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Rev. Biol. Trop. 49 Supl. 1: 111-115, 2001
Redescription of Ammodytoides gilli, the tropical eastern Pacific
sand lance (Perciformes: Ammodytidae)
Bruce B. Collette1and D. Ross Robertson2
1National Marine Fisheries Service Systematics Laboratory,
National Museum of Natural History,Washington DC, 20560-0153, USA. fax 202-357-1896, collette.bruce@nmnh.si.edu
2Smithsonian Tropical Research Institute, Unit 0948, APO A A 34002-0948, USA. fax 507-228-0516,
robertson.ross@naos.si.edu
Abstract:Ammodytoides gilli (Bean, 1895) is the correct name for the tropical eastern Pacific sand lance. Its range is
extended from Cabo San Lucas, Baja California south to Panama, Ecuador, and the Galapagos Islands. Ammodytes
lucasanus Beebe and Tee-Van, 1938 is a junior synonym. Types of both nominal species were re-examined. The species
is redescribed based on 50 specimens (42.3-115 mm SL) from 12 lots and is compared with other known species of
Ammodytoides. Changes in ontogeny from the smallest known specimen (42.3 mm SL, illustrated) are detailed including
reduction in the posterior dorsal fin lobe and development of branched dorsal and anal fin rays.
Key words: Ammodytidae, Ammodytoides, sand lance, eastern tropical Pacific.
While collecting fishes for a book on
eastern tropical Pacific fishes (Allen and
Robertson, 1994), Gerald R. Allen and D.
Ross Robertson collected a small sand lance
off the Panama laboratory of the Inter-Ameri-
can Tropical Tuna Commission in May 1990,
well south of its known mainland range in
Baja California. A recently collected series of
sand lance by Robertson from Isla Montu-
osas, Panama stimulated completion of this
project. The purpose of this paper is to
redescribe this poorly-known sand lance from
fresh material and assess its relationships to
other members of the genus Ammodytoides as
defined by Ida et al. (1994), including three
additional species described by Ida and Ran-
dall (1993), Randall et al. (1994), and Collette
and Randall (2000).
MATERIALS AND METHODS
Institutional abbreviations follow Levi-
ton et al. (1985). CAS includes material origi-
nally deposited at Stanford University, SU,
and the New York Zoological Society, NYZS.
Measurements follow Ida and Randall (1993)
and Randall et al. (1994). Generic nomencla-
ture follows Ida et al. (1994). A 42.3-mm
specimen was illustrated (Fig. 1, 2) because it
was the first fresh eastern Pacific specimen
collected and it was in the best condition. It
appears to be one of the smallest specimens of
Ammodytoides known, providing information
on ontogenetic changes.
Ammodytoides gilli (Bean, 1895)
Figs. 1, 2
Bleekeria gilli Bean, 1895: 629-630 (original
description, “Dr. Stimpson’s collection from the Pacif-
Received: 17-III-2000 Corrected: 23-XI-2000 Accepted: 8-XII-2000
R E V I S TA DE BIOLOGIA T R O P I C A L112
fin length 12.8-16.0% SL; caudal fin concav-
ity 6.3-7.8% SL; pectoral fin length 9.9-
11.1% SL.
Scales small, thin, and cycloid, arranged
in straight diagonal rows; head naked, no row
of small scales on upper part of opercle;
scales extending anteriorly to supratemporal
lateral line canal; about 10-12 rows of pre-
dorsal scales; fins naked except caudal fin
with scales extending about three-fourths dis-
tance to posterior margin; lateral line high on
body, ascending from gill opening to a point
three scales below origin of dorsal fin, pass-
ing posteriorly parallel to upper margin of
body, with pored scales ending above mid-
lateral line, 2-5 scales from the caudal base.
Suborbital lateral line canal interrupted, with
four preorbital and four postorbital pores (as
in A. pylei, Randall et al., 1994:fig. 2A).
Freshly collected fish translucent grey-
white, belly silvery white; large fish with
upper two-thirds of head and nape yellow; 1-
5 small black dots evenly spaced along ante-
rior half of outer margin of dorsal fin. Speci-
mens brown in preservative. Dorsal, anal, and
caudal fins with many tiny black spots (Fig.
1); 1-5 larger black blotches in dorsal fin of
three larger specimens (69.5-84.4 mm SL),
absent in smaller specimens (42.3-62.2 mm
SL). Caudal fin with increased numbers of
melanophores in small specimens, forming
distinct V-shaped blotch when caudal fin is
partially folded.
