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A New Species and Geographical Variation in the Telmatochromis Temporalis Complex (Teleostei, Cichlidae) from Lake Tanganyika

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A New Species and Geographical Variation in the Telmatochromis Temporalis Complex (Teleostei, Cichlidae) from Lake Tanganyika Journal of Fish Biology (2003) 63, 593–616 doi:10.1046/j.1095-8649.2003.00173.x,availableonlineathttp://www.blackwell-synergy.com A new species and geographical variation in the Telmatochromis temporalis complex (Teleostei, Cichlidae) from Lake Tanganyika M. HANSSENS* AND J. SNOEKS Africa Museum, Vertebrate Section, Cichlid Research Unit, Leuvensesteenweg, 13, B-3080 Tervuren, Belgium (Received 28 January 2002, Accepted 11 June 2003) Telmatochromis brachygnathus sp.n. is described from the southern and central parts of Lake Tanganyika. It can be distinguished from the similar T. temporalis mainly by its smaller mouth. Morphological distinct populations were found in both species. # 2003 The Fisheries Society of the British Isles Key words: Cichlidae; distribution; geographical variation; Lake Tanganyika; taxonomy; Telmatochromis brachygnathus. INTRODUCTION The systematics of the cichlids of the East African Lakes are complicated and taxonomic knowledge is poor (Coulter et al., 1986; Greenwood, 1991; Rossiter, 1995; Snoeks, 2000). Even the relatively well-known Lake Tanganyikan ichthyo- fauna turns out to be problematic in view of the results of some recent morphological and molecular studies (Verheyen et al., 1996; Stiassny, 1997; Ru¨ber et al., 1999; Hanssens et al., 1999). An ongoing study of the taxonomy and distribution patterns of the lampro- logine cichlids from Lake Tanganyika (Snoeks et al., 1994) revealed several problems within the genus Telmatochromis Boulenger. Hanssens & Snoeks (2001) addressed the confused status of the nominal species Telmatochromis burgeoni Poll and Julidochromis macrolepis Borodin and confirmed the pro- posed synonymy of Telmatochromis lestradei Poll with Telmatochromis temporalis Boulenger. In the present study, a new Telmatochromis species, morphologically similar to T. temporalis is described; both are referred to as the T. temporalis complex (Hanssens & Snoeks, 2001), because of their high degree of similarity and obvious close relationships. *Author to whom correspondence should be addressed. Tel.: þ32 2 7695628; fax: þ32 2 7695642; email: [email protected] 593 # 2003 The Fisheries Society of the British Isles 594 M. HANSSENS AND J. SNOEKS MATERIALS AND METHODS The following specimens were examined: the holotype and 36 paratypes of T. brachygnathus; the lectotype, two paralectotypes and 57 other specimens of T. temporalis (including the holotype and four paratypes of T. lestradei and the holotype of T. burgeoni, both junior synonyms of T. temporalis). A list of specimens is presented in the type list and in the Appendix. For the description and analyses, 24 morphometric and 16 meristic characters were used, all as described by Snoeks (1994), except for the upper jaw length and the number of enlarged outer row teeth in the oral jaws, which were described by Hanssens & Snoeks (2001). The measurements and meristics taken are listed in