Front Cover
 Front Matter
 Materials and methods
 List of collection stations
 Zoogeographic implications
 The tismiche
 Discussion of the Tortuguero fish...
 Annotated list of species
 Back Matter
 Back Cover

Group Title: Bulletin of the Florida State Museum, Biological Sciences ( Vol. 16 ; No. 1 )
Title: Fishes of the Tortuguero area, Caribbean Costa Rica
Full Citation
Permanent Link: http://ufdc.ufl.edu/UF00027867/00001
 Material Information
Title: Fishes of the Tortuguero area, Caribbean Costa Rica
Series Title: Bulletin of the Florida State Museum, Biological Sciences ( Vol. 16 ; No. 1 )
Physical Description: Book
Language: English
Creator: Gilbert, Carter Rowell
Kelso, Donald P.
Publisher: University of Florida
Publication Date: 1971
 Record Information
Bibliographic ID: UF00027867
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.

Table of Contents
    Front Cover
        Front Cover
    Front Matter
        Front Matter
        Page 1
        Page 2
        Page 3
    Materials and methods
        Page 4
        Page 5
    List of collection stations
        Page 6
        Page 7
        Page 8
    Zoogeographic implications
        Page 9
        Page 10
        Page 11
        Page 12
    The tismiche
        Page 13
        Page 14
    Discussion of the Tortuguero fish fauna
        Page 15
        Page 16
        Page 17
        Page 18
        Page 19
        Page 20
    Annotated list of species
        Page 21
        Page 22
        Page 23
        Page 24
        Page 25
        Page 26
        Page 27
        Page 28
        Page 29
        Page 30
        Page 31
        Page 32
        Page 33
        Page 34
        Page 35
        Page 36
        Page 37
        Page 38
        Page 39
        Page 40
        Page 41
        Page 42
        Page 43
        Page 44
        Page 45
        Page 46
        Page 47
        Page 48
        Page 49
        Page 50
        Page 51
        Page 52
        Page 53
        Page 54
    Back Matter
        Back Matter
    Back Cover
        Back Cover
Full Text

of the
Biological Sciences
Volume 16 1971 Number 1


Carter R. Gilbert
Donald P. Kelso





SCIENCES, are published at irregular intervals. Volumes contain about 300 pages and
are not necessarily completed in any one calendar year.

FRED G. THOMPSON, Assistant Editor

Consultants for this issue:

Communications concerning purchase or exchange of the publication and all manu-
scripts should be addressed to the Editor of the Bulletin, Florida State Museum, Mu-
seum Road, University of Florida, Gainesville, Florida 32601.

Publication date: 4 June, 1971

Price: $0.85



SYNOPSIS: This paper summarizes fish collections taken from 1955 to 1969 in the
vicinity of Tortuguero, Costa Rica. Most collections were made in the estuary
during the summer rainy season, others in the tributary streams, ocean beach, and
open ocean. The 111 species represented nearly double the total reported by Caldwell,
Ogren, and Giovannoli (1959). Nine are additions to Miller's (1966) checklist
of Central American freshwater fishes, and one species is described as new. Of
the 80 freshwater species collected, six (7.5%) are primary division, 18 (22.5%)
secondary division, and the remainder peripheral division forms, according to
Myers' (1938) classification of freshwater fishes based on degree of salt tolerance.
A list of collection localities is included, with pertinent ecological data, together
with an annotated list of species and a summary table. The paper also summarizes
previous literature pertaining to Central American freshwater and Caribbean shore
fishes, discusses the zoogeographic relationships of the Tortuguero fish fauna, de-
scribes the geographical and ecological characteristics of the Tortuguero area, and
discusses the assemblage of larval fishes and invertebrates (tismiche) periodically
observed in the estuary.

Introduction ................... ................. .........................
Acknowledgements ....................................................3.
Materials and Methods ................................................. ./
List of Collection Stations ................................................
Zoogeographic Implications ...............................................
Description and Characteristics of the Tortuguero Area .......................//
The Tismiche .......................................................... /3
Discussion of the Tortuguero Fish Fauna .................................. X.T
Annotated List of Species ................................. ............... 2
Bibliography .................. ............ ....... ...... ...... .... .7

Carter R. Gilbert is Assistant Curator in Ichthyology, Florida State Museum, and As-
sistant Professor, Department of Zoology, University of Florida, Gainesville, Florida
32601. Donald P. Kelso is Assistant Professor, Division of Biology, George Mason Col-
lege, University of Virginia, Fairfax, Virginia 22030.

Gilbert, Carter R. and Donald P. Kelso. 1971. Fishes of the Tortuguero Area, Carib-
bean Costa Rica. Bull. Florida State Mus., Biol. Sci., Vol. 16, No. 1, pp. 1-54

570.8 o-1

vI ,G


The Central American fish fauna, between the Isthmus of Tehuantepec
and Colombia, remains inadequately known. Collections are uneven, and the lit-
erature consists mostly of descriptions and/or taxonomic reviews of the freshwater
species and faunal reports. Meek (1903, 1904) discussed the zoogeography of
Mexican freshwater fishes, and Regan (1906-08) expanded this to cover all of
Central America. Miller (1966) updated the last work, and in addition pre-
sented a checklist, with ranges, of the Middle American freshwater species known
from the Isthmus of Tehuantepec southward. Loftin's (1965) unpublished work on
the zoogeography of Panamanian freshwater fishes represents the only intensive
study of its type for a specific area of Central America, although similar studies,
by William A. Bussing on Costa Rican fishes and Michael Martin on Honduran
Fishes, are currently in progress. Finally Darlington (1957) and Myers (1966) pre-
sented contrasting theories on the derivation of the freshwater fish fauna. These
are discussed in a subsequent section of this paper.
Some of the above papers treat marine fishes that enter fresh water, al-
though little has been published on the strictly marine species occurring in the
Caribbean that are closely restricted to beach or estuarine situations. The most
comprehensive works dealing with western Caribbean marine fishes are those by
Meek and Hildebrand (1923, 1925, 1928). Chickering (1930) and Hildebrand
(1937, 1939) published on the possible effects of the Panama Canal on trans-isth-
mian movement of both marine and freshwater fishes. Other papers, more limited
in scope, are by Bean (1890); Fowler (1903, 1916, 1923); Caldwell, Ogren, and
Giovannoli (1959); Caldwell (1963); Caldwell and Caldwell (1964); and Birdsong
and Emery (1968). Also pertinent to this discussion are several works dealing
with the marine fishes from the adjacent coast of South America: Fowler (1953)
on Colombia; and Schultz (1949), Mago (1965), and Cervigon (1966, 1968) on
Although many fish collections exist from Central America, these are by no
means uniformly distributed. Certain areas have received inadequate attention, of
which the "Mosquito coast" of Nicaragua and Honduras is the most extensive
geographically. Obviously surveys of the freshwater and marine faunas of this and
other critical regions are vital to our knowledge of the distribution and ecology of
Middle American fishes. Unfortunately problems of transportation and communi-
cation make such areas difficult to reach, and the lack of adequate field facilities
make long-term projects even more difficult.
The establishment of research and living facilities near the small Caribbean
village of Tortuguero, Costa Rica, has resulted in an unusually favorable oppor-
tunity for studying and collecting the biota of a small section of the Central American
coast. For the past 15 years this village has been the site of Archie F. Carr's exten-
sive investigations on the ecology and life history of the green turtle, Chelonia
mydas. During this time various individuals associated with the project have made a
number of fish collections that have formed the basis for several papers (Bihlke,
1958; Caldwell, 1958, 1962; Bohlke and Caldwell, 1961; Gilbert, 1966a, 1966b,
1968; Gilbert and Caldwell, 1967; Collette, 1968). Other papers in preparation,
in which Tortuguero specimens are involved, are by B. B. Collette (families Exo-


coetidae [subfamily Hemiramphinae] and Belonidae) and by C. R. Gilbert and J. E.
Randall (genus Gobionellus, family Gobiidae). Caldwell, Ogren, and Giovannoli
(1959) summarized the earlier Tortuguero fish collections, but as these collec-
tions were incidental to the main turtle work, with most of the specimens com-
ing from the readily accessible lagoon, this checklist was incomplete, and the
fishes were mostly typically brackish-water forms. Collections by the authors in 1963
and 1964, supplemented by a few in 1962, 1965, and 1969 from previously un-
sampled habitats, have more than doubled the number of species recorded from the
area. Despite this, many forms remain to be discovered. Most of these probably
will be marine species that seldom, if ever, venture into the estuary, whereas a
lesser number may also be found in the interior streams. Nearly all previous work
has been done during July, August, and September, at the height of the rainy
season, when collecting success in the streams is generally poor. In addition, the
lagoon at this time has a very low salinity (from 0 to 1.5 ppt at the surface), and
many marine species that are present in the estuary during the dry season (when
salinity is high) move out. The only dry-season collections from the Tortuguero
area were made by the junior author in late April and early May 1964, and most of
these were from the lagoon. Obviously additional collections during this time of
year would be especially desirable. The need for more shore collecting along the
entire Caribbean coast of Central America, regardless of season, is pointed up by the
fact that two species (the dactyloscopid Dactylagnus peratikos Bbhlke and Cald-
well, 1961, and the microdesmid Microdesmus carri Gilbert, 1966) are still known
only from Tortuguero material.
The present paper is an outgrowth of a section of a master's thesis by the
junior author (Kelso, 1965). Its purpose is 1) to update the earlier checklist of
Caldwell, Ogren, and Giovannoli (1959); 2) to provide additional taxonomic, eco-
logical, and distributional data for the included fish species; and 3) to indicate
the availability of this material in the Florida State Museum fish collection. All these
factors assume added importance with the proposal to build a sea-level canal in
southern Panama. Because of the gene interchange that will presumably occur
between many geminate species presently confined to the opposite coasts of Middle
America, comparisons of these closely related forms are especially desirable. In
this regard three papers, in which the Atlantic material has been largely or entirely
collected at Tortuguero, have so far been published (Bohlke and Caldwell, 1961;
Gilbert, 1966a; Gilbert and Caldwell, 1967).


Many individuals have aided in this study. We particularly wish to thank Archie F.
Carr, University of Florida, without whom this work would not have been possible, and
who provided financial assistance from grants by the National Science Foundation (G-1684,
G-5479, G-10717, and GB-86), the Office of Naval Research (ONR 580 (12) ), and the
Caribbean Conservation Corporation. Sr. Guillermo Cruz, San Jose, Costa Rica, was of
invaluable help in various logistical matters. Larry Ogren, Key Eoff, Lenton O.
Rowland, Harry F. Hirth, and Leonard Giovannoli made a number of collections
from 1955-1962; Gene Holmes made several collections in 1969; and Archie F.


(Chuck) Carr III, Randy Kaufman, David Flack, Emory Pierce, Stephen Carr, J.
Richard Moore, and Leo Martinez aided during the collections from 1963-1965.
Thomas H. Fraser, Rosenstiel School of Marine and Atmospheric Sciences, Uni-
versity of Miami, provided information on the present status of the genus Centropo-
mus in the western Atlantic. Robert R. Miller and Reeve M. Bailey, Museum of
Zoology, University of Michigan; Roy Irwin, Tulane University; and William A.
Bussing, Departamento de Biologia, Universidad de Costa Pica, San Jose, provided
information on the systematic status of several Costa Rican freshwater fishes. Russell
Parks, Florida State Museum, took the photographs, and Margaret Estey, Florida State
Museum, drew the map. We extend our sincere appreciation to all.

Fishes were collected by various methods. Most collections from the lagoon and tri-
butary streams were made with rotenone-based toxicants (Chem Fish), although some
were taken with a 15' x 4' /4" bag seine and/or an 8' x 4' 4" non-bag seine. All
collections from the ocean beach were made with the bag seine, to which a metal
chain was attached to the lead line. Experimental gill nets were also used, mostly at
night, both in the lagoon and the streams. Collections from the open ocean were
made, during the day, with a small try net pulled by a catamaran at a maximum
depth of 25-30 feet. Finally some specimens were caught by hook and line.
All specimens were preserved in 10% formalin and later transferred
to 40% isopropyl alcohol. These are deposited in the fish collections of the Florida
State Museum, University of Florida, and (1969 collections only) Florida State
University. Synoptic series of some of the more common species and some paratypes
of Hyphessobrycon tortuguerae, Gobionellus pseudofasciatus, and Microdesmus carri have
been distributed to certain other museums.
All measurements are expressed in standard length (SL), except for the chond-
richthyans (Carcharhinidae and Dasyatidae), which are given in total length
(TL). Species in the annotated list that were collected only in the ocean are indicated
by an asterisk (*). Primary, secondary, and peripheral freshwater fishes (Myers,
1938, 1951) are indicated by roman numerals I, II, or III, respectively. Peripheral
freshwater fishes were so designated if they were ever taken in Tortuguero lagoon
or farther upstream during the rainy season (when the water was either fresh or of very
low salinity), even though in some cases (as with Larimus breviceps and Echeneis
naucrates) it is obvious that these fish stray into fresh water only occasionally. Species
not reported in the earlier list (Caldwell et al., 1959) are indicated by the words
"new record." Vernacular names are listed when known. Both "official" names
(those included in Bailey et al., 1970) and local names are included, the latter
indicated by quotation marks.
The procedures followed in listing collections and in the individual species' accounts
generally follow those of Caldwell et al. (1959) except that 1) catalogue numbers
are not used; and 2) certain collection stations are lumped together (see below). For
each station the geographical locality is followed by the date of collection, pertinent
ecological data, time of day, method of collection, names of collectors, field number,
and species obtained. Under each species account is listed the station where collected
and the number of specimens and their ranges in length from each locality. All
collections made from the ocean or ocean beach are given separate station numbers,
as are those from the tributary steams. Also all estuarine collections the junior
author made during April and May 1964 are listed separately, inasmuch as these are
the only ones made from Tortuguero lagoon during the dry season. The numerous

Vol. XVI No. 1


collections made in the estuary during July, August, and September (various years)
have been combined for the following reasons: 1) Some from the lagoon were made
incidentally, comprise one or a few specimens, and have no precise locality data;
2) many were made from the same or closely adjacent localities and thus are largely
repetitious; and 3) estimated distances from the village or from the lagoon entrance
vary individually and different collections might record the same locality dif-
ferently. Estimates of exact distances were particularly difficult for stream collections,
although in view of the relatively few stream collections made and the large expanse
from which they could have come, that any two were from precisely the same spot is
unlikely. For these reasons no attempt was made to pinpoint exact collection localities
on the drainage map (Fig. 1). Because of pronounced ecological differences (see
subsequent discussion), estuarine collections made from July to September were
separated into the following categories: 1) Those made on the far (west) side of the
lagoon (all combined as station 1); 2) from the near (east) side (station 2); and
3) gill-net and hook-and-line collections away from shore (station 3).
Table 2 shows, in a general way, habitat preferences among the various fish species.
Six major habitats are recognized: The streams, ocean beach, and open ocean, in
addition to the three habitats within the lagoon itself (see preceding paragraph).
The preparation and composition of this table is discussed in greater detail in a later
section of this paper (pp. 17-20).
A problem has arisen regarding material collected in the boca (inlet) area. Collections
made on the two sides of the sand spit that separates the estuary from the ocean were
usually combined. Fortunately field notes taken for the two 1963 collections
so involved (field nos. G 63-16 and G 63-21) show where each species was encountered,
and these collections have thus been divided into "A and B" subcollections; but this
is not true for other years. Consequently the remarks on ecological separation of the
species occurring in this general area are based entirely on data from the 1963 collections.
Observations and discussion of the tismiche are largely by the junior author. All
fish identifications are by the senior author, unless otherwise noted. Various papers,
too numerous to mention, were used for this purpose, with those by Meek and Hilde-
brand (1923, 1925, 1928) being especially useful for the marine species, and those
by Meek (1914) and L6pez-Sanchez (1968) being primarily used for the freshwater
forms. Only the ariid catfishes proved troublesome, the available keys either
being outdated, inaccurate, or incomplete. Specific determinations of the two species
of Ariidae found at Tortuguero are therefore based entirely on the geographical dis-
tributions of the species included in Miller's (1966) checklist of Central American
freshwater fishes. Nomenclature used in this paper follows Miller (1966). All nonfish
identifications and stomach analyses are by the junior author.
The spelling of patronymics has recently been the subject of considerable debate
among ichthyologists. If the present (post-1963) rules of zoological nomenclature are
followed, patronymics must be spelled according to the original orthography, both
with regard to the body of the word and the endings (e.g., i, ii, or with no ending in
the case of masculine patronymic, though this does not apply in those occasional
cases that call for a change in gender). The rationale for this is that it is not always
possible to determine if a person's name is spelled correctly, particularly for species
described during the early days of binomial nomenclature. In earlier papers Gilbert
(1966b, 1967a, 1967b) followed current rules, which many others, including Miller
(1966), have chosen not to follow. In the present paper, we have not followed current
rules, primarily to avoid confusing the reader with patronymics in Miller's (1966)
checklist, but in all cases where the spelling of a name is so corrected the
original spelling is also given.


