al., 2012).
To date this protocol has resulted in the identification
of larvae of approximately 170 Caribbean fish species.
A few larvae had no species-level matches in the
Smithsonian DNA database or the Barcode of Life
Database (BOLD; http://www.boldsystems.org/views/
login.php) and are identified only to genus in the
figure legends. Taxonomic coverage was increased
by examining images of marine fish larvae from
specimens collected off the east coast of South Africa,
Florida Straits, Hawaii, and Cozumel, as well as
several Indo-Pacific species reared from aquarium
specimens. Connell?s (2007) website of early life-
history stages of South African marine fishes is
impressive in scope, and although few of his images
are reproduced here, numerous references to his
site and uniform resource locators (URLs) to specific
pages are provided herein. All photo editing was
carried out by the author. Images of most larvae were
cut from their original photographic backgrounds and
placed on a uniform background using the MaskPro 4
plug-in for Adobe Photoshop with the aid of a Wacom
table and stylus. In many cases, the transparent fins
of larvae were difficult to see, and shapes of fins may
be approximations. Photo credits are provided in
figure legends, and affiliations for contributors other
than the author are as follows: Allan Connell, South
African Institute of Aquatic Biology; Donald Griswold,
formerly Smithsonian Institution; Cedric Guigand,
University of Miami, Rosensteil School of Marine
Science; Joshua Lambus, J. Lambus Photography,
Hawaii; Michael Miller, The University of Tokyo;
Julie Mounts, formerly Smithsonian Institution;
Christopher Paparo, The Long Island Aquarium;
David Smith, Smithsonian Institution; Lee Weigt,
Smithsonian Institution; and Matthew Wittenrich,
University of Florida. No professional affiliations are
available for two additional contributors, Matthew
D?Avella and Donald Hughes. Localities from which
specimens were collected or photographed are pro-
vided in figure legends: BLZ refers to Belize, and it is
followed by a four- or five-digit number that repre-
sents the Smithsonian DNA number. DNA barcodes of
Belizean fish larvae are publicly available on BOLD
under the project names APG, BATHY, BZLWA,
BZLWB, BZLWC, BZLWD, BZLWE, CORY, PHAE,
and RYP. GenBank accession numbers for COI
sequences are as listed in Baldwin et al. (2009a,
2009b, 2011), Tornabene et al. (2010), Baldwin &
Weigt (2012), and Weigt et al. (2012). Not all images
examined were reproduced in this paper. A complete
list of larval-fish images examined for colour patterns
is given in Appendix, which also includes URLs for
some of the supplementary images examined.
In the descriptions of nonmelanistic chromatophore
patterns, ?yellow pigment? and ?xanthophores? have
been used interchangeably, as have ?orange pigment?
(or ?red pigment?) and ?erythrophores?. The presence or
absence of yellow and orange pigment was equated
with the presence or absence of xanthophores and
erythrophores, respectively, and the presence or
absence of light-reflecting pigment was equated with
the presence or absence of iridophores. Assessments
of types of nonmelanistic chromatophores were based
solely on examination of fresh specimens and colour
photographs, not histological examination.
The classification of Wiley & Johnson (2009) was
selected for use in this study. This classification was
chosen over other more commonly used fish classifi-
cations because it is a Linnaean classification based
on monophyletic groups. It was essential in this work
to examine the distribution of nonmelanistic chro-
matophores in fish larvae among major teleost groups
from an evolutionary perspective, and constructing a
teleost phylogeny from the Wiley & Johnson (2009)
synapomorphy-based classification was easily accom-
plished (Fig. 1).
RESULTS
DISTRIBUTION OF NONMELANISTIC CHROMATOPHORES
IN BASAL MARINE TELEOSTS AND
NEOTELEOSTS (FIG. 1)
Larvae of most basal teleosts ? elopiforms, albuli-
forms, anguilliforms, and clupeiforms ? do not exhibit
xanthophores, erythrophores, or iridophores (Figs 2,
3). Exceptions occur within the anguilliform families
Ophichthidae, Muraenidae, Congridae, and Nettasto-
matidae. Miller, D?Avella & Tsukamoto (2010: figs 1,
2) described xanthophores along the bodies of two
unidentified ophichthid leptocephalus larvae vide-
otaped swimming at night off Hawaii (Fig. 4A, B). An
image of the ophichthid Neenchelys (Miller, 2009:
fig. 57A) has yellow pigment on the snout, anterior
portion of the oesophagus, and on the gut swellings
(Fig. 5A). Another ophichthid leptocephalus, Myrich-
thys breviceps, captured off Belize bears similar con-
spicuous xanthophores on the gut swellings (Fig. 4C).
Tawa et al. (2012: fig. 2) published a colour image of a
muraenid leptocephalus (Strophidon ui) that exhibits
xanthophores in front of and behind the eye, and
Miller (2009) provided images (Fig. 5B, E, F herein) of
muraenid leptocephali with similar yellow pigment
adjacent to the eye (and in one case also scattered
on the head). Miller (2009) noted the presence of
yellow pigment on the dorsal surface of the eye in
some congrids and ophichthids (Fig. 5D) and yellow
pigment on the snout and anterior portion of the
oesophagus in the nettastomatid Saurenchelys
(Fig. 5C). Identification of more anguilliform larvae
is needed to determine the taxonomic distribution of
498 C. C. BALDWIN
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Figure 1. Teleost phylogeny based on the classification of Wiley & Johnson (2009). Only marine taxa for which colour in
larvae was examined are included. The number of squares associated with each teleost order is equivalent to the number
of species examined. Open squares indicate the absence of xanthophores, erythrophores, and iridophores. Yellow squares
indicate the presence of one or more of these types of chromatophores. The shaded rectangle denotes Johnson &
Patterson?s (1993) Euacanthopterygii.
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Figure 2. Elopomorpa. A, Elops saurus, 31 mm Standard Length (SL), BLZ 8327. B, Albula vulpes, 54 mm SL, BLZ 8420.
C, Megalops atlanticus, 23 mm SL, BLZ 5457. D, Myrophis punctatus, 58 mm SL, Belize. E, Aprognathodon platyventris,
75 mm SL, BLZ 5322. F, Myrophis platyrhynchus, 67 mm SL, BLZ 8392. G, Ahlia egmontis, 70 mm TL, BLZ 7174. H,
Gymnothorax moringa, 71 mm SL, BLZ 8469. Photos A, C, F by Lee Weigt and Carole Baldwin; B, G, H by Julie Mounts
and David Smith; D, E by Julie Mounts and Carole Baldwin.
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xanthophores, but the presence of yellow pigment on
gut swellings in ophichthids, on the snout and ante-
rior oesophagus in ophichthids and nettastomatids, in
front of and behind the eye in muraenids, and dorsal
to the eye in congrids and ophichthids might repre-
sent diagnostic patterns and therefore warrant addi-
tional study. Most leptocephali collected off Belize
lack yellow pigment, yet many are members of fami-
lies discussed above that have it. Anguilliform lepto-
cephali from Belize that lack yellow pigment (Fig. 2)
include Gymnothorax moringa (Muraenidae), Morin-
gua edwardsi (Moringuidae), Chilorhinus seunsoni
(Chlopsidae), and Ahlia egmontis, Aprognathodon
platyventris, Myrophis punctatus, and Myrophis pla-
tyrhynchus (Ophichthidae). Based on the absence of
xanthophores in larval albuliforms and elopiforms, it
is reasonable to assume that their absence is ances-
tral for anguilliforms. The absence of yellow pigment
in leptocephali of Moringua edwardsi and Synapho-
branchidae (Miller, 2009) provides corroborative evi-
dence based on the basal positions of Moringuidae
and Synaphobranchidae in the molecular anguilli-
form phylogeny of Tang & Fielitz (2012). Anguilliform
taxa that exhibit yellow pigment in the leptocephalus
stage ? some Congridae, Nettastomatidae, Ophichthi-
dae, and muraenine Muraenidae ? occupy more distal
phylogenetic positions in the order (Tang & Fielitz,
2012), but they do not constitute a monophyletic
assemblage. It seems likely that xanthophores in
larvae evolved independently within the various fami-
lies of Anguilliformes that exhibit them.
Little information is available on the presence or
absence of nonmelanistic chromatophores in larvae of
basal marine neoteleosts (Fig. 1). Recently hatched
larvae of one phosichthyid stomiatiform from off
South Africa lack erythrophores and xanthophores,
whereas a preflexion larva of a melanostomiatid
has yellow pigment on the head and body (Connell,
2007; see links to images in Appendix). Two aulopi-
form families (Synodontidae and Giganturidae) also
have larvae that lack orange or yellow coloration
(Figs 6A?C, 7A), but an unidentified aulopiform larva
has vibrant erythrophores and xanthophores on
enlarged pectoral fins as well as on the dorsal and
caudal fins (Fig. 7B). A preflexion myctophiform larva
from off South Africa has yellow pigment on the head
and body, but yellow pigment is not as evident in
a larger preflexion larva (Connell, 2007; Appendix). A
Figure 3. Otomorpha (Clupeiformes). A, Anchoa sp., 26 mm Standard Length (SL), BLZ 7162. B, Harengula clupeola,
15 mm SL, BLZ 8419. C, Jenkensia lamprotaenia, 15 mm SL, BLZ 8417. Note: the red coloration behind the head in
J. lamprotaenia is associated with the circulatory system, not chromatophores. Photos by Julie Mounts and David Smith.
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postflexion larval myctophiform from the Florida
Straits lacks erythrophores and xanthophores
(Fig. 7C). In gadiforms, preflexion larvae of an uni-
dentified gadid from South Africa have numerous
xanthophores on the head and body (Connell, 2007;
Appendix), whereas the bregmacerotid Bremaceros
from Belize lacks orange and yellow pigment
(Fig. 6D). Both lampridiform larvae examined (a tra-
chipterid and Lampris) have nearly the entire body
and some fins covered with erythrophores (Fig. 8).
The single stephanoberyciform larva examined, the
bizarre ?mirapinnid? larva of the whalefish family
Cetomimidae (Johnson et al., 2009), has yellow
pigment on the body and fins (Fig. 9), and the single
zeiform examined, Zeus faber (Connell, 2007; Appen-
dix), also has a small bit of yellow pigment in both
pre- and postflexion stages that is mixed with and
largely occluded by melanophores. All preflexion and
postflexion beryciform larvae have erythrophores,
xanthophores, iridophores, or some combination of
those chromatophores (Fig. 10).
To summarize the comparative information for
basal marine teleosts and neoteleosts, several basal
neoteleost orders are similar to anguilliforms in
having some larvae that exhibit nonmelanistic chro-
matophores and others that do not. Xanthophores are
the only types of chromatophores observed in larval
anguilliforms and in the single larval stomiatiform
examined that has nonmelanistic chromatophores.
Xanthophores are lacking in larval elopiforms, albu-
liforms, and the few marine clupeiforms examined.
Xanthophores are not lacking in all otomorphs,
however, as they have been documented in freshwater
Cyprinidae (e.g. Johnson et al., 1995; Parichy et al.,
2000; Parichy, 2003). Erythrophores are first observed
phylogenetically in marine fish larvae in aulopiforms
and are prominent in lampridiforms and some bery-
ciforms. Possibly erythrophores in larvae are phylo-
genetically significant at the level of Eurypterygia
(Aulopiformes and above, Fig. 1). Iridophores first
appear phylogenetically in larval beryciforms and
could provide corroborative evidence for Johnson &
Figure 4. Elopomorpha. A, B, in situ images of an ophichthid leptocephalus off Hawaii captured from video by Matthew
D?Avella, Kona, Hawaii (B previously published in Miller et al., 2010, reproduced here with permission of the copyright
holder). C, Myrichthys breviceps, 116 mm Standard Length, BLZ 8467, photo by Julie Mounts and David Smith.
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Patterson?s (1993) Euacanthopterygii (Beryciformes
and above, shaded rectangle in Fig. 1). Iridophores
are lacking in larvae of the marine elopomorphs,
otomorphs, and other basal neoteleosts examined;
however, they are present in some recently hatched
freshwater Otomorpha (e.g. Danio ? Johnson et al.,
1995; Parichy et al., 2000; Parichy, 2003). As dis-
cussed below, an ontogenetic transition involving
iridophores in atheriniforms, beloniforms, gasterostei-
forms, and mugiliforms, which are members of the
Figure 5. Elopomorpha. A, Neenchelys sp. (Ophichthidae). B, E, F, Muraenidae. C, Saurenchelys sp. (Nettastomatidae).
D, Ophichthidae. Modified from Miller (2009) with the permission of the copyright holder.
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percomorph series Smegmamorpharia, may help diag-
nose that group. Within percomorphs (Mugiliformes
and above, Fig. 1), erythrophores, xanthophores, iri-
dophores, or some combination of them are present in
larvae of every group and in almost every species
examined, and the remainder of this paper is devoted
to descriptions of percomorph colour patterns and
discussions of their potential phylogenetic signifi-
cance at various taxonomic levels.
PERCOMORPHACEA ? SMEGMAMORPHARIA
Johnson & Patterson?s (1993) Smegmamorpha
comprise seven orders of percomorph fishes, but the
Figure 6. Neoteleostei (Aulopiformes, Myctophiformes) and Acanthomorphata (Gadiformes). A, Synodus synodus, 39 mm
Standard Length (SL), Belize. B, Saurida sp., 26 mm SL, BLZ 8329. C, Saurida sp., 32 mm SL, BLZ 8398. D, Bregmaceros
sp., 12 mm SL, BLZ 4242. Note: the red coloration on the head and body in the Bregmaceros image appears to be
associated with the circulatory system, not dermal pigment. Photo A by Julie Mounts and Carole Baldwin; B Lee Weigt
and Carole Baldwin; C Julie Mounts and David Smith; D Lee Weigt and David Smith.
504 C. C. BALDWIN
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monophyly of the group has been questioned. For
example, Springer & Orrell (2004) did not find
support for it based on extensive analysis of the
gill-arch musculature and skeleton. Smegmamorph
orders for which larvae were examined in this study
are Atheriniformes, Beloniformes, Gasterosteiformes,
and Mugiliformes. Larval mugilids and exocoetids
examined share a striking ontogenetic transition in
colour pattern. Small larvae are covered with pale
orange/yellow pigment and melanophores but also
bear some conspicuous iridophores (Fig. 11A, B).
