UNIVERSITY OF AMSTERDAM. A. Ploeg. Abstract. Crenicichla. Ploeg, 1986a) four species are recorded from

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1 Beaufortia INSTITUTE OF TAXONOMIC ZOOLOGY (ZOOLOGICAL MUSEUM) UNIVERSITY OF AMSTERDAM Vol. 37, no. 5 December, 87 Review of the cichlid genus Crenicichla Heckel, 40 from Surinam, with descriptions of three new species (Pisces, Perciformes, Cichlidae) A. Ploeg Institute of Taxonomic Zoology (Zoologisch Museum), University of Amsterdam, P. 0. Box 5, 00 HC Amsterdam, The Netherlands Abstract The Surinam representatives of the cichlid genus Crenicichla Heckel, 40 are reviewed. Seven species, among which three new to science, have been found, viz.: C. albopunctata Pellegrin, 04, C. coppenamensis sp. n., C. nickeriensis sp. n., C. saxatilis (L., 58), C. sipaliwini sp. n., C. lugubris Heckel, 40, and C. multispinosa Pellegrin, 03. The first five species are compared with each other, with C. alta Eigenmann,, from Guyana, and with C. labrina (Spix & Agassiz, ) from the Tocantins River. Crenicichla saxatilis was recently redescribed and figured (Ploeg, 86a); the other six species are described and figured in the present paper. INTRODUCTION Boeseman, 52; Wheeler, 58; Nijssen, 66, 67a, 67b; Richter & Nijssen, 80; While identifying the many Crenicichla Ploeg, 86a) four species are recorded from specimens assembled by Dr. M. Boeseman, Dr. G. F. Mees, and Dr. H. Nijssen during the Brokopondo Research Project (6367), it Surinam, viz.: C. alta, C. saxatilis (including C. saxatilis albopunctata), C. lugubris, and C. multispinosa. appeared to be difficult to interpret the variability in C. saxatilis sensu lato. The intraspecific variability of C. saxatilis from the METHODS Surinam River system appeared to be greater than the interspecific variability of several All measurements were taken as shown in closely related Crenicichla species (Ploeg, 86a). figure 1, and as in Ploeg 86a, b and c. From this study was concluded that several Abbreviations of characters are explained in 'saxatilislike' species exist, which is the subject table VIII. Vertebrae were counted from of this paper. radiographs. Differences between the five saxatilislike species are visualized with scatter In the literature (Gronovius, 54; Linnaeus, 58; Bloch, 2; Pellegrin, 03, diagrams. To avoid ontogenetic influences all 04; Regan, 05, ; Steindachner, ; measurements are expressed as percentages of 73

2 Thanks are due to the abovementioned curators, to Mr. J. Zaagman (ZMA) for the drawings, to Mr. L. A. van der Laan (ZMA) for the photographs, to the University of Amsterdam for the facilities, and to Drs. H. Nijssen and I. J. H. Isbriicker for their criticism, their encouragement, and for editing the manuscript. The Netherlands Organisation for the Advancement of Pure Research (Z.W.O.) supplied me with grants to visit the Natural History Museums of London, Paris and Vienna. Crenicichla Heckel, 40 Fig. 1. Way of taking measurements. Abbreviations are listed in table VIII. Crenicichla Heckel, 40: 44 (original diagnosis; type species, by subsequent designation of Eigenmann & the standard length (SL). Comparisons with two closely related viz. species, C. alta (material listed in Ploeg, in and C. press) labrina (material Bray, 4: 6: C. macrophthalma Heckel, 40). Batrachops Heckel, 40: 432 (original diagnosis; type species, by subsequent designation of Eigenmann & Bray, 4: 6: B. reticulata Heckel, 40). listed in Ploeg, 86b) are included. Local names of the species are from Heyde (86) and from H. Nijssen (pers. comm.). Lists of material are arranged as follows: register Local names in Surinam: Datra Fisi (all species), Adoenko (C. albopunctata, C. coppenamensis, C. saxatilis, and C. sipaliwini), number, numberof specimens, locality, date of collecting, and collector. Djonkodjo and Donfo (C. nickeriensis). Shared characters in the Surinam species ACKNOWLEDGEMENTS Body elongate. Fleshy lips. Lower jaw prognathous. Orbit in upper half of head. Two Rich material present in the Zoologisch welldeveloped lateral lines, continuing on Mueum Amsterdam (ZMA), (Dr. H. Nijssen) was used as a basis for the present study. Additional material was made availably by the Institut Royal des Sciences Naturelles de Belgique (ISNB), Brussels P. the (Dr. J. Gosse), Museum National d'histoire Naturelle (MNHN), Paris M. L. the (Dr. Bauchot), Naturhistorisches Museum Wien (NMW), caudal fin with one to three scales; dorsal part of head naked; scales on cheek, operculum, suboperculum, anterior part of back, and belly cycloid; scales on sides ctenoid; caudal fin with basal and interradial scales; dorsal pectoral, and anal fins without basal scales, although base sometimes covered with skin. Dorsal fin acute posteriorly. Anal fin below first origin just Vienna (Dr. B. Herzig), the Naturhistoriska soft dorsay ray, fin posteriorly acute, three anal Riksmuseet (NRM), Stockholm (Dr. S. O. Kullander), the Rijksmuseum van Natuurlijke Historie (RMNH), Leiden (Drs. M. P. J. van and the Oijen), National Museum of Natural History (USNM), Washington D.C. R. (Dr. P. Vari). Ventral fin spines. acute; V 1.5; fin pectoral rounded. Caudal fin with primary rays. Teeth conical and recurved, outer teeth series fixed, often than inner teeth considerably larger series. Gill rakers on first gill arch denticulated externally. 74

3 Crenicichla Key to the Surinam species 6a. Humeral spot round with a white ring; la. Below lateral lines 00 scales 2 lb. Below lateral lines 52 scales 3 2a. scales below lateral lines, dark spot just posterior to pectoral fin base, nostrils nearer to postlabial skinfold than to orbit Crenicichla lugubris Heckel, 40 2b. 05 scales below lateral lines, no dark spot posterior to pectoral fin; nostrils almost halfway between skinfold postlabial and orbit (Crenicichla multispinosa Pellegrin, 03 3a. Series of dark blotches lateral lines in along males; humeral spot much longer than deep in females.. nickeriensis sp. n. males with yellowish dots (Crenicichla coppenamensis sp. n. 6b. Humeral spot with a dorsoanterior notch; males with silverywhite dots Crenicichla saxatilis (Linnaeus, 58) Crenicichla albopunctata Pellegrin, 04 (Figs. 2 and 3; Pl. 1 Tabs. I and upper; VIII) Crenicichla saxatilis var. albopunctata Pellegrin 04: 374 (original description; in part:, French Guiana). Crenicichla albopunctata; Ploeg, 86c: 4 [restriction of type locality (Surinam, Marowijne River system, Lawa River, Maka Creek, km S. of Stoelmanseiland); designation of the lectotype]. 3b. No series of dark blotches along lateral lines in males; humeral spot in females not or MATERIAL EXAMINED only slightly longer than deep 4 4a. Silvery dots present on operculum and See Ploeg, 86c: 4 (253 specimens, all from cheek in both sexes Marowijne river system). ( Crenicichla albopunctata Pellegrin, 04 4b. Silvery dots absent 5 5a. Maxilla not reaching orbit, or anterior DESCRIPTION margin of orbit; ground colour of body grey Crenicichla sipaliwini sp. n. 5b. Maxilla reaching well below orbit; ground Based on a 7 mm SL male topotype (morphometric and meristic data are presented in colour of body tan 6 tables I and VIII). Table I. Morphometric and meristic data for twelve specimens of Crenicichla albopunctata Pellegrin, 04. Abbreviations are listed in table VIII. CRENICICHLA ALBOPUNCTATA PELLE6RIN. 04 SI CFL HL HO HW DCP LCP SN UJL LJL LLDS LLAS VFL PFL PN PN2 ORB Wl LI L2 L1/L2 L2C LL SCDL DSP DSR 6R ASR P ASN MILLIMETERS PERCENTAGES MEAN , SO ,4 0, ,4 3.2, , , ,4 36, , , /ZMA 0, Surinam. Marowijne River system, LewaRiver, km South ofstoelmanseiland, IV66, H. Nijssen , ,.0..2,0..1, MEAN SO ,