To facilitate identification of the small fishes collected in the tismiche, a number of
specimens were cleared and stained by the enzyme technique Taylor described (1967).
As identification of the various species of Cichlidae (particularly small
specimens) can present problems, Table 4 was prepared, in which dorsal, anal, and
pectoral fin-ray counts are given (except for Cichlasoma dowi) for a maximum of 15
specimens (all from Tortuguero) of each species. Photographs are included of
small specimens of all species of Cichlasoma found at Tortuguero (Figs. 2-4). In addition,
photographs of large specimens of Cichlasoma spilurum from Costa Rica and
Guatemala (Figs. 5a-b) illustrate differences between populations from the two areas.
The drainage map (Fig. 1) was drawn from topographic maps prepared by the Instituto
Geografico Nacional, San Jose, Costa Rica.

Station 1. (C.O.G. stas. 5, 7, 9, 14). Tortuguero lagoon, west (far) side. A
total of 17 collections made on the following dates: 28 July 1956; 12, 24 July 1957;
11 July 1958; 19 August 1958; 3 September 1958; 12-14, 18-21, 24-25 August 1963;
26 August 1964; 13, 16 August 1969. Extensive shallow bays of 4 ft. or less in depth,
some areas with open bottom and others with some submerged aquatic vegetation and
heavy growths of filamentous algae. Rotenone, seines, and cast nets. Various collectors.
Field nos. Ogren48,225,422-424,461,475; G63-17A,G63-19,G63-26,G63-27,G63-35,
G 63-36, G 63-39; H 6902, H 6903, H 6904. Species: 3, 13, 14, 18, 19, 20, 28, 30,
31, 32, 33, 34, 35, 36, 37, 38, 40, 41, 42, 43, 49, 50, 52, 56, 57, 58, 60, 61, 64,
65, 66, 67, 69, 70, 74, 82, 83, 85, 86, 93, 94, 95, 96, 98, 99, 100, 101, 102, 103, 106,
107, 111.
Station 2. (C.O.G. stas. 3, 4, 6, 8). Tortugero lagoon, east (near) side. A total of
40 collections made on the following dates: 25, 27 July 1955; 22 August 1955; 13, 15,
17, 24 July 1956; 25, 28 August 1956; 1-3, 9 September 1956; 31 August-1 Sep-
tember 1957; 9, 16-18, 24, 27, 30 July 1958; 2, 30 August 1958; 10, 13-14, 16-20,
24 August 1963; 18, 25 August 1964; 24 July 1969; 2, 16 August 1969. Mostly
narrow shoreline with bottom usually sloping sharply into deeper water and with
shallow, open areas lacking in all but a few places; heavy submergent, emergent, and
floating vegetation lining much of shore. Rotenone, seines, cast nets. Various
collectors, Field nos. Ogren 1-5, 7, 25, 131, 141, 152, 155, 157, 175, 233, 277, 420,
430-431, 434, 441-442. 448, 452, 455; G 63-14, G 63-16B, G 63-20, G 63-21B,
G 63-34; H 6901, H 6905, H 6906. Species: 14, 19, 21, 25, 28, 30, 31, 33, 35, 37,
38, 40, 43, 44, 45, 49, 50, 52, 56, 57, 61, 64, 65, 67, 68, 69, 82, 83, 85, 86, 91, 92, 93,
94,95,96,98,99, 100, 101,103, 104, 111.
Station 3. Tortuguero lagoon, nonshoreline areas. A total of 10 collections
made on 12 July 1957, from 11-20 August 1963; and ? July 1965. Bottom uniformly
flat, from 12-15 ft. deep, with mud and detritus, no submergent vegetation. Experimental
gill nets, hook and line. All but 1957 collection by C. R. Gilbert, D. P. Kelso, A. F.
Carr III, D. Flack and R. Kaufman. Field nos. Ogren 205, G 63-15, G 63-17B, G
63-18A & B, G 63-22, G 63-25, G 63-29, G 63-30. Species: 1, 5, 6, 24, 25, 57, 58, 66,
70, 73, 76, 103, 108.
Station 4. (C.O.G. sta. 1). Caribbean Sea, open beach near Tortuguero village,
Beach consisting of black volcanic sand sloping sharply into water; heavy surf
almost always present.
Station 4A. 14 August 1955. Fish driven onto beach by predators. Giovannoli.
Species: 8, 10, 12, 15, 42, 46.

Vol. XVI No. 1


Station 4B 15 July 1956. Fish found dead on beach. Ogren, Species: 77.
Station 4C. 30 July 1958. Fish scooped up as it buried itself in sand at surf line
as wave receded. Ogren. Ogren no. 451. Species: 105.
Station 5. (C.O.G. sta. 2). Just inside inlet, where Tortuguero lagoon enters
Station 5A 26 August 1955. Water almost fresh from recent rains. Hook and
line. Giovannoli and A. F. Carr. Species: 1, 104.
Station 5B. 29 July 1958. Jigs and baited hooks. Ogren and Eoff. Ogren no.
449. Species: 1, 49, 52, 104.
Station 6. (C.O.G. sta. 10). Near confluence of Rio Tortuguero, Lagunas,
Penitencia, and lagoon. Silty bottom, water hyacinths present.
Station 6A. 13 July 1957. Night. Fish shaken from hyacinth roots. Ogren.
Ogren no. 208. Species: 92.
Station 6B. 10 August 1957. Night. Hook and line, with shrimp bait. Ogren.
Ogren nos. 247-248. Species: 83, 85.
Station 7. (C.O.G. sta. 11). Caino Mora, ca. 5 miles from inlet. 9 September
1956. Creek steep-banked; water deep, fresh. Afternoon. Hook and line. Ogren.
Ogren no. 176. Species: 85.
Station 8. (C.O.G. sta. 12). Caiio Palacio, ca. 6 miles from inlet and just
above junction with Lagunas Penitencia. 26 August 1956. Fish found dead near
river. Ogren. Ogren no. 136. Species: 16.
Station 9. (C.O.G. sta. 13). Caflo Palacio, ca. 8 miles from inlet and ca. 11/2
miles above junction of Rio Palacio. 26 August 1956. Water fresh, highly turbid,
brown-stained; river in flood, current 2-3 mph; river running through a river-swamp
forest (no hardwoods), ca. 30-40 yds wide at point of collection; spiny palm most
abundant shore vegetation. Fish collected near shore, cichlids under hyacinths taken
with hook and line and shrimp bait, Eleotris shaken from hyacinth roots. Mid-after-
noon. Ogren. Ogren no. 134. Species: 83, 86, 93.
Station 10. Caribbean Sea, open beach SE of Tortuguero village at "Mile 21/2."
1 August 1962. Black sand beach. Fish found alive at water's edge. Kelso. Species: 109.
Station 11. Rio Tortuguero, ca. 2-3 miles from Tortuguero village.
Station 11A. 10 August 1963. Sidewater tributary, a short distance off main stream.
Bottom of very soft mud, with some submerged branches and logs; no aquatic veg-
etation where collection made, but much emergent vegetation and water hyacinths
close by; water turbid (ca. 1 ft. visibility), depth 4 ft. maximum where collection made;
no current where collection made, but a good current present a short distance away.
3:00-4:00 P.M. Rotenone. Gilbert, Kelso, Carr, Flack, Kaufman. Field no. G 63-13A.
Species: 19, 22, 23, 35, 85, 86, 87, 88, 90.
Station 11B. 10 August 1963. Same locality as above, but in main stream.
Sand bottom grading to mud near shore; heavy emergent and overhanging vegetation;
current swift in center of stream, slower near shore; water 3 ft maximum where
collected. 4:00-5:00 PM. Bag and nonbag seines. Gilbert, Kelso, Carr, Flack, Kaufman.
Field no. G 63-13B. Species: 19, 20, 27, 29, 35, 40, 84, 94.
Station 12. Mouth of Tortuguero lagoon, on seaward side of sand spit. 12, 13,
24 August 1963. Bottom consisting entirely of shifting sand; many small floating


rafts of water hyacinths; collection made at maximum depth of about 4 ft. Specimens
of Phallichthys amates pittieri and Cichlasoma nigrofasciatum found among
hyacinth roots. 10:30-12:00 AM. Bag seine with chain attached to lead line. Gilbert
and Kelso. Field no. G 63-16A. Species: 30, 42, 87, 105.
Station 13. Caribbean Sea, open beach just south of Tortuguero lagoon in-
let. 15 August 1963. Black sand beach, heavy surf. Collection made at maximum
depth of about 4 ft. 4:00-5:00 P.M. Bag seine with chain attached. Gilbert and Pierce.
Field no. G 63-21A. Species: 34,42, 64.
Station 14. Benjamin Creek, sidewater area, a short distance above mouth
in Rio Tortuguero, ca. 21/2 miles above Tortuguero village. 17 August 1963. Mud and
silt bottom with many submerged logs and limbs, no aquatic vegetation. Maximum
depth of collection 3 ft. 10:00-11:00 AM. Rotenone. Gilbert, Kelso, Carr, Flack, Kauf-
man. Field no. G 63-23. Species: 19, 21, 27, 28, 30, 40, 81, 82, 85, 86, 87, 88, 93, 99,
Station 15. Deadwater, just above mouth in Cairo Mora, ca. 11/2-2 miles from
Tortuguero village. 17-18 August and 1 September 1963; September 1964. Stagnant
backwater, with very soft mud bottom, many limbs and logs and heavy mats of water
hyacinths; water darkly stained. Rotenone collection made at maximum depth of ca. 4
ft, gill net and spear collections made in 6-7 ft. 11:00-11:45 AM (poison collec-
tion), overnight (gill-net). A specimen of Lepisosteus tropics speared in September
1964. Gilbert, Kelso, Carr, Flack, Kaufman, Martinez. Field no. G 63-24. Species:
4, 14,27,40,43, 57, 82, 83, 85, 87, 93, 111.
Station 16. Caribbean Sea, up to a mile up and out from Tortuguero inlet.
19-21, 23, 25, 31 August 1963. Black sand and silt bottom. Small (12 ft. wide) try
net, pulled by a catamaran, fishing at a maximum depth of 25-30 ft. Specimen of
Scomberomorus maculatus caught with hook and line. Gilbert et al. Field no. G 63-28.
Species: 2, 7, 9, 11, 39, 46, 47, 48, 55, 62, 63, 71, 72, 73, 78, 79, 80.
Station 17. Canio Servulo, ca. 10 stream miles from Tortuguero village, ca.
1-2 miles above mouth in Lagunas del Tortuguero. 22 August 1963. Soft mud bottom
with much leafy detritus and many limbs and branches. Afternoon. Rotenone. Gilbert.
Field no. G 63-31. Species: 27, 28, 40, 57, 82, 83, 85, 86, 87, 88, 92, 93, 95, 101.
Station 18. Caribbean Sea, shore in front of turtle camp, ca. 2 miles SE of
Tortuguero inlet. 22 August 1963. Open black sand beach with heavy surf. Late
afternoon. Bag seine with chain attached. Specimen of Porichthys plectrodon found,
partially eaten, on beach at night. Gilbert and Carr. Field no. G 63-32. Species: 8, 13,
14, 33, 34,42,45,46,49, 52, 53, 54,62,63,64,69, 73, 75, 80, 110.
Station 19 Rio Palacio, ca. 2 miles above in Cario Palacio. 23 August 1963.
Mud bottom with a number of limbs and logs; no vegetation; water white, not stained.
Collection made in a tiny side tributary, the only place found where collecting was
feasible. 11:30-3:00 PM. Nonbag seine. Gilbert and Kelso. Field no. G 63-33. Species:
19, 20, 27, 28, 29, 30, 31, 35, 38, 40,41,66,81, 84,85,88,89,97, 101, 102.
Station 20. Rio Tortuguero, ca. 3 miles upstream from Tortuguero village. 26
August 1963. Sand bottom with mud near banks; water white, turbid, with good current
present; dense, low, submergent vegetation in shallow areas near shore; width of
stream ca. 10-15 ft; depth ca. 4 ft maximum. 11:00-11:30 AM. Nonbag seine. Gil-
bert, Carr, Flack, Kaufman, Field no. G 63-37. Species: 19, 20, 22, 27, 28, 30, 69, 81,
84,85,87,90,93,95, 101.

Vol. XVI No. 1


Station 21. Rio Tortuguero, ca. 4 miles upstream from Tortuguero village. 26
August 1963. Sand bottom, with some mud near bank; water white, rather turbid,
with strong current; no aquatic vegetation; width of stream ca. 25-30 ft; depth of
capture up to 1 ft, with some deeper spots where current funneled into a chute.
1:00-3:00 PM. Nonbag seine; hook and line. Gilbert, Carr, Flack, Kaufman. Field no.
G 63-38. Species: 19, 20, 22, 27, 29, 35, 40, 41, 43, 44, 64, 84, 85, 93, 94, 95.
Station 22. Tortuguero lagoon, west side, shore ca. 2 miles from inlet, "across
from Leo's." 28 April 1964. 7:00 PM. Dipnet. Kelso. Species: 35, 43, 57, 86, 92, 93.
Station 23. Tortuguero lagoon, at turtle camp, ca. 2 miles above inlet. 28 April
1964. Kelso. Species: 43,45.
Station 24. Tortuguero lagoon, at turtle camp, ca. 2 miles above inlet. 2 May
1964. Kelso. Species: 51.
Station 25. Tortuguero lagoon, bay across from airstrip, ca. 3/ mile above inlet.
5 May 1964. 5:00-6:00 PM. Rotenone. Kelso et al. Species: 17, 18, 34, 35, 36, 37, 40,
45, 50, 52, 56, 57, 61, 64, 69, 82, 86, 93, 94, 95, 98, 100, 101, 106, 107.
Station 26. Flats across from Tortuguero village. 6 May 1964. 5:00 PM.
Bottom dredge. Kelso. Species: 16.
Station 27. Caribbean Sea, beach in front of turtle camp. 6 May 1964.
7:00 PM. Fish driven ashore by predators. Kelso et al. Species: 8.
Station 28. Tortuguero lagoon, shore across from village, ca. 3 miles from
inlet. 7 May 1964. 4:00-5:00 PM. Rotenone. Kelso et al. Species: 35, 43, 50, 56, 57, 58,
64, 65, 70.
Station 29 Tortuguero lagoon, inlet area. 7 May 1964. Kelso. Species: 36, 42,
45,46,50,54,62,69, 105.
Station 30 Inlet of Tortuguero lagoon (tismiche collection). 7 August 1964. 10:00-
11:00 PM. Plankton net. Kelso et al. Species: 36, 40, 93, 94, 97, 101, 106.
Station 31. Creek off Rio Aqua Fria, ca. 8 miles from Tortuguero village. 25
August 1964. Rotenone. Kelso et al. Species: 16, 19, 20, 21, 26, 27, 28, 29, 30, 35, 36,
38, 56,64, 69,81,83,87,88, 89,93, 94, 95, 98, 100, 101, 102, 103.
Station 32. Canal off Rio Aqua Fria, ca. 8 miles from Tortuguero village. 25
August 1964. Rotenone. Kelso et al. Species: 22, 23, 26, 84, 88.
Station 33. Benjamin Creek, 1 mile from mouth. 17 August 1969. 11:00-
12:00 AM. Rotenone and seine. Gene Holmes and Rudolfo Martinez. Field no. H
6907. Species: 14, 19, 22, 27, 30, 31, 40, 43, 81, 83, 84, 85, 86, 87, 92, 93, 94, 95,
101, 102.
Station 34. Benjamin Creek, 2 miles from mouth. 18 August 1969. 10:30-
12:00 AM. Rotenone and seine. Holmes, Martinez, and A. F. Carr III. Field no. H 6908.
Species: 20, 22, 28, 56, 82, 83, 85, 86, 87.
Station 35. Benjamin Creek, 134 miles from mouth. 18 August 1969.
12:30-2:00 PM. Seine. Holmes and Martinez. Field no. H 6909. Species: 19, 20, 22, 26,
27,28,30,31,35,43,44,69,70,81,82,85,86, 88, 93,95,98, 101,111.