In larger larval specimens, the orange colour is no
longer present, and the fishes are silvery and covered
almost entirely with iridophores (Fig. 11C, D). This
ontogenetic transition was not observed in any non-
smegmamorph fishes and would appear to provide
corroborative evidence for a close relationship
between Mugiliformes and Atherinomorpha, which
comprises Atheriniformes and Beloniformes. Stiassny
(1990, 1993) proposed a sister-group relationship
between mugiliforms and atherinomorphs despite
strong evidence also suggesting that mugilids are
closely related to perciforms. This same colour tran-
sition occurs in hemirhamphids and belonids but
apparently at a later stage of development. Larvae
of Platybelone and Hemirhamphus are covered with
erythrophores and xanthophores, respectively, and
bear scattered iridophores (Fig. 12A, B, F). A juvenile
Hemirhamphus still bears xanthophores, but the
abdominal region is silvery (Fig. 12C), and adults of
both genera are entirely silver. Larval gasterostei-
forms also have yellow/orange chromatophores and
melanophores covering most of the body as in young
mugilids and beloniforms (Fig. 12D, E), and although
they never undergo the transition to a silvery body,
at least some gasterosteiform larvae have silvery
pigment on the abdomen (Fig. 12E, G). Other gaster-
osteiform larvae known only from early preflexion
stages ? Fistularia, Aulostomus, Aeoliscus ? also
exhibit numerous xanthophores mixed with melano-
phores (Connell, 2007; Appendix), but later larval
stages are needed to determine whether or not they
develop iridophores. Some larval beloniforms may
have a different larval trajectory in that they appear
to lack the early yellow/orange stage. For example, a
5.6 mm Notochord Length (NL) larva of Oxyporham-
phus micropterus from off South Africa is mostly pale,
has melanophores and iridophores along the entire
dorsal margin of the body and along a portion of the
ventral midline, and has a silvery/blue gut (Connell,
2007; Appendix). Possibly there is a small bit of
yellow pigment on the oesophagus and gut, but it was
difficult to determine if this is dermal pigment or part
of the gut contents. It is also possible that the yellow/
orange stage develops after notochord flexion, as all
mugilid, beloniform, and gasterosteiform larvae in
which that colour phase was observed had under-
gone flexion. A 13-mm Standard Length (SL) Hirun-
dichthys has only melanophores and iridophores
(Fig. 13A), but younger larvae are unknown. Larval
atherinids have few or no orange or yellow chromato-
phores (smallest specimen examined 7.0 mm SL ?
Fig. 13B) and have a dense covering of iridophores on
the gut. Note the striking similarity between a 14-mm
Figure 7. Neoteleostei (Aulopiformes, Myctophiformes) A, Gigantura sp., Florida Straits, photo by Cedric Guigand
(previously published in Pineda, Hare & Sponaugle, 2007). B, unknown aulopiform, Hawaii, photo by Joshua Lambus. C,
unknown myctophiform, Florida Straits, photo by Cedric Guigand.
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Figure 8. Acanthomorphata (Lampridiformes). Top, unknown (probably Trachipteridae), Hawaii, photo by Joshua
Lambus. Bottom, Lampris guttatus, Florida Straits, photo by Cedric Guigand.
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SL atherinid larva and an unidentified beloniform
larva in general appearance and the presence
of iridophores on the gut that reflect bright blue
(Fig. 13C, D).
PERCOMORPHACEA ? INCERTAE SEDIS
Acanthuriformes
Acanthuriformes are represented among the larvae
examined by two species of Acanthurus surgeonfishes
from Belize (Fig. 14) and an earlier stage of an uni-
dentified species of Acanthurus from the Florida
Straits (Fig. 15). The specialized, pelagic ?acronurus?
larval stage (e.g. Leis & Richards, 2004) is mostly
transparent with a band of silver- or blue-reflecting
iridophores from the dorsum to the pelvis that encom-
passes the orbit and gut. There is little if any orange
or yellow pigment in the acronurus stage, but an
image of preflexion larvae of Acanthurus mata
(Connell, 2007; Appendix) shows xanthophores on the
upper jaw and an internal streak of xanthophores
beneath the anterior section of the notochord. Rela-
tionships of acanthuriforms are unclear, but a close
relationship with tetraodontiforms has been proposed
(e.g. Tyler, 1980; Rosen, 1984). Tetraodontiform larvae
(see ?Tetraodontiformes? below) typically have irido-
phores on the gut and sometimes over much of the
body, but they are otherwise very different from acan-
thuriforms in having much more colour (xanthophores
or bronze iridophores), melanophores (sometimes in
striking patterns), or both.
Blenniiformes
Blenniiformes are represented in the Belize material
by larval chaenopsids and labrisomids. Larvae of both
families have an elongate body and share the follow-
ing chromatophore pattern: melanophores on the
ventral midline associated with the anal-fin base and
internally along the dorsal margin of the vertebral
column, orange or yellow pigment on the temporal
region of the head, internal orange pigment along the
vertebral column, and orange chromatophores at the
base of the caudal fin (Fig. 16). Colour patterns vary
little among some genera (e.g. Acanthemblemaria of
the Chaenopsidae, Labrisomus of the Labrisomidae ?
Fig. 16A?D), but the single larval specimen of Para-
clinus (Labrisomidae) differs in having prominent
orange chromatophores mixed with the melanophores
along the anal-fin base (Fig. 16E). Additional Paracli-
nus species are needed to determine whether or not
this is a generic pattern. The chromatophore pattern
on the body could not be interpreted well in an in situ
image of a larval fish from Hawaii identified by G. D.
Johnson (pers. comm.) as a blenniid (image not repro-
duced here), but the fish is unlike other blennioids in
having enlarged pectoral fins with xanthophores on
the membranes between fin elements.
Caproiformes
Caproiformes are represented in the images exam-
ined only by preflexion Antigonia rubescens from
South Africa (Connell, 2007; Appendix). Recently
Figure 9. Acanthomorphata (Stephanoberyciformes). Cetomimidae, Mexico. Photo by Donald Hughes.
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Figure 10. Acanthomorphata (Beryciformes). A, Berycidae. B?D, Holocentridae. All from Florida Straits. Photographs by
Cedric Guigand. A, B, and D previously published in Cowen et al. (2007), Thorrold, Zacherl & Levin (2007), and Gaines
et al. (2007), respectively.
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hatched larvae have xanthophores on the body, but no
information is available for larger specimens.
Carangiformes
Carangiformes are not common in larval-fish collec-
tions off Belize, but an 8.0-mm SL Trachinotus
(Carangidae) has most of the body covered with
orange chromatophores and silver-reflecting irido-
phores (Fig. 17). A colour image of a young carangid,
Selene (Fig. 18C), shows yellow/orange chromato-
phores on a somewhat silvery body as well as on most
fins. A 5.1-mm preflexion larva of the coryphaenid,
Coryphaena hippurus, has xanthophores mixed with
melanophores from the tip of the snout posteriorly
to the base of the incipient caudal fin (Fig. 18A).
A postflexion Coryphaena hippurus larva (Fig. 18B)
and a reared postflexion rachycentrid, Rachycentron
canadum (D. Benetti, pers. comm.), have some yellow
pigment mixed with melanophores on the body, but
the solid yellow ground coloration present in preflex-
ion Coryphaena is absent. Colour patterns are not
known for other carangiforms (Nematistiidae, Eche-
neididae), and no conclusions can be drawn regarding
the phylogenetic significance of colour for the order or
for families, genera, and species within. The bars of
pigment in postflexion Coryphaena (Fig. 18B) resem-
ble those of Selene in having some yellowish colora-
tion mixed with melanophores, and this character
could be significant at the ordinal level. Phylogenetic
analysis of both mitochondrial (Miya, Satoh &
Nishida, 2005) and nuclear (Little, Lougheed &
Moyes, 2010) data suggested a close relationship
between carangiforms and pleuronectiforms, but
there is nothing obvious in the colour patterns
of larvae to support this (see ?Pleuronectiformes?
below). Nuclear DNA data have also suggested a
relationship between carangids and istiophorids
(Little et al., 2010), which is discussed under ?Scom-
briformes? below.
Gobiesociformes
Gobiesociformes comprise the Gobiesocidae, Callio-
nymidae, and Draconettidae. Colour information in
larvae is available for several callionymids and one
gobiesocid. Wittenrich et al. (2010) described morpho-
logical development of laboratory-reared Synchiropus
splendidus and noted a complete colour transition
from yellow in early-stage larvae to nearly solid
orange at 8 days posthatching (Fig. 19A, B). Connell
(2007) provided a colour image of a 1.8-mm NL larva
of Callionymus sp. from South Africa (Fig. 19C) and
one of a 1.6-mm NL larva of Draculo (Appendix)
that are nearly identical to the similar-sized (2 days
posthatching) larva of Syn. splendidus (Fig. 19A) in
having the head and body covered in prominent xan-
thophores except for the posterior fin fold. The colour
pattern in larger larvae of the South African Calliony-
mus is unknown, but postflexion larvae of Calliony-
mus bairdi from Belize (Fig. 19D) are nearly solid
orange like Synchiropus. Additional material is
needed to determine whether the observed yellow-to-
orange colour transition is diagnostic of callionymids
or if it is also present in other gobiesociform families.
A 4.0-mm SL larva of the gobiesocid Acyrtops from
Belize has yellow/orange coloration on the head and
scattered over most of the body (Fig. 19E), possibly a
very similar pattern as in the callionymids but with
the erythrophores contracted. Larger and smaller
Figure 11. Percomorphacea (Mugiliformes and Beloniformes). A, Mugil cephalus, 3.5 mm Standard Length (SL), BLZ
6045. B, Prognichthys occidentalis, 11 mm SL, BLZ 7186. C, Mugil sp., 8.5 mm SL, BLZ 7069. D, Exocoetidae, 16 mm SL,
BLZ 6014. Photos A, D by Lee Weigt and Carole Baldwin; B, C by Julie Mounts and David Smith.
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Figure 12. Percomorphacea (Beloniformes, Gasterosteiformes). A, Platybelone argalus, 15 mm Standard Length (SL),
BLZ 6131. B, Hemirhamphus brasiliensis, 19.5 mm SL, BLZ 7071. C, Hemirhamphus balao, 34 mm SL, BLZ 7304.
D, Cosmocampus albirostris, 8.5 mm SL, BLZ 6414. E, Penetopteryx nanus, 19 mm SL, BLZ 8337. F and G, close-up
photographs of Platybelone argalus (A) and Penetopteryx nanus (E), respectively, showing abdominal iridophores. Photos
A, D, E by Lee Weigt and Carole Baldwin; B, C by Julie Mounts and David Smith.
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Figure 13. Percomorphacea (Beloniformes, Atheriniformes). A, Hirundichthys affinis, 13 mm Standard Length (SL), BLZ
4217. B, Atherinomorus stipes, 7 mm SL, BLZ 6051. C, Atherinidae, 14 mm SL, Belize, USNM 353871. D, Beloniformes
(Hemirhamphidae?), 14 mm SL, BLZ 4113. Photo A by Lee Weigt and David Smith; B by Lee Weigt and Carole Baldwin;
C by David Smith; D by Julie Mounts and Carole Baldwin.
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specimens of these taxa are not available. The only
other percomorph larvae examined that were as
extensively covered by erythrophores as postflexion
callionymid larvae are some apogonids and the goby
Priolepis (see ?Perciformes? and ?Gobiiformes? below).
Apogon aurolineatus is particularly similar to Cal-
lionymus bairdi in having a bright orange body with
yellow first dorsal and pelvic fins. The presence
of similar chromatophore patterns in these presum-
ably distantly related taxa is best interpreted as
convergence.
Pigment in larvae does not appear to provide any
evidence for a close relationship between gobiesoci-
forms and blenniiforms as proposed by Miya et al.
(2003) and Springer & Orrell (2004). The extensive
coverage of yellow/orange pigment in the gobiesoci-
forms examined is very different from the more
restrictive colour pattern observed in blenniiforms;
however, information on colour is needed for larvae of
more taxa, especially those blenniiforms hypothesized
by Stepien et al. (1997) to be more basal members of
the group than labrisomids and chaenopsids.
Gobiiformes
Gobiiformes examined include members of the Gobii-
dae, Eleotrididae, and Microdesmidae. Larvae of all
exhibit erythrophores or xanthophores, or both, and
some have iridophores, but there are no obvious diag-
nostic patterns for the order, major clades within the
order, or the highly diverse Gobiidae. Preliminary
comparative data suggest that colour patterns may be
most informative at intergeneric, generic, and species
levels.
Larvae of the microdesmid genera Microdesmus
and Cerdale have nearly identical patterns of orange/
yellow chromatophores (Fig. 20A?C). Hoese (1984)
united Microdesmus, Cerdale, and three other micro-
desmid genera with Ptereleotris and its allies in
an expanded family Microdesmidae, but Smith &
Thacker (2000) recognized the Microdesmidae and
Ptereleotrididae as separate families. Thacker?s
(2003, 2009) molecular phylogenetic analyses did
not recover a monophyletic group comprising micro-
desmids and ptereleotridids, but the molecular analy-
sis of Thacker & Roje (2011) did, with ptereleotridids
paraphyletic with respect to microdesmids. Superfi-
cially, the larval Ptereleotris (Fig. 20D) is quite differ-
ent from larval Microdesmus and Cerdale in that
it does not have an elongate body with an obvious
series of erythrophores internally along the vertebral
column and above the gut. The available colour
images of Ptereleotris, however, are of specimens that
are mostly opaque (vs. transparent), and although
internal erythrophores are present above the gut and
along the lateral midline posteriorly, it is not possible
to determine the extent of these series. Colour pat-
terns in the three genera are otherwise quite similar,
with all having a series of erythrophores along the
ventral midline (except directly beneath the swim
bladder), orange or yellow chromatophores in a series
along the dorsal midline posteriorly, and at least some
orange pigment externally on the central portion of
the caudal peduncle. This pattern was not observed in
other gobiiforms, and may provide corroborative evi-
dence for a monophyletic group comprising the two
families. The hypothesis that Coryphopterus gobies
are more closely related to Microdesmus and Cerdale
than microdesmids are to Ptereleotris (Thacker, 2003)
would not appear to be supported by colour patterns
in larvae. Coryphopterus gobies have a distinctive
pattern of erythrophores on the trunk and lack the
chromatophore pattern described above for microde-
smids (see last paragraph of Gobiiformes section,
below). Coryphopterus shares with microdesmids the
presence of erythrophores mixed with melanophores
along the anal-fin base, but this pigment is also
present in Ptereleotris and even the distantly related
eleotridids.
Figure 14. Percomorphacea (Acanthuriformes). A, Acanthurus bahianus, 26 mm Standard Length (SL), BLZ 8441.
B, Acanthurus chirurgus, 28 mm SL, BLZ 8442. Note: the internal red coloration behind the head is associated with the
circulatory system, not chromatophores. Photos by Julie Mounts and David Smith.
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Miller (1998) synonymized the eleotridid genera
Eleotris and Erotelis, and colour patterns in larvae
of the two genera are strikingly similar and distinc-
tive among gobiiforms (Fig. 20E, F). Thacker?s (2003)
molecular analysis did not recover the two as a mono-
phyletic group; rather, they are successive sister
groups to all other gobiiforms excluding the Odonto-
butidae. The more comprehensive molecular phy-
logeny of Thacker (2009) corroborates a common
ancestry for the two genera, with Erotelis embedded
within the Eleotris clade. In addition to having
similar patterns of erythrophores/xanthophores, Eleo-
Figure 15. Percomorphacea (Acanthuriformes). Acanthurus sp., Florida Straits. Photo by Cedric Guigand.
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Figure 16. Percomorphacea (Blenniiformes). A, Labrisomus cricota, 21.0 mm Standard Length (SL), BLZ 7006.
B, Labrisomus bucciferus, 19.0 mm SL, BLZ 7253. C, Acanthemblemaria greenfieldi, 13.0 mm SL, BLZ 6386.
D, Malacoctenus triangulatus, 18.0 mm SL, BLZ 8439. E, Paraclinus fasciatus, length not recorded, BLZ 6071. Note: in
all images, internal red coloration in the thoracic cavity is associated with the circulatory system, not chromatophores.
Photos A?D by Julie Mounts and David Smith; E by Lee Weigt and Carole Baldwin.
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tris and Erotelis are the only two gobiiform larvae
examined that have a swath of blue-reflecting irido-
phores at the base of the caudal fin.