4 the smallest specimens; lower jaw with maximum four teeth series, in smaller specimens fewer, only two in the smallest specimens; outer teeth series approximately the same size as inner teeth; teeth of upper jaw somewhat smaller than teeth of lower jaw. Number of vertebrae in one specimen (ZMA 3): +. COLOUR PATTERN IN ALCOHOL Male: tan, dorsal half darker than caudal half. Mouth greyish; preventral area dirty whiteyellow; cheek yellowish. Preoperculum greyishbrown. Preorbital stripe present; suborbital stripe very faint, occasionally absent; postorbital stripe sharply delimited. Black lateral band, running from postorbital stripe to hind edge of Fig. 2. Distribution of Crenicichla albopunctata Pellegrin, caudal fin; this band becoming more vague on 04, C. coppenamensis n. sp. and C. lugubris Heckel, 40 flanks, and very faint on caudal fin. Humeral in Surinam. spot on lateral band, just posteriorly to operculum. Silvery dots irregularly distributed on sides in dorsal twothird of body, on head, oper Predorsal contour straight; preventral con culum and cheek; on the head these dots are tour straight. Snout somewhat rounded in present below and above the postorbital stripe, lateral aspect, rounded from above. Maxilla on the preoperculum these dots extend to ven extending to below orbit. Orbital diameter tral part; humeral spot with dots. about half of head depth at orbit level. Nostrils Dorsal fin grey with white dots, smaller than slightly nearer to orbit than to postlabial dots on sides of body; most dots in posterior skinfold. part. Anal fin grey with dots only in posterior Dorsal fin origin just dorsal to pectoral fin part of fin. Ventral and pectoral fins much base; spines increasing in length to about lighter grey, without dots. Caudal fin grey with eighth, from there equal in length; fifth from same dots as in dorsal fin, most dots in dorsal last ray the longest, reaching to 4/5 of caudal fin in adult males, to a fourth of caudal fin in adult part of caudal fin; caudal spot on lateral band at base of caudal fin, just dorsal to lateral line females, to caudal fin base in juveniles. Anal fin scales, surrounded by eight dots; white caudal tip reaching to caudal fin base; fourth from last fin margin, except where lateral band ends. ray the longest. Second ray of ventral fin the Female: ground colour of body as in males. longest. Caudal fin rounded, with two to three Dorsal fin with dark dorsal margin; some basal scales at lateral line level, reduced to one females show two ocelli (black, white ring) over dorsally and ventrally. Interradial scales to a fourth of caudal fin. Four, rarely three (one) th and th spine; absent, occasionally in other females ocelli are ocelli from first dorsal fin scales between dorsal fin and most posterior spine to last ray; ocelli exclusively in females. scale of upper lateral line. with maximum Upper jaw five teeth series in Caudal fin with dark margin, except in the middle. Anal fin with dark lower margin. Ventral adults, in smaller specimens fewer, only two in with dark spine. 76

5 Juvenile (based on a 33 mm SL specimen, ZMA 0): conspicuous lateral band from Guiana do not show this difference. C. albopunctata occurs sympatrically with C. saxatilis tip of snout to end of caudal fin; above and downstream of Tabbetje below lateral band a tanish humeral zone; spot on lateral band; dark, ocellated caudal spot; small whitish ocelli in dorsal fin; dots on yellow both sides of lateral band, on lateral line scale series (these are present in females larger than Hede in the Marowijne basin (approx 'N). Crenicichla albopunctata can be distinguished from C. sipaliwini by its caudal fin length (CFL, fig. 3h). about 0 mm SL, and are white in adult males); caudal fin dorsally with a dark submargin with white edges. Crenicichla coppenamensis sp. n. (Figs. 2, 3, 4, 5, 6; Pl. 1 second and middle; Tabs. II and VIII) Crenicichla alta; (not Eigenmann, ) Boeseman, 52 DISTRIBUTION 8 (Saramacca River). In Surinam Crenicichla albopunctata occurs in the MATERIAL EXAMINED Marowijne river system only. The species is present almost everywhere in this river system Holotype: ZMA 41, mm SL male, (Ploeg, 86c). Samples were examined from the following rivers: Marowijne, Tapanahony, Surinam, left bank tributary to Linker Coppename River, V67, H. Nijssen. Paloemeu, Oelemari and Lawa. Outside Paratypes: Coppename River system: ZMA Surinam the species occurs in western French (42)/ZMA 42 (one), same data as Guiana, where the is from species present Marowijne River eastwards to Approuague River. In the Oyapock River the species seems to be absent. ZMA holotype; bank tributary to linker Coppename River (03 51'N, (), right 56 45'W), V67, H. ZMA Nijssen; (), left bank tributary to Coppename River (03 52'N, 56 55'W), V67, H. Nijssen; ZMA (), right bank REMARKS tributary to Coppename River (03 52'N, 56 53'W), V67, H. Nijssen. Although the silverywhite dots on the oper Saramacca River system: ZMA (), culum enables quick identification it seems left bank tributary to Kleine Saramacca River, useful to make comparisons with reminiscent km E.S.E. of confluence with Saramacca species occurring in the region. River, 2867, H. Nijssen; ZMA 5.6 Crenicichla albopunctata is distinct from C. alta (36), right bank tributary to Kleine Saramacca by the number of gillrakers (GR, fig. 3a); from C. coppenamensis by the number of scales in the River, km E.S.E. of confluence with Saramacca River, 67, H. Nijssen; upper lateral line (LI, fig. 3b), and by the ZMA.848 (one), Toekoemoetoe Creek, S. number of scales between the lateral lines of Poesoegroenoe Village, tributary to (L1/L2, fig. 3c); from C. labrina by the head Saramacca River, III64, M. Boeseman. length (HL, fig. 3d); from C. nickeriensis by the number of soft anal rays (ASR, fig. 3e); from C. saxatilis by the pectoral fin length (PFL, fig. DESCRIPTION 3f), and by the number of scales below the lateral lines (LL, fig. 3g), although the latter Based on holotype (morphometric and meristic depends on the sample that is used for com data are presented in tables II and VIII). parison. Specimens of C. saxatilis from the upper Suriname River and from French Predorsal contour straight; preventral contour straight. Snout somewhat rounded in 77

6 Plate I. Upper: Male of Crenicichla albopunctata Pellegrin, 04. ZMA 3, 2 mm SL. Second: Holotype of C. coppenamensis C. coppenamensis n. ZMA sp., 42, 1 sp. n., ZMA 41, mm SL, male. Middle: Female of SL. Fourth: mm Holotype of C. nickeriensis n. sp., ZMA 43, 1 mm SL, male. Lower: Female of C. nickeriensis n. sp., ZMA 44, 88 mm SL. Photographs L. A. van der Laan.

