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Two new species of Stigmatochromis (Teleostei: Cichlidae) from Lake Malawi, Africa

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Species of Stigmatochomis are haplochromine cichlids endemic to Lake Malawi, and the genus currently includes four species, S. woodi (type species), S. modestus, S. pholidophorus, and S. pleurospilus. The latter two are known from only their holotypes. Herein, we describe two additional species, Stigmatochromis macrorhynchos and Stigmatochromis melanchros from southern Lake Malawi.
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Two new species of Stigmatochromis (Teleostei: Cichlidae) from Lake
Malaw
ˆi, Africa
Jay R. Stauffer, Jr., Rachel M. Cleaver-Yoder, and Adrianus F. Konings
(JRS, RMC-Y) School of Forest Resources, The Pennsylvania State University, University
Park, Pennsylvania 16802, U.S.A.;
(AFK) Cichlid Press, P.O. Box 13608, El Paso, Texas 79913, U.S.A.
Abstract.—Species of Stigmatochomis are haplochromine cichlids endem-
ic to Lake Malaw
ˆi, and the genus currently includes four species, S. woodi
(type species), S. modestus, S. pholidophorus,andS. pleurospilus. The latter
two are known from only their holotypes. Herein, we describe two
additional species, Stigmatochromis macrorhynchos and Stigmatochromis
melanchros from southern Lake Malaw
ˆi.
Keywords: haplochromine cichlids, sand-dwelling cichlids
Historically, the Lake Malaw
ˆi sand-
dwelling cichlid genera have been diag-
nosed primarily on the basis of their
melanin pattern (Eccles & Trewavas
1989). Members of Stigmatochromis pos-
sess a supra-pectoral, supra-anal, and a
pre-caudal spot; the first two spots do not
extend to the base of the dorsal fin, as in
Hemitilapia, Trematocranus, and Tramiti-
chromis intermedius. The spot or blotch at
the nape and a series of small spots along
the base of the dorsal fin distinguish
Stigmatochromis from the spotted species
of Copadichromis, which lack such pig-
mentation. Currently, Stigmatochromis is
comprised of four species: S. woodi (type
species), S. modestus, S. pholidophorus,
and S. pleurospilus. Species of Stigmato-
chromis over 60 mm SL differ from those
of Otopharynx by a snout that is longer or
about as long as the post-orbital head
length, while the snout in Otopharynx is
always shorter. Stigmatochromis is further
characterized by a prognathous lower jaw
and by numerous unicuspid teeth in the
outer series of the oral jaws. In three of
the four species (not in S. pleurospilus)
there are 50 to 74 teeth in the outer row of
the upper jaw (Eccles & Trewavas 1989).
The unique holotype of Stigmatochromis
pleurospilus does not comply with this
diagnosis and we agree with Eccles &
Trewavas (1989) that it probably repre-
sents a juvenile of another species in the
genus. We have no doubt that the type is
a juvenile specimen, which is not only
suggested by the relatively large eye, but
also by its overall size. The purpose of
this paper is to describe two additional
species of Stigmatochromis from Lake
Malaw
ˆi.
Materials and Methods
Fishes were collected in Lake Malaw
ˆi
by chasing them into a monofilament
block net while SCUBA diving. All fishes
were anesthetized with clove oil, eutha-
nized in 1%formalin, pinned in trays so
that the bodies were flat and the fins
erect, preserved in 10%formalin, and
placed in permanent storage in 70%
ethanol. Pigmentation patterns and colors
were recorded in the field via direct
observation, photography, and videogra-
phy. Counts and measurements follow
Stauffer (1991) and Stauffer & Konings
(2006).
Morphometric data were analyzed us-
ing a sheared principal component anal-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
124(3):212–222. 2011.
ysis, which factors the covariance matrix
and restricts size variation to the first
principal component (Humphries et al.
1981, Bookstein et al. 1985). Meristic data
were analyzed using a principal compo-
nent analysis in which the correlation
matrix was factored. Differences among
species were illustrated by plotting the
sheared second principal components
(SPC2) of the morphometric data against
the first principal components (PC1) of
the meristic data (Stauffer & Hert 1992).