Darlington (1957) and Myers (1966) presented contrasting theories regarding


the derivation of the Central American freshwater fish fauna. The latter tendered con-
vincing evidence to refute previous ideas that Central America served as an avenue by
which the characins, catfishes, cichlids, and possibly other groups moved down from
North to South America during the late Mesozoic. Myers thus believes that the
bulk of the South American freshwater fauna came directly from Africa, presumably
before these continents separated and began to drift apart during mid-Mesozoic. The
Central American freshwater fauna is characterized by the dominance of certain autoch-
thonous secondary families or genera (primarily the Poeciliidae and the cichlid genus
Cichlasoma), relatively large numbers of species of marine affinities, and a paucity
of species and lack of diversity in the few primary families (Characidae, Gymnotidae,
and Pimelodidae) occurring north of central Panama. Penetration by the North
American freshwater fauna has been very limited, only one species of Ictaluridae and
one species of Catostomidae having reached the Isthmus of Tehuantepec. Penetration by
the South American fauna has been more extensive, two species (a characin and a
cichlid) having moved north to southern Texas. Geological studies indicate that 1) a
sizeable core of middle Central America, comprising Honduras, El Salvador, and
large adjacent parts of Guatemala and Nicaragua, has remained above the
sea since the Paleozoic; 2) to the north, especially in the Tehuantepec region, con-
siderable marine transgression occurred during the Cenozoic; 3) an oceanic con-
nection existed across southern Nicaragua during the Oligocene and Miocene, and
perhaps at other times; and 4) if the Panama interocean passage (i.e., the area now
occupied by eastern Panama) was ever bridged by dry land prior to the Pliocene, that
bridging was far back in the Mesozoic (Myers, 1966: 771). -Thus no nondisjunct
Central American landmass is believed to have existed between the Jurassic (if then)
and late Pliocene. It should be noted that Savage (1966), in discussing the zoogeo-
graphy of the Central American herpetofauna, hypothesized an Isthmian con-
nection of Central and South America during the Paleocene. Finally, as yet no paleon-
tological evidence documents the presence of catfishes (of South American type),
characins, or cichlids in North or Middle America during the Mesozoic or throughout
most of the Cenozoic. Considering all the above, it seems highly unlikely that Central
America ever harbored a primary freshwater fish fauna till after the final (Pliocene)
emergence of the Panamian isthmus.
The freshwater fish fauna of the Tortuguero area Miller (1966: 782) considered to
belong to the San Juan faunal province, as defined by Regan (1906-08). This province
encompasses lakes Managua and Nicaragua, their outlet the Rio San Juan, and closely
adjacent river systems to the south. Of the 42 species of primary and secondary
freshwater fishes found there, 17 (2 Poeciliidae, 4 Characidae, and 11 Cichlidae) are
endemic or nearly so, although only one endemic genus (the monotypic cichlid
Herotilapia multispinosa) apparently is present. In addition one species each of the
peripheral freshwater families Clupeidae and Atherinidae appears to be so restricted.
Of the 19 endemic fish species occurring in the San Juan faunal province, only 6
(the cichlids Cichlasoma centrarchus, C. citrinellum, C. dowi, C. rostratum and
Herotilapia multispinosa; and the characid Hyphessobrycon tortuguerae) were taken at
Tortuguero. Although further collecting undoubtedly will increase this list, the com-
paratively small percentage of provincial endemics found there indicates that the
freshwater fishes of Tortuguero constitute a somewhat diluted outpost of the main

Vol. XVI No. 1


San Juan fauna. The presence of certain other species, which apparently do not
range north to the Great Lakes-San Juan drainage (e.g., the cichlids Cichlasoma
alfaroi and C. spilotum, the poeciliid Brachyrhaphis parismina, and the pimelodid
Rhamdia wagneri), suggests that the Tortuguero area might best be considered a
region of transition between the San Juan and Isthmian provinces (Miller, 1966:
The marine fishes found at Tortuguero belong almost entirely to the southern
Continental fauna, as defined by Robins (in press) and Gilbert (in press). This fauna
occurs along the South and Central American coasts northward to the tip of Yucatan,
where an abrupt transition occurs between it and the northern Continental fauna. The
amount of overlap between these faunas is apparently small, although more inshore
collections along the coasts of Nicaragua, Honduras, and the eastern Yucatan
peninsula are needed to determine to just what degree this occurs. That some overlap
does exist is shown by the presence at Tortuguero of the northern sciaenids
Menticirrhus americanus and M. littoralis, and by the recent collection of a southern
sciaenid (Bairdiella sanctaeluciae) on the southeast Florida coast (specimens in
University of Florida and Cornell University collections).
No species belonging to the Insular fauna (see Robins (in press) and Gilbert
(in press)) were taken at Tortuguero and probably none occurs there, inas-
much as the conditions these forms require (clear water and a stable environment)
are absent. Yet it should be noted that a diluted Insular fauna does occur in the Lim6n
area, about 50 miles to the south (Caldwell, 1963).

Tortuguero is a small sawmill and turtle-fishing village approximately 52 miles
(84 km) northwest of Puerto Lim6n and 17 miles (27 km) southeast of Barra de
Colorado, on the Caribbean coast of Costa Rica (Carr and Giovannoli, 1957). The
village is on a barrier island separated from the mainland by the brackish Lagunas
del Tortuguero (Fig. 1), formed by the confluence of three streams: Rio Suerte (Caiie
Palacio), Rio Tortuguero, and Rio Sierpe. Rio Suerte has three main tributaries, De-
senredo, Penitencia, and Palacio, and Rio Tortuguero has two, Rio Agua Fria
and Cano Chiquera. Lagunas del Tortuguero extends along the coast south of Tortu-
guero for several miles, and connects with Rio Reventaz6n via the Caiio Negro. The
Suerte-Tortuguero-Sierpe watershed extends inland for about 25 miles (40 km), and
is separated from the drainage of the foothills of the Cordillera Central by Rio Toro
Amarillo, a tributary of Rio San Juan. The watershed is bounded on the north by
tributaries of Rio Colorado and on the south by tributaries of Rio Reventaz6n. Al-
most the entire area is swamp and rain forest, with many interlacing distributaries
connecting the various streams. The main channel of Rio Reventaz6n enters the ocean
at Parismina, 21 miles (34 km) southeast of Tortuguero. Rio Suerte is connected to Rio
Colorado by Cafio Palmas and Laguna Sim6n, which parallels the coast north of
Tortuguero. The main channel of Rio Colorado enters the ocean at Barra de Cdlo-
rado. No direct confluence of the Colorado system exists with Rio San Juan, the
major river in this part of Central America, although temporary connections may
exist during periods of high water.
In the lower part of the Suerte-Tortuguero-Sierpe system the rain runoff water


FIGURE 1. Drainage map of Tortuguero area. Two streams from which collections
were made, Deadwater and Benjamin Creek, are not shown. The former is a back-
water of Caiio Mora and is located a short distance above the junction of that stream
with Caiio Chiquero. Benjamin Creek flows into Rio Tortuguero from the west and is
located just south of Deadwater.

collects first in shallow, mud-bottom pools. These are fairly clear and quiet and, be-
cause they are heavily shaded by palms or hardwoods, are devoid of aquatic plants.
They drain into deep flood-channels, which are more turbid, but are likewise still and
without vegetation, with the bottom ranging from solid to soft mud, sand and sticks.
From these the water moves into narrow, deep, swift-flowing streams, such as Rio
Tortuguero and Rio Palacio. Along the woodland streams patches of rooted aquatic
grasses and water hyacinths (Eichhornia crassipes) sometimes completely cover the

VoL XVI No. 1


surface. To what extent the drainage has been modified by logging is not known, al-
though logs 3 to 4 feet in diameter are floated down the streams during timbering
operations, and the loggers keep the waterways fairly clear of obstructions at these
Lagunas Penitencia, Lagunas del Tortuguero, and Rio Tortuguero join slightly
over a quarter of a mile inland near Tortuguero village. From here the estuary extends
northward parallel to the coast for 3 miles, then turns sharply and enters the ocean.
The estuary is from 400 to 1200 feet wide and width of the peninsula separating the
estuarine basin from the ocean varies from 300 to 1000 feet.
The coast, which extends generally north-northwestward and south-southeastward
in a broad concave arc, is one of the last two major nesting sites in the western Atlantic
for the Green turtle, Chelonia mydas. The beach of black volcanic-sand is completely
open, without offshore bars or coral reefs. Topographically the area is generally a flat
coastal plain, with a few low hills 4 miles inland from the coast south of the estuary
and with one lone prominence, Cerro Tortuguero, rising close to the beach at the
northwest side of the estuarine pass. The foothills of the Cordillera Central begin 30
to 40 miles inland.
The yearly climatic cycle of the region has two dry and two wet seasons (Hirth,
1963). September through November and February through June are dry; July-
August and December-January are rainy, but as Hirth points out, no month has less
than 50 mm of rain. Portig (1965) placed Tortuguero in a higher rainfall class than
Hirth's data indicate. Hirth measured 3882.5 mm from June 1960 through May
1961, whereas Portig estimated 5000-6000 mm as the average annual rainfall. Inland
in the Tortuguero watershed area, lower rainfalls (4000-5000 mm) can be expected.
The watershed is unaffected by rainfall in the mountains. The summer rainy season
appears to begin earlier in the inland watershed area; when the junior author was
at Tortuguero in late April 1964, the rains had already started, whereas on the
coast it was still dry.
The Atlantic coast of Costa Rica was generally classed a "hot zone" by Sapper (1932).
This zone is characterized by an annual mean temperature of 23-260C.

An important phenomenon in Tortuguero estuary is the arrival of the tismiche,
a diverse assemblage of post-larval fishes and crustaceans that appear in the lagoon at
intervals not yet determined. The name tismiche was suggested by Sr. Dilio
Fuentes, Director of the Estaci6n de Biologia Pesquera of Campeche, Mexico, who saw
the phenomenon at Tortuguero in October 1964, and was struck by its similarity to
mixed larval aggregations that occur regularly in the basin of the Rio Papaloapam,
Veracruz, Mexico. The name is evidently of Indian origin, suggesting that the
phenomenon is of regular occurrence in that region. According to Sr. Fuentes,
tismiches occur in Veracruz twice a year, usually in May and December, at which time
the local inhabitants exploit them for food. They appear to consist of several species of
shrimp and fishes, some of which are probably the same as those found at Tortuguero.
Similar assemblages are called seti in Puerto Rico where they consist primarily of
larval gobiids (mostly of the genus Sicydium) and are caught commercially.
Tismiches are not restricted to the New World tropics. Herre (1927: 15-19)


discusses a similar phenomenon in the Philippines, in which the fry of various gobies
and eleotrids (six species predominantly) that have hatched in the sea move into
the estuaries and streams. The natives call these fry ipon; and they form the
basis for an important commercial fishery at certain times of the year. Herre says
nothing about invertebrates in these schools, although some presumably are present.
Ipon migrations occur between October and March, and, though of general occurrence
throughout the islands, seem to be particularly common along the north and northwest
coasts of Luzon. They are commonest for about 3 days after the full moon each month,
when tides are highest.
Although tismiches have been seen off and on since work began at the Tortuguero
turtle camp in 1955 and are well known to local fishermen, their polyspecific
character was not realized until recently. Because all larval aggregations at Tor-
tuguero are not of mixed composition, it has been impossible to determine the season-
ality of the event from old records or from conversations with fishermen.
Large concentrations of larval to post-larval shrimp were seen near the inlet by
Larry Ogren on 3 September 1958 and by the junior author at the tagging camp on 21
August 1963. Individuals from the 1963 tismiche were about 10 mm in
length and were identified as a species of the palaemonid shrimp genus Macrobrach-
ium. The latter concentration was observed for the next 9 days, during which time
the shrimp exhibited a definite pattern of activity, resting by day on the bottom along
shaded sections of the bank, and at night swimming in the upper half of the water
column close to shore. The onset of a strong tidal current would cause the shrimp to
disappear, only to return the following night. The stomach of a Centropomus pecti-
natus, taken at the camp on 31 August 1963, was packed with these shrimp and
nothing else.
Other aggregations of young shrimp, seen on 4 May and 19 July 1964 at the
tagging camp, consisted mostly of atyid shrimp (genera Jonga, Potimirim, and
Micratya) about 5 mm long, with many 10 mm Macrobrachium also present. On 7
August 1964 collections were made from a tismiche near the inlet. In addition to the
above shrimp, which formed the bulk of the collection, many crab megalops (ca. 2
mm across the carapace) and post-larval fishes were also present, including small num-
bers of Microdesmus carri (Microdesmidae) and Awaous tajasica (Gobiidae), and
larger numbers of Gobionellus fasciatus (Gobiidae) and Eleotris sp. (probably both
E. amblyopsis and E. pisonis) (Eleotridae). Very likely other species of fish were also
present, though in smaller numbers. In addition, many sniall (67-85 mm) pipefish
(Oostethus lineatus) were scattered throughout the concentration. These were swim-
ming upstream against the current, the only indication of orientation by any com-
ponent of the tismiche. Many fish-eating bats (Noctilio) were seen fishing at the
surface. Large numbers of adult mullet (genus Mugil) were also present, although
their stomachs proved to be empty when examined, but the stomachs of four large
Pomadasys crocro were packed with small shrimp and fish. This tismiche was observed
both at the inlet and the turtle camp for nearly 3 weeks (till 26 August), during which
time it fluctuated considerably in abundance. Much larger concentrations of fishes
and crab megalops were seen at the pass than at the turtle camp, and on the eight
nights collections were made at the camp only five fishes and no crabs were taken.
Obviously more study is needed to determine what correlation, if any, exists

Vol. XVI No. 1


between the tismiche and various environmental phenomena (such as lunar or
tidal changes), and whether the appearance of the tismiche follows a predictable

Of the 111 fish species collected at Tortuguero, 80 were taken at least once
in fresh, or nearly fresh water. Of these, 56 species (in 25 families) may be classed as
peripheral, of which 9 (Citharichthys uhleri, Coleotropis blackburni, Oligoplites
palometa, Larimus breviceps, Menticirrhus americanus, Bathygobius soporator,
Gobionellus pseudofasciatus, Echeneis naucrates, and Microdesmus carri) are addi-
tions to Miller's (1966) checklist. Eighteen species, belonging to 4 families (22.5%),
are secondary freshwater fishes, and only 6 species (3 families) are primary freshwater
fishes. The percentage of fishes belonging to the last group (7.5%) is thus
considerably less than the percentage (22.8%) of primary species Miller lists for
Central America as a whole, although the latter figure includes a large number of
South American species not recorded beyond eastern Panama.
The 111 species represents an increase of 56 over the number Caldwell, Ogren, and
Giovannoli (1959) recorded from Tortuguero. As one of the species reported in the
1959 paper (Centropomus ensiferus) was misidentified, this number should really
be 57, but the increase actually should only be 54, for three species reported as new in
the present paper (Strongylura timucu, Centropomus undecimalis, and Gobionellus
boleosoma) were present in the 1955-1959 collections, but were not distinguished
from S. marina, C. parallels, and G.fasciatus, respectively.
In addition to the above, the 1959 record of Anchoa lamprotaenia merits comment.