7 Fig. 3ah. Relation between standard length (SL) and other characters for comparison of C. albopunctata with C. alta, C. coppenamensis, C. labrina, C. nickeriensis, C. saxatilis, and C. sipaliwini; see also methods. 7

8 1 Qf 7 U Mn c«n Table II. Morphometric and meristic data for twelve specimens of Crenicichla coppenamensis n. sp. Abbreviations are listed in table VIII. CRENICICHLA COPPENAMENSISSP.N. SI CFl HL HO HW DCP LCP SN UJL LJL LLDS LLAS VFL PEL PN PN2 ORB Wl LI L2 / L2C LL SCOl DSP OSR 6R ASR P ASN MILLIMETERS HI , , , , b , , PERCENTAGES MEAN SO 1,00 0, , ZMA 7 841, HOLOTYPE, Surinam,Coppename River system creek at left bank ofleftcoppename River (03 54'N, 56 46'W), V 67, H Nljssen 2b /ZMA,PARATYPES, Surinam, Coppename River system, iii , creek at left bank ofleftcoppename River (03 54'N, 56 46'W), I JO /, n. Nljssen MEAN , SO I.I I lateral aspect, rounded from above. Maxilla COLOUR PATTERN IN ALCOHOL reaching to below orbit. Orbit in upper part of head, about half as as deep head at orbit level. Nostrils nearer to orbit than to postlabial skinfold. Male (based on holotype): tan; dark on the back, lighter on the belly; head greyish; preventral area light, cheek yellowish. Dorsal fin origin just posterior to hind edge of operculum; spines increasing in length to about ninth, from where equal in length; sixth Black and suborbital preorbital, postorbital, stripe. Lateral band on sides faint, prominent on caudal fin; between lateral line scale running ray from last the longest, reaching to a third of series. Occasionally a second band present, caudal fin, occasionally to end of fin in adult running from below pectoral fin to halfway anal males, to caudal fin base in females and fin. Lightyellow dots which are smaller than juveniles. Anal fin extending to caudal fin base flankscales; dots scattered on flanks, more with fifth ray from last the longest. Second ray numerous along lateral lines. Black humeral of ventral fin the longest. Caudal fin rounded spot posterior to operculum, four scales long, with three to four basal scales at lateral line ocellated by a silvery ring; spot in adults with level, diminishing to one dorsally and ventrally. Interradial scales to a fourth of caudal fin center lateral on line, in below lateral juveniles line. length. Pectoral rounded. Dorsal fin grey and scattered with whitish Upper jaw with maximum five teeth series, dots, which are smaller than the dots on flanks; all teeth equal in length; lower jaw more dots on posterior part of fin. Anal fin grey with maximum four teeth series, outer series with little scattered with hyaline dots. Caudal fin grey larger teeth; in specimens smaller than 55 mm SL only two teeth series in both jaws. with series of small whitish dots, arranged in Number of vertebrae in one specimen (ZMA whitish lines. Black, ocellated caudal spot dorsally on caudal fin, just posterior to basal scales. 41): +. Many specimens (absent in holotype) show a 80

9 small dark spot at end of caudal fin. Pectoral scales between the lateral lines (L1/L2, fig. 5a), and ventral fins much lighter grey, almost the number of spines in the dorsal lin (DSP, fig. white. Female (based on 1 mm SL paratype, ZMA 42): somewhat lighter than males; lateral band on head and ring around humeral spot less conspicuous than in males. No ring around Crenicichla be 5b). coppenamensis can distinguished from C. nickeriensis by the ventral fin the number of scales length (VFL, fig. 5c), between the lateral lines (L1/L2, fig. 5d), and the angle of the snout (ASN, fig. 5e); from C. caudal spot; occasionally a horizontal band saxatilis by the depth of the caudal peduncle along abdominal side in as some males. Yellow dots absent. Dorsal fin much than in lighter males, dark without margined; dots. Anal fin (DCP, fig. 5f), the length of the last dorsal spine (LLDS, fig. 5g), the ventral fin length (VFL, fig. 5h), the number of scales below the lateral without dots and much lighter than in males. lines (LL, fig. 6a), and the number of dorsal Sexual dimorphism only in specimens larger than about 0 mm SL. Caudal fin without spines (DSP, fig. 6b). Crenicichla sipaliwini can be distinguished by the ventral fin length (VFL, dots; often a small dark spot at end of lateral fig. 6c), by the more slender body shape and by band; dark margin along dorsal edge, occasionally a white margin instead, together with a the much larger humeral spot (the latter two not in a figure). dark submargin. Females of C. coppenamensis do not show ocelli Juvenile (based on a 25 mm SL paratype, in their dorsal fin, as females of the related ZMA ): three horizontal zones: brown species C. saxatilis and C. albopunctata. upper part, a darker band from tip of snout to The position of the humeral spot shows caudal fin, and a tan ventral part; preventral ontogenetic variation: adults have the center of area light. Humeral spot on lateral band, but the spot on the lateral line (above the lateral well below the lateral line, ocellated by a light band; in specimens smaller than about 0 mm brown ring. Dorsal fin darkmargined. Dots SL the center of the spot is entirely below the absent. lateral line (still above the lateral band when larger than 40 mm). This variation is shared DISTRIBUTION with C. coppenamensis, C. sipaliwini, and C. saxatilis. In Surinam Crenicichla coppenamensis is present in the Coppename and Saramacca River systems. No material is available from the lower Coppename River. Crenicichla nickeriensis sp. n. (Figs. 3, 5, 6, 7, 8; Pl. I fourth and lower; Tabs. III and VIII) Local names: Datra fisi, djonkodjo mantuari, donfo. REMARKS Crenicichla coppenamensis is distinct from C. alta MATERIAL EXAMINED by the ventral fin length (VFL, fig. 4a), the number of scales below the lateral lines (LL, Holotype: ZMA 43, 1 mm SL male, fig. 4b), the number of scales between the Surinam, right bank tributary to Nickerie lateral lines (L1/L2, fig. 4c), and the number of River, km W.S.W. of Stondansie Fall, 5IV gillrakers (GR, fig. 4d); from C. labrina by the 67, H. Nijssen. head length (HL, fig. 4e), the depth of the Paratypes: Nickerie River system: ZMA caudal peduncle (DCP, fig. 4f), the snout (four), Stondansie Fall, 5IV67, H. Nijssen; length (SN, fig. 4g), the number of scales below ZMA 06 (36), Fallawatra River, 5 km the lateral lines (LL, fig. 4h), the number of S.S.W. of Stondansie Fall, 6IV67, H. 81

10 Fig. 4ah. Relation between standard length (SL) and other characters for comparison of C. coppenamensis with C. alta and C. labrina; see also methods. 82

11 Fig. 5ah. Relation between standard length (SL) and other characters for comparison of C. coppenamensis with C. labrina, C. nickeriensis and C. saxatilis; see also methods. 83

12 Fig. 6ah. Relation between standard length (SL) and other characters for comparison ofc. coppenamensis with C. saxatilis and C. sipaliwini, and of C. nickeriensis with C. alta, C. labrina, and C. saxatilis; see also methods. 84

