Neolamprologus cancellatus, a new cichlid fish from Lake Tanganyika, Africa Mitsuto Aibara1*, Tetsumi Takahashi2, and Kazuhiro Nakaya1 1
Marine Biology and Biodiversity (Systematic Ichthyology), Division of Marine Bioresource and Environmental Science, Graduate School of Fisheries Sciences, Hokkaido University, 3-1-1 Minato-cho, Hakodate, Hokkaido 041-8611, Japan (e-mail: MA,
[email protected]; KN,
[email protected]) 2 Laboratory of Animal Ecology, Department of Zoology, Graduate School of Science, Kyoto University, Kitashirakawa-Oiwakecho, Sakyo, Kyoto 606-8502, Japan (e-mail:
[email protected]) Received: July 21, 2004 / Revised: May 25, 2005 / Accepted: May 27, 2005
Ichthyological Research ©The Ichthyological Society of Japan 2005
Ichthyol Res (2005) 52: 354–359 DOI 10.1007/s10228-005-0296-0
Abstract Neolamprologus cancellatus, a new cichlid species, is described on the basis of eight specimens from the Zambian coast of Lake Tanganyika. The new species is characterized by a subtruncate caudal fin and a body that is slender (depth 22.3–25.2% in standard length) and easily distinguishable from its congeners by having 7–8 anal fin spines, 34–37 scales in longitudinal line, 33 total vertebrae, and a gridlike body pattern on a pale brownish body color. This species is only known to inhabit the shallow (2–7 m depth), rocky bottom of Wonzye Point in the southern part of the lake. Key words Neolamprologus cancellatus · New species · Cichlidae · Lake Tanganyika · Africa
L
ake Tanganyika, one of the Great Rift Valley lakes in central east Africa, is famous for high levels of endemism among cichlid fishes and some other animal groups (e.g., Copepoda, Decapoda, Gastropoda, Ostracoda) (Coulter, 1991; Fryer, 1991). The cichlid genus Neolamprologus Colombe and Allgayer, 1985 is the largest genus among Lake Tanganyikan cichlid fishes, currently comprising 47 valid species (Büscher, 1997; Stiassny, 1997; Schelly et al., 2003). Even recently, many new species from the lake have been discovered and described (e.g., Büscher, 1991b, 1992a,b, 1993, 1995a,b, 1997; Nakaya and Gashagaza, 1995; Schelly et al., 2003). Hence further taxonomic research is absolutely necessary for a better understanding of the species diversity of the lake. During a scientific survey along the Zambian coast of Lake Tanganyika in 2000, specimens of a previously unidentified species of Neolamprologus were collected from the shallow waters of the southern part of the lake. They are described herein as a new species.
Materials and Methods Specimens were collected at Wonzye Point (8°43¢29≤ S, 31°07¢59≤ E), by gill net, at the rocky bottom in depths of 2– 7 m (Fig. 1). The type specimens were fixed in 10% formalin and thereafter maintained in 50% isopropanol. Counts and proportional measurements mainly followed Barel et al. (1977). The lengths of the caudal peduncle, predorsal, dorsal fin base, anal fin base, and upper jaw are those of Hubbs and Lagler (1958). The two posteriormost dorsal- and anal fin soft rays were counted as two, rather than as a single branched ray. Counts of upper and lower
lateral line scales were those from the anteriormost scale with a canal to the posteriormost one, including the scales without a canal. The term “scales in longitudinal line” was used instead of “scales in lateral line” (sensu Barel et al., 1977). Scales with a canal, which are located behind the posterior margin of the hypural plate, were not counted as scales in longitudinal line and lower lateral line. The uppermost pectoral fin ray, which is short, unbranched, and not segmented, was counted. The innermost and outermost soft rays of pelvic fin were measured from its base to tip. All measurements were made with dividers and rounded to the nearest 0.1 mm. Vertebral numbers were determined from radiographs. Sex was determined by gonad examination. Fresh color notes were based on photographs. Institutional abbreviations are as listed in Leviton et al. (1985).
