ArticlePDF Available

Descriptions of five new species in the genus Metriaclima (Teleostei: Cichlidae) from Lake Malaŵi, Africa

Authors:

Abstract and Figures

Five new species of rock-dwelling cichlids from Lake Malaŵi are described. All five species phenotypically resemble Me-triaclima aurora based on the absence of a black band in the dorsal fin and occupying the rock-sand interface of the lake. Differences in overall shape, meristics, and coloration distinguish these new species from each other and from previously described species within this group. The new species Metriaclima glaucos, M. mossambicus, M. nkhunguensis, and M. xanthos originate from the Mozambique coast of the lake, while M. sciasma is restricted to Tanzanian waters.
Content may be subject to copyright.
Accepted by J. Friel: 15 Dec. 2010; published: 12 Jan. 2011
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2011 · Magnolia Press
Zootaxa 2738: 125 (2011)
www.mapress.com/zootaxa/Article
1
Descriptions of five new species in the genus Metriaclima (Teleostei: Cichlidae)
from Lake Malaŵi, Africa
PATRICK J. CICCOTTO1, ADRIANUS KONINGS2 & JAY R. STAUFFER, JR.3
1Department of Biological Sciences, University of Maryland Baltimore County, Baltimore MD, 21250, USA.
E-mail: c76@umbc.edu
2Cichlid Press, El Paso, TX, 79913, USA. E-mail: info@cichlidpress.com
3School of Forest Resources, The Pennsylvania University, University Park, PA 16802, USA. E-mail: vc5@psu.edu
Abstract
Five new species of rock-dwelling cichlids from Lake Mala i are described. All five species phenotypically resemble Me-
triaclima aurora based on the absence of a black band in the dorsal fin and occupying the rock-sand interface of the lake.
Differences in overall shape, meristics, and coloration distinguish these new species from each other and from previously
described species within this group. The new species Metriaclima glaucos, M. mossambicus, M. nkhunguensis, and M.
xanthos originate from the Mozambique coast of the lake, while M. sciasma is restricted to Tanzanian waters.
Key words: Cichlids, Lake Mala i, Metriaclima
Introduction
The rock-dwelling haplochromine cichlids of Lake Mala i, collectively referred to as mbuna, are a diverse group
of fishes consisting of twelve genera. The mbuna genus Metriaclima was originally described by Stauffer et al.
(1997) to accommodate species in the Pseudotropheus zebra complex (Ribbink et al. 1983). The characters of the
genus Metriaclima that distinguish it from other mbuna genera include a moderately-sloped ethmo-vomerine block
with a swollen rostral tip, the presence of bicuspid teeth in the anterior portion of the outer row of both jaws, the
lower jaw at a 45° angle to a line from the tip of the snout to the middle of the caudal peduncle (Stauffer et al.
1997), the lower jaw is often slightly longer and thicker than the upper jaw, a large part of the upper dental arcade
is normally exposed when the mouth is closed, the tips of the teeth in the premaxilla and dentary are in a V-shaped
line with the anteriormost in upper and lower jaw furthest apart, the placement of the bicuspid teeth in the outer
row along the sides of the jaws does not follow the contour of the jaw bone, and the jaws are abducted to a near
180° angle during feeding so that the body is aligned at a perpendicular angle to the substrate to remove diatoms
and loose algal strands from algae attached to the substrate (Konings & Stauffer 2006).
In addition to expanding the diagnosis of Metriaclima, Konings & Stauffer (2006) recognized three phenotypi-
cally similar assemblages within the genus: 1. a Blue-Black barred group associated with M. zebra, 2. a Black Dor-
sal group associated with M. flavifemina, and 3. an Aurora group associated with M. aurora. Both the Black Dorsal
group and the Aurora group are found in the intermediate habitat along the coast of Lake Mala i, where the transi-
tion between rocks and sand occurs and are distinguished from one another by the presence or absence of a black
band in the dorsal fins of males (Konings 2001). Males hold territories mainly over the sand near rocks and exca-
vate beneath rocks to construct their spawning burrow. These species are most commonly found at depths of 3–20
m. The members of the Aurora group are characterized by their habitat preference (sand-rock interface), the
absence of a black submarginal band in the dorsal fin, and by a light brown coloration of most females exhibiting
no (or only faintly present) vertical bars and a yellow margin in the dorsal, anal, and caudal fins. Five described
species were initially recognized as part of the Aurora group: Metriaclima aurora, M. chrysomallos, M. benetos, M.
hajomaylandi, and M. barlowi (see Konings & Stauffer 2006), while putative members were recognized at several
localities along the eastern shore of Lake Mala i in Mala i, Mozambique, and Tanzania (Fig. 1).
CICCOTTO ET AL.
2 · Zootaxa 2738 © 2011 Magnolia Press
FIGURE 1. Map of Lake Mala i showing the locations of species discussed throughout the text.
Species in the genus Metriaclima, as well as species of other mbuna genera, are often differentiated based on
coloration and morphology (Stauffer et al. 1997; Genner & Turner 2005). Coloration has been shown to be a major
factor in mate recognition and selection among the mbuna (Fryer 1959; Stauffer & Van Snik 1996; Knight &
Turner 2004) and unique color patterns are recognized to be sufficient to diagnose valid species (Barlow 1974;
Barel et al. 1977; Stauffer et al. 1995). It is difficult to determine whether allopatric populations that differ in color
are hetero- or conspecific, if similarities in behavioral, ecological, and morphological characteristics are observed.
Multivariate statistical analyses of morphometric and meristic characters have generally been accepted in further
distinguishing between species similar in morphology (Bowers & Stauffer 1993). The purpose of this paper is to
describe five new species in the genus Metriaclima identified as several populations of the Aurora group (M. sp.
“aurora blue”, M. sp. “aurora lumbaulo”, M. sp. “aurora north”, M. sp. “blue reef”, M. sp. “aurora chinuni”, M. sp.
“aurora yellow”, and M. sp. “aurora blacktail”; Konings 2007) through an examination of morphometric and meris-
tic data.
Material and methods
Fishes were collected while SCUBA diving, using monofilament nets for capture. The pigmentation patterns of
males and females were recorded in the field prior to preservation. Variations of color patterns for each population
collected are reported in the descriptions by placing a slash between two colors to designate the pigmentation is
Zootaxa 2738 © 2011 Magnolia Press · 3
FIVE NEW CICHLID FISHES FROM LAKE MALA I, AFRICA
either one of the two reported colors or intermediate between the two. Fishes were anesthetized with clove oil, pre-
served in 10% formalin with fins pinned erect, and later transferred into 70% ethanol for permanent storage. Exter-
nal measurements and counts follow Barel et al. (1977) and Konings & Stauffer (2006). In the pectoral fin, all rays
were counted including the small splinter on the upper edge. Measurements and counts were made on the left side
of the fish, with the exception of gill-raker counts. The size of specimens is reported as standard length (SL) in mm.
Institutional abbreviations follow Leviton et al. (1985), except UMBC, University of Mala i, Bunda College.
To examine differences in body shape among neighboring populations or species, sheared principal component
analysis of the morphometric data was used in which the covariance matrix was factored (Humphries et al. 1981;
Bookstein et al. 1985). The meristic data were analyzed through a principal component analysis, in which the cor-
relation matrix was factored. The first principal components of the meristic data were plotted against the sheared
second principal components of the morphometric data. If the minimum polygon clusters did not differ along one
axis (p<0.05), independent of the other, a MANOVA in conjunction with a Hotelling-Lawley trace was used to
determine if the clusters were significantly different. If however the mean multivariate scores were significantly
different (p<0.05) along one axis independent of the other, a Duncan’s Multiple Range Test was used to determine
differences among the clusters (Stauffer et al. 1997).
Cichlids are substrate-oriented fishes that will not cross open water or large tracts of unsuitable habitat, limit-
ing their exploratory movements along coastlines. It is thus assumed that when genetic exchange exists between
allopatric populations it is between neighboring populations and not significantly between populations that are sep-
arated by geographically large distances or separated by populations of congenerics in similar habitats. We have
therefore limited our comparisons to populations of the Aurora group in which genetic exchange may have
occurred and disregarded those in which such is highly unlikely. Therefore, it was only necessary to compare the
new species with Metriaclima aurora and M. chrysomallos populations.
Comparative material: Metriaclima benetos. PSU 3058, holotype; PSU 3059, 9, USNM 349125, 3; ANSP
176199, 3; MFU 28, 3; 59.7–81.4 mm SL; Mala i: Lake Mala i: Manzinzi Reef.
Metriaclima callainos. PSU 2542.1, PSU 2542.2, MFU 2, USNM 322426, 60.5–80.1 mm SL; Mala i: Lake
Mala i: Nkhata Bay, May 1988.
Metriaclima aurora (Burgess 1976)
Fig. 2A&B
Coloration. Lateral body coloration blue dorsally in males from Likoma Island, Mara Point North, and N’kolon-
gwe, but brown dorsally in males from Mbweca and Tumbi Point, with anterior portion of scales sky-blue and 6
faint blue bars below dorsal fin; caudal peduncle dark blue; yellow belly and breast fading to light blue towards
anal fin. Head sky-blue with preorbital, ventral half of cheek, operculum, and preoperculum yellow; dark yellow/
green opercular spot; gular yellow. Dorsal fin yellow with submarginal sky-blue band and sky-blue blotches
throughout. Caudal-fin rays yellow with light-blue membranes. Anal fin pale blue to pale yellow with one ocellus.
Pelvic fin with light-blue leading edge, remaining yellow. Pectoral-fin rays pale yellow and membranes clear.
Female lateral ground coloration pale blue with center of scales orange/brown; belly white. Head brown; cheek
and operculum with purple highlights; gray/black opercular spot; white gular. Dorsal fin brown with brown lappets.
Caudal-fin rays brown with clear membranes. Anal fin gray proximally and brown distally with single small
orange/brown ocellus. Pelvic fin with white leading edge; first two rays and membranes brown, remainder clear.
Pectoral fin clear.
Remarks. Morphometric and meristic data for populations from Likoma Island, Mara Point, Mbweca, Tumbi
Point, and N’kolongwe in Table 1. The lack of a black submarginal band in the dorsal fin, a yellow ventral half of
the head, breast, and dorsal fin, and the light-brown colored females that have yellow fin-margins distinguish
Metriaclima aurora from most members of the genus, except M. hajomaylandi, M. chrysomallos, and M. xanthos.
Metriaclima aurora differs from M. hajomaylandi and M. chrysomallos by fewer (faint) vertical bars below the
dorsal fin (6 vs. 7–9) and from M. xanthos by a shorter lower jaw (mean 31.2, range 28.4–33.5 % HL vs. mean
35.4, range 34.6–36.7).
CICCOTTO ET AL.
4 · Zootaxa 2738 © 2011 Magnolia Press
TABLE 1. Morphometric and meristic values of Metriaclima aurora from Likoma Island, Mara Point, Mbweca, Tumbi Point,
and N’kolongwe.