Ontogenetic changes: The 42.3-mm
specimen (Fig. 1, 2) appears to be one of the
ic”, USNM 45384). Seale, 1940: 36 (first record from
Galapagos Is., [CAS 6881]). Ida, 1973: 71 (fig. 3, radi-
ograph of head, extra ossicles in oral region). Winterbot-
tom et al., 1989: 62 (range erroneously given as “Cali-
fornia to Hawaii”, Bleekeria renniei Smith possibly con-
specific with B. gilli). Allen and Robertson, 1994: 256
(description, range extended to Panama [USNM
326833], color fig.).
Ammodytes lucasanus Beebe and Tee-Van, 1938:
306-308 (original description [NYZS 25249A = CAS
SU 46501]; Cape San Lucas, Baja California, from stom-
achs of several species of fishes and a comorant).
Ammodytoides gilli. Ida and Randall, 1993:151
(eastern Pacific species). Ida et al. 1994:252 (ANSP
specimens; generic placement). Randall et al., 1994:85
(eastern Pacific). Grove and Lavenberg, 1997:542-544
(description, Galapagos Islands CAS 6881, LACM
44139-1; Ammodytes lucasanus a junior synonym), fig.
304. Collette and Randall, 2000:398 (comparison with
other known species of Ammodytoides).
Ammodytoides lucasanus. Ida et al. 1994:252
(generic placement).
Diagnosis: A species of Ammodytoides
with dorsal-fin rays 44-47 (mean 46.1); anal-
fin rays 22-24 (mean 23.0); pectoral-fin rays
14 or 15; pelvic fins absent; lateral line
incomplete, pored lateral-line scales 85-94 +
2-5 unpored scales = 88-98 (mean 94.4); gill
rakers on first arch (5-7) + (21-25) = 26-31
(mean 28.7); vertebrae (31-33) + (23-26) =
55-58, including hypural plate (mean 56.5).
Description: Body elongate, body depth
9.5-11.2% SL, body width 5.9-7.8% SL; head
length 25.4-27.8% SL; snout length 6.9-7.9%
SL; orbit diameter 4.1-5.1% SL; fleshy
interorbital distance 3.8-4.9% SL; upper jaw
length 7.8-9.5% SL; least caudal peduncle
depth 4.8-5.4% SL; caudal peduncle length
3.6-6.0% SL; predorsal distance 25.1-26.9%
SL; preanal distance 62.5-65.3% SL; caudal
Fig. 1. Ammodytoides gilli. USNM 326833, juvenile, 42.3 mm SL, Panama, Azuero Peninsula,drawn by Keiko Hiratsuka Moore.
113I N T E R N AT I O N A L J O U R N A L OF T R O P I C A L B I O L O G Y AND CONSERVAT I O N
vs. 2.8-3.7%), longer upper jaw (7.8-9.5% SL
vs. 6.6-7.9%), shorter caudal peduncle (3.6-
6.0% SLvs. 8.4-12.0%), longer predorsal dis-
tance (25.1-26.9% vs. 21.8-24.7%), longer
caudal fin (12.8-16.9% SL vs. 10.5-12.9%),
and a longer pectoral fin (9.9-11.1% SL vs.
8.4-9.5%).
Material examined: 50 specimens (42.3-115 mm
SL) from 12 lots, arranged from north to south. USNM
45384 (88.5), lectotype of Bleekeria gilli, herein select-
ed; probably “Dr. Stimpson’s collections from the Pacif-
ic” (Bean, 1895); D 47, A 23, P114, lateral-line scales 90
+ 4 = 94, gill rakers 6 + 22 = 28, vertebrae 33 + 24 = 57.
USNM 357762 (8, 78.5-91.9) and MCZ 35968 (1, 115),
paralectotypes; out of USNM 45384; in poor condition.
CAS SU 46501 (1, 99.0), Baja California, Cape San
Lucas; April 25, 1936; from stomach of Euthynnus
“alleterata” (= E. lineatus), originally NYZS 25249-A;
holotype of Ammodytes lucasanus. CAS SU 46502 (1,
100), Baja California, Cape San Lucas; May 6, 1936;
from stomach of Seriola colburni ; paratype of A .