Morphological and molecular data support a close
relationship between Ctenogobius and Gnatholepis
(Harrison, 1989; Thacker, 2003, 2009). As noted by
Baldwin & Smith (2003), larval Ctenogobius saepepal-
lens and Gnatholepis thompsoni are similar in having
a prominent, narrow bar of orange pigment on the
body just posterior to the anal-fin base that is lacking
in other gobiids (Fig. 21A, B). Baldwin & Smith
(2003) also noted that the orientation of this bar is
slightly different in the two species and that it may
not be homologous. Gobionellus oceanicus has a wide
bar of pale orange pigment on the caudal peduncle
that, because of the long anal fin in this species, is
located at the posterior base of the anal fin as the
narrow bar is in Ctenogobius and Gnatholepis
(Fig. 21C). The phylogenetic significance within the
Gobiidae of orange pigment at the posterior base of
the anal fin extending dorsally from the ventral
midline is unclear. Observations of colour patterns in
larvae of more genera of Gobionellus-group gobies
(Birdsong, Murdy & Pezold, 1988) is needed, includ-
ing Evorthodus, which Thacker (2003, 2009) proposed
as the sister group of Ctenogobius. Postlarval Cte-
nogobius boleosoma (Wyanski & Targett, 2000) shares
with Ct. saepepallens an erythrophore at the tip of the
lower jaw, a vertical reddish-orange streak between
the thoracic and abdominal regions, and the orange
bar posterior to the anal-fin base. Identification of
other Ctenogobius larvae is needed, but the combina-
tion of these pigment characters may be diagnostic of
the genus.
Nes and Psilotris are seven-spine gobies of the
?Gobiosoma group? (Birdsong et al., 1988) that share
with Varicus, Chriolepis, and Gobulus the absence
of head pores (B?hlke & Robins, 1968). R?ber, Van
Tassell & Zardoya (2003) recovered this group as
monophyletic, minus Varicus, which was not included
in their analysis. As noted by Smith & Baldwin (1999),
larvae of Nes and Psilotris are so similar that at first
these authors did not recognize them as distinct taxa.
The chromatophore pattern is nearly identical in the
two genera and distinctive among gobiids examined
(Fig. 21D, E). Identification of additional goby larvae,
including Gobulus and Chriolepis, is needed to deter-
mine whether the pattern is unique to Nes and Psilo-
tris or perhaps to a larger group. R?ber et al. (2003)
hypothesized that the relationships of Nes are as
follows: (Nes(Gobulus(Chriolepis + Psilotris))).
At the generic level, colour patterns in larvae are
useful in diagnosing Bathygobius and Coryphopterus.
Bathygobius larvae have orange/yellow chromato-
phores on the dorsal and ventral portions of the
trunk (in association with melanophores) that ex-
tend toward, and often meet at, the lateral midline
(Fig. 22A?C). Although the colour (yellow or orange)
Figure 17. Percomorphacea (Carangiformes). Trachinotus falcatus, 8.0 mm Standard Length, BLZ 5428. Photo by Lee
Weigt and David Smith.
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Figure 18. Percomorphacea (Carangiformes). A, Coryphaena hippurus, South Africa, photo by Allan Connell. B, C. hip-
purus. C, Selene vomer. B and C, Florida Straits, photos by Cedric Guigand, previously published in Fogarty & Botsford
(2007) and Cowen et al. (2007), respectively. Note: the internal pink coloration in the thoracic region of B is associated
with the circulatory system, not chromatophores.
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and density of pigment differ among species, the
net effect is distinctive (Baldwin & Smith, 2003;
Tornabene et al., 2010). All known Coryphopterus
larvae have diagonal bars of orange pigment on the
trunk, the height of the bars and how far the series
extends anteriorly varying among species (Baldwin &
Smith, 2003; Fig. 22E?H). Colour patterns in larvae
of genera hypothesized to be closest to Bathygobius
(Glossogobius, Istigobius, and Callogobius; Tornabene
& Pezold, 2011, and references therein; Thacker,
2009) and Coryphopterus (Lophogobius; Thacker,
2003, 2009) are unknown. The unique generic pat-
terns described above could diagnose larger groups.
Monophyly of Thacker?s (2003) clade IIIA, which
includes Bathygobius and Priolepis, would not appear
to be supported by larval colour patterns. Larval
Priolepis (Fig. 22D) is distinctively orange.
Labriformes
Labriformes comprise the Cichlidae, Embiotocidae,
Labridae, Odacidae, Pomacentridae, and Scaridae.
The monophyly of the group is questionable (Wiley &
Johnson, 2009, and references therein), and molecu-
lar data suggest that scarids are embedded within
the Labridae (Westneat & Alfaro, 2005; Choat et al.,
2012). Marine labriforms for which colour patterns
were assessed for larvae are pomacentrids, labrids,
and scarids. All exhibit nonmelanistic chromato-
phores, usually erythrophores, but the patterns are
not diagnostic of the order. Based on existing com-
parative material, colour patterns appear useful in
diagnosing some families and genera. All scarid
larvae examined are united in having a linear series
of erythrophores along the ventral midline of the
trunk from beneath the operculum to the anus, a
linear series of erythrophores along the anal-fin base,
a roughly linear series of erythrophores above the
anal-fin base that curves dorsally on the caudal
peduncle where it is continuous with erythrophores
on the lateral midline, and erythrophores on the
caudal fin (Fig. 23). This pattern is present in Cryp-
totomus, Scarus, and Sparisoma, which differ from
one another in (1) the extent of orange coloration
on the caudal fin (with streaks of bright orange chro-
matophores on the ventral lobe in Cryptotomus,
Fig. 23A, vs. orange pigment more scattered or paler
in the other genera); (2) organization of erythrophores
and melanophores above the anal-fin base (somewhat
Figure 19. Percomorphacea (Gobiesociformes). A, B, Synchiropus splendidus, reared specimens. C, Callionymus marleyi,
South Africa. D. Callionymus bairdi, 7.0 mm Standard Length (SL), BLZ 4315. E, Acyrtops beryllinus, 4.0 mm SL, BLZ
6231. Photos A, B by Matthew Wittenrich; C by Allan Connell; D by Lee Weigt and David Smith; E by Julie Mounts and
David Smith.
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Figure 20. Percomorphacea (Gobiiformes). A, Cerdale floridana, 21.0 mm Standard Length (SL), Belize. B, Microdesmus
bahianus, 17.5 mm SL, Belize. C, Microdesmus carri, 22 mm SL, BLZ 8322. D. Ptereleotris helenae, 12.0 mm SL, BLZ
7323. E, Eleotris pisonis, 13.0 mm SL, BLZ 7101. F, Erotelis smaragdus, 13 mm SL, Belize. Note: the internal red
coloration in the thoracic region of P. helenae is associated with the circulatory system, not chromatophores. Photos A, B,
F by Julie Mounts and Carole Baldwin; C by Lee Weigt and Carole Baldwin; D, E by Julie Mounts and David Smith.
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Figure 21. Percomorphacea (Gobiiformes). A, Ctenogobius saepepallens, 10.0 mm Standard Length (SL), BLZ 7387.
B, Gnatholepis thompsoni, 11.0 mm SL, Belize. C, Gobionellus oceanicus, 12.0 mm SL, BLZ 5473. D, Nes longus, 12.0 mm
SL, BLZ 7183. E, Psilotris sp., 12.0 mm SL, BLZ 7187. Photo A by Lee Weigt and Carole Baldwin; B by Julie Mounts and
Carole Baldwin; C by Lee Weigt and David Smith; D, E by Julie Mounts and David Smith.
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haphazard in Scarus ? Fig. 23B, linear in Cryp-
totomus and Sparisoma ? Fig. 23A, C, D); and (3) the
configuration of erythrophores along the ventral
midline on the anterior portion of the trunk (forming
an almost continuous line of orange pigment in
Cryptotomus and Sparisoma, more widely spaced
in Scarus). A generic-level phylogeny of scarids pre-
sented by Kazanciog?lu et al. (2009) suggests that
Cryptotomus and Sparisoma are sister groups, and
(2) and (3) above, if apomorphic, would lend extra
support to this hypothesis. The anterior extension of
the midlateral series of erythrophores almost to the
head in Sparisoma atomarium (Fig. 23D) and Spari-
soma radians (Fig. 23E) is unique among labriforms
and may indicate that these species are more closely
related to one another than either is to Sparisoma
chrysopterum (Fig. 21C).
Larval pomacentrids examined have silvery irido-
phores on the gut and at least some erythrophores or
xanthophores on the trunk and fins (Fig. 24). Colour
patterns of larvae of the four genera examined are
distinctive. Stegastes larvae have a swathe of orange
pigment on the trunk from just behind the eye to
the anterior-most part of the caudal peduncle,
where the swath ends abruptly in a near vertical
line (Fig. 24A?C). Differences among Stegastes
species include the presence or absence of erythro-
phores on the pectoral fin and along the spinous
dorsal-, pelvic-, and anal-fin bases. Colour informa-
tion was available for only one species of Chromis
(Fig. 24D) and one of Abudefduf (Fig. 24E), but pat-
terns in these genera are distinct from one another
and from Stegastes. Chromis lacks colour on the fins
and has erythrophores restricted to the posterior
portion of the trunk and caudal peduncle, whereas
Abudefduf is distinctive in having xanthophores
on most of the trunk (mixed with melanophores)
and conspicuously yellow first dorsal and pelvic fins.
Like Stegastes, pigment on the trunk in Abudefduf
ends abruptly on the anterior portion of the caudal
Figure 22. Percomorphacea (Gobiiformes). A, Bathygobius curacao, 5.5 mm Standard Length (SL), BLZ 7305. B,
Bathygobius lacertus, 6.0 mm SL, BLZ 7370. C, Bathygobius soporator, 6.0 mm SL, BLZ 6072. D, Priolepis hipoliti,
10.0 mm SL, Belize. E, Coryphopterus kuna, 7.5 mm SL, BLZ 5134. F, Coryphopterus tortugae, 7.0 mm SL, BLZ 5227.
G, Coryphopterus personatus, 8.0 mm SL, BLZ 10007. H, Coryphopterus venezuelae, 8.5 mm SL, BLZ 5392. Photos A, B
by Julie Mounts and David Smith; C by Lee Weigt and Carole Baldwin; D-F, H by Julie Mounts and Carole Baldwin;
G by Donald Griswold and Carole Baldwin.
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Figure 23. Percomorphacea (Labriformes). A, Cryptotomus roseus, 9.0 mm Standard Length (SL), BLZ 10005. B, Scarus
iseri, 7 mm SL, Belize. C, Sparisoma atomarium, 10.0 mm SL, BLZ 7312. D, Sparisoma chrysopterum, 9.0 mm SL, BLZ
6383. E, Sparisoma radians, 11.0 mm SL, BLZ 7289. Photo A by Donald Griswold and Carole Baldwin; B by Julie Mounts
and Carole Baldwin; C-E by Julie Mounts and David Smith.
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peduncle. Reared larvae of Amblyglyphidodon terna-
tensis (Fig. 24F) are similar to Abudefduf in having
prominent yellow pigment on the dorsal and pelvic
fins. They are distinctive in having yellow pigment
covering the dorsal portion of the head, including the
dorsal portion of the orbit, and also the anterior
portion of the anal fin. The molecular phylogeny of
Cooper, Smith & Westneat (2009) suggests that Ste-
gastes, Chromis, Abudefduf, and Amblyglyphidodon
are members of four distinct evolutionary assem-
blages (Stegastinae, Chrominae, Abudefdufinae,
Pomacentrinae, respectively), a hypothesis that is
not contradicted by colour patterns in larvae. Larval
Abudefduf and Amblyglyphidodon are the most
similar of the four in terms of coloration, and Abu-
defdufinae and Pomacentrinae are sister groups
according to Cooper et al. (2009). Acquisition of
colour information for larvae of additional pomacen-
trids is needed to determine whether or not the
colour patterns identified herein characterize the
subfamilies (or some subset of them) and whether
aspects of the colour pattern in Abudefduf and
Amblyglyphidodon represent a synapomorphy of
Abudefdufinae and Pomacentrinae.
Figure 24. Percomorphacea (Labriformes). A, Stegastes partitus, 11.5 mm Standard Length (SL), BLZ 8454. B, Stegastes
variabilis, 10.0 mm SL, BLZ 4523. C, Stegastes planifrons, 10.0 mm SL, BLZ 6008. D, Chromis cyanea, 14.0 mm SL, BLZ
8451. E, Abudefduf saxatilis, 10.0 mm SL, BLZ 10214. F, Amblyglyphidodon ternatensis, reared aquarium specimen.
Photos A, D by Julie Mounts and David Smith; B by Lee Weigt and David Smith; C by Lee Weigt and Carole Baldwin;
E by Donald Griswold and Carole Baldwin; F by Matthew Wittenrich.
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All genera of labrid larvae examined (Anampses,
Halichoeres, Lachnolaimus, Thalassoma, Xyrichtys)
except Doratonotus have prominent orange pigment
on the tip of the upper jaw and usually also on the tip
of the lower jaw (Fig. 25). Otherwise, colour patterns
are distinctive among genera or groups of genera.
Anampses (A. Connell, pers. comm.), Halichoeres
(Fig. 25A?D), and Thalassoma (Fig. 25E) larvae
have at least a small cap and sometimes a broader
covering of erythrophores on the gut and orange spots
or blotches on the posterior portion of the head.
Anampses and Halichoeres also have an orange blotch
on the ventral portion of the trunk just posterior to
the anal-fin base. Species-specific features within
Halichoeres include the presence of several orange
blotches on the dorsal and mid-lateral portions of the
trunk (Halichoeres maculipinna, Fig. 25C) and pres-
ence of a vertical line of erythrophores on the caudal-
fin base (Halichoeres poeyi, Fig. 25D). Colour patterns
in Halichoeres bivittatus (Fig. 25A) and Halichoeres
garnoti (Fig. 25B) are extremely similar, but these
species can be separated by the pattern of melano-
phores on the dorsal and anal fins. Rocha, Pinheiro &
Gasparini (2010) presented a preliminary molecular
phylogeny of New World Halichoeres, a relevant
aspect of which is the placement of Ha. maculipinna
in a clade distinct from one comprising Ha. poeyi,
Ha. garnoti, Ha. bivittatus, and several other Hali-
choeres species (Rocha et al., 2010: fig. 4). Larvae
of H. maculipinna differ from those of Ha. poeyi,
Ha. garnoti, and Ha. bivittatus in having much more
orange coloration on the body and more prominent
black blotches on the dorsal and anal fins posteriorly
(Fig. 25). The larvae of Ha. poeyi, Ha. garnoti, and
Ha. bivittatus are very similar. The molecular phyl-
ogeny of labrids by Westneat & Alfaro (2005) did
not include Ha. maculipinna, but it suggests that
Halichoeres is paraphyletic without the inclusion of
numerous other genera, including Thalassoma and
Anampses. The presence of erythrophores in the three
genera on the upper jaw and gut, combined with the
presence of distinct dark markings on the dorsal and
anal fins, constitute a unique pattern within labri-
forms that may support a close relationship among
these genera.
Xyrichtys larvae are elongate and consistently have
erythrophores on both jaws and behind the eye. The
rest of the body is pale except for a blotch of colour
on the caudal peduncle (Fig. 26A?C). In Xyrichtys
novacula (Fig. 26A) this blotch is always yellow,
whereas in Xyrichtys splendens (Fig. 26B) and Xyrich-
tys martinicensis (Fig. 26C) it is orange. Differences
in size and shape of the orange blotch distinguish
these two species. There is little if any intraspecific
variation in chromatophore pattern among the indi-
viduals of each Xyrichtys species examined.
