l__b

MVf

Number 488
19 April 2002

Contributions
IN Science

Larval Blennies erom the Galapagos and
Cocos Islands: Families Tripterygiidae,
Dactyloscopidae, and Chaenopsidae
(Perciformes, Blennioidei)

Guillermo A. Herrera and
Robert J. Lavenberg

OF Los Angeles County

Natural History Museum

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Larval Blennies erom the Galapagos and
Cocos Islands: Families Tripterygiidae,
Dactyloscopidae, and Chaenopsidae
(Percieormes, Blennioidei)

Guillermo A. Herrera^ and
Robert J. Lavenberg^

ABSTRACT. Postflexion larvae of ten eastern Pacific blennioid species from the Galapagos Islands and
Cocos Island are described. Three families are treated: Tripterygiidae, Dactyloscopidae, and Chaenopsidae.
The identifications were based on meristic data, and on comparisons of morphology between adults and
large larvae. The larvae from the Galapagos Islands are Lepidonectes corallicoia (Tripterygiidae); Dacty-
loscopus lacteus, Myxodagnus sagitta, Platygillellus rubellulus, and GHlellus semicinctus (Dactyloscopidae);
and Acanthemblemaria castroi and Chaenopsis schmitti (Chaenopsidae). The larvae from Cocos Island are
Dactyloscopus pectoralis fallax (Dactyloscopidae); and Acanthemblemaria atrata and Stathmonotus sp.
(Chaenopsidae).

All larvae described herein possess the following characters: a relatively long and slender body; a me-
lanophore anterior to the tip of the cleithral symphysis; ventral midline melanophores between pterygio-
phores of the anal fin; and a small head with a short and generally rounded snout (except in dactyloscopids
and in the chaenopsid Chaenopsis schmitti). Larvae of Lepidonectes corallicoia (Tripterygiidae) have a
specific arrangement of melanophores on the pterygiophores of the third dorsal fin and at the base of spines
of the second dorsal fin; pigment is found ventrally on the caudal peduncle and on the posterior margin
of the hypural plates. Larvae of the dactyloscopids {Dactyloscopus lacteus, Myxodagnus sagitta, Platygil-
lellus rubellulus, and Gillellus semicinctus) have large and pointed heads, short preanal length (30-35%
standard length (SL)), and prominent dorsal and anal fins. The dactyloscopid larvae can be identified by
specific combinations of characters: presence, number, and size of head melanophores; presence of pig-
mentation along dorsal margin of the body; presence of melanophores on the hypural borders; and fin
structure and meristics. Most chaenopsid larvae possess a large and elongate melanophore in the midline
along the basipterygium, a melanophore on the jaw angle, pigment dorsally to the anus, and a long preanal
length (40-50% SL). Characters described herein suggest that some larval attributes may be informative
for the elucidation of phylogenetic relationships within each family.

RESUMEN. Se describen las larvas en estado de postflexion de diez blenioideos del pacifico este de las
Islas Galapagos y de Isla del Coco. Las familias consideradas son Tripterygiidae, Dactyloscopidae y Chaen-
opsidae. Las identificaciones se basaron en informacion meristica y en comparaciones de morfologia de
adultos con larvas avanzadas. Las larvas de las islas Galapagos son Lepidonectes corallicoia (Tripterygi-
idae); Dactyloscopus lacteus, Myxodagnus sagitta, Platygillellus rubellulus y Gillellus semicinctus (Dacty-
loscopidae); y Acanthemblemaria castroi y Chaenopsis schmitti (Chaenopsidae). Las larvas de Isla del Coco
son Dactyloscopus pectoralis fallax (Dactyloscopidae), y Acanthemblemaria atrata y Stathmonotus sp.
(Chaenopsidae).

Caracteristicas presentes en las larvas de todas las especies son: cuerpo relativamente largo y elongado,
un melanoforo anterior de la sinfisis de los cleitros, una hilera ventral de melanoforos entre los pterigioforos
de la aleta anal, y una cabeza pequeha con un hocico corto y redondeado (excepto en los dactiloscopidos
y el chaenopsido Chaenopsis schmitti). Las larvas del blenido de tres aletas, Lepidonectes corallicoia, tienen
una configuracion espedfica de melanoforos en los pterigioforos de la tercera aleta dorsal y en la base de
algunas espinas de la segunda aleta dorsal; tienen ademas pigmento ventral en el pedunculo caudal y en
el margen posterior de las placas hipurales. Los dactiloscopidos {Dactyloscopus lacteus, Myxodagnus sag-
itta, Platygillellus rubellulus y Gillellus semicinctus) tienen larvas con una cabeza grande y puntiaguda,
una distancia preanal corta (30-35% longitud estandar (LE)) y aletas dorsal y anal prominentes. Las
especies pueden identificarse mediante combinaciones especificas de caracteres tales como presencia, nu-
mero y tamano de melanoforos cefalicos; presencia de pigmento dorsal sobre el cuerpo y en el borde
posterior de las placas hipurales; y caracteres meristicos de las aletas. Las mayoria de las larvas de los
chaenopsidos poseen un melanoforo grande y elongado ventral al basipterigio, un melanoforo en el angulo
mandibular, pigmento dorsal sobre el ano, y una mayor distancia preanal (40-50% LE). Los caracteres
larvales descritos en este trabajo sugieren que algunos atributos pueden ser informativos para la elucidacion
de relaciones filogeneticas dentro de cada familia.