Institutional abbreviations follow Leviton
et al. (1985).
Results
The diagnoses and descriptions of S.
modestus,S. woodi (Table 1), and S.
pleurospilus (Table 2) were summarized
by Cleaver et al. (2009). Morphome-
tric and meristic data are given below
for all other Stigmatocrhomis species,
including the description of two addition-
al taxa.
Stigmatochromis pholidophorus
(Trewavas)
Fig. 1A, Table 2
Holotype.—BMNH 1935.6.14.1544,
85.7 mm SL, collected by Christy from
northern Lake Malaw
ˆi.
Diagnosis.—The presence of a supra-
pectoral and supra-anal spots, which do
not extend to the base of the dorsal fin,
plus a precaudal spot and a series of small
spots along the base of the dorsal fin
places this species in Stigmatochromis.
Stigmatochromis pholidophorus is further
diagnosed by the presence of mostly
unicuspid teeth in the outer rows of the
lower jaws. The short head length (33.3
vs. 36.3–39.8%SL) and the distance
between snout and dorsal-fin origin
(34.5 vs. 34.8–43.2%SL) distinguish this
species from all other Stigmatochromis
species.
Description.—Jaws prognathous; teeth
on jaws in 3 rows; teeth in outer row
unicuspid; 22 teeth in outer row of left
lower jaw; scale rows on cheek 4; lateral-
line scales 30; 1 pored scale posterior to
hypural plates. Gill rakers on first cerato-
branchial 11, with 4 on epibranchial.
Principal morphometric ratios and meris-
tics in Table 2.
Stigmatochromis macrorhynchos,
new species
Fig. 2, Table 3
Holotype (Fig. 2A).—PSU 4774, 128.2
mm SL, off Domwe Island, Lake Malaw
ˆi,
Malaw
ˆi, Africa 13u59.569S, 34u51.269E;
108 m deep, 7 Mar 1995, collected by the
Malaw
ˆi Fisheries Department.
Paratypes.—PSU 4775, 4, 116.6–124.8
mm SL, data as for holotype.
Diagnosis.—The presence of supra-pec-
toral and supra-anal spots, which do not
extend to the base of the dorsal fin,
combined with a snout length longer than
the post-orbital head length distinguishes
this species as a member of Stigmatochro-
mis. It is further diagnosed by the
presence of mostly unicuspid teeth in the
outer rows of the lower jaws. Stigmato-
chromis macrorhynchos has a narrower
interorbital width than S. melanchros
(13.7–16.2 vs. 19.3–20.5%HL) and a
shorter snout to pelvic-fin origin (38.4–
40.3%SL) than S. melanchros (42.2–
44.6%SL) and S. modestus (42.2–46.9%
SL). The distance between the dorsal fin
origin and the posterior anal fin insertion
is shorter in S. macrorhynchos (51.2–
51.8%SL) than in all other Stigmato-
chromis species (53.5–57.2%SL). The
distance between the anterior dorsal fin
to pelvic-fin origin of S. macrorhynchos
(27.7–30.0%SL) is shorter than in S.
modestus (30.3–33.8%SL), S. woodi
(31.9–38.1%SL) and S. melanchros
(35.7–36.1), and longer than in S. pholi-
dophorus (26.8%SL) and S. pleurospilus
(27.5%SL). The snout length of S.
macrorhynchos (38.0–41.2%HL) is longer
than in S. modestus (33.5–37.4%HL) and
S. pholidophorus (37.1%HL). The smaller
VOLUME 124, NUMBER 3 213
head depth of S. macrorhynchos (59.8–
63.6%HL) further distinguishes it from
S. woodi (64.1–75.8%HL) and S. melan-
chros (67.8–76.8%HL).
Description.—Jaws strongly progna-
thous (Fig. 2A); teeth on jaws in 3 rows;
teeth in outer row unicuspid; 21–26 teeth
in outer row of left lower jaw; scale rows
Table 1.—Morphometric and meristic values of Stigmatochromis modestus and Stigmatochromis woodi.
