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Accepted by M. R. de. Carvalho: 8 Oct. 2007; published: 14 Nov. 2007 39
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2007 · Magnolia Press
Zootaxa 1638: 39–49 (2007)
www.mapress.com/zootaxa/
The first case of aggressive mimicry among lamprologines in a new species of
Lepidiolamprologus (Perciformes: Cichlidae) from Lake Tanganyika
ROBERT SCHELLY1, TETSUMI TAKAHASHI2, ROGER BILLS3 & MICHIO HORI2
1Department of Ichthyology, American Museum of Natural History, Central Park West at 79th St., New York, NY 10024, USA.
E-mail: schelly@amnh.org
2Department of Zoology, Graduate School of Science, Kyoto University, Kitashirakawa-Oiwakecho, Sakyo, Kyoto 606-8502, Japan.
E-mail: tetsumi@terra.zool.kyoto-u.ac.jp; hori@terra.zool.kyoto-u.ac.jp
3South African Institute for Aquatic Biodiversity, Private Bag 1015, Grahamstown 6140, South Africa. E-mail: r.bills@ru.ac.za
Abstract
Lepidiolamprologus mimicus n. sp. is described from material collected along the Zambian coast of Lake Tanganyika. It
is distinguished from congeners by its unique color pattern of bright yellow fins, a brownish-tan flank coloration and
large, dark brown spots along the flanks, in addition to a series of meristic and morphometric characters. Lepidiolampro-
logus mimicus n. sp. exhibits an interesting feeding ecology, in which individuals blend into schools of their prey, yel-
low-finned cyprichromines, with the aid of similar coloration. This is the first instance of aggressive mimicry reported
for lamprologines.
Key words: Lepidiolamprologus mimicus new species; Lamprologini; taxonomy; Lake Tanganyika; aggressive mimicry
Introduction
Lamprologine cichlids, recognized by Takahashi (2003) as one of 16 tribes constituting the Lake Tanganyika
cichlid fauna, exhibit a broad range of morphological, ecological, and behavioral diversity (Stiassny 1997).
New species of lamprologines from Tanganyika and associated rivers continue to be discovered (Hanssens &
Snoeks 2003; Schelly et al. 2003; Aibara et al. 2005). With about 80 described lacustrine species, lamprolog-
ines comprise roughly half of the cichlid species in Lake Tanganyika, where the group is most species rich. In
addition to those of Lake Tanganyika, eight lamprologine species are known from the Congo River to the
west, and one from the Malagarasi River to the east. Lamprologines are not known from outside of the greater
Congo Drainage. Various lamprologine classifications (e.g. Pellegrin 1904; Regan 1920; Colombe & Allgayer
1985; Poll 1986) have dealt with mostly superficial anatomy, and the group is in need of a classification based
on a thorough phylogenetic analysis. Lamprologines are currently assigned to eight genera: Altolamprologus
Poll 1986, Chalinochromis Poll 1974, Julidochromis Boulenger 1898, Lamprologus Schilthuis 1891, Lepidi-
olamprologus Pellegrin 1904, Neolamprologus Colombe and Allgayer 1985, Telmatochromis Boulenger
1898, and Variabilichromis Colombe and Allgayer 1985. This paper deals with a new species in the genus
Lepidiolamprologus, considered by Poll (1986) to comprise the following superficially similar, elongate,
predatory fishes characterized by high lateral line scale counts: L. elongatus (Boulenger 1898), L. cunningtoni
(Boulenger 1906), L. attenuatus (Steindachner 1909), L. profundicola (Poll 1949), L. kendalli (Poll & Stewart
1977), and L. nkambae (Staeck 1978). More recently, analyses based on molecular data (Sturmbauer et al.
1994; Schelly et al. 2006) and morphology (Stiassny 1997; Schelly, in press) have provided evidence that L.
cunningtoni is not closely related to the remaining species assigned to Lepidiolamprologus.
SCHELLY ET AL.
