PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 
111(4):936-941. 1998. 
A new species of the genus Bellator (Pisces: Triglidae), with 
comments on the trigloids of the Galapagos Islands 
William J. Richards and John E. McCosker 
(W.J.R.) NOAA Fisheries, Southeast Fisheries Science Center, 75 Virginia Beach Drive, 
Miami, Florida 33149, U.S.A.; 
(J.E.McC.) California Academy of Sciences, Golden Gate Park, 
San Francisco, California 94118, U.S.A. 
Abstract. — A new, endemic species, Bellator farrago, is described from the 
Galapagos Islands. It differs from its congeners by a combination of characters 
including absence of a supplemental preopercular spine, short first dorsal fin 
spine, produced rostral spines, and lack of scales on the breast and interpelvic 
area. The Galapagos trigloid fauna comprises: B. farrago, Prionotus miles, Pr. 
stephanophrys, and Peristedion crustosum. 
The fishes of the New World genus Bel- 
lator were revised by Miller & Richards 
(1991); they treated four Atlantic species 
and three eastern Pacific species. Recent 
collecting in and around the Galapagos by 
the submersible Johnson Sea-Link revealed 
the presence of an undescribed species. The 
submersible captured one specimen and an- 
other was found in the California Academy 
of Sciences Fish Collection. Prior to this 
study only one Bellator was known from 
the Galapagos based on Gruchy (1970), 
who added Prionotus loxias {= Bellator lox- 
ias) to the Galapagos fauna. A re-exami- 
nation of the Gruchy specimens reveals that 
they also are this undescribed species and 
not B. loxias. This new species is diagnosed 
and described, and details of its habitat are 
provided based on observations made from 
the submersible. The submersible also cap- 
tured three specimens of Peristedion crus- 
tosum Garman which is a new record of this 
species from the Galapagos. 
Methods. — Counts and measurements 
follow Miller & Richards (1991). 
Bellator farrago, new species 
(Figs. 1-3, Tables 1-3) 
[non] Prionotus loxias: Gruchy 1970:526 
(misidentification). Miller & Richards 
1991:646 (distribution, following Gruchy 
1970). Bussing 1995:1646 (distribution). 
Material examined. — Holotype: CAS 
54562 (103.9 mm SL), southeastern Pacific, 
Ecuador, Galapagos Islands, Isla Santa 
Cruz, Academy Bay, RA^ Te Vega Field 
No. TV24-VIII-68. 24 Aug 1968. Para- 
types: CAS 86564 (96.2 mm SL), south- 
eastern Pacific Ocean, Ecuador, Galapagos 
Islands, Isla Genovesa (Tower). 0°21.8'S, 
89°58.2'W, JSL dive 3974, 462 m, coll. J. 
E. McCosker et al., 24 Nov 1995. NMC 69- 
78, 2 (75.5-112.4 mm SL), southeastern 
Pacific Ocean, Ecuador, Galapagos Islands, 
Isla Isabella, mouth of Tagus Cove, coll. 
Ban- et al. 8 Mar 1968. 
Diagnosis. — A species of Bellator with 
opercular spine short, breast and interpelvic 
area lacking scales, belly scaled, first dorsal 
spine shorter than second, supplementary 
preopercular spine weak or absent, anal 
rays 11, pored lateral line scales 50-52. 
Description. — Morphometric data are in 
Table 1, meristic data in Table 2, and com- 
parison with other eastern Pacific Bellator 
in Table 3. Head moderate with produced 
rostrum with small spines on distal edge; 
small spines on lateral edge of first, second, 
and third infraorbital bones; no nasal spine. 
VOLUME 111, NUMBER 4 
937 
Fig. 1. Lateral view and dorsal view of head of Bellator farrago, new species, paratype CAS 86564, 96 mm 
SL. Illustration by Molly Brown. 
although all head bones with very small 
spines giving rough texture; prominent 
spines on anterior and posterior edge of or- 
bit; prominent nuchal, parietal, opercular 
and preopercular spines, but supplemental 
preopercular spine absent in three speci- 
mens and very weakly developed in one. 
Cleithral spine short, not prominent. First 
spine of the first dorsal fin shorter than sec- 
ond spine, serrate anteriorly. Base of first 
ray of second dorsal fin with serrate anterior 
edge. Teeth present on premaxillaries, den- 
taries, head of vomer, extending posteriorly 
from head of vomer on palatines. Nape, 
prepectoral area, opercle, breast, interpelvic 
area scaleless. Belly with scales. Lateral 
line with 50-52 scales bearing pores, body 
scales small, with cteni, with 10 rows above 
the lateral line and 24-34 rows below. Gill 
rakers on first arch include small rudiments 
on epibranchial and hypobranchial with 
short rakers on epibranchial and ceratobran- 
chial (Table 2). First dorsal, second dorsal 
and anal fins with 1 1 elements each. Pec- 
toral fin with 12-13 connected rays, 3 free 
rays. (Connected pectoral rays difficult to 
count.) Vertebrae 26 (9 + 17 on holotype 
and paratype CAS 86564). Swimbladder 
with extrinsic and intrinsic musculature. 