Caldwell, Ogren,
& Giovannoli, 1959
Subspecies name deleted:
Harengula p. caribbaea
Subspecies name added:
Phallichthys pittieri
Hyporhamphus roberti

Specific name changes:
Gobionellus claytonii

Trinectes maculatus

Generic name changes:
Thryinops chagresi
Garmannia spes

Generic and specific name changes:
Molliensia sphenops

Microeleotris mindii

Present paper

H. pensacolae

P. amates pittieri
H. r. hildebrandi

G.fasciatus (Tortuguero
material only)
T. paulistanus (Tortuguero
material only)

Melaniris chagresi
Gobiosoma spes

Poecilia mexicana (Tortu-
guero' material only)

Reference or source

Rivas, 1963

Rosen & Bailey, 1959
B. B. Collette, pers.

Robins & Lachner,
G. C. Miller (fide
C. L. Hubbs), pers.

Miller, 1966
Bohlke & Robins, 1968

Miller, 1966; and
in litt.
Miller, 1966


Generic name changes and/or
specific name additions:
Carcharhinus sp.

Anchoviella sp.

Coleotropis sp.

Eugerres sp.

Dactylagnus sp.

Species to be deleted from list:
Centropomus ensiferus

Complexes of species:
Strongylura marina
Centropomus parallels

Gobionellus claytonii

Table 1. Continued

Carcharhinus leucas

Cetengraulis edentulus

Coleotropis blackburni

Diapterus plumieri

Dactylagnus peratikos

C. parallels

S. marina and S. timucu
C. parallels and C.
G.fasciatus (see above)
Sand G. boleosoma

Specimens examined
by authors
Specimen reexamined
by Gilbert
Gilbert & Caldwell,
Miller, 1966; speci-
mens examined by
Bohlke & Caldwell,

Specimen reexamined
by Gilbert

Collette, 1968
Specimens reexamined
by Gilbert
Specimens reexamined
by Gilbert

The senior author reexamined the 290 specimens on which this record was based, and
Robert R. Miller a lesser number; all proved to be Anchoviella elongata. As a few
(10) specimens of Anchoa lamprotaenia were collected in 1962 and 1963, Caldwell,
Ogren, and Giovannoli's (1959) record for this species is valid, though based on mis-
identified material. The present paper includes, in addition to the new species resulting
from recent collections, a number of name changes, both at the genus and species levels.
Some of these are of a nomenclatural nature, whereas others are taxonomic. These
are listed in Table 1.
Table 2 is intended to show the general ecological distribution of fishes around
Tortuguero by listing the actual number of specimens collected from each of the six
major habitats (ocean, offshore; ocean beach; estuary, west side; estuary, east side;
estuary, center; streams). While habitat preference might be indicated better by
listing the species as rare, common or abundant in each, only for the shore areas of the
lagoon are sufficient collections available to provide a reasonably objective idea of
relative abundance. Obviously the abundance of a particular stream species cannot be
determined accurately from a dozen widely scattered collections, most of which were
made in quiet sidewaters during periods of high water at one season of the year.
Another problem arises regarding the distinction between collections made
during the wet and dry seasons. Increased penetrance of salt water into the est-
uary during the wet season allows certain marine species to move into the
estuary that otherwise would be unable to do so. Because the number of dry-season
collections are too few to permit accurate comparison, both Table 2 and the following
discussion are based solely on material taken during the wet season.
The table also does not take into account 1) size of the specimens tabulated;

Vol. XVI No. 1


2) differences in numbers of collections made from the different habitats; 3)
different collecting methods (which vary in effectiveness for different species); 4)
unusually large concentrations of specimens in one collection (e.g., Pomadasys crocro
and Microdesmus carri); or 5) that many specimens of the larger and commoner
species such as Cichlasoma friedrichsthali, C. maculicauda, and Sphoeroides testudineus
were discarded in the field.
Despite these shortcomings the table does provide some useful information,
particularly with regard to differences in species composition between the ocean and
estuary, and between the estuary and tributary streams. For example, the atherinids
Coleotropis blackburni and Melaniris chagresi were collected together only once,
although they occur commonly within 100 feet of one another at the mouth of the
lagoon (Coleotropis on the ocean side, Melaniris on the estuary side). The pomada-
syids Conodon nobilis and Pomadasys corvinaeformis were never encountered in the
estuary, whereas P. crocro was found in the ocean, the lagoon, and also in the tributary
streams. The sciaenids Bairdiella ronchus and Micropogon furnieri were found only in
fresh and/or brackish water, whereas all other sciaenids were taken only in the ocean,
with the exception of one specimen of Larimus breviceps gill-netted in the estuary.
Differences in salinity tolerance offer one explanation for the above, although un-
doubtedly other factors are also involved. Certainly the fact that such species as
Citharichthys spilopterus, C. uhleri, Achirus lineatus, Trinectes paulistanus and
Sphoeroides testudineus were found only in the lagoon has no relationship to their
ability to live in salt water. Much more needs to be known regarding the ecology of
these species before such problems can be answered.

1. Carcharhinus leucas 6
2. Carcharhinus limbatus 1
3. Himantura schmardae 1
4. Lepisosteus tropics 1
5. Elops saurus 6
6. Megalops atlantica 5
7. Chirocentrodon bleekerianus 1
8. Harengula pensacolae 19
9. Odontognathus compressus 67
10. Opisthonema oglinum 14
11. Pellona harroweri 13
12. Sardinella anchovia 9
13. Anchoa lamprotaenia 2 1
14. Anchoviella elongata 1 345 2 39
15. Cetengraulis edentulus 1
16. Synbranchus marmoratus 2
17. Anguilla rostrata -
18. Myrophis punctatus 178
19. Astyanaxfasciatus 32 59 164
20. Brycon guatemalensis 1 41
21. Hyphessobrycon tortuguerae 7 11


Table 2. Continued

22. Roeboides guatemalensis
23. Gymnotus cylindricus
24. Arius melanopus
25. Bagrefilamentosus
26. Rhamdia wagneri
27. Alfaro cultratus
28. Belonesox belizanus
29. Brachyrhaphis parismina.
30. Phallichthys amates pittieri
31. Poecilia mexicana
32. Strongylura marina
33. Strongylura timucu
34. Hyporhamphus roberti hildebrandi
35. Citharichthys spilopterus
36. Citharichthys uhleri
37. Achirus lineatus
38. Trinectes paulistanus
39. Symphurus plagusia
40. Oostethus lineatus
41. Pseudophallus mindii
42. Coleotropis blackburni
43. Melaniris chagresi
44. Agonostomus monticola
45. Mugil curema
46. Polydactylus virginicus
47. Sphyraena guachancho
48. Scomberomorus maculatus
49. Caranx hippos
50. Caranx latus
51. Chloroscombrus chrysurus
52. Oligoplites palometa
53. Trachinotus carolinus
54. Trachinotus goodei
55. Vomer setapinnis
56. Centropomus parallelus
57. Centropomus pectinatus
58. Centropomus undecimalis
59. Epinephelus (?) sp.
60. Lutjanus griseus
61. Lutjanusjocu
62. Conodon nobilis
63. Pomadasys corvinaeformis
64. Pomadasys crocro
65. Diapterus olisthostomus
66. Diapterus plumieri
67. Diapterus rhombeus
68. Eucinostomus argenteus
69. Eucinostomus pseudogula


4 3

1 5

4 26
5 18

154 2


1 7

2 8


51 30

1 13


9 20 9



1 1


15 98 16

Vol. XVI No. 1


Table 2. Continued
00 OB

70. Bairdiella ronchus
71. Bairdiella sanctaeluciae
72. Cynoscion jamaicensis
73. Larimus breviceps
74. Menticirrhus americanus
75. Menticirrhus littoralis
76. Micropogonfurnieri
77. Ophioscion costaricensis
78. Ophioscion panamensis
79. Stellifer colonensis
80. Umbrina broussoneti
81. Cichlasoma alfaroi
82. Cichlasoma centrarchus
83. Cichlasoma citrinellum
84. Cichlasoma dowi
85. Cichlasoma friedrichsthali
86. Cichlasoma maculicauda
87. Cichlasoma nigrofasciatum
88. Cichlasoma rostratum
89. Cichlasoma spilotum
90. Cichlasoma cf spilurum
91. Herotilapia multispinosa
92. Dormitator maculatus
93. Eleotris amblyopsis
94. Eleotris pisonis
95. Gobiomorus dormitor
96. Leptophilypnusfluviatilis
97. Awaous tajasica
98. Bathygobius soporator
99. Evorthodus lyrics
100. Gobionellus boleosoma
101. Gobionellusfasciatus
102. Gobionellus pseudofasciatus
103. Gobiosoma spes
104. Echeneis naucrates
105. Dactylagnus peratikos
106. Microdesmus carri
107. Batrachoides gilbert
108. Batrachoides surinamensis
109. Porichthys pauciradiatus
110. Porichthys plectrodon
111. Sphoeroides testudineus


6 20 11
120 30 15
24 36 105
251 9 26



many 70
many 7

7 2
many 61

81 17

The six habitats are: Ocean, offshore (00); Ocean beach (OB); estuary, west side (EW); estuary, east side (EE); estuary,
center (EC); streams (S).
Two species (Anguilla rostrata and Chloroscombrus chrysurus) were not collected during the wet season, but are included
for the sake of completeness.

Table 2 also shows that not all species were collected in equal numbers on both
sides of the estuary. For example the predominance of shallow, open mud-sand flats


on the west side apparently offers a more favorable habitat for a burrowing species
such as Myrophis punctatus; this is reflected by the fact that all 199 specimens of this
species were taken there, although it probably occurs along the east shore in the
relatively few places where open flats exist. A comparison of the relative numbers of
the cichlids Cichlasoma centrarchus, C. citrinellum, and C. maculicauda taken on the
two sides shows that most individuals of the first species were collected along the
east bank, whereas the other two were encountered in much greater numbers on the
opposite shore. This appears to be related to a preference by C. centrarchus for heavy
submergent vegetation (not including filamentous algae), such as one finds along much
of the east bank, as opposed to a preference by C. citrinellum and C. maculicauda for more
open situations. Gobiosoma spes was encountered in greatest numbers where heavy
growths of filamentous algae were present; the number of specimens of this species
collected along the west shore, where algae was very abundant in certain places, out-
numbered those taken elsewhere by over 20 to 1. All but two specimens of Dormitator
maculatus collected in the estuary were taken on the east side. This species was
found exclusively among water hyacinth roots, and the differential collecting success
is related to the much heavier concentrations of hyacinths along the east shore.
A number of species found in the tributary streams were never taken in the
estuary, and other more or less common stream inhabitants were collected in the
estuary only once or twice. Although a substantial percentage of the total specimens (7
(the types) out of 25) of Hyphessobrycon tortuguerae and the only two individuals of
Herotilapia multispinosa taken during this study were found in the estuary (in 1955
and 1956 respectively), both are probably stream inhabitants, as evidenced by their
absence from the many subsequent estuarine collections. These and other species
occasionally appearing in the lagoon probably are carried down, in most cases, under
floating mats of water hyacinths. The only specimen of Cichlasoma nigrofasciatum
taken in the lagoon was found under such a mat.
Of the four species ofcharacins collected during this study, one (Astyanax fasciatus)
appears to be more or less ubiquitous, a second (Hyphessobrycon tortuguerae) is
thought to be basically a stream inhabitant (see above), and the other two (Brycon gua-
temalensis and Roeboides guatemalensis) obviously prefer a stream environment. The
various cichlid species collected only in streams were nearly always found in sidewaters
where current was completely lacking. On the basis of this admittedly incomplete evi-
dence, it appears that some other factor, or factors, are involved in the distribution of
the various cichlid species. Of the cichlids found at Tortuguero, two (C. citrinellum and
C. maculicauda) were more common in the estuary, six (C. alfaroi, C. dowi, C. nigrofas-
ciatum, C. rostratum, C. spilotum, and C. the exception of the one specimen of C. nigrofasciatum mentioned earlier), and two
(C centrarchus and C. friedrichsthali) were equally common in both areas. The ecologi-
cal status of Herotilapia multispinosa has been discussed previously.
Finally, of the five species of Poeciliidae encountered during this study, Poecilia
mexicana was found much more commonly in the estuary, whereas the other four
were either taken exclusively in the tributaries (Alfaro cultratus and Brachyrhaphis
parismina) or were much more common there (Belonesox belizanus and Phallichthys
amates pittieri).

Vol. XVI No. 1



1. Carcharhinus leucas (Valenciennes). Bull shark. (III)
Stations 2, 3, 5A, 5B (teeth only saved).
The bull shark is common in Tortuguero lagoon. Many have been caught in the area,
mostly by hook and line, a few in gill nets. The 13 individuals (6 females and 7 males)
collected in 1963-1964 ranged from 1016 to 1930 mm total length, and weighed from
42 to 110 pounds.
Age-length data given by Clark and von Schmidt (1965) indicate the above speci-
mens are all of subadult size, but as Springer (1960: 27) notes, the average size of
adult C. leucas from Trinidad is appreciably less than for adults from the Gulf of Mexi-
co. Inasmuch as the pelvic claspers on male C. leucas taken in Tortuguero lagoon were
well developed, it appears likely that the Tortuguero and Trinidad individuals belong
to the same population.
The stomachs of several bull sharks from Tortuguero contained crabs (Callinectes
bocourti); numerous fishes of various sizes, including one 350 mm Centropomus sp.
(probably undecimalis); bird remains; a small trunkback turtle (Dermochelys); and
the remains of a butchered Hawksbill turtle (Eretmochelys).
Local residents occasionally remove and dry the fins of C. leucas, and then sell
them for soup ingredients.
It is not known how far this shark moves up the tributaries in the Tortuguero area,
although judging from the distances the species travels up the neighboring Rio San
Juan (Thorson, 1966) it probably is considerable. Its ability to migrate upstream is
probably limited more by its size than by its physiology.
2. Carcharhinus limbatus (Valenciennes). Blacktip shark. (*)
Station 16 (1, 867); new record.
The above specimen, a subadult male, was caught by hook and line 23 August 1963,
About a half-mile offshore in about 30 feet of water. This species is probably fairly
common in the area, although it seldom, if ever, enters the lagoon.


3. Himantura schmardae (Werner). (III)
Station 1 (1, 535); new record.
The specimen is an immature male, caught on the flats across from the village
25 August 1963 (field no. G 63-36). No other rays were seen or collected, although
local residents say they sometimes are fairly common.


4. Lepisosteus tropics (Gill). Tropical gar; "alligator fish." (II)
Station 15 (1, 695); new record.
Several individuals were seen in Deadwater; the specimen was collected in Sep-
tember 1964. Deadwater, as the name implies, is a stagnant, darkly stained, weed-choked
I For explanation of symbols see page 4.


backwater, one of the few such places in the immediate vicinity of Tortuguero; it is
the only place where Lepisosteus is definitely known to occur.