Stigmatochromis modestus (n57) Stigmatochromis woodi (n512)
Holotype X
¯Range Lectotype X
¯Range
Standard length, mm 120.0 93.6 67.3–124.1 173.2 151.0 104.0–173.2
Head length, mm 45.2 34.7 26.6–47.6 64.8 56.7 38.6–65.7
Percent standard length
Head length 37.7 38.8 37.7–39.8 37.4 37.5 36.4–38.3
Snout to dorsal-fin origin 39.6 41.2 39.2–43.2 39.8 39.7 34.8–41.8
Snout to pelvic-fin origin 42.2 44.8 42.2–46.9 38.5 40.5 38.5–44.6
Dorsal-fin base length 51.4 51.9 49.5–53.3 51.1 50.8 50.0–52.5
Anterior dorsal to anterior anal 45.1 43.8 42.0–45.9 43.3 45.5 43.0–48.2
Anterior dorsal to posterior anal 55.9 55.4 53.6–57.2 55.0 54.8 53.5–56.9
Posterior dorsal to anterior anal 27.6 27.3 25.9–28.0 28.6 28.7 26.3–30.0
Posterior dorsal to posterior anal 15.0 14.4 13.9–15.0 14.8 15.2 14.1–15.9
Posterior dorsal to ventral caudal 18.2 17.6 15.8–19.1 18.8 19.4 18.4–21.1
Posterior anal to dorsal caudal 18.7 19.3 18.5–21.7 20.9 21.6 20.8–23.5
Anterior dorsal to pelvic-fin origin 31.9 32.2 30.3–33.8 33.9 35.2 31.9–38.1
Posterior dorsal to pelvic-fin origin 48.1 49.0 47.2–50.7 50.8 50.5 49.5–51.9
Caudal-peduncle length 12.9 13.4 12.8–14.5 14.7 15.2 14.4–15.7
Least caudal-peduncle depth 12.2 11.8 11.4–12.3 10.4 11.0 10.1–11.6
Percent head length
Snout length 37.4 35.5 33.5–37.4 40.2 39.6 37.9–42.0
Postorbital head length 38.5 35.9 33.3–38.6 36.1 35.6 34.0–38.6
Horizontal eye diameter 27.9 30.2 26.7–33.8 28.3 28.8 27.0–32.6
Vertical eye diameter 25.6 27.5 23.6–31.1 24.6 24.8 22.8–28.1
Preorbital depth 22.9 19.3 17.1–22.9 26.5 24.8 22.6–26.5
Cheek depth 24.8 20.4 17.5–24.8 24.6 24.3 21.6–27.2
Lower-jaw length 48.3 46.4 45.2–48.3 42.1 43.6 42.1–47.0
Head depth 69.0 66.9 63.3–70.0 64.1 72.2 64.1–75.8
Interorbital width 19.5 17.0 14.7–19.5 15.8 16.5 14.5–20.0
Premaxillary pedicel length 27.4 25.2 23.3–27.4 23.9 23.9 21.3–26.2
Percent eye diameter
Preorbital depth/vertical eye diameter 89.5 71.2 55.2–89.5 98.4 100.4 80.4–112.0
Preorbital depth/horizontal eye diameter 82.0 64.6 50.8–82.0 89.3 86.6 69.3–94.9
Counts Mode Range Mode Range
Dorsal-fin spines 17 16 16–17 16 16 15–16
Dorsal-fin rays 11 11 10–12 10 10, 11 10–11
Anal-fin spines 3 3 3 3 3 3
Anal-fin rays 9 9 8–10 9 9 9
Pectoral-fin rays 13 13 12–13 13 13, 14 13–14
Pelvic-fin rays 5 5 5 5 5 5
Lateral-line scales 29 30 29–31 31 31 30–32
Pored scales posterior to lateral line 1 2 0–2 1 2 1–2
Scale rows on cheek 3 3 3 3 3 2–4
Gill rakers on first ceratobranchial 11 11 10–12 9 11 9–12
Gill rakers on first epibranchial 3 4 3–5 4 4 3–5
Teeth in outer row of left lower jaw 28 27 24–28 21 20 16–24
Tooth rows on upper jaw 3 3 3–4 3 3 2–3
Tooth rows on lower jaw 3 3 2–3 3 3 2–3
214 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Table 2.—Morphometric and meristic values of holotypes of Stigmatochromis pholidophorus and
Stigmatochromis pleurospilus.
