(Teleostei: Cichlidae) from Lake Malawi, Africa
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DUELLMAN AND OCHOA-NEW PERUVIAN BUFO 141 Ministerio de Agricultura, Lima, Peru. The tion, and dispersal. W. E. Duellman (ed.). Monogr. drawings were executed by A. M. Musser, and Mus. Nat. Hist., Univ. Kansas 7:1-485. D. 1975. A revision of the Andean the photograph in Figure 4 was taken by L. LYNCH, J. lep- Trueb. todactylid frog genus Phrynopus. Occas. Pap. Mus. Nat. Hist., Univ. Kansas 35:1-51. STEBBINS,R. C. 1954. Amphibians and reptiles of LITERATURE CITED western North America. McGraw-Hill Book Co., Inc., New York, New York. CEI,J.M. 1977. Serologicalrelationships of the Pata- Tosi, J. A., JR. 1960. Zonas de vida natural en el gonian toad Bufovariegatus (Gunther). Serol. Mus. Peru. Bol. Tecn. Inst. Interamer. Cien. Agric. 5: Bull. 52:2. 1-271. DUELLMAN,W. E. 1987. Two new species of mar- TROLL, K. 1959. Die tropischen Giberge. Bonn. supial frogs (Anura: Hylidae) from Peru. Copeia Geogr. Abhandl. 25:1-93. 1987:903-909. - ANDT. H. FRITTS. 1972. A taxonomic review , (WED) MUSEUM OF NATURAL HISTORY AND DE- of the southern Andean Occas. marsupial frogs. PARTMENT OF SYSTEMATICS AND ECOLOGY, Mus. Nat. Hist., Univ. Kansas 9:1-37. Pap. THE UNIVERSITY OF - ANDL. TRUEB. 1988. of KANSAS, LAWRENCE, , Cryptic species hylid KANSAS66045 AND DEPARTAMENTO marsupial frogs in Peru. J. Herpetol. 22:159-179. (OOM) DE UNIVERSIDAD NACIONAL DE SAN FORMAS,J. R. 1979. La herpetofauna de los bosques BIOLOGIA, temperados de Sudambrica, p. 341-369. In: The ANTONIO ABAD DEL CUZCO, APARTADO 921, South American herpetofauna: its origin, evolu- Cuzco, PERU.Accepted 9 April 1990. Copeia, 1991(1), pp. 141-147 Description of a Facultative Cleanerfish (Teleostei: Cichlidae) from Lake Malawi, Africa J.R. STAUFFER,JR. A new species of the cichlid fish genus Pseudotropheusfrom Lake Malawi, Africa, is described. The new species superficially resembles Pseudotropheus lanisticola and P. livingstonii, but is clearly distinguished by head shape. The behavior pattern of cleaning ectoparasites from other cichlids is unique among all known species of Pseudotropheus. HE cichlid fishes of Lake Malawi a occupy METHODS AND MATERIALS wide variety of habitats and exhibit di- verse behavioral adaptations. Ribbink et al. Fishes were collected by chasing them into a (1983) published a survey of the rock-dwelling monofilament net (7 m x 1 m x 1.5 cm) while cichlids ("mbuna"), documented the distribu- SCUBA diving. Standard length is used tion of many of the mbuna, and commented on throughout. External counts and measure- their behavioral adaptations. Mbuna primarily ments follow Barel et al. (1977), except that the inhabit the rocky shores and rock outcroppings following distances were recorded: anterior in- of Lake Malawi; however, some representatives sertion of dorsal fin to anterior insertion of anal of this occur over sand group substrate or over fin, posterior insertion of dorsal fin to posterior the rock/sand interface. These species are Pseu- insertion of anal fin, anterior insertion of dorsal P. dotropheuselegans Trewavas, livingstonii (Bou- fin to posterior insertion of anal fin, posterior lenger), and P. lanisticola Burgess. The purpose insertion of dorsal fin to anterior insertion of of this paper is to describe a new species that anal fin, posterior insertion of dorsal fin to ven- superficially resembles P. lanisticola and P. liv- tral origin of caudal fin, posterior insertion of ingstonii, and inhabits the rock/sand interface. anal fin to dorsal origin of caudal fin, anterior 1991 the American ? by Society of Ichthyologists and Herpetologists 142 COPEIA, 1991, NO. 1 1 cm Fig. 1. Pseudotropheuspursus, holotype, USNM 307339, adult male, 60.6 mm. insertion of dorsal fin to pelvic-fin origin, and Paratypes.-USNM 307340, 12 (46.9-60.1 mm), posterior insertion of dorsal fin to pelvic-fin or- BMNH 1990.2.14:2-6, 5 (42.1-57.1 mm), MFU igin. These measurements were recorded so that 1 (Malawi Fisheries Unit, Monkey Bay, Malawi, a truss network could be completed, as de- Africa), 3 (43.7-48.2 mm); data as for holotype. scribed by Humphries et al. (1981). Except for MFU 2, 3 (31.9-49.6 mm); Songwe Hill, Lake gill-raker counts, which were recorded from the Malawi, Africa. right side, all counts and measurements were made on the left side of the fish. Vertebral counts cichlid of the Pseudotro- were made from Diagnosis.