5. Elops saurus Linnaeus. Ladyfish; "bone fish." (III)
Station 3 (2, 365-430); new record.
A number of specimens were caught by hook and line at the inlet during Julyand
August of 1963-64, and two other individuals (432-508 mm) were taken at the tagging
camp in early May 1964, during the dry season. The stomachs of the last two specimens
contained six 15-25 mm crabs, three 25-mm river shrimp, and four small clupeiform
fishes. The 508-mm specimen contained an undetermined number of eggs in its
ovaries, averaging about 0.75 mm in diameter.
6. Megalops atlantica Valenciennes. Tarpon; "sabalo." (III)
Station 3 (1 skeleton saved); new record.
Only mature tarpon were caught in Tortuguero lagoon, ranging in length from
about 1000 to 1300 mm and in weight from 30 to 46 pounds. All specimens examined
were males, and all of the seven caught during the summer of 1964 were ripe. The
stomachs of the latter individuals were examined, but all were empty. The apparent
absence of young stages of tarpon at Tortuguero may perhaps be related to the lack of
small brackish and freshwater ponds, which elsewhere have been found to be a
favored habitat of the young (Wade, 1962: 591).
Some workers (Greenwood, 1970) place this species in a separate genus, Tarpon.


7. Chirocentrodon bleekerianus (Poey). (*)
Station 16 (1, 41.5); new record.

8. Harengula pensacolae Goode and Bean. Scaled sardine; "sprat." (*)
Stations 4A (14, 69-102), 18 (3, 95-100), 27 (2, 79-94.5).
9. Odontognathus compressus Meek and Hildebrand. (*)
Station 16 (67, 48-65); new record.
This collection represents the northernmost record for this species along the
Central American coast (Berry, 1964: 730).
10. Opisthonema oglinum (Lesueur). Atlantic thread herring. (*)
Station 4A (14, 66-98).
This species has not been collected at Tortuguero since 1955.
11. Pellona harroweri (Fowler). (*)
Station 16 (13, 29-45); new record.
This represents the northernmost record for this species along the Central
American coast (Berry, 1964: 729).
12. Sardinella anchovia Valenciennes. Spanish sardine. (*)
Station 4A (9, 67-83).
This species has not been collected at Tortuguero since 1955.

Vol. XVI No. 1



13. Anchoa lamprotaenia Hildebrand. Longnose anchovy (III)
Stations 1 (1, 41), 18 (2, 40-44).
In addition to the above, seven other individuals (34-37 mm) were collected in an
unspecified area of the lagoon in August 1962. Gill-raker and anal-ray counts for three
specimens were 16 + 19, 17 + 19, and 18 + 20; and 21, 22, and 22, respectively. As
mentioned previously, all individuals identified by Caldwell, Ogren, and Giovannoli
(1959) as this species have proved, upon reexamination, to be Anchoviella elongata.
14. Anchoviella elongata (Meek and Hildebrand). (III)
Stations 1(345, 39-77), 2 (2, 58-65), 15 (6,44-45), 18 (1, 49), 33 (43, 33-45); new
In addition to the various morphological and meristic characters by which this
species can be distinguished from Anchoa lamprotaenia, it also apparently reaches a
larger size.
15. Cetengraulis edentulus (Cuvier). (*)
Station 4A (1, 72.5).
Caldwell, Ogren, and Giovannoli (1959: 15) list this specimen as Anchoviella sp.
Reexamination shows their gill-raker count (37 + 45 =82) to be in error, the apparent
result of breakage of the upper limb of the outer gill arch; the corrected count should
be 49+45=94. This count, together with the number of dorsal and anal-fin
rays (14 and 24, respectively), agrees closely with counts given by Hildebrand (1963:
245) for Cetengraulis edentulus. Identification of the Tortuguero specimen as this
species is further confirmed by a careful comparison with Hildebrand's (1963: fig. 59)
illustration and by other morphological features appearing in his diagnosis. The only
apparent character of importance he does not mention are the sheaths of large scales
covering the basal half to two-thirds of the anal fin and the lowermost part of the
dorsal fin. The thin membrane connecting the gill covers across the isthmus (a key
generic character) is torn, but can be seen by careful scrutiny.
A second specimen of this species, 88 mm SL, from Cayenne, French Guiana, has
47 + 51 = 98 gill rakers, 13 dorsal rays, and 25 anal rays. The gill membrane in this
specimen also is no longer intact.


16. Synbranchus marmoratus Bloch. (II)
Stations 8 (1, 150), 26 (1, 152), 31 (1, 265).
This is a burrowing species that has the ability to live for long periods of time in
habitats of low oxygen concentration. In addition it sometimes moves overland (as
does Anguilla rostrata) after rains. Carr and Giovannoli (1950) found individuals in
Honduras several feet from water, both under rocks and buried in the mud. Sterba
(1962: 839) reports the maximum total length of this species as 1500 mm.


17. Anguilla rostrata (Lesueur). American eel. (III)
Station 25 (1,460); new record.


18. Myrophis punctatus Liitken. Speckled worm eel. (III)
Stations 1 (178,61-115), 25 (23,67-160).
As mentioned previously, this species was taken only on the mud flats on the far
(west) side of the estuary. It apparently does not grow as large at Tortuguero as it
does elsewhere. Florida individuals are known to reach lengths of at least 285 mm.
19. Astyanaxfasciatus aeneus (Giinther). (I)
Stations 1(32, 17-76),2 (59,21-78), 1A (1, 54.5), 11B (8,23-31.5), 14 (9,16.5-73),
19 (31, 18-100),20 (20,20.5-67.5), 21 (37,18-79), 31 (39,23-72.5), 33 (6,22.5-35.5), 35
(13, 17.72).
This species was extremely common during the wet season in both the streams and
estuary, but was not taken in the latter area during the dry season. Anal-ray counts
recorded by Caldwell, Ogren, and Giovannoli (1959: 16) for 59 specimens are as
follows: iv, 24 (3); iv, 25 (17); iv, 26 (16); iv, 27 (17); iv, 28 (3); iv, 29 (3). The
following color notes are taken from the same paper: "the caudal, anal,
and pelvic fins were brick red, with the upper and lower portions of the first and the
leading edges of the latter two fins even brighter (and darker) than the rest of the fin.
The dorsal and pectoral fins, while red, were much less intensely so. The dorsal portion
of the body was metallic black, and the dark stripe on the caudal pedunce very con-
20. Brycon guatemalensis Regan. "Guapote." (I)
Stations 1 (1, 146), 11B (6, 30-37), 19 (1, 34.5), 20 (1, 23.5), 21 (20, 23-290), 31
(2, 26-38.5), 34 (6, 79-152), 35 (3, 89-225); new record.
This large characin is common above Cuatro Esquinas, and a few stray into the
estuary during the wet season. A 146-mm specimen was seined on the flats across
from the village, and a 390-mm, 2.2-pound individual was gill-netted at the tagging
camp, both in August 1964; neither was saved. This species seems to prefer more
rapidly-flowing water than any of the other species of Characidae found in the area.
21. Hyphessobrycon tortuguerae Bohlke. (I)
Stations 2 (7, 24-39), 14 (2, 21-22.5), 31 (9,18-19).
Although the seven type specimens of this species were taken in the lagoon, the
absence of subsequent material from there indicates that H. tortuguerae is basically a
stream inhabitant. Since its original description, the known range of this species has
been extended outside the Tortuguero area (Miller, 1966: 785; W. A. Bussing, in
22. Roeboides guatemalensis (Giinther). (I)
Stations 11A (5, 20-74), 20 (6,25-49), 21 (2, 61-83), 32 (38,27.5-87.5), 33 (1,33),
34 (2, 72.5-87), 35 (2, 66.5-70); new record.
This species was never found in the main channel of the streams, as was Brycon
guatemalensis. It was the only characin never taken in the lagoon proper.
23. Gymnotus cylindricus LaMonte. (I)
Stations 11A (1, 140), 32 (2, 238-240); new record.

Vol. XVI No. 1


Miller (1966:786) gives the range of Gymnotus cylindricus as "Atlantic slope from
Rio Motagua, Guatemala, to northwestern Honduras," and the range of G. carapo?
as "Both slopes of Costa Rica and Nicaragua (Myers) and the Atlantic slope of extreme
western Panama (Behre, 1928: 310); northern limit of range of this South American
species very uncertain." Despite this, the Tortuguero specimens are identified as
G. cylindricus on the basis of both morphological characters (see below) and the
authority of Bussing (1967), who referred all his Costa Rican material to this species.
In so doing, Bussing (1967: 221) said, "Dr. Robert M. (sic) Miller (in litt.) states that
specimens of Gymnotus from the Atlantic slope of Costa Rica are definitely not
G. carapo."
In the original description (LaMonte, 1935), G. cylindricus was said to differ from the
closely related G. carapo in having a shorter head (9.4 (vs. 6.9) in body length),
shorter snout (3.8 (vs 3.1) in head), deeper head (1.4 (vs. 1.9) in body depth), and
three rows of teeth instead of one. No differences in body coloration were noted.
Nevertheless, R. R. Miller informs me (Gilbert) that color pattern provides the
best means for separating the two species, carapo being marked by oblique stripes and
cylindricus having a uniformly dark coloration. Tortuguero specimens are uniformly
pigmented. Proportional measurements for these individuals (140, 238, and 240 mm)
are as follows (shortest to longest specimens, respectively): body length/head length
8.2, 8.8, and 9.6; head length/snout length 4.3, 4.4, and 3.8; body depth/head depth
1.2, 1.9, and 1.6. These ratios, on the whole, agree more closely with those given by
LaMonte (1935) for G. cylindricus than for G. carapo. Only one row of teeth
can be seen in both the upper and lower jaws of each specimen.
24. Arius melanopus (Giinther). (III)
Station 3 (1, 173); new record.
The identifications of this and the next species are based solely on range
statements given in Miller's (1966: 795) checklist. This specimen was collected by
gill net on the night of 19-20 August 1963, and was the only one seen in Tortuguero
25. Bagrefilamentosus (Swainson). (III)
Stations 2 (1, 280), 3 (5, 218-360); new record.
Twenty-four additional specimens, ranging in length from 218 to 457 mm, and in
weight from 0.5 to 4.6 pounds, were also collected in the estuary in 1963 and 1964,
four of them during the dry season.
Of 19 stomachs examined, 11 contained food. The contents consisted mostly of
small crabs, as well as river and burrowing shrimps, small fishes, vegetable matter, and
some large tarpon scales. Darnell (1958) reports Bagre marinus to feed primarily on
blue crabs and penaeid shrimp, although it also takes fishes and various invertebrates.
One 457-mm female collected 29 April 1964 contained well-developed ovaries,
with eggs measuring from 1 to 17 mm in diameter. A 415-mm female, caught 29
July 1964, contained eggs up to 15 mm in diameter. McLane (1955) states that
in the St. Johns River, Florida, Bagre marinus spawns in brackish water from July
through September. Gunter (1945) says B. marinus in Texas breeds in early May and
carries the eggs and young orally until July.



26. Rhamdia wagneri (Ginther). (I)
Stations 31 (1, 41), 32 (3, 170.5-187), 35 (3, 30.5-66); new record.
The genus Rhamdia is badly in need of revision. Tortuguero specimens were identi-
fied, with Lopez-Sanchez' (1968) key as R. wagneri on the basis of the relatively long
adipose fin (less than 4.3 times in standard length), deeply bifurcated caudal fin, long
occipital process (measuring twice in distance from occipital base to dorsal origin), and
long maxillary barbel (extending past origin of anal fin). R. M. Bailey (pers. comm.)
informs me that this species may not be specifically distinct from R. guatemalensis


27. Alfaro cultratus (Regan). (II)
Stations 11B (2,24-28), 14 (6,21.5-49), 15 (2, 33-39), 17 (2, 15-27.5), 19 (89,15.5-
63.5), 20 (11, 14-52.5), 21 (3, 24-50), 31 (33, 24.5-60.5), 33 (11, 29.5-64), 35 (6, 20.5.
48); new record.
All specimens were collected in quiet sidewaters of tributary streams. Most were
taken in open water, and none were found in thick mats of vegetation. Females at-
tained a maximum length of 63.5 mm, whereas males were no more that 46.5 mm;
females outnumbered males by about five to one.
28. Belonesox belizanus Kner. Pike killifish. (II)
Stations 1 (7, 45-77.5), 2 (1, 32), 14 (2, 48-75.5), 17 (2, 45.5-64), 19 (4, 26-63),
20 (1, 98.5), 31 (2, 39.5-64), 34 (1, 99.5), 35 (3, 37.5-79).
According to R. R. Miller (pers. comm.), Sterba's (1962: 558) statement that females
of this species reach a maximum total length of 200 mm is in error. The largest specimen
present in the University of Michigan collection is about 140 mm, and females at
least as small as 73 mm have been found with embryos.
29. Brachyrhaphis parismina (Meek). (II)
Stations 11B (5, 21.23.5), 19 (6, 13.5-22), 21 (1, 36), 31 (5, 15-42.5); new record.
All but three of the above specimens were females. The three males (one from
station 11B and two from station 31) all measured between 18 and 20 mm. In general
Brachyrhaphis seemed to prefer somewhat weedier situations than Alfaro wherever
the two coexisted.
30. Phallichthys amates pittieri (Meek). (II)
Stations 1 (1, 34.5), 2 (6, 27-43), 12 (1, 28.5), 14 (3, 44.5-51), 19 (4, 17-45), 20
(3, 16.5-18), 31 (14,14-48), 33 (8, 34-48), 35 (2,24-26).
Several of the specimens collected in the lagoon came from under floating rafts of
water hyacinths. Although all specimens from the tributaries were taken in quiet
water, Meek (1914) gives the habitat of this species as small rocky streams, and
Alfaro collected specimens at Tiribi and Descampos, Costa Rica, at altitudes of 1200 m.
Apparently this species is able to live in a variety of habitats.
31. Poecilia mexicana Steindachner. (II)
Stations 1 (23, 21.5-82.5), 2 (136, 14-78), 19 (3, 27-45), 33 (1,40), 35 (1,46).
This is the only poeciliid that occurs regularly throughout the estuary, and in fact

Vol. XVI No. 1


it appears to be rather scarce in the tributary streams (only three collections (five
specimens) coming from the latter). It was present in the lagoon during both
seasons, 42 specimens (not listed above) being taken in one dry-season collection near
the lagoon mouth. This fish is often seen foraging along the shore in company with
Sphoeroides testudineus.
Caldwell, Ogren, and Giovannoli (1959) listed this species as Mollienesia sphenops,
but R. R. Miller, who is studying mollies, has reidentified the Tortuguero material as
P. mexicana. Poecilia mexicana and P. sphenops superficially are virtually
identical, but differ consistently in the nature of the inner tooth rows; these teeth
are unicuspid in mexicana, multicuspid in sphenops (Schultz and Miller, 1971).