Stigmatochromis pholidophorus Stigmatochromis pleurospilus
Standard length, mm 85.7 40.3
Head length, mm 28.5 13.5
Percent standard length
Head length 33.3 33.5
Snout to dorsal-fin origin 34.5 36.1
Snout to pelvic-fin origin 38.8 39.4
Dorsal-fin base length 51.6 53.0
Anterior dorsal to anterior anal 41.9 43.4
Anterior dorsal to posterior anal 54.4 56.5
Posterior dorsal to anterior anal 28 27.5
Posterior dorsal to posterior anal 14.6 14.3
Posterior dorsal to ventral caudal 18.1 18.1
Posterior anal to dorsal caudal 21.4 21.1
Anterior dorsal to pelvic-fin origin 26.8 27.5
Posterior dorsal to pelvic-fin origin 49.5 46.4
Caudal-peduncle length 16.6 17.0
Least caudal-peduncle depth 11.2 10.7
Percent head length
Snout length 37.1 20.7
Postorbital head length 33.1 35.8
Horizontal eye diameter 33.9 42.6
Vertical eye diameter 29.5 36.7
Preorbital depth 22.2 14.9
Cheek depth 16.7 15.9
Lower-jaw length 41.6 37.0
Head depth 69.8 73.5
Interorbital width 20.8 19.8
Premaxillary pedicel length 29.1 24.1
Percent eye diameter
Preorbital depth/vertical eye diameter 75.4 40.5
Preorbital depth/horizontal eye diameter 65.6 35.0
Counts
Dorsal-fin spines 16 15
Dorsal-fin rays 11 12
Anal-fin spines 3 3
Anal-fin rays 10 9
Pectoral-fin rays 13 14
Pelvic-fin rays 5 5
Lateral-line scales 30 29
Pored scales posterior to lateral line 1 1
Scale rows on cheek 4 3
Gill rakers on first ceratobranchial 11 9
Gill rakers on first epibranchial 4 3
Teeth in outer row of left lower jaw 22 17
Tooth rows on upper jaw 3 4
Tooth rows on lower jaw 3 3
VOLUME 124, NUMBER 3 215
on cheek 3; holotype with 33 lateral-line
scales; paratypes with 32–34; pored scales
posterior to hypural plates 1–2. Gill
rakers on first ceratobranchial 10–12,
with 4 on epibranchial. Principal mor-
phometric ratios and meristics in Table 3.
Males (Fig. 2B) with dark gray head
anteriorly fading to light gray/silver pos-
teriorly; pale yellow outline of lower jaw.
Laterally dark blue/gray dorsally, fading
to silver ventrally with 9 dark gray bars.
Scales with blue highlights. Dorsal fin pale
yellow with light yellow/white markings;
blue/white lappets with dark submarginal
band. Caudal fin light gray with blue/white
markings. Anal fin with black spines and
membranes; dark gray rays and mem-
branes with light red and yellow ocelli.
Pelvic fins black and pectoral fins clear.
Females with dark gray interorbital
and white/silver cheeks and operculum.
Laterally, silver with yellow highlights.
Dorsal fin clear with faint orange mar-
ginal bar and yellow highlights. Caudal
fin clear with yellow markings in mem-
branes. Anal fin clear with several yellow
Fig. 1. A, Stigmatochromis pholidophorus; adult individual (approximately 13 cm TL) at Chiofu Bay,
Malaw
ˆi in Lake Malaw
ˆi. B, territorial male Stigmatochromis woodi (approximately 22 cm TL) guarding his
sand bower at Luwala Reef, Malaw
ˆi, Lake Malaw
ˆi.
216 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
ocelli. Pelvic fin clear with white leading
edge; pectoral fin clear.
Etymology.—Specific epithet a noun in
apposition from Greek makros meaning
‘‘long’’ and rhynchos meaning ‘‘snout,’’
referring to the elongated snout of this
species.
