Bijdragen tot de Dierkunde, 56 (2): 221-231 — 1986

The fishes of the cichlid genus Crenicichla in

(Pisces, Perciformes, Cichlidae)

by

A. Ploeg

Institute of Taxonomic Zoology (Zoblogisch Museum), University of Amsterdam,

P.O. Box 20125, 1000 HC Amsterdam, The

Abstract which included the area around the Mana

River, including the French Guianan border of A review of the of the cichlid Crenicichla species genus the River. Heckel, 1840 in French Guiana is presented. Five species

C. Seven of Crenicichla have been are encountered, viz.: C. saxatilis (Linnaeus, 1758), species

albopunctata Pellegrin, 1904, C. multispinosa Pellegrin, recorded in the literature in French as present 1903, C. ternetzi Norman, 1926, and C. johanna Heckel, Guiana, viz.: C. brasiliensis (Bloch, 1792); C. 1840. Their geographic distribution is indicated. The johanna Heckel, 1840 [including C. j. var. variation ofmorphometric, meristic, and pigment charac- carsevennensis examined. The of these characters is Pellegrin, 1905]; C. lugubris ters are variability

compared with the variability within Crenicichla saxatilis Heckel, 1840; C. multispinosa Pellegrin, 1903;

from the River, Surinam 1986). C. saxatilis C. (Ploeg, (Linnaeus, 1758) [including s.

ternetzi var. albopunctata Pellegrin, 1904]; C. Résumé Norman, 1926; and C. vaillanti Pellegrin, 1904.

On les du de Cichlidés For the the of these passe en revue espèces genre present study occurrence

II de Crenicichla Heckel, 1840, de Guyane française. s’agit species in French Guiana is reinvestigated and cinq espèces, à savoir: C. saxatilis (Linnaeus, 1758), C. all material for that of C. type (except albopunctata Pellegrin, 1904, C. multispinosa Pellegrin, brasiliensis) has been examined. The variability 1903, C. ternetzi Norman, 1926, et C. johanna Heckel, within the various species has been and 1840. Leur distribution géographique est discutée, ainsi studied,

la variabilité de leurs is with data available from Surinam que caractères morphométriques, compared

de leur La variabilité de méristiques et pigmentation. ces (Ploeg, 1986, and in prep.). Methods used for caractères est comparée à celle de Crenicichla saxatilis du counts and measurements are as in Ploeg fleuve Suriname, du Surinam (Ploeg, 1986). (1986: 48).

Material of Brazilian tributaries to the

INTRODUCTION River, and of Surinam tributaries to

the Maroni ( = Marowijne) River, is included in

order allow better about the Recently only Le Bail et al. (1984) treated the to a understanding

distributionof the within this The cichlid genus Crenicichla in French Guiana, species area.

be in future although previously much scattered informa- latter material will treated a

tion this the Crenicichla of has been published on subject publication on species

(Vaillant, 1900; Pellegrin, 1903, 1904, and Surinam.

1905; Regan, 1905, 1913; Norman, 1926;

Puyo, 1949). ACKNOWLEDGEMENTS In solution of the the past (up to the French-

Brazilian border in French Rich material in Institute of Taxonomic dispute 1897) present the

the Brazilian of Amsterdam Guiana and present state Zoology (Zoologisch Museum) (ZMA), (Dr.

H. was used basis for the constituted Nijssen) as a present study. Amapa "Guyane frangaise" or Additional material was made available by the British "" (nowadays the name of the capital Museum (Natural History) (BMNH), London (Mrs. B. and of a river in French the Guiana). Formerly Brewster), the Institut Royal des Sciences Naturelles de of the named western part country was Mana, Belgique (ISNB), Brussels (Dr. J. P. Gosse and Mr. L.

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National d'Histoire Naturelle 4a. White dots and Walschaerts), the Museum present on operculum cheek; humeral

(MNHN), Paris (Dr. M. L. Bauchot and Mr. F. spot without notch Crenicichla albopunctata

b. White absent d'Aubenton), the Rijksmuseum van Natuurlijke Historie dots on operculum and cheek; humeral

with notch at dorsoanterior side (RMNH), Leiden (Drs. M. J. P. van Oijen), the Natur- spot

historiska Riksmuseet (NRM), Stockholm (Dr. S. O. Crenicichla saxatilis

Kullander), the Naturhistorisches Museum (NMW),

Vienna (Dr. B. Herzig and Mr. H. Ahnelt), and the Crenicichla saxatilis (Linnaeus, 1758) Office de la Recherche Scientifique et Technique Outre table Paris (Figs. 1, 2; I) Mer (ORSTOM), (Dr. P. Y. Le Bail and Mr. P.