13 Table III. Morphometric and meristic data for twelve specimens of Crenicichla nickeriensis n. sp. Abbreviations are listed in table VIII. CRENICICHLA NICKERIENSIS SP.N. SL CFL HL HO HW DCP LCP SN UJL UL LLDS LLAS VFL PFL PN PN2 ORB Wl LI L2 L1/L2 L2C LL SCDL DSP DSR 6R ASR P ASN MILLIMETERS PERCENTAGES MEAN SO , ZMA 43, HOLOTYPE, Surinam, Nickerle River, right tributary 25, , creek tonickerie River, km WSW of StondansieFall, 5IV 67, H. Nijssen 2/ZMA 7843, PARATYPES, Surinam, Nickerie River, right tributary creek to Nickerie River, km WSW of StondansieFall. 5IV67. H Nijssen MEAN SO , Nijssen; ZMA 05/44(), same data as holotype; RMNH (35), right tributary creek to Fallawatra River near confluence with Nickerie River, 2II71, M. Boeseman; Matappi Creek V (05 01'N, 57.5'W), 80, R. P. Vari; USNM 60 (one), Kapoeri Creek, ca. 4 km N from intersection with Corantijn River (05 'N, 57 'W), RMNH 30554/ZMA (nine), right bank tributary to Nickerie River, 3 km below Stondansie Fall, 1II71, M. Boeseman; RMNH 2855/30555 (five), right bank tributary to middle course of Maratakka River, 2871, M. Boeseman; RMNH (two), tributary to middle course of Maratakka River, 1 III71, M. Boeseman; RMNH 2851/2852 (), right bank tributary to middle course of Maratakka River at second savanna, 61 71, M. Boeseman; RMNH 2853/2854 (), Bigi Birri Creek to Maratakka River, above Cupido, 8III71, M. Boeseman; RMNH (one), ditches around Wageningen, 171, M. Boeseman. Corantijn River system: USNM 60 (two), Steen Creek, (04 4'N, 45 'W), IX 80, R. P. Vari; USNM 603 (one), near Anjoemara (04 50'N, 57 26'W), IX80, R. P. Vari; USNM 6044 (six), a small creek close to Mataway (04 5'N, 57 42'W), V80, R. P. Vari; USNM 6028 (one), Fig. 7. Distribution of C. multispinosa Pellagrin, 03 and C. nickeriensis n. sp. in Surinam. 85

14 IX80, R. P. Vari; USNM 5848 (six), Corantijn River along mud flats at Wakay, just below Amerindian camp boatlanding (05 'N, 57 'W), 7IX80, R. P. Vari; RMNH (), Sand Fall, near Avanavero separating line; mouth and head grey; preventral area light, cheek tan, operculum grey. Faint preorbital stripe; black postorbital stripe; chain of irregularly shaped black blotches along lateral lines, from operculum to Fall in Kabalebo River, IX65, G. F. caudal fin spot, from where a faint stripe runs Mees; ZMA (two), right bank to end of caudal fin. Short suborbital stripe, tributary to Kabalebo River, just downstream triangular. Pink dots, irregularly distributed on of Avanavero Fall, IV65, G. F. Mees. darker part of flanks, posteriorly more concentrated than just beyond operculum; dots DESCRIPTION smaller than flankscalesize. Dorsal fin grey with pink dots, similar to Based on holotype (morphometric and meristic those on flanks, in posterior part. Anal fin grey data are presented in tables III and VIII). scattered with dots, becoming smaller caudad; dots ranked in lateral series showing horizontal Predorsal contour straight; preventral contour somewhat convex. Snout rounded in lines. Ventral and pectoral fins lighter grey, immaculate. Caudal fin grey with dots like lateral aspect and from above. Lower jaw a little prognathous, in other specimens more prognathous; maxilla reaching to anterior margin those on flanks; occasionally a small dark spot in end of faint horizontal band. Ocellated of orbit. Orbital diameterabout a third of head caudal spot dorsally on caudal fin base, depth at orbit level. Nostrils nearer to orbit than to postlabial skinfold. encircled by about eight pink dots. Female (based on 88 mm SL paratype, ZMA Dorsal fin origin just posterior to vertical 44): Ground colouration of and fins body from posterior edge of operculum; spines as in male; preorbital and postorbital stripe, the increasing in length to about th, from where latter conspicuous, dorsally and ventrally equal in length; fifth ray from last the longest, margined by a white zone; humeral spot elongated horizontally, as long as eye, half as in males reaching to a third of caudal fin, occasionally to halfway, in females to caudal spot, in juveniles up to about 50 mm SL reaching to deep as long, situated on lateral line, ocellated by a light brown zone. Chain of blotches along caudal fin base. Anal fin reaching to caudal fin lateral lines absent. base; fifth from last ray the longest. Caudal fin Dorsal fin somewhat lighter than in males; rounded, with 24 basal scales at lateral line from edge of fin an alternating pattern of dark, level, diminishing to one dorsad and ventrad. white and dark spots between the rays. Black or Interradial scales to a fourth of caudal fin white in dorsal spot fin than ocelli in C. (smaller length. Pectoral fin rounded. Upper jaw with maximum six teeth series, all teeth equal in length; lower jaw with maximum saxatilis), from only one spot in the last spines to up occasionally on every interradial membrane along the dorsal margin of the fin. Occa four teeth series, labial teeth series little larger; sionally no spots at all. Anal fin with dark inner teeth equal in length; external teeth series margin; in last three rays six transversal lines, fixed, internal series depressible. Number of vertebrae in one specimen (ZMA 43): +. COLOUR PATTERN IN ALCOHOL forming an alternating grey and white pattern. Ventral and pectoral fin white. Caudal fin pale dorsally with dark and white submargin, ventrally a dark margin only; three margin faint dark bars, dots absent. Round, black caudal spot, ocellated by a lighter ring, situated Male (based on holotype): Brownish, dark on back, tan on belly, without conspicuous axial just above extending fin. lateral line scales in caudal 86

15 Juvenile (based on 48 mm SL paratype, ZMA 05). Dark lateral band running from tip of snout to caudal peduncle; lateral band becoming more vague with increasing age. Humeral spot round, smaller than not eye; ocellated, gests that the females remain smaller than the males. The females show a flattened humeral spot, which is remarkable; normally the spot is more or less round in C. alta and C. (e.g. coppenamensis). Small juveniles of C. nickeriensis are situated below lateral line, on lateral band. not distinguishable from juveniles of C. saxatilis Dorsal fin greyish, with white margin and dark submargin; small white dots in posterior part of and C. coppenamensis. dorsal fin, arranged in series. Anal fin grey Crenicichla saxatilis (Linnaeus, 58) without dots. Ventral and fins much pectoral lighter than other fins. Caudal fin grey; dorsal part of caudal fin with and margin dark submargin; three white blotches in a series caudad. Ocellated dark caudal spot. (Figs. 3, 5, 6, 8,, ; Tabs. IV and VIII) Sparus saxatilis Linnaeus, 58: 8 (original description; Surinam). Perca saxatilis; Bloch 2: 7, fig. 30 (description; Surinam). Crenicichla saxatilis; Heckel, 40: 432 (Surinam); Regan 05: (Surinam); Steindachner, : DISTRIBUTION (description; Marowijne River); Wheeler 58: 6 (description; photograph of holotype of Scarus pavoninus Gronovius, 54); Richter & Nijssen, 80: 5 In Surinam C. nickeriensis is known from Nickerie, Kabalebo, Maratakka, and Corantijn River. Available material from Essequibo River, described by Eigenmann,, does not include this species, which suggests C. nickeriensis is restricted to the above four river systems. (Surinam, Brokopondo Reservoir); Nijssen, 67: 6 (Surinam); Kullander, 82: 655 (discussion on types); Ploeg, 86a: 47 [description; designation of lectotype and restriction of the type locality (Suriname River, Carolina Creek)] and 86c: 2 (Marowijne River system). Scarus pavoninus Gronovius, 54: 63 (Surinam) Crenicichla saxatilis albopunctata Pellegrin, 04: 374 (in part; Surinam). Crenicichla alta; (not Eigenmann, ) Nijssen, 66: 5 REMARKS (Brokopondo Reservoir); Nijssen, 67: 6 (Suriname River system). Crenicichla nickeriensis can be distinguished from C. alta by the head width (HW, fig. 6d); from MATERIAL EXAMINED C. labrina by the head length (HL, fig. 6e), and the number of gillrakers (GR, fig. 6f); from C. saxatilis by the number of gillrakers (GR, fig. Lectotype: NRM 5583; paralectotypes: NRM 6g) and by the angle of the snout (ASN, fig and BMNH 53..: ; 6 specimens from the Suriname, Commewijne, and from C. 6h); sipaliwini by the caudal fin length (CFL, fig. 8a) and by the angle of the snout (ASN, fig. 8b). Crenicichla nickeriensis can be distinguished from all discussed species by the Marowijne River systems (Ploeg, 86a: 4 and 86c: 2). Additional material: BMNH 81.:, Nickerie River, Camp MacClemen, no chain of blotches along the lateral lines in adult date, R. P. Varietal. BMNH 81.: 0 males. The dots on the abdomen in adult males are, Nickerie River, Koekwie Creek, no date, R. P. Vari et al. not as conspicuous as in C. saxatilis. In C. saxatilis these dots are very bright silvery, in C. nickeriensis pale pink. DISTRIBUTION Although many specimens in the RMNH and ZMA collections exceed 0 mm SL, none In Surinam Crenicichla saxatilis is present in the of the females exceeds 0 mm SL, which sug Nickerie, Suriname and Commewijne River 87

