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New Species of Petrotilapia (Teleostei: Cichlidae) from Lake Malawui,Africa

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New Species of Petrotilapia (Teleostei: Cichlidae) from Lake Malawui,Africa KENDALL AND MEARNS-NOVUMBRA DEVELOPMENT 695 their characters, p. 11-22. In: Ontogeny and sys- ROSEN,D. E. 1974. Phylogeny and zoogeography of tematicsof fishes. ASIH, Special Publ. No. 1, Allen salmoniform fishes and relationships of Lepidoga- Press, Lawrence, KS. laxiassalamandroides. Bull. Am. Mus. Nat. Hist. 153: KNIGHT,A. E. 1963. The embryonic and larval de- 265-326. velopment of the rainbow trout. Trans. Am. Fish. RYDER,J. A. 1886. The development of the mud- Soc. 92:344-355. minnow. Am. Nat. 20:823-824. MARTIN, F. D. 1984. Esocoidei: development and SCHULTZ, H.-P., AND G. ARRATIA. 1989. The com- relationships, p. 140-142. In: Ontogeny and sys- position of the caudal skeleton of teleosts (Actin- tematicsof fishes. ASIH, Special Publ. No. 1, Allen opterygii:Osteichthyes). Zool.J. Linn. Soc. 97:189- Press, Lawrence, KS. 231. MATARESE,A. C., ANDE. M. SANDKNOP.1984. Iden- SOIN,S. G. 1980. Types of development of salmon- tification of fish eggs, p. 27-31. In: Ontogeny and iform fishesand their taxonomic importance.J. Ich- systematics of fishes. ASIH, Special Publ. No. 1, thyol. (Engl. Transl. Vopr. Ikhtiol.) 20:49-56. Allen Press, Lawrence, KS. WILSON, M. V. H., AND P. VEILLEUX.1982. Com- , A. W. KENDALLJR., D. M. BLOOD,AND B. M. parative osteology and relationshipsof the Umbri- VINTER. 1989. Laboratoryguide to early life stages dae (Pisces:Salmoniformes). Zool. J. Linn. Soc. 76: of northeastPacific fishes. US Dept. Comm. NOAA 321-352. Tech. Rep. NMFS 80. MCPHAIL,J. D. 1969. Predation and evolution of a (AWK) ALASKA FISHERIES SCIENCE CENTER, NA- stickleback(Gasterosteus). J. Fish. Res. Board Can- TIONAL MARINE FISHERIES SERVICE, NATIONAL ada 26:3183-3208. OCEANIC AND ATMOSPHERIC ADMINISTRA- MELDRIM,W. 1968. The J. ecological zoogeography TION, 7600 SAND POINT WAY NE, BIN of the Olympic mudminnow (Novumbrahubbsi, C 15700, SEATTLE, WASHINGTON98115-0070; Schultz).Unpubl. Ph.D. diss., Univ. of Washington, Seattle. AND (AJM) NATIONAL OCEAN SERVICE, OFFICE OF OCEAN CONSERVATION RESOURCES AND AS- NELSON, J. S. 1994. Fishes of the world. 3d ed. John Wiley and Sons, New York. SESSMENT,HAZARDOUS MATERIALS RESPONSE PECKHAM,R. S., ANDC. F. DINEEN. 1957. Ecology of AND ASSESSMENT DIVISION, NATIONAL OCEA- the central mudminnow,Umbra limi (Kirtland). Am. NIC AND ATMOSPHERICADMINISTRATION, 7600 Midi. Nat. 58:222-231. SANDPOINT WAY NE, BIN C 15700, SEATTLE, PENAZ,M. 1975. Earlydevelopment of the grayling WASHINGTON 98115-0070. Submitted: 5 Thymallusthymallus (Linnaeus 1758). Acta. Sci. Nat. March 1995. 31 Oct. 1995. Section Acad. Sci. Bohemoslov. Brno. 9:1-35. Accepted: editor: J. R. Gold. POTTHOFF,T. 1984. Clearing and staining tech- niques, p. 35-37. In: Ontogeny and systematicsof fishes. ASIH Special Publ. No. 1, Allen Press, Law- rence, KS. Cope/a, 1996(3), pp. 695-702 New Species of Petrotilapia (Teleostei: Cichlidae) from Lake Malawui,Africa JAY R. STAUFFER JR. AND ELLEN S. VAN SNIK A rock-dwelling cichlid endemic to Chinyamwezi and Chinyankwazi islands, Lake Malawi, Malawri, Africa is described. The new species is placed in the genus Petrotilapia based on the presence of predominantly tricuspid teeth on the major dentigerous areas of the jaws and because the teeth are exposed when the jaws are closed. It is distinguished from congeners by the gold coloration of nonter- ritorial males and females. Cheek depth, as expressed as percent of head length, is typically much smaller in the new species than in previously described mem- bers of the genus. THE haplochromine cichlid fishes inhabit- genetic similarity of the African haplochrom- ing lakes Victoria, Tanganyika, and Ma- ines makes it extremely difficult to delimit both laCwirepresent an outstanding case of explosive allopatric and sympatric species. Genetic studies evolution in modern vertebrates. The high di- of sympatric color morphs of the Lake Malawi versity, recent speciation, and mophological and cichlid fishes Pseudotropheus zebra (Boulenger) ? 