40 · Zootaxa 1638 © 2007 Magnolia Press
Material and methods
Collections and observations of the fish were made using SCUBA at depths of 10–30 m. To capture the fish,
they were first driven into monofilament (1 cm stretched mesh) standing nets (2 m X 20 m) and then retrieved
by the diver with hand nets. Specimens were fixed in 10% formalin and preserved in either 50% isopropanol
or 75% ethanol. Detailed ecological observations were made with SCUBA by MH on a daily basis during the
month of November between 1995 and 2006. Meristic counts and morphometric measurements were made on
the left side of specimens with digital calipers to 0.1 mm, and followed Barel et al. (1977) with the following
additions: dorsal and anal fin base length were measured along the dorsum or ventrum from the insertion of
the first spine to the insertion of the last ray in the respective fin, pelvic fin length was measured from the
insertion of the longest ray to its tip, pre-dorsal distance was measured from the premaxillary symphysis to the
insertion of the first dorsal spine, pre-pectoral distance was measured from the premaxillary symphysis to the
insertion of the middle pectoral ray, and pre-pelvic distance was measured from the mandibular symphysis to
the insertion of the interior pelvic ray. Specimens were cleared and counterstained following Dingerkus and
Uhler (1977). Some specimens, including the holotype, were radiographed to determine vertebral counts and
fin elements. For comparison with the new species, portions of the type series of Lepidiolamprologus kendalli,
L. nkambae and L. profundicola were examined.
Abbreviations
AMNH American Museum of Natural History, New York, USA
FAKU Kyoto University Museum, Kyoto, Japan
IRSNB Institut Royal des Sciences Naturelles de Belgique, Bruxelles, Belgium
MCZ Museum of Comparative Zoology, Cambridge, USA
MRAC Musée Royal de l'Afrique Centrale, Tervuren, Belgium
SAIAB South African Institute for Aquatic Biodiversity, Grahamstown, South Africa
Zm Private collection of M. Hori, Kyoto University, Japan
BD body depth
HL head length
IW interorbital width
NLF neurocranial lateral line foramina
SL standard length
Results
Lepidiolamprologus mimicus new species
Figs. 1–5, Table 1
Holotype: SAIAB 41257, Chituta Bay, Zambia, at Chituta Bay cliffs, at 10–15 m depth along the rocky slope,
Nov. 17, 1992, R. Bills. (approximate coordinates: 08°43.82 S, 31°09.41 E)
Paratypes: SAIAB 58308 (n=1; cleared and stained), collected with holotype; FAKU 95137 (male, 123.1
mm SL), FAKU 95138 (female, 121.7 mm SL), Kasenga, Zambia, 14 m depth, 7 Nov. 2005, gill net, M. Hori.
FAKU 95139 (male, 159.7 mm SL), Kasenga, Zambia, 13–16 m depth, 3 Dec. 2005, gill net, M. Hori. FAKU
95140 (male, 119.4 mm SL), Kasenga, Zambia, 2–15 m depth, 30 Oct. 1996, gill net, M. Hori. FAKU 95141
(male, 129.6 mm SL), Kasenga, Zambia, 15 m depth, 12 Dec. 1996, gill net, H. Ochi. FAKU 95142 (male,
76.9 mm SL), Kasenga, Zambia, 19 m depth, 1 Nov. 1995, M. Hori. FAKU 95143 (male, 101.7 mm SL), off
Zootaxa 1638 © 2007 Magnolia Press · 41
NEW TANGANYIKAN CICHLID
Mtondwe Is., Zambia, depth unknown, 1 Nov. 1995, gill net, M. Hori. FAKU 95144 (female, 133.6 mm SL),
Kasenga, Zambia, 14–19 m depth, 13 Nov. 1995, gill net, M. Hori. FAKU 95145 (male, 147.0 mm SL), FAKU
95146 (female, 130.7 mm SL), Kasenge, Zambia, 15 m depth, 11 Jan. 1994, gill net, M. Hori.
FIGURE 1. Lepidiolamprologus mimicus n. sp., holotype, SAIAB 41257, 144.8 mm SL, drawing by Elaine Heemstra.