I wish for their Planquette). to express my gratitude help. Sparus saxatilis Linnaeus, 1758: 278 (original diagnosis;

Thanks are also due to Mr. J. Zaagman for the drawings, type-locality: Surinam).

to Mr. L. A. van der Laan for the photographs, to the Crenicichla saxatilis; Vaillant, 1900: 125 (Carnot,

University of Amsterdam for the facilities, and to Dr. H. Carsevenne and Guyane frangaise); Pellegrin, 1904:

Nijssen and Dr. I. J. H. Isbriicker for their criticism and 373 (Cayenne); Pellegrin, 1905: 589 (Guyane fran-

encouragement. gaise); Steindachner, 1915: 572 (Maroni River); Puyo,

1949: 243-244 (Mana, Mahury, Oyapock, Mont-

siraery, Cayenne); Ploeg, 1986: 47-59 (Suriname, Crenicichla 1840 Heckel, Marowijne and Commewijne River systems).

Crenicichla saxatilis 1904: 374 var. albopunctataPellegrin, (in

Crenicichla Heckel, 1840: 416-417 (original diagnosis). part).

Type-species, by subsequent designation of Eigen-

Material. — Surinam and French Guiana:

mann & Bray, 1894: 620: C. macrophthalma. Marowijne River system: Marowijne River at Tabbetje Batrachops Heckel, 1840: 432-433 (original diagnosis). Hede: ISNB 19567, 8 specimens, 24-XI-1966; Nengre Boggiania Perugia, 1897: 148 (original diagnosis). Creek into Marowijne River near Albina: ISNB 19568, 2

specimens, 26-XI-1966; ZMA 119.374 and NRM Diagnosis. — Elongate; lower jaw somewhat unreg., Crique Vampire, creek tributary to MarowijneRiver, St.- prognathous; orbit in upper half of head; Laurent du Maroni, 4 specimens, 8-X-1979; ZMA preoperculum serrated. Two well-developed 119.733, Crique Cascade, creek tributary to Canal St.- lateral lines with scales than larger adjacent Jean (Maroni) near St.-Jean, 2 specimens, 21-1-1980;

fin ZMA 119.373 and NRM Balate, 5 scales, continuing on caudal with one or two unreg., Crique specimens, 9-X-1979. scales; forehead scales naked; on head, oper- Mana River system: ZMA 119.376, Crique St.-Anne, culum, suboperculum, anterior part of back, 1 specimen, 22-IX-1978. and belly cycloid; caudal fin with basal and Counamama River system: ZMA 119.377, Pri-Pri Yi- interradial fins without basal scales, other Yi, along the Route National 1, between Corossony and

2 VII-1983. scales. Dorsal fin acute posteriorly; caudal fin , specimens,

River ZMA lower rounded; anal fin origin just below first soft system: 119.380, Bayou, Sinnamary River, of village Sinnamary, 1 dorsal three anal upstream ray, posteriorly acute, always specimen, VI-1981; ZMA 119.381, Petit Saut, Sin- fin acute, V 1.5; fin spines; pelvic pectoral the level of 1 namary River, at Crique Coeur-Maroni, rounded. Teeth conical and inner recurved, specimen, 3-II-1983. teeth series often depressible, outer series fixed. River system: ZMA 119.378, Crique

Gill rakers ceratobranchial Papinabo, creek tributary to Kourou River, 1 specimen, externally on first 24-1-1983. denticulate. Mahury River system: ZMA 119.372 and NRM

creek unreg., Crique Blanche, tributary to Orapu River, KEY TO THE SPECIES OF FRENCH GUIANA 2 ZMA 119.375 and NRM specimens, 3-X-1979; unreg.,

Scales la. on sides ctenoid 2 Crique Gabrielle, creek tributary to Mahury River

b. Scales sides ZMA on cycloid Crenicichla johanna upstream of River, 8 specimens, 5-X-1978;

2a. More than 75 scales below lateral humeral creek lines; spot 119.379, Crique Magnan, tributary to Comte

absent 3 River, 1 specimen, 5-X-1978.

b. Less than 76 scales below humeral River ZMA Kaw lateral lines; spot Approuague system: 111.255,

present 4 Guisanbourg, near outlet of Approuague River, 1

3a. Caudal fin with dark spot; white dots absent on sides specimen, VIII-1971.

in males Crenicichla ternetzi River ZMA of nuptial Oyapock system: 107.845, , state

b. Caudal fin without dark white dots Savanna Creek Youminan spot; present on Amapa, Igarape Jumina, at sides in nuptial males Crenicichla multispinosa (04°03'N 51°39'W), 1 specimen, 20-IX-1977; ZMA

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Fig. 1. Crenicichla saxatilis (Linnaeus, 1758) from the Oyapock River, Gabaret Creek, ZMA 107.644, 167 mm SL, male.

and 3 teeth series less in the 107.644, Gabaret Creek near its confluence with Oyapock topotypes), (c) or River, 11/14-XI-1976; ZMA 107.697, Saut 1 specimen, lateral of the in and 2 part upper jaw adults, Maripa and downstream of Pied Saut, falls in Oyapock teeth series in the lateral of the lower part jaw River, 1 specimen, 5-XI-1976; ISNB 20.084, teeth (up to 5 and up to 4 series, respectively, River, 1 specimen, 30-XI-1969; ISNB 20.085, Alikene in adult Creek, right creek tributary to Camopi River, 5 topotypes).

specimens, 1 -XII-1969; ISNB 20.086, Adjoumba Creek,

between Sikini and Camopi, right creek tributary to

Oyapock River, 7 specimens, 6-XII-1969; ZMA 107.734,

Euleupousing Creek near its confluence with Oyapock

River, 1 specimen, 6-X-1967; ZMA 107.728, Mare Ipa

Trois Sauts in 3 near upper Oyapock River, specimens,

8-X-1967.