lucasanus. CAS SU 46266 (7, ca. 80-85), Cape San
Lucas; originally NYZS 25541; stomach contents; in
poor condition. CAS SU 46737 (9, ca. 50-54), Cape San
Lucas; originally NYZS 25429, 25541; in poor condi-
tion. ZMH 18287 (2, 52.5-56.3), Cape San Lucas; April
25, 1936. USNM 377763 (13, 55.0-84.5) and USNM
377764 (3, 51.6-67.7), Panama, Isla Montuosa, 7° 50’N,
82° 25’W; 25 m on fine sand bottom; D. R. Robertson;
Dec. 12, 1998. USNM 326833 (1, 42.3), Panama, Azuero
Peninsula, Playa Venao, small lagoon near Tuna Com-
mission Laboratory, approx. 7° 24.5’N, 80° 11’W; rock
and sand bottom in 2-3 m; G. Allen and D. R. Robertson
station 32; April 29, 1990. CAS 6881 (1, 59), Galapagos
Is., Floreana I., Black Beach; Jan. 14, 1934. LACM
44139-1 (1, 63), Galapagos Is., Fernandina I., Punta
Espinosa; dried. AMNH 16044 (1, 75.7), 10 mi off north-
western coast of Ecuador, 1° 07’ N, 79° 55’ W; Askoy
Expedition; April 17, 1941.
Two additional specimens could not be located.
SIO 59-209 (2, 47-60), Cabo San Lucas, 22° 52.38’ N,
109° 53.6’ W; C. Limbaugh and R. Rosenblatt; March
20-22, 1959.
smallest known specimens of any species of
A m m o d y t o i d e s, providing information on
ontogenetic changes.
Reduction in the length of the expanded
posterior dorsal fin lobe found in the smallest
juvenile is indicated by the decrease in num-
ber of scales between the end of the depressed
fin and the origin of the caudal fin. The tip of
the last dorsal fin ray is separated from the
upper origin of the caudal fin by 6-9 rows of
scales in specimens 62-100 mm SL, 4-6 rows
of scales in specimens 55-60 mm SL, 2-3
rows in specimens 51.2-51.6 mm, and by
only a single scale in the smallest specimen
examined (42.3 mm SL).
Adult A m m o d y t o i d e s have branched
dorsal and anal fin rays, starting with the third
ray in each fin. However, smaller specimens
of A. gilli, 42.3-57.6 mm SL, lack branched
rays in the dorsal and anal fins. Between 58.0
and 58.8 mm, the fin rays are not branched
but the tips of the fin rays are blunt, not point-
ed as in a simple soft ray. By 59.2 mm, the
third to at least the tenth dorsal and anal fin
rays are branched as in the largest specimens
examined.
Comparisons:Ammodytoides gilli has
fewer lateral-line scales (88-98) than other
species of Ammodytoides or Bleekeria (99-
123) and fewer dorsal-fin rays (45-47) than
the other species of Ammodytoides. There is
no row of small scales on upper part of oper-
cle as is present in A. pylei (Randall et al.,
1994). Ammodytoides gilli has a longer head
than either A. pylei or A. kimurai (25.4-27.8%
SL vs. 22.0-24.6%), larger eye (4.1-5.1% SL
Fig. 2. Ammodytoides gilli. USNM 326833, juvenile, 42.3 mm SL, Panama, Azuero Peninsula, photograph by Gerald R. A l l e n .
R E V I S TA DE BIOLOGIA T R O P I C A L114
In addition to Ammodytoides gilli,
known species of the genus include the type-
species, A. vagus (McCulloch and Waite,
1916) from Lord Howe Island, A. rennei
(Smith, 1957) from South Africa, A. kimurai
Ida and Randall, 1993 from the Ogasawara
Islands, A. pylei Randall et al. 1994 from the
Hawaiian Islands, and A. leptus Collette and
Randall, 2000 from Pitcairn Island.
Comparative material examined: Complete data
for the comparative material is presented in Collette and
Randall (2000). Ammodytoides kimurai Ida and Randall,
1993: paratype, USNM 324610 (1, 121). A. leptus Col-
lette and Randall, 2000: holotype, USNM 360076 (1,
96.2), paratypes, BPBM 16949, MCZ 157036, AMS
I.39856-001, USNM 360077-78 (8, 75.8-89.8). A. pylei
Randall, Ida, and Earle, 1994: paratype, USNM 316514
(1, 137). A. rennei (Smith, 1957): RUSI 8440 (1, 56.4);
ROM 41487 (1, 60.7). A. vagus (McCulloch and Waite,
1916): holotype, AMS I-9272 (143 mm SL). Bleekeria
mitsukurii Jordan and Evermann, 1902: USNM 59599
(2, 78.5-151); UW21253 (2, 91.6-107). B. viridianguilla
(Fowler, 1931): paratypes, ANSP 53462-5 (4, 116-137).