Neolamprologus cancellatus sp. nov. (Fig. 2, Tables 1, 2) Holotype. HUMZ 174278, male, 44.1 mm in standard length (SL), Wonzye Point, Zambia, 2 m depth, gill net, rocky shore, 11 Dec. 2000, collected by M. Aibara. Paratypes. Seven specimens (5 males, 2 females). HUMZ 174243, male, 38.4 mm SL, Wonzye Point, 2 m depth, gill net, rocky shore, 12 Dec. 2000, collected by M. Aibara; HUMZ 173921, female, 33.6 mm SL, MRAC 2004.41.P.1, female, 46.4 mm SL, MRAC 2004.41.P.2, male, 46.6 mm SL, MRAC 2004.41.P.3, male, 41.0 mm SL, Wonzye Point, 2 m depth, gill net, rocky shore, 18 Dec. 2000, collected by M. Aibara and T. Takahashi; HUMZ 174069, male, 54.8 mm SL, Wonzye Point, 7 m depth, gill net, rocky shore, 4 Dec. 2000, collected by M. Aibara; HUMZ 174070, male, 57.1 mm SL, Wonzye Point, 7 m depth, gill net, rocky shore, 4 Dec. 2000, cleared and stained for observation of pectoral fin and labial ligament.
A new cichlid from Lake Tanganyika
Fig. 1. Lake Tanganyika, showing type locality of Neolamprologus cancellatus sp. nov.
Diagnosis. A new species of Neolamprologus with light brownish body color with gridlike stripes on lateral side of body, slender body shape (depth 22.3–25.2% SL), subtruncate caudal fin, 19–20 dorsal fin spines, 7–8 anal fin spines, 34–37 scales in longitudinal line, 5–7 gill rakers on lower limb of 1st gill arch, and 33 total vertebrae. Description. Counts and measurements are shown in Table 1. The holotype is described below, and data of the seven paratypes are given in parentheses only when different from those of the holotype. Body slender and cylindrical; depth less than head length. Dorsal profile of head moderately curved. Snout isognathous; length a little larger than eye diameter. Interorbital region broadly rounded. Mouth terminal; posterior margin of premaxilla almost reaching to level of anterior margin of eye. Suborbital narrow, width about half of eye diameter. Dorsal fin larger posteriorly; 1st dorsal spine shortest; spine length progressively increasing posteriorly to 7th
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(7th–9th) spine; soft rays all branching; longest ray almost reaching to caudal fin base when depressed (extending the base in larger paratypes). Anal fin spines longer posteriorly; soft rays all branching except for last one; longest ray reaching to the caudal fin base. Pectoral fin reaching to the level of 11th dorsal spine (10th–12th); uppermost ray short and unsegmented; uppermost two and lowermost two rays unbranched and others branched. First pelvic fin ray spinous, soft rays all branched; 1st soft ray longest, reaching between anus and origin of anal fin. Caudal fin subtruncate. Eight (6– 11) canine teeth present anteriorly on premaxilla and 8 (6– 9) on dentary; conical teeth on these bones in many rows, outer teeth larger, inner ones smaller. Scales on body weakly ctenoid; those between upper lateral line and posterior part of spinous dorsal fin base smaller and rather irregular in distribution. Opercle with a few scales. Cheek and dorsal side of head naked. Dorsal and anal fins not scaled. Base of anterior part of spinous dorsal fin with small scales immersed in epidermis. Thoracic region naked. Caudal fin scaly from base to its middle part, naked distally. Gill rakers on lower limb clavate and slender, those of the anterior part a little reduced. Labial cartilage on the ligament not ossified. Coloration in fresh specimens.