FIGURE 2. Photographs of Lepidiolamprologus mimicus n. sp. collected from Kasenga, Zambia before fixation. (upper)
Zm 05521-1, paratype, 123.1 mm SL, male. (lower) Zm 05521-2, paratype, 121.7 mm SL, female. Photo by TT.
SCHELLY ET AL.
42 · Zootaxa 1638 © 2007 Magnolia Press
FIGURE 3. Lepidiolamprologus mimicus n. sp., paratype, SAIAB 58308, 96.1 mm SL: (a) first gill arch, (b) premaxilla
and dentary with anguloarticular, labial ossification in gray, (c) lower pharyngeal jaw ventral view, (d) lower pharyngeal
jaw dorsal view.
Differential diagnosis. From the Lepidiolamprologus sensu Poll (1986), Lepidiolamprologus mimicus n.
sp. is distinguished from L. cunningtoni by, among numerous other neurocranial and gill arch characters, pos-
session of a sesamoid bone in the labial ligament, 2 pores at NLF0, and a color pattern based on three rows of
irregular spots, whereas L. cunningtoni lacks a labial ossification, has a single, coalesced NLF0 pore, and a
color pattern based on a series of faint bars. With the other five members of Poll’s Lepidiolamprologus, L.
mimicus n. sp. shares a whole suite of neurocranial, gill arch, and coloration characters, but is still easily dis-
tinguishable. Of these species, it is unique in having bright yellow fins and very dark, large spots on the
flanks, without additional small spots or a complex pattern of spots and worm lines on the head (as in L. nka-
mbae and L. kendalli). Lepidiolamprologus mimicus n. sp. is somewhat deeper-bodied than L. nkambae, L.
kendalli, and L. attenuatus (BD 23.4–26.2% SL, vs. max 23.3% SL in L. nkambae), has a more elongate head
than L. attenuatus (HL 31.0–33.7% SL, vs. 28.2–30.5% SL in L. attenuatus), has one more vertebra than both
L. nkambae and L. kendalli (34 vs. 33), and has a wider interorbital region (IW 17.4–24.5% HL) than L. nka-
mbae (IW 14.3–16.9% HL) and L. kendalli (IW 14.3–17.8% HL). In addition, it usually has one more dorsal
fin ray than both L. nkambae and L. kendalli (11 vs. 10). The longitudinal line scale count of L. mimicus n. sp.
Zootaxa 1638 © 2007 Magnolia Press · 43
NEW TANGANYIKAN CICHLID
is almost always higher (73–79) than in L. attenuatus and L. elongatus (both 66–73), L. kendalli (67–70) and
L. profundicola (66–74). Finally, L. mimicus n. sp. generally has fewer gill rakers (10–12) on the first cerato-
branchial than L. elongatus (11–14) and L. attenuatus (13–15).
FIGURE 4. Lepidiolamprologus mimicus n. sp., paratype, SAIAB 58308, 96.1 mm SL: (a) neurocranium (b) lachrymal
and dermosphenotic.
Description. Counts and measurements for the holotype and 12 paratypes given in Table 1. An elongate,
moderately shallow-bodied species (BD 23.4–26.2, mean 24.7% SL), with a large mouth and sleek form char-
acteristic of an open-water piscivore. Greatest body depth at about base of fifth dorsal fin spine. Head length
31.0–33.7, mean 32.4% SL. Head profile slightly convex, rising smoothly to dorsal fin base.
Fins. Dorsal fin XVIII 10–11, anal fin V 8–9. Spines in the dorsal fin increase gradually in length posteri-
orly until about the 7th spine, after which they remain constant. Anal fin spines gradually increase posteriorly
through entire series. Dorsal and anal fins come to a point and terminate at about the end of the caudal pedun-
cle; no rays are produced. Caudal fin large and emarginate, with 14 branched rays. Pectoral and pelvic fins
short, terminating well ahead of anus. First ray in pelvic fin is longest.