Stigmatochromis melanchros, new species
Fig. 3, Table 3
Holotype (Fig. 3A).—PSU 4944, 160.73
mm SL, Mazinzi Reef, Lake Malaw
ˆi,
Malaw
ˆi, Africa; 13u539S, 34u539E, 10
Mar 1996, J. Stauffer.
Paratypes.—PSU 4945, 1, 154.15 mm
SL, Mazinzi Reef, Lake Malaw
ˆi, Malaw
ˆi,
Africa, 23 Mar 1996, J. Stauffer; PSU
4947, 1, 154.68 mm SL, Mazinzi Reef,
Lake Malaw
ˆi, Malaw
ˆi, Africa, 7 Mar
1995, J. Stauffer; PSU 4609, 1, 114.5 mm
SL, Mazinzi Reef, Lake Malaw
ˆi, 5 Mar
2002, J. Stauffer.
Diagnosis.—The presence of supra-pec-
toral and supra-anal spots, which do not
extend to the base of the dorsal fin, plus a
precaudal spot and a series of small spots
along the base of the dorsal fin, combined
with a snout length longer than the post-
orbital head length distinguishes this
species as a member of Stigmatochromis.
It is further diagnosed by the presence of
mostly unicuspid teeth in the outer rows
of the lower jaws. Stigmatochromis mel-
anchros is distinguished from S. macro-
Fig. 2. Stigmatochromis macrorhynchos. A, PSU4774, holotype, 128.2 mm SL, off Domwe Island,
Malaw
ˆi, Lake Malaw
ˆi. B, non-territorial male (approximately 12 cm TL) at Mumbo Island, Malaw
ˆi,
Lake Malaw
ˆi.
VOLUME 124, NUMBER 3 217
Table 3.—Morphometric and meristic values of Stigmatochromis macrorhynchos (n55) and
Stigmatochromis melanchros (n54).
Stigmatochromis macrorhynchos Stigmatochromis melanchros
Holotype X
¯Range Holotype X
¯Range
Standard length, mm 128.2 121.6 116.6–128.2 160.7 146 114.5–160.7
Head length, mm 46.5 45.6 44.5–46.5 60.9 54.7 43.4–60.9
Percent standard length
Head length 36.3 37.5 36.3–38.4 37.9 37.5 36.8–37.9
Snout to dorsal-fin origin 36.6 38.2 36.6–40.0 34.8 36.1 34.8–37.2
Snout to pelvic-fin origin 38.4 39.5 38.4–40.3 44.6 43 42.2–44.6
Dorsal-fin base length 49 48.8 48.4–49.3 50.3 50.3 49.4–51.4
Anterior dorsal to anterior anal 39.5 41 39.5–42.2 44.4 46.5 44.4–47.7
Anterior dorsal to posterior anal 51.3 51.5 51.2–51.8 54.2 55 54.2–56.6
Posterior dorsal to anterior anal 27.1 26.8 26.0–27.4 29.7 29.5 29.0–29.7
Posterior dorsal to posterior anal 14 14.3 14.0–14.8 14.8 15.2 14.8–15.5
Posterior dorsal to ventral caudal 20.4 19.4 19.0–20.4 18.4 19.7 18.4–21.1
Posterior anal to dorsal caudal 23.3 22.2 21.4–23.3 22.3 22.6 22.2–23.5
Anterior dorsal to pelvic-fin origin 27.7 28.7 27.7–30.0 35.7 36 35.7–36.1
Posterior dorsal to pelvic-fin origin 49.1 48.2 46.7–49.1 50.1 50.4 49.4–51.6
Caudal-peduncle length 18.1 17 16.0–18.1 15.1 15.1 13.5–16.2
Least caudal-peduncle depth 11 10.7 10.3–11.1 10.7 11.3 10.7–11.9
Percent head length
Snout length 38.9 39 38.0–41.2 38.7 38.6 36.2–40.2
Postorbital head length 35 34.1 32.4–35.0 37.5 37.7 37.0–38.3
Horizontal eye diameter 29.1 30.6 29.1–31.4 26 26 25.2–26.8
Vertical eye diameter 26.2 26.8 26.2–27.9 21 22.2 21.0–23.7
Preorbital depth 22 21.9 20.5–23.3 22.6 23 22.6–23.4
Cheek depth 20.8 19.8 18.6–20.9 23.5 24.8 23.5–26.3