Doratonotus and Lachnolaimus are monotypic
genera, and their larvae are clearly distinct from one
another and other labrids (Fig. 26D, E). Doratonotus
larvae have internal erythrophores along the verte-
bral column and along the myosepta of the posterior
third of the trunk and, as noted above, lack erythro-
phores on the jaws. Lachnolaimus larvae are very
different from those of other labrids in having almost
the entire body covered with orange/yellow chromato-
phores ? mostly xanthophores anteriorly and eryth-
rophores posteriorly.
Lophiiformes
Lophiiformes comprise more than a dozen perco-
morph families, but larvae of only one species from
Belize, Antennarius pauciradiatus (Antennariidae),
have been identified (Fig. 27A). Images of colour
patterns in several unidentified lophiiform larvae
from the Florida Straits were used for comparative
purposes (Fig. 27B?D). In Ant. pauciradiatus the dis-
tended skin forming the characteristic lophiiform
?balloon? around the head and body is lightly covered
with erythrophores in the lower jaw and gular
regions. The three unidentified lophiiform larvae
have different colour patterns from Ant. pauciradia-
tus and one another, two of them exhibiting xantho-
phores (Fig. 27C, D) and one of them seemingly
lacking xanthophores/erythrophores and exhibiting
only blue iridophores (Fig. 27B). Lophiiforms, which
form part of Rosen & Patterson?s (1969) and
Patterson & Rosen?s (1989) Paracathopterygii, were
hypothesized to be percomorph fishes in the molecu-
lar phylogeny of Miya et al. (2003). The ?paracan-
thopts? examined for colour patterns in larvae are
gadids (Fig. 6D), lophiiforms, and ophidiiforms (see
below). With the limited amount of material avail-
able, the only observation relevant here is that unlike
the gadid Bregmaceros, which lacks xanthophores,
erythrophores, and iridophores, all lophiiforms (and
ophidiiforms) examined have one or more of those
types of chromatophores. Like numerous other acan-
thomorph orders, lophiiforms are currently consid-
ered incertae sedis within the Percomorphacea
(Fig. 1), but the hypothesis of Holcroft & Wiley (2008)
that lophiiforms are closely related to tetraodon-
tiforms is discussed below (see Tetraodontiformes).
Ophidiiformes
Ophidiiformes were also hypothesized to be perco-
morphs rather than paracanthopterygians by Miya
et al. (2003, 2005). Wiley & Johnson (2009) noted that
no convincing evidence for the monophyly of the order
(ophidioids + bythitoids) exists. Colour information in
larvae is known for two ophidioids from Belize, Cara-
pus bermudensis (Carapidae) and Parophidion schmi-
dti (Ophidiidae), both of which have conspicuous
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Figure 25. Percomorphacea (Labriformes). A, Halichoeres bivittatus, 12.5 mm Standard Length (SL), BLZ 6426.
B, Halichoeres garnoti, 15.0 mm SL, BLZ 7085. C, Halichoeres maculipinna, 17.0 mm SL, BLZ 7124. D, Halichoeres poeyi,
14.0 mm SL, BLZ 6102. E, Thalassoma bifasciatum, 11.0 mm SL, BLZ 8334. Photos A-C by Julie Mounts and David
Smith; D, E by Lee Weigt and Carole Baldwin.
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Figure 26. Percomorphacea (Labriformes). A, Xyrichtys novacula, 12.0 mm Standard Length (SL), BLZ 5394. B, Xyrich-
tys splendens, 17.0 mm SL, BLZ 7019. C, Xyrichtys martinicensis, 15.0 mm SL, Belize. D, Doratonotus megalepis, 7.0 mm
SL, Belize. E, Lachnolaimus maximus, 7.0 mm SL, BLZ 7373. Photos A, C, D by Julie Mounts and Carole Baldwin;
B, E by Julie Mounts and David Smith.
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Figure 27. Percomorphacea (Lophiiformes). A, Antennarius pauciradiatus, 6.0 mm Standard Length (SL), BLZ 6043.
B-D, unknown lophiiforms, Florida Straits. Note: the internal pink coloration in the images appears to be associated with
the circulatory system, not chromatophore patterns. Photo A by Lee Weigt and Carole Baldwin; B-D by Cedric Guigand,
B and C previously published in Cowen et al. (2007) and Fogarty & Botsford (2007), respectively.
526 C. C. BALDWIN
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orange/yellow chromatophore patterns (Fig. 28). In
situ images of two Hawaiian ophidiids that appear to
be larval Brotulataenia and Lampogrammus (Fig. 29)
show strikingly beautiful larvae with numerous xan-
thophores on the body and fins, as well as erythro-
phores on the fins in the former. Erythrophores in
Parophidion are confined to the head and trunk and do
not extend onto the median fins. No conclusions about
the potential phylogenetic significance of colour pat-
terns in larval ophidioids can be drawn at this time,
and colour patterns, if any, in young bythitoids are
unknown.
Perciformes
Perciformes remain a conglomerate of families and
incertae sedis genera that are not united by synapo-
morphies (Wiley & Johnson, 2009), and larvae of the
group exhibit a diverse array of chromatophore pat-
terns. At the interfamilial level, the Chaetodontidae
and Pomacanthidae have larvae that are covered
with iridophores ? silver/bronze in Holacanthus,
Chaetodon, and Pomacanthus arcuatus, silver/blue in
Pomacanthus paru (Fig. 30). Although not as con-
spicuous in Holacanthus (Fig. 30A) and Pomac. paru
(Fig. 30D) as in Chaetodon and Pomac. arcuatus,
Figure 28. Percomorphacea (Ophidiiformes). A (and inset), Carapus bermudensis, 152 mm Standard Length (SL), BLZ
8462. B, Parophidion schmidti, 36.5 mm SL, BLZ 8459. Photos by Julie Mounts and David Smith.
COLOUR IN MARINE TELEOST LARVAE 527
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Figure 29. Percomorphacea (Ophidiiformes). Top, Brotulataenia sp. and bottom, Lampogrammus sp., Hawaii. Photos by
Joshua Lambus.
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Figure 30. Percomorphacea (Perciformes). A, Holacanthus ciliarus, 16.0 mm Standard Length (SL), BLZ 8477. B,
Chaetodon capistratus, 12.0 mm SL, BLZ 8436. C, Pomacanthus arcuatus, 10.0 mm SL, BLZ 10101. D, Pomacanthus paru,
10.0 mm SL, BLZ10213. Photos A, B by Julie Mounts and David Smith; C, D by Donald Griswold and Carole Baldwin.
COLOUR IN MARINE TELEOST LARVAE 529
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larvae of all specimens of all three genera have xan-
thophores, erythrophores, or both, minimally on the
jaws, snout, base of dorsal fin, and caudal peduncle.
It does not appear that chaetodontids and pomacan-
thids resemble smegmamorphs in transitioning
from an early stage featuring predominantly
xanthophores/erythrophores to a later stage featur-
ing primarily iridophores, as preflexion Chaetodon
marleyi and Pomacanthus rhomboides have few if
any xanthophores or erythrophores (Fig. 31). The
preflexion Chaetodon and Pomacanthus are quite
similar in having a broad band of iridescent silvery
blue iridophores around the trunk. Chaetodontids
and pomacanthids have been considered to be so
closely related in the past that until Burgess? (1974)
publication, pomacanthids were classified as a
subfamily of the Chaetodontidae. The families were
still considered closely related by Tyler et al. (1989),
who delineated a monophyletic group comprising
chaetodontids, pomacanthids, and Drepane (Drepa-
neidae) based on configuration of the ethmoid bone.
Colour in larval Drepane is unknown, but the pres-
ence in larvae of that genus of a colour pattern
comprised largely of iridophores with xanthophores/
erythrophores positioned as described above for
chaetodontids and pomacanthids could provide cor-
roborative evidence for the monophyly of the group.
Recent molecular studies have challenged the
hypothesis of a close relationship between chaeto-
dontids and pomacanthids. Using mitochondrial
genes, Bellwood, van Herwerden & Konow (2004)
hypothesized that chaetodontids are more closely
related to scatophagids than to pomacentrids, and
Holcroft & Wiley (2008) hypothesized that chaeto-
dontids and scatophagids are part of a large group
also comprising acanthuroids, lophiiforms, and
tetraodontiforms but not pomacanthids. Colour infor-
mation for larval scatophagids is not available, but
colour in larvae would not appear to support a closer
relationship between chaetodontids and acanthurids
(Figs 14, 15) than between chaetodontids and poma-
canthids (Fig. 30). Larval chaetodontids bear little
resemblance to larval lophiiforms (Fig. 27), but
larval tetraodontid tetraodontiforms examined are
similar to chaetodontids and pomacanthids in having
the body covered with bronze/gold iridophores and
the abdominal region silvery (see ?Tetraodontiformes?
below).
At the familial level, larval Apogonidae have nearly
the entire body covered with erythrophores. With
the exception of Priolepis gobies and Callionymus
dragonettes, no other larvae examined are covered
so completely with erythrophores, and this feature
may be synapomorphic for the family or some subset
of it. Colour patterns in western Atlantic larval
Apogon, Astrapogon, and Phaeoptyx (Fig. 32) have
Figure 31. Percomorphacea (Perciformes). Top, preflexion larva of Chaetodon marleyi, 3.8 mm Notochord Length (NL).
Bottom, preflexion larva of Pomacanthus rhomboides, 3.9 mm NL. Both images are of specimens from off South Africa.
Photos by Allan Connell.
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Figure 32. Percomorphacea (Perciformes). A, Apogon aurolineatus, 9.0 mm Standard Length (SL), BLZ 10001. B,
Astrapogon puncticulatus, 13.0 mm SL, BLZ 7125. C, Phaeoptyx xenus, 10.0 mm SL, BLZ 10220. Photos A, C by Donald
Griswold and Carole Baldwin; B by Julie Mounts and David Smith.
COLOUR IN MARINE TELEOST LARVAE 531
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been described (Baldwin et al., 2009a; 2011). Patterns
of melanophores, in conjunction with orange/yellow
chromatophores, diagnose the three genera. Within
Apogon, colour patterns delineate several species
groups that may be meaningful in the generic classi-
fication of the group (Baldwin et al., 2011).
Larval Gerreidae examined lack erythrophores and
xanthophores on most of the body, but some pale
yellow pigment is usually present over the gut and
swimbladder and sometimes on the dorsal portion of
the head (Fig. 33). Determining whether this pattern,
combined with the distinctive arrangement of mela-
nophores at the bases of the median fins, character-
izes Eucinostomus (larvae of four species identified) or
the entire family requires additional material.
Late-stage larvae of the Lutjanidae are easily rec-
ognized by the cap of silver iridophores over the gut
and opercular region, a mostly clear trunk, and xan-
thophores associated with the dorsal fin (or its base)
and sometimes caudal peduncle (Fig. 34). The pattern
of xanthophores appears species specific within
Lutjanus. Presence of the same general pattern of
iridophores and xanthophores in Ocyurus chrysurus
(Fig. 34F) may lend support to the hypothesis based
on morphological (including larval) and molecular
data that Ocyurus is a synonym of Lutjanus (Domeier
& Clarke, 1992; Chow, Clarke & Walsh, 1993; Clark,
Domeier & Laroche, 1997), but information on colour
patterns in larvae of other lutjanid genera is needed
to determine at what taxonomic level the colour
pattern in Lutjanus is significant. In general appear-
ance, including the presence of melanophores dorsally
on the head, a silvery gut, pigment associated with
the pelvic-fin spine, and usually orange or yellow
chromatophores on the caudal peduncle, larval Lut-
janus is very similar to larvae of the grouper genus
Mycteroperca (see ?Scorpaeniformes? below).
At the generic level, larval Haemulon (Haemulidae)
has a stripe of xanthophores/erythrophores mixed
with dark melanophores on the posterior half of the
trunk (Fig. 35A, B). Among perciforms examined,
larval Calamus is most similar in having a swathe of
xanthophores/erythrophores on the posterior portion
of the trunk mixed with melanophores (Fig. 35C, D).
There are no chromatophores on the posterior portion
of the trunk in the haemulid Anisotremus (Fig. 35E),
but several species of the haemulid genus Pomadasys
are very similar to Haemulon (e.g. Pomadasys com-
mersonnii, Connell, 2007; Appendix). There is no con-
sensus based on morphological or molecular data for
inter-relationships of haemulids and sparids among
perciforms.
A larva tentatively identified as an opistognathid
(Fig. 36A) and the sciaenid Odontoscion (Fig. 36B)
lack orange/yellow chromatophores on the trunk and
have xanthophores confined to the head and gut
region. A larval Mullidae, Upeneus parvus, does not
resemble any other perciform larvae examined in that
it is covered with melanophores and blue iridophores
(Fig. 36C). Based on mitochondrial and nuclear
DNA data, Smith & Craig (2007) suggested a close
relationship between mullids and a nonperciform
family, Dactylopteridae (Dactylopteriformes are incer-
tae sedis in percomorphs in the classification of Wiley
& Johnson, 2009). Although colour information is
lacking, dactylopterid larvae have huge head spines
that are lacking in Upeneus. Possibly the dense cov-
ering of iridophores in Upeneus larvae will be of value
in identifying its closest relatives in the future. Pre-
flexion larvae of Oplegnathus (Oplegnathidae), Pem-
pheris (Pempheridae), Neoscorpis (Scorpididae), and
several other perciform families from off South Africa
have a considerable amount of yellow pigment on the
trunk (Connell, 2007; Appendix), but larger speci-
mens are needed for comparisons with other postflex-
ion perciform larvae.
Pleuronectiformes
Pleuronectiformes are united as a monophyletic group
in part based on ontogeny (transformation from
a bilaterally symmetrical larva to an asymmetrical
adult), and one or more elongate anterior dorsal-
fin elements characterize larvae of several families
(Hensley & Ahlstrom, 1984). Colour patterns in
larvae should prove useful phylogenetically at some
levels. The presence of bright orange erythrophores
on the median fins and fin bases, pelvic fin, head, and
lateral midline (where they are small to minute) in
the bothids Bothus maculiferus and Bothus ocellatus,
combined with the absence of melanophores, may be
synapomorphic for the genus (Fig. 37A, B). Superfi-
cially different from Bothus larvae because of the
presence of numerous melanophores, larvae of the
paralichthyid genus Syacium (e.g. Fig. 37C-F) are
quite similar to Bothus in nonmelanistic colour
pattern. Larvae of Syacium papillosum and three
unidentified species have orange/yellow pigment on
the median fins and fin bases, pelvic fin, head, and
lateral midline, but also in a distinct patch on the
dorsal portion of the gut. A single, small Citharichthys
larva (Paralichthyidae) appears to have a similar
pattern, but erythrophores are tiny (or contracted),
and it is difficult to discern their precise distribution
(Fig. 37G). Bothids and at least some paralichthyids
(including Citharichthys and Syacium) consistently
appear as sister groups or components of a slightly
larger monophyletic group based on morphology and
molecules (Cooper & Chapleau, 1998; Berendzen &
Dimmick, 2002; Pardo et al., 2005; Azevedo et al.,
2008), and the general colour pattern described above
may help define a clade that includes bothids and
those paralichthyids. However, the more distantly
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related Cynoglossidae (one Symphurus examined) has
a similar colour pattern, with orange pigment present
on the median fins and fin bases, head, and lateral
midline (Fig. 37I). The single Achiridae examined
(Trinectes) is heavily covered with melanophores,
largely obscuring the nonmelanistic colour pattern,
but pale orange pigment is visible on the median and
pelvic fins (Fig. 37H). An in situ image of an uniden-
tified pleuronectiform larva from off Hawaii, possibly
a bothid, has the same general colour pattern
Figure 33. Percomorphacea (Perciformes). A, Eucinostomus gula, 9 mm Standard Length (SL), BLZ 10107. B, Eucinos-
tomus jonesi, 9.2 mm SL, BLZ 7257. C, Eucinostomus harengulus, 12 mm SL, BLZ 7345. D, Eucinostomus melanopterus,
15 mm SL, BLZ 10228. Photos A, D by Donald Griswold and Carole Baldwin; B, C by Julie Mounts and David Smith.