Jaws and Teeth (Fig. 2). Lower jaw prognathous, with both outer and inner row teeth pointed unicuspids
in both jaws. Single series of enlarged, recurved, procumbent caniniform teeth (8 premaxillary and 6 dentary)
situated on anterior third of both jaws, with largest teeth furthest from symphysis. Posterior to large teeth sin-
gle rows of slightly enlarged caniniform teeth run almost whole length of dentigerous arms of both dentary
and premaxilla, outer row on premaxilla with 49–60 teeth. Small, caniniform inner teeth in five to nine irregu-
lar rows nearest symphysis, tapering to single row posteriorly and running most of length of dentigerous arms
of premaxilla and dentary.
SCHELLY ET AL.
44 · Zootaxa 1638 © 2007 Magnolia Press
TABLE 1. Morphometric and Meristic data for Lepidiolamprologus mimicus n. sp.
Gill rakers (Fig. 3a). Slender, elongate, and non-denticulate, with up to four subsidiary branches. Thir-
teen to 16 gill rakers along outer row of first gill arch. No rakers present on hypobranchial, 10–12 rakers along
ceratobranchial, sometimes single raker in angle of arch, and 3–4 rakers on epibranchial.
Lower pharyngeal jaw (Fig. 3c, d). Longer than wide, with slight interdigitation along ventral suture.
About 24 teeth in posterior tooth row. All teeth slender and beveled or hooked.
Scales. Flank scales small, ctenoid, and regularly imbricating. Lateral line scales 73–79, with 51–67 in
upper and 26–45 in lower lateral line branch. Considerable overlap (16–20 scales) between upper and lower
branches of lateral line. Cheek usually naked, but 4–5 cheek scales observed in three of 13 individuals, small
scales on belly and small, deeply embedded scales on chest. Opercle and subopercle heavily scaled. Caudal
fin with scales extending more than half its length.
Vertebrae. 34; 16+18.
Additional osteology (Figs. 3, 4). Infraorbital series comprised of broad, plate-like lachrymal with 6 large
sensory canal pores, lacking infraorbitals adjacent to lachrymal. Dermosphenotic present. Labial ossification
present. Single supraneural present. Supraoccipital crest well-developed and increasing in depth posteriorly,
with sharply-angled posterior edge. No frontal ridge present, two distinct pores at NLF0. Frontal shelf well-
developed and extending anteriorly beyond NLF2. Pre-orbital process of lateral ethmoid rotated postero-
medially. Post-orbital process of sphenotic followed by a large, spoon-shaped wing. Frontal and ethmoid-
vomer continuous in profile, without abrupt drop-off.
Character holotype n mean min max SD
Standard length (mm)
Percent of standard length
Body depth
Head length
Caudal peduncle depth
Caudal peduncle length
Anal fin base length
Dorsal fin base length
Pelvic fin length
Pectoral fin length
Pre-dorsal distance
Pre-pectoral distance
Pre-pelvic distance
Percent of head length
Lower jaw length
Eye diameter
Snout length
Interorbital width
144.8
25.4
33.1
10.2
14.0
20.1
55.4
21.6
17.7
33.4
33.4
36.3
46.6
18.7
39.7
24.5
13
13
13
12
13
13
13
13
13
13
13
13
13
13
13
13
121.3
24.7
32.4
10.4
16.4
19.8
54.3
22.0
18.1
32.4
33.1
36.9
53.6
24.0
36.2
20.5
76.9
23.4
31.0
10.1
14.0
17.7
51.5
21.2
17.1
31.1
30.2
33.8
46.6
18.7
31.6
17.4
159.7
26.2
33.7
10.7
19.4
21.4
56.6
23.4
19.4
34.7
34.9
38.4
56.5
28.3
39.7
24.5
23.8
0.98
0.69
0.19
1.33
1.03
1.45
0.75
0.73
0.98
1.24
1.29
3.27
2.24
2.17
1.96
COUNTS
Total lateral line scales
Upper branch lateral line
Lower branch lateral line
Dorsal fin spines and rays
Anal fin spines and rays
Gill rakers—lower arm of 1st arch
Vertebrae
75
52
37
XVIII 11
V 9
10
16+18
73 (1), 74 (2), 75 (3), 76 (2), 77 (2), 78 (1), 79 (2)
51 (1), 52 (1), 53 (2), 54 (1), 56 (1), 57 (1), 58 (1), 60 (1), 63 (3), 67 (1)
26 (2), 29 (1), 30 (1), 31 (2), 32 (1), 34 (2), 36 (1), 37 (1), 40 (1), 45 (1)
XVIII 10 (2), XVIII 11 (11)
V 8 (5), V 9 (8)
10 (5), 11 (5), 12 (3)
16+18 (n=3)
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NEW TANGANYIKAN CICHLID
FIGURE 5. Photographs showing coloration of Lepidiolamprologus mimicus n. sp. 1) The coloration of an adult at non-
foraging time, 2) that of an adult at foraging time with females of Paracyprichromis brieni in background, 3) that of sub-
adult with young of P. brieni in background, and 4) that of young with juveniles of P. brie ni in background.