Lower-jaw length 44 44.2 42.4–47.7 42.7 42.7 40.7–44.4
Head depth 62.9 62.1 59.8–63.6 76.8 72.8 67.8–76.8
Interorbital width 16.2 14.7 13.7–16.2 19.8 19.9 19.3–20.5
Premaxillary pedicel length 22.1 20.2 19.0–22.1 21.2 21.7 20.5–23.7
Percent eye diameter
Preorbital depth/vertical eye diameter 84.2 81.8 73.5–86.9 107.6 103.8 95.8–109.3
Preorbital depth/horizontal eye diameter 75.6 71.8 65.3–76.4 86.9 88.4 84.7–92.9
Counts Mode Range Mode Range
Dorsal-fin spines 16 16 15–16 16 16 15–16
Dorsal-fin rays 10 10 10–11 10 10, 11 10–11
Anal-fin spines 3 3 3 3 3 3
Anal-fin rays 8 9 8–9 8 8, 9 8–9
Pectoral-fin rays 13 13 13 13 13, 14 13–14
Pelvic-fin rays 5 5 5 5 5 5
Lateral-line scales 33 32, 33 32–34 32 31, 32 31–32
Pored scales posterior to lateral line 2 2 1–2 2 2 2–3
Scale rows on cheek. 3 3 3 3 3 3
Gill rakers on first ceratobranchial 11 11 10–12 12 12 9–12
Gill rakers on first epibranchial 4 4 4 4 4 3–5
Teeth in outer row of left lower jaw 21 21, 22,
23,
25, 26
21–26 13 13, 17, 18,
19
13–19
Tooth rows on upper jaw 3 3 2–3 4 3, 4 3–4
Tooth rows on lower jaw 3 3 2–3 3 3 3
218 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
rhynchos by a wider interorbital width
(19.3–20.5 vs. 13.7–16.2%HL) and from
S. pholidophorus by a larger preorbital
depth in relation to the vertical eye
diameter (95.8–109.3%vs. 75.4%). Stig-
matochromis melanchros is distinguished
from S. modestus by a shorter lower jaw
(40.7–44.4%HL vs. 45.2–48.3%HL) and
by fewer teeth in the outer row of the left
lower jaw (13–19 vs. 24–28). Stigmato-
chromis melanchros has a smaller hori-
zontal eye diameter (25.2–26.9%HL)
than S. woodi (27.0–32.6%HL).
Description.—Jaws prognathous (Fig.
3A); teeth on jaws in 3–4 rows; teeth in
outer row unicuspid; teeth in outer row of
left lower jaw 13–19; scale rows on cheek
3; holotype with 32 lateral-line scales;
paratypes with 31–32; pored scales poste-
rior to hypural plate 2–3. Gill rakers on
first ceratobranchial 9–12, with 3–5 on
epibranchial. Principal morphometric ra-
tios and meristics in Table 3.
Males (Fig. 3B) with anterior portion
of head dark gray and posterior portion
light gray/silver with green highlights.
Laterally, dark blue/black with scales
outlined in blue/green. Dorsal fin dark
gray with blue/green highlights and
orange and white tips. Caudal fin dark
gray with blue/white markings. Anal fin
black with pale yellow/white ocelli. Pelvic
fins black with white leading edge.
Pectoral fins with gray rays and clear
membranes.
Females (Fig. 3C) with gray interorbit-
al and gray/white cheeks and operculum.
Laterally, silver with pale orange outline
Fig. 3. Stigmatochromis melanchros. A, PSU4944, holotype, 160.7 mm SL, Mazinzi Reef, Malaw
ˆi, Lake
Malaw
ˆi. B, male (approximately 19 cm TL). C, female (approximately 17 cm TL) at Mazinzi Reef, Malaw
ˆi,
Lake Malaw
ˆi.
VOLUME 124, NUMBER 3 219
of scales; three lateral spots and a series of
black markings along base of dorsal fin.