Notes. — For a detailed description of

Crenicichla saxatilis see Ploeg (1986), concerning

C. saxatilis from its restricted type-locality:

Surinam, Carolina Creek, Suriname River

system.

In Surinam and French Guiana C. saxatilis

occurs in the rivers of and between the

Surinameand Oyapock River systems. In these

rivers the species exhibits a considerable

variability in colour pattern, not only between

the rivers, but also within one river. Puyo

2. Distribution of Crenicichla saxatilis (1949: 243-244) pointed out that variation in Fig. (Linnaeus, dots 1758). Single more than one sample. colour is influenced by different habitats: "la may represent

coloration de l'eau (boueuse, claire, verdatre,

etc.)". Morphometric and meristic variability Variability within one river could not be

can be explained in particular by the influence observed (as found in the Suriname River of different water temperatures and currents (Ploeg, 1986: 56)), because in French Guiana

(Ploeg, 1986: 57). Specimens from French C. saxatilis is not known from both lower and

Guiana differ from those of the Carolina Creek of the upper parts same river (except for the

in: from however (topotypes) (a) having occasionally a less Oyapock River, which no series

humeral in than conspicuous spot (always present larger one to three specimens were

10 rakers in topotypes), (b) or more gill (8-9 available, showing no noteworthy intraspecific

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Table I

Morphometric and meristic data of C. saxatilis, C. albopunctata, C. johanna, C. ternetzi, and C. multispinosa. Data marked with

* are based on the lectotype and two paralectotypes only (MNHN A 9497).

Species C. saxatilis C. albopunctata C. johanna C. ternetzi C. multispinosa

Number of specimens 25 25 25 25 11

Total length (mm) 46-149 98-169 66-283 57-245 124-266

Standard length (mm) 37-121 78-139 53-235 47-203 104-225

Head length (% SL) 31-36 32-35 29-34 30-38 32-34

Head depth (% SL) 15-24 18-23 17-21 14-18 14-16

Head width (% SL) 12-16 12-14 14-18 11-16 13-15

Depth caudal peduncle

(% SL) 11-14 10-12 10-12 10-12 10-12 Length caudal peduncle

(% SL) 10-14 8-11 10-13 11-14 11-12*

Snout length (% SL) 8-10 9-12 8-11 8-13 11-14

Upper jaw length (% SL) 9-14 10-14 10-12 8-13 9*

Lower jaw length (% SL) 14-18 15-19 14-17 10-16 13-14*

Length last dorsal spine

(% SL) 9-13 10-12 9-11 7-10 8-9*

Orbital diameter (% HL) 19-29 20-25 16-22 15-26 15-20

Width of interorbital (% HL) 21-35 15-24 27-38 20-30 20-29

Scales first lateral line 21-25 22-25 24-28 26-28 26-29

Scales second lateral line 8-13 8-11 13-15 11-14 12-15

Scales below lateral lines 52-68 57-65 110-125 90-100 98-110

Number of dorsal spines 15-20 17-21 20-22 23-25 23-25

Number of soft dorsal 13-15 11-15 16-19 13-15 13-15 rays

Number of soft anal 8-11 9-11 11-13 9-11 9-10 rays

Material included ZMA 111.255; ZMA 107.839, ZMA 107.658; ISNB 15.885, RMNH 18.961;

ISNB 20.086 107.840, ISNB 15.868, 15.887, 20.089, MNHN A 9497,

106.509 20.069, 20.070; 20.091, 20.114 9542

MNHN 1899.113,

1981.539

In the Maroni River C. Material.— Lectotype, by designation variability). upper sax- present MNHN 149 98-59, mm SL male, Guyane frangaise. atilis is not found; from the Mana and Sin-

material is available namary River systems no Surinam and French Guiana:

Maroni River ZMA 107.839, Lawa River, from upper regions. system: Maka 10 km south of Stoelmans Island, 1 River show Creek, Oyapock specimens a slightly dif- specimen, 21-IV-1966; ZMA 107.840, Lawa River, in adult ferent pattern of dots on the sides Maka Creek, 10 km south of Stoelmans Island, 1 males: fewer and somewhat The larger. specimen, 21-IV-1966; ZMA 106.507, right bank has the notch ventrad than humeral spot more tributary to Marowijne River, east to Mooisantitabbetje,

in specimens from the Suriname River. 36 km north of Stoelmans Island, 12 specimens, 19-IV- 1967; ZMA 106.508, left bank tributary of Marowijne

River, west to Manbarie fall, 6 km north of Stoelmans

Island, 19 specimens, 20-IV-1967; ZMA 106.509, Lawa Crenicichla albopunctata Pellegrin, 1904 River, Maka Creek, 10 km south of Stoelmans Island, 53 3-5; table (Figs. I) specimens, 21 -IV-1966; ZMA 106.510, Soea kisi Creek,

left bank tributary to River, 12 km

Crenicichla saxatilis var. albopunctata Pellegrin, 1904: 374 southwest ofStoelmans Island, 49 specimens, 22-IV-1967;

(original description). Syn type-localities: Guyane ZMA 106.511, Kamaloea, right bank tributary to

anglaise, Surinam, Guyane frangaise. Marowijne River, 9 km southeast of mouth of Gran

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Fig. 3. Crenicichla albopunclata Pellegrin, 1904 from French Guiana, lectotype MNHN 98-59, 149 mm SL, male.