16 Fig. 8ah. Relation between standard length (SL) and other characters for comparison of C. nickeriensis with C. sipaliwini and of C. saxatilis with C. alta, C. labrina and C. sipaliwini; see also methods. 88

17 Table IV. Morphometric and meristic data for twelve specimens of Crenicichla saxatilis (Linnaeus, 58). Abbreviations are listed in table VIII. CRENICICHLA SAXATILIS (LINNAEUS, 58) 51 CFL H HO HW DCP LCP SN UJL LJL LLOS LLAS VFL PEL PN PN2 ORB Wl LI L2 L1/L2 L2C LL SCDL DSP DSR GR ASR P ASN MILLIMETERS so so , ' PERCENTAGES ,2.5 MEAN SO , , /ZMA 02,Surinam, Surinwne River, Carolina Creek, topara River, XI166, Coll. H. Nijssen ofpara, 30XI64. Coll. S.M Leopold III & J P Oosse , MEAN ! ,5 SO ,86 0, systems, and in the Marrowijne River system Gronovius, 54; the caption presents this downstreams of Tabbetje Hede. Outside specimen as the holotype of Crenicichla saxatilis. Surinam the species is present in French Because Linnaeus' description was based on Guiana and possibly in Guyana. this specimen and on two specimens present in REMARKS A detailed description of Crenicichla saxatilis is presented by Ploeg (86a: 50). For comparison morphometric and meristic data are presented in tables IV and VIII). Crenicichla saxatilis differs from C. alta by the depth of the caudal peduncle (DCP, fig. 8c), the pectoral fin length (PFL, fig. 8d), the number of scales between the upper lateral line and the dorsal fin (SCDL, fig. 8e), and by the number of gillrakers (GR, fig. 8f); from C. labrina by the head and from length (HL, fig. 8g); C. sipaliwini by the depth of the caudal peduncle the number of scales (DCP, fig. 8h), below the lateral lines (LL, fig. a), and by the number of scales between the upper lateral line and the dorsal fin (SCDL, fig. the number of dor b), sal spines (DSP, fig. c), and by the number of pectoral fin rays (P, fig. lod). Wheeler (58: 6) redescribed and figured (pi. 30, fig. 2) the holotype of Scarus pavoninus Fig.. Distribution of C. saxatilis (Linnaeus, 58) and C. sipaliwini n. in Surinam. sp. 8

18 * Crenicichla sipaliwini sp. n. (Figs. 3, 6, 8,, and ; Pl. II upper; Tabs. V and VIII) MATERIAL EXAMINED his own collection, the Gronoviusspecimen is one of the three syntypes of C. saxatilis. Wheeler states that, according to the Lin Lonnberg, naean specimen was not among those still preserved in the University of Upsala, and therefore presumably no longer in existence. However, according to Fernholm & Wheeler All from Corantijn River system, Sipaliwini River near Sipaliwini airstrip. (83: ), the Linnaean material is preserved in the collection of the NRM. The Holotype: RMNH 30561, 0 mm SL, prob material consists of two specimens: NRM 5583 ably female, 6II66, G. F. Mees. and A of the first of these two photograph is presented by Ploeg (86a), who designated this of Crenicichla saxatilis. specimen as lectotype All from same RMNH Paratypes: locality: 2856 (), 6II66, G. F. RMNH Mees; 2857/ZMA (nine), 166, G. F. The second specimen may represent the Mees; RMNH 2858 (five), II66, G. F. specimen figured by Linnaeus (54: 65, fig.!) Mees; RMNH 285 (one), 6II66, G. F. Mees; ZMA 46 (eight), /26166, G. Very conspicuous in Crenicichla saxatilis is the F. Mees. dorsoanteriornotch in the humeral spot. Occasionally a second notch is present dorsally. C. saxatilis never shows a circular spot as present in DESCRIPTION C. alta. The position of the humeral spot is variable. In larger upstream specimens the spot Based on holotype (morphometric and meristic is situated on the lateral line; in upstream data are presented in tables V and VIII). juveniles and most downstream specimens it is situated below the lateral line; in very young Predorsal contour straight; preventral contour straight. Snout pointed in lateral aspect, specimens it is situated well below the lateral rounded from above. Maxilla not reaching to line and on the lateral band. orbit, or to anterior margin of orbit. Orbit Table V. Morphometric and meristic data for twelve specimens of Crenicichla sipaliwini n. sp. Abbreviations are listed in table VIII. CRENICICHLA SIPALIWINI SPN SL CFL HI HO HW DCP LCP SN UJL LJL LLDS LLAS VFL PFL PN PN2 ORB Wl LI L2 L1/L2 L2C LL SCDL DSP OSR GR ASR P ASN MILLIMETERS PERCENTAGES , ,5.1 MEAN , SD , 0.4 0, , , RMNH 30561, HOLOTYPE, Surinam, CorantijneRiver system, , Sipaliwini River nearsipaliwini airstrip, 666, flees.6 34, , /ZMA 1.683/7846, PARATYPES,Surinam,CorantijneRiver system, SipaliwiniRiver nearsipaliwini airstrip, /26166, Mees , , MEAN.1 36, , SD ,55 0,

19 Fig. ah. Relation between standard length (SL) and other characters for comparison of C. saxatilis with C. sipaliwini and of C. sipaliwini with C. alta, and C. labrina; see also methods. 1

20 Plate II. Upper: Holotype of Crenicichla sipaliwini n. RMNH Heckel, 40, sp., 30561, 0 SL. Second: C. mm lugubris ZMA.7, 4 mm SL. Third: Male of C. multispinosa Pellegrin, 03, ZMA.8, 5 mm SL. Lower: Female of C. multispinosa Pellegrin, 03, ZMA.8, 7 mm SL. Photographs L. A. van der Laan. about onethird as deep as head at orbit level. Scales on cheek remarkably smaller than on operculum. Nostrils nearer to orbit than to skinfold. postlabial Dorsal fin origin just anterior to hind edge of operculum; spines increasing in to about length ninth, from where equal in sixth length; ray from last the longest, reaching to a third of caudal fin, occasionally to end of fin in adults, to caudal fin base in Anal fin juveniles. extending to caudal fin base with fifth ray from last the longest. Ventral fin with first soft ray the 2