Protammodytes sarisa(Robins and Böhlke, 1970): holo-
type, ANSP 113091 (115 mm SL); paratype, ANSP
113092 (99.2).
ACKNOWLEDGMENTS
We thank Gerald R. Allen for giving the
first Panamanian specimen to the first author
thereby initiating this study. John Earle assist-
ed in field work which was supported by
National Geographic Society Grant 5831-96
to the second author. Curators at the institu-
tions listed above made comparative material
available. Keiko Hiratsuka Moore drew Fig.
1. Ruth Gibbons X-rayed specimens. Drafts
of the manuscript were read by Hitoshi Ida,
Thomas A. Munroe, and John E. Randall.
Hitoshi Ida provided especially useful com-
ments.
RESUMEN
Ammodytoides gilli (Bean, 1895) es el nombre cor-
recto para la “lanceta arenera” del Pacífico tropical ori-
ental que habita desde Baja California sur a Panamá,
Ecuador, y las islas Galápagos. Ammodytes lucasanus
Beebe and Tee-Van, 1938 es un sinónimo. Se redescribe
la especie con base en 50 ejemplares y se le compara con
otras especies de .Ammodytoides. Se indican los cambios
ontogénicos a partir de un ejemplar de 42 mm de longi-
D i s t r i b u t i o n: Ammodytoides gilli i s
presently known from the tropical eastern
Pacific from Cabo San Lucas, Baja Califor-
nia, south to Panama, Ecuador, and the Gala-
pagos Islands. Previous records from the
Hawaiian Islands apply to the recently-
described A. pylei (Randall et al., 1994).
Type Locality: In the original descrip-
tion, Bean (1895) wrote only that the speci-
mens were from “Dr. Stimpson’s collection
from the Pacific.” After examining all avail-
able material from the eastern Pacific and
material of all other described and known
undescribed species of A m m o d y t o i d e s, it
seems clear that the name gilli is the first
name applied to the sand lance found in the
tropical eastern Pacific Ocean. Direct com-
parison with the types of A m m o d y t e s
lucasanus shows this name to be a junior syn-
onym of Ammodytoides gilli.
Ecology: Like other members of the
family, A. gilli is usually found over sand bot-
toms at moderate depths, 2-25 m. A school of
about 500 were seen group-spawning at 4 PM
on Dec. 13, 1998 in 7 m of water over sand in
the lagoon on the south side of Isla Montuosa
in the Gulf of Chiriqui by the second author.
This sand lance is preyed upon by a variety of
fishes such as the tuna Euthynnus lineatus
and the amberjack Seriola colburni as well as
sea birds such as comorants.
Discussion:Ammodytoides gilli and the
three additional species described by Ida and
Randall (1993), Randall et al. (1994), and
Collette and Randall (2000) fit the definition
of the genus as given by Ida et al. (1994).
Species of this genus differ from species of
B l e e k e r i a in the following characters: no
teeth in jaws; infraorbital canal interrupted;
pored lateral-line scales ending high on cau-
dal peduncle instead of curving downward
and continuing onto base of caudal fin; two
predorsal bones present; olfactory rosettes
absent; neural and haemal spines on four pos-
terior caudal vertebrae expanded and flat-
tened; dorsal-fin rays 44-50 (one fewer than
the previously recorded range), and anal-fin
rays 21-25.
115
Ida, H. & J. E. Randall. 1993. Ammodytoides kimurai, a
new species of sand lance (Ammodytidae) from
the Ogasawara Islands. Japan. J. Ichthy. 40:
147-151.
Ida, H., P. Sirimontaporn & S. Monkolprasit. 1994. Com-
parative morphology of the fishes of the family
Ammodytidae, with a description of two new gen-
era and two new species. Zool. Stud., Tokyo 33:
251-277.
Leviton, A. E., R. H. Gibbs, Jr., E. Heal & C. E. Dawson.
1985. Standards in ichthyology and herpetology.
Part I. Standard symbolic codes for institutional
resource collections in herpetology and ichthyolo-
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Randall, J. E., H. Ida & J. L. Earle. 1993. Ammodytoides
pylei, a new species of sand lance (Ammodytidae)
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Seale, A. 1940. Report on fishes from Allan Hancock
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tud, incluyendo reducción del lóbulo de la aleta dorsal
posterior y el desarrollo de rayos ramificados en las ale-
tas anal y dorsal.
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