All Likoma Island Mara Point
n=17 n=20
mean holotype mean range mean range
Standard length, mm 70.4 80.5 76.7 69.2–83.8 66.5 53.4–81.7
Head length, mm 21.7 24.9 24 21.9–25.8 20.4 16.3–25.5
Percentage of standard length
Head length 30.8 30.9 31.3 29.9–32.7 30.6 29.3–32.3
Snout to dorsal 33.8 32.9 33.6 32.0–35.1 33.7 31.3–35.9
Snout to pelvic 38.1 40.3 39.4 37.6–42.2 38.3 36.6–40.1
Greatest body depth 32.6 30.4 31.3 29.5–33.2 33.2 31.2–36.0
Caudal peduncle length 14.3 15.4 13.9 12.2–15.4 14.2 13.2–15.6
Least caudal peduncle depth 11.7 11.1 11.9 10.8–13.0 11.8 10.9–12.8
Dorsal-fin base length 60.1 61.5 60 57.3–61.5 60 57.1–61.9
Anterior dorsal to anterior anal 52.4 53.2 52.9 50.1–56.5 52.7 51.1–54.9
Anterior dorsal to posterior anal 63.6 63.8 63.5 60.6–66.6 63.7 61.3–65.8
Posterior dorsal to anterior anal 29.4 29.9 29 27.0–30.7 29.5 27.5–30.9
Posterior dorsal to posterior anal 15.0 16.3 14.4 12.5–16.3 15.1 13.5-16.4
Anterior dorsal to pelvic-fin origin 36.0 34.4 34.6 32.5–36.7 36.5 33.4–39.9
Posterior dorsal to pelvic-fin origin 57.4 53.2 54.3 49.5–56.2 58 56.1–60.6
Percentage of head length
Horizontal eye diameter 37.0 36.6 37.6 35.5–39.8 36.8 35.1–38.1
Vertical eye diameter 36.8 36.4 37.2 34.9–40.3 36.5 33.3–38.2
Snout length 26.9 29.8 26.4 23.7–29.8 25.5 23.4–27.9
Postorbital head length 39.3 39 38.5 37.2–39.8 39.1 36.9–41.6
Preorbital depth 21.8 26.4 21 18.6–26.4 20.7 18.6–23.7
Lower-jaw length 31.2 30.2 30.7 29.2–32.5 32 30.5–33.5
Cheek depth 23.1 23.8 24.1 20.2–26.2 22.1 18.8–25.4
Head depth 86.8 85.2 82.5 75.8–87.2 87.9 79.9–94.5
Meristics mode mode range mode range
Dorsal-fin spines 18 17 18 17–18 18 17–19
Dorsal-fin rays 9 9 9 8–10 9 8–10
Anal-fin spines 3 3 3 3–3 3 3–3
Anal-fin rays 8 8 8 7–8 8 7–8
Pelvic-fin rays 5 5 5 5–5 5 5–5
Pectoral-fin rays 14 14 14 14–15 14 13–14
Lateral line scales 31 31 30, 31 29–32 31 30–33
Pored scales posterior to lateral line 2 2 0 0–2 2 0–2
Cheek scales 4 4 4 4–5 4 4–5
Gill rakers on first ceratobranchial 12 11 12 10–13 11 10–13
Gill rakers on first epibranchial 3 3 3 3–4 3 2-4
Teeth in outer row of left lower jaw 20 17 22 17–26 18 12–25
Teeth rows on upper jaw 3 3 3 3–4 3 3–4
Teeth rows on lower jaw 3 4 4 3–4 3 3–4
Zootaxa 2738 © 2011 Magnolia Press · 5
FIVE NEW CICHLID FISHES FROM LAKE MALA I, AFRICA
continued.
Mbweca Tumbi Point N'kolongwe
n=19 n=20 n=20
mean range mean range mean range
Standard length, mm 70.7 55.3–79.7 71.3 60.8–79.3 67.8 59.2–75.2
Head length, mm 21.8 16.8–24.6 21.8 18.3–24.5 21 18.3–22.7
Percentage of standard length
Head length 30.8 29.5–31.9 30.5 28.9–32.4 31 29.7–33.0
Snout to dorsal 33.5 31.3–34.9 33.7 31.9–36.3 34.6 33.1–37.4
Snout to pelvic 37.3 34.1–41.4 38.1 36.2–40.3 37.8 36.4–39.6
Greatest body depth 33.2 31.2–35.5 32.6 30.2–34.9 32.7 30.2–35.6
Caudal peduncle length 14.8 13.7–16.2 14.2 12.3–16.5 14.4 12.6–15.8
Least caudal peduncle depth 11.8 11.2–12.5 11 10.5–11.7 11.9 11.3–13.0
Dorsal-fin base length 60.5 58.7–62.6 60.3 57.3–62.0 59.8 56.8–61.4
Anterior dorsal to anterior anal 52.1 50.7–54.2 51.5 48.8–54.3 53.1 51.4–55.4
Anterior dorsal to posterior anal 63.7 61.9–65.6 63.5 60.5–65.3 63.6 60.2–65.7
Posterior dorsal to anterior anal 29.6 28.3–30.9 29.5 27.7–30.7 29.4 27.5–31.4
Posterior dorsal to posterior anal 15 14.3–16.0 14.9 13.7–16.0 15.7 14.6–17.1
Anterior dorsal to pelvic-fin origin 36.2 33.6–39.5 36 33.0–38.4 36.6 34.5–39.3
Posterior dorsal to pelvic-origin 57.7 54.0–59.7 58.4 55.7–59.9 58 55.3–60.2
Percentage head length
Horizontal eye diameter 38 35.6–40.3 36.8 34.8–38.9 36 32.3–39.6
Vertical eye diameter 38.1 35.1–39.8 36.7 34.5–38.7 35.9 33.0–38.8
Snout length 27.6 25.4–30.1 27.7 25.1–30.5 27.3 23.7–29.9
Postorbital head length 38.3 36.4–40.0 38.9 37.0–40.7 41.5 39.0–44.2
Preorbital depth 23.3 20.4–26.0 23 20.7–25.7 20.8 19.0–23.8
Lower-jaw length 30.9 28.4–32.1 31.2 28.9–33.1 31.2 28.6–32.6
Cheek depth 23.2 19.8–25.9 22.8 21.1–25.1 23.4 20.2–26.2
Head depth 85.4 81.7–90.6 90.4 84.3–95.4 87.3 80.3–94.1
Meristics mode range range mode range
Dorsal-fin spines 18 17–18 18 17–18 18 17–19
Dorsal-fin rays 8,9 8–10 9 8–10 8 8–9
Anal-fin spines 3 3–4 3 3–4 3 3–3
Anal-fin rays 8 7–8 8 7–8 8 7–8
Pelvic- fin rays 5 5–5 5 5–5 5 5–5
Pectoral- fin rays 14 14–15 14 13–15 14 13–15
Lateral line scales 31 30–33 30 30–32 31 29–32
Pored scales posterior to lateral line 2 0–2 2 0–2 2 0–2
Cheek scales 5 3–5 4 4–6 5 4–7
Gill rakers on first ceratobranchial 12 10–13 12 11–13 12 10–14
Gill rakers on first epibranchial 3 2-5 3 2–4 3 2–4
Teeth in outer row of left lower jaw 20 16–25 18, 19, 21 17–23 20 16–25
Teeth rows on upper jaw 3 3–4 3 3–4 3 3–4
Teeth rows on lower jaw 3 3–4 3 3–4 4 3–4
CICCOTTO ET AL.
6 · Zootaxa 2738 © 2011 Magnolia Press
FIGURE 2. A. Metriaclima aurora, male in habitat at Likoma Island, Mala i; approx. 80 mm SL; not preserved; B. M.
aurora, female in habitat at Likoma Island, Mala i; approx. 70 mm SL; not preserved; C. M. chrysomallos, male in habitat at
Nametumbwe, Mala i; approx. 80 mm SL; not preserved; D. M. chrysomallos, male in habitat at Gome, Mala i; approx. 80
mm SL; not preserved.
Distribution. Metriaclima aurora is distributed at Likoma Island (type locality) and along the coast of
Mozambique at Mara Point, Mbweca, Tumbi Point, and a relatively isolated location to the south near N’kolongwe
(Fig. 1). Mbweca and Tumbi Point are the only localities where the brown-barred males are found. Populations
have been transplanted to Thumbi West Island, where the species has spread to several locations around Cape
Maclear (Stauffer & Hert 1992; Konings 2001).
Material examined. USNM 215292, holotype; USNM 215293, 16; 69.2–83.8 mm SL; Mala i: Lake Mala i:
probably Likoma Island; J. Freiberg, 5 May 1976. — PSU 4480, 20; 53.4–81.7 mm SL; Mozambique: Lake
Mala i: Mara Point North, 12° 11.254' S, 34° 41.968' E; A.F. Konings & J.R. Stauffer, 17 Feb 2002. — PSU 4481,
19; 55.3–79.7 mm SL; Mozambique: Lake Mala i: Mbweca, Lake Mala i, 12° 17.513' S, 34° 42.392' E; A.F.
Konings & J.R. Stauffer, 28 Feb 2006. — PSU 4482, 20; 60.8–79.3 mm SL; Mozambique: Lake Mala i: Tumbi
Point, 12° 20.297' S, 34° 41.853' E; A.F. Konings & J.R. Stauffer, 17 Feb 2002. — PSU 4483, 20; 59.2–75.2 mm
SL; Mozambique: Lake Mala i: N’kolongwe, 12° 47.671' S, 34° 47.159' E; A.F. Konings & J.R. Stauffer, 16 Feb
2002.
Metriaclima chrysomallos Stauffer, Bowers, Kellogg & McKaye 1997
Fig. 2C&D
Coloration. Lateral body coloration of breeding males from populations at type locality (Mumbo Island) and from
Mala ian east coast between Nsinje River and Makanjila Point light blue with gold highlights and 7–9 faint bars
below dorsal fin, fading to white ventrally; breast orange fading to white at pelvic fin. Dorsal half of head, opercu-
lum, and cheek light blue; ventral portions orange/yellow; snout orange. Dorsal fin pale blue, posterior membranes
orange distally. Caudal-fin rays orange with clear membranes. Anal-fin rays blue/gray with 2 orange ocelli. Distal
portion of pelvic fin orange, remaining clear. Pectoral fin clear.
Females brown/gray with light-blue highlights laterally; belly white. Head dark brown with blue/green high-
lights, gray/black opercular spot, and white gular. Dorsal fin brown. Caudal-fin rays brown and membranes clear.
Anal-fin rays brown with clear membranes. Leading edge of pelvic fin white, rays brown; membranes clear. Pelvic
fins clear.
Zootaxa 2738 © 2011 Magnolia Press · 7
FIVE NEW CICHLID FISHES FROM LAKE MALA I, AFRICA
TABLE 2. Morphometric and meristic values of Metriaclima chrysomallos from Nametumbwe and Gome.
all Nametumbwe Gome
n=20 n=20
mean mean range mean range
Standard length, mm 67.5 65.1 53.6–71.9 69.9 58.4–79.6
Head length, mm 21.2 20.6 16.7–22.6 21.8 18.7–23.9
Percentage of standard length
Head length 31.4 31.6 30.0–32.8 31.3 29.9–32.8
Snout to dorsal 33.4 33.4 30.7–35.7 33.3 31.1–34.8
Snout to pelvic 37.9 37.9 36.5–39.8 38.0 35.5–40.1
Greatest body depth 33.1 33.5 31.8–36.0 32.7 30.7–35.2
Caudal peduncle length 13.6 13.5 12.1–14.6 13.7 12.5–15.1
Least caudal peduncle depth 12.3 12.1 11.2–13.1 12.5 11.5–13.2
Dorsal-fin base length 61.4 61.4 58.9–63.0 61.4 58.8–64.5
Anterior dorsal to anterior anal 53.0 52.9 50.5–54.6 53.1 51.0–55.0
Anterior dorsal to posterior anal 64.2 64.0 62.3–65.9 64.3 61.5–66.7
Posterior dorsal to anterior anal 29.6 29.5 28.4–31.2 29.7 27.8–31.4
Posterior dorsal to posterior anal 15.0 14.7 13.9–16.0 15.3 14.1–16.1
Anterior dorsal to pelvic-fin origin 36.5 37.2 35.1–39.2 35.9 33.3–38.2
Posterior dorsal to pelvic-fin origin 57.9 58.5 56.3–60.1 57.2 54.7–59.0
Percentage of head length
Horizontal eye diameter 35.5 36.9 35.1–39.1 34.2 31.7–36.7
Vertical eye diameter 35.4 36.9 34.0–39.4 33.8 31.8–35.9
Snout length 25.7 24.5 20.3–27.3 26.9 23.5–29.9
Postorbital head length 39.0 38.4 37.0–39.9 39.7 37.8–41.5
Preorbital depth 20.3 19.6 16.5–22.2 20.9 18.4–24.2
Lower-jaw length 32.1 32.0 29.6–33.8 32.1 29.6–34.2
Cheek depth 23.9 22.7 17.8–25.4 25.1 20.6–28.4
Head depth 86.4 86.2 80.4–91.0 86.6 79.9–93.3
Meristics mode mode range mode range
Dorsal-fin spines 18 18 17–19 18 16–19
Dorsal-fin rays 9 9 8–10 9 8–9
Anal-fin spines 3 3 3–3 3 3–3
Anal-fin rays 8 8 7–8 8 7–9
Pelvic-fin rays 5 5 5–5 5 5–5
Pectoral-fin rays 14 14 14–15 14 14–15
Lateral line scales 31 31 30–32 31 30–32
Pored scales posterior to lateral line 2 0, 2 0–2 2 0–2
Cheek scales 4 4 3–6 4, 5 4–5
Gill rakers on first ceratobranchial 12 12 10–12 12 9–13
Gill rakers on first epibranchial 3 3 2–3 3 2–4
Teeth in outer row of left lower jaw 22 21, 22 15–24 22 19–25
Teeth rows on upper jaw 3 3 3–4 3 3–4
Teeth rows on lower jaw 3 3 3–4 3 3–4
CICCOTTO ET AL.
