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Western Atlantic Species of the

Genus Holocentrus

Loren P. Woods

Curator, Division of Fishes

Among the fishes collected in the Gulf of Mexico by the United
States Fish and Wildlife Service M/V Oregon during the years
1950-53, there were 17 specimens of Holocentrus taken from the
bottom in otter trawls in depths from 29 to 60 fathoms. These
could not be determined by using the key of Jordan and Evermann
(1896, p. 848) or that of Parr (1930, pp. 34-35) or by comparing
them with identified specimens in the Chicago Natural History
Museum collection. It was therefore necessary to review the litera-
ture, especially of species that have long been in synonymy, and
re-examine all available specimens. These studies have resulted in
the reduction of two species to synonymy (osculus and sanctipauli) ,
recognition of one previously synonymized species (rufus), and
description of one new species (bullisi).

METHODS

Counts and measurements have been made according to standard
practice as outlined by Hubbs and Lagler (1947, pp. 8-15), except
when noted otherwise. All rudiments of gill rakers are included in
the counts given. The length of the lower jaw was measured from
the posterior edge of the angular to the anterior tip of the dentary,
that of the upper jaw, with mouth closed and bones in normal
position, from the anterior tip of the premaxillary to the posterior
margin of the maxillary. The head length was measured from the
anterior tip of the premaxillary to the tip of the longest opercular
spine, that of the preopercular spine from its tip to the dorsal angle
with the anterior margin of the preopercle, that is, to the margin of
the scales at the lower posterior corner of the cheek.

92 FIELDIANA: ZOOLOGY, VOLUME 37

DISCUSSION

Dr. Edward M. Nelson, Assistant Professor in Anatomy, Stritch
School of Medicine, Loyola University, simultaneously has been
studying the structure and relations of the air bladder to the auditory
bullae in the entire family Holocentridae (Nelson, this volume, pp.
121-137) . His observations have been referred to below, but without
details or figures. It is considered not entirely inconsistent with
Nelson's data to recognize the four lines of development that can
be distinguished within the genus Holocentrus as subgenera as
follows, although on the basis of internal structures Nelson has
distinguished only two lines, Holocentrus and Adioryx.

Flammeo Jordan and Evermann

Flammeo Jordan and Evermann, 1898, Bull. U. S. Nat. Mus., no. 47,
p. 2871 (type, Holocentrus marianus).

The species referred to Flammeo have the dorsal fin spines X,
I, the last spine of the dorsal usually longer than the tenth, separated
from it although attached at its base by a membrane and more or
less closely joined with the first dorsal soft ray, the body elongate,
and the lower jaw extending beyond the upper. This subgenus
contains the species marianus, sammara, laevis, opercularis, and
scythrops.

Sargocentron Fowler

Sargocentron Fowler, 1904, Proc. Acad. Nat. Sci. Phila., 55: 235 (type, Holo-
centrus leo).

Sargocentron includes species of large size, the body deep and
squarish, the spiny dorsal fin high. The species grouped here are
leo Cuvier and Valenciennes (=spinifer), tieroides, spinosissimus,
praslin, and ensifer.

Adioryx Starks

Adioryx Starks, 1908, Science, 28: 614 (type, Holocentrus suborbitalis).

Adioryx includes the intermediate residue of species, some
grading toward Flammeo as do xantherythrus, coruscus, and bullisi,
and others forming a group more typical for Adioryx, such as subor-
bitalis, vexillarius, punctatissimus, lacteoguttatus, microstomus, Here,
and hastatus.

Holocentrus Scopoli

Holocenthrus Scopoli, 1777, Int. Hist. Nat., p. 499 (misprint for Holocentrus
after Gronow's Holocentrus maxilla).

WOODS: SPECIES OF HOLOCENTRUS 93

Holocentrus contains only the species ascensionis and rufus. It
is characterized externally by having the dorsal spines more nearly
of equal length, produced anterior dorsal and anal soft rays, very
elongate caudal lobes, and the upper lobe much longer than the
lower; internally by the tubular structure of the auditory bulla with
its opening directed posteriorly and the swim bladder with a mem-
branous area on either side on its anterior face (Nelson, this volume,
pp. 121-137).

Three of these groups are found in the Atlantic: subgenera
Holocentrus, Adioryx, and Flammeo. The three in the Pacific are
Adioryx, Flammeo, and Sargocentron.

Mr. Norman Wilimovsky has suggested that a number of generic
names take priority over Adioryx Starks, 1908, type Holocentrus
suborbitalis Gill. I have re-examined the descriptions, wherein
numerous names have been conferred on larval and post-larval
Holocentridae and others, and find as follows:

Rhynchichthys Valenciennes (Cuvier and Valenciennes, 1831,
p. 504), type R. pelamidis Valenciennes, from the Indian Ocean,
dorsal X, I, 12. The type is obviously a larval fish and we have at
hand larval specimens of Myripristis occidentalis with preopercular
spines almost as well developed as those figured for R. pelamidis
(Cuvier and Valenciennes, 1831, pi. 208) . Further, it is characteristic
of Myripristis to have the dorsal X, I. The count, anal IV, 12,
given by Valenciennes could not refer to any known species of
Holocentrus, but the number 12 is characteristic of Myripristis.
Rhynchichthys pelamidis Valenciennes is therefore referred to syn-
onymy of the genus Myripristis.

Rhinoberyx Gill (1862, p. 237), type Rhynchichthys brachyrhynchus
Bleeker, from Amboina. This species is figured in Bleeker's Atlas
(9, pi. 357, fig. 1). It has the appearance of a small Myripristis.
It has no preopercular spine, the dorsal is X, I, 13 or 14 and the
anal is IV, 11 or 12, the anal spines are more like those of Myripristis
in size and proportion, and the number of anal rays is applicable
only to species of Myripristis. Rhinoberyx chryseus Cope (1870, p.
464, fig. 2) is a synonym of Myripristis jacobus.

Rhamphoberyx Gill (1863, p. 87), type R. poecilopus Gill, from
Cape San Lucas, Baja California, Mexico. Specimens on which
this genus and species were based were "1M to 2% inches long."
From the description it is evident that the specimens were larval
or were just transforming. The spine at the angle of the preoper-
culum is "not essentially enlarged, but simply forms the angle at

94 FIELDIANA: ZOOLOGY, VOLUME 37

the preoperculum." Counts given are dorsal X, I, 14; anal IV, 11.
No species of Holocentrus known to me has 11 anal rays. Gilbert
and Starks (1904, p. 65) have placed poecilopus in the genus Myri-
pristis, and Hildebrand (Meek and Hildebrand, 1923, p. 296), who
had access to Gill's types, apparently agreed with Gilbert and Starks,
although Meek and Hildebrand had no specimens of poecilopus from
Panama. The other species placed in this genus by Gill (1863, p.
88), Rhamphoberyx leucopus Gill, has dorsal X, I, 14; anal IV, 12.
No preopercular spine is mentioned in the original description.
Hildebrand (Meek and Hildebrand, 1923, p. 295) has placed leucopus
in the synonymy of Myripristis occidentalis.