21 longest. Caudal fin rounded with three basal REMARKS scales at lateral line level, diminishing to one dorsally and ventrally. Interradial scales to a Crenicichla sipaliwini is distinct from C. alta by fourth of caudal fin length. Pectoral fin rounded. Upper jaw with maximum four teeth series, all teeth equal in length; lower with maxi jaw the number of scales below the lateral lines (LL, from fig. loe); C. labrina by the head length (HL, fig. 1 Of), the snout length (SN, fig. log), and the number by of scales below the lateral mum four teeth series, outer series with little lines (LL, fig. loh). larger teeth. The relative position of the humeral spot, as No radiographs available in C. coppenamensis, shows ontogenetic variation: adults have the centre of the spot on the COLOUR PATTERN IN ALCOHOL lateral line (above the lateral band); in smaller specimens the centre of the spot is entirely Because of lack of conspicuous below the lateral line. sexual dimorphism male and female are not described No conspicuous sexual dimorphism was separately. Juveniles only differ in the position detected in Crenicichla sipaliwini. The holotype and size of the humeral spot and are therefore not treated separately. shows an ocellus in the dorsal fin as present in females of for instance C. saxatilis and C. Description based mainly on holotype. albopunctata. However, it also shows white dots Yellowgrey on belly, dorsally more greyyellow; dorsal part of head dark grey, sides of in the dorsal and anal fins as in males of C. saxatilis and C. albopunctata. The last dorsal and head greyish, ventral part yellowishgrey; anal fin rays are not extremely elongated as in preventral area and cheek yellowish. In conspicuous black preorbital and postor C. saxatilis and C. albopunctata. This may indicate that either females are at only hand, or bital stripes, suborbital stripe absent. Vague that sexual dimorphism is lacking. Because of lateral band, running between lateral line scale series (in larger specimens such as holotype the ocellus in the dorsal fin, I suppose the holotype is a female. absent). Black humeral spot, occasionally with Crenicichla sipaliwini is more elongated than dorsoanterior notch. Humeral spot in adults with its center on lateral line, in juveniles below other Surinam saxatilislike species. This may be due to the habitat: fast running water. lateral line, in the latter relatively larger. Dorsal fin grey. However, often without (as Crenicichla lugubris Heckel, 40 (Figs. 2; Pl. II second; Tabs. VI and VIII) Crenicichla lugubris Heckel, 40: 4 (original description: Rio Negro); Eigenmann, : 5 (Essequibo); Pellegrin, 04: 383 (Orinoco, Tonantins, Iga, Guyana); Ploeg, 86b: 72 (Tocantins River); Regan, 05: 5 (River Capin, River Cupai, Essequibo River); LoweMcConnell, 6: 6 (Essequibo River, Demerara River, Upper Cuyuni River). Crenicichla adspersa Heckel, 40: 4 (original description; Mato Grosso). in holotype) a black margin. Holotype with one black ocellus between first two soft dorsal rays. Vague dots in posterior part of fin, arranged in rows from base of to rays more caudal distal margins. Anal fin grey with dots arranged as in dorsal fin. Caudal fin grey with series of same kind of dots as in dorsal fin, arranged in light bars. Black, ocellated caudal spot dorsally on caudal fin, to basal scales. Small just posterior dark spot at the centre of caudal fin end. Pectoral fin hyaline, ventral fin white. Crenicichla funebris Heckel, 40: 4 (original description; Rio Guapore); Giinther, 62: 307 (River Capin, DISTRIBUTION Guyana). Crenicichla johanna var. B. strigata Giinther, 62: 306 Crenicichla sipaliwini is known only from the (original description; River Capin, River Cupai); LoweMcConnell, 6: 6 (River Capin, Manari, Sipaliwini River, Corantijn River sytem. Rupununi). 3

22 MATERIAL EXAMINED Predorsal contour slightly convex; preventral Syntype: NMW 61.8, 1 mm Brazil, Rio Negro. SL from contour anteriorly slightly convex, below orbit straight. Snout in lateral pointed aspect, rounded from above. Maxilla extending to SURINAM, Corantijn River system: USNM 60 (one), Matapi creek, about 1 km from intersection with Corantijn River (05 00'N, 57 'W), IX80, R. P. Vari; USNM 6030 (one), Creek about 2 km N. of Matapi (05 03'N, 57 'W), IX80, R. P. Vari; USNM 60 stream (one), entering Corantijn River at approximately km 385, slightly N. of Tiger Falls IX (04 00'N 58 02'W), 80, R. P. Vari et al.; USNM 6032 (one), Matappi creek V (05 01'N, 57.5'W), 80, H. M. Madari; USNM 6033 (three), small creek close to Mataway (04 5'N, 57 42' W), III80, R. P. Vari et al. ; ZMA.7 (one)/rmnh 373 Kabalebo (one), River, Avanavero Fall, Zandvallen, IX 65, G. F. Mees. anterior margin of orbit. Orbit large, diameter about half head depth at orbital level. Nostrils nearer to postlabial skinfold than to orbit. Dorsal fin origin just dorsal to hind edge of operculum; spines increasing in length to fourth, from which equal in length; fifth from last ray longest and reaching to caudal fin base. Anal fin extending to caudal fin base; fourth ray from last longest. Ventral fin rounded; first soft ray the Caudal longest. fin straight, with up to six basal scale series. Upper jaw with maximum eight teeth series, lower with maximum five teeth series. jaw In both jaws outer teeth the largest. DESCRIPTION COLOUR PATTERN IN ALCOHOL Based on ZMA.7. No sexual Based principally on ZMA.7 (morphometric and meristic data are presented in tabs. VI and VIII). Number of vertebrae in one specimen (syntype: NMW ): dimorphism in colour pattern found. Ground colour of body brown, countershaded. No preorbital, Table VI. Morphometric and meristic data for ten specimens of Crenicichla lugubris Heckel, 40. Abbreviations are listed in table VIII. CRENICICHLA LUGUBRIS HECKEL. 40 SL CFL HL HO HW DCP LCP SN UJL LJL LLDS LLAS VFL PFL PN PN2 ORB Wl LI L2 LI/ L2C LL SCDL DSP DSR 6R ASR P ASN MILLIMETERS ! ' PERCENTAGES MEAN.0, , , SO ,0 4, BMNH 81.:08.Surinam. Nickerle River. 80,Varl etal USNM 6030, Surinam. CorantijneRiver. Creek about2 km ,3.5 North of Matapi (05.03'N, 57 W), IX80, Veri et el ,8.7, /6USNM 6033, Surinam,CorantijneRiver,Small creek close to Mataway (05 0IN, 57 'W), 0.Varietal , RMNH Unreg,Surinam. Corantijne River. KabaleboRiver, AvanaveroFall, Zandvollen, IX65, Mees MEAN, , 8ZMA 7, Surinam, Corantijne River.KabaleboRiver, SO , ,61 0, AvanaveroFall. Zandvallen, IX65. Mees USNM 60, Surinam, Corantijne River, Matapi Creek, ca 1 km from intersection with CorantijneRiver (05.00'N, 57.'W), IX80, Veri et el USNM 60. Surinam, Corantijne River. Stream entering CorantijneRiver at approximately km 38, slightly North of Tiger Fells (00'N, 52'W), IX80, Vari et el. 4