Coloration. In life, background body color brownish-tan, with three broken white to silvery-blue stripes
composed of irregular blotchy spots. Superimposed over this pattern are three rows of seven to nine large,
dark-brown spots stretching from above opercle to caudal flexure. Large spots irregular, with intervening
space between them horizontally about as large as the spots. Vertically, some spots merge together partially,
though some are staggered irregularly resulting in misalignment between horizontal rows. Largest spots in
lower row, which is superimposed over the lower branch of lateral line at about center of flank, while middle
row of slightly smaller spots is superimposed over upper lateral line branch. Top row of spots sits along dorsal
fin base, with spots stretching partially onto dorsal fin. Opercle with a patch of lavender and a black blotch
posteriorly, belly white, eyes with bright yellow dorsal and ventral bordering. Bluish-white, thin line running
below eye from lachrymal to whitish patch on cheek. All fins with bright yellow; dorsal, anal, and caudal fins
with rows of large yellow maculae interspersed with small patches of white. Yellow maculae blend together
vertically to form series of yellow bands. Pectoral fin pale yellow on the base. Preserved coloration a uniform,
light yellow-tan with brown spots in pattern described above. No sexual dimorphism in coloration.
Diet. Rocky littoral, open-water piscivore.
Ecology and behavior. (Fig. 5). Ecology and behavior of the holotype and paratypes was not recorded.
However, these aspects of the biology of Lepidiolamprologus mimicus n. sp. were studied at Kasenga from
1993 to 2006. Underwater observation during the period and analysis of stomach contents of samples revealed
that this species was exclusively piscivorous and mainly preyed on young and sub-adults of cyprichromine
cichlids. In the littoral region of Kasenga, four species of cyprichromine (Cyprichromis leptosoma, C. zona-
SCHELLY ET AL.
46 · Zootaxa 1638 © 2007 Magnolia Press
tus, C. coloratus, and Paracyprichromis brieni) form mixed-species schools composed of various subgroups
of age or sex of each species in open water 1–4 m above the rocky substrate (Takahashi et al. 2002, 2006).
Usually L. mimicus n. sp. stalked solitarily near or mingled in the school at a depth of 10 m or more, and
attacked mainly young and juveniles of the school. At least 30–40 individual hunts were observed. When
hunting, adults of L. mimicus n. sp. changed their body coloration markedly from dark-brown to pale beige;
their characteristic dark-brown spots disappeared and rows of fine silver dots on the flank faded, but the yel-
low anal fin and black dotted line at the tip of the dorsal fin remained. This coloration generally looked like
that of female cyprichromines and specifically very much resembled that of female Paracyprichromis brieni.
The degree of resemblance in coloration of L. mimicus n. sp. to its prey during hunting was greater in young
and sub-adults than adults (Fig. 5–3, -4), and the predominant prey were young and juveniles of Paracyp-
richromis brieni, with small Cyprichromis making up the balance. These phenomena indicate both that L.
mimicus n. sp. is highly specialized to forage on cyprichromine fishes and also that its coloration, especially
during hunting, is a form of aggressive mimicry.