8 · Zootaxa 2738 © 2011 Magnolia Press
At Gome, lateral body coloration of breeding males light blue, with 7–9 faint bars below dorsal fin. Head light
blue with gray highlights; interorbital area gray with 1 blue bar; gular white with yellow blotches. Dorsal fin gray
proximally, fading to white distally. Caudal-fin rays clear with blue membranes. Anal-fin rays gray with light blue/
white membrane and 2 orange ocelli. Leading edge of pelvic fin white, rays clear; first 2 membranes gray, remain-
ing clear. Pectoral fin clear. Females same as in other populations.
Remarks. Morphometric and meristic data for Nametumbwe and Gome populations in Table 2. The absence
of a black submarginal band in the dorsal, a lack of distinct vertical bars, and the light-brown colored females that
have yellow fin-margins distinguish this species from most other Metriaclima, except M. benetos, M. glaucos new
species, and M. mossambicus new species. Male M. chrysomallos from Mumbo Island and Nametumbwe are dis-
tinguished from those of the latter three species, except the Lumessi population of M. mossambicus, by a sky-blue
nape and a yellow/orange ventral half of the head. Metriaclima chrysomallos is distinguished from males of the
Lumessi population of M. mossambicus by a blue dorsal fin, which is yellow in latter species. Males of the Gome
population of M. chrysomallos (which are all light blue and have a white-blue pectoral-fin base) are distinguished
from those of M. glaucos and M. benetos and those of M. nkhunguensis by the absence of orange/yellow pigment in
the base of the pectoral fin. In addition, males of M. benetos possess some degree of black pigment between the
eyes and in the iris. On the basis of coloration, females of a number of species in the Aurora group are indistin-
guishable from each other, but those of M. aurora have fewer and broader bars below the dorsal (6 vs. 7–9) and
those of M. glaucos generally have fewer teeth in the outer row of the lower left jaw compared to those of M. chry-
somallos (mode 16, range 14–19, vs. mode 21/22, range 15–24).
FIGURE 3. Plot of the first principal component of the meristic data (PC1) and the sheared second principal component of the
morphometric data (PC2) of Metriaclima chrysomallos from Nametumbwe ( ) and M. aurora from N’kolongwe (x).
Zootaxa 2738 © 2011 Magnolia Press · 9
FIVE NEW CICHLID FISHES FROM LAKE MALA I, AFRICA
In the populations at Mumbo Island and Nametumbwe, males possess the orange coloration on the ventral por-
tions of the head and breast to some extent. At Gome, Metriaclima chrysomallos males lack the orange coloration
on the head, breast, and posterior portion of the dorsal fin and at Ntekete, south of the Nsinje River, all-blue males
occur sympatrically with those with orange ventral heads and with those that have varying degrees of yellow color-
ation.
The population of Metriaclima chrysomallos at Nametumbwe is similar in male color pattern to M. aurora
from Likoma Island with the main difference being the number of vertical bars below the dorsal fin (7–9 vs. 6) and
the yellow dorsal fin in M. aurora. In Figure 3 the Nametumbwe population of M. chrysomallos is compared to the
nearest population of M. aurora (N’kolongwe) in a principal component analysis. The minimum polygon clusters
formed by M. aurora and M. chrysomallos overlap, but are significantly (p<0.05; MANOVA) different.
Size accounts for 81.7% of the observed variance and the second principal component accounts for 5.4%. Vari-
ables with the highest loadings on the sheared second principal components are snout length (-0.40), caudal pedun-
cle length (-0.38), and lower jaw length (0.32). Snout length, expressed as percent head length, is generally larger
in M. aurora from N’kolongwe (mean 27.3, range 23.7–29.9) compared to M. chrysomallos from Nametumbwe
(mean 24.5, range 20.3–27.3). The first principal component of the meristic data accounts for 20.5% of the total
variance. Variables with the highest loadings on the first principal component are pored scales posterior to the lat-
eral line (0.27), dorsal spines (0.24), and teeth rows on the lower jaw (0.24).
Distribution. Stauffer et al. (1997) originally described this species from Mumbo Island, but it is also found
between Makanjila Point (Mala i) and Meponda (Mozambique) in the southeastern portion of the lake (Fig. 1).
Material examined. PSU 4485, 20; 58.4–79.6 mm SL; Mala i: Lake Mala i: Gome Rock, 12° 13.30.744' S,
34° 52.021' E; A.F. Konings & J.R. Stauffer, 12 Feb 2001. — PSU 4486, 20; 53.7–72.9 mm SL; Mala i: Lake
Mala i: Nametumbwe, 13° 37.001' S, 34o 55.385' E; A.F. Konings & J.R. Stauffer, 25 Jan 2007.
Metriaclima glaucos, new species
Fig. 4A–C; Table 3
Metriaclima sp. "aurora blue": Konings 2001: 155.
Holotype. PSU 4487, 69.2 mm SL; Mozambique: Lake Mala i: Cobwé, 12° 8.243' S, 34° 45.391' E; A.F. Konings
& J.R. Stauffer, 18 Feb 2002.
Paratypes. PSU 4488, 16; UMBC 15, 1; AMNH 246003, 2; 53.0–67.4 mm SL; data as for holotype.
Diagnosis. A moderately-sloped head and bicuspid teeth in the outer row of the jaws place this species in
Metriaclima. The absence of a black submarginal band in the dorsal fin in conjunction with a yellow gular region
and branchiostegal membranes, and the light-brown colored females that have yellow fin margins distinguish M.
glaucos from other members of Metriaclima, except M. chrysomallos and M. benetos. There are generally fewer
teeth in the outer row of the lower left jaw (mode 16, range 14–19) of M. glaucos compared to M. chrysomallos
(mode 22, range 15–25) and more compared to M. benetos (mode 12, range 9–13). On the basis of color pattern,
females of M. glaucos cannot reliably be distinguished from those of most other members of the Aurora group, but
they differ from those of M. xanthos new species by a shorter lower jaw (mean 30.8, range 28.8–32.8 % HL vs.
mean 35.4, range 34.6–36.7 % HL) and from those of M. aurora by 8–11 vertical bars below the dorsal fin vs. 6 in
M. aurora.
Description. Morphometric and meristic data in Table 3. Dorsal snout profile slightly concave to straight;
mouth cleft slightly downward to horizontal; jaws isognathous. Teeth on dentary in 2-3 rows, on premaxilla in 2 or
3 rows; outer row teeth typically bicuspid anteriorly and unicuspid posteriorly, middle rows when present tricuspid,
innermost row unicuspid; lower pharyngeal jaw with numerous slender teeth with teeth in posterior row slightly
larger. Portion of upper dental arcade normally exposed when mouth closed. Tips of teeth in premaxilla and dentary
in V-shaped line with anteriormost in upper and lower jaw furthest apart and not touching in closed mouth. Lateral
scales ctenoid.
Breeding males light blue/gray laterally with 7 dark gray bars below dorsal fin and white belly. Head light
blue/gray with preorbital, cheek, and operculum light blue; darker blue opercular spot; yellow gular region and bra-
chiostegals. Dorsal fin blue with white lappets. Caudal-fin rays gray with white/blue membranes. Anal fin gray
with 1–4 yellow ocelli. Pelvic fin with white leading edge, first ray gray and remainder clear. Pectoral fin clear.
CICCOTTO ET AL.
10 · Zootaxa 2738 © 2011 Magnolia Press
TABLE 3. Morphometric and meristic values of Metriaclima glaucos from Cobwé. Mean and range include holotype.
Metriaclima glaucos
n=20
holotype mean range
Standard length, mm 69.2 61.4 53.0–69.2
Head length, mm 21.0 18.7 16.1–21.0
Percentage of standard length
Head length 30.3 30.5 29.4–31.9
Snout to dorsal 32.7 33.4 31.7–34.6
Snout to pelvic 37.4 38.2 35.9–40.4
Greatest body depth 31.9 32.8 31.8–34.0
Caudal peduncle length 14.6 14.2 12.5–15.5
Least caudal peduncle depth 10.9 11.0 9.9–12.3
Dorsal-fin base length 64.4 61.9 59.8–64.4
Anterior dorsal to anterior anal 52.5 53.6 50.8–55.8
Anterior dorsal to posterior anal 66.3 64.4 62.9–66.5
Posterior dorsal to anterior anal 29.6 29.3 27.1–30.9
Posterior dorsal to posterior anal 14.9 15.1 14.4–16.5
Anterior dorsal to pelvic-fin origin 36.6 37.2 35.2–39.5
Posterior dorsal to pelvic-fin origin 59.1 58.7 55.6–60.7
Percentage of head length
Horizontal eye diameter 30.8 33.7 30.7–36.6
Vertical eye diameter 32.0 33.7 31.0–36.9
Snout length 28.1 28.9 25.4–30.8
Postorbital head length 42.0 41.8 38.5–43.3
Preorbital depth 21.5 21.9 19.2–23.6
Lower-jaw length 29.3 30.8 28.8–32.8
Cheek depth 25.0 23.8 21.4–26.7
Head depth 92.5 91.0 86.5–95.9
Meristics mode range
Dorsal-fin spines 18 18 17–19
Dorsal-fin rays 8 9 8–10
Anal-fin spines 3 3 3–3
Anal-fin rays 8 8 7–8
Pelvic-fin rays 5 5 4–5
Pectoral-fin rays 14 14 14–15
Lateral line scales 30 30 29–32
Pored scales posterior to lateral line 1 2 0–2
Cheek scales 6 6 4–7
Gill rakers on first ceratobranchial 11 11 10–11
Gill rakers on first epibranchial 3 3 2–4
Teeth in outer row of left lower jaw 16 16 14–19
Teeth rows on upper jaw 3 3 2–3
Teeth rows on lower jaw 3 3 2–3
Zootaxa 2738 © 2011 Magnolia Press · 11
FIVE NEW CICHLID FISHES FROM LAKE MALA I, AFRICA
FIGURE 4. A. Metriaclima glaucos, holotype, PSU 4487, 69.2 mm SL; Mozambique: Lake Mala i: Cobwé. B. M. glaucos,
male in habitat at Cobwé, Mozambique; approx. 65 mm SL; not preserved. C. M. glaucos, female in habitat at Cobwé, Mozam-
bique; approx. 55 mm SL; not preserved.