Contributions in Science, Number 488, pp. 1-15
Natural History Museum of Los Angeles County, 2002

INTRODUCTION

The blennioids are oviparous fishes (except for
three viviparous genera) that lay their eggs attached
to a nest substrate, and exhibit male parental care.
The larvae are elongate and hatch with pigmented
eyes and sparse pigmentation (Matarese et ah,
1984; Stepien, 1986; Watson, 1996). Common fea-
tures are the presence of large melanophores dor-
sally on the swimbladder and gut and small mela-
nophores along the ventral margin of the trunk, be-
tween the pterygiophores of the long anal fin (Wat-
son, 1996; Cavalluzzi, 1997).

The sizes of larvae at hatching and flexion vary
depending on the species (Matarese et ah, 1984),
and the larvae do not show striking pigment pat-
terns or morphological adaptations for pelagic life.
Aside from some Blenniidae, the larvae of most
blennioid families are largely limited to coastal wa-
ters (Watson, 1996). Indeed, some larvae have been
observed to remain actively in coastal areas as they
school in groups around kelp and other algae (Ste-
pien et ah, 1997).

Few characters are shared by all blennoid larvae,
and most of them have been summarized by Ma-
tarese et al. (1984), Watson (1996), and Cavalluzzi
(1997). These are: body moderately elongate; pre-
anal distance between 30 and 50% of the standard
length (SL); large swimbladder; melanophores on
the ventral margin of the trunk (between pterygio-
phores of the anal fin); a melanophore on the tip
of the cleithral symphysis (Herrera and Lavenberg,
1999); and six branchiostegal rays.

Matarese et al. (1984) summarized the larval
characters of some Blennioidei, at a time when not
many descriptions were available. The best known
larvae are those of the families Blenniidae and Trip-
terygiidae, probably because of their wider distri-
butions and greater species numbers. The knowl-
edge of the larvae of the other four families, which
are predominantly from the New World, has in-
creased in recent years with the descriptions of lar-
vae from the Gulf of California (Brogan, 1992), the
California Current region (Watson, 1996), and the
western Atlantic (Cavalluzzi, 1997). The larvae of
species from the southeastern Pacific are less well
known (Balbontin and Perez, 1979; Perez, 1979).

Nineteen Blennioidei are known from the Gala-
pagos Islands, twelve of which are endemic (Grove
and Lavenberg, 1998). They belong to the families
Tripterygiidae (one species), Dactyloscopidae (five
species), Blenniidae (three species), Labrisomidae
(seven species), and Chaenopsidae (three species).
No species of the family Clinidae are known from
the archipelago. In this paper, field-collected speci-
mens mainly from the Galapagos Islands, obtained
from plankton samples collected during the Allan
Hancock expeditions, are described. Additionally,

1-2. Natural History Museum of Los Angeles County,
Research and Collections, 900 Exposition Boulevard, Los
Angeles, California 90007.

2 ■ Contributions in Science, Number 488

the larvae of three species from Cocos Island, Costa
Rica, are included. The material considers postflex-
ion larval stages and some juveniles. Larvae of the
following species are described: Tripterygiidae,
Lepidonectes corallicola (Kendall and Radcliffe,
1912); Dactyloscopidae, Dactyloscopus lacteus
(Myers and Wade, 1946), D. pectoralis fallax
(Dawson, 1975), Myxodagnus sagitta (Myers and
Wade, 1946), Platygillellus rubellulus (Kendall and
Radcliffe, 1912), and Gillellus semicinctus (Gilbert,
1890); and Chaenopsidae, Acanthemblemaria cas-
troi (Stephens et al., 1966), A. atrata (Hastings and
Robertson, 1999), Chaenopsis schmitti (Bohlke,
1957), and Stathmonotus sp. The larvae of D. pec-
toralis fallax, A. atrata, and Stathmonotus sp. were
collected in Cocos Island. Larvae of six species of
the family Labrisomidae (from the seven species cit-
ed for the archipelago) were described separately
(Herrera and Lavenberg, 1999).

MATERIALS AND METHODS

Most of the plankton samples were collected by the RW
Velero III, during the Allan Hancock Pacific expedition
cruises to the eastern Pacific and Galapagos Islands. Sam-
pling sites included Espanola, Santa Maria, Santa Cruz,
Isabela, Baltra, and Genovesa Islands (Eraser, 1943). A
single sample was collected in Chatham Bay, Cocos Island,
Costa Rica. Samples were normally collected at night (at
anchorage), using an electric light and dip nets. The larvae
are housed in the larval fish collection at the Natural His-
tory Museum of Los Angeles County (LACM). The spec-
imens were measured to the nearest 0.1 mm and illustrat-
ed with a camera lucida. The size range of the series il-
lustrated was determined by the available material. Al-
though the larval series are not complete because of the
sporadic nature of the sampling, they still contain valuable
ontogenetic information. Meristic data were obtained
from collection specimens and from the literature. Mea-
surements and definitions follow those of Leis and Rermis
(1983).

