20ol.J. Linn. SOL.,48, pp. 255-302. With 3figures
May 1969
The cichlid fishes of Guyana, South America, with notes on their
ecology and breeding behaviour
ROSEMARY H. LOWE-McCONNELL, F.L.S.
Streatwick, Streat, Hassocks, Sussex
Accepted for publicationJanuary 1969
Field studies in Guyana, north-eastem South America, from 1957 to 1962 included observations on the ecologyand behaviour of cichlids in natural waters. Collections made were examined
at the British Museum (Natural History) together with other cichlid material from Guyana.
This paper records these observations and gives keys for the field indentification of these cichlids.
Fifteen genera comprising at least 35 species occur in Guyana, mostly species with a wide
distribution in tropical South America. In the predatory CichIa ocellariswhich grows larger than
other South American cichlids, growth continues long after the fish matures. Males tend to
be larger than associated females. The behaviour of Cichla ocellaris and Cichlasoma festivum
when guarding their young suggests that the caudal ocellus has a vital rBle in the orientation of
young to parent. Most of these cichlids in which breeding habits are known are substratumspawners. An attachment gland, producing viscous threads, was found on the heads of recently
hatched Geophagusjurupari young, although this species carries the young in the mouth.
CONTENTS
.
.
Introduction
.
. .
Studies on Guyana cichlids
.
. .
Ecological conditions in the areas studied .
. .
General behaviour of South American cichlids .
.
.
Field key to the genera of cichlid fishes found in Guyana
Notes on the status, occurrence, and natural history of Guyana cichlid species
Cichla Schneider .
Key to Cichla species found in Guyana
.
. .
Chaetobranchus Heckel .
Acaroma Myers
.
Aequidats Eigenmann & Bray .
Field key to Aequidens species found in Guyana .
. .
Acarichthys Eigenmann .
Geophagus Heckel .
Key to Geophagus species found in Guyana.
.
Biotodoma Eigenmann & Kennedy
Cichlasoma Swainson
.
- . .
Field key to Cichlasoma species found in Guyana
. .
Petenia Gunther .
Crenicara Steindachner
. . .
Batrachopr Heckel
. .
Crm'cichhHeckel
PtemphyllumHeckel
Dwarfcichlids
Acknowledgements
References
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PAGE
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256
ROSEMARY
H. LOWE-MCCONNELL
INTRODUCTION
Studies on Guyana cichlids
The percoid family Cichlidae includes such excellent food fishes as Tilapia which are
now widely distributed throughout the tropics. With the exception of one genus in
India and Ceylon (Eutroplus) and five endemic genera in Madagascar (Kiener &
MaugC, 1966) cichlids are indigenous only to South and Central America and to Africa
and the Near East. Many of the African species support important fisheries and their
ecology has been much studied in recent years, but very little is yet on record about the
ecology and behaviour of South American species in natural waters of northern South
America. The life histories of some of the small species are well known to aquarists,
for cichlids are characteristically still-water fishes which can be bred in aquaria, but
little is known of their natural history in the field ;the larger species, though often an
important source of food for local peoples, have been even less studied.
Field studies made whilst I was living in Guyana (formerly British Guiana), in the
north-east of South America, from 1957 to 1962, included some observations on the
ecology and behaviour of cichlids in natural waters. Collections made during this work
were brought to the British Museum (Natural History) London, where they were
examined together with cichlid material from Guyana already in the collections, some
of which formed the basis of Regan’s taxonomic studies of South American cichlids
(1905-1913), and material collected since, mainly by Carter in 1934. The aim of this
present paper is to record these field observations and to give keys for the field identification of Guyana cichlids which it is hoped will stimulate further work on these fishes.
These studies were mainly on the larger species, food fishes and potential food fishes,
so little attention is paid here to the ‘dwarf cichlids’ which can be studied in aquaria,
and which are greatly in need of taxonomic revision.
Previous studies of South American cichlids have been mainly taxonomic or observations on small species in aquaria. In addition to Regan’s papers (1905-1913), taxonomic
studies of Guyana cichlids are included in Eigenmann (1912) and Fowler (1914), and
of those from neighbouring territories by Schultz (1949) from T‘enezuela, Boeseman
(1952, 1954, 1956) from Surinam and Puyo (1949) from French Guiana; Fowler
(1954) gives a check list of cichlids from Brazil. Puyo includes notes on the mores of
some species and Eigenmann on ecological groupings, but these other papers are
purely taxonomic. Descriptions of colours, and in some cases of breeding habits, of
cichlids in aquaria are given in aquarium handbooks such as Axelrod & Schultz (1955),
and by Sterba (trans. Tucker, 1962). Most of the cichlid species found in Guyana are
widely distributed in South America and some have been studied by departments
interested in fish culture in neighbouring countries (Fontenele, 1950; Braga, 1953).
The British Museum (Natural History) has many of the types described by Regan
and co-types of Eigenmann’s material. Eigenmann’s (1912) studies were made mainly
on fishes from the Essequibo River and its tributary the Potaro below and above the
800-foot-high Kaieteur Falls, and from waters of the coastal plain. Carter collected
from the great rivers of the forest area, the Cuyuni and Mazaruni, both tributaries of
the Essequibo, and also from the Demerara River (Fig. 1.) The present studies were
CICHLID
FISHES
OF GUYANA
257
'2
FIGURE
1 . Map of Guyana, showing places mentioned in the text.
258
ROSEMAF~Y
H. LOWE-MCCONNELL
concentrated in two areas :(1) the Rupununi savanna district in the south-west bordering
Brazil, where the Essequibo and Amazon drainage systems are in contact (LoweMcConnell, 1964), and (2) on the freshwater trenches and swamps of the coastal plain
behind Georgetown. These collectionswere therefore complementaryto the earlier ones.
Rather scattered observationswere alsomade during travels to other parts of the country.
The fish fauna of much of Guyana is still unexplored.
Much of the work on African cichlids has been on the systematics of the large flocks
of endemic species found in the Great Lakes of East and Central Africa (Fryer, 1959a,
b; Greenwood, 1964). The cichlids found in Guyana, on the other hand, are probably
none of them endemic to Guyana, and most of them are representatives of species and
genera widely distributed over north-eastern tropical South America. In this they
resemble the cichlid faunas of rivers of Africa, which have far fewer and more generalized and widely distributed species than do the lake faunas. The riverine faunas of
Guyana are, however, richer in numbers of cichlid genera (about fifteen) than are the
riverine faunas of Africa-where the Congo for instance carries about ten, the Zambezi
about five genera.
Regan (1906 : 238) in a diagram summing up his view of the probable relationships
of the South American cichlid genera indicates that Cichluand Chaetobranchusprobably
came from a basic stock leading to an Acara ( = A e p i k ) group. From this group several
evolutionary lines with three anal spines diverged : one leading through Crenicara to
the long-bodied Batrachops and Crenicichla, all with finely serrated preoperculars ;
another line to Geophagus and Heterogramma (=Apistogramma) with lobes on the first
gill arch ; and three distinct lines to Nannacara, to Acaropsis (=Acaronia), and to
Astronotus. Regan envisaged that from the Acara group, with its three anal spines, arose
the Cichlaoma group with more than three anal spines, and that from Cichlasoma six
independent lines diverged, one to Petenia, ahother (on the C. severum line) to Pterophyllum and Symphysodon (both genera present or likely to be found in Guyana), and
other lines to Uaru, to Heterotilapia, to Iierichthys and to Neotroplusand Paraneotroplus
(genera not represented in Guyana).
Regan (1920) later considered the character of most importance in the classification
of cichlid genera to be the structure of the skull apophyses supporting the upper
pharyngeal bones. The African Cichlidae may be divided into those with the upper
pharyngeal apophysis formed by the parasphenoid only (Tilapia type), and those in
which the apophysis is formed by the parasphenoid in the middle and basioccipital at
the sides (Haplochromistype). According to Regan (1920: 34) all the American cichlid
genera with the exception of Cichlu have the apophysis formed by the parasphenoid
only, as in Tilapia and the condition in Cichla resembles Haplochromis.
Wickler (1963) has questioned the importance of the structure of the apophysis
since groupings based on types of breeding behaviour do not altogether conform to
groupings based on this character, but Greenwood (1965) produced evidence in
aquarium-raised Astatoreochromis that the ba’sic structure of a ‘Haplochromis type’
apophysis persisted despite considerable atrophy of the pharyngeal apopophysis
(presumably due to relative disuse and altered diet). Greenwood also pointed out that
the examples of convergence in breeding behaviour are well known in the family. For
example, oral incubation must have arisen several times-the genus Tilapiu has both
CICHLID
FISHES
OF GUYANA
259
substratum-spawning and oral-brooding members, and this is probably also the case
within the South American genus Geophagus.
As this present study was restricted to cichlid species present in Guyana, no attempt
is made here to reconsider the evolutionary relationships of the genera ; for this,
consideration of the whole South American cichlid fauna would be needed. Material
from other parts of South America is available at the British Museum (N.H.) for many
of the species considered here, but was only examined when Guyana material was
scanty. Species described from neighbouring territories and not yet recorded from
Guyana but which are likely to be found there when the rivers are further explored, are
referred to after the keys based on Guyana species.
Future collectors must beware of aquarium escapes ; aeroplane loads of tropical
fish from Brazil en route to aquarists in North America and elsewhere now touch down
in Guyana for the water in the fish boxes to be changed, and the possibility now exists
that exotic species from much further south may find their way into the waters of
Guyana in this manner.
Ecological conditions in the areas studied
The ecological conditions prevailing in the waters of the Rupununi savanna district
have already been described (Lowe-McConnell, 1964). On the coastal plain the fresh
waters studied included the complex of man-made trenches near Georgetown (6"50' N,
58"W) used for irrigation and drainage of the sugar and rice fields ;trenches generally
2 m wide and 1 m to 1.5 m deep, supplied with water from the coastal rivers diverted into
vast swampy water conservancies. The aquatic plant communities of these trenches
have been described by Fanshaw (1954) ;many of them have a Hydrocleis- Utricularia
community with floating Salvinia, Azolla, Pistia, Neptunia, Eichhornia, several species
of Nymphaea waterlilies and submerged Cabomba, Elodea and Nitella. Besides cichlids
these trenches carry a rich fauna of characoids, gymnotoids and some nematognath
catfishes.
Wiremesh traps set throughout the year April 1957 to May 1958 in these trenches
took 529 fishes, of which 73% were cichlids belonging to six species: Cichlasoma
bimaculatum (lOS), Acaronia nassa (lol), Cichlasoma festivum (81), Cichla ocellaris (71),
Crenicichla saxatilis (8), and Geophagus jurupari (1). The non-cichlids trapped with
them included the characoids Hoplias malabaricus (75 of this large predatory species),
Acestrorhynchus microlepis (8), Metynnis sp. and Pygopristis denticulatus (lo), Serrasalmus rhombeus (13) and S . 'niger' (27), also the catfishes Hypostomus (=Plecostomus)
sp. (27), Hoplosternum littorale (13) and Rhamdia sebae (13).
When Eigenmann (1912) drained one of these trenches for the first time after 20
years, the 23 fish species present included only four species of cichlid : Cichlasoma
bimaculatum (17), C .festivum (12), Geophagus jurupari (1) and Crenicichla saxatilis (3)
(no Cichla ocellaris or Acaronia nassa) ; the majority of the 2254 fishes taken were small
characoids, many of them of a size which would not have been retained in the wiremesh
traps used in our experiments, but three species of gymnotoid fish, a nandid and six
species of catfish were also present (Eigenmann, 1912: 88). The predatory characoid
Pirai or Piranha, Serrasalmus, have evidently gained access to the fresh waters of the
260
ROSEMARY
H. LOWE-MCCONNELL
coastal plain since Eigenmann’s day, reputedly when the high floods of the 1930’s
allow them to enter the Lama water conservancy which feeds these trenches from the
Mahaica River.
Carter (1934,1935) has described some of the ecological conditions in the areas from
which his collections came ;he stressed the differences between the ‘white’ and ‘black’
water rivers of Guyana, which are so different chemically, and also the problems of
de-oxygenation which so many fishes have to face here. Recent studies suggested that
cichlids living in small pools are particularly exposed to such problems (LoweMcConnell, 1964). Carter (1935) noted that some Aequidens (especially A. potaroensis
Eig. and A . tetramerus (Heckel)) were common in swamps in Guyana, together with
occasional Crenicichla lugubris (Heck.) and C . alta Eig. When studying the respiratory
adaptations of these South American fishes in Guiana and Paraguay, Carter did not
mention any special accessory respiratory organs in cichlids, but in Paraguay Carter
and Beadle (193 1) noted that Acara (=Aequidens)portulagrensisHeckel was among those
species which have the habit of lying just below the water surface for many minutes at
a time drawing water over the gills from the very shallow oxygenated layer at the surface.
During the present studies the hardiness of these South American cichlids when
out of water was often noted. Cichlasoma were often still alive on market slabs; these
fishes and Crenicichla species remained alive for half an hour or so out of water in a
damp cloth. These fishes often had the stomach distended with air ;the stomach walls
are well vasculated, and it is suspected that the stomach can be used as an accessory
respiratory organ in species such as Cichlasoma bimaculatum. This needs experimental
verification. Stomachs were often empty of food when the fish were crowded into small
pools at the end of the dry season, the time when an accessory respiratory organ was most
needed, and when food might be scarce.
General behaviour of South American cichlids
Cichlids generally breed in still, rather than running, water. They are diurnal fishes,
with large eyes and in which the colours and colour patterns have been shown to be
important for species recognition and in breeding behaviour (Baerends& Baerends-van
Roon, 1950), and often hiding away motionless by night (Lowe-McConnell, 1964).
They all, as far as is known, practice some form of parental care of the eggs and young
and they have been the subjects of much behaviour work in aquaria in many parts of
the world. Typically the cichlids of the African Great Lakes are mouth-brooders in
which one parent (more often the female) broods eggs and young (Lowe, 1959 ;Greenwood, 1964), although in a few species both parents may guard the eggs and young.
The American cichlids, on the other hand, are characteristicallysubstratum-spawners,
in which eggs and newly hatched young are attached to a substratum and guarded by
both, or one, parent. This pattern is also shown by the Indian genus Eutroplus (Sterba,
1962: 724), the wide distribution of this pattern suggesting that it is the more basic
pattern of behaviour. A few American species are known to carry the young in the
mouth, but adaptations for oral-brooding are not as complete as in the African species ;
in Geophagusjurupari studied here, for example, the recently hatched young had an
attachment gland on the head, as in substratum-spawners. Sexual dimorphism or
CICHLID
FISHES
OF GUYANA
261
dichromatism is generally much less extreme in these ‘guarding’ species than in the
African mouth-brooders, in which the female remains drab while the male may attain
a striking breeding dress associated with defence of a spawning territory. T h e males
are, however, often slightly larger than the females with which they spawn in both
cases (Fig. 3) and may have elongated dorsal and anal fins. I n the guarders the very
young have cement glands on the head with which they are attached to the substratum
(Fontenele, 1952, and pers. obs.).