Neomyripristis Castelnau (1873, p. 99), type A7. amaenus Cas-
telnau, from Torres Strait, Australia, is described as being very near
to Myripristis but with different teeth; preopercular without a
spine; dorsal X, I, 15; anal IV, 14; opercle with one long spine,
below which are two short spines. All of these characters are those
of Myripristis rather than Holocentrus.

The original description of the genus Neoniphon Castelnau
(1875, p. 4), type A7. armatus, from Cape York, Australia, is stated
to have counts as follows: dorsal XI, I, 12; anal III, 9; pectoral 15;
pel vies 7; scales over 38-40. The color, with a large black spot on
membranes between the first three dorsal spines, and numerous
round spots on cheek and body, forming irregular lengthwise lines;
the proportions; and the counts, except for the number of anal
spines, along with the mention of lower jaw being larger than upper,
all agree closely with Holocentrus sammara Forskal. The chief
difficulty is that the size is stated to be nearly 17 inches (470 mm.)
and the largest sammara of which I can find a record is 267 mm.
(about 10% inches). Neoniphon hasta De Vis (1885, p. 537), from
Queensland (dorsal XI, I, 12; anal III, 7; lateral line 40), has two
long spines at the angle of the preopercle; the black blotch on the
dorsal membranes seems to be identical with that of armatus.
Both species have been recognized by McCulloch (1929, pp. 134-
135). It appears likely that both of these species are synonyms
of H. sammara, a species ranging from the Red Sea to the Tuamotus
and abundant in the Sunda area and New Guinea. H. sammara is
not reported from Australia although it should certainly occur along
the northern shore and the northern portion of the eastern shore of
Australia. At any rate it is fairly certain that both species described
under Neoniphon belong to the Flammeo group and if so the name
Neoniphon has priority over Flammeo. However, it appears to be

WOODS: SPECIES OF HOLOCENTRUS 95

more sensible to continue to use the name Flammeo Jordan and
Evermann (1898, p. 2871), based as it is on a well-known species
widely used in the literature, rather than to replace it with a name
based on a species whose status is uncertain.

Harpage De Vis (1884, p. 447), type H. rosea De Vis, from the
South Sea Islands. Dorsal XII, I, 14; anal III, 11, lateral line scales
42^, "preopercle scarcely spiniferous, with short bifid spine at
angle." The short preopercular spine, the structure of the dorsal
and the number of the anal rays, and the premaxillary overhanging
the mandible relate this species to, or identify it with, Holotrachys,
as suggested by Jordan and Seale (1906, p. 223). H. lima has dorsal

XII, 15; anal IV, 11; lateral line scales 43^; and bears a short
spine at the angle of the preopercle. The short first anal spine may
have been overlooked by De Vis. Nelson has found H. lima to be
a specialized form of the Myripristis branch.

Beanea Steindachner (1902, p. 337), type B. trivittata Stein-
dachner, from the Red Sea, may not even belong to the family

Holocentridae. "D. VII-I, 9; A. IV, 8; P^ I, 5; 1. lat. 25^." All
members of this family of which I can find a description have X, I,
XI, or XII dorsal spines and 29 or more lateral line scales; further,
all have more than 5 soft rays in the pelvic fin. The type of Beanea
was a small specimen (larval?) 30.6 mm. in length. It is described
as having the body form of Myripristis, and the "border of the
preopercle and lower horizontal part of the forward arm of the same
bone entirely sharply toothed." No mention is made of a spine
at the angle of the preopercle that is characteristic of Holocentrus.

Sargocentron Fowler (1904, p. 235) was proposed as a subgenus
for Holocentrus leo Cuvier, a species that I believe to be a synonym
of H. spinifer Forskal. The name Sargocentron is used in this paper
in the sense I believe it was proposed, namely, as a subgenus to
include large, rectangular-bodied species with "margin of preopercle
coarsely serrated." (See p. 92 for list of such species.)

A recently proposed genus Beloholocentrus Fowler (1944, p. 100),
type B. atractus Fowler from Courtown Key (12° 24' N., 81° 28' W.),
Caribbean Sea, appears to be based on young that have been de-
scribed by Bean (1906, p. 42, fig. 2) as Holocentrus meeki (= Holo-
centrus rufus) and the "Rhynchichthys" stage of H. ascensionis.

Various names proposed for subgenera by Whitley (1933, pp.
68-69), namely, Cephalofarer (type H. siccifer), Faremusca (type
H. punctatissimus) , and Kutaflammeo (type H. tahiticum) do not

FIELDIANA: ZOOLOGY, VOLUME 37

enter into the question of priority of names used herein. H. sicci-
fer is a synonym of H. vexillarius, which along with punctatissimus
fits very well into the subgenus Adioryx.

The remaining genera of this family, Myripristis, Corniger, Holo-
trachys, Plectrypops, Ostichthys, and the fossil genera are considered
well enough known or far enough removed from this group to have
no bearing on the divisions within the genus Holocentrus and require
no further discussion.

Fig. 14. Heads of Holocentrus coruscus (a), H. bullisi (b), and H. vexillarius
(c), to illustrate differences in spination of snout and supraorbital regions.

KEY TO WESTERN ATLANTIC SPECIES OF HOLOCENTRUS

la. Dorsal spines X, I (subgenus Flammeo Jordan and Evermann); scales in
lateral line with tubes 46 or 47; dorsal X, 1, 12 or 13; anal IV, 8 or 9; pectoral
ii, 12; head pointed; mouth large, lower jaw protruding beyond upper;
opercular spines short, upper slightly larger; third anal spine exceedingly
long, reaching well beyond base of caudal fin. . .Holocentrus marianus Cuvier

WOODS: SPECIES OF HOLOCENTRUS 97

16. Dorsal spines XI.

2a. Scales in lateral line with tubes 46 to 57; gill rakers on lower limb of

first arch 14 to 17 subgenus Holocentrus Scopoli

3a. Length of upper jaw 6.60 to 7.40, of lower jaw 5.10 to 5.85, both in
standard length; scales in lateral line with tubes 46 to 51.