32. Strongylura marina (Walbaum). Atlantic needlefish. (III)
Station 1 (2, 129-141).
Of the 14 specimens of this species Caldwell, Ogren, and Giovannoli (1959) list,
only two are actually S. marina; the remainder are S. timucu. These two individuals
were collected 11 July 1958 over the flats across from the village, together with
seven S. timucu. All subsequent collections of needlefish from the Tortuguero area
have proved to be timucu. Neither species was encountered in the tributary streams,
although both probably occur there.
Studies by Darnell (1958) indicate that small ashes and shrimp form the bulk of
the diet of S. marina.
33. Strongylura timucu (Walbaum). Timucu; "long guard." (III)
Stations 1 (26, 64-275), 2 (3, 168-204), 18 (4, 53-76); new record.
Although Berry and Rivas (1962) synonymized Strongylura timucu with S.
marina, Collette (1968) has shown timucu to be a valid species, differing
from marina in having 1) both left and right gonads developed instead of only the
right one, and 2) fewer predorsal scales (120-185 vs. 213-304). In addition timucu
typically is darker than marina: Melanophores usually extend to the ventral margin of
the orbit in timucu but not past the middle of the orbit in marina; the preorbital
bone is densely pigmented in timucu but only slightly so in marina; and melanophores
are present along the lateral line in timucu but are usually absent in marina.
As indicated above, timucu is much the commoner of the two species at


34. Hyporhamphus roberti hildebrandi Jordan and Evermann. (III)
Stations 1 (18, 70-186), 13 (1, 38), 18 (4,44-62), 25 (5,114-119).
This species shows a distribution at Tortuguero similar to that of S.'...\ 1. ,a


35. Citharichthys spilopterus Giinther. Bay whiff; sandfishh." (III)
Stations 1 (46, 13.5-95), 2 (8, 30-91), 11A (1, 76), 11B (2,44.5-57), 19(12, 36-39),


21 (3, 37.5-45), 22 (1, 64.5), 25 (19, 30-68.5), 28 (4, 46.5-63.5), 31 (6, 34.5-60.5),
35 (2,62-103).
This flatfish was extremely common in the estuary and tributary streams during
both wet and dry seasons. Reid (1955) found the food of C. spilopterus in Texas to
consist primarily of copepods.
36. Citharichthys uhleri Jordan. (III)
Stations 1 (5, 11-33.5), 25 (4, 26.5-32.5), 29 (1, 12), 30 (1, 13), 31 (13, 14-30); new
Citharichthys uhleri has, in the past, frequently been confused with C. spilopterus
(Erdman, 1967: unnumbered pl. 27), but may be distinguished from that species by a
number of characters. Adult uhleri have numerous diffuse whitish blotches on the
pigmented side of the body that are absent in spilopterus. Very small specimens of
uhleri (ca. 20 mm or less) are characterized by 6 dark, broad, irregular, interrupted
lines on the ocular side; 6 well defined, regularly-situated dark spots on both dorsal
and ventral parts of the blind side, with a 7th spot situated on the mid-side between
the posteriormost spots; and 5 dark spots on the middle of both the dorsal and anal
fins; spilopterus specimens of comparable size lack such markings. Finally, the two
species differ in both dorsal and anal-ray counts (Table 3).

TABLE 3. DORSAL AND ANAL FIN-RAY COUNTS IN Citharichthys spilopterus AND
C. uhleri
Dorsal rays
70 71 72 73 74 75 76 77 78 79 80 81 82 83 N
C. spilopterus 4 7 7 9 5 2 1 35 78.4
C. uhleri 2 3 7 4 1 17 71.9

Anal rays
50 51 52 53 54 55 56 57 58 59 60 61 62 N 5
C. spilopterus 1 7 10 8 5 4 35 59.6
C. uhleri 2 5 6 2 1 1 17 51.9

The failure of workers to differentiate between uhleri and spilopterus probably
accounts in large part for the presumed rarity of the former in collections ("known
only from the type," according to Gutherz (1967: 35). C. uhleri is very close to, and
perhaps not specifically distinct from C. arenaceus, which is also rare in collections.
Should these two prove to be the same, the name uhleri would have priority.
C. uhleri is an addition to Miller's (1966) checklist of Central American fresh-
water fishes.


37. Achirus lineatus (Linnaeus). Lined sole. (III)
Stations 1 (42, 12-56.5), 2 (1, 38.5), 25 (2, 38.5-88).
38. Trinectes paulistanus Miranda-Ribeiro. Southern hogchoker. (III)
Stations 1 (9, 17-27), 2 (2, 13-18), 19 (2, 43-53), 31 (2, 15.5-16).
This species appears to be less common than Achirus lineatus at Tortuguero.
Based on present collections, as well as overall knowledge of their comparative

Vol. XVI No. 1


ecologies, T. paulistanus occurs in completely fresh water much more frequently
than does A. lineatus. This species is found from southeastern Yucatan to Brazil,
where it is the allopatric replacement of the more northern T. maculatus. T. paul-
istanus differs primarily from maculatus in usually having five instead of four pelvic
rays on each side.


39. Symphurus plagusia (Bloch and Schneider). (*)
Station 16 (1, 79); new record.

40. Oostethus lineatus (Valenciennes). Opossum pipefish. (III)
Stations 1 (5,48-119), 2 (9,67.5-90), 11B (2,127), 14 (1,120), 15 (1,109), 17 (1,
126), 19 (1, 166.5), 21 (2, 122-145), 25 (2, 110-130), 30 (20,67-85, 33 (1, 130).
The authorship of this specific name has been questioned. Jordan and Evermann
(1896: 773) credit it to Valenciennes, based on the listing by Kaup (1856: 59) in
the original description of the species as "Dorichthys lineatus (Valenciennes
MS)." Caldwell, Ogren, and Giovannoli (1959:18) followed this decision; others
(Hubbs, 1929; Bailey et al, 1970: 23) attribute the species to Kaup. We follow Jordan
and Evermann for reasons Gilbert (1964: 129-130) has previously discussed.
0. lineatus, which ranges from Panama through the Greater Antilles to South
Carolina (Miller, 1966: 796), is more closely restricted to fresh and/or brackish
water than most species of northern-ranging pipefish (i.e., those occurring north of
the Yucatan Straits). Males with well-developed embryos were found both in
Tortuguero estuary and well up the tributary streams, indicating that the young
probably hatch in fresh water. However, the capture on the night of 7 August 1964
(sta. 30) of 20 small individuals swimming against the current at the mouth of the
lagoon suggests that early development of the species may occur in the ocean. McLane
(1955) found 0. lineatus only in the lower reaches of the St. Johns River, Florida, and
reported gravid females (males?) from there during the months of July and
Several small (65-80 mm) individuals of 0. lineatus were found in the stomach of a
254-mm jack (Caranx sp.) that had been feeding on the mass of small fishes and
invertebrates concentrated at the mouth of Tortuguero lagoon the night of 13 August
Caldwell, Ogren and Giovannoli (1959:18) report one specimen of this species
from Tortuguero (UF 5833), collected in 1957, to have only 18 tail rings, whereas the
other five individuals examined had 22 or 23. Subsequent counts on a total
of 34 Tortuguero specimens, plus four other specimens from Florida and Cuba,
are as follows: 22(8), 23(24), 24(6). Close examination of the single atypical specimen
shows no obvious malformation, and no other morphological characters could be found
to separate the two types. Additional material must be examined to clarify this
41. Pseudophallus mindii (Meek and Hildebrand). (III)
Stations 1 (4, 62-91), 19 (1, 90), 21 (1, 96).



42. Coleotropis blackburni Schultz. (III)
Stations 1 (2, 48-66.5), 4A (1, 76), 12 (1, 68.5), 13 (23, 31-65), 18 (94, 19-94.5),
29 (35, 15.5-25).
All but two specimens of this species were taken in the surf in water not over 1
or 2 feet deep. It was encountered only once in the estuary, at a spot not far from the
lagoon entrance. Two specimens were collected 13 August 1969 (during the wet
season) together with six individuals of Melaniris chagresi, the only time these two
species were found sympatrically.
Coleotropis colecanos, which Caldwell (1962) described from a single Tortuguero
specimen, has recently been synonymized with the present species (Gilbert and Cald-
well, 1967).
43. Melaniris chagresi (Meek and Hildebrand). (III)
Stations 1 (125, 28-109), 2 (147,25-103), 15 (1, 19), 21 (3, 53-58), 22 (1,80), 23
(1, 89.5), 28 (4, 48-62), 33 (2, 29-30.5), 35, (6,46.5-64.5).
This is one of the commonest species in the estuary and was also collected several
times in the tributary streams. One of the largest specimens, a female taken in late
April 1964, contained eggs averaging 1 mm in diameter. Individuals were commonly
seen swimming slowly at the surface along the shore both at night and during
the day. Ogren saw fish-eating bats (Noctilio) preying on it (Caldwell, Ogren, and
Giovannoli, 1959:23).

44. Agonostomus monticola (Bancroft). Mountain mullet; "califavor." (III)
Stations 2 (2, 28-31),.21 (2, 45-60), 35 (5, 30.5-37.5); new record.
Adult A. monticola characteristically occur in mountain torrents. Only small
individuals were found in Tortuguero lagoon, thus indicating that this species
spawns in the ocean, with the young later moving into fresh water, where they stay
until sexually mature. Anderson (1957) suggested a similar mode of reproduction,
also based on indirect evidence. Although this seems likely, it is also possible that the
species spawns in fresh water and the eggs are carried into the ocean, where they
45. Mugil curema Valenciennes. White mullet, "lisa." (III)
Stations 2 (8, 16-27), 18 (34, 22-36). 23 (4, 71-83), 25 (3, 23.5-95), 29 (1, 16).
This species is common in Tortuguero estuary throughout the year. Both young
and adults were present, the largest specimen seen (but not saved) measuring 369
mm. One large female (length not recorded) caught in May contained well developed
ovaries 150 X 20 mm. An unusually large number of mullet were seen at the pass
during the tismiche, but the stomachs of the several specimens examined were
empty. Darnell (1958) states that most Mugil species feed upon plankton, fila-
mentous algae, diatoms, and other minute vegetable matter, or upon organic de-
tritus and other such nutritive material as can be filtered from the bottom mud.

46. Polydactylus virginicus (Linnaeus). "Barbu;" "gourd fish." (III)

Vol. XVI No. 1


Stations 4A (1, 45), 16 (93, 50-122.5), 18 (79, 46-76.5), 29 (3, 45.5-48).
In addition to the above, three specimens (145-159 mm SL) were caught in the
estuary at the tagging camp in July 1963, a 178-mm female was taken at the camp in
May 1964, and three specimens 120-230 mm SL were taken at the boca, also in May
1964. The 178-mm female contained tiny eggs. Stomachs of five of the large speci-
mens were examined, of which three were empty and the other two contained a small
unidentified fish and three small river shrimp. Reid (1955) found the food of Polydac-
tylus octonemus in Texas to consist primarily ofpenaeid shrimps.

47. Sphyraena guachancho Cuvier. Guachanche. (*)
Station 16 (3, 56-62.5); new record.
Chuck Carr caught a 25-pound barracuda, possibly this species, in the lagoon on
23 April 1965.


48. Scomberomorus maculatus (Mitchill). Spanish mackerel. (*)
Station 16 (none saved); new record.
One specimen was caught by hook and line in the ocean in August 1963. Many
others were seen in the area.


49. Caranx hippos (Linnaeus). Crevalle jack; "jurel." (III)
Stations 1 (7, 31-48), 2 (5,255-310), 5B (2, 260-297), 18 (1, 39.5).
This species and the next are common in the estuary, where they seem to be
concentrated near the mouth. The stomach of a large (590 mm SL; 9.6 pounds)
individual contained some large fish vetebrae, sticks, and several large Macrobra-
chium shrimp. A 20-pound individual caught in the ocean in front of the tagging
camp contained a 254-mm Centropomus sp. Previous studies (Darnell, 1958) list
the food of C. hippos as mostly fishes and crabs, with some squid, shrimp, and other
50. Caranx latus Agassiz. Horse-eye jack; "jurel." (III)
Stations 1 (5, 43.5-51), 2 (6, 45-190), 25 (1, 110), 28 (2, 47.5-56.5), 29 (1, 94).
This was the only species of Caranx taken in the estuary during the wet season.
51. Chloroscombrus chrysurus (Linnaeus). Bumper. (III)
Station 24 (5, 170-191); new record.
The largest specimen contained eggs averaging 0.5 mm in diameter.
52. Oligoplites palometa (Cuvier). "Leathercoat." (111)
Stations 1 (8, 18-38.5), 2 (1,27), 5B (3, 284-323), 18 (32, 15-29), 25 (1, 70).
Stomachs of seven specimens, ranging from 284 to 385 mm SL, were examined.
Four of these were empty, one contained a large crab (probably Callinectes), another
a 50-mm Sphoeroides testudineus, and the other nine small shrimp. Females caught
in July 1964 contained eggs measuring from 0.5 to 1.0 mm in diameter.
0. palometa is an addition to Miller's (1966) checklist of Central American
freshwater fishes.


53. Trachinotus carolinus (Linnaeus). Pompano. (*)
Station 18 (2, 75-92); new record.
54. Trachinotus goodei Jordan and Evermann. Palometa. (*)
Stations 18 (7, 73-144), 29 (1, 111); new record.
This species was called Trachinotus glaucus (Bloch) until recently.
55. Vomer setapinnis (Mitchill). Atlantic moonfish. (*)
Station 16 (9, 27.5-36); new record.

56. Centropomus parallels Poey. Fat snook; "robalo."
Stations 1, (10, 65-104), 2 (4, 19-103), 25 (29, 41-69.5), 28 (1, 66.5), 31 (1, 60.5),
34 (1, 89).
Neither C. parallels nor C. undecimalis seems to be quite so common at Tortu-
guero as C. pectinatus, although neither is rare. The young (up to 100 mm) of all
three species occur over the estuarine flats (from July to September anyway), and
they have been taken in the same collection at least once (field no. G 63-26);
young of parallels and pectinatus also have been collected well up the
tributaries. In contrast to the above, adults of these species show distinct ecological
preferences. Adult pectinatus were found throughout the estuary as well as in the
streams, and many were taken with gill nets; the species seemed to become in-
creasingly more common away from the inlet. Neither parallels nor undecimalis
were caught in gill nets, nor were large specimens seen or collected outside the
boca area. Of the last two species, all positively identified individuals caught by
hook and line were undecimalis, and as all other specimens so collected were sub-
stantially longer than the largest individual (580 mm) of parallels recorded by
Rivas (1962: 60), it is likely that these were undecimalis also.
Tagging studies on undecimalis in Florida (Volpe, 1959) indicate that the
species is nonmigratory, and one may assume that this is true of other kinds of
Centropomus as well. If so, however, it is surprising that no intermediate-sized indi-
viduals of any of the three species were seen at Tortuguero. Local Tortuguero resi-
dents say that large numbers of mature snook move into the pass at certain times,
and Archie F. Carr (pers. comm.) observed such an "arrival" at the beginning of
the dry season in September 1964. This was concurrent with the arrival of schools
of clupeids upon which the snook were feeding. The specimen of C. ensiferus re-
ported by Caldwell, Ogren, and Giovannoli (1959: 24) proves, upon reexamination,
to be C. parallels.
57. Centropomus pectinatus Poey. Tarpon snook; "big bone." (III)
Stations 1 (41, 21.5-100), 2 (16, 37.5-148), 3 (6, 240-390), 15 (2, adults; not
saved), 17 (1, 93), 22 (1, 57.5), 25 (27, 38.5-78), 28 (1, 76).
Stomachs of 71 specimens gill-netted at night were examined, of which 36 con-
tained food. This consisted (in order of abundance) of river shrimp, small fish (up
to 80 mm SL), and crabs. Large numbers of parasites were found in the coeloms of
most large specimens of C. pectinatus. Dr. G. W. Hunter, III, identified these as the
larval stage of a tapeworm of the genus Gymnorhynca. An earlier stage lives in
crustaceans, and the adults parasitize elasmobranchs. What appeared to be the
same parasite was also found in Bairdiella ronchus.

Vol. XVI No. 1


58. Centropomus undecimalis (Bloch). Snook; "robalo." (III)
Stations 1 (9, 34.5-61), 3 (2, 385-485), 28 (5, 123.5-202.5); new record.
Caldwell, Ogren, and Giovannoli (1959) did not distinguish undecimalis from
parallels although they had both species in their collections. The two super-
ficially are very similar, but are readily distinguished by the presence, in undeci-
malis, of fewer total gill rakers (12 or 13 vs. 16 to 18, excluding rudiments), and
fewer scales in the first row above the lateral line (69 to 71 vs. 80 to 92) (above
counts based on Tortuguero specimens only). In addition, C. undecimalis grows
substantially larger.
Volpe (1959) says that Florida undecimalis spawn in saline passes during late
spring and early summer, an observation confirmed by Marshall (1958), who adds
that not all the eggs may ripen simultaneously. Females of undecimalis with well-
developed ovaries containing eggs 0.5 mm in diameter were caught at Tortuguero
on 19 July 1964, 8 August 1964, 12 August 1963, and 2 September 1963. Adult males
and females taken in the estuary in September 1964 were all in breeding condition.