ISNB Lawa River ZMA Creek, 35 specimens, 24-IV-1967; 20816, Mahury system: 119.386, Crique

River, Maripasoela, Daouda Creek near airstrip, 3 Gabrielle, creek tributary to Mahury River upstream of specimens, 12-XI-1969; ISNB 20817, River, Saut Roura, 1 specimen, 5-X-1978.

Sonelle, 3 specimens, 16-XI-1969; ISNB 20818, Bois

Blanc bank creek to Inini 5 Creek, right tributary River, Notes. — A detailed description of Crenicichla specimens, 16-XI-1969; ISNB 20819, Quaqui River, albopunctata will be included in a future article right bank tributary to Tampoc River, Saut Bali, 3 on the Crenicichla species of Surinam. specimens, 18-XI-1969; ISNB 20820, Quaqui River,

of Saut 5 ISNB The syntype-localities "Guyane anglaise, upstream Bali, specimens, 18-XI-1969;

20821, rapids in Paloemeu River, upstream of confluence Surinam, Guyane frangaise" cover an area far

1 with Walimeroe Creek, specimen, 3-XI-1966; ISNB larger than the distribution of the species

20822, right bank tributary to Marowijne River near Tab- recorded above. Therefore, the type-locality betje Hede, 6 specimens, 24-XI-1966; ISNB 20823, small needs to be restricted. Because of the brooklet in Oelemari 3 13-XI- upper River, specimens, to determinefrom where the lecto- ISNB small brooklet in Oelemari impossibility 1966; 20824, upper I herewith restrict the River, 5 specimens, 14-XI-1966; ISNB 20825, Paloemeu type originated, type-

River 2 l-XI-1966; ISNB at Papadronsoela, specimens, locality arbitrarily to the Maroni River system, 20826, right bank tributary to Paloemeu River between Lawa River, Maka Creek, 10 km S. of Trombeka-North and Trombeka-South, 2 specimens, 8- Stoelmans Island (ZMA 106.509, 119.839, XI-1966; ISNB 20827, right bank tributary to

119.840: a series of , downstream of confluence with large well-preserved

Paloemeu River, near Paloemeu airstrip, 3 specimens, 9- specimens).

XI-1966; ISNB 20828, right bank tributary to Oelemari The five of syntypes C. saxatilis var. albopunc-

River near Oelemari airstrip, 15 specimens, 10-XI-1966; tata represent at least two species. Only one of ISNB 18966, Oelemari near airstrip, 5 specimens, no MNHN the specimens, 98-59, represents the ISNB small brooklet in Oelemari date; 20829, upper

described here as C. and River, 3 specimens, 14-XI-1966; ISNB 20830, rapids in species albopunctata, therefore is the Para- Oelemari River near Oelemari airstrip, 6 specimens, 15- designated lectotype.

XI-1966; ISNB 20831, to Lawa River near tributary lectotype MNHN A-9502 can be identified as Anapaikekondre, 4 specimens, 28-XI-1966; ISNB 18959, C. saxatilis by the shape of its humeral spot (a Paloemeu near Walimeroe Creek, 1 specimen, no date; notch at the dorsoanterior side). Paralectotype ISNB 19567, Marowijne River near Tabbetje Hede, 5 MNHN 4083 is also ZMA 119.385 and NRM Cri- C. saxatilis, specimens, no date; unreg., probably

creek Canal 2 because of its que Cascade, tributary to St.-Jean, origin "Surinam" (most likely

21-1-1980. specimens, originating from the Suriname River; in this Mana River ZMA 119.384 and NRM system: unreg., river no other C. saxatilis-like species occur). Sub-Acarouany, creek tributary to lower Mana River, Unfortunately, its colour pattern is faded. opposite to village Acarouany, 3 specimens, 3-III-1980.

Paralectotypes MNHN A-9498 are two Approuague River system: ZMA 119.383 and NRM

basin, 4 IV-1983. from Guiana. One of the unreg., Approuague specimens, specimens (British)

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Fig. 4. Crenicichla albopunctata Pellegrin, 1904 from the type-locality, ZMA 107.840, 124 mm SL, female.

shows humeral in the Crenicichla 1840 two a spot as C. saxatilis, johanna Heckel, other is faded. (Figs. 5, 6; table I)

The syntype of C. vaillanti (MNHN 4050) Crenicichla johanna Heckel, 1840: 425-426 (original descrip- originates from French Guiana: "Mana". It is tion), type-locality: Guapore River. bleached and a specimen. Morphometric C. johanna; Regan, 1913: 504 (name only); Le Bail et al., meristic data with both C. saxatilis and C. agree 1984: 43 (Guyane frangaise).