T h e present field studies indicated the importance of the caudal ocellus or black
spot in helping the young to orientate to the parent in at least two species-Cichla
ocellarisand Cichlasomafestivum. Such a caudal spot is present in many South American
cichlid genera (e.g. Cichla, Cichlasoma, Aequidens, Crenicichla, Acaronia) in which the
young are, or may be, guarded by, and follow, the parents, but the caudal spot is
absent or reduced in most mouth-brooders. (In the African Tihpia which are guarders,
T . zillii and T. melanopleura, the ‘Tilapia spot’ on the hind end of the dorsal fin is
particularly well developed in breeding fish and may perhaps serve the same function.)
This requires experimental verification.
T h e behaviour of the cichlids at night, hiding away motionless against a rock face,
generally rather near the surface, or amongst vegetation, has been described (LoweMcConnell, 1964); the fish appear to be sleeping and are not disturbed by light. They
could be lifted out of the water with a dipnet by night, staying completely motionless
till the last minute, then shooting away with a sudden tremendous spurt unless captured,
the fishes next to the disturbed fish remaining motionless. Amerindian fishermen take
advantage of this habit to catch cichlids by night. T h e fishes’ complete immobility
may help to conceal them from the large nocturnal predators such as catfish which
probably hunt by detecting water vibrations and chemical disturbances.
Many of these South American fishes, including cichlids, make noises ;for example
Geophagus surinamensis could be heard, and felt, grating its pharyngeal teeth. Myrberg
(1965) studied sound production in the cichlids Hemichromis bimaculatus, Cichlasoma
nigrofasciatim and Pterophyllum sp. in aquaria and concluded that sound is associated
with aggressiveness in these species.
FIELD KEY TO THE GENERA OF CICHLID FISHES FOUND IN
GUYANA
I n the following keys scale counts are taken along the body below the upper lateral
line. Fish lengths quoted are standard lengths (without caudal fin) unless otherwise
stated; lengths given thus ‘24/26 cm’ are standard/total lengths, to bring data into line
with those used in fishery investigations. Many of the fishes in these South American
waters have pieces bitten from the caudal fin, possibly the work of Serrasalmus, which
renders total lengths unsatisfactory here. ‘Material available’ signifies specimens now
in the British Museum (Natural History), and collected personally during these
studies unless otherwise stated. Only the major references to synonyms in the Guyana
area are indicated for each species. T h e general appearance and significant markings
of some of these fishes are shown in Fig. 2.
262
ROSEMARY
H. LOWE-MCCONNELL
M
3
FIGURE
2. The general appearance and significant markings of some Guyana cichlids: A,
Cichla ocellaris; B, Acaronia nassa; C , Cichlasoma bimaculatum; D,Cichlasoma festivum; E,
Batrachqps punctulatus; F,Crenicichla alta; G , Crenicichla shigata; H, Aequidens tetramerus;
I, Aeqtcidens vittatus; J, Aequidens potarom’s; K,Aequidm geayi; L,Acarichthys heckelii; M,
Geqphagus jurupari; N, Biotodoma cupido. (For sizes see text.)
CICHLID
FISHES
OF GUYANA
263
Kgt
la. First dorsal and anal branched rays prolonged; body deep and much
. Pterophyllum (p. 297)
compressed .
b. First dorsal and anal branched rays not especially prolonged .
. 2
2a. Gill rakers very long and slender, more than 50 on lower half of first gill
arch
.
.
Chaetobranchus (p. 271)
b. Gill rakers not very long and slender, less than 25 on lower half of first
.
3
gill arch
3a. Preoperculum finely serrate
b. Preoperculum entire
.
.
.
.
4
6
4a. Jaws equal anteriorly; short-bodied fishes, scales 26-29, those of lateral
line of same size as those above and below it
. Crenicaru (p. 289)
b. Lower jaw projecting; long-bodied fishes, scales more than 38, those of
lateral line larger than those above or below it
.
. 5
5a. Inner teeth depressible; scales 38-130
b. Inner teeth not depressible; scales 55-70
.
.
. Crenicichla (p. 291)
. Batrachops (p. 290)
6a. Gill rakers large, stiff, perch-like, mouth large; dorsal fin notched
between spiny and soft parts; upper and lower lateral lines continuous
in young
.
. Cichla (p. 265)
b. Gill rakers minute or short; no notch between spiny and soft parts of
dorsal fin; upper and lower lateral lines not continuous
. 7
7a. Upper part of first gill arch with downwardly-projectinglobe
b. Upper part of first gill arch without downwardly-projecting lobe
.
.
8
11
8a. Upper lateral line running within a scale or scale and a half below dorsal ;
small 'dwarf' cichlids
.
. .Apistogramma (p. 298)
b. Lateral line well separated from dorsal for most of its length (at least two
and a half scales away) ;larger-growing species .
.
9
9a. Gill rakers at base of downwardly-projectinglobe Acarichthys (p. 277)
b. Gill rakers running at edge of lobe on first gill arch
.
. 10
10a. Preorbital twice as deep as eye in adult ;eye three times as far from snout
as from posterior margin of operculum in adult; lateral line generally
forked at the caudal peduncle
Geophagus (p. 278)
.
.
ROSEMARY
H. LOWE-MCCONNBLL
264
b. Preorbital little if any wider than eye; eye a little nearer to posterior
margin of operculum than to snout ;lateral line not continued on caudal
.
Biotodoma (p. 281)
lla. Three anal spines
.
b. More than three anal spines
.
.
.
12
14
12a. Mouth large, freely protractile, end of maxillary exposed .
. Acaronia (p. 272)
b. Mouth small, not greatly protractile, maxillary not exposed .
. 13
13a. Lateral line running close to dorsal ;head densely scaled ;dwarf cichlid
.
.
Nannacara (p. 299)
b. Lateral line well separated from dorsal ;larger-growing species
.
.
Aequidens (p. 273)
14a. Premaxillary processes shorter than head ;end of maxillary concealed
.
Cichlasoma (p. 282)
b. Premaxillary processes as long as head ; end of maxillary exposed .
. Petenia (p. 289)
In addition to cichlid genera shown in the key, certain other genera from neighbouring countries which may one day be found in Guyana are :
(1) Chaetobranchopsis Steindachner, 1875 (p. 133); a genus with numerous long
slender gill-rakers as in Chaetobranchusbut with more than three anal spines (contrast
three in Chaetobranchus).Two species, distinguishable by colour patterns, C. orbicularis
(Steindachner) and C. bitaeniutus E. Ahl, 1936, are known from the Amazon.
(2) Astronotus Swainson, 1839 (p. 229) ;a genus with three anal spines, preoperculum
entire, without a downwardly-projecting lobe on the first gill arch, and with the soft
vertical fins (dorsal, ventral and caudal) covered with scales. Astronotus ocellatus
(Cuvier), a magnificent and well-known aquarium fish (Sterba 1962, fig. 877) occurs
in the Rio Negro, and A. orbicularis Haseman is another Amazon species.
(3) Symphysodon Heckel 1840 (p. 332) ; characteristically disk-shaped fishes with
strongly compressed bodies and long anal fins with seven or more spines. Also striking
aquarium fishes (Sterba 1962, figs 951,952), the young attach themselves to the bodies
of the parents where they are said to feed on a skin secretion (Herald, 1961:199),though
they can be reared in the absence of the parents (Sterba, 1962: 782). S. discus Heckel
occurs in the Rio Negro and various subspecies of S. aequijasciatus Pellegrin in the
Amazon.
(4) Uaru Heckel, 1840 (p. 330); another aquarium fish (Sterba 1962: 723 and figs
1106, 1107), with slender compressed teeth, rather small scales (lateral line scales
larger than others), and more than three anal spines U. imperialis (Steindachner) and
U.amphiacanthoides Heckel are known from Rio Negro and Amazon respectively (the
former considered synonymous with the latter by Regan, 1905c: 439).
CICHLID
FISHES
OF GUYANA
265
NOTES ON THE STATUS, OCCURRENCE AND NATURAL HISTORY
OF GUYANA CICHLID SPECIES
Cichla Schneider
Cichla Bloch & Schneider, 1801: 340.
(Type species :Cichla ocellaris Bloch & Schneider, 1801 : 340, by subsequent designation of Eigenmann & Bray, 1894 :611 .)
Perciform, spinous and soft dorsal fins separated by notch. Soft dorsal, caudal and
anal fins scaled; three anal spines. Gill rakers long and heavy. Lateral line continuous
in young, usually interrupted in adult, forked at base of caudal.
Many species of Cichla were named by Schomburgk (1852) which, as Eigenmann
(1912) pointed out, appeared to be based on different growth stadia. Cichla spp. are
widespread in tropical South America; the colours vary very much from different
waters, and colour patterns change with the size of fish. Eigenmann (1912 :509) thought
that there were three species of the genus in South America, but was uncertain whether
or not there were two species in Guyana; he considered only C . ocellaris Bloch &
Schneider, after pointing out that some of his small specimens (138 mm long and
figured as plate 69, fig. 2) did not agree well with Steindachner’s figures of either
C . ocellaris or C. temensis Humboldt of comparable size.
Since Eigenmann’s day two species have become widely recognized in Brazil,
C . ocellaris and C . tememk, which are both cultivated in fish ponds (Allsopp, unpublished
report ;Fontenele, 1950 ; Braga, 1952), though Schultz (1949) synonymized C. ternemis
with ocellaris when considering Venezuelan fishes. Sawaya & Maranhao (1946) reported
that C. ocellaris and ternensis hybridize readily in ponds, and they recorded a third, still
undescribed species, from Brazil (known locally as ‘Tucunark putanga’ and growing
only to 30-40 cm long, as does C . ternensis). Young of C. ocellaris and temensis (about
70 mm long) brought from Brazilian ponds by Allsopp were examined in Guyana,
and the colour patterns agreed well with the C. ocellaris and C . ternensis figured by
Steindachner (1883, plate I, figs 2 and 3 ) ; C. ocellaris young have three vertical dark
bars and a horizontal dark band only from the last vertical bar to the caudal, whereas
C . ternensis young have the horizontal band from snout to caudal but no vertical bars,
and more rounded lobes to the caudal fin. These two colour patterns were quite distinct
(though there is some evidence that there may have been some confusion over which
species was which in some fish culture work). These two species are also well known to
sport fishermen; McCormick (1949) comments that for adult fish the colour patterns
provide the main difference, the more widely encountered C . ocellaris (TucunarC-assu)
being ‘brighter and more beautiful’ but with a stubbier and less humped outline than
C . temensis (Tucunart-branco or Tucunark-tinga).
During the present field studies all the Cichla collected in Guyana with one exception
appeared to be C. ocellaris. This one specimen of C. ternensis, from the Amazon-drainage
side of the Rupununi District at Pirara, shot with an arrow when the fish were running
back to the Takatu River after the rains, appeared without doubt to be a different species,
differing mainly in colour (green with light yellow spots in horizontal lines along the
body and without the vertical dark bars of C. ocellaris), but also in its cleaner lines and
less deep body, sharper angle to the lower jaw, and more distinctly lobed caudal fin.
266
ROSEMARY
H. LOWE-MCCONNELL
Regan (1906) recognized two species, C . temensis having smaller and more numerous
scales. In the British Museum collection all the material previously collected from
Guyana is C. ocellaris, but C. ternensis were examined from neighbouring countries;
these appeared to be a distinct species, and the small specimens (53/63 mm long)
agreed with Steindachner’s figures of young C. temmsis.
It seems therefore that two species of Cichla do occur in Guyana, but that C. temensis
is rare and may be confined to the Amazon (or Amazon and Orinoco) drainage areas.
(This leaves in abeyance the question of Eigenmann’s doubtful fig. 2 specimens, from
Gluck Island in the Essequibo River of Guyana.) C. ocellarisvariesvery much in colour
from river to river and in the degree to which the vertical bars become ocellated in
older fish.
Key to Cichla species found in Gayana
la. Colour brownish or olivaceous with three or more broad dark vertical
bars on the upper part of the body and an ocellus on the upper part of the
caudal base; vertical bars tend to be replaced by large ocelli in large specimens ;young with a horizontal dark stripe extending back from the hindermost vertical stripe or blotch to caudal fin; scales 83-102
C . ocellaris
b. Colour greenish with distinct light yellowish spots horizontally aligned
in adult ;without vertical dark bars (though with an ocellus on upper part
of caudal base) ;horizontal stripe in young along whole length of body from
eye (or snout) to end of caudal fin; caudal fin more distinctly lobed; scales
smaller, 104-121 along body
.
C. temensis
Cichla t m m i s Humboldt
Common names. Dark or Deer Lukanani ;TucunarC-branco, TucunarC-tinga.
Cichla temensis Humboldt, 1883 : 169 (Orinoco, Rio Negro, Amazonas); Heckel, 1840:
413; Gunther, 1862: 304; Steindachner, 1883 : 3, pl. I fig. 3 (juvenile).
Material aoailable: One (221 mm), Rupununi District, Pirara stop-off, Amazon
drainage, 30 Sep. 1957.
The only specimen seen during recent field studies was collected when it was running
down Pirara Creek which drains to the Takatu River; it had been damaged by a
SerrasQlmus bite; it is here known locally as Deer or DarC Lukanani because of its
dappling with light spots, and it is said to grow large. As already mentioned young
from fishponds in Brazil were examined in Guyana, and from Santarem at the British
Museum (N.H.), together with adult specimens from Brazil.
Cichla ocellaris Schneider
Common names. Lukanani ;TucunarC-assu.
Cichla ocellaris Schneider, 1801 : 340, pl. 66 (‘India oriental’: S. America); Gunther,
1862: 304 (Demerara); Steindachner, 1883: 3, pl. I fig.2 (juvenile); Eigenmann, 1912:
509, pl. 69, figs 1-4; Puyo, 1949: 242, fig. 127.
CICHLID
FISHES
OF GUYANA
267
Material available. Two (183-340 mm) British Guiana, coll. Schomburgk;four (253335 mm) Berbice, coll. Matthey; 20 (77-176 mm) Rupununi District (11 from Amazon
drainage, nine from Essequibo drainage) ;two (125-242 mm) Georgetown trenches.
Material examined inJield. 250 (197 from Rupununi District, 53 from coastal plain
fresh waters).
Distribution. Widely distributed in tropical waters of South America. Common
throughout Guyana in rivers, savanna pools and coastal plain swamps and freshwater
trenches. One of the best-liked food fishes, C. ocellaris is caught on hooks (on handlines
and by trolling), shot with bow and arrow, trapped as the fish are returning to the main
rivers after the rains, seined and poisoned from ponds, trapped in coastal trenches.
Colour. Well known from aquarium books (Sterba 1962, fig. 1069); this is a very
colourful species but varies very much with the growth stage and from river to river.
The brownish green ground colour, darker on the back, grades to silvery white on the
belly. The longitudinal band on the hind half of the body of young fish becomes disrupted into isolated blotches as the vertical dark bars develop, and as the fish grows
these vertical bars become broken up into dark blotches which are increasingly ocellated
with silver and yellow pigment, particularly under the soft dorsal. From an early age
a deep black spot on the upper half of the caudal fin base is ocellated with bright yellow ;
this becomes a particularly brilliant ‘bull’8-eye’in breeding fishes. The lower part of the
head becomes bright yellow and the lower parts of the belly, lower half of the caudal
fin and the pelvics all deepen in colour, from yellow to orange or brick red, in breeding
fishes. The iris is bright orange-red. The male develops a distinctive adipose hump on
the nape just before spawning.
Growth. C. ocellaris is a predatory species which grows much larger than any other
Guyana cichlid, and unlike many of the other cichlids continues to grow long after it
has reached maturity. The sizes seen, the relative sizes of males and females and the
sizes at which they mature, are indicated in Fig. 3.