FIGURE 6. Collection localities of Lepidiolamprologus mimicus n. sp. Holotype locality represented by open circle.
Mating and breeding of L. mimicus n. sp. have not been observed despite the long-term underwater survey
from 1993 to 2006. However, on two occasions, pairs were observed roaming together for several days around
the same area at a depth of 20–25 m. In both cases, the pairs were wandering around a boundary between
rocky and sandy substrate, and sometimes inspected holes or crevices among boulders, but did little foraging.
Zootaxa 1638 © 2007 Magnolia Press · 47
NEW TANGANYIKAN CICHLID
After the pairs were caught by gill-net underwater, gentle pressure on their abdomens made the males eject
semen and females eject mature eggs. These observations suggest that L. mimicus n. sp. is a monogamous
species like L. elongatus and L. attenuatus, and not a harem species like L. profundicola (for breeding habit of
Lepidiolamprologus, see Kuwamura 1997). It also seems that they breed in rather deep water (more than 30
m) beyond the reach of a diving survey. Mature males and females were usually caught with gill-nets in deep
water from 20 m to 50 m depth.
Distribution. (Fig. 6). Zambian coast of Lake Tanganyika from Kasenga to Kapembwa. The west side of
Chituta Bay, the type locality, is characterized by a steep rocky shore starting in the south at the west side of
the mouth of the Lunzua River, with usually poor visibility (<10 m). From the river mouth, where the rocks
run to about 5 m depth, the depth of the rocky slope increases steadily until the point at Kasenga village where
it is approximately 50 m deep. Specimens of Lepidiolamprologus mimicus n. sp. were collected as solitary
individuals wandering across the steep sloped rocky habitat, comprised of large rocks and rock slabs (some
several cubic meters). Other Lepidiolamprologus occurring sympatrically in Chituta Bay are L. cunningtoni,
L. kendalli, L. elongatus, L. attenuatus, and L. profundicola. Only L. nkambae is absent. According to Kon-
ings (1988; page 188 and photo on page 121), Lepidiolamprologus mimicus n. sp. also occurs in central Tan-
zania.
Etymology. The specific name, mimicus, is from the Latin, in reference to the specific feeding ecology
involving imitative coloration described herein.
Remarks. Lepidiolamprologus mimicus n. sp. exhibits all of the features that Poll (1986) used to charac-
terize Lepidiolamprologus: a high number of ctenoid scales (>60) in the longitudinal line; more than 20 scales
in the lower lateral line; dorsal fin formula XVII–XXI, 10–12; anal fin formula IV–V, 7–9; and large size
(140–305 mm TL). In Poll’s (1986) classification, Lepidiolamprologus included the following elongate, pred-
atory species: L. elongatus, L. attenuatus, L. profundicola, L. kendalli, L. nkambae, and L. cunningtoni. To the
exclusion of L. cunningtoni, L. mimicus n. sp. and the remaining five species of Poll’s Lepidiolamprologus all
possess an ossification in the labial ligament, two distinct pores at NLF0, forked or stellate gill rakers, scales
on the caudal fin to approximately 80% of caudal length, and three series of spots along the body, situated at
the mid-line, upper lateral line, and dorsal fin base.
Mimicry in its various forms is fairly widespread in marine fishes (Randall 2005; Moland et al. 2005). A
common form is aggressive mimicry, in which a predatory species resembles a harmless species, allowing it to
avoid detection when approaching prey (Wickler 1968). In addition to the numerous marine examples, several
tropical freshwater characids, citharinids, and cichlids have been recognized as aggressive mimics (Sazima
2002). Within Lake Tanganyika, aggressive mimicry has been described in scale-eating perissodines (Yanag-
isawa et al. 1990; Hori & Watanabe 2000), but this is the first case observed in lamprologines. Interestingly,
the abundant schools of brightly-colored Lake Tanganyika cyprichromines are subject to aggressive mimicry
from two independent cichlid lineages. Color morphs of Perissodus microlepis with yellow anal fins insinuate
themselves into schools of cyprichromines to feed on scales, while yellow-finned Lepidiolamprologus mim-
icus n. sp. blend into schools to pick off whole fish.