—Dorsal part of head and body pale brownish. Ventral part of head pale yellowish (pale in alcohol). Ventral part of body whitish. Two brown lateral bands on body; dorsal medial one paler, running from dorsal side of head to caudal fin along dorsal fin base; midlateral one denser, from upper lip to caudal fin base via eye. Two brown blotches on head; anterior one on interorbital region and posterior one on nape. Nine obscure vertical stripes on body; 1st to 5th stripes below spinous dorsal fin, 6th and 7th below soft-ray part of dorsal fin, and 8th and 9th on caudal peduncle; these stripes extending from dorsal side of body to midlateral band. Crossing areas of bands and stripes much darker. Dorsal fin light yellowish (pale in alcohol), with seven brown markings. Anal fin light yellowish (pale in alcohol). Lappets of dorsal and anal fin membrane light yellowish (pale in alcohol). Small pale-brown maculae absent on caudal fin membrane (present in alcohol). Distribution. Known only from Wonzye Point, Zambia (Fig. 1), shallower than 7 m depth. Etymology. The species name cancellatus is derived from Latin adjective for gridlike, in reference to its gridlike color pattern on body. Remarks. The new species belongs to the tribe Lamprologini (sensu Poll 1986), in having more than three anal fin spines and conical teeth on premaxilla and dentary. This species was considered to belong to the genera Lamprologus and Neolamprologus at first, because it has following characteristics: all teeth conical, scales less than 50 in longitudinal line, a genital lappet that is not enlarged in males, and dorsal spines fewer than 22. According to Poll (1986), Neolamprologus differs from Lamprologus by the presence of scales on head, operculum, thoracic region, and abdomen (extremely reduced or absent in Lamprologus, with exception of L. meleagris Büscher, 1991 and L. stappersi Pellegrin, 1927), and a pelvic fin with the 1st soft ray being longest (the 2nd or 3rd ray longest in
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Fig. 2. Neolamprologus cancellatus sp. nov., holotype, HUMZ 174278, 44.1 mm SL, collected by gill net in Wonzye Point, southern part of Lake Tanganyika, Zambia
Lamprologus), indicating this species can be assigned to Lamprologus on scales, while to Neolamprologus on pelvic fin. We tentatively assign the new species to Neolamprologus because the character of the pelvic fin is more clear and reliable in recognition of the genus (e.g., Poll, 1986; Maréchal and Poll, 1991a, b), while the presence of scales is quite varied in Lamprologus, even with some exceptions. According to Stiassny (1997), the genus Lamprologus cannot be diagnostically separated from the genus Neolamprologus at present. Because the taxonomy of the two genera is not stable, we compare Neolamprologus cancellatus sp. nov. with all species of both Lamprologus and Neolamprologus. All the Lamprologus species are clearly distinguished from N. cancellatus by the characters mentioned in the diagnosis. Within the genus Neolamprologus, N. cancellatus is characterized by a subtruncate caudal fin and a slender body (depth 22.3–25.2% SL). Among 47 valid species, because the caudal fin of 15 species shows elongated, this species is clearly distinguishable from them. Within the remaining 32 species with nonelongated caudal fin (i.e., rounded to truncate caudal fin), 21 species are distinguished from N. cancellatus by their deep bodies (depth consistently more than 26% SL, whereas it is 22.3–25.2% SL in N. cancellatus). The new species and the remaining 11 species below have slender body (depth usually less than
26% SL); Neolamprologus cylindricus Staeck and Seegers, 1986, Neolamprologus devosi Schelly, Stiassny, and Seegers, 2003, Neolamprologus fasciatus (Boulenger, 1898), Neolamprologus hecqui (Boulenger, 1899) [body depth more than 26% SL in Poll (1978), but those of comparative specimens in this study were less than 26% SL], Neolamprologus longior (Staeck, 1980), Neolamprologus meeli (Poll, 1948), Neolamprologus nigriventris Büscher, 1992b, Neolamprologus pectoralis Büscher, 1991, Neolamprologus pleuromaculatus (Trewavas and Poll, 1952), Neolamprologus schreyeni (Poll, 1974), and Neolamprologus tetracanthus (Boulenger, 1899). Neolamprologus cancellatus can be clearly distinguished from these 11 species by the meristic characters (Table 2): 34–37 scales in longitudinal line from N. hecqui (45–55 scales), N. meeli (42–46), and N. pleuromaculatus (47–64); 7– 8 anal fin spines from N. cylindricus (5 spines), N. devosi (5– 6), N. fasciatus (9–11), N. longior (5–6), N. nigriventris (5–6), N. schreyeni (5), and N. tetracanthus (4); 33 total vertebrae from N. pectoralis (34–35 vertebrae). Additionally, this new species pale brownish or whitish body coloration clearly distinguishes it from N. cylindricus, N. pectoralis, and N. schreyeni (blackish body color), N. devosi (light beige brownish body), N. longior (yellowish body), N. nigriventris (blackish abdomen and whitish dorsal body), and N. tetracanthus (grayish body); and the presence of brown