and meristic data Crenicichla brasiliensis vittata albopunctata. Morphometric var. (non Heckel, 1840);

of C. but Pellegrin, 1904: 383 (Carsevenne (Guyane frangaise)). are reminiscent mostly albopunctata, do

Crenicichla johanna var. carsevennensis Pellegrin, 1905: 168- with C. saxatilis. the not exclude synonymy For 169 (original description of this variety). Entre les French moment I consider the Guiana syntype rivieres Carsevenne et Cachipour (conteste franco- be with C. The to conspecific albopunctata. sec- bresilien). ond syntype, MNHN 2993, from the Essequibo

Material. — French Guiana and Brazil: needs with Crenicichla River, comparison River Suivelaleu Oyapock system: ZMA 107.658,

from the River. Creek its confluence with River specimens Essequibo near upper Oyapock

(02°40'N 52°33'W), 1 specimen, 13-X-1976; ISNB

15.868, Igarape Cumuri, left bank of tributary to

Oyapock River, upstream offirst rapids of Grande Roche,

2 specimens, 5-XII-1962; ISNB 15.869, Igarape

Taparabu, right creek tributary to Oyapock River,

downstream of St.-Georges, 3 specimens, 8-XII-1962;

ISNB 20.069, Crique Alikene, left creek tributary to

Camopi River, 3 specimens, l-XII-1969; ISNB 20.070,

Crique Sikini, left creek tributary to Oyapock River, 1

specimen, 6-XII-1969; MNHN 01-461, Camopi River, 1

specimen; MNHN 1981-539, Crique Gabaret, 1

specimen.

Calgoene River system: MNHN 1899.113, Calgoene

C. River, 1 specimen (paratype of johanna var. carseven-

nensis).

Description. — Body behind head laterally

compressed; head somewhat flattened; predor-

sal contour straight, preventral contour convex;

angle between predorsal and preventral contour

lateral about 35°; snout blunt, both in aspect Fig. 5. Distribution of Crenicichla albopunctata Pellegrin, and from mouth wider than 1904 and Crenicichla 1840. dots above; lips thick; johanna Heckel, Single may

more one orbit in half of orbital represent than sample. interorbital; upper head,

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1/3 of head orbit — No sexual diameter about depth at level; Colour pattern. dimorphism

in colour of nostrils nearer to postlabial skinfold than to colour pattern found. Ground body orbit, distance between nostrils and orbit about brown, countershaded. Dorsal, pectoral,

orbital maxilla From as long as diameter; reaching to pelvic, and anal fins light. tip of snout to anterior of orbit. end of caudal fin brown about 1/4 margin a dark band,

below Scales cycloid; some scales on anterior part of of body depth, covers the body, running

fused in vertical number of lower lateral body direction; upper lateral line and through scales below lateral lines (in 13 specimens): 110 line. In the caudal fin this band is rather faint.

111 113 115 117 brown (2), (1), 112 (3), (1), (1), (1), Just below dorsal spines a second, lighter

in 118 (2), 119 (1), 125 (1); number of scales in lateral band is present. More conspicuous

lines: and lateral 24/13 (1), 24/15 (1), 25/15 (2), this area more caudad, are about 12 dark

26/13 (1), 26/14 (3), 27/13 (2), 27/14 (1), 28/13 vertical bars between dorsal fin and upper

(2); 5-6 horizontal scale series between the lateral line. Small brown dots, forming dark

line caudal of head. The lateral lines; lateral continuing on lines, are present on dorsum fin with interradial scales 1/3- caudal fin shows lateral band in the one or two scales; a vague

2/3 of caudal fin length. middle, and two light bands, forming a V with

anteriad the In lateral part of mouth two teeth series in the open end directed and point both jaws; teeth of outer series largest and directed backwards. Anal fin with distinct dark fixed, inner teeth series depressible. Externally margin. Neither humeral spot, nor spot on

caudal fin on first ceratobranchial 9-12 rakers. present.

Dorsal fin origin just dorsal to pectoral fin; spines increasing in length to about tenth; soft Notes. — Unfortunately, the holotype of C.

of dorsal fin fifth from last the not available due to part acute, ray johanna (NMW 16.431) was longest, reaching to just behind caudal fin base; the current policy of the Vienna Museum. Mr.

D XX. XX.19 XXI.17 XXI.18 Ahnelt with 18 (2), (1), (3), kindly provided me a good

(2), XXII. 16 (2), XXII. 17 (3). Caudal fin photograph of this specimen. rounded; about 23% of SL. Anal fin acute, The label of the Crenicichla specimen with third from the MNHN 1899-113 indicates that it ray posterior longest; spines register no.

A III.10 III.11 III.13 is the of C. thick; (1), (3), III.12 (7), holotype johanna var. carsevennensis,

Ventral fin first soft the which is This recorded (2). acute, ray longest, not true. specimen was and reaching only to one third of the distance to by Pellegrin (1904: 383) as C. brasiliensis var. anal fin less than half vittata. the of origin; spine as long as According to original description first soft Pectoral fin about 1.5 C. var. in which one ray. rounded; johanna carsevennensis, only times ventral fin. the of this as long as specimen was mentioned, holotype

carsevennensis Fig. 6. Crenicichla johanna Heckel, 1840, paratype of C. j. var. Pellegrin, 1904 from the Calçoene River,

MNHN 1899.113, 175 mm SL, sex unknown.