Comparative materials.
Lepidiolamprologus attenuatus: AMNH 98043 (alcohol and 1 C&S); MCZ 49254 (alcohol and 1 C&S);
SAIAB 41253; Zm 031-a, 031-b, 05018, 05097-2, 05193, 25041, 840-a.
Lepidiolamprologus cunningtoni: SAIAB 42470 (alcohol and 1 C&S); MCZ 32596 (1 C&S); Zm 959-
1~8.Lepidiolamprologus elongatus: AMNH 97219; SAIAB 44646 (alcohol and 1 C&S); Zm 04049-2~3,
04085-1, 04085-3, 04284-1~2, 04284-4~5, 04305.
Lepidiolamprologus kendalli: MRAC 77-36-P-1 (paratype, male, 129.9 mm SL), N. W. of Mutondwe Isl.,
Zambia, 40 m depth, 16 Nov. 1972; SAIAB 46848 (1 alc); SAIAB 46874 (2 alc); Zm 04088-1~2, 04111-1~2,
04188, 04306, 04621-1~2, 04636.
SCHELLY ET AL.
48 · Zootaxa 1638 © 2007 Magnolia Press
Lepidiolamprologus mimicus n. sp.: AMNH 237601 (n=1; tissue taken), Kapembwa, Zambia, 08°37.37'S,
30°50.87'E, Mar. 10, 2004, R.Bills and R. Schelly.
Lepidiolamprologus nkambae: MRAC 77-49-P-1 (holotype, female, 113.0 mm SL), Nkamba bay, Sumbu
National Park, Zambia, Apr. 1975; AMNH 216075 SW (9 C&S); Zm 04658-1~3, 05696-1~2, 97651-a~b,
98138.
Lepidiolamprologus profundicola: MRAC 114.160 (holotype, female, 211.3 mm SL), 114.161 (paratype,
female, 222.2 mm SL), Stat: 68, Baie au Sud immidiat du cap Tembwe, Congo, 15 Jan. 1947; MRAC 114.162
(paratype, male, 246.9 mm SL), Stat: 140, Autour de l’ile de Kabimba, 23 Feb. 1947; MRAC 82.12.P.518-519
(1 C&S); IRSNB 345 (paratype, male, 167.9 mm SL), collected with holotype; SAIAB 41255; Zm 04004,
04050, 04402, 04463, 04533, 04560-1, 04565, 04570-1, 04597-1~2, 04622-1, 04629-2.
Acknowledgments
We are very grateful to the Fisheries Department, Agriculture and Natural Resources, Government of Zambia,
for their full cooperation and permission to make collections. For assistance in the field, we are also grateful
to L. M. Mwape and H. Phiri, and other staff of the Lake Tanganyika Research Unit in Mpulungu, and to Alex
Chilala, Lackson Kachali, Ant and Belinda Forster, and John Lundberg. TT and MH express our sincere
thanks to H. Kawanabe, and M. Kohda for the opportunity to conduct this study; to J. Snoeks, P. Miguel, G.
Lenglet, and P. Campbell for assistance to TT during the examination of type specimens; and to H. Ochi, who
provided support and encouragement. RS and RB thank Sally Terry, Barbara Brown, Damaris Rodriguez,
Radford Arrindell, and Scott Holtz for assistance with collections. Thanks to Elaine Heemstra for her drawing
of the holotype. Funding was provided by a Grant-in-Aid for JSPS Fellows (No. 20188) and Grant-in-Aid for
Special Purposes (No. 18779002) for TT, and Grant-in Aid for Scientific Research on Priority Areas (No.
14087203) and Biodiversity Research of the 21st COE (A14) for MH. Additional funding was provided by the
Axelrod Ichthyology fund, a Doctoral Dissertation Improvement grant (NSF DEB-0508686), and a Biotic
Surveys and Inventories grant (NSF DEB-0542540) for RS and by the All-Catfish Species Inventory (NSF
DEB-0315963) for RB.
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