The larvae of the three species of Blenniidae from Gal-
apagos, Hypsoblennius brevipinnis (Gunther, 1861),
Ophioblennius steindachneri (Jordan and Evermann,
1898), and Plagiotremmus azalea (Jordan and Bollman,
1890), were also found together with the larvae described
here and in a previous work (Herrera and Lavenberg,
1999). These three species have a wide distributional
range in the eastern Pacific and their larvae have been
described from the California region (Watson, 1996). Lar-
vae of another blennioid, the dactyloscopid Dactylagnus
mundus (Gill, 1862), were not found in the samples.

RESULTS

The following descriptions include general charac-
terizations of morphology, dorsal- and anal-fin de-
velopment, and pigmentation. The larval series are
limited by the available material collected. For
those species represented by a few individuals the
descriptions are restricted to a usually narrow size
range. Meristic data and information about preanal
distance, a helpful diagnostic character of larval
blennioid families, are summarized in Table 1.

Herrera and Lavenberg: Larval blennies from Galapagos

Table 1. Selected meristic and morphologic data for species of Tripterygiidae, Dactyloscopidae, Blenniidae, and Chaenopsidae included in this study.

Pigmentation

Dorsal Dorsal Hyp. Ventral Preanal

Species Vertebrae DorsaP" AnaP' PT'" head trunk border gut distance (%) Reference

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Contributions in Science, Number 488

Herrera and Lavenberg: Larval blennies from Galapagos I 3

t Not found in samples but present in the Galapagos.

# Collected from Cocos Island.

References: (1) Bussing (1991); (2) Dawson (1976); (3) Brogan (1992); (4) Dawson (1975); (5) Dawson (1974); (6) Dawson (1977); (7) Watson (1996); (8) Rosenblatt and McCosker
(1988); (9) Stephens (1963); (10) Stephens et al. (1966); (11) Hastings and Springer (1994).

DESCRIPTION OF LARVAE
Family Tripterygiidae

A single triperygiid occurs in the Galapagos Islands,
the endemic Lepidonectes corallicola. In general,
the larvae of the Tripterygiidae are elongate and
slightly compressed; have small short and rounded
heads with no spination; have dorsal, anal, and
pectoral fins that develop early (shortly after flex-
ion); and have the spines of the first dorsal and on
the anal fins that form last (Leis and Rennis, 1983).

Lepidonectes corallicola
(Kendall and Radcliffe, 1912)

Figure 1

MATERIAL EXAMINED. From 2,281 speci-
mens (6.2-18.3 mm SL), five are illustrated; LACM
45614-6 (8.3, 10.0, 13.0, and 15.1 mm SL) and
LACM 45621-17 (17.2 mm SL).

MORPFIOLOGY. The body is moderately elon-
gate (preanal length 41-”43% body length (BL))
with a small head and rather small and rounded
snout. Flexion completed by 8.0 mm. Anal fin,
third dorsal fin, and pelvic fins are apparent by 8.0
mm. First and second dorsal fin apparent by 13.0
mm SL. Pores of the preopercular and circumor-
bital sensory canal systems are evident by 17.2
mm SL.

PIGMENTATION. In small larvae (6.2 mm) a
single dorsal melanophore is present on the nape;
this melanophore becomes embedded with growth.
Other cephalic pigmentation includes a pair of me-
lanophores on the hindbrain (Figs, la-b), which in-
creases to three pairs with the appearance of pairs
on the midbrain and forebrain (Fig. le). Gut pig-
ment consists of a single melanophore located ven-
troanteriorly (behind insertion of the pelvic fins).

By 8.3 mm SL three to five melanophores are
present on the ventral midline of the caudal pedun-
cle, and the posterior margins of the upper and
lower hypural plates each possess a single mela-
nophore. By 10.0 mm, dorsal midline melano-
phores are associated with the posterior portion of
the third dorsal fin, and the posterior margin of the
upper and lower hypural plates each possess two
melanophores. By 15.1 mm SL, the second dorsal
fin has a single melanophore at the base of spines
1, 3, and 7-11 (rarely associated with spines 2 and
6), the third dorsal fin has a single melanophore at
the base of each soft ray, the posterior margins of
the dorsal and ventral hypural plates have a wide
band of pigment, two melanophores are present on
the upper jaw, and small melanophores pepper the
membrane between rays of the caudal fin, mainly
in the lower half. By 17.2 mm SL, the juvenile stage

is observed. Additional pigment includes a lateral
band of small melanophores extending from the
pectoral-fin base to the end of the caudal peduncle.

REMARKS. Below 10.0 mm SL, the larvae are
similar in appearance to some labrisomids, espe-
cially those species lacking preopercular spines.
However, pigmentation, meristics, and morpho-
metries can differentiate them. In L. corallicola dor-
sal midline melanophores develop from posterior to
anterior (opposite for Malacoctenus zonogaster, La-
brisomidae), length to the origin of the third dorsal
fin is 60-63% SL in L. corallicola (68-71% SL in
M. zonogaster), and preanal distance is 41-43% SL
in L. corallicola (38-41% in M. zonogaster). After
10.0 mm SL, the presence of three distinct dorsal
fins in L. corallicola separates this species from the
labrisomids.