Holocentrus ascensionis Osbeck
36. Length of upper jaw 7.70 to 9.25, of lower jaw 6.10 to 6.82, both in
standard length; scales in lateral line with tubes 50 to 57.

Holocentrus rufus Walbaum
26. Scales in lateral line with tubes 45 or fewer; gill rakers on lower limb

of first arch 9 to 12 subgenus Adioryx Starks

4a. Pectoral axil black (or dusky in very large specimens); dorsal rays
13 or 14; anal rays 9; snout smooth, no spinules between narial
opening and premaxillary groove, antero-dorsal orbital margin

flanged and denticulate (fig. 14) Holocentrus vexillarius Poey

46. Pectoral axil always pale; dorsal rays 11 or 12; anal rays 7 or 8;

snout with patch of spinules between narial opening and premaxillary

groove; antero-orbital margin not flanged, entire.

5a. Pectoral rays ii, 10 or ii, 11; a large black oval spot on spiny

dorsal membrane distally between first 3 or 4 spines, rest of

membranes dusky in very small specimens; snout spinules

prominent, a spine extending into posterior narial opening (fig. 14) .

Holocentrus coruscus Poey
56. Pectoral rays ii, 12; no large oval spot on first three or four
spiny dorsal membranes; only a small spot on distal portion of
membrane in specimens 58 to 76 mm.; snout spinules smaller,
limited to rim of premaxillary groove (fig. 14) with snout merely
rugose between this groove and nares. Holocentrus bullisi sp. nov.

Table 1.— FIN RAY COUNTS IN WESTERN ATLANTIC SPECIES
OF THE GENUS HOLOCENTRUS

Dorsal Rays

Species X, I XI 11 12 13 14 15 16

ascensionis 24

rufus 19

vexillarius 28 . . . . 23

bullisi 17 1 16

coruscus 22 3 19

marianus .... 5 . . 1 4

Table 2.— FIN RAY COUNTS IN WESTERN ATLANTIC SPECIES
OF THE GENUS HOLOCENTRUS

Species IV

ascensionis 24

rufus 19

vexillarius 28

bullisi 17

coruscus 22

marianus 5

* Both pectorals of each fish were counted and tabulated separately.

Anal Rays

7 8 9 10

11
1?

Pectoral Rays*
13 14 15 16
.. .. 4 41

1 17

.... 7 25

1 26 1

17 . .

.. 2 35 2

.32 2 . .
37

2 20

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WOODS: SPECIES OF HOLOCENTRUS 99

Subgenus FLAMMEO

Holocentrus marianus Cuvier. Figure 15.

Holocentrum marianum Cuvier, 1829, in Cuvier and Valenciennes, Hist. Nat.

Poissons, 3: 164 (219) — Martinique.
Holocentrum rostratum Poey, 1860, Memorias . . . , 2: 157 — Cuba (name

preoccupied).

Counts and description are based on 7 specimens, including
cotypes of Poey's rostratum, 74 to 124 mm., from the Bahamas,

Fig. 15. Holocentrus marianus Cuvier, AMNH no. 19076; 103 mm. Haiti.

Cuba, Haiti, Glover Reef, Isle of Pines and Jamaica. These speci-
mens belong to the Museum of Comparative Zoology, Natural
History Museum, Stanford University, the American Museum of
Natural History, and Bingham Oceanographic Collection.

Description. — Dorsal fin rays X, I, 13 (12 in one specimen);
anal rays IV, 9 (8 in one specimen); pectoral rays ii, 12; pelvics I,
7; scales in lateral line bearing tubes 46 or 47, 3^ rows between
lateral line and origin of dorsal fin, 8 between lateral line and anus;
gill rakers 6 or 7 on upper limb, one at angle, 12 on lower limb.

Depth of body 2.93-3.23, length of head (to tip of longest oper-
cular spine) 2.59-2.86, length of snout 9.46-10.6, diameter of eye
6.47-8.15, width of interorbital 11.5-14.8, length of upper jaw 5.88-
6.72, length of lower jaw 4.64-4.86, depth of caudal peduncle 11.4-
12.4, length of caudal peduncle 5.43-6.21, length of fourth dorsal
spine 6.07-10.1, length of third anal spine 3.36-4.40, length of pec-

100 FIELDIANA: ZOOLOGY, VOLUME 37

toral fin 3.60-4.97, length of pelvic fin 4.47-5.53, length of upper
caudal lobe 3.49-3.73, length of lower caudal lobe 3.53-3.99, all in
standard length.

Third anal spine very long, its tip reaching from base of caudal
rays to fork of caudal; lower jaw projecting beyond upper; maxillary
reaching from anterior margin of pupil to middle of pupil; last dorsal
spine nearer to first dorsal soft ray than to penultimate spine;
margin of premaxillary groove smooth; opercular spines short and
heavy, of equal length or with upper spine slightly enlarged; profile
nearly straight.

Color completely faded in all available specimens.

Remarks. — Mrs. M. M. Dick, Museum of Comparative Zoology,
has very kindly re-examined Poey's types of H. rostratum, MCZ
10969, two specimens, 142 and 150 mm. standard length. These
specimens are apparently correctly referred to the synonymy of
marianus as was done by Howell-Rivero (1938, p. 183). Their fin
ray and scale counts are in agreement with this species; the only
differences noted are the slightly shorter maxillary (2.52 and 2.54)
— but the maxillary reaches to the center of the pupil — and the
shorter third anal spine (2.45 to 2.48). This latter difference is
probably because of a difference in method of measuring. The
other characters of serrae and spines on head bones, especially oper-
cular spines, and projecting lower jaw are characteristic of marianus.

Dr. Edward M. Nelson has found no differences in the structure
of the swim bladder and auditory bulla in species referred to Flammeo
(F. sammara, F. laevis, F. scythrops, F. opercularis) and Adioryx
suborbitalis or Holocentrus coruscus, vexillarius and numerous Pacific
species referred to the genus Holocentrus. It is therefore proposed
to recognize Flammeo as a subgenus of Holocentrus on the basis of
its peculiar dorsal (X, I) and protruding lower jaw.

Subgenus HOLOCENTRUS

Holocentrus ascensionis Osbeck. Figure 16.

Perca ascensionis Osbeck, 1771, Iter Chinensis, p. 388 — Ascension Island.
Holocentrum longipinne Cuvier (part), 1829, in Cuvier and Valenciennes,

Hist. Nat. Poissons, 3: 138 (185)— Brazil; Martinique; Porto Rico; St.