One papilla present on each eye at one 
o'clock on the following specimens: CAS 
86564, NMC 69-78 (right eye only of 1 12.4 
mm SL specimen). No papillae on eyes of 
holotype. Mandible with flat ridge about % 
its length. 
Coloration. — Two prominent dark sad- 
dles below each dorsal fin and less distinct 
dark area on dorsal surface of caudal pe- 
duncle. Small, irregular, dark spots on dor- 
sal surface of head. Dark areas on margin 
of interspinous membrane of first dorsal fin; 
dark margin on anterior edge of membrane 
of second dorsal fin. Trunk area below lat- 
eral line, and anal and caudal fins lack any 
dark areas, being basically pale. Pectoral fin 
with middle rays dark from base to tip. 
From color photograph of a specimen ap- 
proximately 1 hour after death (Fig. 2): 
specimen bright red with dorsal saddles ap- 
pearing as deep red, margin of first few in- 
terspinous membranes of first dorsal black, 
as well as ventral edge of caudal peduncle 
extending onto lower caudal rays. Dorsal 
fins and anal fin red with scattered yellow 
938 
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 
Figs. 2-3. 2, Photograph of a paratype of Bellator farrago, new species (CAS 86564), taken approximately 
one hour after death. 3, Photograph of Peristedion crustosum taken form submersible Johnson Sea-Link at 486 
m, seamount SE of Isla San Cristobal, Galapagos. 
spots on second dorsal fin. Dark spots not 
visible on head but faint indication of yel- 
low spots. Medial rays of caudal also yel- 
lowish. Pectoral fin with black medial rays 
extending to tip. Lower flank and bottom of 
head white. 
Etymology. — From the Latin farrago, a 
medley or mixture, in reference to the new 
species' combination of its congeners' char- 
acters, here considered a noun in apposi- 
tion. 
Discussion. — This new species is endem- 
VOLUME 111, NUMBER 4 
939 
Table 1 .— Morphometric characters of Bellator farrago, new species. 
CAS 54562 
Speci 
CAS 54564 
mens 
NMC 69-78 
NMC 69-78 
Character 
(mm) 
%SL 
(mm) 
%SL 
(mm) 
%SL 
(mm) 
%SL 
Standard length 
103.9 
96.2 
112.5 
75.5 
Head length 
37.5 
36.1 
33.7 
35.0 
38.8 
34.5 
24.8 
32.8 
Rostral length 
3.1 
3.0 
2.8 
2.9 
1.9 
1.7 
2.1 
2.8 
Rostral width 
4.0 
3.8 
3.9 
4.1 
3.6 
3.2 
3.1 
4.1 
Snout length 
16.5 
15.9 
15.8 
16.4 
14.3 
12.7 
9.3 
12.3 
Premaxillary length 
15.5 
14.9 
12.7 
13.2 
15.3 
13.6 
9.8 
13.0 
Orbit length 
10.4 
10.0 
8.3 
8.6 
11.3 
10.0 
7.0 
9.3 
Orbit depth 
10.9 
10.5 
7.5 
7.8 
11.6 
10.3 
6.0 
7.9 
Interorbital width 
3.9 
3.8 
3.8 
4.0 
4.9 
4.4 
3.8 
5.0 
Cheek height 
8.7 
8.4 
8.5 
8.8 
8.9 
7.9 
5.6 
7.4 
Opercular spine length 
10.1 
9.7 
9.8 
10.2 
11.3 
10.0 
7.2 
9.5 
Preopercular spine length 
9.9 
9.5 
9.0 
9.4 
9.2 
8.2 
7.2 
9.5 
Cleithral spine length 
8.3 
8.0 
7.9 
8.2 
7.6 
6.8 
7.0 
9.3 
1st Dorsal spine length 
8.3 
8.0 
8.2 
8.5 
10.4 
9.2 
8.0 
10.6 
2nd Dorsal spine length 
13.3 
12.8 
12.2 
12.7 
13.2 
11.7 
10.5 
13.9 
3rd Dorsal spine length 
14.4 
13.9 
13.8 
14.3 
17.3 
15.4 
11.2 
14.8 
2nd Dorsal fin base length 
25.5 
24.5 
25.7 
26.7 
28.4 
25.2 
18.5 
24.5 
Anal fin base length 
30.9 
29.7 
29.3 
30.5 
33.2 
29.5 
22.6 
29.9 
Pectoral fin length 
34.7 
33.4 
31.8 
33.1 
34.5 
30.7 
24.2 
32.1 
1st Free pectoral ray length 
32.8 
31.6 
24.2 
25.2 
32.5 
28.9 
19.1 
25.3 
2nd Free pectoral ray length 
28.1 
27.0 
21.5 
22.3 
28.6 
25.4 
16.9 
22.4 
3rd Free pectoral ray length 
25.4 
24.4 
19.0 
19.8 
24.5 
21.8 
13.2 
17.5 
Pelvic fin length 
30.6 
29.5 
28.1 
29.2 
32.2 
28.6 
21.9 
29.0 
Body depth 
28.9 
27.8 
23.6 
24.5 
27.2 
24.2 
19.1 
25.3 
Least depth of caudal peduncle 
7.3 
7.0 
7.1 
7.4 
8.4 
7.5 
6.1 
8.1 
Gill raker length 
2.8 
2.6 
2.4 
2.5 
4.4 
3.9 
1.8 
2.4 
Snout angle (degrees) 
43.0 
28.0 
34.0 
33.0 
Table 2. — Meristic characters of Bellator farrago new species. 