Family Dactyloscopidae

The family Dactyloscopidae is restricted to the New
World, in the Pacific and Atlantic Oceans. A few
descriptions of dactyloscopid larvae from the Gulf
of Galifornia (Brogan, 1992), Baja California (Wat-
son, 1996), and the Atlantic (Cavalluzzi 1997) are
available. From these, the larvae of the sand star-
gazers can be characterized as having an elongate
body, a short and compact gut, long and relatively
high dorsal and anal fins, jugular pelvic fins, 5 + 5
principal caudal rays, and a large head with point-
ed snout and no cirri. The preanal length is short,
compared to that of other families, and ranges from
28-35% SL (see Table 1). The species from the Gal-
apagos show specific arrangements of cephalic, hy-
pural, and trunk pigment. The posterodorsal region
of the swimbladder is pigmented, but it is difficult
to see the melanophores, especially in larger or
more robust larvae (such as Dactyloscopus).

Dactyloscopus lacteus
(Myers and Wade, 1946)

Figure 2

MATERIAL EXAMINED. Only two specimens
collected; LACM 45628-5 (7.7 mm SL) and LACM
45617-2 (9.2 mm SL).

MORPHOLOGY. The head is large and deep
with a pointed snout and a slight projection of the
tip of the lower jaw. Preanal length reaches 31-
34% SL. The larvae lack a notch on the ventral
margin of the caudal peduncle, a character that is
present in adults of many Dactyloscopus species
(Dawson, 1975). At 7.7 mm, the anal-fin rays are
longer and more prominent than the dorsal-fin
rays, indicating that the anal fin develops earlier. At
this size, only the rayed portion of the dorsal fin

Figure 1 Larvae of Lepidonectes corallicola: (a) 8.3 mm (b) 10.0 mm (c) 13.0 mm (d) 15.1 mm (LACM 45614-6)
(e) 17.2 mm (LACM 45621-17)

4 ■ Contributions in Science, Number 488

Herrera and Lavenberg: Larval blennies from Galapagos

Contributions in Science, Number 488

Herrera and Lavenberg: Larval blennies from Galapagos ■ 5

has developed (Fig. 2a). By 9.2 mm all dorsal-fin
spines have formed although they are not yet at
their maximum length and sensory pores have
formed on the preopercular border, the ventral
margin of the lower jaw and below the eye (Fig.
2b).

PIGMENTATION. In addition to the pigmenta-
tion anterior to the tip of the basipterygium, Dac-
tyloscopus lacteus has another small melanophore
on the basipterygium, close to the tip. The two me-
lanophores are very close and usually difficult to dis-
tinguish from each other when expanded. Two large
dendritic melanophores are present; one dorsally on
the head, and the other ventrally on the gut. No
melanophores occur dorsally along the trunk or on
the caudal peduncle. The posterior margins of the
hypural plates are pigmented. This differs from D.
pectoralis (Brogan, 1992), D. byersi (Watson, 1996),
and D. pectoralis fallax (Dawson, 1975).

Dactyloscopus pectoralis fallax
(Dawson, 1975)

Figure 3

MATERIAL EXAMINED. Description of a sin-
gle specimen, LACM 45658 (10.0 mm SL), from
three larvae (all near 10.0 mm SL) collected in Co-
cos Island, Costa Rica.

MORPHOLOGY. The body shape is similar to
that of Dactyloscopus lacteus. The preanal length
is 30-32% BL. At 10.0 mm SL head pores or dorsal
spines have not yet formed, which may differ from
D. lacteus, because the latter species had them by
9.2 mm (see Fig. 2b). The soft dorsal has formed
although it has not reached its maximum size, and
the dorsal spines are small projections.

PIGMENTATION. Dactyloscopus pectoralis fal-
lax has a single dendritic melanophore dorsally on
the head and another ventrally on the gut. The sole
difference between D. pectoralis and D. pectoralis

Figure 3 Larva of Dactyloscopus pectoralis fallax: 10.0 mm (LACM 45658-2)

6 ■ Contributions in Science, Number 488

Herrera and Lavenberg: Larval blennies from Galapagos

Figure 4 Larvae of Myxodagnus sagitta: (a) 7.3 mm (b) 8.8 mm (LACM 45625-7) (c) 11.6 mm (LACM 45623-10)
(d) 13.4 mm (LACM 45625-7)

fallax is the presence in the former of several me-
lanophores ventrally on the gut (Brogan, 1992) that
are not present in the latter. However, more speci-
mens of D. pectoralis fallax should be examined to
determine whether this pigment appears later in the
ontogeny.

Myxodagnus sagitta
(Myers and Wade, 1946)

Figure 4

MATERIAL EXAMINED. From a total eighty-
two specimens collected, in the size range of 7.0-
14.0 mm, four are used for description; LACM
45625-7 (7.3 and 8.8 mm SL), LACM 45623-10
(11.6 mm SL), and LACM 45625-7 (13.4 mm SL).