The largest seen in Guyana, a 50153 cm male weighing about 8 lb (3%kg) came from
Manari Creek (Amazon drainage). All those above 38/43 cm (above about 3 lb, 1.5 kg,
in weight) were males; 75 males to 67 females were recorded. Both sexes matured
around the same size, about 20/24 cm long (weight 200 g), but the males were generally
slightly larger than the associated females when spawning pairs were trapped together
(e.g. males about 20 cm and females 18 cm standard length). No fish larger than 23/27
cm (weight about 300 g) were taken from coastal trenches (though they grew larger in
the ponds). There was no difference in the maturation size in the two areas, which
suggests that heavy fishing may account for small sizes in the coastal trenches.
The Rupununi has only one well-defined wet season (May to September), and
breeding appears to be more seasonal here than on the coast. The length frequencies
of C . ocellaris caught at different times of year in the Rupununi are shown in Fig. 3 F-H.
In September (G),just after the rains, many young-of-the-year were seen. By December
(H)
no fish smaller than 14 cm standard length were caught ;lengths were, however,
scattered over a wide range and numbers were too low to allow estimates of growth
rates from length frequency mode progressions. Some prematuration size C. ocellaris
were also taken in April (F),suggesting that growth between December and April is
slow, a time when food is increasingly scarce as the dry season progresses.
ROSEMARY
H. LOWE-MCCONNELL
268
Young C. ocellaris, 45 mm long caught in a shoal following the parents, were introduced into a'Fisheries Division pond, Georgetown on 2 August 1957, and by January
1958 (five months later) a 17/19 cm 120 g fish was caught (a female with brilliant caudal
Standard length (cm)
2
6 8
10 12 14
16 18 20 22 24 26 28 30 32 34 36 38 40 42 44 46 48 5(
1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1
4
20
10
0
30
20
10
0
10
5
0
50
H
5'
3
2
2
24 -62
:81 hd hi bi 4': hs' %
Standard length (cm)
FIGURE
3. The relative sizes of different cichlid fishes from Guyana waters. The relative sizes
(standard lengths in centimetres) of males (spotted) and females (fine stipple) are shown
€or: A,Acaroma nassa; B,Cichlasoma bimaculatum; C , Cichlasomafestivum; D, Cichla ocellaris
from Georgetown trenches; E,CichZa ocellaris from the Rupununi District. (Unsexed or indeterminate fish shown blank.) The relative sizes of Rupununi Cichla ocellaris caught F in April
G September and H December 1957 are then shown separately (solid black histograms).
ocellus). This suggests that maturation size can be reached in less than one year. In
Brazilian ponds C. ocellaris are said to mature after 11-12 months (Fontenele, 1950).
Breeding s e m m . In the Rupununi district C. ocellaris spawn before and during the
rains which start in May; gonads were quiescent by the end of the rains in September,
but starting to become active again by December (based on observations of the gonads
of 40 fish, 20 male and 20 female, examined in April, of which 29 were ripe or ripening ;
on observations of parents with young when the water was low before the rains in
.
CICHLID
FISHES
OF GUYANA
269
May ;the quiescent gonads of the 3 1 fish examined in September ;and five out of eight
with gonads showing signs of activity in December). Places on the savanna where this
species spawned when the savannas were flooded were shown to me.
On the coastal plain, where there are two rainy seasons a year (April-August and
November-December) breeding may be more evenly distributed over the year. Here
gonad activity was noted in C . ocellaris in April, August, October, November and
January. In the Lama water conservancy a pair were watched guarding a nest hole in
January 1957 ; at Sophia, Georgetown, C . ocellaris were watched leading young in
April 1961 ; at Mon Repos, also near Georgetown, a 22/27 cm female was leading
45 mm long young on 2 August.
In Brazilian ponds, well south of the equator, Fontenele (1950) reports that Cichlu
may spawn at any time of year, but spawning is more frequent from June to December ;
there may be three of four spawnings a year. Before spawning the male develops the
post-orbital nuptial hump and becomes aggressive; in the female the yellow on chest
and fins becomes orange, and spawning is said to start some 7-20 days after the appearance of these secondary sexual characters. As in Guyana, the male is nearly always
larger than the associated female.
Breeding behaviour. Cichla grow too large to breed in aquaria; Sterba (1962) reports
‘nothing known’ of their breeding, and Braker (1962) quotes Magalhaes (1931) as
saying the young are brooded in the branchial cavity. This appears to be quite erroneous.
The spawning habits of Cichla are well known from ponds in Brazil (Sawaya & Maranhiio, 1946; Fontenele, 1950). Here both male and female are said to clear a surface for
spawning and to make several 8-20 cm diameter circular ‘nests’, small depressions,
1.5-6 cm deep (as figured by Sawaya & MaranhPo 1946, pl. I1 for C. ocellaris; Braga,
1952: 275 for C . ternensis) which they dig in the bottom mud or bank with chin and
pectoral fins. They evidently prefer shallow water (20-30 cm deep), and Sawaya &
MaranhPo say that they generally work at night. The male is then said to bounce the
female against the cleared surface on which the eggs are sprayed in rows and fertilized ;
this operation may take up to 23 hours, during which time 6000-10,000 eggs may be
laid. Nest pits may be built even after this. The male and female guard the spawn, the
male generally attacking intruders ;very little feeding takes place during spawning and
incubation. The eggs remain attached to the hard surface till they hatch, after about
70-90 hours, depending on the temperature. The young are then transferred to the
nest pits in the parent’s mouth. The young, which hatch at about 5-6 mm long, have
adhesive organs on the head, described by Fontenele (1952). The transfer of all the
young to a nest pit is said to take about four hours, and they remain at the bottom of the
pit for about four days before swimming up and around. During this time they are
guarded by both parents, who may move them from one pit to another if they are
disturbed. Parent Cichlu are not good fighters, leaving the nests if danger occurs, but
returning very quickly when it is over. The yolk sacs are resorbed and the young are
able to feed five to six days after spawning. The shoal then swims out guarded by both
parents. Fontenele says that the young are guarded till about 35 mm long; Sawaya &
MaranhPo state that a month after spawning the male only continues to guard, chasing
the female away, and after another month the young are independent.
During these field studies the lentil-likeyellow eggs of C . ocelluris were found attached
18
270
ROSEMARY
H. LOW-MCCONNELL
to sticks in the savanna ponds ; in the Lama water conservancy a pair of C. ocellark
were watched guarding a hole several centimetres deep and 10-12 cm across with a
hard surface nearby; and parents were often watched guarding their shoals of freeswimming young. Young 45 mm long, in a shoal of over 1250 young, were caught following the 22/27 cm female at Mon Repos in August.
Observations made on parent C. ocellak guarding their young in May 1961 at
Karanambu on the Rupununi River, suggested that the caudal ocellus, which becomes
so brilliant in the breeding fish, has an important function in enabling the young to
orientate themselves to the parent. The river was low, and the water rather cloudy at
this time before the rains started, and five pairs of C. ocellaris, each with a cloud of
young above them, could be seen round the rocks in a small bay. I n each case the two
parents were of comparabie size (about 36 standard/41 total cm length as in a fish
caught here on a troll). The young in each group were of similar size (probably between
20-40 mrn long). There appeared to be several thousand young in each bunch, which
kept in a shoal about 0-5m in diameter, the small fishes weaving into the shoal, spreading
out and turning back, always above the parents. The shoal seemed to spread by the
small fish from the back rushing forward ;the back ones also seemed to be the first to
start diving when at times they all dived and the shoal sank out of sight, but the water
was too murky to see if or how they were ‘called’down. One shoal moved progressively,
another stayed in the same place for 20 minutes or so. Each shoal seemed to be orientated
by the parent’s tail with its well-marked dark ocellus showing up clearly in the murky
water, waved like a flag in front of them. From above, from where the young ones also
got their view, the caudal ocellus, a bull’s eye black on yellow, stood out very clearly
on the whitish grey (lower lobe) and yellowish (upper lobe) background on each side
of the tail. The black bars along the sides of the parent’s body, seen from above as a
double row of blobs, were also very clear, as were the white crescents on top of the
parent’s eyes. The parent fish was orientated at about 45” with head down and the tail
waving in the water like a flag. The body of the parent was so large that it was mostly
hidden in the cloudy water, so that all that the young could probably see would be the
tail. At intervals plops at the surface suggested the second parent was close by. The
young seemed to have no fear of a large Arapaima gigas which rolled near them when
it surfaced to breathe every eight to ten minutes.
Food. The stomach contents of 106C. ocellariswere examined (71 from the Rupununi
District, 35 from Sophia, Georgetown, trenches). Of these 48 were empty, 48 contained
fish remains, four contained palaemonid shrimps, and three bottom debris including
a bolus of algae, pieces of chitin, and invertebrate eggs. The fish remains in the stomachs
included other cichlids in six cases (Cichlasoma festivum in 3, C. bimaculatum in 1,
Crenicichla saxatilis in 1, and Geophagus or Aequidens in 1); characoids in 11 cases
(including Creatochanescaudomaculatusin 1,Charax sp. in 1,Curimatella in 2, Astyanax
sp. in 3 ) ; one stomach contained both a small characid and a small cichlid, another
contained a Corydoras sp. They evidently take whatever small fish they can catch,
rather than being very selective. The palaemonid shrimps were found in rather small
C. ocelluris (8-13 cm fish). Young ocellaris 4 cm long, caught while still with their
parents, contained planktonic Crustacea and algal filaments;they already had nematodes in the gut.
CICHLID
FISHES
OF GUYANA
271
Predators andparasites. Small C. ocellariswere found in the stomachs of the characid
Hoplias malabaricusat Sophia. Nematode worms were generally abundant in C. ocellaris.
Chaetobranchus Heckel, 1840
Chaetobramhus Heckel, 1840: 401.
(Type species :ChaetobranchusJlavescensHeckel, 1840:402, by subsequent description
by Eigenmann, 1912: 483.)
Readily distinguished from all other Guyana cichlids by the numerous long setiform
gill rakers. There is only one species in Guyana, C.JEavescens.
Chaetobranchusfravescens Heckel
ChaetobranchusjlavescensHeckel, 1840: 404 (Rio Negro) ;Eigenmann, 1912:483.
Material available. One (268 mm) South Akerica, coll. Schomburgk (type of C.
robustus Gunther, synonymized with C.JEavescensby Regan, 1906:235) ;17(56-165 mm)
Karanambu area, near Rupununi River.
Material examined in$eld. 72 (55/70- -1280 mm) Rupununi District.
Distribution : Guianas, Amazonia, Peru, Bolivia. Found only in the Rupununi
District during present studies, where it was seined from drying pools in the North
Savannas only, though Eigenmann (1912) records it from Rockstone on the Essequibo
River and the Lama Stop-off.
Colour. Tilapia-like in shape, in life this a colourful species. The general body colour
is grey-green with a spotted effect due to darker areas at the centre of each scale, a dark
lateral blotch, dark red spots and vertical stripes on the caudal, dorsal and anal fins.
The large eye and large mouth are linked by two peacock green stripes, separated by a
yellow-ochre stripe down the preorbital; chin white, chest pearly grey. Adult males
have elongated caudal rays and the lower half of the caudal is red. There is no caudal
ocellus. In the young the vertical fins are striped in dark red. Iris cream.
Growth, The size distribution of 37 adults examined from Karanambu ponds,
April 1957 was as follows:
Totallength(cm)
No. males
No. females
22 23 24 25 26 27 28 29 30
0
2
1
6
5
4
4
5
1
1
6
0
2
0
0
0
0
0
Total
19
18
Thus themales tended to be larger than the females. This species matured when about
17/22 cm long.
Breeding seasons. Most of the females had ripening gonads in April 1957, just before
the rains. In September 1957, after the rains, 33 young of 5-7 cm standard length were
found ;young collected in January 1960were 10/13 cm (45 g weight). So the indications
are that breeding is seasonal and occurs during the rains. Nothing is known of breeding
habits (the lack of a caudal ocellus suggests it may be a mouth brooder).
Food. The long slender gill rakers and weak teeth suggest that this is a planktonfeeder, but fish examined in April from the drying ponds had little food, stomachs
were very small and empty or contained only grey slime ;the liver was huge and there
272
ROSEMARY
H. LOW-MCCONNELL
was much fat along the coiled intestines. This species seems to be restricted in distribution to the larger bodies of water, places where it might be expected that plankton would
be richest in the wet season, the main feeding time in this environment.
Acaronia Myers, 1940
Acaropsis Steindachner, 1875 : 20 (preoccupied).
Acaronia Myers, 1940a: 170. (Substitute name as Acaropsis preoccupied by an arachnid). (Type species: Acara nmsa Heckel, 1840, by monotypy of subgenus Acaropsis
Steindachner, raised to genus by Eigenmann & Bray, 1894: 613, and retained for
Acaronia.)
A genus with the characters of Aequidens, but with the mouth large, premaxillaries
very protractile and the end of the maxilla exposed when the mouth is closed. One
species only in Guyana, A. nassa.
Acaronia nasa (Heckel)
Acara nasa Heckel, 1840:353 (Rio Guapore ;Miiller & Trotschel in Schomburgk, 1848
624 (L. Tapacuma, British Guiana).
Acmopsis nassa, Eigenmann & Bray, 1894: 613 ;Eigenmann, 1912 : 485 ;Puyo, 1949 :
240.
Material available. One (40 mm) Demerara R., coll. Beckford (Regan, 1905); two
(50 & 130 mm) Maduni Creek, coll. Eigenmann; three (c. 115-140 mm) Demerara R.,
coll. Rosenberg; 12 (46-98 mm) Rupununi District Amazon drainage; eight (24-108
mm) Rupununi District Essequibo Drainage.
Material examined infield. 82 from Sophia trenches, Georgetown; large numbers of
small ones from Rupununi District. The ecological data came mainly from Sophia.
Distribution.Guianas, Amazon, Orinoco. In Guyana widely distributed and common
in fresh waters of coastal plain and Rupununi District.
Colour. A large-scaled fish with characteristically large and protrusibIe mouth, very
narrow preorbital and large eye, in life the general body colour is grey with a goldenbronze sheen dorsally and pearl-white ventrally; dark lateral blotch and small dark
bar on the upper part of the caudal peduncle ; the head has chalky white streaks on
preoperculum and cheek, the angle of each preoperculum has a dark spot ocellated with
silvery gold, and there is a second dark spot immediately below each eye. Caudal, soft
dorsal and anal fins spotted with darker grey, anal with chalky white spots anteriorly,
pelvics chalky white, pectorals immaculate lemon yellow. The large eye with its huge
pupil has a yellow-orange iris.
Growth. The size range seen and relative sizes of males and females were as follows:
Standardlength(cm):
7
8
No. males
No. females
1
1
1
8
9 10 11 12 13 14 15 16 17
4
1
1
1
3
8
6
7
5
2
6
1
5
0
1
0
0
0
0
0
Total
42
38
When pairs are trapped together the male is generally the larger fish (e.g. 11/13 cm
75 g female, 13/15 cm 105 g male; 10/13 cm female, 12/15 cm male). The largest seen
CICHLID
FISHES
OF GUYANA
273
were males, 14/18 cm 145 g and 15 cm standard length. Females matured when only
8/10 cm long (30 g); maturation size was harder to determine in the male.