COLOUR IN MARINE TELEOST LARVAE 533
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Figure 34. Percomorphacea (Perciformes). A, Lutjanus analis, 16.5 mm Standard Length (SL), BLZ 8466. B, Lutjanus
griseus, 13.5 mm SL, BLZ 8427. C, Lutjanus synagris, 18.0 mm SL, BLZ 7150. D, Lutjanus vivanus, 26 mm SL, BLZ 8399.
E, Lutjanus mahogani, 23.0 mm SL, BLZ 5453. F, Ocyurus chrysurus, 17.5 mm SL, BLZ 7052. Images in column on right
are enlarged views of the dorsal fin of each image in left column. Photos A-C, F by Julie Mounts and David Smith;
D by Lee Weigt and Carole Baldwin; E by Lee Weigt and David Smith.
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observed in most pleuronectiforms ? i.e. yellow/orange
pigment on the median fins (dorsal and anal fins
almost completely covered with pigment), dorsal- and
anal-fin bases, pelvic fin, head, and lateral midline
(Fig. 38).
Species differences in the material examined are
evident in the configuration of erythrophores, Bothus
serving as an excellent example (Fig. 37A, B). In the
absence of melanophores, preserved larvae of Bo. ocel-
latus and Bo. maculiferus lack diagnostic pigment
patterns, but fresh specimens are easily distinguished
by the pattern of erythrophores (Bo. ocellatus with
more orange markings along the dorsal- and anal-fin
bases than Bo. maculiferus and with erythrophores
on the base of the caudal fin, in dashes along the
dorsal and ventral body margins posteriorly, and in
lines between those dashes and the orange markings
along the bases of the dorsal and anal fins vs. none of
these markings in Bo. maculiferus). As noted under
?Carangiformes?, a relationship between pleuronecti-
forms and carangiforms as proposed by Little et al.
(2010) would not appear to be supported by colour
patterns in larvae, nor would a proposed relationship
(Smith & Craig, 2007) with Xiphias gladius (see
?Scombriformes? below).
Scombriformes
Scombriformes are represented in the material exam-
ined by larvae of the Gempylidae, Istiophoridae,
Scombridae, Sphyraenidae, and Xiphiidae. Early
postflexion larvae of Xiphias (Fig. 39A) and Sphy-
raena (Fig. 39B) have extremely similar colour pat-
terns ? yellow extending from the snout to the caudal
peduncle with numerous melanophores mixed in. A
larval scombrid examined, by contrast, is mostly pale,
with only a small amount of yellow pigment over
the gut and pale orange pigment along the anal-fin
base and lateral midline (Fig. 39C, see also http://
vertebrates.si.edu/fishes/larval/perci.html for an
image of a preflexion Thunnus that is similar). A
postflexion Auxis from off South Africa has some
reddish pigment over the gut but is otherwise simi-
larly pale (Connell, 2007; Appendix). A larval gem-
pylid is similar to the scombrids examined in having
a mostly pale head and trunk, but the spinous dorsal
fin is densely pigmented with melanophores and
bronze iridophores (Fig. 39D). A larval istiophorid
examined does not resemble any of the other scom-
briform larvae in that nearly the entire body is
covered with blue-reflecting iridophores (Fig. 39E).
The relationship of billfishes to scombrids has been
the subject of much controversy (e.g. Collette et al.,
1984; Johnson, 1986; Orrell, Collette & Johnson,
2006; Little et al., 2010), and larval colour patterns do
not appear to shed much light on the matter. The
most recent molecular hypotheses (Orrell et al., 2006;
Little et al., 2010) suggest that billfishes are not the
closest relatives of scombrids. Orrell et al. (2006:
fig. 3) proposed a sister-group relationship between
Sphyraena and Xiphias + Istiophoridae, and, if apo-
Figure 35. Percomorphacea (Perciformes). A, Haemulon sciurus, 8.0 mm Standard Length (SL), BLZ 7369. B, Haemulon
plumieri, 7.0 mm SL, BLZ 6204. C, Calamus sp., 5.0 mm SL, BLZ 6022. D, Calamus sp., 9.0 mm SL, BLZ 7256. E,
Anisotremus virginicus, 5.5 mm SL, BLZ 7346. Photos A, D, E by Julie Mounts and David Smith; B, C by Lee Weigt and
Carole Baldwin.
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morphic, the similar colour patterns in larval Sphy-
raena and Xiphias could provide further support for
this hypothesis.
Scorpaeniformes
Scorpaeniformes traditionally have comprised the
scorpaenids and their mail-cheeked relatives (e.g.
Nelson, 2006), but Wiley & Johnson (2009) followed
Imamura & Yabe (2002) in placing scorpaenoids,
platycephaloids, and the Serranidae in the Scorpaeni-
formes. Some molecular data conflict with this
hypothesis and the proposed monophyly of the com-
ponent suborders ? Scorpaenoidei and Serranoidei
(Smith & Wheeler, 2004; Smith & Craig, 2007). All
scorpaeniforms for which colour in larvae has been
examined ? the scorpaenids Scorpaena, Scorpaenodes,
Dendrochirus; the peristiid, Peristedion; and the ser-
ranids Diplectrum, Serranus, Hypoplectrus, Gonio-
plectrus, Mycteroperca, Bathyanthias, Liopropoma,
Pseudogramma, Rypticus, and Pseudanthias ? have
orange/yellow chromatophores on the body. The three
scorpaenid genera have erythrophores or xantho-
phores on an enlarged pectoral fin, the placement and
extent of the colour aiding generic recognition: in
Figure 36. Percomorphacea (Perciformes). A, Opistognathidae?, 9.0 mm Standard Length (SL), BLZ 6394. B, Odonto-
scion dentex, 6.0 mm SL, BLZ 10153. C, Upeneus parvus, 10.5 mm SL, BLZ 4505. Note: internal red coloration in the
thoracic region of A is associated with the circulatory system, not chromatophores. Photo A by Julie Mounts and David
Smith; B by Donald Griswold and Carole Baldwin; C by Lee Weigt and David Smith.
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Figure 37. Percomorphacea (Pleuronectiformes). A, Bothus maculiferus, 13.0 mm Standard Length (SL), BLZ 4219.
B, Bothus ocellatus, 19.0 mm SL, Belize. C, Syacium sp., 17.0 mm SL, Belize. D, Syacium sp., 15.5 mm SL, BLZ 7078. E,
Syacium sp., 12.0 mm SL, BLZ 6010. F, Syacium sp., 13.0 mm SL, BLZ 8463. G, Citharichthys sp., 9.5 mm SL, BLZ 6006.
H, Trinectes sp., 5.0 mm SL, BLZ 10161. I, Symphurus sp., 11 mm SL, BLZ 7779. Photo A by Lee Weigt and David Smith;
B, C by Julie Mounts and Carole Baldwin; D, F by Julie Mounts and David Smith; E, G, I by Lee Weigt and Carole
Baldwin; H by Donald Griswold and Carole Baldwin.
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Scorpaenodes nearly the entire fin is covered with
xanthophores and melanophores with sometimes
the very proximal area clear (Fig. 40A); in Scorpaena
yellow or orange chromatophores ? and sometimes
dense melanophores ? cover the proximal portion of
the fin, and the distal portion is clear (Fig. 40B?D);
and in Dendrochirus xanthophores form a band
across the central portion of the fin (Fig. 40E). The
pectoral fin superficially looks larger in Scorpaenodes
than in the other genera because the distal portion is
heavily pigmented. Within Scorpaena, the colour of
the chromatophores on the pectoral fin and whether
or not they are mixed with melanophores allow
species recognition (Fig. 40B?D). In Peristedion, the
pectoral fin is also enlarged, the upper rays extremely
so, and there appear to be erythrophores mixed with
melanophores on the upper elongate rays (Fig. 41).
Diplectrum, Serranus, and Hypoplectrus (serranine
serranids) likewise have erythrophores or xantho-
phores, always mixed with melanophores, on the pec-
toral fin and are further similar to Scorpaena larvae
in having yellow/orange chromatophores on the head
and on the anterior and midlateral portions of the
trunk (Fig. 42). The pectoral fin is not enlarged in the
serranines, but it is enlarged and highly pigmented
in some epinepheline serranids, including the genus
Rypticus (Fig. 43A?C). Distinct patterns of orange/
yellow chromatophores and melanophores on the
enlarged fin characterize genetic lineages/species of
Rypticus. Larvae of another epinepheline serranid
examined, Pseudogramma, have an enlarged pectoral
fin, but have only pale orange or yellow coloration vs.
bright orange/yellow as in Rypticus (Fig. 43D). The
presence of an elongate, pigmented pectoral fin is
unusual among larval percomorphs examined and
may be phylogenetically significant in uniting scor-
paenids and serranids at the ordinal level.
Larval Pseudogramma also share with at least
some Rypticus a similar pattern of prominent orange/
yellow chromatophores on the posterior portion of the
trunk: one blotch is present centrally on the caudal
peduncle, and two blotches precede it, one on the
dorsal portion of the trunk and one on the ventral
portion (Fig. 43B?D). Pseudogramma and one species
Figure 38. Percomorphacea (Pleuronectiformes). Unknown pleuronectiform, Hawaii. Photo by Joshua Lambus.
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Figure 39. Percomorphacea (Scombriformes). A, Xiphias gladius. B, Sphyraena barracuda, 8.0 mm Standard Length,
Belize. C, Scombridae. D, Gempylidae. E, Istiophoridae. Photos A, C-E by Cedric Guigand (specimens from Florida
Straits), B by Julie Mounts and Carole Baldwin.
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Figure 40. Percomorphacea (Scorpaeniformes). A, Scorpaenodes carribaeus, 9.0 mm Standard Length (SL), BLZ 6019.
B, Scorpaena inermis, 7.0 mm SL, Belize. C, Scorpaena grandicornis, 7.5 mm SL, BLZ 10215. D, Scorpaena bergi, 10 mm
SL, BLZ 10232. E, Dendrochirus brachypterus, South Africa. Photo A by Lee Weigt and Carole Baldwin; B by Julie Mounts
and Carole Baldwin; C, D by Donald Griswold and Carole Baldwin; E by Allan Connell.
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of Rypticus (Fig. 43B, D) have additional orange/
yellow chromatophore blotches further anteriorly.
An image of the Pacific Pseudogramma polyacantha
shows a nearly identical colour pattern to that
observed in the Atlantic Pseudogramma gregoryi
except that the chromatophores are xanthophores
rather than erythrophores (Fig. 44). Additional speci-
mens of Pseudogramma polyacantha are needed to
determine whether the colour is always yellow, but in
multiple colour images of Pseudogramma gregoryi
examined from Belize larval-fish collections, the
colour is always orange.
Larvae of other epinepheline serranids examined ?
the Spanish flag Gonioplectrus, the grouper Myctero-
perca, and the liopropomin basses Bathyanthias and
Liopropoma (Figs 45, 46) ? do not have enlarged
pectoral fins. Gonioplectrus has orange pigment on and
at the base of the elongate second dorsal-fin spine, in a
large blotch over the opercular region and base of
pectoral fin, and along the entire length of the elongate
pelvic-fin spine (Fig. 45). The pelvic-fin spine is also
orange in settlement-stage Mycteroperca bonaci larvae
(Fig. 45). Gonioplectrus has only melanophores in a
blotch on the caudal peduncle, whereas Mycteroperca
has erythrophores in a similar arrangement and posi-
tion. Larval Bathyanthias from the Florida Straits and
reared Liopropoma rubre larvae have two extremely
elongate dorsal-fin spines, the anterior of which has
pigmented swellings along its length (Fig. 46). In
Bathyanthias, the swellings are small and orange,
whereas in Liopropoma they are very large, and each
is encircled with a prominent band of yellow pigment.
Larvae of both genera have a stripe of erythrophores
from the snout to the eye and additional erythrophores
on the tip of the lower jaw and on the head behind the
eye. Bathyanthias has orange pigment on the second
dorsal, caudal, anal, and pelvic fins, but this pigment
is not evident in available photographs of Liopropoma.
Larvae of one anthiine serranid, Pseudanthias cooperi,
do not greatly resemble other serranids in colour
pattern: they are mostly pale, with a bright orange
blotch at the posterior end of the lower jaw, pale
xanthophores on the head, and pale xanthophores and
erythrophores on the trunk (Fig. 47).
Stromateiformes
Stromateiformes are represented in the Belize
material by one 5.0-mm SL unidentified specimen
that resembles an illustration of a 5.1-mm SL larva
of Cubiceps pauciradiatus (Lamkin, 2006) in pattern
Figure 41. Percomorphacea (Scorpaeniformes). Peristedion sp., Florida Straits. Photo by Cedric Guigand.
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of melanophores. The DNA barcode for the Belize
larva is approximately 3% different from that of
Cu. pauciradiatus, which is greater than typical
intraspecific divergence (Weigt et al., 2012). Further
study is needed, but the DNA data clearly identify
the larva as a nomeid. It is mostly pale, but has
xanthophores mixed with chromatophores on the top
of the head and in a series over the swimbladder
and gut (Fig. 48). Two species of Stromateidae from
South Africa (Connell, 2007) have xanthophores
mixed with melanophores in preflexion stages, and
one postflexion specimen has most of the head and
Figure 42. Percomorphacea (Scorpaeniformes). A, Serranus baldwini, 9.5 mm Standard Length (SL), BLZ 8400.
B, Diplectrum bivittatum, 12 mm SL, BLZ 7318. C, Hypoplectrus sp., 5.0 mm SL, BLZ 4588. Photos A, B by Julie Mounts
and David Smith; C by Julie Mounts and Carole Baldwin.
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Figure 43. Percomorphacea (Scorpaeniformes). A, Rypticus sp., 12.0 mm Standard Length (SL), Belize. B, Rypticus sp.,
11.5 mm SL, Belize. C, Rypticus bistrispinus, 8 mm SL, BLZ 7715. D, Pseudogramma gregoryi, 11 mm SL. Photos A, B,
D by Julie Mounts and Carole Baldwin; C by Lee Weigt and Carole Baldwin.
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trunk covered with xanthophores and large dark
spots.