23 postorbital, or suborbital spot/stripe. Five bars between dorsal fin and lateral line. Dorsal, anal, ventral, and pectoral fins grey. Dorsal upper fin dark Caudal fin margined. with triangular black blotch above middle of base. just part, Surinam); Regan, 05: 4 (description, bibliography; Surinam); Ploeg, 86c: 0 [restriction of type locality (Marowijne River, S. of Gran Creek) Surinam, French Guiana], Crenicichla lugubris (not Heckel, 40), Richter & Nijssen, 80: 4 (Suriname River). Humeral spot just posterior to pectoral fin. DISTRIBUTION MATERIAL EXAMINED In Surinam C. lugubris is found in the Corantijn Lectotype, designated by Ploeg (86c: 0): River system only. From it is recorded Guyana MNHN 542, 7 mm SL, French Guiana, by Eigenmann (, Essequibo River system). The species is also in the northern present Amazonian branches in Brazil. Hitherto C. Melinon. Paralectotypes: MNHN 542 (two), MNHN 47 (three), French Guiana, Melinon. lugubris is not recorded from French Guiana. Marowijne River system: 4 specimens (see Ploeg, 86c: 0). REMARKS Suriname River system: ZMA 04 (one), Awara Creek, right bank tributary to Suriname This is the first record of genuine C. lugubris from Surinam. Richter & Nijssen (80: 5) erroneously recorded the from the species Suriname basin, based on a misidentification (now C. multispinosa). Crenicichla 03 multispinosa Pellegrin, (Fig. 7; Pl. II third and lower; Tabs. VII and VIII) Crenicichla multispinosa Pellegrin, 03: 4 (original description; Guyane Pellegrin, 04: 380 (in River, 1 km S. of Botopasi, III67, H. Nijssen; RMNH (), Awaradam, VII65, G. F. Mees; ZMA 67 (five), Brokopondo Reservoir, 1 km E. of former Mamadam Rapid, /XII78, C. J. J. Richter; ZMA 5.5 Gran MAu (three), Creek, right bank tributary to Gran Rio, 1 km N.E. of Dombaai km (=Bendi Watra), S.W. ofdjoemoe, 30167, H. Nijssen; ZMA (four), Jenjee Creek, bank right tributary to Suriname River, km N. of Table VII. Morphometric and meristic data for twelve specimens Pellegrin, 03. Abbreviations are listed in table VIII. of Crenicichla multispinosa CRENICICHLA MULTISPINOSA PELLE6RIN. 03 SL CFL HI HD HW DCP LCP SN UJL LJL LLDS LLAS VFL PFL PN PN2 ORB Wl LI L2 L1/L2 L2C LL SCDL DSP DSR 6R ASR P ASN MILLIMETERS t PERCENTAGES ,.6.4, MEAN,4, , , SD ,4 4,52 0, , , , , ,0,.0, /ZMA 8 Surinam, SurinameRiver, SopoFoll (=Mamadem), IX 66, H. Nijssen RMNH 30562, Surinam, SurinameRiver in andbetween falls at , Mamadam, / MEAN SO

24 Snouttip Botopasi, III67, H. Nijssen; RMNH (), Mamadam, in and between falls, /164, M. Boeseman; RMNH 2874 (one), Mamadam, under just Grand Fall, /164, M. Boeseman; RMNH 374 near (one), Brokopondo, 30XII63, M. Boeseman; RMNH 375 (three), near (=Mamadam) Falls, /164, M. Boeseman; ZMA (three), Suriname River, 1 km S. of Botopasi, 167, H. ZMA Nijssen; (five), Suriname River, km N. of Botopasi, III67, H. Nijssen; MNHN A 45 (one), no date, Melinon. Brokopondo, 4164, M. Boeseman; RMNH 376 (nine), near Brokopondo, 28164, M. Boeseman; RMNH 377 (five), near DESCRIPTION Brokopondo, 7II64, M. Boeseman; RMNH 378 (one), near Brokopondo, 2II Based on 5 mm SL male, ZMA.8 64, M. Boeseman; ZMA (seven), (morphometric and meristic data are presented Parwapa (=Paba) Creek, left bank tributary in tables VII and VIII). to Suriname River, 2.5 km N. of Botopasi near Foetoenakaba(= Voetoekaba), 167, H. Nijssen; ZMA 5.5 (five), right bank Head depressed; predorsal contour straight; preventral contour straight. Snout pointed in lateral rounded from above. aspect, Lower jaw tributary to Gran Rio, 4 km N.E. of Awadam slightly prognathous only; maxilla not reaching ( = Awaradam) rapids, 167, H. Nijssen; anterior margin of orbit. Orbital diameter ZMA 5.8(three), Sopo Fall (= Mamadam) about half as deep as head at orbit level. in Suriname River, IX66, H. Nijssen; Nostrils nearer to orbit than to postlabial ZMA 5. (one), Sopo Fall (= Mamadam) in Suriname River, S.W. tributary of Brokopondo Reservoir, IX66, H. ZMA.8 Nijssen; (two), Sopo skinfold. Origin of dorsal fin just posterior to hind edge of operculum; spines increasing in length to about ninth, from there equal in length; Table VIII. Ranges in morphometric and meristic characters in Crenicichla albopunctata Pellegrin, 04, C. coppenamensis n. C. nickeriensis C. saxatilis sp., n. (Linnaeus, 58), C. sipaliwini C. lugubris Heckel, 40, and sp., n. sp., C. multispinosa Pellegrin, 03. Abbreviations in parentheses behind the characters. ALBOPUNCTATA COPPE NAME NSIS NICKERIENSIS SAXATIL1S SIPALIWINI LUGUBRIS MULTISPINOSA StandardLength (SL) CaudalFin Length (CFL) Depth Caudal Peduncle (DCP) Length Caudal Peduncle (LCP) SnoutLength (SN) Upper Jaw length (UJL) Lower Jaw Length (LJL) Length Last DorsalSpine (LLDS) Length Last AnalSpine (LLAS) Ventral Fin Length (VFL) Pectoral Fin Length (PFL) DistanceOr bit Nostr (PN) D istance Nostri 1 (PN2) Orbital Width (ORB) interorbital Width (Wl) ScalesAnterior Lateral Line (LI) Scales Posterior Lateral Line (L2) Scales Between Lateral Lines (L1L2) Lateral LineScales on Caudal Fin (L2C) Scaleseries Below Lateral Lines (LL) Scales Between Dorsal Fin and L1 (SCDL) Dorsal Spines (DSP) 25 Dorsal SoftRays (DSR) Gill Rakers (GR) Anal Spines (ASR) Pectoral Fin Rays (P) Angle Snout (ASN)