Breeding seasm. Ripe A. m s a females were caught at Sophia every month from
April to November ; ripening ones were taken in February and March ;no data were
available for December and January. I n September more fish seemed to have quiescent
gonads. Thus the breeding season seems to be very protracted, breeding occurring
both in the dry season before the rains and during the rains. The gonads regress very
thoroughly in this species between breeding periods. The ova are large and dark ochre
in colour. Nothing is known of breeding habits.
Food. The stomach and intestinal contents of 50 Sophia-trapped A . nmsa were
examined. Palaemonid shrimp remains were found in 22, insect remains in 25, and
fish remains in 12. Five were empty, three contained chitin, one vegetable debris,
one pieces of water plants, one desmids, one mush. Several items were sometimesfound
in one fish, often with a gap in the intestine between them indicating that they had
been eaten at different times (palaemonid shrimps and Odonata nymphs were a
common combination for alternate meals). The insect remains were Odonata
nymphs in 13 cases (two with wings, evidently taken just as the adults were
emerging), water beetles in three cases, aquatic Hemiptera in one, aerial insects in
two. The 12 A. nmsa containing fish included two with cyprinodonts, four with small
characids, one with a cichlid (C. bimaculatum ?)
The mouth is opened suddenly and the food appears to be swept into the large mouth
with the inrushing water. A . nassa lurk near the bottom of the trenches ;they are solitary
fish.
Predators. A nassa was found inside Hoplias malabaricus on one occasion.
Aequidens Eigenmann & Bray, 1894
Aequidens Eigenmann & Bray, 1894 : 6 16.
(Type species :Acara tetramerus Heckel, 1840, by subsequent designation of Eigenmann
& Bray, 1894 : 617.)
Acara, Regan, 1905 a.
Eigenmann & Bray gave the new name as no type was indicated for Acara until
Gill (1858) restricted it to Heckel’s Acara crassipinnis, a species since determined to be
synonymous with Lobotes ocellatus Agassiz, the type of Astronotus Swainson, 1839 ;
Eigenmann & Bray considered that this made Acara a synonym of Astronotus. Regan
(1905a) considered this argument unsound, but it has been accepted by more recent
authors.
Gills with minute rakers and no lobe; mouth small, but not greatly protractile,
anal with three spines ; lateral line well separated from dorsal ;preoperculum entire ;
scales of lateral line of same size as those above and below it.
During these field studies Aequidens were collected from the Rupununi District
together with the much commoner Acarichthys heckelii (considered by some authors
to be an Aequidens, see below). The young of Aequidensgeayi (Pellegrin) were very
common in stony pools of the drying-up bed of the Rupununi River in the South
Savannas, and a few A . tetramerus (Heckel) were collected from leafy but open pools
274
ROSEMARY
H. LOWE-MCCONNELL
in the North Savannas at the end of the rains. No other species of Aeqvlidens were found
in this area.
Later studies on museum collections showed A. potmoensis Eigenmann to be the
common species in the forest rivers of the interior, and A. wittatus (Heckel) in the coastal
rivers. There is some overlap in distribution as Carter’s collections included both these
species and A . tetramerus all taken from near the Penal Settlement on the Mazaruni
River. The relationships of these species and A. gu&&
are complex, as is discussed
below. A. maronii Steindachner was seen in various rivers, but not collected during
these studies.
Gonad states and stomach contents of A e q u i k were not examined in the field;
most of the specimens caught were small and were preserved whole as their specific
identification was uncertain when field studies were made. Significant markings are
shown in Fig. 2.
Field key to Aequidens species found in Guyana
la. No caudal ocellus or black spot at base of upper half of caudal fin
b. Ocellus or black spot above lateral line at base of caudal fin .
.
.
2
3
2a. Dorsal and anal fins densely scaled at base; large dark ocellated blotch
below soft dorsal, mainly above lateral line but with a less dark extension,
making a keyhole shape of it, below lateral line; black band from nape
A. rnaronii
through angle of preoperculum ;rounded profile .
b. Dorsal and anal fins naked; dark lateral stripe from middle of dorsal fin
obliquely backward to vent, a second stripe from nape through eye to angle
of preoperculum (best defined on angle of preoperculum below cheek), the
stripes of the two sides forming saddles when viewed from above, one over
. A. geayi
the nape, the other over the back below the spinous dorsal
3a. Dorsal and anal fins naked (or with only one scale at base of each membrane
in old specimens)
.
. 4
b. Dorsal and anal scaly at base
.
.
A. guianensis
4a. A dark band from eye semiobliquely to base of hind dorsal rays, this band
generally continued forward from eye to (and sometimes along) upper lip ;
black band downwards from eye to preopercular angle ;a black spot below
the middle of the upper lateral line (on the dark band), and another, well
rounded, at the base of the upper caudal rays; indications of about six
vertical stripes, sometimes darkened where these cross the lateral bands.
Soft dorsal and anal in adult sometimes with a single scale on membrane
A. vktatus
b. Dark lateral band, if present, running more medianly through or from
median lateral spot to spot or bar at base of upper caudal rays, and continued
over the head between the eyes (obscured in dark fish) .
, 5
5a. A dark band from eye to preopercular angle; caudal spot tends to be elongated into a short bar across the lateral line; ill-defined vertical stripes,
colour deepened where these cross the lateral band .
. A. potaroensis
CICHLID
FISHESOF GUYANA
275
b. A dark spot below and behind eye; distinct median lateral spot and no
lateral band ; caudal spot round and ocellated, above lateral line at base of
caudal .
. A . tetramerus
Acara guianensis Regan (1905: 341) appears to be known only from the type, a
fish 80/110 mm long, locality ‘Guiana’, received from ‘Berlin Museum’ (1851.5.2.8).
The colours are faded but the distinctive caudal spot puts it into the vittatus-potaroensis-tetramerus group, and traces of band from eye to preoperculum suggest A. vittatus
and A . potaroensis. It differs from these, however, in having soft dorsal and anal scaly
at the base ;in large specimens of A . vittatus there may be a scale on each membrane,
but otherwise dorsal and anal are scaleless in A . wittutus and A. potaroensis. It seems
therefore that A.guianensis Regan is zdistinct species. It isnot knownfromwhich Guiana
it was collected. Another species of this group recently described from French Guiana
is A . itanyi Puyo, 1943, distinguished from A. wittatus mainly by the colour pattern
(the vertical stripes forming distinct blotches where they cross the lateral band running
from upper lip to between last dorsal rays and top of the caudal peduncle) but which
also has the soft dorsal and anal slightly scaly at their base. A . itanyi also appears to be
known only from the types, two specimens 70 and 114 mm, which I have not seen.
More material is needed to determine the status of these two species, and whether
A . itanyi is a synonym of A. guianensis.
Other Aequidens species not yet recorded from Guyana but occurring in neighbouring
countries are : A . pulcher (Gill), A . mariae Eigenmann, and A . duopunctata Haseman.
(1) A. pulcher (Gill) from Venezuela and Trinidad; a species readily recognizable
by about ten vertical dark bars across the body, the first from the nape through eye to
angle of the preoperculum, one through the lateral blotch and the last at the base of the
caudal fin.
(2) A . mariae Eigenmann known from Brazil. This species has an oblique lateral
stripe to the hind end of the soft dorsal fin (as in A . vittatus),but instead of continuing
forwards through the eye to the upper lip, the lateral stripes of the two sides meet
over the nape behind the eye; there is also a dark stripe from eye to angle of
preoperculum.
(3) A . duopunctata Haseman recorded from Manaus, which has colouring very
similar to A . tetramerus and was considered by Regan (1913a :282) to be synomymous
with tetramerus, though material collected from Manaus since then suggests that they
are distinct species.
Many Aequidens species have been bred in aquaria (including A . maronii, A . tetramerus, A . curviceps (E. Ahl) from the Amazon, A . Zutifrons (Steindachner) from
Columbia, and A. portulugrensis (Hensel) from southern Brazil). These are all said to
breed in ‘typical cichlid fashion’ (Sterba, 1962), that is they are substratum spawners
in which male and female guard the eggs and young.
Aequidens maronii Steindachner. Keyhole cichlid
Acara maronii Steindachner, 1882: 141, pl. 2 fig. 4 (Maroni R., Guiana).
Aequidens maronii: Eigenmann, 1912: 489 (Demerara R., Barima R.).
276
ROSEMARY
H. LOWE-MCCONNELL
Material available. One (77 mm) Demerara, coll. Beckford; one (34 mm) Kumaka,
British Guiana, coll. Eigenmann ; one (73 mm) Georgetown, coll. Rosenberg ; two
(53,54 mm) Mackenzie, Demerara R., coll. Carter.
Material examined in field. None, though this species was seen in the river at
Mackenzie.
Eigenmann (1912) records the species from Demerara, Barima R. and Kumaka.
This species is exported as an aquarium fish, a ‘peaceful species’ which breeds in
aquaria.
Aequidens geayi (Pellegrin)
Acarageayi Pellegrin, 1902: 417 (copied); 1903 : 178, pl. 4 fig.3.
Aequidensgeayi:Eigenmann, 1912: 493 (Potaro and Essequibo Rivers).
Material availuble. One (84 mm) Essequibo, coll. Ehrhardt ; one (90 mm) Amatuk,
coll. Eigenmann; one (86 mm) Upper Cuyuni, coll. Carter; one +small ones
R. Cuyuni rock pools below Camair a Falls, coll. Carter; 75 (21-99mm) Rupununi
River, rocky pools in south savannas.
Specimens from French Guiana (from Oyapock R., Approuague R. & Mataroni
Creek, coll. Ternetz (43-93 mm)) appear to have steeper profiles than the Rupununi
specimens.
This species might make a good aquarium fish.
Aequidens vittatus (Heckel)
Acara vittatus Heckel, 1840: 346 (Cujuba).
Aequidens vittatus : Eigenmann, 1912 :489 (Demerara River).
Material available. Two (39, 52 mm) Lama, coll. Eigenmann; two (79, 99 mm)
Hyde Park, Demerara R., Coll. Rosenberg; four (60-84 mm) Mackenzie, Demerara
R., coll. Carter; one (56 mm) Mazaruni R., coll. Carter; one (99 mm) Bootooba,
Demerara R. ;one (66 mm) Lama Stop-off.
Specimens from Approuague R. French Guiana, coll. Ternetz, were also examined.
Aequidens guianensis (Regan)
Acaraguianemis Regan, 1905 : 341 (Guiana).
Material available. Type only (80 mm) Guiana, Berlin Museum.
Aequidens tetramerus (Heckel)
Acara tetramerus Heckel, 1840: 341 (Rio Branco).
Aequidens tetramerus: Eigenmann, 1912: 491 (Rupununi, Essequibo at Rockstone and
Gluck I., Lama Stop-off).
Materid available. Five (90-181 mm) South America, coll. Schomburgk; one
(112 mm) Essequibo R., coll. Ehrhardt; six (66-120 mm) Hyde Park, Demerara R.,
coll. Rosenberg; one (62 mm) Upper Cuyuni R., coll. Carter; one (68 mm) Potaro
CICHLID
FISHES
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277
R., coll. Carter; 20 (46-95 mm) Karow Creek, Mazaruni R. near Penal Settlement,
coll. Carter; one (62 mm) British Guiana, coll. Graham; five (46-54 mm) open pools
near Karanambu Rupununi R.
Aequidens potaroensis Eigenmann
Aequidenspoturoensis Eigenmann, 1912: 490, pl. 64, fig. 2 (Potaro R., British Guiana,
co-types from Ireng River). Note : the Potaro is in the Essequibo drainage, the Ireng
in the Amazon drainage.
Material available. Two (79, 114 mm) British Guiana, coll. Eigenmann (co-types) ;
one (106 mm) Chinapoon R., Upper Potaro, coll. Bovallius; nine (64-106 mm) Mazaruni
R. creek near Penal Settlement, coll. Carter; one (66 mm) Mackenzie, Demerara R.,
coll. Carter; five (48-113 mm) Potaro R., COIL Carter, one (116 mm) Cuyuni R., coll.
Carter; nine (25-1 10 mm) Upper Cuyuni R., coll. Carter; one (93 mm) Mazaruni R.,
coll. Allderidge.
Acarichthys Eigenmann
Acarichthys Eigenmann, 1912: 500.
(Type species : Acara heckelii Miiller & Trotschel, by subsequent designation by
Eigenmann, 1912: 500.)
The genus Acurichthys has the dorsal lobe of the first gill arch very feebly developed,
the rakers about two, along its base. This genus was created by Eigenmann for heckelii,
a species in many ways intermediate between Aequidens and Geophagus, and included
in one or other of these genera by other authors. The mouth is small, not greatly protractile; the eye in the posterior half of the head; the caudal emarginate, scaled at
base only. The genus is allied to Retroculus, known only from much further south and
not represented in Guyana.
Acarichthys heckelii were common in Rupununi pools ;in body shape and colour they
were very like Aequidens geayi, but from their general behaviour they were thought to
be a Geophagus (until the gill lobe was examined). The local fishermen often referred
to them as ‘Shau’, the Wapishiana name for Geophagus, or ‘Deburu’, their name for
A e q u i d k , though heckelii was also given the distinctive Wapishiana name ‘Parubai’.
Acarichthys heckelii (Miiller & Trotschel). Parubai
Acara heckelii Muller & Trotschel in Schomburgk, 1848 : 624 (swamps and savannas
of British Guiana).
Geophagus thayeri Steindachner, 1875 :48, pl. 3 fig. 2 (varioustributaries of the Amazon).
(Synonymy given by Eigenmann and recently confirmed by Dr F. Terofal.)
Acara subocularis Cope, 1878: 696 (Peruvian Amazon). (Species on which Regan,
1905 : 557, redescribed this species reexamined at British Museum.)
Acarichthys heckelii: Eigenmann, 1912 :500.
Muterialavailable.Two (81,102mm), Obidos Brazil, coll. Agassiz (the specimens on
which Regan, 1905 : 557, redescribed Acura subocularis Cope) ; 30 (53-112 mm)
278
ROSEMARY
H. LOWE+MCCONNELL
Rupununi District pools, both North and South Savannas, Amazon and Essequibo
drainage.
Material examined in the field. This was the commonest Aequidens-like species in
the Rupununi pools ;thought to be a Geophagus until the gill lobe was examined. There
was great variation in the development of the gill lobe, in some specimens it was hardly
developed at all, suggesting that the erection of a new genus for this species is questionable, though the Geophagus-like appearance and habits distinguish it clearly from
other Aequidens.
Distribution.Widely distributed, and as far as is known the only species of the genus.
CoZour. Grey with a large black spot near the middle of the sides, a black spot on the
anterior dorsal spines; a narrow dark bar from the nape through the eye to the angle
of the preoperculum; dorsal, anal and caudal with hyaline spots.
Breeding habits. Not known, but of a pair caught together from a muddy pool in
the Sandcreek, Rupununi, the ripening male was 10/13 cm long and the ripe female
9/11 cm long; the female contained dark green ova; no difference was noted in colour
between the two sexes.
Food. No stomachs were examined in the field. Preserved specimens contained
bottom debris, chironomid larvae and vegetable debris.
Geophagus Heckel, 1840
Geophagus Heckel, 1840: 383.
(Type species: Geophagus surinamensis Bloch, 1791, as Geophagus alt@ons Heckel,
1840 (one of Heckel's typical species) = Sparus s u r i n a h Bloch, 1791, designated
by Eigenmann & Bray, 1894:621.)