Tetraodontiformes
Tetraodontiformes are represented in the material
examined by larval monacanthids and ostraciids
(Balistoidei), and tetraodontids and a molid
(Tetraodontoidei). Larval Canthigaster rostrata and
Sphoeroides (Tetraodontidae) have strikingly similar
colour patterns in that most of the body is covered
with iridophores that reflect bronze/gold/blue colours
(Fig. 49A?C). A similar shimmering colour pattern is
present in larval Ranzania, a molid (Fig. 49D). Colour
pattern in larvae of these tetraodontoids, which is
very different from that observed in larval ostraciids
and monacanthids, may provide corroborative evi-
dence for the monophyly of the Tetraodontoidei (sensu
Holcroft, 2005) or a less inclusive clade within that
Figure 44. Percomorphacea (Scorpaeniformes). Top, posterior region of larval Pseudogramma gregoryi, Florida Straits,
photo by Cedric Guigand. Bottom, posterior region of larval Pseudogramma polyacantha, South Africa, photo by Allan
Connell.
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Figure 45. Percomorphacea (Scorpaeniformes). Top, Gonioplectrus hispanus, Florida Straits, photo by Cedric Guigand.
Bottom, Mycteroperca bonaci, 21.0 mm Standard Length, Belize, photo by Julie Mounts and Carole Baldwin.
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Figure 46. Percomorphacea (Scorpaeniformes). Top, Liopropoma rubre, reared, photo by Christopher Paparo. Bottom,
Bathyanthias sp., Florida Straits, photo by Cedric Guigand.
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Figure 47. Percomorphacea (Scorpaeniformes). Pseudanthias cooperi, South Africa. Photo by Allan Connell.
Figure 48. Percomorphacea (Stromateiformes). Nomeidae (Cubiceps?), 5.0 mm Standard Length, BLZ 6047. Photo by
Lee Weigt and Carole Baldwin.
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Figure 49. Percomorphacea (Tetraodontiformes). A, Canthigaster rostrata, 12.0 mm Standard Length (SL), Belize. B,
Sphoeroides spengleri, 7.0 mmSL, BLZ 10114. C, Sphoeroides testudineus, 5 mmSL, BLZ 10147. D, Ranzania laevis, Florida
Straits. Photo A by Julie Mounts and Carole Baldwin; B, C by Donald Griswold and Carole Baldwin; D by Cedric Guigand.
548 C. C. BALDWIN
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assemblage. Monocanthus ciliatus larvae are mostly
yellow, with xanthophores present on the head,
trunk, and spinous dorsal and pelvic fins, and larval
Aluterus is less colourful but has some yellow pig-
ment on the head, trunk, and spinous dorsal and
caudal fins (Fig. 50A, B). Larval ostraciids have a pale
orange/yellow coloration overlain by dark dots or
large dark spots (Fig. 50C, D). Relationships of tetrao-
dontiforms to other acanthomorph fishes are unre-
solved, but taxa hypothesized to be close relatives
include zeiforms, acanthuriforms, and lophiiforms
(Wiley & Johnson, 2009, and references therein).
There is little resemblance between tetraodontiform
larvae examined and Z. faber (Connell, 2007; Appen-
dix), and, as noted above (see Acanthuriformes), there
is little similarity between acanthurid and tetraodon-
tiform larvae. There is at least superficial resem-
blance between tetraodontid and chaetodontid/
pomacentrid larvae in that the pigment is primarily
in the form of iridophores (Figs. 30, 49).
The tetraodontiform/lophiiform hypothesis (Holcroft
& Wiley, 2008; Miya et al., 2010) is intriguing in light
of colour patterns and other aspects of larval morphol-
ogy: compare a preflexion larval ostraciid from South
Africa with an unidentified lophiiform (Fig. 51). In
both, but in larvae of no other teleosts examined,
the trunk is contained within an inflated, three-
dimensional sac that is covered with xanthophores.
Aboussouan & Leis (2004) noted the presence of a
?dermal sac? in tetraodontids, diodontids, molids,
ostraciids, and balistids but not in triacanthodids,
triacanthids, and monacanthids. Pietsch (1984: 320)
described lophiid and other lophiiform larvae as
having the epidermal layer of the head and body
?greatly distended by transparent, gelatinous connec-
tive tissue?. The inflated condition in tetraodontiforms
appears to be restricted to the preflexion stage,
whereas that in lophiiforms persists after flexion
(compare Figs 50C, D and 51; see also Connell, 2007).
Trachiniformes
Trachiniformes, which currently comprise the
fishes traditionally included in the Trachinoidei
(Chiasmodontidae, Champsodontidae, Pinguipedidae,
Cheimarrhichthyidae, Trichodontidae, Creediidae,
Percophididae, Leptoscopidae, Trachinidae, and
Uranoscopidae) and Ammodytidae, were considered
likely to be paraphyletic by Wiley & Johnson (2009).
Among trachinoid larvae from off South Africa
(Connell, 2007; Appendix), an early-stage larval
uranoscopid bears little resemblance to an early-stage
chiasmodontid and a champsodontid. The last two are
similar in having only melanophores and lacking
obvious nonmelanistic colour in the preflexion stage,
whereas the larval uranoscopid has distinctive
blotches of xanthophores. Mito (1962) described a
reared series of Champsodon snyderi and noted
that the species has only melanophores throughout
larval development. Information on colour patterns
in postflexion larvae of other champsodontids might
be useful in commenting on the proposed relation-
ship between champsodontids and scorpaeniforms
(Mooi & Johnson, 1997). As noted (see ?Scorpaeni-
formes? above), the latter have distinctive patterns of
erythrophores/xanthophores.
DISCUSSION
The first broad comparative survey of nonmelanistic
colour in marine teleost larvae presented herein has
revealed a striking array of colour patterns that may
inform phylogeny at multiple taxonomic levels. The
absence of xanthophores, erythrophores, and irido-
phores in pelagic larvae of most basal marine teleosts
examined, and their presence in many basal neotele-
osts, in all percomorph orders examined, and in
nearly every percomorph species examined, suggest a
phylogenetic trend (Fig. 1). It is premature to make
specific hypotheses about the evolution of the three
types of chromatophores in larvae of marine teleosts,
but in the limited material examined, xanthophores
are the only type of nonmelanistic chromatophore
present in larval marine anguilliforms and stomiati-
forms. Erythrophores appear at the level of Euryp-
terygia, and iridophores first appear in larval
euacanthopterygians. The situation in Anguilliformes
and basal neoteleosts, in which larvae of some
members of an order exhibit nonmelanistic pigment
but others do not, suggests that nonmelanistic chro-
matophores have arisen independently (or were lost)
multiple times. Considerably more information on
colour patterns in more larvae is needed to determine
the taxonomic distribution of the various types of
chromatophores among teleosts, as is investigation of
the cellular basis of observed chromatophore types. In
this study, presence or absence of chromatophores
was assessed from visual examination of fresh speci-
mens and colour photographs, but histological studies
are needed to confirm that the presence or absence of
a particular nonmelanistic pigment corresponds to
the structural presence or absence of the vertebrate
chromatophore unit that typically contains that
pigment. Biological and physical factors that might
affect the development or display of chromatophores
also warrant study. For example, thyroid hormones
are known to have a role in regulating larval-fish
development, including pigment (e.g. Brown & Kim,
1995, and references therein). However, experimental
manipulation of exogenous hormones suggests they
may affect timing of pigment development and
density/shape of pigment features but not the types of
chromatophores that develop (Reddy & Lam, 1992;
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Figure 50. Percomorphacea (Tetraodontiformes). A, Monacanthus ciliatus, 8.0 mm Standard Length (SL), BLZ 10223.
B, Aluterus schoepfi, 7.5 mm SL, Belize, USNM 353531. C, Ostraciidae, 10.0 mm SL, Belize. D, Lactophrys trigonus,
6.0 mm SL, BLZ 8446. Photo A by Donald Griswold and Carole Baldwin; B by David Smith; C by Julie Mounts and Carole
Baldwin; D by Julie Mounts and David Smith.
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Brown & Kim, 1995; Clement, Lichtenbert &
Kohler, 2001). It seems unlikely that physical factors
related to habitat ecology have much bearing on the
presence or absence of nonmelanistic chromato-
phores, as larvae of most marine teleosts inhabit
a similar planktonic realm. In the material examined
for this study, fish larvae such as those of clupeids,
engraulids, elopiforms, and anguilliforms that
lack nonmelanistic chromatophores were collected
in the same plankton samples as larvae that have
them.
Apomorphic characters of chromatophore patterns
rather than the presence or absence of chromatophore
types may have the greatest phylogenetic potential in
teleosts. Pigment patterns of adults have been incor-
porated into phylogenetic studies of fishes at subge-
neric, generic, and familial levels (e.g. Mabee, 1995;
Ayache & Near, 2009; Layman & Mayden, 2012), but
nonmelanistic colour patterns of marine fish larvae
have not. Among the percomorphs examined, colour
patterns in marine fish larvae may have phylogenetic
significance at several taxonomic levels: interordi-
nal ? Mugiliformes/Beloniformes, Tetraodontiformes/
Lophiiformes; interfamilial ? Chaeotodontidae/Poma-
centridae, Bothidae/Paralichthyidae, Chaenopsidae/
Labrisomidae, Callionymidae/Gobiesocidae; familial ?
Apogonidae, Eleotrididae, Microdesmidae, Scaridae,
Scorpaenidae, Tetraodontidae; intergeneric ? Abu-
defduf/Amblyglyphidodon, Anampses/Halichoeres/
Thalassoma, Ctenogobius/Gnatholepis/Gobionellus,
Nes/Psilotris, Lutjanus/Ocyurus; and generic ? Bathy-
gobius, Coryphopterus, Scorpaena, Rypticus, Xyrich-
tys. Slight differences in generic colour patterns of
larvae ? including whether the pattern comprises
xanthophores or erythrophores ? often distinguish
component species (e.g. those of Xyrichtys, Pseudo-
gramma, Scorpaena).
Matsumoto (1965) noted that vesicles that contain
yellow and red pigments are sometimes found in the
same cell in freshwater Xiphophorus, suggesting that
the distinction between xanthophores and erythro-
phores is artificial. The presence in some marine
genera of species with nearly identical patterns of
pigment except for the colour of the chromatophores
(orange or yellow) suggests that xanthophores and
erythrophores are distinct. Erythrophores in fishes
reportedly are red/orange because of carotenoids
ingested in the diet, whereas the yellow pigment of
xanthophores (from pteridines) is metabolized endog-
enously (Grether et al., 2004, and references therein).
Whether the origins of yellow and orange pigments in
marine fish larvae also are endogenous and exog-
enous, respectively, does not appear to have been
investigated. It seems remarkable that nearly identi-
cal patterns of pigment that differ only in colour (e.g.
Xyrichtys, Fig. 26A, C; Pseudogramma, Fig. 44) would
have different biochemical origins. Future studies
that investigate the biochemistry of pigments in
marine fish larvae might also compare the chemistry
of pigments in basal teleosts such as eel leptocephali,
basal neoteleosts such as whalefish, and perco-
morphs. Similarities or differences in the biochemis-
try of the pigments may shed light on the homology
and evolutionary history of larval-fish pigments
among marine teleosts.
Kendall et al. (1984) and Mabee (1995) noted that
individual variation and convergent evolution of some
patterns are considered obstacles for phylogenetic use
of pigment characters because of difficulties in estab-
lishing hypotheses of homology. Numerous specimens
of many species were examined in this study (Appen-
dix), and individual variation in nonmelanistic chro-
matophore patterns is low. Differences frequently
appear to reflect the expanded or contracted state of
Figure 51. Percomorphacea (Tetraodontiformes). Left, unknown lophiiform, Florida Straits, photo by Cedric Guigand
(previously published in Fogarty & Botsford [2007]). Right, Ostraciidae, South Africa, photo by Allan Connell.
COLOUR IN MARINE TELEOST LARVAE 551
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the chromatophores rather than their pattern
(Fig. 52). Assessing homology of nonmelanistic chro-
matophores in larvae among major groups of marine
teleosts through phylogeny is complicated by the
absence of a comparable, transient, pigment phase in
most freshwater fishes. Larvae of marine Clupei-
formes examined, for example, lack nonmelanistic
chromatophores, but most Otomorpha are freshwater
inhabitants. The presence of xanthophores and irido-
phores in larval cypriniforms such as freshwater
Danio is well documented (Johnson et al., 1995;
Parichy et al., 2000; Parichy, 2003, 2006). What is the
relationship between the pigment pattern of larval
freshwater fishes and that of pelagic marine fish
larvae? As shown in the ontogenetic series of numer-
ous marine teleosts photographed by Connell (2007),
the pigment pattern of fully developed marine fish
larvae is preceded by a very different pigment phase
in recently hatched larvae. For example, a reared
myctophid series shows yellow pigment on the head
and body in yolksac larvae, but all yellow pigment has
disappeared by 4.3 mm NL. Acanthurid acronurus
larvae examined have iridophores but no erythro-
phores or xanthophores, yet preflexion Acanthurus
mata exhibit xanthophores (see ?Acanthuriformes?
above). Studies on aquarium-reared callionymids
revealed a colour transformation in larval Syn. splen-
didus from yellow in the preflexion stage to bright
orange after flexion (Fig. 19; Wittenrich et al., 2010).
Possibly the early pigment pattern in marine fish
larvae is the ontogenetic counterpart of the larval
pigment pattern in freshwater fishes. Assembling
ontogenetic series of marine fishes that incorporate
chromatophore patterns from recently hatched to
adult stages, investigating the development of the
pattern at each stage, and comparing the develop-
ment of pigment among marine and freshwater fishes
would shed light on the homology of pigment patterns
among life-history stages and among related taxa.
Such ontogenetic series also warrant further study
as a potential source of phylogenetic characters.
Mabee (1995) highlighted the phylogenetic signifi-
cance of pigment-pattern development in the evolu-
tion of centrarchid sunfishes, which are freshwater
fishes that exhibit direct development of the adult
pigment pattern from the early larval stage. The
ontogeny of pigment patterns in marine fishes may be
an even riper source of phylogenetic information yet
to be tapped. In addition to the colour transitions
in myctophids, acanthurids, and callionymids men-
tioned above, the ontogenetic transition in mugilids
and some beloniforms from a stage dominated by
xanthophores, erythrophores, and melanophores to
one dominated by iridophores is an example. This
transition also suggests that the presence of irido-
phores may be homologous in the two groups because
the condition was derived from a similar ontogene-
tic transformation. Incorporating developmental
studies of colour patterns in future phylogenetic
treatments of the topic could thus provide both crucial
Figure 52. Intraspecific variation in pattern of erythrophores in Halichoeres bivittatus. A, 12.5 mm Standard Length
(SL), BLZ 6426. B, 12.5 mm SL, BLZ 6424. Note that the pattern of orange pigment is nearly identical in the two
specimens but that in BLZ 6426 the erythrophores in the vertical series behind the head appear to be contracted. Photos
by Julie Mounts and David Smith.
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information on character homology and new phyloge-
netic characters.