Lateral coloration of females gray dorsally and white ventrally. Head gray with gray cheeks, blue/gray opercu-
lum with green highlights, gray/black opercular spot, and white gular region. Dorsal fin gray with yellow/orange
lappets. Caudal-fin rays gray with clear membranes and faint yellow/orange spots. Anal fin gray proximally and
yellow distally without ocelli. Pelvic fin with white leading edge and black submarginal band, membranes yellow
to clear. Pectoral fins clear.
Distribution. Metriaclima glaucos was collected at Cobwé, Mozambique, and is only known from the type
locality (Fig. 1).
Etymology. The specific epithet, glaucos, is Greek for bluish gray, in reference to the blue-gray body and fin
coloration. It is treated as a noun in apposition.
Metriaclima xanthos, new species
Fig. 5A–C; Table 4
Metriaclima sp. "aurora lumbaulo": Konings, 2001: 155.
Holotype. PSU 4494, 44.3 mm SL; Mozambique: Lake Mala i: Lumbaulo, 11° 54.024' S, 34° 54.307' E; A.F.
Konings & J.R. Stauffer, 19 Feb 2002.
Paratypes. PSU 4495, 4; 44.6–49.3 mm SL; data as for holotype.
Diagnosis. A moderately-sloped head and bicuspid teeth in the outer row of the jaws place this species in
Metriaclima. The brown and blue/gray alternating lateral bars in conjunction with the yellow dorsal and dark anal
fin of breeding males distinguish M. xanthos from males of all other members of Metriaclima. Female M. xanthos
are yellow/brown, usually more yellow than females of any other member of the Aurora group, but not all female
M. xanthos can be distinguished according to the color pattern. Metriaclima xanthos is further distinguished from
CICCOTTO ET AL.
12 · Zootaxa 2738 © 2011 Magnolia Press
M. aurora males from the Mbweca and Tumbi Point populations that are similar but lack dark pigment in the anal
fin, and from the neighboring M. glaucos by the lower jaw length, which is longer in M. xanthos (mean 35.4, range
34.6–36.7 % HL) than in M. aurora (mean 31.2, range 28.4–33.5 % HL) and in M. glaucos (mean 30.8, range
28.8–32.8 % HL).
Description. Morphometric and meristic data in Table 4. Dorsal snout profile slightly concave to straight;
mouth cleft at slightly downward angle; jaws isognathous. Teeth on dentary in 2 or 3 rows, on premaxilla in 2 or 3
rows; outer row teeth typically bicuspid anteriorly and unicuspid posteriorly, inner rows tricuspid, innermost row
unicuspid; lower pharyngeal jaw with numerous slender teeth with teeth in posterior row slightly larger. Portion of
upper dental arcade normally visible in closed mouth. Tips of teeth in premaxilla and dentary in V-shaped line with
anteriormost in upper and lower jaw furthest apart and not touching in closed mouth. Lateral scales ctenoid.
FIGURE 5. A. Metriaclima xanthos, holotype, PSU 4494, 44.3 mm SL; Mozambique: Lake Mala i: Lumbaulo. B. M. xan-
thos, male in habitat at Lumbaulo, Mozambique; approx. 70 mm SL; not preserved. C. M. xanthos, female in habitat at Lum-
baulo, Mozambique; approx. 50 mm SL; not preserved.
Lateral body coloration of breeding male blue/gray with 6 or 7 brown bars below dorsal fin, yellow ventrally,
yellow caudal peduncle with flecks of blue. Head gray with 2 blue/gray interorbital bars; ventral third of operculum
yellow with gray opercular spot; gular region yellow. Dorsal fin yellow with blue lappets. Caudal-fin rays yellow
with blue membranes. Anal fin dark gray/brown with light blue distal edge and 4 orange ocelli. Leading edge of
pelvic fin white, first ray brown, remainder yellow. Pectoral fin clear.
Female yellow/brown laterally with 5 faint brown bars, yellow breast, and white belly. Head orange/brown;
gular region yellow; gray/black opercular spot. Dorsal fin yellow/brown with orange lappets. Caudal-fin rays
orange with brown membrane. Anal fin orange with 1–4 small yellow ocelli. Pelvic fin orange. Pectoral fin clear.
Remarks. We compared the minimum polygon clusters formed by plotting the second sheared principal com-
ponent of the morphometric data against the first principal component of the meristic data for Metriaclima aurora,
M. glaucos, and M. xanthos (Fig. 6). Size accounted for 90.9% of the observed variance and the second principal
component accounted for 2.4%. Variables with the highest loadings on the sheared second principal components
were horizontal eye diameter (0.59), vertical eye diameter (0.55), and snout length (-0.28). The first principal com-
ponent of the meristic data accounted for 14.3% of the total variance. Variables with the highest loadings on the
first principal component were tooth rows on lower jaw (0.34), tooth rows on upper jaw (0.32), and dorsal spines
Zootaxa 2738 © 2011 Magnolia Press · 13
FIVE NEW CICHLID FISHES FROM LAKE MALA I, AFRICA
TABLE 4. Morphometric and meristic values of Metriaclima xanthos from Lumbaulo. Mean and range include holotype.
Metriaclima xanthos
n=5
holotype mean range
Standard length, mm 44.3 45.7 44.3–49.3
Head length, mm 13.9 14.7 13.9–15.6
Percentage of standard length
Head length 31.4 32.2 31.4–32.8
Snout to dorsal 35.4 34.7 32.4–36.5
Snout to pelvic 36.4 36.9 36.1–37.4
Greatest body depth 33.9 33.0 30.8–35.1
Caudal peduncle length 13.2 14.2 12.8–15.3
Least caudal peduncle depth 11.4 11.9 11.4–12.4
Dorsal-fin base length 62.8 61.6 58.6–63.9
Anterior dorsal to anterior anal 52.6 51.3 49.6–53.1
Anterior dorsal to posterior anal 65.6 64.1 62.2–65.6
Posterior dorsal to anterior anal 30.3 29.8 28.3–31.1
Posterior dorsal to posterior anal 15.9 15.4 14.8–16.7
Anterior dorsal to pelvic-fin origin 37.2 35.9 33.3–37.4
Posterior dorsal to pelvic-fin origin 60.7 57.8 55.3–60.7
Percentage of head length
Horizontal eye diameter 36.8 37.1 36.7–37.6
Vertical eye diameter 33.7 35.9 33.7–37.1
Snout length 27.5 26.2 24.1–27.5
Postorbital head length 43.7 40.0 38.4–43.7
Preorbital depth 21.4 20.9 19.2–21.7
Lower-jaw length 34.6 35.4 34.6–36.7
Cheek depth 20.1 20.8 18.7–22.6
Head depth 92.2 87.8 80.8–92.2
Meristics mode range
Dorsal-fin spines 17 17 17–18
Dorsal-fin rays 9 8, 9 8–10
Anal-fin spines 3 3 3–3
Anal-fin rays 9 8 8–9
Pelvic-fin rays 5 5 5–5
Pectoral-fin rays 13 14 13–14
Lateral line scales 29 29, 31 29–32
Pored scales posterior to lateral line 0 1, 2 0–2
Cheek scales 4 4 3–5
Gill rakers on first ceratobranchial 12 11, 12 11–13
Gill rakers on first epibranchial 3 3 2–3
Teeth in outer row of left lower jaw 21 21 15–23
Teeth rows on upper jaw 3 3 2–3
Teeth rows on lower jaw 3 2 2–3
CICCOTTO ET AL.
14 · Zootaxa 2738 © 2011 Magnolia Press
(0.24). The minimum polygon clusters formed by M. aurora and M. glaucos did not overlap. Metriaclima aurora
had a larger horizontal eye diameter (mean 37.0) and vertical eye diameter (mean 36.8) than M. glaucos (mean 33.7
for horizontal and vertical eye diameters). Metriaclima xanthos was intermediate between these two species, how-
ever a MANOVA in conjunction with a Hotelling-Lawley trace demonstrated that the three minimum polygon
clusters were significantly (p<0.05) different.
Distribution. Metriaclima xanthos was only collected at Lumbaulo, Mozambique, and is not known to occur
elsewhere (Fig. 1).
Etymology. The specific epithet, xanthos, is Greek for yellow, in reference to the yellow belly and dorsal fin of
breeding males. It is treated as a noun in apposition.
FIGURE 6. Plot of the first principal component of the meristic data (PC1) and the sheared second principal component of the
morphometric data (PC2) of Metriaclima aurora (, Likoma Island; x, Mara Point; , Mbweca; +, Tumbi Point; , N’kolon-
gwe), M. xanthos (), and M. glaucos ( ).
Metriaclima sciasma, new species
Fig. 7A–C; Table 5
Pseudotropheus sp. "kingsizei north": Konings 2001: 159.
Metriaclima sp. "aurora north": Konings 2007: 177.
Holotype. PSU 4490, 63.8 mm SL; Tanzania: Lake Mala i: Magunga, 10° 19.344' S, 34° 34.849' E; T.D. Kocher,
A.F. Konings, & J.R. Stauffer, 11 Feb 2005.
Zootaxa 2738 © 2011 Magnolia Press · 15
FIVE NEW CICHLID FISHES FROM LAKE MALA I, AFRICA
Paratypes. PSU 4491, 17; UMBC 16, 1; AMNH 246002, 2; 48.4–71.2 mm SL; data as for holotype. — PSU
4492, 20; 46.9–66.9 mm SL; Tanzania: Lake Mala i: Ngwazi, 10° 10.490' S, 34° 32.927' E; T.D. Kocher, A.F.
Konings, & J.R. Stauffer, 10 Feb 2005. — PSU 4493, 20; 53.6–68.5 mm SL; Tanzania: Lake Mala i: Makonde, 9°
56.862' S, 34° 27.296' E; T.D. Kocher, A.F. Konings, & J.R. Stauffer, 8 Feb 2005.
FIGURE 7. A. Metriaclima sciasma, holotype, PSU 4490, 63.8 mm SL; Tanzania: Lake Mala i: Magunga. B. M. sciasma,
male in habitat at Magunga, Tanzania; approx. 60 mm SL; not preserved. C. M. sciasma, female in habitat at Magunga, Tanza-
nia; approx. 50 mm SL; not preserved.
Diagnosis. A moderately-sloped head and bicuspid teeth in the outer row of the jaws place this species in
Metriaclima. A black pelvic fin, a dark-blue cheek and preorbital area, and a blue dorsal fin without a black sub-
marginal band in breeding males and the light-brown colored females that have yellow fin-margins distinguish M.
sciasma from all other members of Metriaclima. Breeding males of M. sciasma are distinguished from those of
some populations of M. zebra by the presence of a single interorbital bar (2 in M. zebra) and by black pigment on
the anterior membranes of the anal fin. Female M. sciasma cannot reliably be distinguished from those of most
other members of the Aurora group, but differ from those of M. aurora by more vertical bars below the dorsal fin
(7–10 vs. 6).
Description. Morphometric and meristic data in Table 5. Dorsal snout profile slightly concave to straight;
mouth cleft with slightly downward angle; jaws isognathous. Teeth on dentary in two to three rows, on premaxilla
in two to four rows; outer row teeth typically bicuspid anteriorly and unicuspid posteriorly, inner rows tricuspid,
innermost row unicuspid; lower pharyngeal jaw with numerous slender teeth with teeth in posterior row slightly
larger . Portion of upper dental arcade normally visible in closed mouth. Tips of teeth in premaxilla and dentary in
V-shaped line with anteriormost in upper and lower jaw furthest apart and separate in closed mouth. Lateral scales
ctenoid.