MORPHOLOGY. Among the larval dactylos-
copids studied, this species has the most slender and
elongate body, the smallest head, and the shortest

preanal distance (28-30% BL). The anal- and dor-
sal-fin rays have developed by 7.0 mm (Fig. 4a),
whereas the dorsal spines develop later (Fig. 4c)
The head is deep in small larvae, as in most dac-
tyloscopids, but becomes depressed with develop-
ment. The anterior projection of the lower jaw is
slight in small larvae, and increases with size. Pelvic
fins develop at an early stage (Fig, 4a), but they
remain smaller than in all other species of the fam-
ily.

PIGMENTATION. The cephalic pigmentation
pattern of Myxodagnus sagitta resembles that of
Dactyloscopus lacteus, with a large dendritic me-
lanophore above the head and below the gut. Fur-
thermore, a melanophore is present at the base of
each dorsal ray, except in the last one to two rays.
A small melanophore develops on the ventral mar-
gin of the caudal peduncle; these melanophores are
lacking in M. opercularis (Brogan, 1992; Watson,

Contributions in Science, Number 488

Herrera and Lavenberg: Larval blennies from Galapagos ■ 7

a

Figure 5 Larvae of Platygillellus rubellulus: (a) 8.9 mm (LACM 45614-11) (b) 11.1 mm (LACM 45621-10)

1996). In smaller larvae, the posterior edge of each
hypural plate has one small melanophore on the
border; these melanophores increase with develop-
ment forming bands of pigment. In some specimens
a small melanophore develops directly anterior to
the anus (Fig. 4d).

Platygillellus rubellulus
(Kendall and Radcliffe, 1912)

Figure 5

MATERIAL EXAMINED. Two specimens, out
of fifteen collected (size range 8.0-11.1 mm SL) are
described; LACM 45614-11 (8.9 mm SL) and
LACM 45621-10 (11.1 mm SL).

MORPHOLOGY. The body shapes of Platygil-
lellus rubellulus and Dactyloscopus lacteus larvae
are similar in having large heads and deep bodies
anteriorly, but differing in cephalic pigmentation.
The preanal length reaches 33-35% BL, which is
the longest among the studied dactyloscopids. At
11.1 mm, three pores occur above the eye and one
occurs on the margin of the lower jaw. The dorsal
fin is notched anteriorly, with the third spine short-
er than the adjacent ones, anterior and posterior. In
addition, the third and fourth spines are more sep-
arated from one another than other dorsal spines.
The last four dorsal spines decrease in size and first
four dorsal-fin rays increase progressively in size.
The relative sizes of spines and rays resembles a
doubly notched dorsal fin (Fig. 5b).

PIGMENTATION. Pigment pattern is the same in
both illustrated larvae. Two pairs of melanophores

8 ■ Contributions in Science, Number 488

occur dorsally on the head; a large pair above the
midbrain, and a small pair over the forebrain. A sin-
gle melanophore is present on the nape. Except for
a small single melanophore before the anus, no ven-
tral melanophores are found on the gut, A single and
small melanophore occurs on the ventral margin of
the caudal peduncle just behind the last anal-fin ray.
Another ventral midline melanophore is found im-
mediately anterior to the ventral hypural plate. This
melanophore varies in position from the space be-
tween fourth and fifth caudal rays (lower hypural),
on the fifth ray, in the space between the last two
procurrent caudal rays, and slightly in front of the
first procurrent caudal ray.

Gillellus semicinctus (Gilbert, 1890)

Figure 6

MATERIAL EXAMINED. Of the forty speci-
mens collected, three are used for description:
LACM 45614-19 (6.5, 10.0, and 12.4 mm SL).

MORPHOLOGY. The larvae are more elongate
than those of Dactyloscopus lacteus and the head
is proportionally smaller. The preanal length reach-
es 30-32% SL. Figure 6 shows the approximate
formational sequence of the dorsal- and anal-fin
spines and rays, and the subtle projection of the
lower jaw with development. At 6.5 mm flexion is
complete, the anal fin is beginning to develop, and
the dorsal fin has not formed (Fig. 6a). In the larg-
est larva, 12.4 mm, the first three dorsal spines,
which are separated from the following spines in
the form of a finlet, are present. Their lengths de-

Herrera and Lavenberg: Larval blennies from Galapagos

Figure 6 Larvae of Gillellus semicinctus-. (a) 6.5 mm, (b) 10.0 mm, and (c) 12.4 mm (LACM 45614-19)

crease progressively from the first to the third (Fig.
6c). Posteriorly, the dorsal spines and rays are ap-
proximately the same size, and the border of the
dorsal fin is straight.

PIGMENTATION. Pigment pattern does not
change in the size range studied. The larvae of Gil-
lellus semicinctus have two or three small melano-
phores ventral to the gut. One (or two in some lar-
vae) is always just posterior to the insertion of the
pelvic-fin rays, and the second is located more pos-
teriorly next to the anus. A ventral and internal
melanophore is also present behind the cleithra.
Pigment is absent dorsally on the head, which is an
uncommon feature in the family that has been also
reported in larvae of two other species of Gillellus
from the western Atlantic, G. jacksoni and G. ur-
anidea (Cavalluzzi, 1997). This lack of pigment
may be a reductive specialization characterizing the
genus.