Thomas; San Domingo; Havana.
Holocentrum productum Poey, 1868, Synopsis, p. 300 — Cuba.
Holocentrum sancti pauli Gunther, 1880, Rep. Voy. Challenger, Zool., 1, pt.

6, Rept. Shore Fishes, p. 4, pi. 1, fig. A — St. Paul's Rocks.

101

102 FIELDIANA: ZOOLOGY, VOLUME 37

Measurements are based on 25 specimens from Bermuda, St.
Croix, and Brazil, 58 to 191 mm.

Description. — Dorsal fin rays XI, 15 (rarely 14 or 16) ; anal rays
IV, 10; pectoral rays ii, 13 to 15 (usually ii, 14); pelvics I, 7; scales
in lateral line bearing tubes 46 to 51, 3 3/2 between lateral line and
origin of dorsal, 8 between lateral line and anal opening; gill rakers
on first arch 8 on upper limb, one at angle, 14 or 15 on lower limb,
total 23 or 24.

Depth of body 2.69-3.13, length of head 2.80-3.17, length of
snout 11.8-15.3, diameter of eye 6.35-9.45, width of interorbital
14.0-19.2, length of upper jaw 6.62-7.37, length of lower jaw 5.11-
5.84, length of preopercular spine 16.0-18.7, depth of caudal pe-
duncle 11.5-13.9, length of caudal peduncle 5.1-6.31, length of
fourth dorsal spine 6.06-7.2, length of third anal spine 5.23-6.52,
length of pectoral fin 4.32-4.93, length of pelvic fin 3.01-3.60, length
of upper caudal lobe 2.60-3.93, length of lower caudal lobe 2.97-4.03,
all in standard length.

Body deeper than in H. rufus except in small specimens; mouth
large, upper jaw reaching posteriorly to below center of pupil or
beyond; lower jaw generally not extending anterior to upper in
small specimens, in large specimens even with or slightly protruding;
anterior soft rays of dorsal and anal greatly elongate; upper caudal
lobe considerably longer than lower.

Color. — In alcohol, essentially as in H. rufus (q.v.) except spiny
dorsal membranes lacking distinct white spot distally, distal portion
instead with a wedge pale or dusky colored (green to greenish
brown in living specimens).

Remarks. — This species may usually be distinguished from rufus
by the lower number of tube-bearing scales in the lateral line. Since
ascensionis occasionally has 51 scales, it is necessary to rely on
measurements. H. ascensionis always has a larger mouth (measure-
ments of both upper and lower jaws) and in all except specimens
70 mm. or less a much shorter preopercular spine. When specimens
of the same size of both ascensionis and rufus are measured and
compared, there is no difficulty in distinguishing all specimens (see
tables and comparative measurements under H. rufus) . The species
are very similar in characters but are not geographic subspecies since
ascensionis ranges from Brazil, Panama, and the West Indies to
Bermuda, while H. rufus is also found in all of these localities except
Brazil. I have not been able to examine specimens of ascensionis
from West Africa or Ascension Island. Osbeck's original description

WOODS: SPECIES OF HOLOCENTRUS 103

did not give the number of lateral line scales (at least in the 1771,
English edition, available to me). Giinther's description of H.
sanctipauli records 48 lateral line scales and this is in accordance
with my findings on specimens from Brazil.

We have one small specimen from St. Croix that has the preoper-
cular spine forked on one side as in sanctipauli; on the other side
the spine is broken.

One larval specimen of ascensionis (61 mm.) from St. Croix was
examined; it is similar to "meeki" in most respects but is deeper-
bodied and has 48 and 49 tubed lateral line scales. We have also
one specimen from Puerto Rico completely transformed at 52
mm. This specimen has 47 scales and in proportions closely re-
sembles larger ascensionis.

H. productum Poey was based on a specimen 68 mm. long. The
counts given in the original description (dorsal XI, 15; anal IV, 10)
agree with ascensionis.

I have examined a specimen, USNM 37428, 56 mm. standard
length, one of Poey's specimens and labeled as the type of productum.
There are, however, a number of discrepancies between this specimen
and the original description quoted above. The National Museum
specimen has: Dorsal rays XI, 11; anal IV, 7; pectoral 13; scales in
lateral line 42. All of these counts fit coruscus. There is a spot
distally on the membranes between the first and fourth dorsal
spines. Since the snout was broken the measurements of the head
and body which were made are considered not reliable and are not
given. At the time this type was examined (1947) no attention
was paid to the presence or absence of spinules on the snout present
in coruscus.

Since there are such differences in the counts given by Poey
and the specimen labeled type in the National Museum it appears
possible that there is an error in labeling and that USNM 37428 is
not Poey's type of productum. At any rate it is believed USNM
37428 is certainly a specimen of coruscus.

Studies by Nelson show that the structure of the swim bladder
and its relation to the auditory bullae as well as the structure of
the bullae themselves are similar in ascensionis, rufus, and nominal
meeki (= rufus). The form and relation of these two structures are
different from all the other species of Holocentrus that were exam-
ined by Nelson (23 species, but not including the nominal osculus
and sanctipauli).

104 FIELDIANA: ZOOLOGY, VOLUME 37

Holocentrus rufus Walbaum. Figure 17.

Perca rufa Walbaum, 1792, Art. Pise, p. 351 — Bahamas (after Perca marina

rubra of Catesby).
Sciaena rubra Schneider, 1801, in Bloch and Schneider, Syst. Ichthy., p. 82

(after Catesby) (ref. copied).
Holocentrum osculum Poey, 1860, Memorias, 2: 156 — Cuba; Regan, 1904,

Ann. Mag. Nat. Hist., (7), 13: 259-260— St. Thomas, Virgin Islands.
Holocentrum perlatum Poey, 1860, Memorias, 2: 157 — Cuba.
Holocentrus meeki Bean, 1906, Proc. Biol. Soc. Wash., 19: 31 — St. David's

Island, Bermuda.

Measurements are based upon 18 specimens, 70 to 180 mm.,
from Bermuda, Florida, Panama, Colombia, St. Croix, and Glover
Reef, British Honduras. The counts include many additional
specimens.

Description. — Dorsal fin rays XI, 14-16; anal rays IV, 10 (rarely
9 or 11); pectoral rays ii, 13-15 (usually ii, 14); pelvics I, 7; scales
in lateral line bearing tubes 50-57, 33^ between lateral line and origin
of dorsal, 7 or 8 between lateral line and anal opening; gill rakers
on first gill arch 8 or 9 on upper limb, one at angle, 15-17 on lower
limb, total 24-26.