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with brooklet left of subspecies is a specimen register no. Igarape Cumuri, Oyapock River, of first rapids of Grande Roche, 3 specimens, 5- MNHN 1899-112. The type-locality of the upstream XII-1962; ISNB 19.678, Camopi River, Pauwe Jean-Jean variety is "Entre les rivieres Carsevenne et (downstream of Saut Mauvais), 1 specimen, 30-XI-1969; franco-bresilien)". The Cachipour (conteste ISNB 20.089, Camopi River, Pauwe Jean-Jean named Carsevenne River is nowadays Cal- (downstream of Saut Mauvais), 9 specimens, 30-XI-1969;

is written ISNB 20.090, Crique Alikene, left creek tributary to goene River; Cachipour presently Camopi River, 6 specimens, l-XII-1969; ISNB 20.091, Cacipore. Saut Alikote (upstream of Camopi village), 122 The distribution of C. johanna in specimens, 4-XII-1969; ISNB 10.114, upstream of St. is The in the interesting. species occurs of Kumalawa, upstream Camopi village, 1 specimen, 27- and in the Esse- Oyapock, Amazon, Branco, VII-1982; MNHN 01-462, Camopi River, 1 specimen, quibo River systems. Its distribution surrounds no date. the centre of the Guianas from the Corantijne

the from where — Based and 15 basin to Approuague system, Description. on lectotype the species is not known. According to Lowe- other specimens (ISNB 20.091). Very elongate,

McConnell (1964: 105) the Essequibo River slightly laterally compressed; head more or less

River and Branco systems are in contact depressed; predorsal contour either slightly

and Tacutu (through the Rupununi River) convex or straight; preventral contour straight;

the in flooded between and during rainy season wet years by angle preventral predorsal contour areas. A similar situation exists between bran- about 35°; snout pointed from the side, blunt ches of the Oyapock River and branches of the from above; mouth as wide as interorbital

River and the maxilla anterior of Jari systems can explain presence diameter; not reaching edge

between orbit and of C. johanna in these systems. orbit; nostrils halfway

In the material studied the number of scales postlabial fold of skin; orbital diameter about below with the the lateral lines is compared half head depth at orbit level; orbit in upper

mentioned 98 half caudal about original description. Heckel of head; peduncle as deep as

whereas I found This be scales, 110-125. can long. due to the of but also with ctenoid scales behind way counting, to Occasionally just intraspecific variation (Heckel's specimens pectoral fin; scales on body becoming larger

with 93 scales came from the Rio Guapore in western Brazil). posteriorly; lectotype below

labeled 94 A specimen as "paratype" (there are lateral lines, in 13 other specimens 90, (4),

in the collec- 95 96 with no paratypes) of C. johanna present (3), (2), 97, 99, 100; lectotype 27/12

is scales in lateral in 15 tion of Vienna (NMW 77.770) a C. vittata. lines, other specimens:

26/12, 26/13 (4), 26/14 (2), 27/12, 27/13 (5),

28/11 lateral line scales 3 4 times Crenicichla ternetzi Norman, 1926 (2); or larger

table than adjacent scales; 3-4 scales in horizontal (Figs. 7, 8; I) series between lateral lines; lateral line continu-

Crenicichla ternetzi Norman, 1926: 97 (original description). ing onto caudal fin with 2 scales; 3-4 basal scale

French Guiana: Oyapock River, Sant Cafesoca; Le series on caudal fin; interradial scales reaching

Bail et al., 1984: 43 (Guyane frangaise). to half caudal fin length.

Material. — Dorsal fin behind of Lectotype, by present designation, BMNH origin just upper edge

163 French Guiana: 1926.3.2:860, mm SL, male, operculum; spines increasing in length to about Oyapock River, Sant [=Saut] Cafesoca. Paralectotype: seventh, remaining spines equal in length; soft

BMNH 1926.3.2:861, 161 mm SL male, same data as sixth from last the part posteriorly acute, ray lectotype. longest, reaching to half caudal fin length; French Guiana and Brazil: lectotype: D XXIII. 13, in 15 other specimens: Oyapock River system: ISNB 15886, creek tributary to XXIII. 13, XXIII. 15, XXIV. 13, XXIV. 14 brooklet left of Igarape Ciparini, tributary to Oyapock (12). Caudal fin rounded, 17-22% of SL. Anal River upstream of Clevelandia, State of Amapa, 2

ISNB fin fourth from last the specimens, 4-XII-1962; 15887, French Guiana: posteriorly acute, ray

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Fig. 7. Crenicichla ternetzi Norman, 1926 from the Oyapock River at Saut Cafesoca, lectotype BMNH 1926.3.2:860, 163

mm SL, male.