59. Epinephelus (?) sp. (*)
New record.
A large grouper, 1340 mm SL and weighing 170 pounds, was caught at the pass
on 31 July 1964. The specimen was tentatively identified by Kelso as Epinephelus
itajara (Lichtenstein), but was not preserved and was not seen by the senior author.
The stomach contained several blue crabs (Callinectes), measuring 100 mm
across the carapace.


60. Lutjanus griseus (Linnaeus). Gray snapper. (II!)
Station 1 (7, 26.5-119); new record.
61. Lutjanusjocu (Bloch and Schneider). Dog snapper. (III)
Stations 1 (30, 18-150), 2 (5, 18.5-138), 25 (1, 67.5).
In addition to the small specimens present in most rotenone collections, large
individuals are commonly caught by hook and line in deeper water. Stomachs of
three specimens (560-620 mm SL and weighing from 9.5 to 12.4 pounds) caught in
July 1964 were examined, of which one was empty and two contained small crabs.
Stomachs of five smaller specimens (140-210 mm SL) poisoned near the airstrip
on 26 August 1964 contained many small Macrobrachium, one 25-mm river shrimp,
and several different species of fishes (Citharichthys spilopterus, Eucinostomus
pseudogula, Strongylura timucu, and small specimens of an unidentified goby).
Mature ovaries containing eggs 0.5 mm in diameter were found in a 380-mm female
collected 26 August 1964.


62. Conodon nobilis (Linnaeus). Barred grunt. (*)
Stations 16 (10, 55-143), 18 (6, 48.5-50.5), 29 (1, 58.5); new record.


63. Pomadasys corvinaeformis (Steindachner). (*)
Stations 16 (4,48-76), 18 (39, 65-91); new record.
This species, unlike P. crocro, does not appear to enter brackish or fresh water
around Tortuguero.
64. Pomadasys crocro (Cuvier). Burro grunt. (III)
Stations 1 (30, 32-50), 2 (2689, 13.5-154.5), 13 (50, 18-21.5), 18 (1, 19), 21
(3, 33-310), 25 (1, 54), 28 (2, 59-59.5), 31 (2, 7-12).
This is a very common species, both in the estuary and tributary streams. P. crocro
appears to spawn from July to September in the Tortuguero area, probably in the
estuary. Tremendous numbers of young encountered in the lagoon near the airfield
on 15 August 1963 probably had hatched from 2 to 4 weeks previously. Four ripe
females (275-380 mm SL) were collected between 17 and 26 August 1964 at the pass
and at the tagging camp. Stomach contents of four large specimens collected at the
pass 8 August 1964 included mostly post-larval shrimp, as well as some small fish
and crabs.


65. Diapterus olisthostomus (Goode and Bean). Irish pompano. (III)
Stations 1 (2, 25-89), 2 (49-56), 28 (1, 54.5).
This and the following four species have previously been referred to the family
Gerridae. Inasmuch as that family name is preoccupied in the class Insecta (order
Coleoptera), emendation of the name to Gerreidae is necessary (Bailey and Moore,
66. Diapterus plumieri (Cuvier). Striped mojarra. (III)
Stations 1 (3,210-265), 3 (1, not saved), 19 (1, 53).
Caldwell, Ogren, and Giovannoli (1959: 25) list this species as Eugerres sp.
67. Diapterus rhombeus (Cuvier). (III)
Stations 1 (12, 38-60), 2 (20, 29-62).
68. Eucinostomus argenteus Baird and Girard. Spotfin mojarra. (III)
Station 2 (1, 56): new record.
The only specimen was collected in the estuary near the airstrip 15 July 1962.
E. argenteus lacks the large black blotch found at the tip of the dorsal fin in E.
pseudogula, and has only seven (instead of eight) gill rakers on the lower limb of
the outer arch.
69. Eucinostomus pseudogula Poey. Slender mojarra. (III)
Stations 1 (98, 11.5-71.5), 2 (16, 22-62), 18 (15, 14.5-19), 20 (1, 14.5), 25
(3, 20.5-49.5), 29 (6,14-57), 31 (4, 18-25.5), 35 (1, 53.5).
This species may not be specifically distinct from E. melanopterus (Bleeker),
which was described from the eastern Atlantic (Miller, 1966: 797). If the two are the
same, the name melanopterus would have priority.

70. Bairdiella ronchus (Cuvier). "Pis-pis." (III)
Stations 1 (20,79-118.5), 3 (14,89.5-230), 28 (1, 122), 35 (3,92.5-119).
The absence of young individuals from Tortuguero collections may indicate that

Vol. XVI No. 1


this species either spawns outside the estuary and the young do not enter until
they have attained a certain size, or else it spawns in late fall or winter when no
collections were made. Of 37 stomachs examined, 14 were empty and the others
contained small fishes and river shrimp.
71. Bairdiella sanctaeluciae (Jordan). Striped croaker. (*)
Station 16 (1, 104.5); new record.
This essentially Caribbean species has recently been found along the southeastern
coast of Florida (UF 12031, CU 43321), and thus is one of the few southern conti-
nental fishes known north of the Yucatrn Straits (Robins, in press; Gilbert, in press).
72. Cynoscionj amaicensis (Vaillant and Boucourt). (*)
Station 16 (1, 65.5); new record.
Counts on this specimen are as follows: Dorsal rays, X-I, 26; anal rays, II-9; gill
rakers (not including rudiments), 2 + 8 = 10; pectoral rays, 17-16; lateral-line scales not
countable, but apparently relatively few in number (70-80).
73. Larimus breviceps Cuvier. (III)
Stations 3 (1, 115), 16 (242,49-132), 18 (34, 22-43.5); new record.
This species was the one most commonly collected in offshore trawling, but was
taken only once in the estuary. L. breviceps is another addition to Miller's (1966)
Central American freshwater fish checklist.
74. Menticirrhus americanus (Linnaeus). Southern kingfish. (III)
Station 1 (1, 40); new record.
This species, collected a short distance from the lagoon entrance, is an addition
to Miller's (1966) freshwater checklist.
75. Menticirrhus littoralis (Holbrook). Gulf kingfish. (*).
Station 18 (5, 19.5-135); new record.
These specimens were identified by Roy Irwin, who is revising the genus. M.
littoralis and M. americanus are two of the few northern continental species (Robins,
in press; Gilbert, in press) found south of the Yucatan Straits.
76. Micropogonfurnieri (Desmarest). (III)
Station 3 (7, 189-285).
The absence of young individuals from collections may indicate that spawning
occurs outside the estuary.
77. Ophioscion costaricensis Caldwell. (*)
Station 4B (2, 110-117).
This species, which was described from the above specimens, is very close to
0. brasiliensis Schultz, and may not be specifically distinct. Nevertheless a cursory
comparison by the senior author of the two types of 0. costaricensis with speci-
mens of 0. brasiliensis from South America seems to confirm Caldwell's (1958)
observation of a significantly smaller eye and wider interorbital space in costaricensis.
78. Ophioscion panamensis Schultz. (*)
Station 16 (14, 31-90.5); new record.
79. Stellifer colonensis Meek and Hildebrand. (*)
Station 16 (12, 42-130.5); new record.


80. Umbrina broussoneti 'Cuvier. (*)
Stations 16 (6, 86.5-129.5), 18 (1, 95); new record.
Gilbert (1966b) recently removed this species from the synonymy of U. coroides


81. Cichlasoma alfaroi2 Meek. (II) Figure 2a
Stations 14 (1, 48), 19 (2, 24.5-28.5), 20 (1, 28), 31 (5, 26-80.5); new record.
This species has been referred to in most publications as C. lethrinus Regan (see
Miller, 1966: 792). It was found only in the tributary streams, as were C. dowi, C.
rostratum, C. spilotum, and C. cfspilurum.
C. alfaroi is easily confused with rostratum, the two having very similar fin-
ray counts (Table 4) and pigmentation patterns (Figure 2a-b). They are most
readily distinguished by caudal-fin shape emarginatee in alfaroi, rounded in
rostratum), maximum size (alfaroi is smaller), and (in smaller specimens) by distribu-
tion of pigment at the base of the caudal fin. In alfaroi the pigment is concentrated
on the upper half of the peduncle, whereas rostratum has it more diffuse and evenly
distributed along the entire caudal base;

82. Cichlasoma centrarchus (Gill and Bransford). (II) Figure 3a
Stations 1 (6, 19-68), 2 (20, 14-80), 14 (2, 56-104), 15 (2, 54-99), 17 (3, 45-52.5),
25 (1,36), 34 (1,51.5), 35 (3, 68-93).
C. centrarchus appears to be the least common of the four cichlid species regularly
found in the estuary.

83. Cichlasoma citrinellum (Ginther). "Crana;" "mountain tuba." (II) Figure 2c
Stations 1 (120, 20.220), 2 (30, 21.5-92), 6B (1, 139), 9 (1, 111), 15 (1, 29), 17
(4, 38.5-105), 31 (1, 57), 33 (6, 24-52), 34 (1, 89).
Although collected in the tributaries, citrinellum appears to be much more
common in the estuary. Of the cichlids it ranked second only to maculicauda in
abundance. Stomach contents of 19 specimens caught in May 1964 near the airstrip
included "orange slime," well-mascerated plant material, mud, sand, and one small eel.

84. Cichlasoma dowi3 (Gunther). "Guapote." (II) Figures 4a-c
Stations 11B (2, 20-48). 14 (1, 14) (?), 19 (1, 30), 20 (4, 17-27.5), 21 (5, 21-85),
32 (1, 190), 33 (18, 22-29.5); new record.
The small (14 mm) specimen from station 14 (Figure 4c) is questionably identi-

SNew emendation; originally spelled broussonnetii but named for Pierre M. A. Broussonet, French

2 New emendation; originally spelled alfari but named for Dr. A. Alfaro, formerly director of the
National Museum of Costa Rica.

S3New emendation; originally spelled dovii but named for Capt. John M. Dow.

Vol. XVI No. 1


Cichlasoma AND Herotilapia
Anal rays

C. alfaroi
C. centrarchus
C. citrinellum
C. dowi
C. cf dowi
C. friedrichsthali
C. maculicauda
C. nigrofasciatum
C. rostratum
C. spilotum
C. cf spilurum
H. multispinosa


1 21




4 1

12 1

Soft rays

X X XI 6 7 8 9
5 7
1 11 3 1 11 3
14 1
12 19
1 1
1 1 11 3
12 3
1 12 2 6 6 3
6 7
1 2
3 1 2
2 2

C. alfaroi
C. centrarchus
C. citrinellum
C. dowi
C. cf dowi
C. friedrichsthali
C. maculicauda
C. nigrofascatum
C. rostratum
C. spilotum
C. cf spilurum
H. multispinosa

C. alfaroi
C. centrarchus
C. citrinellum
C. dowi
C. cf dowi
C. friedrichsthali
C. maculicauda
C. nigrofasciatum
C. rostratum
C. spilotum
C. cf spilurum
H. multispinosa

Dorsal rays
5 7
8 7

4 9

26 2

1 13 1
8 3 2


Soft rays

7 8 9 10 11 12 13 N
12 12
5 10 15
2 10 3 15
5 23, 3 31
1 1
5 9 1 15
8 6 1 15
1 10 4 15
5 8 13
1 2 3
1 2 3
2 2

Pectoral rays (both sides)

12 13



1 20

1 3

14 15 16 N
8 16 24
1 30
13 17 30
40 21 62
29 1 30
19 11 30
9 30
7 17 2 26
3 3 6
5 6

10 N


fied as C. dowi. It agrees closely in;color pattern with C. dowi specimens of compar-
able, or larger, size (Figures 4a-b), but differs in several meristic characters (Table 3).
The most notable of these is number of anal spines, which is two higher (IX) than
found in any of the other specimens examined (range V-VII). Both spinous and
soft dorsal-ray counts are low, but are within the range of variation for the species.
The number of pectoral rays is also low, but because of the difficulty in obtaining
accurate pectoral counts for very small cichlids, the values may actually be higher
than listed. All meristic data are within the range of the closely related C.
friedrichsthali, thus suggesting that the specimen may be that species. The smallest
individual of friedrichsthali examined (19 mm SL) has a color pattern very similar
to that of the 30 mm specimen illustrated in figure 4d, and markedly different from
that of the specimen in question. In friedrichsthali-the bar at the base of the caudal
fin is narrower and more diffuse, the bars along the sides of the body are much
narrower and extend uninterrupted between the bases of the dorsal and anal fins,
and the dark lateral blotch just posterior to the middle of the body is much less
prominent. Nevertheless it is possible that C. friedrichsthali may initially have a
color pattern very similar to that of C. dowi, but which (unlike dowi) undergoes a
marked change early in life (i.e., between 14 and 19 mm SL).
85. Cichlasomafriedrichsthali (Heckel). "Viejita." (II) Figure 4d
Stations 1 (24, 28-128), 2 (36, 21-153), 6B (1, 107), 7 (2, 124.5-168.5), 11A
(6,95-175), 14 (20,40-160), 15 (4,60-111.5), 17 (11,19-135), 19 (1,19, 20 (5,20-24),
21 (3, 54.5-96), 33 (28, 20-52.5), 34 (13, 70.5-146.5), 35 (11, 33-140.5).
This species appears to be equally common in the tributary streams and the
estuary during both dry and wet seasons. The stomachs of 18 individuals caught
near the airstrip in May 1964 were examined, of which 17 were empty and the other
contained a small unidentified fish.
The species name has frequently been misspelled friedrichstahli (R. R. Miller,
pers. comm. )For further remarks concerning this species, see above.
86. Cichlasoma maculicauda Regan. "Tuba." (II) Figure 3c
Stations 1 (251, 10-208), 2 (9, 27-191), 9 (2, 180-182), 11A (14, 21-195), 14
(1, 102), 17 (3,45-205), 22 (10, 10-11), 25 (5, 24.5-39.5), 33 (1,26), 34 (2,65.5-139.5),
35 (3,44-102:5).
C. maculicauda is the most abundant cichlid in the estuary, and is also very
common in the streams. The stomachs of several individuals collected in May 1964
contained plant material, mud, and sand. Concentrations of large individuals were
frequently seen beneath the outhouses bordering the lagoon, where they presum-
ably were feeding on fecal matter.
Numerous small (10-12 mm) individuals were collected in the estuary, and on
three separate occasions fish of this size were observed being accompanied or
guarded by large adults. This indicates that the species probably spawns in the
87. Cichlasoma nigrofasciatum (Ginther). (II) Figure 3b
Stations 11A (7, 14.5-61.5), 12 (1, 14.5), 14 (13, 36.5-76), 15 (5, 42-73), 17
(32, 11.5-63), 20 (1, 16.5), 31 (4, 50-69), 33 (3, 38.5-50.5), 34 (5, 45.5-68.5); new

Vol. XVI No. I



FIGURE 2. A.-Cichlasoma alfaroi, UF 16392, 27.8 mm SL. B.---C. rostratum, UF
11232, 32 mm SL. C.-C. citrinellum, UF 11088, 29 mm SL. D.-C. cf spilurum, UF
16396, 18.4 mm SL. All specimens from Tortuguero area.



FIGURE 3. A.-Cichlasoma centrarchus, UF 11060, 14 mm SL. B.-C. nigrofasciatum,
UF 16395, 16.5 mm SL. C.-C. maculicauda, UF 7170, 29 mm SL. D.-C. spilotum, UF
11173, 30 mm SL. All specimens from Tortuguero area.