Geophagus are readily distinguished by the downwardly-projecting lobe on the
first gill arch, carrying the gill rakers at its margin, the upper lateral line well separated
from the dorsal (at least 24 scales away), lower lateral lines forked at the caudal, and the
deep preorbital (about twice as wide as eye in adult).
Only two species of Geophagus are at present known from Guyana, G . jurupari
Heckel and G. surinclmensis (Bloch), though Biotodoma cupid0 (Heckel) (which has the
gill lobe but not the deep preorbital) is considered to be a Geophagus by some authors.
There are, however, a number of other Geophagus species in neighbouring territories,
which may one day be found in Guyana, G. camopienn's Pellegrin from French Guiana
(greatly resembling young G. surinamensis),G . daemon Heckel and G . acuticeps Heckel,
both from the Amazon, and G . wavrini Gosse, 1963, described from the Upper Orinoco.
A number of other Geophagus species from much further south are among fish being
transported by aquarium dealers which might escape into Guyana's waters.
Key to Geophagus species found in Guyana
la. Dorsal and anal scaled at base in adult ;caudal densely scaled to near its tip ;
a large black spot on middle of sides; D XVII-XIX; gill rakers about ten
on lower part of anterior arch
.
. 6.surinamensif
CHICLID
FISHFS
OF GUYANA
279
b. Dorsal and anal naked in adult; no black spot on sides but usually a small
black spot at base of upper caudal rays; D XIV-XVI; gill rakers 14-16 on
lower part of anterior arch .
.
G.jurupari
Of the species from neighbouring countries G. wavrini Gosse is distinguished by
having four to six gill rakers, and G. daemon and G. acuticeps both have 19-21 gill
rakers on the lower part of the anterior arch, as opposed to 6-17 in the Guyana species
and G. campoensis. G . campoensis is distinguishable from G. surinamensis, which it resembles, by having fewer dorsal spines (D XV-XVI) and less numerous scales (30-31
compared with 33-36 along a median series in surinamensis).
Geophagus species kept by aquarists include G. jurupari, G. surinarnensis (and G.
(=Biotodom) cupido, also G. acuticeps, G . australe Eigenmann and G . gymnogenys
Hensel (southern species) and G. brasiliensis Quoy & Gaimard (from eastern Brazil).
Many of these are bred in aquaria, and the interesting situation exists that some appear
to be mouth-brooders and some not. Reid & Atz (1958) reviewing the literature, think
that both G . jurupari and G. surinamensis are mouth-brooders, but that G. brasiliensis
is not. (B.cupido also appears to be a mouth-brooder). In G.jurupari the female is said
to brood the young (Sterba, 1962: 713), but in G. acuticeps, G. brasiliensis, G . gymnogenys and G.surinamensis both sexes are said to share brood care (Sterba, 1962).Myers
(1939) pointed out how easily a substratum spawner can become a mouth-brooder by
retaining the eggs or young for increasing lengths of time in the mouth when moving
them from place to place.
Geophagusjurupari Heckel
Geophagus jurupari Heckel, 1840: 392 (Rio Negro); Eigenmann, 1912: 504, pl. 67
figs 1-3 (coastal rivers and Rupununi) ;Puyo, 1949 :246.
Geophagus leucostictus Miiller & Trotschel in Schomburgk, 1848: 625 (L. Amucu,
Rupununi savanna swamps).
Geophagus macrolepis Giinther, 1862 : 3 14.
Material available. One (27 mm) Guiana, coll. Schomburgk (type of G. macrolepis
Gunther) ; 18 + small ones (40-144 mm) Rupununi District, Essequibo and Amazon
drainages.
Material examined in jield. About 200 from Rupununi District; four from
Georgetown.
Distribution. Amazon and Guianas. In Guyana this is the commonest Geophagus
in the Rupununi District ;it is said to be fairly common in the coastal rivers where it is
called ‘Creekmouth Patwa’, but it was only once trapped. Eigenmann (1912: 505)
records it from numerous localities. Puyo (1949: 247) says that in French Guiana
G.jurupari is much rarer than G. surinamensis.
Colour. Variable ;general body colour pearly grey, sometimes with iridescent peacock
scales on the back, and with a characteristic small black spot at the base of the upper
caudal rays ;head and operculum tan or chocolate brown spotted with chalky white or
light peacock, and with two or three chocolate brown bands between the eyes and
280
ROSEMARY
H. LOW-MCCONNELL
across the snout; vertical fins immaculate or spotted with translucent spots at base;
pelvics with elongated outer ray cream colour; iris yellow ochre.
Growth. Specimens from 5 to 19 cm standard length were examined and the smallest
ripe fish seen were 15/18 cm; there was little difference in size between the sexes.
Food. Few stomachs were examined in the field and most were empty ; a specimen
trapped at Pirara contained an insect. In aquaria this species works through the bottom
soil for small worms, insect larvae and crustacea, and in the field they can be seen
feeding on the bottom of muddy pools.
Breeding semolfs. Many G. jurupari in Karanambu ponds near the Rupununi River
were ripe at the end of the dry season (April/May), and small fish (50-90 mm) were
taken in September at the end of the rains, suggesting that this species spawns mainly
in the rains.
Breeding behaoiour. G. jwupari breed in aquaria and the female (Sterba, 1962: 713)
or both parents (Reid & Atz, 1958: 79) brood the young in the mouth, for ‘about 14
days’ (Sterba) or ‘up to 37 days’ (Reid & Atz: 85). Eigenmann (1912: 506) collected a
specimen with young in her mouth. This mouth-brooding habit is unusual among
South American cichlids, and not all Geophagus are mouth-brooders. Reid & Atz
state that at least ten different accounts of the breeding of Geophagus brasiliensis in
captivity agree that the eggs and young are cared for by both parents ‘in typical cichlid
fashion’, though occasional reports describe specimens collected in the field with
young in the mouth. Certain features of mouth-brooding in G. jurupariappear unusual,
for instance Reid & Atz, who watched ten spawnings of G. jurupari in aquaria, state
that they waited for practically 24 hours before picking up and incubating the spawn
(instead of picking it up at once as do the African mouth-brooding cichlids). Both
sexes cleaned the surface on which the eggs were laid in batches and fertilized in
sequence by the male; the eggs were guarded by both parents for about a day, then
picked up and incubated orally by both parents or the female alone. One- or two-day-old
young were released into depressions dug into the bottom, and were transferred back
and forth between male and femaIe. Free-swimming young returned to the mouth of
one or other parent (or were gathered by the parents) when disturbed or at night for as
long as 37 days. Unlike other orally-incubating cichlids studied in aquaria, G. jurupari
fed while brooding, possibly supplying food to the young in the mouth; Reid ik Atz
pointed out that G. jurupari young have relatively little yolk compared with young of
the African oral-brooders.
In Guyana Geophaguswereneverobserved spawning, but adult G.jurupariseined from
one of the Fisheries Division ponds in Georgetown (17 April 1958) spewed out yolked
young into a bucket. These young secreted sticky threads from a median small knob
on top of the head, above the eye, which was evidently an attachment gland. It was
puzzling to find these glands, comparable with those known from substratum spawners,
in a fish embryo which is incubated orally, and this seems to be the first record of such
a gland in the young of this species, though very transient, non-functional glands have
been recorded in mouth-brooding Tilapa (Fishelson, 1966). These G. jurupari adults
later proved to be one male and two females (one recently spent) so it was not known
which sex spewed the young from the mouth. Both adults and young were transferred
to a shaded aquarium at 09.45 hours; at 11.00 hours no adults were brooding but a
CICHLID
FISHES
OF GUYANA
281
hole had been made in the bottom sand and some( not all) the young were in this hole.
Shortly after this the adults died, SO it was not possible to observe the normal course of
events.
Geophagus jurupari by the nature of their feeding habits tend to live in pools where
the bottom is of soft deposits, likely to cloud the water when disturbed, which may
have influenced the development of oral incubation in this species.
Geophagus surinamensis (Bloch)
Sparus surinamensis Bloch, 1791 : 277 ; pl. 275 fig. 2 (Surinam).
Geophagus surinamensis: Muller & Trotschel in Schomburgk, 1848 : 625 (Lakes
Tapacuma, Amacu and savanna swamps, British Guiana) ;Eigenmann, 1912: 503, pl.
66 fig. 3 (Demerara, Essequibo and Mazaruni Rivers); Puyo, 1949: 245.
Material available. Two (150,174 mm) Guiana, coll. Schomburgk; five (76-91 mm)
Mazaruni R., coll. Carter ; 14 (60-173 mm) Rupununi District, Essequibo and Amazon
drainages.
Material examined in$eld. About 50, all from Rupununi District.
Distribution .North-eastern South America ;very common in French Guiana (Puyo,
1949). In Guyana Eigenmann collected this species from Demerara, Essequibo and
lower Mazaruni Rivers. It was not seen in coastal plain waters during this recent work,
but was quite common in savanna pools in the Rupununi, in both Essequibo and
Amazon drainage areas.
Colour. One of the most colourful of the larger cichlids. The characteristic black
lateral spot is set on a background of orange-yellow stripes along the body, with peacock blue stripes on a deep red background on pelvics, caudal, hind dorsal and anal
fins. Adult males have blue spots on operculum and cheek, and iridescent green between
the orange stripes along the body. Sterba (1962: 713, fig. 1102) shows a black band from
nape to eye, but this was not noticed in Guyana specimens though these had a black
line down the angle of the preoperculum. The young have the black lateral spot but no
yellow stripes.
In young specimens of G. surinamensis the caudal is not as densely scaled as in the
adults making the young fish resemble more than ever G. camopiensis from French
Guiana.
Breeding. No data obtained. A specimen with young in its mouth was caught at
Rockstone by Eigenmann (1912: 504), and Puyo (1949: 246) records two females with
eggs in the mouth.
Geophagus surinamensis makes a grating noise with its pharyngeal teeth.
Biotodoma Eigenmann & Kennedy
Mesops Gunther, 1862 : 3 11 (preoccupied).
Biotodoma Eigenmann & Kennedy, 1903 :533. (Substitute name for Mesops and taking
the same type species, Geophagus cupid0 Heckel, 1840, but subsequent designation by
Eigenmann & Bray, 1894 : 621 .)
282
ROSEMARY
H. LOWE-MCCONNELL
Eigenmann & Kennedy separated this genus from G q h a g u s by the lateral line not
forked on the caudal, and the preorbital about equal to the eye in adult (contrasted with
lateral line forked at the caudal, and preorbital in adult about twice as wide as eye in
Geophagus). Present studies suggest that the lateral line is not always forked at the
caudal in Geophagus, and the preorbitalleye size relationship changes with growth,
so these are not very good generic characters. Nevertheless, B . cupido is very unlike
Geophagus in general appearance, and might be taken for an Aequidens in the field until
the gill lobe is examined. Further evidence that Biotodoma is distinct from Geophagus
has come from recent egg-structure studies by Wickler (1956), who has shown that the
eggs and egg attachment filaments are of quite different types in B. cupido and G.
brasiliensis. There appears to be only one species of Biotodoma.
Biotodoma cupid0 (Heckel)
Geophqus cupido Heckel, 1840 :399 (Rio Negro).
Mesops cupido : Gunther, 1862 : 3 1 1.
Biotodoma cupido : Eigenmann, 1912 : 501.
Material available. One (75 mm) Essequibo R., coll. Ehrhardt; two (52, 56 mm)
Wismar, Demerara R., coll. Pinkus; two (46, 50 mm) Pirara Creek Stop-off.
Distribution. Middle Amazon basin, western Guiana, now an aquarium species
(Sterba, 1962: 710). Eigenmann (1912) collected this from the Essequibo and Potaro
Rivers and Lama Stop-off. During recent studies it was not distinguished in the field,
and was found only in collections from the Rupununi.
Colour. B. cupido is characterized by a large target-like black-centred ocellus below
the dorsal fin on the hind end of the upper lateral line ;black stripe from nape through
eye to angle of preoperculum.
Breeding habits. Eigenmann & Kennedy (1903) say that the name Biotodoma alludes
to their habit of carrying the young in the gills, i.e. this species is evidently a mouthbrooder.
Cichlasoma Swainson
Cichlaurus Swainson, 1839 : 173 (name used only in key).
Cichlasoma Swainson, 1839 :230.
(Type species : L a b punctatus Bloch, 1792, by monotypy Swainson 1839 : 231 ;
Labrus punctatus Bloch = Cychlasoma bimaculatum (Linn.), Eigenmann & Bray, 1894:
615.)
There has been considerable controversy concerning the correct name of this genus
(e.g. Trevassos, 1953), as on an earlier page of the same work Swainson gave a key
referring to C i c h l a s m as Cichlaurus. When brought to the International Commission,
the Commission’s advisers declared that under the First Reviser Rule 24(a) of the
International Code, Cichlasoma should stand, as the first reviser, Swain (1882) declared
Cichluum unworthy of attention. Bailey (1957) puts forward this view. (Gill’s (1858:
377) redefinition of Cychlasoma, Swainson (Emend.) omitted any reference to
Cichlaurus.)
CICHLID
FISHES
OF GUYANA
283
Cichlasomaare rather like Aequidens in general shape but immediately distinguishable
by having more than three anal spines ;gill rakers are short ;mouth small, premaxillaries
not greatly protractile ;the preoperculum is entire and the lateral line well separated
from the dorsal.
So far only three Cichlasomaspecies are known from Guyana (plus Petenia spectabilis
considered by some authors to be a Cichlasoma).This genus is particularly well represented in Central America.
Field key to Cichlasoma species found in Guyana
la. Pelvics situated in front of origin of dorsal, body increasing in depth to
below posterior part of dorsal fin ; a conspicuous dark band from the snout
to the end of the first dorsal rays, a prominent black ocellus at base of upper
.
.
C. festivum
caudalrays
b. Pelvics inserted behind origin of dorsal
.
. 2
2a. D XIV-XVI 9-11, A IV-VI 8-9; scales 24-25 in median series: colour
becoming dark with age, but with a large dark spot at base of upper caudal
rays and a bold black blotch about the middle of the sides . C . bimaculatum
b. D XV-XVII 13-14, A VII-VIII 12-14; scales 30 in median series; colour
very variable but with a series of seven bands on the sides, the one from
dorsal to end of anal (sixth) the heaviest and continued on bases of dorsal and
anal fins .
C. severum
Two other Cichlasoma species which may one day be found in Guyana are C. coryphaenoides(Heckel) known from the Rio Negro, and C.psittacum (Heckel) from Orinoco
and Brazil. The latter is readily distinguished by having the lateral line scales larger
than those below it on the sides of the body, and a lengthwise band from eye to caudal
base where it forms a spot. C. coryphaenoides (‘Chocolate cichlid’ of aquarists) is
brownish with obscure darker cross bars, a blackish blotch or vertical bar above the
lateral line and below the 11-13th dorsal spines, and blackish fins; D XVI 12-14, A
VI-VII 9-11 ;scales 31-32 in a median series.
Cichlasma bimaculatum (Linnaeus)
Common names. Common or Congo Patwa
Sciaena bimaculata Linn., 1754: 66 (copied).
Labrus bimaculatus Linn., 1758: 285.
Cichlasoma bimaculatum: Pellegrin, 1903: 204; Eigenmann, 1912: 495 ; Puyo, 1949 :
241.