longest, in males reaching to halfway the caudal with some vague light dots in posterior three

behind base of soft ventral fin fin fin length, in females to just rays; yellowish; pectoral

in other less than about 50 caudal fin; lectotype: A III.9, 15 grey; juveniles (SL mm)

specimens: III.9 (2), III. 10 (13). Ventral fin yellow with dark brown horizontal band from

acute; spine halfas long as first soft ray; second snout to end of caudal fin rays; a second

fin. horizontal dark band base of dorsal soft ray longest, little shorter than caudal on fin, con-

Pectoral fin little than ventral the head between the short longer fin, tinuing on eyes; a

rounded. horizontal dark stripe, from dorsal part of orbit

of Externally on first ceratobranchial 11 to 12 to edge operculum.

gill rakers; dorsal four or five rakers

Notes. — Crenicichla ternetzi is reminiscent of knob- boomerang-shaped, remaining ones C. C. multispinosa. However, multispinosa can shaped. easily be distinguished from C. ternetzi by its col-

—The is dull white dots in males of C. Colour pattern. lectotype grey our pattern: (a) many

band. The with only a dark lateral description

is of the colour pattern based on fresh material

(ISNB 20.091).

Ground colour brown; back dark brown, a

still darker lateral ventral of band, part body

yellow-brown to yellow; lateral band running

from below distal tip of lower jaw, nostril, over orbit and below first lateral gill cover, line, on and above second lateral line, to end of caudal

fin, where it gradually becomes faint; vague,

dark suborbital in several stripe males, vague

darker suborbital blotches in females; no dots

with dark on body; dorsal fin in males grey

with faint margin, grey part posteriorly light

females with dorsal fin without dots; grey dots,

with narrow dark margin, thin white edge

fin of basally of this margin; caudal males grey

with white females without these vague dots, Fig. 8. Distribution of Crenicichla ternetzi Norman, 1926,

dots; caudal fin with two white bands forming and Crenicichla multispinosa Pellegrin, 1903. Single dots

V with the end directed males than a open anteriad; may represent more one sample.

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in C. mouth of Paloemeu River, 1 specimen, 16/17-1-1964; multispinosa (absent ternetzi), (b) presence ISNB 18961, Lawa River, Anapaikekondre, 26 caudal in of a fin spot in C. multispinosa (absent specimens, 18-XI-1966; ZMA 106.511, Kamaloea or C. ternetzi), (c) white bands in the caudal fin of Saloea Creek, 9 km south of Gran Creek, 35 specimens, C. ternetzi females in C. (absent multispinosa). 24-IV-1967; ZMA 100.735, Marowijne River, 2

Another distinguishing character is the number specimens, 1952.

of scales below the lateral lines (on the average Notes. — A detailed description of Crenicichla C. this eight more in multispinosa). However, in article multispinosa will be included a future character is of less value because of the known the Crenicichla of Surinam. on species in Crenicichla intraspecific variability some spp. Dots of on posterior part body present only in (cf. Ploeg, 1986). C. multispinosa and C. ternetzi nuptial males, collected during the short rainy do not occur sympatrically. season: December-February. The resemblance of the colour pattern of C. multispinosa seems to be restricted to the adult C. ternetzi and juvenile C. johanna is Suriname and Maroni (Marowijne) River remarkable. Both species occur sympatrically. systems. In these systems the species is collected

in the upper regions only.

Crenicichla multispinosa Pellegrin, 1903 The original type-locality (Guyane frangaise)

table is is in (Figs. 8, 9; I) poorly defined since the species present

small of the Crenicichla multispinosa Pellegrin, 1903: 124 (original de- a part country only. Therefore, I scription), Guyane franchise; Pellegrin, 1904: 380-381 herewith restrict the type-locality to the Maroni (Guyane frangaise, Surinam); Le Bail et al., 1984: 44 Creek material River, South of Gran (based on (Guyane frangaise). listed above). Crenicichla lugubris (not Heckel, 1840); Nijssen, 1967: 246

Lake Richter C. multispinosa and C. ternetzi are more similar (Surinam: Brokopondo); & Nijssen, 1980: than other 125 (Surinam, Lake Brokopondo). to each other both are to Crenicichla

The species. very elongate body shape, more or Material. — Lectotype, by present designation,

less maxilla not MNHN 9542, 137 mm SL, French Guiana. Paralec- depressed head, reaching totypes: MNHN 9542, 2 specimens, French Guiana; anterior edge of orbit, small scales, pointed

MNHN 9496, 3 specimens, French Guiana. dorsal snout, many spines, and the nostrils

between orbit and skin- Surinam and French Guiana: placed just postlabial

Marowijne River system: ISNB 18958, Paloemeu fold, distinguish these two species from the

River, Papadronsoela, 1 specimen, l-XI-1966; ISNB remaining species of the genus. 18959, Paloemeu River, Waloemeroe Creek, 38 specimens, 3-II-1966; ISNB 18960, Oelemari River near

2 RMNH SPECIES ERRONEOUSLY RECORDED FROM FRENCH airstrip, specimens, 15-XI-1966; unreg.,

Tapanahony River, 2 km downstream of airstrip, 14 GUIANA

RMNH specimens, 17-XI-1965; unreg., Tapanahony

Crenicichla brasiliensis — River, 2.5 km downstream of airstrip, 5 specimens, 30- (Bloch, 1792). 1904: 381 used the XI-1965; RMNH 28975, TapanahonyRiver, 2 km below Pellegrin, name C.