Depth of body 3.05-3.61, length of head 3.05-3.43, length of
snout 13.6-16.6, diameter of eye 7.12-9.86, width of interorbital
13.4-17.7, length of upper jaw 7.74-9.25, length of lower jaw 6.12-
6.82, length of preopercular spine 11.7-17.1, depth of caudal peduncle
13.4-15.3; length of caudal peduncle 4.68-5.5, length of fourth
dorsal spine 5.63-7.01, length of third anal spine 4.85-6.10, length of
pectoral fin 4.28-5.35, length of pelvic fin 2.99-4.10, length of upper
caudal lobe 2.47-3.42, length of lower caudal lobe 3.14-3.85, all in
standard length.

Body slender, compressed; mouth small, upper jaw reaching
posteriorly from anterior margin of pupil to below center of pupil,
rarely just beyond; lower jaw not extending to anterior margin of
upper but slightly included; snout smooth, opercular spines two,
upper spine longer and stronger, preopercular spine relatively long
and slender; third anal spine about three-fourths of the length of
first anal rays; anterior soft dorsal rays very much elongated,
reaching beyond end of body when depressed; caudal fin deeply
forked, its lobes long and slender, upper lobe longer than lower.

Color. — In alcohol: lips light orange; premaxillary processes
dusky; upper margin of exposed portion of maxillary white, a band
of white diagonally extending across cheek to base of preopercular

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106 FIELDIANA: ZOOLOGY, VOLUME 37

spine, cheeks rosy; back reddish orange, lower sides yellow. In
formalin-preserved specimens: three distinct brown lines along back
above lateral line; spiny dorsal membrane pink basally, dusky dis-
tally except a distinct white spot on anterior distal edge of each
membrane, dorsal spines brown; soft dorsal, anal, and caudal pale
orange to pink; first three anal spines white, fourth dusky; pectoral
light orange, the axil white; pelvics pale orange, the spine and outer
edge of first ray white, inner edge of first ray dusky. Fresh-frozen
specimen: tips of premaxillaries and mandibles dark red; upper
margin of maxillary white continuous with white diagonal bar across
cheek to angle of preopercle; head and body pale brick red, above
lateral line 3 narrow greenish brown lengthwise lines following
margins of scales narrowing and disappearing posteriorly, centers of
scales pale pink, lower sides paler than upper, belly white, breast
streaked with faint pink, base of pectoral dark red; spiny dorsal
membranes each with small white spot distally just behind spine,
narrow black line along margin of membrane; membranes pink
anteriorly, dark green posteriorly; soft dorsal, caudal and anal fins
red or faint orange red; first three anal spines white; pelvic
spines white, outer rays red, inner rays paler; pectoral rays pale
pink.

Remarks. — This species has been recognized by Jordan and
Evermann (1896, p. 849) as a subspecies of ascensionis and by
Jordan, Evermann, and Clark (1930, p. 236) as a separate species
with the note, "The northern form probably varying into H . ascen-
sionis, which ranges from the Bahamas to Brazil." Specimens in
the Chicago Natural History Museum collection labeled ascensionis
from Bermuda, Bahamas, Puerto Rico, St. Croix, Florida, Mis-
terioso Bank (Lat. 18° 30' N.; Long. 83° 30' W.), Glover Reef,
Panama, Colombia, Venezuela, and Brazil (149 specimens in all)
can readily be separated into two groups based on the number of
tube-bearing scales in the lateral line. Geographically it is apparent
that ascensionis is commonly found from Bermuda throughout the
West Indies to Santos, Brazil, while rufus appears to be rare in
Central America and northern South America and does not occur
in Brazil. Only one rufus was found among 17 specimens of ascen-
sionis from Panama. In the West Indies and Florida the two species
appear to be equally abundant. In Bermuda ascensionis has been
found on the offshore banks and in the passes exclusively; both are
found together along shore near the edge of the reef, while rufus
alone was collected on the lagoon reefs. A lot of 21 specimens from

WOODS: SPECIES OF HOLOCENTRUS 107

Bermuda (CNHM no. 42867) was separated into two groups with
the following differences noted:

rufus ascensionis

8 specimens 13 specimens

Standard length 87.70-168 81.40-191

Head 3.14- 3.32 2.80- 3.17

Depth of body 3.08- 3.35 2.80- 3.09

Length of maxillary 8.08- 8.67 6.62- 7.15

Preopercular spine 13.40- 14.50 16.00- 18.70

All of the above specimens were well preserved and carefully mea-
sured. Specimens of each sex in each group were measured and
compared with the above.

Specimens smaller than 80 mm. that were separated on the basis
of lateral line scales could be distinguished by all characters except
size of eye and length of preopercular spine. In measuring additional
specimens from other parts of the range the proportions occasionally
overlapped but individuals aberrant in one or another of the sepa-
rating characters could always be distinguished. In all cases the
lengths of the upper and lower jaws were different and after the
two groups had been distinguished nearly all individuals could be
sorted by inspection before scales were counted or measurements
made.

I have examined Catesby's figure of Perca marina rubra (1731,
2, pi. 3, fig. 2) on which Walbaum based his Perca rufa. This
figure is not carefully drawn and was made from a specimen with
its mouth open and the jaws protruded forward, the head raised,
and the opercular bones separated; the preopercular spine is not
clearly delineated, and the number of scales and fin rays is not
accurate. Nevertheless, Catesby's figure unmistakably represents
the same kind of fish herein called rufus. The text accompanying
the plate mentions color but otherwise states no characters of any
use in defining the species.

A specimen from St. Thomas, described by Regan (1904, p.
259) with the remark, "I have no doubt it is the little known H.
osculum of Poey," has been received on exchange from the British
Museum. This specimen agrees in all respects with the counts,
measurements, and coloration given here for rufus. The lateral-line
scales number 52, the maxillary extends to below the center of the
pupil, and the spiny dorsal membranes bear white spots distally.

Mrs. M. M. Dick has examined the type of H. perlatum Poey,
MCZ no. 10938, said by Howell-Rivero (1938, p. 183) to be a syno-

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WOODS: SPECIES OF HOLOCENTRUS 109

nym of H. osculus Poey. The measurements, especially of the eye
and the maxillary, and the length of the latter in relation to the
pupil, as well as the color of the dorsal membrane, all support my
opinion that perlatum is a synonym of rufus.