Tables I and II, respectively. Data were explored and analysed using principal component analyses (PCA) and Mann–Whitney U-tests. Some of the meristics were excluded from the statistical analyses. On some specimens part of the scales were missing, so the number of upper lateral line scales could not be determined. Also the total number of outer oral teeth in the oral jaws could not be counted on all specimens. The number of scales around the caudal peduncle is invariable and was therefore also excluded from the analyses. PCA are used here as model-free and distribution-free techniques for exploring the multivariate data set (Marcus, 1990). PCA were carried out on log-transformed measurements and raw meristics. A covariance matrix was used to calculate the factor loadings and scores of the log- transformed measurements. In all analyses of measurements, the loadings of the variables TABLE I. Measurements of the holotype and the holotype plus 36 paratypes of T. brachygnathus Holotype Mean Æ S.D. Range Standard length (LS)58Á554Á9 Æ 8Á342Á0–76Á0 Body depth % LS 29Á927Á9 Æ 1Á924Á8–32Á8 Head length (LH) % LS 30Á930Á5 Æ 0Á828Á8–32Á3 Head width (WH) % LS 44Á244Á1 Æ 1Á340Á5–46Á1 Interorbital width % LH 28Á225Á1 Æ 2Á618Á9–29Á3 Interorbital width % WH 63Á856Á9 Æ 6Á042Á9–66Á7 Snout length % LH 35Á935Á8 Æ 2Á132Á4–40Á2 Lower jaw length % LH 28Á229Á3 Æ 2Á324Á7–33Á9 Upper jaw length % LH 29Á329Á9 Æ 1Á526Á3–32Á9 Premaxillary pedicel length % LH 30Á930Á6 Æ 1Á228Á1–33Á1 Cheek depth % LH 29Á329Á9 Æ 3Á122Á6–34Á8 Eye diameter % LH 26Á026Á7 Æ 1Á424Á4–29Á1 Lachrymal depth % LH 23Á820Á7 Æ 1Á817Á1–25Á6 Lower pharyngeal jaw length (LLP) % LH 22Á722Á4 Æ 0Á920Á6–24Á3 Lower pharyngeal jaw width (WLP) % LLP 122Á0 124Á4 Æ 4Á6 113Á9–134Á5 Dentigerous area length % LLP 58Á560Á3 Æ 3Á952Á4–66Á7 Dentigerous area width (WDeA) % WLP 70Á074Á0 Æ 2Á570Á0–80Á0 Dentigerous area length % WDeA 68Á665Á6 Æ 3Á957Á1–72Á7 Dorsal fin base length % LS 65Á064Á5 Æ 1Á561Á5–67Á8 Anal fin base length % LS 25Á622Á4 Æ 1Á420Á2–25Á6 Predorsal distance % LS 29Á128Á9 Æ 1Á026Á4–31Á2 Preanal distance % LS 64Á164Á9 Æ 1Á762Á1–67Á9 Prepectoral distance % LS 29Á930Á0 Æ 1Á227Á7–33Á3 Preventral distance % LS 33Á333Á8 Æ 1Á731Á1–37Á8 Caudal peduncle length (LCP)% LS 12Á814Á1 Æ 1Á111Á8–16Á9 Caudal peduncle depth % LCP 106Á791Á1 Æ 9Á073Á3–107Á7 # 2003 The Fisheries Society of the British Isles, Journal of Fish Biology 2003, 63, 593–616 # 2003 The Fisheries Society of the British Isles, NEW TABLE II. Meristics of the holotype and the holotype plus 36 paratypes of T. brachygnathus. Frequency indicated between parentheses TELMATOCHROMIS Holotype Number and frequency Enlarged outer teeth upper jaw 16 8–18 (median 13) Enlarged outer teeth lower jaw 17 10–20 (median 15) Outer teeth upper jaw 44 29–61 (median 38) Outer teeth lower jaw 27–44 (median 38, n ¼ 10) Journal of Fish Biology Inner tooth rows upper/lower jaw 5/4 5/4 (f 2), 5/5 (f 12), 6/5 (f 2), 6/6 (f 15), 6/7 (f 1), 7/6 (f 1), 7/7 (f 4) AND GEOGRAPHIC VARIATION Gill raker 3/1/4 2/1/4 (f 1), 2/1/5 (f 1), 3/1/2 (f 4), 3/1/3 (f 10), 3/1/4 (f 8), 3/1/5 (f 3), 3/1/6 (f 1), 4/1/3 (f 2), 4/1/4 (f 2), 4/1/5 (f 2), 4/1/6 (f 1), 5/1/3 (f 1), 5/1/5 (f 1) Dorsal fin formula XXII 7 XX 7 (f 2), XXI 6 (f 1), XXI 7 (f 9), XXI 8 (f 4), XXII 6 (f 1), XXII 7 (f 