9. from French MNHN 137 male. Fig. Crenicichla multispinosa Pellegrin, 1903 Guiana, lectotype 9542, mm SL,

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tria C. b. C. b. LINNAEUS, C., 1758. Systema naturae per regna brasiliensis for var. johanna and var. secundum naturae, classes, ordines, genera, species, lugubris, which he regarded varieties of the same locis. cum characteribus, differentiis, synonymis, from the MNHN collection species. Specimens Tomus I. Editio decima, reformata: 1-824 (L. Salvii, labelled C. brasiliensis re-identified C. are as Holmiae). johanna (MNHN 01-461) and C. ternetzi LOWE-MCCONNELL, R. H.,. 1964. The fishes of the

Rupununi savannadistrict of British Guiana, South (MNHN 01-462). America. Part 1. Ecological groupings of fish species

Crenicichla lugubris Heckel, 1840. — The record and effects of the seasonal cycle on the fish. J. Linn. Soc., (Zool.) 45 (304): 103-144. of Le Bail et al., 1984 is based on a misiden- NIJSSEN, H., 1967. Final remarks and tentative list of fish misidentified tification (Richter & Nijssen, 1980 species. Progr. Rep. biol. Brokopondo Research C. multispinosa as C. lugubris). Project, Surinam, 4: 240-246.

NORMAN, J. R., 1926. Descriptions ofnine new freshwater

fishes from French Guiana and Brazil. Ann. Mag. DISCUSSION nat. Hist., (9) 18: 91-97.

PELLEGRIN, J., 1903. Description de Cichlides nouveaux

rivers in French Guiana de la collection du Museum. Bull. Mus. Hist. nat. Although many are

Paris, 9: 120-125. scarcely sampled, there are striking differences

1904. Contribution a l'etude , anatomique, biologi- C. between the Maroni River system (with des Poissons de la famille des que et taxinomique and C. multispinosa), and the albopunctata Cichlides. Mem. Soc. zool. , 16: 41-402, pis. River and Oyapock system (with C. ternetzi C. IV-VII.

The river share one 1905. Sur deux du Crenicichla de la johanna). two systems , poissons genre

collection du Museum de Paris. Bull. Soc. zool. species only: C. saxatilis. Most rivers between France, 30: 167-169. these two border rivers (Mana, Iracoubo, Sin- PERUGIA, A., 1897. Di alcuni pesci raccolti nell' alto and show namary, Kourou, Cayenne, Orapu) Paraguay dal Cav. Guido Boggiani. Annali Mus.

an influence of the Maroni River the occur- by civ. Stor. nat. Genova, (2) 18 (2): 147-150.

The River rence of C. albopunctata. Oyapock PLOEG, A., 1986. Occurrence and variability of

has its Crenicichla Crenicichla saxatilis (Linnaeus, 1758) in Surinam, system, however, own species and restriction of its type-locality (Pisces, Per- together with some influence of the Amazon ciformes, Cichlidae). Bijdr. Dierk., 56 (1): 47-59. basin (C. johanna). PUYO, J., 1949. Faune de l'empire 12. Poissons

de la Guyane frangaise: 1-280 (Office de la Recher-

che scientifique et technique d'Outre-mer, Paris).

REGAN, C. T., 1905. A revision of the fishes of the South

REFERENCES American cichlid and genera Crenicara, Batrachops,

Crenicichla. Proc. zool. Soc. London, 1905:

P. Y. P. 1984. Cle de 152-168. BAIL, LE, PLANQUETTE & J. GERY,

determinationdes continentaux coders de 1913. A of the cichlid fishes of the poissons et , synopsis genus

Guyane, IV. Especes dulcaquicoles non Crenicichla. Ann. Mag. nat. Hist., (8) 11: 498-504.

Siluriformes. Bull. Liaison Inst. Rech. & H. 1980. Notes the natn. agron. RICHTER, C.J.J. NIJSSEN, on fishery Lab. Hydrobiol., Kourou, 9: 1-97. and fish fauna of the Brokopondo Reservoir

EIGENMANN, C. H. & W. L. BRAY, 1894. A revision ofthe (Surinam). Fish Mgmt., 11 (3): 119-130. American Cichlidae. Ann. New York Acad. Sci., 7: STEINDACHNER, F., 1915. Ichthyologische Beitrage

607-624. (XVIII). Sber. kaiserl. Akad. Wiss. Wien, math.-

HECKEL, J., 1840. Johan Natterer's neue Flussfische naturw. Kl., (1) 124 (8-10): 567-591.

Brasilien's, nach den Beobachtungen und Mit- VAILLANT, L., 1900. Contribution a l'etude de la faune

teilungen des Entdeckers beschrieben (erste ichthyologique de la Guyane frangaise et du conteste

Annln. wien. Mus. franco-bresilien. Nouv. Hist, Abtheilung, die Labroiden). Archs. Mus. nat., (4)

Naturg., 2: 1-470. 2: 123-135, pi. VII.

Received: 22 January 1986

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