Observations by Beebe (1936, p. 198) on specimens of H. meeki,
kept alive in aquaria on Nonsuch Island, Bermuda, are summarized
as follows: "Little by little, in the course of several weeks they
changed from Holocentrus meeki to the common squirrel fish, Holo-
centrus ascensionis. . . . There was no doubt about the fact that
meeki is the immature stage of ascensionis." I have again counted
the lateral-line scales on the types of H. meeki and find 52 and 53
bearing tubes. Other specimens collected by Mowbray in Bermuda
(CNHM no. 49077), certainly the same as the types of meeki, have
51 and 56 tubed scales. This number agrees with rufus rather
than with ascensionis. Also, the number of fin rays and gill rakers
of meeki are within the range of rufus. There are three specimens
from Bermuda (CNHM no. 48723), 70.5 to 71.7 mm. standard
length, that have completely transformed and resemble young rufus
in shape of head and size of eye. The preopercular spine is short,
however, proportionately equal to ascensionis of the same size. The
depth, head, size of eye, and length of maxillary are different from
those of ascensionis.

Types of meeki are 69 and 75 mm. standard length. Three
specimens from Brazil that measure 58, 65, 67 mm. and one from
Puerto Rico that is 52 mm. are completely transformed and resemble
young adults in shape of head and size of eye. The mouth, however,
is small (7.2, 7.31, 7.24) but still larger than the mouth size of
rufus (71.2, 70.7, and 70.0 mm. standard length; length of upper
jaw 9.25, 8.64, 8.76 in standard length repectively) . (The specimen
of ascensionis from St. Croix that resembles meeki in its stage of
development is 61 mm.) I am of the opinion that ascensionis trans-
forms at a smaller size than rufus.

Subgenus ADIORYX

Holocentrus vexillarius Poey

Holocentrum vexillarium Poey, 1860, Memorias, 2: 158 — Cuba.
Holocentrus brachypterus Poey, 1865, Repertorio, 1: 184 — Cuba.
Holocentrum sicciferum Cope, 1871, Trans. Amer. Phil. Soc, 14: 465 — New

Providence, Bahamas.
Holocentrum riparum Poey, 1875, Enumeratio, p. 37 — Cuba.

110 FIELDIANA: ZOOLOGY, VOLUME 37

Six specimens measured, 53.7 to 142 mm., from Bermuda,
Glover Reef, and Panama.

Description. — Dorsal fin rays XI, 13 (rarely 14) ; anal rays IV, 9
(rarely 8 or 10) ; pectoral rays ii, 13 (rarely ii, 12 or ii, 14) ; scales in
lateral line bearing tubes 40-44, 33^ rows between lateral line and
origin of dorsal, 8 between lateral line and anal opening; gill rakers
on first gill arch 6 or 7 on upper limb, one at angle, 11 (rarely 10)
on lower limb, total 17-19.

Depth of body 2.54-2.79, length of head 2.56-3.03, length of
snout 12.5-13.4, diameter of eye 6.17-9.05, width of interorbital
9.37-10.8, length of upper jaw 7.56-8.82, length of lower jaw 5.65-
6.90, length of preopercular spine 10.5-18.0, depth of caudal peduncle
9.84-11.2, length of caudal peduncle 4.93-6.55, length of fourth
dorsal spine 5.37-6.17, length of third anal spine 4.18-5.08, length
of pectoral fin 3.86-4.51, length of pelvic fin 3.97-4.68, length of
upper caudal lobe 3.55-3.91, length of lower caudal lobe 3.99-4.18,
all in standard length.

Body deep, compressed, upper profile angular; mouth small,
premaxillary extending posteriorly to below a point midway between
anterior margin and center of pupil; lower jaw fitting into upper,
not entering profile; no spinules on snout; dorso-anterior margin of
orbit broadened into flange bearing heavy obtuse spines in specimens
of 80 mm. and greater (a specimen 54 mm. had orbit flanged but
spines did not reach margin, which was entire); opercular spines
two, these nearly equal in size (usually lower is longer and sometimes
stronger) ; preopercular spine rugose, extending beyond lower margin
of subopercle; third anal spine equal to or slightly shorter than
anterior anal soft rays, when pressed parallel with lengthwise axis
of body its tip not reaching end of body.

Color. — In alcohol: three broad dark brown lengthwise lines
above lateral line, running between scale rows, centers of scales
pale; head, breast and lower sides with brown puncticulations; sides
below lateral line with faint broad brown lines narrowing and fading
posteriorly (not evident in largest specimens); axil of pectoral jet
black in young, dusky in large individuals; spiny dorsal membranes
dusky distally with vertical brown or black line covering most of
membrane immediately anterior to each spine; membrane between
third and fourth anal spines sometimes faintly dusky distally; rest
of fins pale.

Remarks. — The specimen figured by Fowler (1904, p. 234, fig.
5), stated to be the type of H. sicciferum Cope, is certainly a small

WOODS: SPECIES OF HOLOCENTRUS 111

specimen of H. vexillarius. The counts of dorsal, anal, and pectoral
fin rays given by both Cope and Fowler are within the range of this
species (see table) and this combination of counts does not fit any
other Atlantic species. H. sicciferum is reported by Cope to have 45
lateral-line scales. Three of our specimens have 44 tubed scales
and the one following the last tubed scale is not much smaller and
is perforated so the apparent difference of sicciferum is attributed to
method of counting. In our series of specimens from Bermuda
ranging in size from 38 to 142 mm. there are some whose dorsal
fins are colored as shown in Fowler's figure of the type of sicciferum.
The large eye, small mouth and abruptly down-curved snout as
well as the relatively deep body are all characteristic of this species
at all sizes.

The figure and description in Metzelaar (1919, p. 43) of H.
coruscus are based on small vexillarius. His counts — dorsal XI,
133^-14; anal IV, 9 — and his figure showing the black pectoral axil
could refer to no other species than vexillarius.

One large specimen from Bermuda (142 mm., approx. 175 total
length) agrees with the details of proportions and color pattern
given by Poey for brachypterus, the type specimen of which he stated
to be 195 mm. in length. The only discrepancy is in the number of
soft anal rays, stated to be 8 by Poey. I have found 9 soft rays in
most specimens of vexillarius counted, but one had 8 and another
10 rays.

H. riparum was based on a specimen of 50 mm. If this is total
length (including caudal fin) it is almost certainly a larval stage.
The counts of riparum agree with vexillarius as do the markings on
the dorsal fin membranes described by Poey. Howell-Rivero (1938,
p. 183) examined the type specimens of riparum and listed them as
synonyms of vexillarius.