A new cichlid from Lake Tanganyika
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Table 1. Counts and measurements of Neolamprologus cancellatus sp. nov. Holotype HUMZ 174278
Paratypes n=7
Sex Standard length (SL, in mm) Total length
Male 44.1 51.9
Male (5); female (2) 33.6–57.1 39.9–68.2
Counts Dorsal fin rays
XX, 9
Anal fin rays
VIII, 7
Pectoral fin rays Pelvic fin rays Caudal fin rays Scales in longitudinal line Scales in upper lateral line Scales in lower lateral line Gill rakers Canine teeth (premaxilla) (dentary) Vertebrae (abdominal) (caudal) (total)
14 I, 5 16 37 22 8 5
XIX–XX, 8–10 XIX, 8 [1]; XIX, 10 [1]; XX, 8 [1]; XX, 9 [4] VII–VIII, 6–7 VII, 7 [3]; VIII, 6 [3]; VIII, 7 [1] 14 I, 5 16 34–37 23–29 6–8 5–7
Measurements (% SL) Body depth Head length Snout length Eye diameter Interorbital width Upper jaw length Lower jaw length Caudal peduncle length Caudal peduncle depth Predorsal length Dorsal fin base length Anal fin base length Longest dorsal spine length Longest dorsal soft ray length Longest anal spine length Longest anal soft ray length Pectoral fin length Pelvic fin length (inner) (outer) Caudal fin length
8 8
6–11 6–9
(mean ± SD)
(35.6 ± 1.0) (25.6 ± 2.0) (7.5 ± 0.8) (6.6 ± 0.8) (8.0 ± 1.6) (7.3 ± 1.1)
15 18 33
15 18 33
24.3 31.7 9.8 9.5 5.7 10.0 12.5 16.6 12.0 31.3 60.1 24.5 15.4 15.6 16.6 20.0 17.9
22.3–25.2 29.6–32.4 9.6–10.7 8.3–9.5 5.6–6.5 9.1–10.3 11.7–13.0 15.8–18.8 10.7–12.7 29.2–32.4 55.4–60.8 23.2–26.1 14.4–17.0 15.4–20.4 14.3–17.7 17.3–28.6 17.1–19.3
(23.9 ± 1.2) (31.1 ± 1.1) (10.1 ± 0.4) (8.9 ± 0.5) (6.0 ± 0.3) (9.7 ± 0.4) (12.2 ± 0.5) (17.4 ± 1.1) (11.6 ± 0.8) (31.1 ± 1.3) (58.2 ± 2.1) (24.4 ± 1.0) (15.2 ± 0.9) (18.5 ± 2.0) (15.9 ± 1.2) (21.6 ± 4.1) (18.1 ± 0.7)
12.5 22.7 20.4
10.2–13.5 20.3–27.0 19.5–22.8
(12.2 ± 1.2) (24.4 ± 2.2) (21.2 ± 1.2)
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Table 2. Comparison of meristic characters in Neolamprologus cancellatus sp. nov. and slender-bodied species of Neolamprologus with rounded to truncate caudal fin
Neolamprologus cancellatus sp. nov. N. cylindricusa N. cylindricus N. devosia N. fasciatusb N. fasciatus N. hecquib N. hecqui N. longiora N. meelib N. nigriventrisa N. pectoralisa N. pleuromaculatusb N. pleuromaculatus N. schreyenia N. tetracanthusb N. tetracanthus
n
BD (% SL)
Ds
As
ScLL
Gr
V
8 2 7 17 3 10 53 10 2 6 5 5 49 7 9 53 10
22.3–25.2 22.7–23.3 21.2–24.0 22.5–26.5 25.0–27.0 22.1–27.4 26.3–28.2 23.0–28.3 24.4–24.6 25.0 19.5–23.0 19.6–22.5 20.0–26.3 21.4–24.0 22.8–25.1 25.3–29.9 24.5–31.5
19–20 18–19 19–20 17–19 18–19 18–20 18–19 18–19 19 18 20–21 19–20 17–19 17–18 17–19 18–20 19–20
7–8 5 5 5–6 10 9–11 6–8 6–8 5–6 7 5–6 6–8 5–7 6 5 4 4
34–37 34 32–33 34–35 35 33–37 45–55 45–52 33 42–46 — — 47–60 48–64 32 36–40 37–39
5–7 — 6–10 5–6 11–12 9–12 8–13 7–11 8 11–12 6–8 6–8 9–12 10–12 — 6–8 6–7
33 — 32–33 31–32 — 31–33 — 31–32 — — 34–35 34–35 — 32–34 — — 33–34
Ds, dorsal fin spines; As, anal fin spines; ScLL, scales in longitudinal line; Gr, gill rakers on lower limb of 1st gill arch; V, vertebrae a Data from original discription b Data from Poll (1956) including type specimens
lateral bands distinguishes it from N. fasciatus, N. hecqui, N. meeli, and N. pleuromaculatus (no band).
Key to the species of Neolamprologus with rounded to truncate caudal fin and slender body 1a. 1b. 2a. 2b. 3a. 3b. 4a. 4b. 5a. 5b. 6a. 6b. 7a. 7b. 8a. 8b. 9a.
9b.