Watson (1996) described larvae of a Gillellus
species, tentatively ascribed to semicinctus. Fiow-
ever, in his series the pretransformation specimens
(<13.3 mm) develop preopercular spines and a
ventral series of melanophores at the anal-fin base
that is not continuous. The larvae described here
do not have preopercular spines and a continuous

melanophore series occurs between pterygiophores.
The differences may be due to intraspecific varia-
tion or the larvae may belong to different species.

REMARKS ON DACTYLOSCOPID LARVAE

The relationships between the eight dactyloscopid
genera have been studied by Doyle (1998) using
morphological data. Heteristius is the basalmost
taxon, and Platygillellus is the next basalmost tax-
on. The remaining six genera are divided into two
clades; one of them includes Gillellus (together with
Leurochilus and Sindoscopus) and the other com-
prises Dactyloscopus, Dactylagnus, and Myxodag-
nus. Some of the larval characters presented here
are congruent with the relationships hypothesized
by Doyle (1998). A large, single, medial dendritic
melanophore occurs dorsally on the head and ven-
tral to the gut in the derived clade comprising Dac-
tylagnus, Myxodagnus, and Dactyloscopus. These
genera also share the same general body shape,
with a straight anterior profile of the dorsal fin and
a shorter preanal distance. In the other derived
clade, the larvae of Gillellus lack cephalic melano-
phores, which represents a derived condition. Un-
fortunately, larvae of the other two genera of the

Contributions in Science, Number 488

Herrera and Lavenberg: Larval blennies from Galapagos ■ 9

clade, Leurochilus and Sindoscopus, are not
known.

The larvae of the phylogenetically more primitive
Platygillellus show features that can be considered
plesiomorphic and include small melanophores in
pairs above the brain (vs. large and single), a longer
preanal distance (vs. shorter), a dorsal fin divided
into regions (vs. straight margin), and several small
ventral melanophores on the gut (vs. single and
large).

Family Chaenopsidae

The tube blennies are found in tropical and tem-
perate coastal waters of the New World, in both
the eastern Pacific and western Atlantic. Compared
to other blennies, chaenopsid larvae have a rela-
tively longer and straighter gut with a prominent
dorsally pigmented swimbladder. Most species have
a large and elongate melanophore midventral at the
basipterygium area. Other larval characters include
the presence of pigment dorsally to the anus, pelvic
fins that are inserted close to the base of the pec-
toral fin, rugae in the gut, ventral pigment on gut
and breast, no dorsal pigmentation, and a broad
and serrated premaxilla with a long ascending pro-
cess (Brogan, 1992).

Acanthemblemaria castroi
(Stephens et ah, 1966)

Figure 7

MATERIAL EXAMINED. From the 712 larvae
examined, in the size range of 5.3 mm notochord
length (NL) to 16 mm SL, five were used for de-
scriptions: LACM 45621-1 (5.3 mm NL), LACM
45644-1 (8.4 mm NL), LACM 45614-1 (12.4 mm
SL), LACM 45623-1 (14.0 mm SL), and LACM
45614-1 (16.0 mm SL).

MORPHOLOGY. Larvae are characterized by
elongate body, preanal length of 40-44% BL, small
and rounded head with a short snout, developing
bifid nasal cirri, and bifid orbital cirri. Flexion oc-
curs after 5.3 mm and is complete by 10.0 mm, at
which time the dorsal and anal-fin rays develop.
The dorsal spines begin to develop by 12.0 mm.

PIGMENTATION. In addition to the general
blennioid pattern, the larvae of Acanthemblemaria
castroi have melanophores that become embedded
during development located above the swimbladder
and anus; a melanophore on the mandibular angle
by 12.4 mm SL (Fig. 7c); a single melanophore on
the nape that becomes partially embedded with
growth (Figs. 7b-e); and three to four small mela-
nophores that are present during all stages on the
ventral midline of the caudal peduncle (Fig. 7a). No
pigment occurs ventrally on the gut, or on the pos-
terior border of the hypurals.

Acanthemblemaria atrata
(Hastings and Robertson, 1999)

Figure 8

MATERIAL EXAMINED. Five larvae from a to-
tal of 214 collected in Chatham Bay, Cocos Island,

10 ■ Contributions in Science, Number 488

Costa Rica, in the size range of 4.1-18.4 mm, are
illustrated: LACM 46011-1 (4.1 mm NL, 5.3 mm
NL, 6.9 mm SL, 11.8 mm SL), and LACM 45658-
1 (18.4 mm SL)

MORPHOLOGY. The larvae are similar in mor-
phology and meristics to those of Acanthemble-
maria castroi, but differ slightly in pigmentation.
The largest specimen (18.4 mm SL) has both single
nasal cirri and single orbital cirri; while larval pig-
mentation is still present, heavy cranial spination
appears along with sensory pores of the mandibu-
lar, nasal, occipital, preopercular, and orbital series.
The dorsal- and anal-fin formation is similar to that
of A. castroi.