Holocentrus hastatus Valenciennes, ranging from Cape Verde Is-
lands to Angola, according to Fowler (1936, pp. 549-551) appears
to be a close relative of H. vexillarius. It differs in having dorsal
rays XI, 15, anal IV, 10, scales 48 to 51, and in having the dark
color on spiny dorsal membranes limited to the anterior portion of
the dorsal. No specimens of this species were examined by me;
the above notes are from Fowler (loc. cit.) who also regarded hastatus
as a close relative of sicciferum Cope, here considered a synonym
of vexillarius.

112 FIELDIANA: ZOOLOGY, VOLUME 37

Holocentrus coruscus Poey

Holocentrum coruscum Poey, 1860, Memorias, 2: 158 — Cuba.

Holocentrus tortugae Jordan and Thompson, 1905, Bull. U. S. Bur. Fish., 24:

236, fig. 1 (published in May, 1905)— Garden Key, Tortugas.
Holocentrus puncticulatus Barbour, 1905, Bull. Mus. Comp. Zool., 46: 117,

pi. 2 (published in September, 1905) — Flatts Inlet, Bermuda.

Eight specimens measured, 53 to 100 mm., from Bermuda and
the Bahamas.

Description. — Dorsal fin rays XI, 12 (rarely 11); anal rays IV,
8 (rarely 7); pectoral rays ii, 11; pelvics I, 7; scales in lateral line
bearing tubes 41-45, 3}^ rows between lateral line and origin of
dorsal fin, 6 or 7 between lateral line and anal opening; gill rakers
on first arch 5-7 on upper limb, one at angle, 10 on lower limb,
total 16-18.

Depth of body 3.09-3.70, length of head 2.70-3.08, length of
snout 13.4-15.2, diameter of eye 6.13-8.14, width of interorbital
9.58-13.0, length of upper jaw 6.46-7.68, length of lower jaw 5.53-
5.97, length of preopercular spine 17.4-20.0, depth of caudal peduncle
12.0-13.7, length of caudal peduncle 5.03-5.72, length of fourth
dorsal spine 5.77-7.32, length of third anal spine 4.17-5.41, length
of pectoral fin 4.50-5.41, length of pelvic fin 4.52-5.07, length of
upper caudal lobe 4.54-4.93, of lower lobe 4.67-4.93, all in standard
length.

Body elongate, slender, but sides more rounded than other
Atlantic species of Holocentrus; mouth large, upper jaw reaching
posteriorly to below center of pupil; lower jaw very slightly shorter
than upper, not fitting inside upper; a patch of spinules on snout
between nares and premaxillary groove (fig. 14), one or two spines
extending laterally over posterior narial opening; antero-dorsal mar-
gin of orbit entire; opercular spines two, the upper longer and
stronger; preopercular margin serrate, a strong, keeled spine at its
angle extending beyond spines of ventral portion of subopercle;
third anal spine longer than anterior anal rays, when laid back
falling short of end of body; caudal forked, lobes rounded.

Color-In alcohol: reddish brown with three narrow, white,
lengthwise lines following centers of scale rows above lateral line,
lines broader below lateral line; cheek white with oblique brown
band from orbit to postero- ventral angle, breast silvery; spiny
dorsal fin with prominent black oval spot distally on membranes
between first three spines in large specimens, sometimes extending

WOODS: SPECIES OF HOLOCENTRUS 113

to fourth spine, rest of dorsal membranes faintly dusky distally;
soft dorsal and other fins pale; pectoral axil pale.

Remarks. — Although Parr (1930, p. 36) has stated that Poey's
original description of this species is inadequate for proper identi-
fication, I believe Poey's combination of counts (dorsal XI, 12;
anal IV, 8; pectoral 13) must refer either to coruscus or, possibly, to
bullisi, which has no large black spot on the dorsal. From the
counts given by Jordan and Bollman (1889, p. 550) and Beebe and
Tee Van (1928, p. 81, fig.) it appears that all these authors had
specimens of H. vexillarius with 13 dorsal rays and 9 anal rays.
Two of Beebe and Tee Van's counts give 8 and 8% anal rays.
H. tortugae Jordan and Thompson and H. puncticulatus Barbour
are also certainly the same species. I have compared numerous
Bermuda and Bahama specimens with the original descriptions of
these two species and can find no significant differences. Both
descriptions state 13 dorsal rays and both figures show 12. Both
figures also show the characteristic spinules on the snout, but these
are not mentioned in the descriptions. Longley and Hildebrand
(1941, pp. 54-55) have discussed these spinules and other contrasting
differences between coruscus and vexillarius.

Holocentrus bullisi sp. nov. Figure 18.

Type. — United States National Museum no. 164980, Oregon
Station 222, Campeche Banks, Lat. 22° 32' N., Long. 88° 47' W.,
29 fathoms, January 11, 1951. Collected by Harvey Bullis. Stand-
ard length 116.5 mm.

Paratypes. — CNHM no. 48730, Bermuda, Argus Bank, April 6,
1908. Collected by L. L. Mowbray. Standard length 79 mm.

CNHM no. 46193, Oregon Station 35, 75 miles northwest of
Dry Tortugas, Florida, Lat. 25° 35' N., Long. 83° 46' W., 60 fathoms,
June 26, 1950. Collected by H. H. Shoemaker. Standard length
129 mm.

CNHM no. 59868, Oregon Station 646, Campeche Banks, Lat.
22° 27' N., Long. 89° 59' W., 37 fathoms, October 5, 1952. Collected
by J. B. Siebenaler. One specimen, standard length 71 mm.

CNHM no. 59869, Oregon Station 725, Campeche Banks, Lat.
22° 15' N., Long. 88° 55' W., 25 fathoms, December 13, 1952.
Collected by Loren Woods and Robert Inger. Three specimens,
standard length, 58.8 to 76.5 mm.

CNHM no. 59870, Oregon Station 727, off Cape San Bias,
Florida, Lat. 28° 44' N., Long. 85° 01' W., 30 fathoms, December

114 FIELDIANA: ZOOLOGY, VOLUME 37

16, 1952. Collected by Loren Woods and Robert Inger. Nine
specimens, standard length 105 to 120 mm.

CNHM no. 59871, Oregon Station 892, off Cape San Bias,
Florida, Lat. 28° 55' N., Long. 87° 07' W., 29 fathoms, March 7,
1953. One specimen, 116 mm.