4 spines on anal fin . . . . . . . . . . . . . . . . . .N. tetracanthus More than 4 spines on anal fin . . . . . . . . . . . . . . . . . . . 2 9 to 12 spines on anal fin . . . . . . . . . . . . . . . .N. fasciatus 5 to 8 spines on anal fin . . . . . . . . . . . . . . . . . . . . . . . . . 3 8 to 16 scales in upper lateral line . . . . . . . N. schreyeni 23 to 51 scales in upper lateral line . . . . . . . . . . . . . . . . 4 Body without vertical stripe and blotch . . . . . . . . . . . 5 Body with vertical stripes or blotches . . . . . . . . . . . . . 7 Head small, length 28.9–29.8% SL; pectoral fin long, length 27.7–32.6% SL . . . . . . . . . . . . . . . . . N. pectoralis Head large, length 30.1–32.7% SL; pectoral fin short, length 18.0–24.1% SL . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Abdomen blackish . . . . . . . . . . . . . . . . . . N. nigriventris Abdomen pale . . . . . . . . . . . . . . . . . . . . . . . . N. longior 42 to 64 scales in longitudinal line . . . . . . . . . . . . . . . . 8 32 to 37 scales in longitudinal line . . . . . . . . . . . . . . . 10 One to several blotches on body . . . . . . . . . . . . . . . . . . 9 No blotch, but with many vertical stripes on body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. hecqui Distribution at northernmost area of the lake; body relatively slender, length 20.0–26.3% SL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . N. pleuromaculatus Distribution only at central-west area of the lake (near Katibili Bay, Congo); body relatively deep, length 26.0% SL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. meeli
10a. 7 or 8 spines on anal fin . . . . . . . . N. cancellatus sp. nov. 10b. 5 or 6 spines on anal fin . . . . . . . . . . . . . . . . . . . . . . . . 11 11a. Body blackish with white vertical stripes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. cylindricus 11b. Body light beige brownish with dusky vertical stripes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. devosi Comparative materials examined. N. cylindricus.—HUMZ 122526, 74.1 mm SL, Nkumbula I., Zambia; HUMZ 125008, 81.9 mm SL, Chisanza, Zambia; HUMZ 125671, 60.7 mm SL, Zambia; HUMZ 173746, 173771, 176047, 176295, 4 specimens, 59.4–89.2 mm SL, Kasenga, Zambia. N. fasciatus.—HUMZ 116512, 86.3 mm SL, Bemba, Congo; HUMZ 174244, 174245, 175985, 176417, 176418, 176476, 176666, 176792, 176793, 9 specimens, 33.5–99.8 mm SL, Wonzye Point, Zambia; HUMZ 175985, 80.6 mm SL, Kasenga, Zambia. N. hecqui.—HUMZ 173861, 173881, 173885, 173889, 173890, 176438, 6 specimens, 42.4– 58.3 mm SL, Nkumbula I., Zambia; HUMZ 173904, 173905, 173907, 3 specimens, 43.6–54.5 mm SL, off Nkumbula I., Zambia; HUMZ 176712, 50.5 mm SL, Chituta Bay, Zambia. N. pleuromaculatus.—HUMZ 116781, 76.5 mm SL, Mouth of Ruzizi River, Burundi; HUMZ 127123, 128530, 2 specimens, 68.7–84.2 mm SL, Gitaza, Burundi; HUMZ 127273, 133215–133217, 4 specimens, 42.0–52.5 mm SL, Rumonge, Burundi. N. tetracanthus.—HUMZ 122474, 122564, 2 specimens, 81.0–82.2 mm SL, Nkumbula I., Zambia; HUMZ 123073, 98.9 mm SL, Wonzye Point, Zambia; HUMZ 125473, 56.0 mm SL, Inangu Peninsula, Zambia; HUMZ 125526, 86.9 mm SL, Cape Chikutula or Cape Kachase, Zambia; HUMZ 173737, 175972, 175973, 176052, 4 specimens, 76.0–96.7 mm SL, Kasenga, Zambia; HUMZ 176383, 71.1 mm SL, north of Mtondwe I., Zambia. Acknowledgments We express our sincere thanks to Professor Michio Hori, Kyoto University, and Professor Masanori Kohda, Osaka City University, for giving us an opportunity to conduct taxonomic research on the fishes of Lake Tanganyika. We are also thankful to L.M. Mwape,
A new cichlid from Lake Tanganyika H. Phiri, and other staff members of the Lake Tanganyika Research Unit in Mpulungu, Zambia, for their assistance during the survey, and M. Wald in New York for correction of English. This study was partly supported by the grant awarded to the first author by the Ito Foundation for the Advancement of Ichthyology, Tokyo, Japan.
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