Specimens 
Character 
CAS 54562 
CAS 54564 
NMC 69-78 
NMC 69-78 
Std. length (mm) 
103.9 
96.2 
112.5 
75.5 
1st Dorsal fin rays 
11 
11 
11 
11 
2nd Dorsal fin rays 
11 
11 
11 
11 
Anal rays 
11 
11 
11 
11 
Pectoral fin rays 
13+3 
12+3 
12+3 
12+3 
Gill rakers 
Epibranchials 
2r 
2r 
1 + lr 
1 + lr 
Ceratobranchials 
7+3r 
10+ Ir 
10 
10 
Hypobranchials 
3r 
3r 
3r 
3r 
Total gill rakers 
7 
10 
11 
11 
Squamation 
Lateral line scales 
52 
52 
52 
50 
Scale rows above lateral line 
10 
9 
10 
10 
Scale rows below lateral line 
30 
27 
34 
24 
940 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 
Table 3. — Characters for distinguishing the eastern Pacific species of Bellator. 
Species 
Characters 
gymnostethus 
xenisma 
farrago 
Supplemental preopercular spine present absent present absent 
First dorsal spine shorter than 2nd longer than 2nd longer than 2nd shorter than 2nd 
Breast & interpelvic area naked scaled scaled naked 
Rostral spines produced not produced produced produced 
ic to the Galapagos Islands and is easily 
distinguished from its congeners. Its pos- 
sesses character states present in other Pa- 
cific Bellator (see Table 3); however, in 
combination it differs from each. Its color- 
ation is most like that of B. loxias Jordan 
(in Gilbert 1896) and it has a similar but 
notably different preopercular spine condi- 
tion, but differs in squamation and rostral 
spine condition. The new species' reduced 
first anterior dorsal spine is like that of B. 
gymnostethus Gilbert 1891, from which if 
differs in its coloration and preopercular 
spine condition. Bellator farrago is similar 
to B. xenisma (Jordan & Bollman, 1890) in 
coloration and in its rostral spine condition, 
but differs in its preopercular spine state, its 
first dorsal spine condition, and in lacking 
scales on the breast. Its exact relationship 
to other species of Bellator cannot be de- 
termined at this time as character polarity 
for the genus has not been developed nor 
have cladistic analyses been made. It is pre- 
mature to speculate further. 
A paratype of the new species (CAS 
86564) was collected by the junior author 
while aboard the submersible (JSL). It was 
captured using the submersible 's suction 
device and photographed at the surface 
soon after its death (Fig. 2). The water 
depth at the collection site was 462 m, tem- 
perature 8.58°C, and the bottom was largely 
black volcanic rock overlain with sand 
patches and small boulders partially en- 
crusted with a thin cover of stony corals 
and sponges. 