Lateral body coloration of breeding males blue dorsally with 7–10 gray bars, white ventrally, and blue/gray
caudal peduncle. Preorbital, cheek, and operculum dark blue/gray; blue opercular spot; gray/blue gular region; pre-
maxillary and maxillary blue/gray; blue/gray interorbital bar. Dorsal fin blue with light-blue lappets. Caudal fin
with gray rays and blue membranes. Anal fin blue/gray with 2–7 yellow ocelli. Pelvic fin black with white leading
edge. Pectoral fin clear.
CICCOTTO ET AL.
16 · Zootaxa 2738 © 2011 Magnolia Press
TABLE 5. Morphometric and meristic values of Metriaclima sciasma from Magunga, Makonde, and Ngwazi. Holotype from
Magunga.
all Magunga Makonde Ngwazi
n=21 n=20 n=20
holotype mean range range range
Standard length, mm 63.8 60.9 48.4–71.2 53.6–68.5 46.9–66.9
Head length, mm 18.7 18.3 14.8–21.0 16.5–21.1 14.4–20.6
Percentage of standard length
Head length 29.4 30.1 28.7–31.8 29.0–31.9 29.0–31.4
Snout to dorsal 30.4 31.1 29.3–33.6 29.9–33.1 30.0–33.1
Snout to pelvic 38.6 37.8 35.6–39.7 37.0–43.4 34.6–38.4
Greatest body depth 31.4 31.8 29.1–34.3 30.5–34.1 29.6–33.7
Caudal peduncle length 15.2 14.5 13.7–15.9 12.4–16.2 12.9–16.5
Least caudal peduncle depth 12.4 11.8 10.7–12.4 11.1–12.3 11.3–13.1
Dorsal-fin base length 63.5 61.7 59.6–63.7 59.0–63.6 58.9–65.5
Anterior dorsal to anterior anal 51.6 51.1 47.7–53.8 49.3–54.3 48.8–53.6
Anterior dorsal to posterior anal 65.3 64.2 62.3–66.7 61.8–66.7 60.6–66.9
Posterior dorsal to anterior anal 31.0 29.7 27.7–32.3 26.9–30.7 27.0–32.1
Posterior dorsal to posterior anal 15.6 14.8 13.7–15.6 14.1–15.9 13.4–15.7
Anterior dorsal to pelvic-fin origin 35.6 35.3 31.0–38.3 34.0–38.0 32.8–38.0
Posterior dorsal to pelvic-fin origin 56.8 57.3 54.6–60.7 53.3–58.6 56.1–60.5
Percentage of head length
Horizontal eye diameter 33.9 33.8 31.4–37.1 30.9–35.1 30.4–36.8
Vertical eye diameter 32.7 33.7 31.9–36.6 30.9–35.1 31.5–37.1
Snout length 26.9 26.3 22.8–28.1 23.0–31.5 23.8–28.1
Postorbital head length 41.5 41.0 39.2–42.6 38.8–43.1 38.7–43.3
Preorbital depth 23.7 21.4 18.3–24.6 16.6–28.9 18.0–23.5
Lower-jaw length 33.2 34.3 31.6–35.9 30.7–36.0 32.7–36.9
Cheek depth 26.4 24.5 20.2–28.8 20.9–27.8 18.6–28.7
Head depth 93.5 90.8 80.8–101.6 86.0–96.8 81.9–93.7
Meristics mode range range range
Dorsal-fin spines 17 17 16–18 16–18 17–19
Dorsal-fin rays 9 9 8–10 8–9 8–10
Anal-fin spines 3 3 3–3 3–3 3–3
Anal-fin rays 8 8 7–9 7–8 7–9
Pelvic-fin rays 5 5 5–5 5–5 5–5
Pectoral-fin rays 14 14 14–15 14–15 13–15
Lateral line scales 29 30 29–31 28–30 29–31
Pored scales posterior to lateral line 2 2 0–3 0–2 0–2
Cheek scales 5 5 4–6 3–6 3–6
Gill rakers on first ceratobranchial 12 12 11–13 10–13 11–14
Gill rakers on first epibranchial 3 3 3–4 2–4 2–4
Teeth in outer row of left lower jaw 19 17 13–21 14–23 12–22
Teeth rows on upper jaw 3 3 2–3 2–3 2–4
Teeth rows on lower jaw 3 3 2–3 2–3 2–3
Zootaxa 2738 © 2011 Magnolia Press · 17
FIVE NEW CICHLID FISHES FROM LAKE MALA I, AFRICA
Females brown dorsally and gray/white ventrally. Head brown with blue highlights on operculum and grey/
green opercular spot; white gular region. Dorsal and anal fins yellow/brown. Proximal third of caudal fin dark
brown, distal third light brown/yellow, membranes with faint blue spots. Pelvic fin with white leading edge, first 2
rays and membranes yellow/brown with remaining clear. Pectoral fin clear.
Distribution. Metriaclima sciasma has an extensive distribution north of the Ruhuhu River in the northeastern
part of Lake Mala i (Fig. 1). The river and the sandy shores south of Mbamba Bay form a geographic barrier
between M. sciasma and the other members of the Aurora group.
Etymology. The specific epithet, sciasma, is Greek for shadow, in reference to the black pelvic fins of males. It
is treated as a noun in apposition.
Metriaclima nkhunguensis, new species
Fig. 8A–C; Table 6
Metriaclima sp. "blue reef": Konings 2001: 58.
Holotype. PSU 4603, 72.8 mm SL; Mozambique: Lake Mala i: Nkhungu Reef, 12° 58.801' S, 34° 45.837' E; A.F.
Konings & J.R. Stauffer, 1 Mar 2006.
Paratypes. PSU 4604, 5; 65.3–73.3 mm SL; data as for holotype.
Diagnosis. A moderately-sloped head and bicuspid teeth in the outer row of the jaws place this species in
Metriaclima. Both males and females are light blue, fading to white ventrally, distinguishing them from all other
members of Metriaclima, except M. callainos. Male M. nkhunguensis are distinguished from those of the similarly
colored M. chrysomallos from the Gome population, M. estherae, and M. callainos, the latter two species of the
sediment-free rocky habitat, by the yellow base of the pectoral fin (vs. no yellow pigment in males of the other
three species). Female M. nkhunguensis are light blue, lighter than those of the only other known species with all-
blue females, M. callainos, which are a darker cobalt-blue (female M. estherae are either brown or orange/red).
Metriaclima nkhunguensis is further distinguished from M. callainos by a shorter lower jaw (mean 31.2, range
29.4–34.1 % HL vs. mean 37.5, range 34.9–43.3 % HL).
FIGURE 8. A. Metriaclima nkhunguensis, holotype, PSU 4603, 72.8 mm SL; Mozambique: Lake Mala i: Nkhungu Reef. B.
M. nkhunguensis, male in habitat at Nkhungu Reef, Mozambique; approx. 60 mm SL; not preserved. C. M. nkhunguensis,
female in habitat at Nkhungu Reef, Mozambique; not preserved.
CICCOTTO ET AL.
18 · Zootaxa 2738 © 2011 Magnolia Press
TABLE 6. Morphometric and meristic values of Metriaclima nkhunguensis from Nkhungu Reef. Mean and range include holo-
type.
Metriaclima nkhunguensis
n=6
holotype mean range
Standard length, mm 72.8 69.6 65.3–73.3
Head length, mm 22.3 20.9 19.8–22.3
Percentage of standard length
Head length 30.6 30.1 29.1–31.0
Snout to dorsal 29.8 31.1 29.8–33.1
Snout to pelvic 38.1 37.5 36.5–38.1
Greatest body depth 31.9 32.9 31.6–34.1
Caudal peduncle length 14.3 14.0 12.9–15.3
Least caudal peduncle depth 12.3 12.2 11.7–12.7
Dorsal-fin base length 62.5 61.4 58.5–62.6
Anterior dorsal to anterior anal 53.7 52.6 51.5–53.7
Anterior dorsal to posterior anal 64.9 63.7 62.1–64.9
Posterior dorsal to anterior anal 29.5 29.6 28.0–32.0
Posterior dorsal to posterior anal 15.9 15.2 14.1–16.4
Anterior dorsal to pelvic-fin origin 36.2 36.1 34.6–37.5
Posterior dorsal to pelvic-fin origin 59.8 59.5 58.3–60.2
Percentage of head length
Horizontal eye diameter 32.6 34.0 32.6–35.8
Vertical eye diameter 33.1 34.3 33.1–35.3
Snout length 29.4 27.1 24.6–29.4
Postorbital head length 39.4 38.3 35.6–40.6
Preorbital depth 23.5 21.1 19.0–23.5
Lower-jaw length 29.7 31.2 29.4–34.1
Cheek depth 24.5 24.8 23.0–26.3
Head depth 91.6 92.5 89.4–96.3
Meristics mode range
Dorsal-fin spines 17 17, 18 17–18
Dorsal-fin rays 10 9 9–10
Anal-fin spines 3 3 3–3
Anal-fin rays 7 8 7–8
Pelvic-fin rays 5 5 5–5
Pectoral-fin rays 15 14 14–15
Lateral line scales 31 30, 31 30–31
Pored scales posterior to lateral line 1 1 0–2
Cheek scales 4 5 4–5
Gill rakers on first ceratobranchial 13 13 11–13
Gill rakers on first epibranchial 2 3 2–3
Teeth in outer row of left lower jaw 22 22 19–23
Teeth rows on upper jaw 3 3 3–4
Teeth rows on lower jaw 3 3 3–3
Zootaxa 2738 © 2011 Magnolia Press · 19
FIVE NEW CICHLID FISHES FROM LAKE MALA I, AFRICA
Description. Morphometric and meristic data in Table 6. Dorsal snout profile slightly concave to straight;
mouth cleft with slightly downward angle; jaws isognathous. Teeth on dentary in three rows, on premaxilla in three
to four rows; outer row teeth typically bicuspid anteriorly and unicuspid posteriorly, inner rows tricuspid, inner-
most row unicuspid; lower pharyngeal jaw with numerous slender teeth with teeth in posterior row slightly larger.
Portion of upper dental arcade normally visible in closed mouth. Tips of teeth in premaxilla and dentary in V-
shaped line with anteriormost in upper and lower jaw furthest apart and separate in closed mouth. Lateral scales
ctenoid.
Breeding males pale blue laterally fading to white-blue ventrally. Head entirely pale blue with white gular
region. Dorsal fin light blue with white-blue lappets. Caudal-fin rays clear with blue membranes. Anal fin light
blue and 1–5 orange ocelli. Pelvic fin light blue with white leading edge. Pectoral fin clear with orange base.
Females light blue fading to white ventrally. Head light blue/gray, white gular region. Dorsal fin light blue with
orange lappets. Caudal fin light blue. Anal fin light blue with 1-4 orange ocelli. Pelvic fin light blue/gray with
white leading edge. Pectoral fin clear with white base.
Distribution. Metriaclima nkhunguensis is endemic to Nkhungu and Minos reefs in Mozambique (Fig. 1).
Etymology. The specific name refers to one of the reefs at which the species is endemic.
Metriaclima mossambicus, new species
Fig. 9A–E; Table 7
Metriaclima sp. "aurora chinuni": Konings 2001: 158.
Metriaclima sp. "aurora black tail": Konings 2001: 158.
Metriaclima sp. "aurora yellow": Konings 2001: 158.
Holotype. PSU 4497, 79.7 mm SL; Mozambique: Lake Mala i: Chiloelo, 13° 12.541' S, 34° 48.523' E; A.F. Kon-
ings & J.R. Stauffer, 13 Feb 2002.
Paratypes. PSU 4498, 16; UMBC 17, 1; AMNH 246001, 2; 54.9–77.0 mm SL; data as for holotype. — PSU
4601, 19; 47.8–76.5 mm SL; Mozambique: Lake Mala i: Chinuni, 13°03.753'S 34°47.909'E; A.F. Konings & J.R.