Vol. XVI No. 1


FIGURE 4. A.-Cichlasoma dowi, UF 11171, 30 mm SL. B.-C. dowi, UF 17304, 23.5
mm SL. C.-C. cf dowi, UF 11227, 14 mm SL. D.-C.friedrichsthali, UF 11059, 30 mm
SL. All specimens from Tortuguero area.


FIGURE 5. A.-Cichlasoma cf spilurum, UF 11032, 57.5 mm SL, Tortuguero area.
B.-C. spilurum, UF 13984, 50.5 mm SL, Lake Izabal, Guatemala.

This species appears to be the commonest stream-inhabiting cichlid in the area,
and might well be included in the list of Tortuguero Cichlidae that are exclusively
confined to this habitat (see account of C. alfaroi). The one specimen found in the
estuary (station 12) was obviously a stray that had drifted down under a clump of
water hyacinths.
88. Cichlasoma rostratum (Gill and Bransford). (II) Figure 2b
Stations 11A (4, 17.5-208), 14 (1, 32), 17 (4, 31.5-159), 19 (1, 173), 31 (1, 51),
32 (2, 155.5-173.5), 35 (3, 127-142); new record.
89. Cichlasoma spilotum Meek. (II) Figure 3d
Stations 19 (2, 27.5-30), 31 (1, 59); new record.
90. Cichlasoma cfspilurum (Gunther). (II) Figures 2d, 5a
Stations 11A (1, 57), 20 (2, 14.5-18.5); new record.
These individuals differ from specimens of C. spilurum from Lake Izabal, Guate-
mala (the type locality), in having less distinct bars on the side of the body; a single
large vertically oblong blotch on the mid-side of the body (blotch indistinct or
absent in C. spilurum); and a small well-defined caudal spot (spot larger, lighter, and
more diffuse in C. spilurum) (Figures 5a-b). Fin-ray counts are very similar or
identical, as the following data demonstrate:

Vol. XVI No. 1


Dorsal spines Costa Rica, XVII (2), XVIII (1); Guatemala, XVII (1), XVIII
(3), XIX (1).
Dorsal soft rays Costa Rica, 9 (1), 10 (2); Guatemala, 9 (3), 10 (2).
Anal spines Costa Rica, IX (3); Guatemala IX (5).
Anal soft rays Costa Rica, 7 (1), 8 (2); Guatemala, 7 (4), 8 (1).
Pectoral rays Costa Rica, 13 (1), 14 (5); Guatemala 13 (6), 14 (4).
Either C. spilurum is a variable species with regard to color pattern, or the
Tortuguero specimens represent an undescribed sibling form.
91. Herotilapia multispinosa (Giinther). (II)
Station 2 (2, 80-91).
This species has not been taken at Tortuguero since 1955, despite many collections
from presumably favorable habitats.

92. Dormitator maculatus (Bloch). Fat sleeper. (III)
Stations 2 (35, 10-65), 6A (1, 75), 17 (2, 26.5-36), 22 (2, 11.5-22.5), 33 (1, 38.5).
D. maculatus was much less common in the 1962-1965 collections than in earlier
ones. This probably was the result of less intense collecting among water hyacinth
roots, where this species seems to be largely confined.
According to Sterba (1962: 750), D. maculatus reaches 250 mm TL. This may
be an exaggeration, for the largest specimen seen by us (from Volusia County,
Florida) was 146 mm TL (107 mm SL), and the largest (of 179 specimens) reported
by Meek and Hildebrand (1916: 355) was only 115 mm SL. Sterba (op. cit.) also re-
ported small fishes to be the main item of diet, but McLane (1955) found the
stomachs of Florida specimens to contain mostly plant material.
93. Eleotris amblyopsis (Cope). (III)
Stations 1 (116, 11.5-67), 2 (74, 19-67), 9 (1, 62), 14 (2, 38-46), 15 (13, 14-26.5),
17 (3, 13-27), 20 (6, 28-41.5), 21 (3, 26.5-34), 22 (1, 38), 25 (33, 8-63), 30 (many, ca.
9; see below), 31 (4, 43-51), 33 (28, 22-60), 35 (10, 25.5-54.5).
The young of both E. amblyopsis and E. pisonis were frequently found among
water hyacinth roots. Unlike Dormitator maculatus, these species were taken in other
habitat as well. Larval specimens of Eleotris were the commonest fish component
of the tismiche. Although specific identifications could not be made at this size,
both amblyopsis and pisonis were probably present.
E. amblyopsis was called E. isthmensis by Meek and Hildebrand (1916: 357-360).
It is readily distinguished from E. pisonis by its larger scales (44 to 54 versus 59 to 68
scales in lateral series and 12 versus 18 rows between bases of second dorsal and anal
94. Eleotris pisonis (Gmelin). Spinycheek sleeper. (III)
Stations 1 (1, 29), 2 (2, 87-280), 11B (1, 30.5), 20 (1, 36), 21 (3, 29-108), 25
(1, 110), 30 (many, ca. 9; see above), 31 (1, 73.5), 33 (1, 93).
95. Gobiomorus dormitor Lacepede. Bigmouth sleeper; mudfishh."
Stations 1 (11, 34-222.5), 2 (18, 21-101), 17 (1, 140), 20 (1, 185), 21 (4, 82-104),
25 (3, 35.5-55.5), 31 (2, 171-187.5), 33 (1, 101), 35 (3, 57.5-190).
The stomachs of three specimens collected in May 1964 contained river shrimp
10 to 25 mm long. Gravid females were collected in the estuary the same month.


96. Leptophilypnusfluviatilis Meek and Hildebrand. (IlI)
Stations 1 (2, 31-35), 2 (4, 34-42).


97. Awaous tajasica (Lichtenstein). River goby. (III)
Stations 19 (2, 59.5-137), 30 (7, 12.5); new record.
Although larval individuals of this species were not one of the more common
components of the tismiche, many more probably were present than the above
collections indicate (Sta. 30).
98. Bathygobius soporator (Valenciennes). Frillfin goby. (III)
Stations 1 (13, 22-70), 2 (2, 48-58), 25 (7, 16.5-47.5), 35 (1, 37.5); new record.
This species is an addition to Miller's (1966) Central American freshwater fish
99. Evorthodus lyrics (Girard). Lyre goby. (III)
Stations 1 (11, 9-35), 2 (1, 37), 14 (1, 28).
100. Gobionellus boleosoma (Jordan and Gilbert). Darter goby. (III)
Stations 1 (16, 16.5-33), 2 (1, 15.5), 25 (5, 15.5-19.5), 31 (15, 17.5-28); new
This species was present in pre-1959 collections, but was not distinguished from
Gobionellusfasciatus (= claytonii) by Caldwell, Ogren, and Giovannoli (1959: 28).
101. Gobionellus fasciatus (Gill). (III)
Figure 6b
Stations 1 (55, 11-69), 2 (23, 19.5-49), 14 (7, 25.5-42.5), 17 (2, 23.5-25), 19
(5, 46.5-62.5), 20 (1, 12.5), 25 (6, 32.5-60), 30 (many, ca. 10), 31 (42, 18-46.5), 33
(3, 33-48), 35 (1,25.5).
G. fasciatus was called G. claytonii (Meek) by Caldwell, Ogren, and Giovannoli
(1959: 28). Unpublished studies by the senior author and John E. Randall indicate
that the two are valid, though very closely related, species, apparently differing
only by the presence in fasciatus of a well-defined cheek blotch and more completely
scaled belly. G. claytoni is restricted to the southwestern Gulf of Mexico, whereas
G. fasciatus occurs throughout the southern Caribbean, having been recorded from
Costa Rica to Dominica. Although future collecting may decrease this distributional
hiatus, there seems little doubt that the ranges of the two species are widely
102. Gobionellus pseudofasciatus Gilbert and Randall, new species. (III)
Figure 6a
Stations i (6, 17-35), 19 (1, 44), 31 (2, 27-31.5), 33 (1, 34.5); new record.
A diagnosis is presented here to establish the scientific name of this species. A
more detailed description will appear in a later paper by the senior author and
John E. Randall.
DIAGNOSIS. A species of Gobionellus with large scales (31 to 33 in lateral series),
usually 17 pectoral-fin rays (occasionally 16 or 18), and the combination of 12 total
second dorsal and 13 total anal elements. Differs from G. fasciatus, which it
most closely resembles, in having 1) a dark blotch of pigment extending from
lower posterior corner of cheek anteriorly at about a 45 angle and which, if

Vol. XVI No. 1



FIGURE 6. A.-Gobionellus pseudofasciatus, female, holotype, UF 13516, 34.7 mm
SL, Tortuguero area. B.-G. fasciatus, female, UF 5804, 36.3 mm SL, Tortuguero area.

continued, would pass just anterior to eye (blotch confined to posterior corner of
cheek in fasciatus); 2) small, dark discrete spots in the proximal part of most anal-fin
membranes of males (spots larger and more diffuse in male fasciatus); 3) no spots
in anal-fin membranes of females (diffuse spots present in female fasciatus); 4) no
diagonal line extending from postero-dorsal margin of maxillary to below and be-
hind eye; 5) the pectoral fins with four or five narrow, slightly diagonal bars (bars
absent in fasciatus); 6) the upper half of the body lighter; and 7) a small (in
females) to moderately large (in males) posteriorly-directed canine tooth in anterior
part of each lower jaw (absent infasciatus).
HOLOTYPE. University of Florida (UF) 13516 subadultt female, 35.0 mm SL,
illustrated), west side of Tortuguero lagoon, just below point across from Tortuguero
village, Lim6n Province, Costa Rica (station 1), 25 August 1963, C. R. Gilbert and
R. Kaufman (field no. G 63-36).
PARATYPES (Costa Rica-Tortuguero) UF 13517 (1), University of Miami, Insti-
tute of Marine Science (UMML) 23117 (1), U.S. National Muse m (USNM) 201589
(1) (3, 17.0-25.3), same data as for holotype. UF 13518 (2, 17.2-24.5), same locality.
as for holotype, 18 August 1963, C. R. Gilbert et al (field no. G 63-27). UF 13519
(1, 44.2), station 19. UF 13520 (1), Academy of Natural Sciences of Philadelphia
(ANSP) 109179 (1) (2, 27.1-31.6), station 31. Florida State University (FSU) 17695
(1, 34.5), station 33. (Guatemala) USNM 114338 (10, 29.2-53.0), Jicotea Creek,
tributary of Rio Sarstoon, ca. 14 mi. above mouth, 30 April 1947, Midence, Miller,
and Holloway (field no. M 47-51). (Canal Zone-Mindi Cut) USNM 81824 (1),
USNM 205202 (1) (32.3-35.3), 14 Jan. 1911, Meek and Hildebrand. USNM 105109 (1),
USNM 123264 (1) (29.7-32.2),28 Jan. 1911, Meek and Hildebrand. (Trinidad) Univer-
sity of Puerto Rico (UPR) 2225 (1, 28.5), first irrigation ditch on hwy. south of
Caroni River, 9 May 1964, J. E. Randall.


103. Gobiosoma spes (Ginsburg). (III)
Stations 1 (216, 14-34), 2 (9, 16.5-25), 31 (3, 18.5-23.5).
G. spes was taken in Tortuguero estuary only during the wet season, with
nearly all specimens coming from the west shore. Except for several individuals
taken in shipworm holes in a piling at the tagging camp, all individuals were found
living in mats of filamentous algae or other submergent aquatic vegetation.

104. Echeneis naucrates Linnaeus. Sharksucker. (III)
Stations 2 (1, 174), 3 (5, 110-370), 5A (1, 155), 5B (2, 111-170).
The presence of E. naucrates in fresh water is undoubtedly related to the
ability of Carcharhinus leucas (to which the sharksucker commonly attaches it-
self) to move back and forth between fresh and salt water. This species is an
addition to Miller's (1966) Central American freshwater fish checklist.

105. Dactylagnus peratikos B6hlke and Caldwel,-(*)
Stations 4C (1, 44), 12 (5, 48.5-53.5), 29 (1, 49).
This species is still known only from the above Tortuguero specimens.

106. Microdesmus carri Gilbert. (III)
Stations 1 (2, 36.5-72), 25 (1, 54.5), 30 (34, 28-33.5); new record.
M. carri, like the preceding species, is known only from Tortuguero mater-
ial. The fact that both forms are burrowers, and thus not readily collected by con-
ventional means, probably accounts in large part for their apparent rarity.
All specimens of carri collected during the tismiche (Station 30) are postlarvae;
the other three are adults, or at least show adult characteristics. As the postlarvae
presumably were moving into the estuary at the time of capture, spawning and
early development are hypothesized to have occurred in the ocean (Gilbert, 1966a:
331). M. carri is an addition to Miller's (1966) Central American freshwater fish

107. Batrachoides gilbert Meek and Hildebrand. (III)
Stations 1 (1, 122.5), 25 (4, 31-151); new record.
B. gilbert is readily distinguished from the other two species of Batra-
choides occurring in the southwestern Caribbean (B. surinamensis (Bloch and
Schneider) and B. manglae Cervigon) by number of dorsal and anal rays. Counts
for the above six specimens of B. gilbert are dorsal soft rays 24 (3), 25 (1), and
26 (2); and anal rays 22 (2) and 23 (4). For B. surinamensis and B. manglae,
respectively, dorsal counts are 28 or 29 and 22 or 23, and anal counts 25 to 27 and
20 (data from Meek and Hildebrand, 1928: 914-915; and Cervigon, 1966: 858 re-

Vol. XVI No. 1


108. Batrachoides surinamensis (Bloch and Schneider). (III)
Stations 3 (1, 285); new record.
This specimen was caught by hook and line just inside the lagoon in July
1965. Dorsal and anal ray counts are 29 and 27 respectively.
109. Porichthys pauciradiatus Caldwell and Caldwell. (*)
Station 10 (1, 65.5); new record.
This species, recently described from Panama (Caldwell and Caldwell, 1963),
is known from Costa Rica, Panama, and Brazil. The single Brazilian specimen
differs in several pigmentary features from other known specimens (Gilbert,
1968: 717-719), and may prove to be a nameworthy form.
110. Porichthys plectrodon Goode and Bean. Atlantic midshipman. (*)
Station 18 (1, -); new record.
This specimen was found dead on the beach, the posterior part of the body
partially eaten by crabs.
Until now this species has been called Porichthys porosissimus (Valenci-
ennes). It has recently been found that P. porosissimus is confined to the south-
western Atlantic from southern Brazil to Argentina, and that populations occur-
ring from central Brazil northward are P. plectrodon. The two species are very
similar in color pattern, but differ trenchantly in shape of the sagitta (otolith).
In porosissimus this structure is identical to that shown by Gilbert (1968: 679,
fig. 2d) for the eastern Pacific P. myriaster, whereas the sagitta indicated as be-
longing to "P. porosissimus" (op. cit., fig. 2a) is actually from P. plectrodon. In
addition porosissimus has a significantly higher average gill-raker count (ca. 15 to
13) (op. cit., p. 700, Table 4), although there is considerable overlap. Finray
counts are very close, with the range of counts for plectrodon completely over-
lapping those given for porosissimus (op. cit., Tables 1-3).

111. Sphoeroides testudineus (Linnaeus). Checkered puffer; "mutros;"
"pesisapos." (III)
Stations 1 (81, 5-115.5), 2 (17, 70-124), 15 (1, 83), 35 (1, 100).
This species was most abundant over the mud flats along the west shore. It
was common during both wet and dry seasons, although no specimens taken dur-
ing the dry period were saved. S. testudineus is primarily a scavenger, and
was observed feeding on a wide variety of detritus and garbage. Despite its rela-
tive abundance, only one individual was found in the stomach of a predator
(Oligoplites palometa).


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Vol. XVI No. 1


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Vol. XVI No. 1

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