PIGMENTATION. The larvae of Acanthemble-
maria atrata show the same pigment pattern as A.
castroi, but in addition exhibit a band of melano-
phores on the posterior margin of the hypural
plates. Further, one or two melanophores develop
ventrally on the anus (Figs. 8a-c); these melano-
phores disappear during development (Figs. 8d-e).
A more complete larval series is described for this
species, which allows following of the formation
pattern of the ventral melanophore on the basip-
terygium. Two elongate melanophores develop lat-
erally in early stages, converging anteriorly in an
inverted V-shaped pattern, with the wings of the V
fusing along the midventral margin (Brogan, 1992).

Chaenopsis schmitti (Boelke, 1957)
Figure 9

MATERIAL EXAMINED. Only two specimens
collected; LACM 45614-3 (8.0 and 14.6 mm SL).

MORPHOLOGY. Among ail the blennioids con-
sidered here, this species has the most slender and
elongate body. The preanal length ranges from 40-
44% BL. The snout is pointed and increases in
length with size. At 8.3 mm SL only the posterior
rays of the dorsal fin are developed, whereas anal-
fin rays are just forming. By 14.6 mm, the rayed
portion of both the dorsal and anal fin are formed,
and the dorsal-fin spines are appearing.

PIGMENTATION. The larvae of Chaenopsis
schmitti possess single melanophores at the man-
dibular angle, midlaterally on the preoperculum,
and on the hindbrain. No melanophores are present
on the caudal peduncle.

Stathmonotus sp.

Figure 10

MATERIAL EXAMINED. A single specimen
was collected in Chatham Bay, Cocos Island, Costa
Rica; LACM 45634-15 (8.3 mm SL). Although the
genus has not been reported there, the specimen can
be tentatively ascribed to the species Stathmonotus
culebrai based on meristic data. Also, 5. culebrai is
the only species of the genus from Pacific Central
America (Hastings and Springer, 1994).

MORPHOLOGY. The general shape of the larva
is similar to that of Stathmonotus stahli and S.
hemphilli (Brogan, 1992; Cavalluzzi, 1997), with a

Herrera and Lavenberg: Larval blennies from Galapagos

a

e

Figure 7 Larvae of Acanthemblemaria castroh (a) 5.3 mm (LACM 45621-1) (b) 8.4 mm (LACM 45644-1) (c) 12.4 mm
(LACM 45614-1) (d) 14.0 mm (LACM 45623-1) (e) 16.0 mm (LACM 45614-1)

Contributions in Science, Number 488

Herrera and Lavenberg: Larval blennies from Galapagos ■ 1 1

Figure 8 Larvae of Acanthemblemaria atrata: (a) 4.1 mm (b) 5.3 mm (c) 6.9 mm (LACM 46011-1) (d) 11.8 mm
(e) 18.4 mm (LACM 45658-1)

small and round head, and a prominent swimblad-
der. The preanal length reaches approximately 50%
BL (the longest among chaenopsids); the dorsal fin
originates behind the level of the pectoral-fin base

12 ■ Contributions in Science, Number 488

(vs. anterior to the pectorals in the other three
chaenopsids); and the number of pectoral-fin rays
is low (9-11 vs. 13-14 in other Chaenopsidae). In
adult Stathmonotus all dorsal-fin elements are

Herrera and Lavenberg: Larval blennies from Galapagos

spines (37-41). In the larva, the dorsal fin is com-
posed by two portions, with twenty-one spines and
seventeen soft rays, which indicates that the most
posterior spines of the adult dorsal fin are the result
of secondary ossification.

PIGMENTATION. The specimen appears to be
somewhat bleached and some pigment may be
missing. The larva has a pair of small ventral me-
lanophores immediately behind the insertion of the
pelvic-fin rays, a single ventral melanophore on the
gut, and a melanophore over the anus. No mela-
nophores appear on the trunk, caudal peduncle, or
the hypural margins. The midventral melanophore
at the basipterygium is not visible, but it may be
embedded, as in Stathmonotus sinuscalifornici at a
similar size (Brogan, 1992). The pigmentation of
this specimen lacks the expanded and stellate me-
lanophores along the ventral margin of the trunk
and gut of S. sinuscalifornici (Brogan, 1992) or the
lines of melanophores near the dorsal (of S. hem-
philli) and ventral margin of the trunk of 5. stahli
(Cavalluzzi, 1997).

REMARKS ON CHAENOPSID SPECIES

An interesting character of postflexion larvae of all
species is the presence of an elongate pigment patch
midventral to the basipterygium, observed and il-
lustrated by Brogan (1992) from larvae of the

Chaenopsinae (Hastings and Springer, 1994), such
as Emblemaria, Chaenopsis, and Coralliozetus. The
pigment patch results from the convergence at the
ventral midline of two more lateral and usually
elongate melanophores. Although the ventral me-
lanophore is not visible in large larvae of Neoclinus
and Stathmonotus, it does form in preflexion larvae
and becomes embedded during the development
(Brogan, 1992).