Yasir & Qin (2007) described developmental
changes in pigment patterns of the marine anemon-
efish Amphiprion ocellaris from embryo to larva and
noted that a better understanding of the process of
pigment formation is needed. Embryologically, neural
crest cells, which give rise to all four types of chro-
matophores, migrate to the dermis, and the density
and distribution of the chromatophores are integrated
to produce colour patterns in vertebrates (Hawkes,
1974; Bagnara, 1987; Hall, 1999). Holt (2011) noted
that the pigment patterns of fishes are formed during
embryogenesis and are replaced by the adult pattern
at metamorphosis. Studies of freshwater Danio indi-
cate that the adult colour pattern results from incor-
poration of both embryonic chromatophores and
differentiation of new chromatophores from stem cells
at metamorphosis, and the ratio of contribution
by the two may vary significantly among species
(Parichy, 2003, 2006). The existence of an additional
colour phase (between the recently hatched and adult
phases) in pelagic larvae of marine fishes would
appear to complicate this scenario. Furthermore,
juveniles of many marine fishes exhibit yet another
colour phase that is different from the pelagic larval
and adult phases (e.g. Haemulidae, Labridae, Poma-
centridae, Pomacanthidae). The ontogeny of chro-
matophore patterns has been studied in few marine
fishes, but Nakamura et al. (2010 and references
therein) have investigated development of the adult
pigment pattern in Pleuronectiformes. Two types of
melanophores and xanthophores/erythrophores are
hypothesized to exist in pleuronectiforms ? those that
develop in the larval stage prior to metamorphosis
and those that develop afterwards and form the adult
pigment pattern. Flatfishes exhibit an unusual ontog-
eny in which larvae are bilaterally symmetrical,
including the pigment pattern, but metamorphosis
involves migration of one eye to the other side of the
head resulting in a bilaterally asymmetrical juvenile
and adult. There is typically little or no pigment on
the ?blind? side but the ?eyed? side is well pigmented.
Whether chromatophores of pelagic larvae in other
marine fishes are succeeded ontogenetically by adult
versions of those pigment cells is unknown, but
this would be a reasonable hypothesis based on the
different pigment patterns exhibited in larval and
adult stages. Other features that are present in
pelagic larvae of some marine fishes, such as elongate
fin rays and head spines, disappear or transform
upon metamorphosis to the juvenile stage. Studies
in marine fishes that would explain the origin of
adult pigment patterns (i.e. from existing neural
crest lineages or newly differentiated stem cells) are
lacking.
If the colour patterns exhibited by marine fish larvae
are present only during the pelagic larval period, are
they adaptations that have functional significance?
Grether et al. (2004) discussed the potential signifi-
cance of the different pigments in adult freshwater
fishes, but only one of their hypotheses seems poten-
tially relevant to pelagic marine fish larvae ? ?spectral
fine-tuning?. Grether et al. (2004) noted that orange
and yellow pigments have different absorptive proper-
ties that may enable spectral fine-tuning, which, in
turn, may enhance the ability to blend into the back-
ground. Orange coloration is not restricted to fishes in
a plankton sample: numerous invertebrates such as
the larvae of shrimps and crabs look very similar.
Indeed, the guts of numerous freshly caught marine
fish larvae are orange because of diet (see Figs 27D,
39D, 41, 46). Carvalho, Zuanon & Sazima (2006)
provided examples of transparency and similar colour
patterns in freshwater fishes and crustaceans that
travel as a group and suggested that these organisms
may be utilizing a type of protective association known
as numerical mimicry as a means of avoiding potential
predators. Regarding the bright orange coloration
in larvae of the mandarinfish, Syn. splendidus,
Wittenrich et al. (2010) noted that Lindquist (2002)
and Young & Bingham (1987) hypothesized that
orange may be an aposematic warning colour in the
larvae of some marine organisms.
In summary, colour patterns in marine teleost
larvae emerge as an intriguing new source of poten-
tially valuable phylogenetic information, but consid-
erably more data are needed to fully assess their
significance. An increased global focus on document-
ing colour patterns in larvae prior to preservation
would facilitate incorporating information from larval
colour patterns into more systematic studies, and
developmental, histological, and biochemical studies
that shed light on the ontogeny of colour patterns in
early life history stages would improve our under-
standing of ontogenetic pigment phases, characters,
character transformations, and character homology.
ACKNOWLEDGEMENTS
David Smith and Lee Weigt contributed to the field
work that resulted in the Belize larval-fish images
in numerous ways. The molecular data that enabled
identifications of larvae analysed in this paper were
published by Weigt et al. (2012), and a complete list of
acknowledgements for the Belize field work is pro-
vided in that paper. Julie Mounts contributed greatly
to photographic efforts in Belize. David Smith pro-
vided identifications of several larvae not identified
through DNA analysis. David Johnson identified or
corrected my misidentifications of species represented
in several images by Joshua Lambus and Cedric
COLOUR IN MARINE TELEOST LARVAE 553
? 2013 The Linnean Society of London, Zoological Journal of the Linnean Society, 2013, 168, 496?563
Guigand, and he provided contact information for
those photographers and Donald Hughes. I thank
Allan Connell, Cedric Guigand, Joshua Lambus,
Matthew D?Avella, Michael Miller, Donald Hughes,
Chris Papero, Todd Gardner, Matthew Wittenrich,
Robert Cowen, Daniel Benetti, and Dallas Alston for
allowing me to use their images of fish larvae or
assisting me in locating images. The addition of
numerous fish species not represented in the Belize
material greatly increased the taxonomic scope of the
paper, and many of the additional images are so
stunning that it was a privilege to study and incor-
porate them into this work. Ross Robertson provided
a portable Wacom tablet and provided instructions on
how to remove images from their original back-
grounds. Without this editorial procedure, many of
the Belize larval-fish images would not have been
publishable. Sandra Raredon and Jeff Williams pro-
vided initial guidance on the best photographic equip-
ment and equipment set-up for photography in the
field. Wen-Hsiung Li, Chief Editor of Zoological
Studies, and Aoi Sasaki of TERRAPUB (publisher of
Aqua-BioScience Monographs), allowed the images in
Figure 4B and Figure 5, respectively, to be repro-
duced. Katsumi Aida, Editor in Chief of Aqua-
BioScience Monographs, facilitated communication
with Sasaki. Fabio Di Dario, this journal?s editorial
staff, and anonymous reviewers provided suggestions
that greatly improved the manuscript. This is contri-
bution number 937 of the Caribbean Coral Reef Eco-
systems Program, Smithsonian Institution, supported
in part by the Hunterdon Oceanographic Research
Fund. Publication of the colour figures was made
possible in part through the generous support of
the Herbert R. and Evelyn Axelrod Endowment
Fund for systematic ichthyology and Mrs. Alexandra
S. Baldwin.
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aposematic coloration in larvae of the ascidian Ecteinascidia
turbinate. Marine Biology 96: 539?544.
COLOUR IN MARINE TELEOST LARVAE 557
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APPENDIX
Teleost larvae for which images were examined for colour patterns. ?N? in the ?Colour? column indicates no
xanthophores, erythrophores or iridophores; ?Y? indicates the presence of any of those types of chromatophores
alone or in any combination. The table is colour-coded by teleost cohort: blue indicates Clupeomorpha, green
Elopomorpha, purple Euteleosteomorpha.
Order Family Genus Species Colour Image
Clupeiformes Clupeidae Harengula humeralis N BLZ 5175
Clupeiformes Clupeidae Harengula clupeola N Fig. 3
Clupeiformes Clupeidae Jenkensia lamprotaenia N Fig. 3 and other BLZ images
Clupeiformes Clupeidae Jenkensia sp. N BLZ 8418
Clupeiformes Engraulidae Anchoa cayorum N BLZ 8468
Clupeiformes Engraulidae Anchoa sp. N Fig. 3 and other BLZ images
Albuliformes Albulidae Albula vulpes N Fig. 2 and other BLZ images
Anguilliformes Chlopsidae Chilorhinus seunsoni N Fig. 2 and other BLZ images
Anguilliformes Moringuidae Moringua edwardsi N Fig. 2
Anguilliformes Muraenidae Gymnothorax moringa N Fig. 2
Anguilliformes Muraenidae Strophidion ui Y Tawa et al. (2012)
Anguilliformes Muraenidae Unknown Unknown Y Fig. 5 and Miller (2009)
Anguilliformes Nettastomatidae Saurenchelys sp. Y Fig. 5 and Miller (2009)
Anguilliformes Ophichthidae Aprognathodon platyventris N Fig. 2 and other BLZ images
Anguilliformes Ophichthidae Myrichthys breviceps Y Fig. 4 and other BLZ images
Anguilliformes Ophichthidae Myrophis punctatus N Fig. 2 and other BLZ images
Anguilliformes Ophichthidae Myrophis platyrhynchus N Fig. 2 and other BLZ images
Anguilliformes Ophichthidae Ahlia egmontis N Fig. 2 and other BLZ images
Anguilliformes Ophichthidae Unknown Unknown Y Fig. 4 and Miller et al. (2010)
Anguilliformes Ophichthidae Unknown Unknown Y Fig. 5 and Miller (2009)
Anguilliformes Ophichthidae Neenchelys sp. Y Fig. 5 and Miller (2009)
Anguilliformes Ophichthidae Unknown Unknown Y Fig. 5 and Miller (2009)
Elopiformes Elopidae Elops saurus N Fig. 2
Elopiformes Megalopidae Megalops atlantica N Fig. 2
Stomiatiformes Melanostomiatidae Opostomias micipnus Y http://www.fisheggsandlarvae.com/
FIA2%20Melanostomiidae.htm
Stomiatiformes Phosichthyidae Vinciguerria nimbaria N http://www.fisheggsandlarvae.com/
DIIIA1%20Phosichthyidae.htm
Aulopiformes Synodontidae Saurida sp. N Fig. 6 and other BLZ images
Aulopiformes Synodontidae Saurida sp. N Fig. 6 and other BLZ images
Aulopiformes Synodontidae Synodus foetens N BLZ 6429
Aulopiformes Synodontidae Synodus synodus N Fig. 6 and other BLZ images
Aulopiformes Synodontidae Trachinocephalus myops N BLZ 7061
Aulopiformes Bathypteroidae? Unknown Unknown Y Fig. 7
Aulopiformes Giganturidae Gigantura sp. N Fig. 7
Myctophiformes Myctophidae Unknown Unknown Y http://www.fisheggsandlarvae.com/
CLIIA2%20Myctophidae.htm
Myctophiformes Unknown Unknown Unknown N Fig. 7
Lampridiformes Trachipteridae? Unknown Unknown Y Fig. 8
Lampridiformes Lampridae Lampris guttatus Y Fig. 8
Gadiformes Bregmacerotidae Bregmaceros sp. N Fig. 6 and other BLZ images
Gadiformes Gadidae Unknown Unknown Y http://www.fisheggsandlarvae.com/
LIIIF1%20Gadidae.htm
Beryciformes Berycidae Centroberyx sp. Y http://www.fisheggsandlarvae.com/
EIIA3%20Berycidae.htm
Beryciformes Berycidae Unknown Unknown Y Fig. 9
Beryciformes Holocentridae Myripristis berndti Y http://www.fisheggsandlarvae.com/
EIIIB6A%20Myripristis.htm
Beryciformes Holocentridae Unknown (2) Unknown (2) Y Fig. 9A, B
Beryciformes Holocentridae Unknown Unknown Y Fig. 9
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APPENDIX Continued
Order Family Genus Species Colour Image
Stephanoberyciformes Cetomimidae Unknown Unknown Y Fig. 10
Zeiformes Zeidae Zeus faber Y http://www.fisheggsandlarvae.com/
LIA3%20Zeus%20faber.htm
Acanthuriformes Acanthuridae Acanthurus chirurgus Y Fig. 14 and other BLZ images
Acanthuriformes Acanthuridae Acanthurus bahianus Y Fig. 14 and other BLZ images
Acanthuriformes Acanthuridae Unknown Unknown Y Fig. 15
Acanthuriformes Acanthuridae Acanthurus mata Y http://www.fisheggsandlarvae.com/
LIIIE7%20Acanthuridae.htm
Atheriniformes Atherinidae Atherinomorus stipes Y Fig. 13 and other BLZ images
Atheriniformes Atherinidae Unknown Unknown Y Fig. 13
Beloniformes Belonidae Unknown Unknown Y BLZ, 18 mm SL Photo
2-23-2004.tif
Beloniformes Belonidae Platybelone argalus Y Fig. 12 and other BLZ images
Beloniformes Exocoetidae Unknown Unknown Y BLZ 7036
Beloniformes Exocoetidae Prognichthys occidentalis Y Fig. 11
Beloniformes Hemirhamphidae Hemirhamphus brasiliensis Y Fig. 12
Beloniformes Hemirhamphidae Oxyporhamphus micropterus Y http://www.fisheggsandlarvae.com/
CHIA1%20Oxyporhamphus.htm
Blenniiformes Chaenopsidae Acanthemblemaria aspera Y BLZ 6046, 8449 and others
Blenniiformes Chaenopsidae Acanthemblemaria greenfieldi Y Fig. 16 and other BLZ images
Blenniiformes Labrisomidae Labrisomus cricota Y Fig. 16 and other BLZ images
Blenniiformes Labrisomidae Labrisomus haitiensis Y BLZ 4513, 4360 and others
Blenniiformes Labrisomidae Labrisomus gobio Y BLZ 8447
Blenniiformes Labrisomidae Labrisomus bucciferus Y Fig. 16 and other BLZ images
Blenniiformes Labrisomidae Malacoctenus macropus Y BLZ 8416, 7021 and others
Blenniiformes Labrisomidae Malacoctenus triangulatus Y Fig. 16 and other BLZ images
Blenniiformes Labrisomidae Paraclinus fasciatus Y Fig. 16 and other BLZ images
Caproiformes Caproidae Antigonia rubescens Y http://www.fisheggsandlarvae.com/
LIIIE4%20Caproidae.htm
Carangiformes Carangidae Trachinotus falcatus Y Fig. 17
Carangiformes Carangidae Selene vomer Y Fig. 18
Carangiformes Carangidae Seriola sp. Y http://www.fisheggsandlarvae.com/
EIIA2%20Carangidae.htm
Carangiformes Coryphaenidae Coryphaena hippurus Y Fig. 18
Gasterosteiformes Aulostomidae Aulostomus chinensis Y http://www.fisheggsandlarvae.com/
LIIA3%20Aulostomidae.htm
Gasterosteiformes Fistularidae Fistularia sp. Y http://www.fisheggsandlarvae.com/
HIA3%20Fistularia.htm
Gasterosteiformes Centriscidae Aeoliscus punctulatus Y http://www.fisheggsandlarvae.com/
LIIA5%20Centriscidae.htm
Gasterosteiformes Syngnathidae Unknown Unknown Y BLZ 6408
Gasterosteiformes Syngnathidae Cosmocampus albirostris Y Fig. 12 and other BLZ images
Gasterosteiformes Syngnathidae Cosmocampus elucens Y BLZ 10222, 10160 and others
Gasterosteiformes Syngnathidae Penetopteryx nanus Y Fig. 12 and other BLZ images
Gobiesociformes Callionymidae Callionymus bairdi Y Fig. 19 and other BLZ images
Gobiesociformes Callionymidae Callionymus marleyi Y Fig. 19
Gobiesociformes Callionymidae Draculo celetus Y http://www.fisheggsandlarvae.com/
CDIIIA1%20Callionymidae.htm
Gobiesociformes Callionymidae Synchiropus splendidus Y Fig. 19 and Wittenrich et al. (2010)
Gobiesociformes Gobiesocidae Acyrtops beryllinus Y Fig. 19
Gobiiformes Eleotrididae Erotelis sp. Y Fig. 20 and other BLZ images
Gobiiformes Gobiidae Bathygobius lacertus Y Fig. 22
Gobiiformes Gobiidae Bathygobius soporator Y Fig. 22 and other BLZ images
Gobiiformes Gobiidae Bathygobius curacao Y Fig. 22 and other BLZ images
Gobiiformes Gobiidae Coryphopterus personatus Y Fig. 22 and other BLZ images
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APPENDIX Continued
Order Family Genus Species Colour Image
Gobiiformes Gobiidae Coryphopterus tortugae Y Fig. 22 and other BLZ images
Gobiiformes Gobiidae Coryphopterus glaucofraenum Y BLZ 5225
Gobiiformes Gobiidae Coryphopterus venezuelae Y Fig. 22 and other BLZ images
Gobiiformes Gobiidae Coryphopterus eidolon Y BLZ 5318
Gobiiformes Gobiidae Coryphopterus kuna Y Fig. 22 and other BLZ images
Gobiiformes Gobiidae Ctenogobius saepepallens Y Fig. 21 and other BLZ images
Gobiiformes Gobiidae Gnatholepis thompsoni Y Fig. 21 and other BLZ images
Gobiiformes Gobiidae Gobionellus oceanicus Y Fig. 21 and other BLZ images
Gobiiformes Gobiidae Nes longus Y Fig. 21 and other BLZ images
Gobiiformes Gobiidae Priolepis hipoliti Y Fig. 22 and other BLZ images
Gobiiformes Gobiidae Psilotris sp. Y Fig. 21 and other BLZ images
Gobiiformes Gobiidae Psilotris sp. Y BLZ 6015
Gobiiformes Microdesmidae Cerdale floridana Y Fig. 20 and other BLZ images
Gobiiformes Microdesmidae Microdesmus carri Y Fig. 20 and other BLZ images
Gobiiformes Microdesmidae Microdesmus bahianus Y Fig. 20 and other BLZ images
Gobiiformes Microdesmidae Ptereleotris helenae Y Fig. 20 and BLZ 5444
Labriformes Labridae Anampses lineatus Y Connell pers. comm.