-A genus radiographs. that resembles P. The new was with the ho- pheus superficially lanisticola species compared and P. but differs in of P. and livingstonii shape (Fig. 2). lotype livingstonii(BMNH 63.11.12.22) has a smaller the and two Pseudotropheus pursus generally holotype (USNM 216266) para- cheek and more scale rows on the cheek of P. lanisticola. depth types (BMNH 1976.7.29:1-2) than either of the aforementioned Differences in were species. body shape analyzed using it differs from these two sheared Moreover, behaviorally principal components analysis (PCA) in that it is not associated with snail shells et al., 1981; Bookstein et al., species (Humphries 1985). as are P. and P. This which differ- (Lanistes nyassanus) lanisticola technique, quantifies shape Ribbink et ences the of livingstonii (Burgess, 1976; al., 1983). among populations independent It is distinct from all known in size et was used Stauffer Pseudotropheus (Reyment al., 1984), by color and in that it is a facultative clean- and Boltz to between two pattern (1989) distinguish er. sympatric species from Lake Malawi: P. barlowi McKaye and Stauffer and P. xanstomachusStauf- fer and Boltz. All specimens were collected at Description.-Morphometric ratios and meris- either Songwe Hill (34056'E, 14?00'S) or Kan- tics are presented in Table 1. Jaws isognathous chedza Is. (34056'E, 13'58'S), Lake Malawi, Ma- (Fig. 1); teeth on jaws in three rows (premaxilla lawi, Africa. Institutional abbreviations follow of one paratype had two rows); most teeth in Leviton et al. (1985), except where noted. outer rows bicuspid, those in inner rows tricus- pid; 15 teeth in outer row of left lower jaw of 12-17 in Pectoral fins with Pseudotropheuspursus n. sp. holotype, paratypes. 13 in and 12 14 Figs. 1-4 rays holotype paratypes, rays in 11 paratypes; anal fin with three spines and Holotype.-USNM 307339, adult male, 60.6 mm, seven rays in holotype and 21 paratypes, three Kanchedza Is., Lake Malawi, Africa, 2-3 m, 18 spines and eight rays in remaining paratypes; April 1989. caudal fin emarginate (Fig. 1). Vertebrae of ho- STAUFFER-NEW MALAWI CLEANING CICHLID 143 0.3 0.2 ., 0.1 0.0- -0.1 - -0.2 -0.3 -0.2 -0.1 0.0 0.1 0.2 SHRD PC2 Fig. 2. Plot of individual principal component scores of Pseudotropheuspursus (squares), P. lanisticola (triangles),and P. livingstonii(circle) on the sheared lo cm second and third principalcomponents. Fig. 3. Lower pharyngealbone of Pseudotropheus pursus, holotype, USNM 307339, adult male, 60.6 mm. lotype 13 + 16 (abdominal + caudal), seven paratypes with 13-14 + 15-16. Lower pharyn- geal bone of holotype triangular in outline (Fig. blue stripe; preorbital blue; gular white with a 3); pharyngeal teeth in left posterior row 20, yellow blotch on anterior portion; pectoral and those in left median row 14. Scales along lateral pelvic fins with clear membranes and yellow rays; side ctenoid; holotype with 29 pored lateral-line anal fin clear with two orange ocelli; caudal-fin scales, paratypes with 31-32; pored scales pos- rays and some membranes yellow, other mem- terior to hypural plate 0--3 (Table 1). Seventeen branes blue; dorsal fin clear with yellow lappets. specimens, including holotype, with six scale Females similarly colored but not as intensely; rows on cheek, remaining paratypes with five. ocelli absent on anal fin. Gill rakers simple, first gill arch with 8-11 on ceratobranchial, 2-3 on epibranchial, one be- Etymology.-The name pursus, from the Latin tween epibranchial and ceratobranchial. meaning clean, was chosen to reflect the clean- Sheared PCA of the three species demon- ing behavior of this species. strated that there was no overlap among species when the sheared second principal component Distribution.-Pseudotropheus pursus appears to was plotted against the sheared third principal be restricted to the eastern side of the Nankum- component (Fig. 2). The first principal com- ba Peninsula in Lake Malawi. Numerous col- ponent is interpreted as a size component and lections and SCUBA observations by Ribbink the sheared components as shape, independent et al. (1983), McKaye (pers. comm.), and myself of size (Humphries et al., 1981; Bookstein et throughout other portions of the lake did not al., 1985). After size, which accounts for 85.7% record this species. of the total variance, is removed, the second and third principal components account for 36% Life history.-Pseudotropheus pursus is a faculta- and 23% of the remaining variance. Those vari- tive cleaner. It has been observed to feed on ables that have the highest loadings on the periphyton and to clean parasites from Lethri- sheared second principal component are cheek nops cf. lituris (Cichlidae; Fig. 4). Additionally, depth, snout length, and caudal peduncle length, it has been observed to clean parasites from while those that have the highest loadings on Cyrtocara cf. fenestrata and Cyrtocara cf. picta the sheared third principal component are pel- (Cichlidae). The cleaning of the latter two spe- vic-fin length, caudal-peduncle length, and pre- cies has been recorded on underwater video orbital depth (Table 2). tapes cataloged at The Pennsylvania State Uni- Lateral body coloration of males grey dor- versity Fish Museum (PSU 2273). The host fish sally, white ventrally with 4-6 grey bars; scales approaches P. pursus and positions itself with its in outlined blue; head grey with one interorbital dorsal fin oriented toward the cleaner, and per- 144 COPEIA, 1991, NO.