17), XXII 8 (f 1), XXIII (f 1), XXIII (f 1) Anal fin formula VIII 6 VI 5 (f 1), VI 6 (f 9), VI 7 (f 2), VII 6 (f 17), 2003, VIII 5 (f 2), VIII (f 2) Pectoral fin 14 13 (f 1), 14 (f 30), 15 (f 6) 63, Upper lateral line scales 27 22–29 (median 27) 593–616 Longitudinal line scales 34 33–36 (median 35) Scales around caudal peduncle 16 16 (f 35) 595 596 M. HANSSENS AND J. SNOEKS on the first principal component (PC) were of the same sign and of a similar magnitude, indicating that this axis can be interpreted as a proxy for general size (Jolicoeur & Mosimann, 1960; Humphries et al., 1981; Bookstein et al., 1985). This was confirmed by plotting the first axis v. standard length (LS). The correlation matrix was used for the PCA on the raw meristics. Non-parametric Mann–Whitney U-tests were used for univariate comparisons; they were only performed on samples of similar length class and calculated on the relative measurements (percentages) and meristics. Abbreviations used in the text are: MRAC, Muse´e Royale de l’Afrique Centrale (Tervuren); BMNH, Natural History Museum [formerly British Museum (Natural His- tory), London]; DRC, Democratic Republic of Congo; LH, head length. Further abbre- viations for the measurements are explained in Tables I and II. RESULTS TELMATOCHROMIS BRACHYGNATHUS SP.NOV.[FIGS.1(A), 3AND4] Etymology From the Greek brawnz ‘short’ and gnayoz ‘jaw’, used as a noun in apposi- tion, referring to the relatively small mouth of the species. Diagnosis Telmatochromis brachygnathus is readily distinguished from the ‘three banded’ Telmatochromis species, T. vittatus Boulenger, T. bifrenatus Myers and T. brichardi Louisy, by the lack of dark longitudinal bands and a deeper body (body depth 24Á8–32Á8 v. 23Á5% of LS). Telmatochromis brachygnathus is differentiated from T. dhonti by a smaller head (head length 28Á8–32Á3 v. 31Á4–34Á1% of LS), smaller mouth (upper jaw length 26Á3–32Á9 v. 32Á0–42Á8% of LH; lower jaw length 24Á7–33Á9 v. 35Á8–43Á6% of LH), and by the enlarged outer row oral dentition [enlarged flattened, straight tipped anterior oral teeth v. rounded teeth with recurved tips, Fig. 1(a), (c)]. Telmatochromis brachygnathus is most similar to T. temporalis from which it is differentiated by a smaller mouth (upper jaw length 26Á3–32Á9 v. 29Á8–40Á5% of LH; lower jaw length 24Á7–33Á9% of 31Á8–41Á6% of LH) [Fig. 1(a), (b) and Fig. 2]. Holotype MRAC 78–25-P-720; Cap Chaitika, Zambia; P. Brichard; January 1978; male of 58Á5mm LS. Paratypes Not all paratypes were measured, the number of specimens measured is mentioned between parentheses, when relevant. MRAC 331, Albertville, DRC, Capt. Hecq, 1899; MRAC 112746-747 (1), Baie de Kabimba, rochers, DRC, Explor. Hydrobiol. L. Tang., 10 November 1946; MRAC 113118-119 (1), Kabimba, Nord du lac Tanganyika, DRC, G. Leleup; 18 July 1961; MRAC 112755-759 (2), Stat. 108, M’Toto dans la baie et parmi les rochers au sud de l’entre´e, DRC, Explor. Hydrobiol. L. Tang, 5 February 1947; MRAC 122760-761 (1), Stat. 166, Baie de Vua, plage du fond, # 2003 The Fisheries Society of the British Isles, Journal of Fish Biology 2003, 63, 593–616 NEW TELMATOCHROMIS AND GEOGRAPHIC VARIATION 597 (a) fl 1 mm 1 cm (b) fl 1 mm 1 cm (c) fl 1 cm 1 mm FIG.
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