Description.1 — Dorsal fin rays XI, 12 (XI, usually 12, rarely
11) ; anal rays IV, 8 (IV, 8) ; pectoral rays ii, 12 (ii, 12, rarely ii, 13) ;

S*W

Fig. 18. Holocentrus bullisi sp. nov. Type, USNM no. 164980; 116.5 mm.
Campeche Banks, Yucatan, Mexico.

scales in lateral line bearing tubes 39/40 (39 to 43), 33^ (3j/0 rows
between lateral line and origin of dorsal, 7 (7) between lateral line
and anal opening; gill rakers on first arch 6 (5 or 6) on upper limb,
one at angle, 10 (9 or 10) on lower limb, total 16 (16).

Depth of body 3.20 (2.56-3.19), length of head 2.88 (2.66-3.08),
length of snout 12.0 (10.9-12.2), diameter of eye 7.67 (5.98-7.82),
width of interorbital 13.4 (9.8-14.1), length of upper jaw 7.33 (6.13-
7.18), length of lower jaw 5.84 (4.82-5.67), length of preopercular
spine 15.3 (10.3-17.1), depth of caudal peduncle 11.9 (9.8-13.2),
length of caudal peduncle 5.62 (5.12-6.16), length of fourth dorsal
spine 6.26 (5.16-6.56), length of third anal spine 4.70 (4.04-4.68),
length of pectoral fin 4.42 (4.20-4.73), length of pelvic fin 4.27
(4.06-4.56), length of upper caudal lobe 3.97 (3.82-4.34), length of
lower caudal lobe 4.80 (3.85-4.80), all in standard length.

1 The counts and proportions of the type are given first, with extremes noted
on paratypes in parentheses.

WOODS: SPECIES OF HOLOCENTRUS 115

Body slender, compressed; mouth moderately small, maxillary
extending posteriorly to midway between anterior margin of eye
and middle of pupil in all specimens; lower jaw fitting into upper
in the type, upper and lower jaws equal in paratypes; pre- and sub-
orbitals serrate; a patch of spinules on snout between nares and pre-
maxillary groove (these broken in type but scars remain), no spine
directed over nares as in H. coruscus; no spines on dorso-anterior
border of orbit; interorbital concave with numerous pores; opercular
spines two, the upper longer and stronger; preopercle serrate on both
margins, preopercular spine strong with keel basally, reaching just
beyond longest spine of subopercle in type, longer in paratypes;
dorsal fin almost completely divided, third and fourth dorsal spines
subequal and slightly shorter than soft rays; third anal spine longer
than anterior anal rays, when pressed down reaching to base of
caudal rays; caudal forked, rays broken on type so exact shape of
lobes cannot be determined, lobes angular or rounded, upper slightly
longer in paratypes; 5 more or less free spines on dorsal base of
upper caudal rays, 4 on ventral.

Color. — In alcohol: pale brownish yellow; cheeks pearly white;
back and sides above lateral line darker with three very distinct
narrow white lines from head, fading posteriorly, following centers
of scales; each scale with brown dot on both sides of this line giving
appearance of very narrow brown border to white line, very faint
in largest paratype; below lateral line four broader white lengthwise
lines, narrowing and fading posteriorly, brown border of these lines
scarcely evident on type; fins all pale except for a minute dusky
spot distally immediately behind first dorsal spine on type, on
smallest paratype traces of dusky vertical streaks on each membrane;
pectoral axil pale.

Remarks. — This species appears to be most closely related to
Holocentrus coruscus Poey, from which it differs as follows:

bullisi coruscus

Pectoral rays 14 12 or 13

Lateral line scales 40 and 41 42 to 45

Narial spine Absent Present

Color of dorsal See above Oval black spot

distally between
first 3 spines

From ascensionis, rufus, and vexillarius, bullisi differs in number
of fin rays, scales, and color; from marianus it differs in number of
lateral line scales, structure of dorsal (XI in bullisi, X, I in marianus)
(see table for counts).

116 FIELDIANA: ZOOLOGY, VOLUME 37

Named for Harvey R. Bullis, Jr., of the United States Fish and
Wildlife Service.

ACKNOWLEDGMENTS

I wish especially to thank Mrs. M. M. Dick of the Museum of
Comparative Zoology for having examined the types of two of
Poey's species, Holocentrus perlatum and H. rostratum. Miss M.
Storey, Natural History Museum of Stanford University, Dr. D.
Merriman, Bingham Oceanographic Laboratory, Mr. J. T. Nichols,
American Museum of Natural History, and Mr. N. Marshall, British
Museum (Natural History) have lent specimens for study. Dr.
Robert R. Miller, University of Michigan, has made numerous help-
ful editorial suggestions.

REFERENCES

Bean, T. H.

1906. Catalogue of the fishes of Bermuda, with notes on a collection made
in 1905 for the Field Museum. Field Columbian Mus., Zool. Ser., 7: 21-89,
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Beebe, William

1936. Food of the Bermuda and West Indian tunas of the genera Parathunnus
and Neothunmis. Zoologica, 21, (15), pp. 195-205, pis. 1-3.

Beebe, William and Tee Van, John

1928. The fishes of Port-au-Prince Bay, Haiti. Zoologica, 10, (1), pp. 1-279,
map and text figs, (not numbered).

Bleeker, Pieter

1862-77. Atlas icthyologfque des Indes Orientales Neerlandaises. 9 vols.,
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Castelnau, F. L.

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Soc. Victoria, 2:37-158.
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Catesby, Mark

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Cope, E. D.

1870. Contribution to the ichthyology of the Lesser Antilles. Proc. Amer.
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Cuvier, G. and Valenciennes, A.

1831. Histoire naturelle des poissons. 7, 242 pp., 23 pis. (Strasbourg de luxe
edition).

De Vis, C. W.

1884. Fishes from South Sea Islands. Proc. Linn. Soc. N. S. Wales, 8 : 445-457.

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Fowler, H. W.

1904. New, little known and typical berycoid fishes. Proc. Acad. Nat. Sci.

Phila., 56:222-238.
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(pts. 1 and 2), 1493 pp.
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Gilbert, C. H. and Starks, E. C.

1904. The fishes of Panama Bay. Mem. Calif. Acad. Sci., 4: 1-304, pis. 1-33.

117

118 FIELDIANA: ZOOLOGY, VOLUME 37

Gill, Theodore

1862. Remarks on the relations of the genera and other groups of Cuban
fishes. Proc. Acad. Nat. Sci. Phila., 14: 235-242.

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WOODS: SPECIES OF HOLOCENTRUS 119

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