Stauffer, 14 Feb 2002. — PSU 4602, 18; 54.5–79.1 mm SL; Mozambique: Lake Mala i: Lumessi, 13° 8.196' S,
34° 47.844' E; A.F. Konings & J.R. Stauffer, 14 Feb 2002.
Diagnosis. A moderately-sloped head and bicuspid teeth in the outer row of the jaws place this species in
Metriaclima. Male M. mossambicus from Chiloelo are distinguished from those of all other members of Metria-
clima by the absence of a black submarginal band in the dorsal and by black streaks in the caudal and in the poste-
rior portion of the dorsal fin. Male M. mossambicus from Chinuni are distinguished from those of all other
Metriaclima by the absence of a black submarginal band in the dorsal and by a narrow black submarginal band in
the anal fin. Male M. mossambicus from Lumessi are distinguished from all other Metriaclima, except M. aurora,
M. chrysomallos, and M. xanthos by the absence of a black submarginal band in the dorsal, a yellow ventral half of
the head, breast, and dorsal fin, and a light-blue nape. Male M. mossambicus from Lumessi population differ from
those of M. chrysomallos by a yellow dorsal fin which is light-blue in the latter. Metriaclima mossambicus differs
from M. aurora by more vertical bars below the dorsal, 7–9 vs. 6, and from M. xanthos by a shorter lower jaw
(mean 31.4, range 27.9–33.8 % HL vs. mean 35.4, range 34.6–36.7 % HL). Based on the color pattern, female M.
mossambicus cannot reliably be distinguished from those of most other members of the Aurora group but have 7 or
8 bars below the dorsal fin vs. 6 bars in M. aurora.
Description. Morphometric and meristic data in Table 7. Dorsal snout profile slightly concave to straight;
mouth cleft with slightly downward angle; jaws isognathous. Teeth on dentary in two to four rows, on premaxilla in
two to four rows; outer row teeth typically bicuspid anteriorly and unicuspid posteriorly, inner rows tricuspid,
innermost row unicuspid; lower pharyngeal jaw with numerous slender teeth with teeth in posterior row slightly
larger. Portion of upper dental arcade normally exposed when mouth closed. Tips of teeth in premaxilla and dentary
in V-shaped line with anteriormost in upper and lower jaw furthest apart and not touching in closed mouth. Lateral
scales ctenoid.
CICCOTTO ET AL.
20 · Zootaxa 2738 © 2011 Magnolia Press
TABLE 7. Morphometric and meristic values of Metriaclima mossambicus from Chiloelo, Chinuni, and Lumessi. Holotype
from Chiloelo.
all Chiloelo Chinuni Lumessi
n=20 n=19 n=18
holotype mean range range range
Standard length, mm 79.7 67.4 54.9–79.7 47.8–76.5 54.5-79.1
Head length, mm 23.7 20.8 16.6–23.9 14.5–23.4 16.7–24.2
Percentage of standard length
Head length 29.8 30.9 29.8–32.7 29.3–31.9 29.7–32.0
Snout to dorsal 31.3 33.8 31.3–37.0 32.6–35.2 32.7–36.3
Snout to pelvic 39.4 39.3 37.8–41.0 37.3–41.3 37.4–40.4
Greatest body depth 33.8 34.1 32.6–37.4 31.2–37.0 32.1–35.1
Caudal peduncle length 13.2 13.9 13.1–15.6 12.3–15.2 13.0–15.3
Least caudal peduncle depth 12.4 11.9 11.3–13.0 11.0–13.3 10.7–12.6
Dorsal-fin base length 62.4 61.0 59.5–63.4 59.0–63.8 59.0–62.5
Anterior dorsal to anterior anal 56.6 54.3 51.7–58.4 51.6–57.8 52.2–56.3
Anterior dorsal to posterior anal 66.7 64.8 63.7–67.2 61.8–67.3 63.6–67.1
Posterior dorsal to anterior anal 30.8 29.6 28.8–32.1 27.6–31.2 27.4–31.5
Posterior dorsal to posterior anal 16.5 15.7 15.2–17.4 13.5–16.9 14.6–16.9
Anterior dorsal to pelvic-fin origin 38.8 37.8 35.5–41.8 33.1–40.7 35.8–39.2
Posterior dorsal to pelvic-fin origin 58.9 58.1 55.6–61.7 55.7–61.2 55.6–59.8
Percentage of head length
Horizontal eye diameter 31.4 33.8 31.4–37.5 31.1–35.8 32.1–36.6
Vertical eye diameter 31.0 33.6 31.0–35.9 31.6–35.6 31.9–36.5
Snout length 28.9 28.2 26.6–31.3 26.0–29.7 26.3–30.2
Postorbital head length 42.1 41.7 40.4–44.8 39.3–42.4 40.2–43.0
Preorbital depth 23.4 23.6 19.1–29.0 18.9–26.2 21.3–27.9
Lower-jaw length 31.0 31.4 27.9–33.8 29.0–32.6 29.8–33.1
Cheek depth 31.8 24.9 22.7–31.8 19.0–26.2 21.7–28.2
Head depth 96.9 96.0 89.0–103.0 83.3–100.0 95.1–111.8
Meristics mode range range range
Dorsal-fin spines 18 18 17–18 17–18 16–18
Dorsal-fin rays 9 9 8–10 8–10 8–10
Anal-fin spines 3 3 3–3 3–3 3–3
Anal-fin rays 8 8 7–9 7–9 7–8
Pelvic-fin rays 5 5 5–5 5–5 5–5
Pectoral-fin rays 14 14 14–15 14–15 14–15
Lateral line scales 31 30 29–32 30–32 29–31
Pored scales posterior to lateral line 2 2 0–2 0–2 0–3
Cheek scales 4 5 3–5 3–6 3–6
Gill rakers on first ceratobranchial 12 12 11–13 11–12 10–13
Gill rakers on first epibranchial 3 3 2–4 2–4 2–3
Teeth in outer row of left lower jaw 15 21 15–23 15–24 16–26
Teeth rows on upper jaw 3 3 3–4 3–4 2–4
Teeth rows on lower jaw 3 3 2–4 3–4 3–4
Zootaxa 2738 © 2011 Magnolia Press · 21
FIVE NEW CICHLID FISHES FROM LAKE MALA I, AFRICA
FIGURE 9. A. Metriaclima mossambicus, holotype, PSU 4497, 79.7 mm SL; Mozambique: Lake Mala i: Chiloelo. B. M.
mossambicus, male in habitat at Chiloelo, Mozambique; approx. 75 mm SL; not preserved. C. M. mossambicus, female in hab-
itat at Chiloelo, Mozambique; approx. 50 mm SL; not preserved. D. M. mossambicus, male in habitat at Chinuni, Mozambique;
approx. 75 mm SL; not preserved. E. M. mossambicus, male in habitat at Lumessi, Mozambique; approx. 80 mm SL; not pre-
served.
Breeding males at Lumessi typically with gray/blue ground color with posterior edge of scales gold in dorsal 2/
3 and faint lateral barring; ventral third gold. Ventral third of head gold; cheeks yellow, gular region orange, inter-
orbital light blue/gray with gray/green interorbital bar, and gray/black opercular spot. Proximal 2/3 of dorsal fin
gold, remainder blue/gray. Caudal fin gold with yellow/gray membranes. Proximal 1/3 of anal fin orange, distal 2/3
blue/gray. Pelvic fin orange. Pectoral-fin rays orange with clear membranes.
Breeding males from Chinuni typically with light-blue ground color and 7 gray bars below dorsal fin, caudal
peduncle blue/gray. Head light blue dorsally with 1 dark interorbital bar, purple cheek and operculum, pale yellow
gular region, and gray opercular spot. Dorsal fin blue with orange tips. Caudal-fin rays pale blue with light-orange
membranes. Anal fin gray with a black submarginal band, white lappets, and 2-3 yellow ocelli. Pelvic fin dark gray
with white leading edge and broad black submarginal band. Pectoral fins clear.
Breeding males from Chiloelo with light-blue ground color and 7 or 8 dark blue bars below dorsal fin; belly
white with area between pelvic fins orange. Head light-blue with 1 gray interorbital bar, gular region orange. Dor-
sal fin light-blue with white lappets, posterior third of rayed portion with black membranes. Caudal fin black with
light-blue rays. Anal fin light blue/gray with 2–6 orange ocelli and black submarginal band and white lappets. Pel-
CICCOTTO ET AL.
22 · Zootaxa 2738 © 2011 Magnolia Press
vic fin clear, except for white leading edge and gray/black band. Pectoral fins clear with gray rays and yellow/
orange base.
Females of all three populations of Metriaclima mossambicus indistinguishable. Body gray/brown laterally;
belly white. Head brown, cheek with green highlights, gular region white. Dorsal fin gray with green spots and
orange lappets; orange tips on rays. Caudal fin clear with orange/yellow margin and small blue spots throughout.
Anal fin clear in proximal half, orange/brown in distal half. Pelvic fin clear except for first two orange/brown mem-
branes. Pectoral fin clear.
FIGURE 10. Plot of the first principal component of the meristic data (PC1) and the sheared second principal component of
the morphometric data (PC2) of Metriaclima aurora from N’kolongwe ( ) and M. mossambicus (x, Lumessi; , Chiloelo; +,
Chinuni).
Remarks. Initially, we believed the three populations of M. mossambicus were separate species based on dif-
ferences in male breeding color. The minimum polygon clusters formed by plotting the first principal components
of the meristic data against the sheared second principal components of the morphometric data for the populations
from Chiloelo, Chinuni, and Lumessi completely overlap, however (Fig. 10). These morphologically similar popu-
lations of M. mossambicus are considered conspecific. Variation in male coloration within populations has been
observed, such as the amount of black pigment in the caudal fin in males from Chiloelo or the extent of yellow on
the lateral portions of males from Lumessi. Between Meponda and the Chiloelo River (28 km) no other Aurora
group species have been observed.
In the principal component analysis of M. mossambicus and M. aurora from N’kolongwe, size accounts for
88.1% of the observed variance and the second principal component accounts for 3.4%. Variables with the highest
loadings on the sheared second principal components are preorbital depth (-0.64), vertical eye diameter (0.31), and
head depth (-0.31). The first principal component of the meristic data accounts for 17.9% of the total variance.
Variables with the highest loadings on the first principal component are tooth rows on lower jaw (0.57), dorsal
spines (0.48), and tooth rows on upper jaw (0.38). The minimum polygons for M. aurora and M. mossambicus
overlap, but are significantly (p<0.05; MANOVA) different (Fig. 10).
Distribution. Metriaclima mossambicus is found along the Mozambique shore of the lake, between the
Chiloelo River and Chinuni (Fig. 1).
Etymology. The specific epithet, mossambicus, refers to the area where the species is native. An adjective.
Zootaxa 2738 © 2011 Magnolia Press · 23
FIVE NEW CICHLID FISHES FROM LAKE MALA I, AFRICA
Artificial key to Metriaclima species
An artificial key was constructed for Metriaclima based primarily on color patterns in addition to characters
reported in Stauffer et al. (1997), Konings & Stauffer (2006), and examinations of type material. We follow Kon-
ings (2007) in regarding M. sandaracinos as a synonym to M. pyrsonotos and M. melabranchion a synonym of M.
zebra. Several undescribed species will likely be placed in this genus. Pseudotropheus livingstonii, P. elegans, P.
pursus, and M. lanisticola are the subject of another study in preparation and it appears that the skull of several of
these species does not conform to the typical characters of Metriaclima. We have included in the key M. lanisticola
as this species is without doubt a valid member of the genus.