Material available. 20 (41-156 mm) as listed by Regan, 1905: 69; two (60-86 mm)
Georgetown trenches, coll. Eigenmann; three (10, 11, 14 mm) New Amsterdam, coll.
Matthey; 23 (35-88 mm) Rupununi District.
Material examined in field. 204 from Georgetown area; 175+ from Rupununi
District.
ROSEMARY
H. LOWE-MCCON’NELL
284
Distribution. A robust and widely distributed species found throughout most of
northern South America. I n Guyana this appears to be one of the commonest cichlids,
both in fresh waters of the coastal plain and in the Rupununi.
C o h r . Well known from aquarium literature as the ‘Two-spot cichlid’ (Sterba,
1962: 691, fig. 1072). Colour very variable, becoming darker with age; ground colour
greenish brown with a large bold black blotch about the middle of the side, and another
black spot at the base of the tail, with an indefinite darker band from eye to lateral
blotch, traces of eight or nine vertical dark bars, and a dark spot below the eye ;pectoral
web lemon yellow becoming more intense in the male ;fish becomes brighter at spawning
time, the belly becoming golden yellow. The development of pigmentation in the young,
and changes of colour with emotional state in this species have been studied by Baerends
and Baerends-van Roon (1950: 60, figs 25 and 18).
Growth. C. bimaculatum is a round-bodied solid fish, growing up to 12/14 cm,
110 g weight. The relative sizes of the two sexes trapped in Georgetown trenches
were as follows :
Standardlength(cm)
5
6
No. males
No. females
2
3 10 13 15 16 23 11
7 15 13 16 18 7 0
7
7
8
9 10 11 12
Total
93
83
Males tend to be slightly larger than the associated females, e.g. in a spawning pair
trapped together the male was 12/14-5 cm, 95 g, the female 10/12*5cm 65 g. Females
first mature when 719 cm long (about 20-25 g) ; in males it was difficult to determine
the maturation size.
Most of the Rupununi fish [measured were young of 3-7 cm in September 1957,
probably the product of spawning early in the rains.
Standard lengths are much more reliable than total lengths as many of these fishes
had damaged caudal fins.
Breeding seasons. In the traps set near Georgetown, ripe females were taken every
month of the year except December (when no samples were taken) and February
(when few fish were caught, but these included ripening fish). Too few data were
collected from Rupununi fishes to determine whether breeding there was more
seasonal.
Breeding behaviour. This species was not seen spawning in the wild, though it
spawned in aquaria in Georgetown. It is often bred in aquaria, a 12 cm female lays up
to 700 eggs at a spawning, and both sexes guard the eggs and young.
Food. Stomachs and intestinal contents of 101 C . bimaculatum were examined, all
from the Georgetown area. These contained varied items : eight contained mollusc
shells, four palaemonid shrimps, ten insects (including chironomid larvae in two,
Odonata nymphs in two, Hemiptera in one, and aerial insects-cicada, caterpillar and
beetle in three), four had fish scales and bones, three had vegetable debris; there were
filamentous green algae in 12 and seeds in two. Many of these items could be picked
up as bottom debris. Thus C. bimaculatum appears to be an omnivorous feeder, eating
CICHLID
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285
algal filaments, bottom debris and various invertebrates. Most alimentary canals
contained several of the above items if they were full of food.
Predators and parasites. The caudal fins were often damaged and appeared to have
been bitten off. C. bimaculatum were found in stomachs of Hoplias malabaricus and
Cichla ocellaris. Nematode and trematode worm stages were often abundant in this
species.
Cichlasoma f e s t i w m (Heckel).
Common name. Flying Patwa
Herosfestivus Heckel, 1840 : 376 (Guapore).
Mesonauta festivus: Eigenmann & Bray, 1894: 619; Eigenmann, 1912: 498.
Cichlasoma festivum : Regan, 1905 : 69.
Materialavailable. One (102 mm) Demerara, coll. Beckford; two (26, 70 mm)
Cuyuni R., coll. Carter; one (58 mm) British Guiana, coll. Graham; ten (22-62 mm)
Rupununi District (both Essequibo and Amazon drainages).
Material examined in the jield. 85 from Georgetown trenches. Large numbers were
also watched in these trenches when they were breeding, and small shoals were observed
in the Rupununi pools.
Distribution. A distinctive species widely distributed throughout western Guiana
and the Amazon basin, and very common in Guyana, both in trenches on the coastal
plain and in the Rupununi pools. I t is a very hardy species, surviving half an hour out
of water if kept damp (possibly using the stomach as an accessory respiratory organ).
Colour. Well known from aquarium literature (Sterba, 1962: 697). I n the field a
striking black line runs diagonally from the upper lip through the eye to the extended
rays of the dorsal fin (this line appears pale in colour at night by torchlight) ; there is a
large black spot on a clear yellow background on the upper half of the caudal peduncle ;
the body colour above the diagonal line is yellow-brown, each scale having a darker
centre. Vertical fins are grey spotted with yellow, pectoral fins immaculate, the pelvics
with an elongated lemon yellow outer soft ray (which becomes particularly noticeable
in breeding fish) ;the iris is dark brown dorsally and dark red ventrally. T h e breeding
male has bright blue on the lower lip and on top of the eyes ;in the male the snout appears
elongated and more ‘roman’ in shape than in the female; the head has a general
purplish and yellow-green sheen.
Growth. T h e sizes of C. festivum from Georgetown trenches were :
Standard length (mm)
-
No. males
No. females
40 50 60 70 80 90
100 110
___
0
0
Total
_ _ _ -
0 3 12 18 13
2 9 1 0 4 0
2
0
0
0
48
25
The females were smaller than the males, which probably accounts for the unequal
sex ratio, the smaller females escaping through the meshes of the trap. When pairs
were trapped or seen together, the male generally looked about one third as large
again as the associated female (for example in pairs trapped, female SOjl10 mm 25 g,
19
286
ROSEMARY
H. LOWE-MCCONNELL
male 95/125 mm 35 g ; female 80/100 mm 20 g, male 100/135 mm 45 8). Large males
also tend to develop an elongated snout. The smallest ripe females were 70/95 mm 20 g ;
maturation size was more difficult to determine in the male, but a male of 80/110 mm
had the very long and obvious pelvics of a breeding fish. The young fish wander round
the trenches in small shoals of a dozen or so fish.
Breeding seasons. C.festivum were watched spawning when the water was low before
the main rains, and continued to have broods of young through the rains; ripe or
ripening females were caught every month from March to July and in September,
and ripening females were also taken in January. The breeding behaviour is described
below.
Food. Stomachs examined from 33 C.festivum from Georgetown trenches contained
mainly large algae (Closterium, a desmid), Bosmina-like crustacea, bottom debris and
pieces of plant.
Predators andparasites. C. festivum were found in the stomachs of Hoplias malabaricus
and of Cichla ocellaris.Parasites found in the intestine of C.festivum at Sophia included
a new genus and species of oxyurid nematode worm, Ichthyouris Y O . Inglis, 1962.
Breeding behaviour of C . festivum in Sophia trenches, Georgetown
In May 1957 the water was very clear before the rains started.
9 May, 17.30 hours. Apair of C. festivum were found guarding a sugar-cane stem
sloping into the water from the bank.
10 May, 08.30 hours. A mass of small, fawn-coloured, translucent eggs were fixed to
the upper side of the sugar-cane stem; these were guarded by both parents, who took
it in turns to fan the eggs and chase away intruders or peck at the bottom debris,
seemingly for food. The female was also seen to peck at the anal region of the male,
though the significance of this was not clear. The guarding parents kept within 1 m of
the eggs, shooting out at intruders, but backing away from similar-sized C. festivum
when about 0.7 m away from the eggs. The pelvic fins were kept horizontal in the water
(contrast hanging down when leading the young). The male was considerably larger
than the female and had brighter blue on top of his eyes and on the lower lip, otherwise
their colours were very much the same.
11 May, 15.00hours. About eight of the eggs were white andopaque; there were then
three pairs of C.festivum all with broods of young within about 20 m of the pair with the
eggs12 May, 13.00 hours (ie. two to three days after the eggs were laid). The young had
hatched and were about 3 mm long, black with fawn-grey remnants of yolk sac ; they
were all concentrated into a wriggling mass about 5 x 2.5 cm, heads fixed to the stem
and tails waving, the whole looking like a shaggy sea slug. This bunch was concentrated
at the top of the area formerly occupied by the egg mass and occupied a smaller area
than had the eggs; they were about 10 cm below the water surface, with no cover
above them and with male and female on guard below.
13 May, 09.00 hours. The bunch was even more concentrated, into an area 2.5 x 2
cm, now higher up the stem, within 2 cm of the water surface, the top of the bunch
just below the surface when the wind blew, with the female parent on guard.
CICHLID
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287
14 M a y . After 13-14 hours of heavy rain, the water level had risen in the trenches
and the place where the young had been was now some 20 cm below the surface. It
was hard to see the young, but the parents showed them to be on another, horizontal
piece of suger-cane, above the other and only 5-8 cm below the water surface. The
young were scattered and lying attached to the top of the sugar-cane; they were still
black in colour but now with glistening eyes and small fawn yolksacs. One was seen
in the water between the stick and the parent’s mouth; then the parents (possibly
disturbed by my arrival) moved the young down.
15 M a y , 19.00 hours. Cane cutting was in progress, debris had fallen into the trench,
and this family of fish had disappeared.
Other family parties of C. .festivum were watched in Sophia trenches in May 1957,
June 1959, and in April and June 1961. I n all cases both parents guarded the cloud of
young, which kept above them in the water. These young were made conspicuous by
the iridescent eyespot and air bubble in the swim bladder, which shone as though
painted with luminous paint. There were estimated to be about 100 young in most
broods (100-200 in some, less than 100 in others). These young generally kept in a ball
about 30cmindiameterandfollowedoneoftheparents,whichprecededthemwithspecial
jolting movements of the elongated light-coloured, lemon-yellow, pelvic fins. These
pelvics hung vertically downwards in the water in brooding fish (in contradistinction
with the horizontally held pelvics when fish were guarding eggs). As the young
grew, this ball become larger, the young though fewer in number scattered further
apart. The male and female parents took turns at leading the young and chasing away
intruding fish, of their own or other species. At other times the young seemed to lead
and the parents to follow. Adult C. festivum swim backwards and forwards with almost
equal ease when swimming with their young, which enables the two parents to travel
below them facing in opposite directions. T h e ocellus on the base of the caudal peduncle
is very conspicuous in this species as the parent’s tail is waved gently ahead of the young,
and, as in Cichla ocellaris, it seems that this ocellus may play an important part in
orientating the young to the parent. T h e blue on the tops of the eyes and upper lip was
also easy to see from above, from where the young get their view (but possibly too far
ahead for the young to see).
The young were all the same size in one batch (perhaps about 6 mm long) and the
whole family party of parents and young wandered for some 5 m along the trench,
under and around submerged plants such as Pontaderia and Utricularia. T h e parents
at this stage seemed to be guarding the young quite independently of any territory,
whereas the territories guarded by parents before the young were free-swimming
seemed to be less than 1 m long-the parents backing away from other C. festivum at
this distance from their eggs or attached young.
Up to six pairs with free-swimming young and one pair guarding attached young
were seen together in one trench at one time; there were also several shoals of half (40
mm) and threequarter grown (50-60 mm) C. festiwum in the same trench. T h e trenches
also had young Cichla ocellaris, a predator, at this time, but the C.festivum did not seem
unduly worried by their presence-they appeared to use more energy in chasing off
members of their own species ; solitary Acaronia nassa, another predatory cichlid,
lurked near the bottom of these trenches. Various fish-eating birds fish along these
.
288
ROSEMARY
H. LOWE-MCCONNELL
trenches. When disturbed by a shadow falling on the water, the parent C. festivum fled
and the young leapt forward, but then seemed to seek the parents; finding the female
they stayed by her while the male cruised around.This was repeated several times,
when the fishes were disturbed experimentally, until they gained the shelter of the
water plants. When undisturbed, male and female would take turns to peck at the
bottom or pick algae off the plants. At dusk the parents and young tended to remain
static.
Cichlasoma severum (Heckel). Marat
Heros severus Heckel, 1840; 362 (Rio Negro).
Cichlasoma severum : Regan, 1905:322.
Material available. One (120 mm) British Guiana, coll. Schomburgk; eight (85-142
mm) Rupununi District (from both Essequibo and Amazon drainages).
Material examined in the$eld. 25 from Rupununi District only. Few were dissected
as attempts were being made to transport this species for trials as pond fish and for
clearing water weed, since they appeared to be vegetarians. They proved to be hardy
and easy to transport, but it was difficult to find them in sufficient numbers for these
trials.
Distribution. Northern Amazon basin and Guiana; in Guyana found in Rupununi
District but not on the coastal plain; Eigenmann (1912) also records one specimen from
the Lower Potaro River.
Colours. The saucer-shaped C. severum is known as an aquarium fish (‘Banded
cichlid’, Sterba, 1962:702, fig. 955). In the field the colours are very variable and change
rapidly with emotional states. Basically the dark grey-green body has about eight dark
greeny-black stripes, the penultimate one joining a black spot on the soft dorsal to a
black spot near the hind end of the anal ;this penultimate band is the most pronounced,
persisting when the other bands fade and on death. These vertical stripes, which
appeared and faded, were also present in a young fish of 45/60 mm, which also had
chocolate spots on the anterior half of the body. The males often show these rows of
spots, which may be red in the adult fish; in some aquarium literature (Axelrod &
Schultz, 1955:624) it is said that such spots are characteristic of the male, but one ripe
female had such spots. One dying fish lost these spots, another developed them. The
iris was bright red to dull orange.
Growth. Fishes examined were in the size range 45/60 mm to 140/190mm. Males and
females appeared to be of comparable sizes. The smallest ripe fish seen, a female
containing bright orange avo, was 125/170 mm long.
Breeding seasons. Ripe females were caught from savanna ponds in April 1957 near
Karanambu. Most of the adult fish caught at Pirara when the fish were moving back
to the main river in September had completely regressed, fully quiescent gonads.
Young C. severum (45160 mm) from near Karanambu, and individuals of 85/115 mm
and 105/130 mm at Jacare, were caught in September. This suggests that breeding
occurs in the rainy season which in this district extends from May to September. In
aquaria this species spawns and guards young in ‘typical cichlid fashion’ (Sterba,
1962: 702), which means that both sexes guard the eggs and young.
Food. In September 1957 at Pirara C. severum contained green algae and chewed
CICHLID
FISHES
OF GUYANA
289
vegetable matter ; in April, fish taken from drying ponds where there was very little
food contained only gritty bottom debris, sand grain and bits of dead plant material
and chitin. By September they had well-developed fat ribbons, indicating that the wet
season is the main feeding time.
Petenia Gunther
Petenia Gunther, 1862 : 301.
(Type species : Petenia splendida Gunther, 1862, by monotypy.)
Cichlasoma-like with more than three anal spines, but with premaxillaries very
protractile, their processes about as long as the head ; maxilla considerably exposed ;
cheek and opercular bones scaly ; outer series of teeth enlarged, canines moderate or
strong.
One species in Guyana.
Petenia spectabilis (Steindachner)
Acara (Petenia) spectabilis Steindachner, 1875 : 36. pl. 4 (Amazon).