25 reaching to 2/3 of caudal fin length in adults, to males, though less conspicuous ocellated. Dark a fifth of caudal fin in juveniles; third from last band along middle; margins dark dorsally and soft ray the longest. Anal fin reaching to caudal ventrally. fin base; third from last ray the longest. Ventral fin acute, first soft ray the longest. Caudal fin Juvenile (based on 45 mm SL specimen, RMNH 2874): Body showing three zones: a with free ray tips; five basal scales at lateral line brownish upper zone, a darker zone from orbit level, to two ventrally and dorsally; interradial to caudal peduncle, and a lighter ventral zone; scales to a third of caudal fin. Pectoral fin no bluish part on head; dots on flanks yellowish rounded. instead of silvery, as in adult males; caudal spot Scales on suboperculum, operculum, cheek, ocellated by a light brown ring; dark stripe from anterior part of back, and occasionally just pos caudal spot to end of rays; dark margin in dor terior to base of pectoral fin cycloid, elsewhere sal fin and in anal fin; look like larger juveniles ctenoid. Lateral line scales larger than adjacent adult males, except for the absence of a spot scales. Upper jaw with maximum seven teeth series, below the orbit and pattern the presence of yellowish coloured dots on the sides. lower jaw with maximum four teeth series. Number of vertebrae in three specimens (lectotype and two paralectotypes MNHN DISTRIBUTION 542): + / + / +. Crenicichla multispinosa seems to be endemic in Suriname and Marowijne River systems. In COLOUR PATTERN IN ALCOHOL these river systems the species is present in the middle and upper parts only. Male (based on 5 mm SL specimen, ZMA.8): Brown; ventrally lighter and yellowish; cheek yellow; mouth bluegreyish; preventral area dirty white; a pattern of brown blotches bluish on a below orbit background in (absent some which then show specimens, a REMARKS Ploeg (86c) restricted the type locality of C. multispinosa to Marowijne River, S. of Gran Creek. suborbital stripe). Posterior part of sides with Crenicichla multispinosa is very similar to C. many silvery dots, smaller than flankscalesize. ternetzi Norman, 26, which seems to be Dorsal fin whitish dark grey, margined; whitish dots scattered on dorsal fin. Anal fin whitish, dark margined; few white dots which are similar to those in dorsal fin. Ventral and endemic in the Oyapock River system (French Guiana and Brazil). An easy character to separate the two species first on sight, apart from other characters 86c: is (Ploeg, ), pectoral fin hyaline. Caudal fin with many the absence of the caudal fin spot in C. ternetzi. silvery dots; ground colour somewhat darker than dorsal fin; dark caudal spot, silvery ocellated. Female (based on 1 mm SL specimen, Although C. is named multispinosa after its high number of dorsal spines, the species is certainly not unique in this character: C. lugubris and C. ternetzi also have up to 25 dorsal spines. RMNH 30562): Ground colour as in males. Dark lateral band, running between lateral line REFERENCES scale series. Neither suborbital stripe, nor a spot pattern below orbit; dots absent on flanks and on fins. Dorsal fin somewhat darker than in males, lacking the dark Anal fin less margins. whitish; dark Ventral margined. and pectoral fins in males. Caudal fin with as spot as in BLOCH, M. E., 2. Naturgeschichte der auslandische Fische, 6: 36, 26 pis. (J. Morino & Comp., Berlin). BOESEMAN, M., 52. A preliminary list of Surinam fishes not included in Eigenmann's enumeration of. Zool. Meded. (): 0. 7

26 EIGENMANN, C. H.,. The freshwater fishes of British, 67a. Ichthyological observations: JanuaryMarch Guiana, including a study of the ecological grouping 67. In: Progress Reports, Biological Brokopondo of the species and the relation of the fauna of the Research Project, Surinam, 4: 47. plateau to that of the lowlands. Mem. Carnegie, 67b. Final remarks and tentative list of fish Mus., 5: ixix, 1578, 3 pis. species. In: Progress Reports, Biological EIGENMANN, C. H. & W. L. BRAY, 4. A revision of the Brokopondo Research Project, Surinam, 4: 06. American Cichlidae. Ann. New York Acad. Sci., 7: PELLEGRIN, J., 03. Descriptions de Cichlides nouveaux de la collection du Museum. Bull. Mus. Hist. nat. FERNHOLM, B. & A. WHEELER, 83. Linnaean fish Paris, : 05. specimens in the Swedish Museum of Natural, 04. Contribution a l'etude anatomique, biologi History, Stockholm. Zool. J. Linn. Soc., 78 (3): que et taxinomique des Poissons de la famille des 286. GRONOVIUS, L. T., 54. In J. E. Gray (ed.): Catalogue of fish collected and described by Laurence Theodore Gronow, now in the British Museum (Trustees British Mus. (Nat. Hist.), London), : ivii, 16. GUNTHER, A., 62. Catalogue of the fishes in the British Cichlides. Mem. Soc. zool. France, : 41402, pis. IVVII. PLOEG, A., 86a. Occurrence and variability of Crenicichla saxatilis (Linnaeus, 58) in Surinam, and restriction of its typelocality (Pisces, Perciformes, Cichlidae). Bijdr. Dierk. 56 (1): 475. Museum. Volume fourth. Catalogue, 86b. The cichlid genus Crenicichla from the of the Acanthopterygii pharyngognathi and Anacanthini in the collection of the British Museum, : ixxi, 1534 (Trustees Brit. Mus. (Nat. Hist.), London). HECKEL, J., 40. Johan Natterer's neue Flussfische Brasilien's, nach den Beobachtungen und Mittheilungen des Entdeckers beschrieben (erste Abteilung, die Labroiden). Annln. wien. Mus. Naturg., 2: HEYDE, H., 86. Surinaamse vissen: 14 (Published by the author, Paramaribo). KULLANDER, S. O., 82. Cichlid fishes from the La Plata basin. Part III. The Crenicichla lepidota species group (Teleostei: Cichlidae). Rev. Suisse Zool., 8 (3): 6661, figs. Tocantins River, State of Para, Brazil, with descriptions four of new species (Pisces, Perciformes, Cichlidae). Beaufortia, 36 (5): 5780., 86c. The fishes ofthe cichlid Crenicichla in genus French Guiana (Pisces, Perciformes, Cichlidae). Bijdr. Dierk. 56 (2): 11. REGAN, C. T., 05. A revision of the fishes of the South American cichlid Crenicara, genera Batrachops, and Crenicichla. Proc. zool. Soc. London, 05: 28.,. A synopsis of the cichlid fishes of the genus Crenicichla. Ann. Mag. nat. Hist., (8) : RICHTER, C.J.J. & H. NIJSSEN, 80. Notes the on fishery potential and fish fauna ofthe Brokopondo Reservoir LINNAEUS, C., 54. Museum S:ae R:ae M:tis. Adolphi (Surinam). Fish Mgmt., (3): 0. SPIX, J. B. DE [VON] & [J.] L. [R ] AGASSIZ,. Friderici Regis Svecorum, Gothorum, Vandalorumqve. Haer. Norv. Due. Slesv. Hols. Storm. Dit. Selecta genera et species piscium, quos in itinere per Com. Oldenb. Delmenhorstiae. &c. &c. in quo animalia rariora imprimis, et exotica: quadrupedia, Brasiliam annis MDCCCXVIIMDCCCXX, jussu et auspiciis Maximiliani Josephi I., Bavariae Regis aves, amphibia, pisces, insecta, vermes describuntur Augustissimi, peracto collegit et pingendos curavit et determinantur, Latine et Svetice cum iconibus, : ixxx, 16,, 33 pis. (P. Momma, Holmiae). LINNAEUS, C., 58. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis, 1. Editio decima, reformata: (L. Salvii, Holmiae). LOWEMCCONNELL, R. H., 6. The cichlid fishes of Guyana, South America, with notes on their ecology Dr. J. B. de Spix, [etc.], digessit, descripsit et observationibus anatomicis illustravit Dr. L. Agassiz, praefatus est et ididit itineris socius Dr. F. C. Ph. de Martius, : [vi] iii, 16, iiixvi, 2, 48 pis. (C. Wolf, Monachii). STEINDACHNER, F.,. Ichthyologische Beitrage (XVIII). Sber. kaiserl. Akad. Wiss. Wien, math. naturw. Kl., (1) 4 (8): WHEELER, A. C., 58. The Gronovius fish collection: i and breeding behaviour. Zool. J , 3 pis. Linn. Soc., 48: catalogue and historical account. Bull. Brit. Mus nat. Hist., (hist. Ser.), 1 (5): 7, pis NIJSSEN, H., 66. Ichthyological observations: May and June 66. In: Progress Reports, Biological Brokopondo Research Project, Surinam, 3: 80. Received: October, 87. Institute of Taxonomic Zoology (Zoologisch Museum), University of Amsterdam, P.O. Box 5, 00 HC Amsterdam, The Netherlands. 8

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