1. Females beige to yellow with yellow pigment in anal fin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
- Females brown, dark gray or blue without yellow anal fin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .14
2. Males with black stripe in dorsal fin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3
- Males without black stripe in dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
3. Snout length 27.7–36.6% HL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. flavifemina
- Snout length 37.9–44.7% HL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .M. phaeos
4. Six or fewer bars (distinct or faint) below dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .5
- Seven or more bars (distinct or faint) below dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6
5. Lower jaw length 28.4–33.5 %HL (mean 31.2) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .M. aurora
- Lower jaw length 34.6–36.7 %HL (mean 35.4) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. xanthos (part)
6. Males with distinct vertical bars . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7
- Males without distinct vertical bars . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .10
7. Males with entirely black pelvic fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .M. sciasma
- Males with clear pelvic fin or dark band in pelvic fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .8
8. Males with black submarginal band in anal fin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .9
- Males without black pigment in anal fin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. hajomaylandi
9. Males with yellow dorsal fin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. xanthos (part)
- Males with white/blue dorsal fin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .M. mossambicus (part)
10. Males with yellow nape. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. barlowi
- Males with blue nape. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11. Males with distinct interorbital bar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. benetos
- Males without distinct interorbital bar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .12
12. Males with yellow dorsal fin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .M. mossambicus (part)
- Males with white/blue dorsal fin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .13
13. Fourteen-ninenteen, mode 16 of teeth in outer row of lower left jaw. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. glaucos
- Fifteen-twenty five, mode 22 of teeth in outer row of lower left jaw. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .M. chrysomallos
14. Females blue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .15
- Females brown to blue-gray . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .17
15. Females with distinct vertical bars . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. lombardoi
- Females without distinct vertical bars. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16. Lower jaw length 29–34% HL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. nkhunguensis
- Lower jaw length 35–43% HL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .M. callainos
17. Males without distinct vertical bars . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .18
- Males with distinct vertical bars . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .22
18. Males with orange to red pigment in dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .19
- Males with blue dorsal fin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .20
19. Males with yellow pectoral fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. mbenjii
- Males with clear pectoral fin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. greshakei
20. Males with yellow gular region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .M. xanstomachus
- Males with white gular region. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .21
21. Females light brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .M. zebra (part)
- Females dark gray-blue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. estherae
22. Five or fewer bars below dorsal fin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .M. lanisticola
- Six or more bars below dorsal fin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .23
23. Males with orange to red pigment in dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .24
- Males with blue dorsal fin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .27
24. Anal fin same orange color as dorsal fin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .M. cyneusmarginatus (part)
- Anal fin blue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .25
25. Males with black submarginal band in anal fin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. emmiltos
- Males without black band in anal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .26
26. Base of pectoral fin in males yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. fainzilberi (part)
- Base of pectoral fin in males white to blue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. pyrsonotos
CICCOTTO ET AL.
24 · Zootaxa 2738 © 2011 Magnolia Press
27. Anal fin black and black streaks in caudal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. pulpican
- Anal fin blue and no black streaks in caudal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .28
28. Base of pectoral fin in males yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. fainzilberi (part)
- Base of pectoral fin males white to blue. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .29
29. Males with black stripe in dorsal fin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .30
- Males without black stripe in dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .M. zebra (part)
30. Males with black pigment in anal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .M. zebra (part)
- Males without black pigment in anal fin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .M. cyneusmarginatus (part)
Discussion
We have used the principal component analysis plots to distinguish between neighboring populations and species.
Three of the previously described Aurora group species not examined in these analyses included Metriaclima bene-
tos, M. hajomaylandi and M. barlowi. We disregarded these species because they are geographically separated
from the new species described and none have a neighboring population to any of the newly described species.
Metriaclima sciasma was not included in our PCA analysis because of the geographic distance between their
ranges and that of the other Aurora group species discussed herein as well; moreover its male coloration is rather
distinct and easily distinguished from all other species in the genus. Similarly, M. nkhunguensis was omitted from
the PCA plots because of the blue female coloration, which is rare among mbuna in general, and its isolation on
Nkhungu and Minos reefs.
Although differing in male coloration, the three sampled populations of Metriaclima mossambicus are not con-
sidered separate species because of major overlaps in morphology of neighboring populations, the similarity of
female coloration, and the observed variability in male coloration within each population. Intra-population vari-
ance in color among males, as observed in M. mossambicus and in other mbuna species, e.g., Labeotropheus fuelle-
borni (Arnegard et al. 1999), suggests that specific mate recognition systems are flexible and have not isolated a
precise male color pattern. These neighboring populations may have begun to diverge into separate species. The
three populations of M. mossambicus are considered semispecies sensu Mayr (1963) that have acquired some, but
not all characteristics of species status. While it appears that these three populations have acquired the coloration
differences necessary for reproductive isolation, different evolution trajectories cannot be diagnosed at this time.
The polychromatism in M. mossambicus may also be a function of environmental variability.
Polychromatism in Lake Victoria cichlids (Seehausen et al. 1997; Seehausen & van Alphen 1998) as well as
the Central American cichlid Cichlasoma citrinellum (Barlow 1976) both appear to be correlated with water turbid-
ity. Environmental differences at the localities of the three color-morphs of Metriaclima mossambicus may have
favored the development of one color pattern over the other. Moreover, rocky habitats in between the three col-
lected populations have not been sampled and they may be inhabited by M. mossambicus males of which exhibit
color patterns intermediate to the ones described here. Such a gradual change in male color pattern is observed in
the various populations of M. chrysomallos as commented above. Metriaclima mossambicus is not regarded as
conspecific with M. chrysomallos because the intermediate habitat in between the southernmost population of the
former and the northernmost population of the latter is occupied by a member of the Black Dorsal group, the not
yet scientifically described M. sp. ‘black dorsal chiloelo’. This area is bordered at either end by a river delta, the
Chiloelo River to the north and the Meponda River to the south. These deltas serve as additional barriers to a possi-
ble genetic exchange between the two species.
Members of either the Aurora or the Black Dorsal group completely populate, in alternating fashion, the avail-
able intermediate habitats along the Mozambique shoreline. The only area with ample intermediate rocky habitat
that at first seems to lack a representative of either group is found at Minos and Nkhungu reefs (Fig. 1). Females of
all known species of the Aurora group are characterized by a light, yellow to beige body coloration and yellow
margins of unpaired fins, yet females of the only member of Metriaclima found in the intermediate habitat at these
reefs, M. nkhunguensis, are light blue and lack any yellow fin margin. The reefs are shallow, less than seven meters
deep, and another common member of Metriaclima, M. estherae, occurs in the more rocky sections, but always in
close proximity to M. nkhunguensis. Male coloration of both these species is identical, with the exception of the
orange pectoral fin base of M. nkhunguensis. In addition to the pectoral fin base pigment, these species can only be
identified by their respective positions in their habitat or by a close-up examination of their mouths, which are
wider in M. estherae. The latter characters support our assumption that M. nkhunguensis is a member of the Aurora
group in which the female coloration is different at these reefs compared to other populations in the group.
Zootaxa 2738 © 2011 Magnolia Press · 25
FIVE NEW CICHLID FISHES FROM LAKE MALA I, AFRICA
Acknowledgments
We thank the government of Mala i for providing the permits that allowed this research to be possible. Fishes
were collected and processed under the research permit issued to the Molecular Biology and Ecology Research
Unit (MBERU) University of Mala i and the approval of the Animal Use and Care Committee at Pennsylvania
State University (IACUC #16945; 00R084). This work was funded in part by the NSF/NIH joint program in ecol-
ogy of infectious diseases (DEB 0224958).
Literature cited
Arnegard, M.E., Markert, J.A., Danely, P.D., Stauffer, J.R., Ambali, A.J. & Kocher, T.D. (1999) Population structure and colour
variation of the cichlid fish Labeotropheus fuelleborni Ahl along a recently formed archipelago of rocky habitat patches in
southern Lake Mala i. Proceedings of the Royal Society of London, Series B: Biological Sciences, 266, 119–130.
Barel, C., van Oijen, M., Witte, F. & Witte-Maas, E. (1977) An introduction to the taxonomy and morphology of the haplo-
chromine Cichlidae from Lake Victoria. Netherlands Journal of Zoology, 27, 333–389.
Barlow, G.W. (1974) Contrasts in social behavior between Central American cichlid fishes and coral reef surgeon fishes. Amer-
ican Zoologist, 14, 9–34.
Barlow, G.W. (1976) The Midas cichlid in Nicaragua. In: Thorsen, T.B. (Ed), Investigations of the ichthyofauna of Nicaraguan
lakes. University of Nebraska, Lincoln, pp. 333–358.
Bookstein, F., Chernoff, B., Elder, R., Humphries, J., Smith, G. & Strauff, R. (1985) Morphometrics in evolutionary biology.
The Academy of Natural Sciences of Philadelphia. Philadelphia, Pennsylvania, Special Publication 15, 277 pp.
Bowers, N.J. & Stauffer, J.R., Jr. (1993) New species of rock-dwelling cichlid (Pisces: Cichlidae) from Lake Mala i Africa,
with comments on Melanochromis vermivorous Trewavas. Copeia, 1993, 715–722.
Fryer, G. (1959) The trophic interrelationships and ecology of some littoral communities of Lake Nyasa with special reference
to the fishes, and a discussion of the evolution of a group of rock-frequenting Cichlidae. Proceedings of the Zoological
Society of London, 132, 153–282.
Genner, M.J. & Turner, G.F. (2005) The mbuna cichlids of Lake Malawi: a model of rapid speciation and adaptive radiation.
Fish and Fisheries, 6, 1–34.
Humphries, J.M., Bookstein, F.L., Chernoff, B., Smith, G.R, Elder, R.L. & Poss, S.G. (1981) Multivariate discrimination by
shape in relation to size. Systematic Zoology, 30, 291–308.
Knight, M.E. & Turner, G.F. (2004) Laboratory mating trials indicate incipient speciation by sexual selection among popula-
tions of the cichlid fish Pseudotropheus zebra from Lake Malawi. Proceedings of the Royal Society of London, Series B:
Biological Sciences, 271, 675–680.
Konings, A. (2001) Mala i cichlids in their natural habitat, Third edition. Cichlid Press, El Paso, Texas, 352 pp.
Konings, A. (2007) Mala i cichlids in their natural habitat, Fourth edition. Cichlid Press, El Paso, Texas, 424 pp.
Konings, A.F. & Stauffer, J.R., Jr. (2006) Revised diagnosis of Metriaclima (Teleostei: Cichlidae) with description of a new
species from Lake Mala i National Park, Africa. Ichthyological Explorations of Freshwaters, 17, 233–246.
Leviton, A.E., Gibbs, R.H., Heal, E. & Dawson, C.E. (1985) Standards in herpetology and ichthyology: part I. Standard sym-
bolic codes for institutional resource collections in herpetology and ichthyology. Copeia 1985, 802–832.
Mayr, E. (1963) Animal species and evolution. The Belknap Press of Harvard University Press, Cambridge, Massachusetts, 797
pp.
Ribbink, A.J., Marsh, B.A., Marsh, A.C., Ribbink, A.C. & Sharp, B.J. (1983) A preliminary survey of the cichlid fishes of
rocky habitats in Lake Malawi. South African Journal of Zoology, 18, 149–310.
Seehausen, O., Van Alphen, J.J.M. & Witte, F. (1997) Cichlid fish diversity threatened by eutrophication that curbs sexual
selection. Science, 277, 1808–1811.
Seehausen, O. & Van Alphen, J.J.M. (1998) The effect of male coloration on female mate choice in closely related Lake Victo-
ria cichlids (Haplochromis nyererei complex). Behavioral Ecology and Sociobiology, 42, 1–8.
Stauffer, J.R., Jr., Bowers, N.J., Kellogg, K.A. & McKaye, K.R. (1997) A revision of the blue-black Pseudotropheus zebra
(Teleostei: Cichlidae) complex from Lake Malawi Africa, with a description of a new genus and ten new species. Proceed-
ings of the Academy of Natural Sciences of Philadelphia, 148, 189–230.