Petenia spectabilis: Eigenmann & Bray, 1894 : 615 ;Pellegrin, 1903 :244; Fowler, 1954 :
315.
Cichlasoma spectabile : Regan, 1905 : 339.
Material available. One (150/185 mm) Manari Creek, a tributary of Takatu R.,
Amazon drainage Rupununi District ;no specimens from Guyana previously available
in British Museum, though material available from Venezuela and Amazon.
Material examined infield. Members of this species were watched swimming singly
round rocks in Manari Creek, but proved difficult to catch, disappearing into holes
between the rocks. T h e single specimen obtained was caught on a worm-baited hook.
Petenia spectabilis was not seen elsewhere in Guyana.
Distribution. An Amazon species, found only in the Amazon drainage Rupununi
District of Guyana.
Colour. Brown with a large lateral spot and smaller black spot on upper half of the
caudal peduncle at base of caudal fin.
Breeding. The one specimen caught was a quiescent male. This species has been
bred in aquaria (Sterba, 1962 : 704), where it is said to be quarrelsome and to breed in
‘typical cichlid manner’ (Axelrod & Schultz, 1955 :625) (which means that both parents
guard eggs and young). T h e sexes can be distinguished by the elongated posterior rays
in the dorsal and anal fins in the male.
Food. A predacious species, with highly protrusible mouth.
Crenicara Steindachner
Crenicara Steindachner, 1875 : 39.
(Type species: Crenicara elegans Steindachner, 1875, by original designation and
monotypy .)
Short-bodied cichlids in which the preoperculum margin is serrate; body ovate;
scales of lateral line of the same size as those above and below it; mouth small, jaws
equal. One species in Guyana.
290
ROSEMARY
H. LOWE-MCCONNELL
Crenicarapunctulata (Gunther)
Acara punctulata (part) Giinther, 1863:441 (Essequibo).
Crenicarapunctulatu: Pellegrin, 1903: 169; Regan, 1905: 152; Eigenmann, 1912: 511.
Crenicara elegans Steindachner, 1875:39, pi. 1, fig. 1.
Material available. The type (76/104 mm) from R. Essequibo, coll. Ehrhardt.
Distribution. C .punctulata, recorded from Amazon and Guianas, was not seen during
recent studies. It appears to be rare. Eigenmann (1912) took only one specimen (68
mm), from Gluck Island, Essequibo River.
Colozcr. The type in the British Museum is a brownish fish with a series of darker
blotches on and above the lateral line and another more distinct series below the lateral
line ;a white-edged dark stripe from eye to mouth ;a dark lateral stripe from operculum
continues along the middle of the caudal to its tip : soft dorsal and middle dorsal rays
cross-barred ; anal with a blackish edge.
Biology. Nothing is known of the biology of this species. Another species, Crenicara
maculata (Steindachner) from the Middle Amazon is bred in aquaria (Sterba, 1962:
705, fig. 915).
Batrachops Heckel
Batrachops Heckel, 1840 : 432; Eigenmann, 1912: 512.
(Type species : Batruchops reticulutus Heckel, 1840, by subsequent designation of
Eigenmann & Bray, 1894:620.)
Very similar to Crenicichla,with elongated body, serrated preoperculum, and lateral
line scales larger than those above and below it, but, unlike Crenicichlu, Batrachops
has non-depressible inner teeth.
There appears to be only one species in Guyana, B.punctulatus Regan, though there
is some confusion between this species and B. reticulatus (in part).
Batrachops punctulatus Regan
Crenicichla reticulata (not of Heckel), Pellegrin, 1903:378 (Essequibo).
Butruchopspunctulatus Regan, 1905: 156, pl. 14, fig. 1; Eigenmann, 1912: 512.
Material available. One (117/140 mm) Essequibo R., coll. Ehrhardt (type); three
(57, 121, 125 mm) Upper Cuyuni, coll. Carter; one (57 mm) Cuyuni R., coll. Carter;
two (119, 135 mm )Rupununi River.
Material examined in field. None.
Regan described B. punctulatus from one specimen only, a shallow-bodied fish with
a blackish intramarginal band along the dorsal. Eigenmann (1912) ‘collected two specimens (68, 128 mm) from Crab Falls on the Essequibo which he considered belonged
to this species, and with which he synonymized Pellegrin’s C . reticulata, although
the dorsal had aseries of black spots (as in C . reticulatu but not in Regan’s B. punctulatus
type). Differerences in markings of the dorsal fin might be due to growth stages or a
secondary sexual character (as in Crenicichla alta, see below) ; attempts to determine
the sex and gonad states of Batrachops in the British Museum material were not successful, but the two small specimens collected by Carter showed both an intermarginal
CICHLIDFISHES
OF GUYANA
291
dark band and a horizontal series of dark spots below this band, as well as a few other
spots at the hind end of the fin.
A specimen of ‘B.reticulatus’ from the Rio Negro, type area of B . reticulatus Heckel
(B.M. 1909.8.24.2, 115/138 mm, presented by Sutton) also seems to be identifiable
with B. punctulatus Regan (though it is a slightly deeper-bodied fish and has a series
of black spots on the dorsal). However, I have not examined Heckel’s types of B.
reticulatus; Regan (1905 : 155) synonymizes Crenicichla elegans Steindachner from the
Peruvian Amazon with Heckel’s type (a 250 mm fish), which suggests that both B .
reticulatus Heckel and B. pumtulatus occur in the Rio Negro.
Crenicichla Heckel
Crenicichla Heckel, 1840: 416; Eigenmann, 1912: 513.
(Type species. Craicichla macrophthalma Heckel, by subsequent designation of
Eigenmann & Bray, 1894: 620.)
Elongate fishes with margin of preoperculum denticulate, lower jaw projecting
and teeth of inner series depressible; scales of lateral line larger than those above or
below it.
Eigenmann (1912) listed five species of Crenicichla from British Guiana and for
which he gave a key (p. 513). These are C. saxatilk (Linn.), C. alta Eigenmann, C .
wallacii: Regan, C . lugubris Heckel and C .johanna Heckel. Regan (19134 described another small species, C. nana, included in Eigenmann’s collections of wallacii: (and appearing to differ from this mainly in the height of the dorsal fin) the type appears unique
and this may not be a valid species. Fowler (1914) described C. pterogramma from the
Rupununi District ;this appears to be a colour form of C . alta. Lowe-McConnell(l964)
added to Guyana’s list C. strigata Gunther and C. ornata Regan (uith which C. Zenticulata was then thought to be synonymous). Re-examination suggests that C. ornata
and C . lenticulata are both valid species, but that the ‘ornata’ collected in the Rupununi
may be a new colour form. The many varieties of C . brasiliensis Bloch figured by
Pellegrin (1903 : fig. 42) indicate the complexity of this group of small-scaled species
of Crenicichla.
Field key to Crenicichla species found in Guyana
. 2
b. Scales smooth (cycloid), embedded; no caudal ocellus; scales 97-108 below
la. Scales rough (ctenoid), at least on sides of body below lateral line
lateral line
C. johanna
.
2a. Scales large, 38-72 below lateral line
.
b. Scales smaller, more than 72 (72-130) below lateral line .
.
.
3
6
3a. Dark humeral spot present on or below lateral line ; maxillary extending to
beloweye
.
.
4
b. No humeral spot ;maxillary not extending to vertical from anterior margin
of eye (small species) .
.
5
ROSEMARY
H. LOWE-MCCONNELL
29 2
4a. Humeral spot entirely below the upper lateral line, frequently ocellated ;a
band from eye to humeral spot .
C. saxatilis
b. Centre of humeral spot on lateral line, except in the very young; a band
from chin or eye to tip of operculum (continued to middle caudal rays, running below the humeral spot in some, generally young, specimens) C. alta
5a. Last dorsal spine half head length
.
b. Last dorsal spine scarcely more than one third head length
. C. wallacii
. C . nana
6a. Scales above lateral line ctenoid (except anteriorly) ; soft dorsal 15-17 ;
scales 93-113 .
. 7
b. Scales above lateral line nearly all cycloid ;soft dorsal 17-19 ;scales 112-130 ;
head and chest spotted with black, sides of body unspotted with a series of
narrow vertical bars above lateral line; dorsal, anal and caudal fins edged
with black; caudal with one large black spot, above and touching lateral
line, otherwise unspotted .
. C. ornata
7a. Scales 93-108, 14-16 between spiny dorsal and lateral line ;two dark stripes
along body (outlining dark band in young) ending in a dark spot on base
of caudal, a third stripe along upper lateral line, a fourth above this, along
.
base of dorsal, connecting a series of spots or rings on each side
. C . strigata
b. Scales 106-113, 16 or 17 between spiny dorsal and lateral line; brownish
in adult with a dark spot above pectoral and another at base of caudal
. C. lugubris
From the literature it seems that some of the following species could occur in Guyana :
(A) Large-scaled species
C. macrophthaZma Heckel, known from Amazonia and Venezuela; a species with
62-70 scales and lacking a caudal ocellus; (thought by Regan, 1913 : 502, to include
C. suntaremensis Haseman, 1911, pl. 62, fig. 1).
C.geayi Pellegrin, 1903 (339, pl. 6, fig. 4) from Orinoco and Amazonia ;54-57 scales ;
(Schultz, 1949, thought that Venezuelan specimens should be reexamined to see if
this species is confused with C. macrophthalma, but C. geayi has a distinct caudal
ocellus, and a smaller eye).
C. notophthalma Regan, 1913, a small species from Manaus. Both this and the rather
similar small C. dorsocelZata Haseman, 191 1, from the Middle Amazon, generally
have a distinctive large dark spot or ocellus on the dorsal fin; this is, however, not
present in all the British Museum specimens, and from personal observations made
in Brazilian waters in 1968 it seems that this dorsal spot in Crenicichluis often a juvenile
character.
(B) Small-scaled species
C . multispinosa Pellegrin, 1903, from French Guiana and Surinam ; scales 90-108,
with a blackish ocellus on upper part of caudal base.
CICHLID
FISHES
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293
C. ternetxi Norman, 1926, from French Guiana; scales 83-86, lacking a caudal
ocellus.
C . lenticulata Heckel, from Manaus; scales 112-130; at one time thought to be
synonymouswith C. ornata, but the colour pattern is very different (sides of body heavily
spotted between and around larger dark blotches on and below lateral line; caudal fin
spotted with small black spots in addition to largeblackspot). Regan( 1905)distinguished
C . lenticulata from C . ornata by scale number between first dorsal spine and lateral
line (15 or 16 in C. lenticulata; 18-20 in C. ornata), also the maxillary extending to
below anterior margin of eye in C . lenticulata and a little beyond it in C . ornata.
These colour forms of the lugubris-ornata complex are puzzling, and we may be
dealing here with a polychromatic species. Pellegrin, 1903: 383 fig. 42, figured several
of these (including lugubris, lenticulata and strigata) as well asjohanna all as colour
varieties of C. brasiliensis (Bloch).
Crenicichla saxatilis (Linnaeus)
Sparus saxatilis Linnaeus, 1758:278 (Surinam).
Perca saxatilis: Bloch, 1792: 79, pl. 309 (Surinam).
Crenicichlasaxatilis: Heckel, 1840:432 ;Muller & Trotschel in Schomburgk, 1848:626
(British Guiana, all rivers) ;Eigenmann, 1912: 513.
Material available. 11 (89-226 mm) from various parts of Guiana listed by Regan,
1905 ;three (93-100 mm) lowlandsof Br. Guiana, coll. Eigenmann; seven (85-138 mm)
Demerara R., coll. Rosenberg; five (100-150 mm) Berbice, coll. Matthey; 12 (65-122
mm) Mackenzie, Demerara R., coll. Carter; four (45-55 mm) Tumatumari, Potaro R.,
coll. Carter; one (117 mm) Sophia trenches, Georgetown; two (59, 62 mm) Cajuero,
Rupununi drainage ;four (92-108 mm) Jacare, Rupununi District, Amazon drainage.
Material examined in jield. About 20, mostly from Sophia; (Rupununi data are
excluded as the specific identification in the field was not always reliable in this area).
Distribution. Widely distributed through tropical South America, this is the commonest species of Crenicichla in the fresh waters of the coastal plain of Guyana.
Colours. Greyish with a black humeral spot, often outlined with silver in breeding
fish, entirely below the lateral line ; a dark band from the eye to the humeral spot and
frequently forward to the chin; a caudal spot at the base of upper rays ; sides frequently
with light, silvery or light blue spots which form an incomplete ring around humeral
and caudal spots ;vertical fins spotted with light spots ;a dark spot below the eye ;young
with a faint lateral band. In a breeding pair trapped together, the male had iridescent
silvery scales scattered all over the body (not present in the female) and round the
humeral spot ;belly white, bones below cheek pinkish, pectorals yellow, pelvics pink,
silvery light spots on hind dorsal and anal, and hind rays of these fins much elongated.
In the female the iridescence was restricted to round the humeral ocellus ; the enlarged
broad belly shape was distinctive and the belly was dark red in colour, genital orifice
white; no silver flecks on anal or hind dorsal, but the dorsal had a submarginal white
band above a black band 2 mm from the edge (these bands absent in the male). (However,
an aquarium specimen examined later showed a white submarginal band along the
dorsal in a male with well-developed testes.)
20
294
ROSEMARY
H. LOWE-MCCONNELL
Growth. The C . saxatilis size range in the Sophia traps was 370/460 nun to 180/210
mm (150 g). I n a breeding pair the male was larger than the female (male 170/210 mm
125 g, female 150/180 mm 95 g). Both sexes appear to mature when 150/180 mm long.
The ripe ova are bright orange, elongated and 1-2 mm along the long axis.
Breeding. Ripe fish were trapped in March, before the rains, and in October. Breeding
behaviour was not observed personally ;the parents are reputed to move the eggs and
young up and down with the water level (as observed in C.festiuum,p. 286). In apuaria
spawn is deposited in a shallow pit. Sterba (1962 :708) says breeding in this species is as
in C. dorsocellata, a species in which brood care is undertaken mainly by the male.
Food. Most specimens examined were empty, but items such as palaemonid shrimps
and the elytrae of aquatic Hemiptera were found in some C . saxatilis guts. This is a
very hardy species which will live for at least half an hour out of water if kept damp, and
the indications are that it uses the vascular stomach as an accessory respiratory organ
in the dry season.
Enemies. C. saxatilis were found in the stomachs of Hoplias malabaricus and Cichla
ocellaris.
Crenicichla alta Eigenmann
Crenicichla alta Eigenmann, 1912: 516, pl. 68, fig. 3 (Essequibo and Potaro Rivers).
Crenicichlapterogramma Fowler, 1914 : 281, fig. 20.
Eigenmann (1912: 514) reported that he caught two forms of Crenicichla resembling
saxatilis, one along the coast and inland to Rockstone on the Essequibo, the other
inland of Rockstone; the two forms were only found together at Rockstone (Gluck
Island), the boundary between the two areas. He considered that the coastal form was
undoubtedly C . saxatilis of Bloch, and the new species he described as C. alta. This
latter species is most readily distinguished from C , saxatilis by its having the humeral
spot always on the lateral line (Fig. 2F) (even in the young where a lateral band is
sometimes distinct from the humeral spot and sometimes continuous with it), whereas
in saxatilis the humeral spot, frequently ocellated, is entirely below the lateral line.