DISCUSSION

The Blennioidei exhibit reproductive features that
are generally associated with low dispersal capa-
bilities such as low fecundity, demersal spawning,
some form of parental care, robust larvae, and
short larval period. The larvae of most families, ex-
cluding Blenniidae, have few specializations (Wat-
son, 1996), and the number of larval characters
available to study relationships is low. Cavalluzzi
(1997) described the larvae of two species of Dac-
tyloscopidae and two of Chaenopsidae and dis-
cussed the use of life history characters for eluci-
dating relationships within the suborder Blennioid-
ei. He found no synapomorphies based on larval
morphology that support the monophyly of the
suborder. The chaenopsid and dactyloscopid larvae
described herein, plus the description of the larvae
of six labrisomid species (Herrera and Lavenberg,

Contributions in Science, Number 488

Herrera and Lavenberg: Larval blennies from Galapagos ■ 13

1999), did not help to resolve further synapomor-
phies. However, variation among larvae may help
to elucidate relationships within and among fami-
lies.

Among dactyloscopids, the larvae of the phylo-
genetically primitive genus Platygillellus possess
plesiomorphic characters such as paired cephalic
melanophores and a discontinuous anterior profile
of the dorsal fin. Larvae of the more derived genera
Myxodagnus, Dactyloscopus, and Dactylagnus,
which constitute a clade (Doyle, 1998), possess a
single large melanophore on the head and ventrally
on the gut, and a continuous and straight anterior
dorsal-fin profile. The larvae of Gillellus, another
derived dactyloscopid unrelated to the previous
clade, lack cephalic melanophores and the anterior
margin of the dorsal fin is discontinuous. Unfortu-
nately, the larvae of the other two genera of the
clade {Sindoscopus and Leurochilus) have not been
described. Although the absence of cephalic mela-
nophores may be considered a derived feature with-
in Dactyloscopidae (e.g., also observed in the prim-
itive tripterygiids), it is not possible to establish
whether the loss is derived from a plesiomorphic
condition (small paired melanophores) or a rela-
tively more derived condition (large and single me-
lanophores). Nevertheless, these characters are con-
sistent with the dactyloscopid relationships of
Doyle (1998).

Chaenopsid have larval features that are consis-
tent with phylogenetic relationships obtained by
Hastings and Springer (1994) from adult morphol-
ogy. The phylogenetically more primitive genus
Neoclinus (previously considered a labrisomid), has
larvae that develop several melanophores dorsally
on the head and has a comparatively short preanal
distance (<39% SL). More derived chaenopsids
lack dorsal head pigmentation and have longer pre-
anal distances (40-57%). Also, the phylogenetically
primitive Neoclinus has a well-defined pigment pat-
tern dorsally on the trunk, with melanophores at
the base of the dorsal-fin spines and in between the
pterygiophores of the soft rays (Watson, 1996); this
feature is absent in all remaining and more derived
chaenopsid larvae (Brogan, 1992; Watson, 1996;
Cavalluzzi, 1997), although it is common in labri-
somids and in the basal tripterygiids.

The larvae of all Chaenopsid genera examined so
far possess an elongate melanophore midventrally
on the basipterygium that results from the fusion
of two melanophores that develop laterally and
converge on the ventral midline (Brogan, 1992). An
elongate melanophore on the basipterygium is also
observed in the labrisomid Starksiini Starksia gal-
apagensis (Herrera and Lavenberg, 1999). Based on
molecular evidence, Stepien et al. (1997) considered
the labrisomids a paraphyletic group and hypoth-
esized that the tribe Starksiini was more closely re-
lated to Chaenopsidae than were other Labrisomi-
dae. Although the larval evidence is suggestive, fur-
ther work is needed to confirm the affinities be-
tween Chaenopsidae and Starksiini.

14 ■ Contributions in Science, Number 488

Among blennioid larvae, other melanophore
characters do not seem to be informative about re-
lationships above genus level, although it remains
to be determined whether they are useful at lower
taxonomic level (i.e., between species). For exam-
ple, a band of melanophores is present in the pos-
terior margin of the hypural plates in the species
Hypsoblennius jenkinsi, H. sordidus, Dactylosco-
pus pectoralis fallax, Acanthemblemaria atrata,
and Malacoctenus zonogaster, but it is absent in
their respective congenerics H. brevipinnis, D. lac-
teus, A. castroi, and M. tetranemus (Balbontin and
Perez, 1979; Stevens and Moser, 1982; Herrera and
Lavenberg, 1999).

In conclusion, the phylogenetic significance of
larval pigment characters still remains to be deter-
mined. However, as suggested by Cavalluzzi
(1997), it seems, that pigment characters may be
more important for low taxonomic level (i.e., inter-
or intrageneric) than for higher taxonomic level
comparisons.

ACKNOWLEDGMENTS

We acknowledge and thank the University of Southern
California and the Natural History Museum of Los An-
geles County for continued financial support. We also
wish to acknowledge the National Science Foundation for
their support of larval fish curation at LACM, which
made collections available for study (NSF DEB 8814791).
We thank Richard Feeney (LACM) who assisted us during
our studies of fish larvae from the Galapagos, and C.
Thacker (LACM) for reading the manuscript. We thank
Carol Stepien (Cleveland State University) and Martin
Cavalluzzi (Oregon State University) for their valuable
comments and suggestions on the manuscript.

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Received 29 October 1999; accepted 3 October 2000.

Contributions in Science, Number 488

Herrera and Lavenberg: Larval blennies from Galapagos ■ 15

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