Stauffer, J.R., Jr., Bowers, N.J., McKaye, K.R. & Kocher, T.D. (1995) Evolutionarily significant units among cichlid fishes:
The role of behavioral studies. In J. L. Nielson (Ed), Evolution in the Aquatic Ecosystem: Defining Unique Units in Popu-
lation Conservation. American Fisheries Society Special Publication 17, Bethesda, Maryland, pp. 227–244.
Stauffer, J.R., Jr. & Hert, E. (1992) Pseudotropheus callainos, a new species of mbuna (Cichlidae), with analyses of changes
associated with two intra-lacustrine transplantations in Lake Malawi, Africa. Ichthyological Explorations of Freshwaters,
3, 253–264.
Stauffer, J.R., Jr. & Van Snik, E.S. (1996) New species of Petrotilapia (Teleostei: Cichlidae) from Lake Malawi, Africa.
Copeia, 1996, 695–702.
... specimens) basins (Fig. 1) using a combination of electrofishing and seining by the Maryland Department of Natural Resources' Maryland Biological Stream Survey (MBSS). The small sample sizes of specimens from the Potomac and Patapsco river populations reflect the rarity of P. peltata in these watersheds and throughout Maryland streams (MDNR, 2007; Ciccotto and Stranko, 2011). Fish were anesthetized with MS-222, preserved in 10% formalin, and then later transferred into 70% ethanol. ...
... A principal component analysis in which the correlation matrix was factored was used to analyze the meristic data. A Duncan's Multiple Range Test was then used to determine differences among clusters along each axis with each population considered a separate group and with all populations outside the Choptank River grouped together (Stauffer and van Snik, 1997; Ciccotto et al., 2011). All analyses were conducted in SAS version 9.2 (SAS Institute, Cary, NC). ...
... ). Due to its geographic isolation, morphological distinctness, and its unique habitat, the Choptank River population of P. peltata should be managed separately from the western shore to help preserve this unique component of the biological heritage of the Chesapeake Bay. In the Eastern Piedmont physiographic province in Maryland, P. peltata is threatened mainly due to expanding urbanization (Ciccotto and Stranko, 2011). Different threats likely face the Choptank River populations, where agricultural landscapes are more widespread but human population densities are lower. ...
Article
Full-text available
Morphological variation among populations of Percina peltata (Shield Darter) from streams in Maryland and Pennsylvania was examined, with particular emphasis on comparing a geographically isolated, lowland population on the eastern shore of the Chesapeake Bay (Choptank River basin) that has been excluded from previous systematic works on other upland populations of this species from west and north of the Chesapeake Bay. Statistically significant morphometric differences were observed between the isolated population and other populations examined, with measurements of eye diameter and the distance between the posterior of the first dorsal fin and the anterior and posterior bases of the anal fin explaining the most variation between populations. No differences in meristics were found. The morphological distinctness and geographic isolation of the Choptank River population of P. peltata, in conjunction with the threatened status of the species in Maryland, suggest special emphasis should be given to conserving this unique component of the biodiversity of the Chesapeake Bay.
... in lakes Malaŵi and Victoria have emphasized the importance of male courtship colors in the diagnosis of species (e.g. Ribbink et al. 1983; Seehausen 1996; Turner 1996). However, Knight & Turner (2004, using mate choice experiments, showed that color differences between allopatric morphs is not an absolute character in determining specific status. Ciccotto et al. (2011) have tried to link a difference in color pattern to an additional morphological difference of neighboring populations or morphs, in order to determine or estimate heterospecificity. Lake Malaŵi is well known for its endemic, small, rock-dwelling haplochromine cichlid fishes known as mbuna. Although Joyce et al. (2011) found that they we ...
... It should be noted that several nominal species, once included in Metriaclima are no longer recognized in this genus. Metriaclima melabranchion was considered to be conspecific with M. zebra and therefore a junior synonym (Ciccotto et al. 2011). In Stauffer et al. (1997) Pseudotropheus heteropictus Staeck 1980 was included in Metriaclima on the assumption that it represented the species found at Thumbi West Island, later described as M. flavifemina, but is now again (Konings & Stauffer 2006) referred to as Pseudotropheus heteropictus. ...
Article
Full-text available
Lake Malawi is known for its endemic haplochromine species flock, most notably the rock-dwelling cichlids known as mbuna. One of the larger genera of mbuna is Metriaclima, a group consisting of 31 described species (including the five described herein) and approximately 45 recognized unique populations. Metriaclima is diagnosed by its feeding behavior and several morphological characteristics including the angle of the vomer and the presence of bicuspid teeth in the outer row of both the upper and lower jaws. Metriaclima zebra, the type species for the genus, was described based on a single specimen. While the collection location of this holotype is not known, based on the travel records of its collector, it is likely that the specimen originated from Likoma Island. The holotype was therefore compared to specimens from several localities around this island and was found to be morphologically indistinguishable from some of these. This study includes the morphological analysis of 496 specimens of Metriaclima belonging to 31 collections from Lake Malawi. Morphometric differences were analyzed and the relationships among several distinguishable populations of Metriaclima zebra were investigated. Our study further resulted in the description of the following five new species belonging to the M. zebra species complex: M. pambazuko, M. lundoense, M. midomo, M. tarakiki, and M. nigrodorsalis. These species were distinguished and described based on color patterns, morphometric, meristic, and ecological differences. These new species were compared with and distinguished from nearby populations of Metriaclima having similar pigmentation patterns and/or similar ecological niches. An artificial dichotomous key to the described species of Metriaclima is presented
... Alternatively, some of the geographically close, but strongly diverged populations might represent cryptic species. Such patterns of cryptic diversity are common in the rock-dwelling cichlids, and new species with subtle morphological differences are frequently discovered (Stauffer et al., 1997(Stauffer et al., , 2013Ciccotto et al., 2011). The status of species is difficult to test with genetic data in Lake Malawi cichlids as the intraspecific diversity may be as large as divergence between closely related taxa (e.g., Husemann et al., 2015). ...
Article
Full-text available
Ecological speciation, which relies heavily on selection driving the emergence of new species, has become the primary paradigm through which rapid species radiations are understood. In this way, selection, particularly ecological selection, is assumed to be the driver of most species radiations. However, in many radiations, such as the radiation of Lake Malawi’s cichlids, the assumption of selection as the driver of speciation has rarely been explicitly tested, and drift, often, has completely been ignored as potential factor. In order to understand the forces driving the divergence of Lake Malawi cichlids at the microevolutionary level, we studied the genetic and phenotypic divergence of ten allopatric populations of Maylandia zebra. We estimated effective population sizes as proxy for drift. Further, we compared neutral genetic differentiation to divergence in three phenotypic traits: body size, body shape, and melanophore counts. We found small, yet significant, population differentiation in all the studied traits across most populations. Population sizes were small rendering the potential for drift to be high. However, phenotypic differentiation exceeded neutral expectations for all traits suggesting divergent local selection. Our data suggest that natural, and potentially also sexual, selection may be the dominant force driving population differentiation in Lake Malawi’s rock-dwelling cichlids, despite the potential for drift in small populations.
Article
We estimated the effective population sizes (Ne ) and tested for short term temporal demographic stability of populations of two Lake Malawi cichlids: Maylandia benetos, a microendemic, and Maylandia zebra, a widespread species found across the lake. We sampled a total of 351 individuals, genotyped them at 13 microsatellite loci and sequenced their mitochondrial D-Loop to estimate genetic diversity, population structure, demographic history and effective population sizes. At the microsatellite loci, genetic diversity was high in all populations. Yet, genetic diversity was relatively low for the sequence data. Microsatellites yielded mean Ne estimates of 481 individuals (+/- 99 SD) for M. benetos and between 597 (+/- 106.3 SD) and 1524 (+/- 483.9 SD) individuals for local populations of M. zebra. The microsatellite data indicated no deviations from mutation-drift equilibrium. Maylandia zebra was further found to be in migration-drift equilibrium. Temporal fluctuations in allele frequencies were limited across the sampling period for both species. Bayesian Skyline analyses suggested a recent expansion of M. zebra populations in line with lake level fluctuations, whereas the demographic history of M. benetos could only be estimated for the very recent past. Divergence time estimates placed the origin of M. benetos within the last 100 ka after the refilling of the lake and suggested that it split off the sympatric M. zebra population. Overall, our data indicate that microendemics and populations in less favorable habitats have smaller Ne indicating that drift may play an important role driving their divergence. Yet, despite small population sizes high genetic variation can be maintained. This article is protected by copyright. All rights reserved. This article is protected by copyright. All rights reserved.
Article
Full-text available
A new genus, Metriaclima, is described for members of the Pseudotropheus zebra complex from Lake Malaŵi. The presence of bicuspid teeth in the anterior portion of the outer row of both the upper and lower jaws distinguishes Metriaclima from many of the previously described genera of rock-dwelling cichlids that inhabit Lake Malaŵi, including Cyathochromis, Cynotilapia, Gephyrochromis, Labidochromis, and Petrotilapia. The absence of two horizontal stripes along its flanks, distinguishes it from Melanochromis. The isognathous jaws of Metriaclima delimits it from Genyochromis, which is characterized by having the lower jaw extend in front of the upper jaw. The mouth of Metriaclima is terminal, while that of Labeotropheus is inferior. Within the genus Pseudotropheus as it is now recognized, species of Metriaclima are unique because they have a moderately sloped ethmo-vomerine block and a swollen rostral tip. Ten previously undescribed species that have a slight variation from the characteristic blue/black barring are described. The new species are recognized primarily by their distinctive adult coloring in conjunction with the discontinuity of morphological differences throughout their range.
Article
Full-text available
A rock-dwelling cichlid endemic to Chinyamwezi and Chinyankwazi islands, Lake Malaŵi, Malaŵi, 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 nonterritorial males and females. Cheek depth, as expressed as percent of head length, is typically much smaller in the new species than in previously described members of the genus.
Article
Full-text available
Morphometric and meristic values were obtained from six populations of Lake Malaŵi fishes purported to belong to the species Melanochromis vermivorus Trewavas and compared with data from type material of M. vermivorus. Multivariate statistical analysis of the data revealed differences between the type material and the sampled populations with respect to shape and meristic counts. A previously recognized, but undescribed, species, Melanochromis sp. "chinyamwezi" (endemic to Chinyamwezi Island), is shown to be conspecific with the supposed M. vermivorus populations. We describe a new species of Melanochromis based on the above material.
Article
Full-text available
Multivariate discrimination by shape in relation to size. Syst. Zool., 30:291–308.—The diverse methods for analyzing size-free shape differences tend to be guided by computational expediency rather than geometric principles. We question the use of ratios and ad hoc combinations of spatially unrelated measures. Neither are linear discriminant functions or series of independent regressions helpful to the visualization of shape differences. A bridge is needed between traditional quantitative methods and thegeometrical analysis of shape. In principle any measured transects between landmarks of a form can serve as characters in a morphometric analysis. Systematic studies use a highly non-random sample of these, particularly biased regarding geometrical information. We suggest defining size and shape in terms of factors—estimates of information common to a universe of measured distances. The model presented here calculates a linear combination of variables that quantifies shape differences among populations, independent of size. In analyses in which the first two principal components confound size and shape, size is removed from one axis with shear coefficients derived from regression of general size on principal components centered by group. The general size factor is estimated by the principal axis of the within-group covariancematrix of the log-transformed data. Residuals from the regression of general size onthe transformed axes approximate a shape-discriminating factor that is uncorrelated with size within group and displays the interpopulation shape differences borne by the first two principal components. The results bear a direct and interpretable correspondence to biorthogonal analysis of shape difference.