Material available. Two (96 mm) Nickaparu, coll. Eigenmann (co-types) ; one (62
mm) Rockstone, coll. Eigenmann (co-type); one (94 mm) Gluck I., coll. Eigenmann
(co-type) :one (67 mm) Essequibo, coll. Ehrhardt ; two (139,143 mm) Potaro R., coll.
Bovallius .six (117-127 mm) Potaro R., coll. Carter; six (117-175 mm) Mazaruni R.
coll. Carter; three (129-174 mm) Upper Cuyuni R., coll. Carter; seven (45-168 mm)
Rupununi District (five from Amazon drainage, two from Essequibo drainage).
Material examined infield. None.
Distribution. Eigenmann found very little overlap in distribution between C. alta
and C. saxatilis, C. alta was the inland form, and he drew attention to the need to reexamine Amazonian specimens to see to which species they belonged. Fowler (1914 :
280) recorded one C. alta (163 mm) from the Rupununi, together with C. pterogramma.
During these present studies both C. alta and C . saxatilis were taken in Rupununi
District waters draining to the Amazon (and two probable C . saxatilis from the Essequibo drainage), but no C. saxatilis were found with the C . alta collected by Carter
from the forested rivers of the interior of Guyana. It seems, therefore, that separation
is ecological rather than geographical, C . alta being the species of the heavily forested
CICHLIDFISHES
OF GUYANA
295
rivers, C . saxatilk in waters in more open country. Fowler (1954) omits C . alta from
the list of fishes of Brazil. Schultz (1949: 168) lists one specimen of C . alta ( 5 1 mm)
collected from Caripito, Venezuela by Beebe, and from Surinam Boeseman (1952)
recorded one specimen (114 mm total length) from the Upper Saramacca River.
Crenicichla pterogamma described by Fowler (1914: 281) from one specimen (162
mm) from the Rupununi District, differed from C . alta and C . saxatilis in colour, the
new name referring to the pale submarginal streak on the dorsal fin, so a careful examination was made of the colour of the dorsal fin in all the C. ulta specimens in the British
Museum. This showed that in this species ripening females have a very definite white
submarginal band on the dorsal fin; the breeding male has white spots on the dorsal
fin and no white band. The type of C. pterogramma would appear to be a mature female
C. ulta (though Fowler does not mention the sex).
Many of Carter’s specimens were breeding fishes, still containing the gonads, which
showed that this species matures when about 130 mm standard length; males were
slightly larger than females.
Crenicichla wallacii Regan
Crenicichlawallacii Regan, 1905 : 163, pl. 14, fig. 2 (Essequibo River).
According to Regan (1905e: 163),Wallace had a drawing of a fish from the Rio Negro
which Regan considered to be this species ; the specimens were lost with Wallace’s
other collections before the species was described.
Material awailabk. One (85 mm) R. Essequibo, coll. Ehrhardt (type).
No specimens of this species were seen during recent studies.
Distribution.Eigenmann (1912 :517) records this species from Essequibo and Rupununi ;a small species (25-85 mm) ‘readily distinguished by the presence of, even in the
very young, a distinct caudal spot, and by the short maxillary’.
Crenicichla nana Regan
Crenicichla nana Regan, 1913 :502 (British Guiana).
Regan described this species from two specimens(40,45 mm) received from Professor
Eigenmann as C. wallacii, from which species Regan considered that they differed
especially in the ‘lower’ (i.e. shorter) dorsal spines. Known only from the types (in the
British Museum, Natural History) ;further material is needed to determine the status
of this species and whether it really is distinct from C. wallacii.
Crenicichlajohanna Heckel
Crenicichla johanna Heckel, 1840: 417 (R. Guapore); Eigenmann, 1912: 520 (Potaro
and Demerara Rivers, Lama Stop-off, British Guiana).
Material available. One (90 mm) Upper Cuyuni, ~011.Carter; One (172 mm)
Demerara R., coll. Carter ;two (167,205 mm) Rupununi R. (the smaller one taken from
a crevice in a submerged log).
Material examined infield. Very few.
Distribution. Widespread in tropical South America, Venezuela, Amazon, Peru,
Paraguay, and in Guyana in both coastal rivers (but not trenches) and Rupununi.
296
ROSEMARY
H. LOWE-MCCOWLL
Colour. A very smooth fish with embedded cycloid scales, adult a very uniform grey,
without a caudal ocellus ; young rarely with a submarginal stripe on upper and lower
parts of caudal. Eigenmann (1912 : 520) says ‘young with small black dots on the head
and upper half of the body, arranged into longitudinal lines on the sides ; back with
numerous cross-shades; no caudal spot. Caudal in the young sometimes narrowly
margined with light, then with a broad black crescent, then another light crescent, the
base and the middle of the fin dark’ (description very like that of C . Zugubris and C .
m a t a young except for the absence of the caudal spot).
Crenicichla Zugubris Heckel
CrenicichZa Zugubris Heckel, 1840 : 422 (Rio Negro) ;Eigenmann, 1912 : 5 18.
Materia2 available. Three (150-255 mm) British Guiana and Essequibo, coll.,
Schomburgk (Regan, 1905); four (155-193 mm) Mackenzie, Demerara R., coll.
Carter; one (118 mm) Upper Cuyuni, coll. Carter; three (130-235 mm) Rupununi
District (two from Amazon, one from Essequibo drainage); also two (78, 120 mm)
from Rupununi with colour rather as in C . strigata but possibly this species (see below).
Material examined in$eld. None.
Distribution. A widely distributed species recorded from Amazon, Venezuela, and
Guianas. In Guyana recorded from many rivers (as above), but seen only from Rupununi
District during the present studies.
CoZmr. Adult uniform dark above, somewhat lighter below; a black spot at the base
of the middle caudal rays. Eigenmann (1912: 519) describes young (86-140 mm) as
follows: ‘spots on lower half of cheeks and chin; a narrow line across the chin and
snout to eye and from eye to end of operculum, continued on the side as a broader
but less intense band of black, which may be bordered above and below by light; a
second dark line from the eye along the upper section of the lateral line, a third from
the tip of the snout to between the eyes, then dividing and extending to the end of the
head, from where it is continued as a series of small streaks or spots along either side
of the dorsal ; spots between the second and third stripes on the head ; caudal with a
black spot at the base of the middle rays, surrounded by an arrow-shaped area, this
bordered by a light band above and below, beyond this dark again; base of dorsal
dusky, a submarginal black band’.
However, this description of young C . Zugubris accords very well with four specimens
(86-101 mm) from Manari, Rupununi District Amazon drainage, and believed to be
C . strigata (see below). Also, the smallest of Carter’s specimens of C . Zugubris (1 18 mm)
from Upper Cuyuni, has no trace of a lateral stripe, which seems to support the
possibility that Eigenmann was describing the young of C . strigata, rather than of C .
Zugubris. Eigenmann (1912 : 519) queries Crenicichla johanna strigata Giinther as a
synonym of C . lugubris.
Crenicichla strigata Gunther
CrenicichZa.johanna variety strigata Gunther, 1862. 306 (R. Capin).
CrenicichZa strigata: Regan, 1905 : 165, pl. 15, fig. 1 (R. Amazon).
,
CICHLID
FISHES
OF GUYANA
297
Material available. Two (177, 188 mm), R. Cupin (Amazon), from college of Surgeons (types) ; four (86-101 mm) Manari, tributary of Takatu, Amazon drainage,
Rupununi District; one (78 mm) Morebay Pond, near Rupununi River (possibly C.
lugubris).
Distribution. Amazon drainage ;possibly also in Essequibo drainage (if small specimens described as young C. lugubris by Eigenmann or Morebay Pond specimen are
really C. strigata), contrary to statement of Lowe-McConnell(l964) who then thought
this species restricted to Amazon drainage in Guyana.
Crenicichla ornata Regan
Crenicichla ornata Regan, 1905 : 167, pl. 15, fig. 2 (Rio Negro).
?Crenicichlabrasiliensis var. lenticulata Pellegrin, 1903: fig. 42.
MateriaZavailable. Three (134141 mm) Rio Negro, coll. Antony (types) ;two -t ?one
(172, 202 + 57 mm) Manari, tributary of Takutu R., Amazon drainage, Rupununi
District.
The Manari specimens have 112-121 slightly ctenoid scales below the lateral line,
and 18-20 scales between lateral line and first dorsal spine, as in C. ornata, but the
vertical dark bars above the lateral line are opened out into vertical ellipses, though
the dark colour is still above the lateral line as in C. ornata, not mainly on or below the
upper lateral line as in C. lenticulata.
Pterophyllum Heckel
Pterophyllum Heckel, 1840: 334.
(Type species : Pterophyllum scalare Heckel = Platax scalaris Cuvier & Valenciennes,
by monotypy ;further designated by Ahl, 1928 (2001.
Anz. 76 : 252).)
Plataxoides Castelnau, 1855 : 21, pl. 11, fig. 3 (type species P. dumerdii Castelnau).
Greatly compressed and deep-bodied, with graduated dorsal and anal spines forming
parts of elevated lobes.
The nomenclatural validity of the generic name Pterophyllum has been discussed
by Schultz (1967) following the use of Plataxoides by Gosse (1963:4), although Myers
(1940b : 36) gave reasons why Pterophyllum is the valid name.
According to Schultz (1967) two species occur in Guyana, P . dumerilii (Castelnau)
and P. scalare (Lichtenstein), both occurring in the Rupununi River; a third species
P . altum Pellegrin (possibly only a subspecies of P . scalare) represents the P . scalare
form in the Upper Orinoco basin. Schultz (1967: 7) considers P. eimekeiAh1, 1928,
to be synonymous with P. scalare. There is much overlap in meristic characters of
species and populations, but according to information given by Schultz, the species
may be distinguished as follows :
la. A short black bar from origin of spiny dorsal running ventrally to lateral line,
another short bar halfway between eye and spiny dorsal origin, also a
dorsoventral bar through eye and across cheek
.
. P . dumerilii
b. Black bar through eye extends dorso-posteriorly to origin of spiny dorsal
fin (replacing the two middle predorsal vertical bars on P . dumerilii) . 2
298
ROSEMARY
H, LOW-MCCONNELL
2a. Rarely more than 26 soft dorsal rays
b. Soft dorsal rays 27-31
.
.
. P. scalare
. P. altum
P. scalare occurs in the Amazon basin and in the Rupununi and Essequibo Rivers of
Guyana (with a single record in French Guiana) ; P . dumerilii has been found in the
Amazon basin and Rupununi River; P . altum appears to be confined to the Upper
Orinoco basin.
During these field studies Pterophyllum were seen in the rivers on many occasions,
but the only one trapped came from Crane Pond Creek draining to the Rupununi
River near Karanambu ;this was identified as P. eimekei Ah1 (=P.
scalare).
P. scalare, the Angelfish, is a well-known aquarium fish. I n aquaria this species
spawns on broad-leaved water plants; the eggs are cared for by both parents, which
fan them continuously. Eggs hatch after 24-36 hours (at 26-30°C), the young are
chewed out of the egg shells by the parents and spat ,onto the leaves, where they hang
by short threads. Later the young are brought down into shallow pits on the bottom;
they start to swim after four to five days, and the brood is led out by the parent after a
further two days (Sterba, 1962: 718); rival or courting males are said to emit loud
creaking noises, made by the jaws.
Dwarf cichlids
My field studies were primarily concerned with potential food fishes, so little attention was paid to the dwarf cichlids, many of which have been studied by aquarists. They
were, however, found to be fairly abundant in the leaf litter of certain river pools in
the Rupununi District at the end of the rains. Specimens brought back to the
British Museum (N.H.) proved to be mostly Apistogramma species.
In the dwarf cichlids the upper lateral line runs close to the dorsal fin, a character
which enables them to be readily distinguished from the young of the larger-growing
species. Two genera of dwarf cichlids are known from Guyana, Apistogramma, in
which the first gill arch has a downwardly-projecting lobe with feeble gill rakers along
its margin, and Nannacara, which lacks this lobe and in which the head is densely
scaled.
Apistogramma Regan
Heterogramtnu Regan, 1906 (preoccupied).
Apistogramma Regan, 1913 : 282 (substitute name for Heterogramma (preoccupied by
a lepidopteran) and therefore carrying the same type species Mesops taeniatus Giinther,
1862, designated type species in Eigenmann, 1912: 506).
The following Apistogramma are recorded from Guyana :
A. steindachneri Regan, 1908 : 370 (type locality Georgetown, British Guiana).
A . ortmanni Eigenmann, 1912: 506, pl. 68, fig. 1 (type locality Essequibo R).
A. ortmanni m p n u n i Fowler, 1914: 277, fig. 19 (Rupununi River).
A. matipinnis E. Ahl, 1936: 141 (type locality western Guiana); according to Ah1
this species includes steindachwi (non-Regan) of Eigenmann, 1912: 508, pl. 68,
fig. 2.
CICHLID
FISHES
OF GUYANA
299
Other Apistogramma species from north-eastern South America are known to aquarists, but the genus is in need of revision (Hoedeman, 1951).
The commonest dwarf cichlids in the leaf litter samples from the Rupununi appear
to be A. ortmanni rupununi (25f specimens of 30+ mm standard length) but one or
two other (possibly new) species were present; this material awaits further work on
the genus.
Nunnucara Regan
Nannacara Regan, 1905 : 344.
(Type species :Nannacara unomala Regan, 1905, by monotypy.)
Nannacara are minute cichlids allied to Aequidetls but with the lateral line running
obliquely upwards to the spinous dorsal, from which it is separated by only half or one
series of scales for most of its length.
A few small specimens of Nannacara were found in leaf litter from Karanambu
Creek and Morebay Pool (near Karanambu), Rupununi River (one possibly a new
species). Two species are at present known from Guyana :
Nannacaru anornula Regan, 1905 : 344 (type locality R. Essequibo) ; Eigenmann,
1912: 487, pl. 65, fig. 9, Lama, Maduni Creek, Aruka River.
Nunnucara bimaculata Eigenmann, 1912: 487, pl. 66, fig. 1 (Erukin, Potaro,
Essequibo Rivers), similar to N . anornula but readily distinguishable by its lateral
and caudal spots.
N . anornula from Western Guiana is now an aquarium species (the ‘Goldeneyed
Dwarf Cichlid’). It grows to about 8 cm (Sterba, 1962: 713) ;the colour is very changeable (Sterba, 1962 :788). In aquaria the eggs are said to be laid on a substrate previously
cleaned by either or both parents, and guarded by the female; the young hatch after
two to three days and are gathered by the female into a pit where they remain for about
five days on the bottom (Sterba, 1962: 714).
Another species N . tuenia Regan, 1912, from the Amazon, known as the ‘Lattice
Dwarf Cichlid’ is also kept by aquarists and bred in aquaria, but is not known from
Guyana.
ACKNOWLEDGEMENTS
The Trustees of the British Museum (Natural History) very kindly provided working
facilities and a grant towards expenses during the examination of the material at the
British Museum (Natural History). 1 am very grateful to my colleagues in the Fish
Section there for many helpful discussions during the course of this work and
especially to Dr. P. H. Greenwood for reading the manuscript. In Guyana the field
studies were made when attached to the Fisheries Division, Department of Agriculture, Georgetown, as an Honorary Research Officer, and I am very grateful for the
facilities provided there and to all who helped to collect the material; in the Rupununi
the help and hospitality of Mr & Mrs E. McTurk and many others were invaluable.
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