As a result of our studies, we conclude 
that the Galapagos trigloid fauna comprises: 
Bellator farrago, Prionotus miles, and Pr. 
stephanophrys of the family Triglidae and 
Peristedion crustosum Garman, 1899 of the 
Family Peristediidae. As stated in the intro- 
duction, the previous record of B. loxias (as 
Prionotus loxias, Gruchy 1970) from the 
Galapagos was based on material which we 
describe herein as B. farrago. Bellator lox- 
ias reaches offshore to the Cocos Island 
area as noted by Bussing (1995) and a spec- 
imen collected by the Albatross (MCZ 
30779, 81.0 mm SL) which was identified 
by the senior author. The species of Prio- 
notus reported from the Galapagos also de- 
serve explanation. Prionotus miles Jenyns, 
1842 is an endemic species and has been 
captured at several locations throughout the 
islands. Galapagos records of P. albirostris 
Jordan & Bollman (1890) (Gruchy 1970, 
Bussing 1995, Grove & Lavenberg 1997) 
are based on misidentifications of P. ste- 
phanophrys Lockington (1881), as con- 
firmed by the senior author after examina- 
tion of Gruchy's specimens. Other errone- 
ous records of Galapagos triglids are ex- 
plained by Gruchy (1970:526): including 
Jordan & Evermann's (1896:2172) fisting 
of P. /zorre/25 Richardson (1845) from Gal- 
apagos and Teague's (1951:26) mistake in 
listing the type locality of P. quiescens Jor- 
dan & Bollman (1890) as the Galapagos Is- 
lands rather than the Gulf of Panama. 
Grove & Lavenberg (1997:302-303) re- 
ported upon a poorly preserved specimen 
(LACM 20838, 138.5 mm SL) of Periste- 
dion crustosum Garman 1899 taken off 
Daphne Minor, Galapagos, in 1938 by the 
Allan Hancock Expedition. They suggested 
that ''Peristedion crustosum may prove to 
be a mainland species, which would make 
the population in the Galapagos a new spe- 
cies." Four specimens of Peristedion were 
VOLUME 111, NUMBER 4 
941 
collected by the junior author on JSL dives 
and after comparison with the type material 
of the two eastern Pacific Peristedion by the 
senior author, we find that the JSL speci- 
mens are P. crustosum. The specimens are: 
CAS 86565, 151.6 mm SL, Galapagos Is- 
lands, Isla Genovesa (Tower Island), 
0°21.8'S, 89°58.2'W, JSL Dive 3974, sand 
bottom strewn with small volcanic rocks, 
372 m; and CAS 86562, 103.1 and 108.8 
mm SL, Galapagos Islands, 1°06'S, 
89°12.2'W, seamount SE of Isla San Cris- 
tobal, JSL Dive 3934, volcanic rock and 
boulder bottom with sand channels, 486 m 
(Fig. 3). Other specimens were observed at 
Isla Femandina, off Cabo Douglas (JSL 
Dive 3957, 0°17.5'S, 91°38.9'W), at 431 m 
resting on a 60° sand slope, and off Isla 
Floreana (JSL Dive 3944, 1°13'S, 90°23'W), 
above a flat sand bottom at 305 m, the sin- 
gle specimen collected and deposited at the 
Instituto Nacional de Pesca in Quayaquil. 
Collection records for both species of 
Peristedion are unknown other than the ma- 
terial upon which Garman based his origi- 
nal description. Peristedion crustosum was 
illustrated by Garman in color, but the other 
species, P. barbiger, was not. The two spe- 
cies can be easily separated by the nature 
of the chin and lip barbels and rostral spine 
shape. In P. barbiger the barbels are un- 
usually thick and closely spaced giving a 
thick bearded appearance. In P. crustosum 
the barbels are slimmer and fewer in num- 
ber. Peristedion crustosum has a well de- 
veloped nasal spine but this spine is weak 
or absent in P. barbiger. The striking dif- 
ference is the form of the rostral spines. In 
P. crustosum these spines project forward 
nearly parallel, wherein P. barbiger these 
spines project forward in a convergent di- 
rection, though their tips do not touch. 
There were indications of thickening of 
these bones which possibly could be hy- 
perostosis, a condition found in a few trig- 
loid fishes. 
Acknowledgments 
The late Andre de Roi generously donat- 
ed the holotype of the new species. We 
thank R. Grant Gilmore and the staff of 
Harbor Branch Oceanographic Institution 
for the use of the submersible Johnson Sea- 
Link. For assistance and permission to study 
in Ecuador, we thank: Oscar Aguirre, Sub- 
director de Pesca de Galapagos; Harold 
Miiller, Franklin Ormaza-Gonzalez, and 
Carlos Villon, Instituto Nacional de Pesca; 
Arturo Izurieta Valery and Eduardo Ama- 
dor, Parque Nacional Galapagos; Alejandro 
Villacis, Captain de Puerto de Isidro Ayora; 
and Chantal Blanton, Director de Estacion 
Cientifica Charles Darwin. Sylvia Lafram- 
boise (NMC) kindly loaned us the Gruchy 
(1970) specimens and Karsten Hartel 
(MCZ) kindly loaned us the Peristedion 
types and B. loxias. Molly Brown prepared 
the illustration. We also thank the David 
and Lucile Packard Foundation and the Dis- 
covery Channel for grants and other assis- 
tance. 
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