1996 by the American Society of Ichthyologists and Herpetologists 696 COPEIA, 1996, NO. 3 and Petrotilapia tridentiger Trewavas found sig- (1991) and were taken from the left side of the nificant differences in allelic frequencies in sev- body, except for gillraker meristics, which were eral of the polymorphic loci but failed to dis- taken from the right side. The number of scales tinguish any fixed differences (McKaye et al., in the overlapping portion of the upper and 1982; 1984). Similarly, the morphological char- lower lateral lines was not counted; pored scales acters of many sympatric and allopatric forms posterior to the hypural plate were recorded overlap (Marsh, 1983). Despite the lack of fixed separately. The posterior simple ray of the anal genetic and morphological differentiation, spe- fin was not counted (Stauffer, 1994). Standard cific status of many forms has been confirmed length (SL) is used throughout, and morpho- by observation of assortative mating (Marsh, metric values are expressed as percent SL for 1983). Lewis (1982) suggested that, in the case measurements which extend past the opercu- of shallow water species, behavioral characters lum of the fish and percent head length (HL) should be used as taxonomic characters. Stauf- for measurements which do not extend past the fer et al. (1993) interpreted bower shape as a operculum. Institutional abbreviations follow manifestation of a behavioral character and used Leviton et al. (1985), except where noted. it to delimit three Copadichromisspecies from The new species was compared with the type Lake Malawi. Among rock-dwelling haplo- series of P. tridentiger (BMNH 1935.6.14.244- chromine species, coloration has been shown to 246), P. genalutea (BMNH 1981.2.2.222-224), be important in mate recognition and selection and P. nigra (BMNH 1981.2.2.207-209), all within a species (Fryer, 1959; Schroder, 1980; from Monkey Bay. In addition, specimens of P. Stauffer and Boltz, 1989), and in many cases tridentiger were collected from Songwe Hill different male color patterns are the primary (34056'E, 14?00'S; n = 11, PSU 3021), Mazinzi trait used to delimit species (Marsh, 1983). Reef(34?58'E, 14?07'S; n = 2, PSU 3019, 3020), Trewavas (1935) diagnosed the genus Petro- and Crocodile Rocks (35?10'E, 14?17'S; n = 9, tilapia when she described the type species, P. PSU 3018). tridentiger, as having all tricuspid jaw teeth. Meristic differences were analyzed by prin- Marsh (1983:2) described Petrotilapia genalutea cipal component analysis (PCA) in which the Marsh and Petrotilapia nigra Marsh and modi- correlation matrix was factored (SAS 6.07). Body fied the diagnosis "to include those Lake Ma- shape was assessed by analyzing the morpho- lawi cichlids with predominantly tricuspid teeth metric data with a sheared PCA (Humphries et on the major dentigerous area of the jaws ex- al., 1981; Bookstein et al., 1985), in which the cluding the posterior sides of the premaxilla and covariance matrix was factored (SAS 6.07). This dentary." The other character deliminating the procedure restricts size variation to the first genus is the visibility of teeth when the jaws are principal component; thus, subsequent com- closed. ponents are strictly shape related. Comparisons Certainly, as it is now diagnosed, Petrotilapia among species were made by plotting the consists of a complex of sibling species (Marsh sheared second principal component of the et al., 1981). The type material for all of the morphometric data with the first principal com- described Petrotilapia species is from Monkey ponent of the meristic data. A multivariate anal- Bay. Marsh (1983) described the three distinct ysis of variance (MANOVA) was used to test sympatric color forms as separate species based differences among the minimum polygon clus- on behavioral observations that demonstrated ters formed by each species in the above plot. assortative mating of the three forms. His de- cision was the of et supported by study McKaye chrysosn. al. which found differences Petrotilapia sp. (1982), frequency Ribbink et al., 1983:211 in seven loci the three Petrotilapia "gold" polymorphic among sym- Figures 1-2; Table 1 patric color forms. The purpose of this paper is to describe a new allopatric species of Petro- Holotype.-PSU 2726, adult male, 116.9 mm tilapia that is endemic to Chinyankwazi (35?00'E, SL, Chinyamwezi Island, Lake Malawi, Africa, 13?53'S) and Chinyamwezi (35?00'E, 13?56'S) 1-3 m, 13 March 1991. islands. These two islands are known for their high degree of endemism (Stauffer, 1988, 1993; Paratypes.-PSU 2727, 16 (84.9-131.3 mm), Stauffer et al., 1995). data as for holotype, PSU 2728, 3 (106.5-117.4 mm), Chinyamwezi Island, Lake Malawi, Afri- 1-3 28 Feb. PSU 4 MATERIALS AND METHODS ca, m, 1988; 2729, (94.2- 112.2 mm), Chinyamwezi Island, Lake Malawi, Fishes were collected by chasing them into a Africa, 1-3 m, 4 May 1991;PSU 2730,8 (83.7- monofilament gill net while SCUBA diving. Ex- 116.3 mm), Chinyamwezi Island, Lake Mala,wi, ternal counts and measurements follow Stauffer Africa, 1-3 m, 18June 1991; PSU 2731,5 (88.4- STAUFFER AND VAN SNIK-NEW PETROTILAPIA 697 TABLE 1. MORPHOMETRICAND MERISTICVALUES OF THE TYPE SERIESOF Petrotilapia chrysos(n = 78). Mean, standarddeviation (SD), and range include the holotype. Holotype Mean SD Range Standardlength, mm 116.9 105.9 10.8 79.0-131.3 Head length, mm 36.5 33.6 4.3 22.7-42.1 Percent of standard length Head length 31.2 31.7 1.6 28-35 Snout to dorsal-finorigin 36.6 36.9 1.9 31-41 Snout to pelvic-finorigin 36.3 35.7 2.1 31-40 Pectoral-finlength 32.7 29.0 4.3 18-39 Pelvic-fin length
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