Labriformes Labridae Doratonotus megalepis Y Fig. 26 and other BLZ images
Labriformes Labridae Halichoeres bivittatus Y Fig. 25 and other BLZ images
Labriformes Labridae Halichoeres poeyi Y Fig. 25 and other BLZ images
Labriformes Labridae Halichoeres garnoti Y Fig. 25 and other BLZ images
Labriformes Labridae Halichoeres maculipinna Y Fig. 25 and other BLZ images
Labriformes Labridae Lachnolaimus maximus Y Fig. 26 and other BLZ images
Labriformes Labridae Thalassoma bifasciatum Y Fig. 25 and other BLZ images
Labriformes? Labridae Xyrichtys martinicensis Y Fig. 26 and other BLZ images
Labriformes Labridae Xyrichtys splendens Y Fig. 26 and other BLZ images
Labriformes Labridae Xyrichtys novacula Y Fig. 26 and other BLZ images
Labriformes Pomacentridae Abudefduf saxatilis Y Fig. 24
Labriformes Pomacentridae Chromis cyanea Y Fig. 24
Labriformes Pomacentridae Stegastes planifrons Y Fig. 24 and other BLZ images
Labriformes Pomacentridae Stegastes variabilis Y Fig. 24 and other BLZ images
Labriformes Pomacentridae Stegastes diencaeus Y BLZ 8452, 8404
Labriformes Pomacentridae Stegastes adustus Y BLZ 7046
Labriformes Pomacentridae Stegastes leucostictus Y BLZ 4585, 8472 and others
Labriformes Pomacentridae Stegastes partitus Y Fig. 24 and other BLZ images
Labriformes Pomacentridae Amblyglyphidodon ternatensis Y Fig. 24
Labriformes Scaridae Cryptotomus roseus Y Fig. 23 and other BLZ images
Labriformes Scaridae Scarus iseri Y Fig. 23 and other BLZ images
Labriformes Scaridae Sparisoma radians Y Fig. 23 and other BLZ images
Labriformes Scaridae Sparisoma chrysopterum Y Fig. 23 and other BLZ images
Labriformes Scaridae Sparisoma atomarium Y Fig. 23 and other BLZ images
Lophiiformes Antennariidae Antennarius pauciradiatus Y Fig. 27
Lophiiformes Unknown Unknown Unknown Y http://www.fisheggsandlarvae.com/
DIIIA4%20Lophiiformes.htm
Lophiiformes Unknown Unknown Unknown Y Fig. 27
Lophiiformes Unknown Unknown Unknown Y Fig. 27
Lophiiformes Unknown Unknown Unknown Y Fig. 27
Mugiliformes Mugilidae Mugil sp. Y BLZ 4506,
Mugiliformes Mugilidae Mugil sp. Y Fig. 11 and other BLZ images
Mugiliformes Mugilidae Mugil cephalus Y Fig. 11
Mugiliformes Mugilidae Liza tricuspidens Y http://www.fisheggsandlarvae.com/
LIIB9%20Mugilidae.htm
Ophidiiformes Carapidae Carapus bermudensis Y Fig. 28 and other BLZ images
Ophidiiformes Ophidiidae Parophidion schmidti Y Fig. 28 and other BLZ images
Ophidiiformes Ophidiidae Brotulataenia sp. Y Fig. 29
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APPENDIX Continued
Order Family Genus Species Colour Image
Ophidiiformes Ophidiidae Lampogrammus? sp. Y Fig. 29
Perciformes Apogonidae Apogon aurolineatus Y Fig. 32 and other BLZ images
Perciformes Apogonidae Apogon binotatus y BLZ 6331 and others (Baldwin
et al., 2011)
Perciformes Apogonidae Apogon sp. 1 Y BLZ 5260 and others (Baldwin
et al., 2011)
Perciformes Apogonidae Apogon phenax Y BLZ 6361 and other (Baldwin
et al., 2011)
Perciformes Apogonidae Apogon townsendi Y BLZ 6329 and others (Baldwin
et al., 2011)
Perciformes Apogonidae Apogon maculatus Y BLZ 7717 and others (Baldwin
et al., 2011)
Perciformes Apogonidae Apogon mosavi Y BLZ 7713 and others (Baldwin
et al., 2011)
Perciformes Apogonidae Astrapogon puncticulatus Y Fig. 32 and others (Baldwin et al.,
2009a)
Perciformes Apogonidae Astrapogon alutus Y BLZ 6040 and others (Baldwin
et al., 2009b)
Perciformes Apogonidae Astrapogon stellatus Y BLZ 6038 and others (Baldwin
et al., 2009b)
Perciformes Apogonidae Phaeoptyx pigmentaria Y BLZ 7013 and others (Baldwin
et al., 2009b)
Perciformes Apogonidae Phaeoptyx conklini Y BLZ 5039 and others (Baldwin
et al., 2009b)
Perciformes Apogonidae Phaeoptyx xenus Y Fig. 32 and others (Baldwin et al.,
2009a)
Perciformes Chaetodontidae Chaetodon capistratus Y Fig. 30 and other BLZ images
Perciformes Chaetodontidae Chaetodon marleyi Y Fig. 31
Perciformes Gerreidae Eucinostomus jonesi Y Fig. 33 and other BLZ images
Perciformes Gerreidae Eucinostomus harengulus Y Fig. 33 and other BLZ images
Perciformes Gerreidae Eucinostomus gula Y Fig. 33 and other BLZ images
Perciformes Gerreidae Eucinostomus melanopterus Y Fig. 33 and other BLZ images
Perciformes Haemulidae Haemulon aurolineatum Y BLZ 10013
Perciformes Haemulidae Anisotremus virginicus Y Fig. 35 and other BLZ images
Perciformes Haemulidae Haemulon plumieri Y Fig. 35 and other BLZ images
Perciformes Haemulidae Haemulon sciurus Y Fig. 35 and other BLZ images
Perciformes Haemulidae Haemulon flavolineatum Y BLZ 10221 and other BLZ images
Perciformes Haemulidae Pomadasys commersonnii Y http://www.fisheggsandlarvae.com/
EIIIB3%20Haemulidae.htm
Perciformes Scorpididae Neoscorpis lithophilus Y http://www.fisheggsandlarvae.com/
FIIB1%20Neoscorpis.htm
Perciformes Lutjanidae Lutjanus mahogoni Y Fig. 34
Perciformes Lutjanidae Lutjanus analis Y Fig. 34 and other BLZ images
Perciformes Lutjanidae Lutjanus synagris Y Fig. 34 and other BLZ images
Perciformes Lutjanidae Lutjanus vivanus Y Fig. 34
Perciformes Lutjanidae Lutjanus apodus Y BLZ 8429 and others
Perciformes Lutjanidae Lutjanus griseus Y Fig. 34 and other BLZ images
Perciformes Lutjanidae Ocyurus chrysurus Y Fig. 34 and other BLZ images
Perciformes Opistognathidae? Unknown Unknown Y Fig. 36 and other BLZ images
Perciformes Oplegnathidae Oplegnathus robinsoni Y http://www.fisheggsandlarvae.com/
EIIIA2%20Oplegnathidae.htm
Perciformes Oplegnathidae Oplegnathus conwayi Y http://www.fisheggsandlarvae.com/
FIIA7%20Oplegnathidae.htm
Perciformes Pempheridae Pempheris schwenkii Y http://www.fisheggsandlarvae.com/
LIIA2%20Pempheridae.htm
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APPENDIX Continued
Order Family Genus Species Colour Image
Perciformes Pomacanthidae Holacanthus ciliarus Y Fig. 30 and other BLZ images
Perciformes Pomacanthidae Pomacanthus arcuatus Y Fig. 30 and other BLZ images
Perciformes Pomacanthidae Pomacanthus paru Y BLZ 10213
Perciformes Pomacanthidae Pomacanthus rhomboides Y Fig. 31
Perciformes Sciaenidae Atractoscion aequidens Y http://www.fisheggsandlarvae.com/
LIIA6%20Atractoscion.htm
Perciformes Sciaenidae Equetus punctatus Y BLZ 10121
Perciformes Sciaenidae Odontoscion dentex Y Fig. 36
Perciformes Scorpididae Neoscorpis lithophilus Y http://www.fisheggsandlarvae.com/
FIIB1%20Neoscorpis.htm
Perciformes Sparidae Calamus? sp. Y Fig. 35
Perciformes Sparidae Calamus? sp. Y Fig. 35 and other BLZ images
Pleuronectiformes Achiridae Trinectes sp. Y Fig. 37 and other BLZ images
Pleuronectiformes Bothidae Bothus maculiferus Y Fig. 37 and other BLZ images
Pleuronectiformes Bothidae Bothus ocellatus Y Fig. 37 and other BLZ images
Pleuronectiformes Bothidae Unknown Unknown Y Fig. 38
Pleuronectiformes Paralichthyidae Citharichthys? sp. Y Fig. 37 and other BLZ images
Pleuronectiformes Paralichthyidae Syacium sp. Y Fig. 37
Pleuronectiformes Paralichthyidae Syacium sp. Y Fig. 37 and other BLZ images
Pleuronectiformes Paralichthyidae Syacium sp. Y Fig. 37 and other BLZ images
Pleuronectiformes Paralichthyidae Unknown Unknown Y BLZ 7086
Pleuronectiformes Cynoglossidae Symphurus sp. Y Fig. 37 and other BLZ images
Pleuronectiformes Soleidae Solea turbynei Y http://www.fisheggsandlarvae.com/
MIIIA5%20Solea%20bleekeri.htm
Scombriformes Scombridae Auxis rochei Y http://www.fisheggsandlarvae.com/
LIIIA11%20Auxis.htm
Scombriformes Sphyraenidae? Unknown Unknown Y http://www.fisheggsandlarvae.com/
EIIIA2A%20Sphyraenidae.htm
Scombriformes Sphyraenidae Sphyraena barracuda Y Fig. 39 and other BLZ images
Scombriformes Gempylidae Unknown Unknown Y Fig. 39
Scombriformes Istiophoridae Unknown Unknown Y Fig. 39
Scombriformes Scombridae Unknown Unknown Y Fig. 39
Scorpaeniformes Peristediidae Peristedion sp. Y Fig. 41
Scorpaeniformes Scorpaenidae Scorpaena inermis Y Fig. 40 and other BLZ images
Scorpaeniformes Scorpaenidae Scorpaena bergii Y Fig. 40 and other BLZ image
Scorpaeniformes Scorpaenidae Scorpaena grandicornis Y Fig. 40 and other BLZ images
Scorpaeniformes Scorpaenidae Scorpaenodes caribbaeus Y Fig. 40
Scorpaeniformes Unknown Dendrochirus brachypterus Y Fig. 40
Scorpaeniformes Serranidae Diplectrum bivittatum Y Fig. 42 and other BLZ images
Scorpaeniformes Serranidae Pseudanthias cooperi Y Connell pers. comm.
Scorpaeniformes Serranidae Gonioplectrus hispanus Y Fig. 45
Scorpaeniformes Serranidae Hypoplectrus sp. Y Fig. 42 and other BLZ images
Scorpaeniformes Serranidae Mycteroperca bonaci Y Fig. 45 and other BLZ images
Scorpaeniformes Serranidae Pseudogramma gregoryi Y Fig. 43, 44, and other BLZ images
Scorpaeniformes Serranidae Pseudogramma polyacanthum Y Fig. 44
Scorpaeniformes Serranidae Rypticus sp. Y Fig. 43 and other BLZ images
Scorpaeniformes Serranidae Rypticus sp. Y Fig. 43
Scorpaeniformes Serranidae Rypticus sp. Y Fig. 43
Scorpaeniformes Serranidae Bathyanthias sp. Y Fig. 46
Scorpaeniformes Serranidae Liopropoma rubre Y Fig. 46
Scorpaeniformes Serranidae Serranus tigrinus Y BLZ 7730, 5352
Scorpaeniformes Serranidae Serranus baldwini Y Fig. 42 and other BLZ images
Stromateiformes Nomeidae Cubiceps? Unknown Y Fig. 48
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APPENDIX Continued
Order Family Genus Species Colour Image
Stromateiformes Stromateidae Centrolophus niger Y http://www.fisheggsandlarvae.com/
FIIA5%20Centrolophus%20
niger.htm
Tetraodontiformes Monacanthidae Monacanthus ciliatus Y Fig. 50 and other BLZ images
Tetraodontiformes Monacanthidae Aluterus schoepfi Y Fig. 50
Tetraodontiformes Ostraciidae Unknown Unknown Y Fig. 50 and other BLZ images
Tetraodontiformes Ostraciidae Lactophrys trigonus Y Fig. 50 and other BLZ images
Tetraodontiformes Ostraciidae Unknown Unknown Y Fig. 51
Tetraodontiformes Tetraodontidae Canthigaster rostrata Y Fig. 49 and other BLZ images
Tetraodontiformes Tetraodontidae Sphoeroides testudineus Y Fig. 49 and other BLZ images
Tetraodontiformes Tetraodontidae Sphoeroides spengleri Y Fig. 49 and other BLZ images
Tetraodontiformes Tetraodontidae Ranzania laevis Y Fig. 49
Trachiniformes Champsodontidae Champsodon capensis N http://www.fisheggsandlarvae.com/
LIIB1%20Champsodontidae.htm
Trachiniformes Chiasmodontidae Chiasmodon niger N http://www.fisheggsandlarvae.com/
LIIA8%20Chiasmodontidae.htm
Trachiniformes Uranoscopidae Unknown Unknown Y http://www.fisheggsandlarvae.com/
CMIA3%20Uranoscopidae.htm
BLZ, Belize. In cases in which it is followed by a four- or five-digit number, this represents the Smithsonian DNA number.
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