Dorsal fin clear with faint orange mark-
ings. Caudal fin clear with gray markings
in membranes. Anal fin clear with several
yellow ocelli. Pelvic fin clear with white
leading edge; pectoral fin clear.
Etymology.—Specific epithet is a noun
in apposition derived from the Greek
melas, meaning ‘‘black’’ and from khros
meaning ‘‘skin’’ or ‘‘surface of body’’ in
reference to the overall black color of the
breeding males.
Remarks.—Stigmatochromis melan-
chros is a piscivore that breeds in rocky
reefs where territorial males defend the
vertical face of a large boulder near the
sandy bottom. This differs from the
sympatric S. woodi; males of which build
large bowers on the open sand away from
any rocks (Fig. 1B).
Discussion
Eccles & Trewavas (1989) originally
diagnosed Stigmatochromis based on the
melanin pattern that consisted of a small
supra-pectoral spot below the upper
lateral line, a small supra-anal spot
between the upper and lower lateral line,
and a spot near the end of the caudal
peduncle. Cleaver et al. (2009) expanded
on this diagnosis and stated that the
anterior two spots of Stigmatochromis do
not extend to the base of the dorsal fin,
which separates Stigmatochromis from
Hemitilapia, Trematocranus, and Tramiti-
chromis intermedius, which have spots
that extend to the base of the dorsal fin.
Additionally, the absence of a series of
small spots along the base of the dorsal
fin distinguishes Stigmatochromis from
spotted Copadichromis, which lack such
spots. Finally, they stated that in mem-
bers of Stigmatochromis greater than
60 mm SL, the snout is longer or equal
to the postorbital-head length, which
differentiates them from those of Oto-
pharynx, in which the snout length is
always shorter than the postorbital-head
length.
When the sheared second principal
components of the morphometric data are
plotted against the principal components of
the meristic data, the minimum polygon
clusters of the five species of Stigmatochro-
mis do not overlap apart from that of S.
woodi and S. melanchros (Fig. 4). The
latter two species occur sympatrically and
have distinguishable breeding strategies.
Males of S. woodi construct large spawn-
ing platforms in the sand (Fig. 1B), away
from the rocky substrate, whereas those of
S. melanchros defend the vertical face of a
large boulder that sits on the sand, usually
at the edge of a rocky reef.
Stigmatochromis pleurospilus is prob-
lematic as reported earlier (Cleaver et al.
2009). Eccles & Trewavas (1989) placed
this species in Stigmatochromis based on
pigment patterns. Examination of the
holotype reveals that it has bicuspid teeth
in the outer row of the lower jaw, which
we also found in some juveniles of S.
woodi and S. melanchros (non-type
material). A detailed examination of
the holotype (BMNH 1935.6.14.1475) re-
vealed that many of its scales are reversed,
possibly by maltreatment of the specimen
or that a deformed individual was pre-
served. We have retained S. pleurospilus
in Stigmatochromis, because at this time
we cannot justify its placement in any of
the other Lake Malaw
ˆi genera.
Comparison Material
Stigmatochromis woodi. Lectotype:
BMNH 1921.9.6.139–144, 173.1 mm SL;
Paralectotypes: BMNH 1921.9.6.139–144,
5, 103.99–170.2 mm SL; PSU 4610, 2,
171.7–174.0 mm SL, Mazinzi Reef, Lake
Malaw
ˆi, Malaw
ˆi, Africa, J. Stauffer, 7
Mar 1995; PSU 4946, 2, 147.3–162.6 mm
SL, Mazinzi Reef, Lake Malaw
ˆi, J.
Stauffer, 5 Mar 1996; PSU 4948, 2,
142.5–177.4 mm SL, Mazinzi Reef, Lake
Malaw
ˆi, J. Stauffer, 5 Mar 2002.
220 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Stigmatochromis modestus.Holotype:
BMNH 1893 1.17.5, 120.0 mm SL, Lake
Malaw
ˆi, Williams, 1891; PSU 4466, 6,
67.3–124.1 mm SL; Malaw
ˆi: Lake Malaw
ˆi:
Nametumbwe, 13u38.2909S, 34u51.3349E,
Stauffer & Konings, 25 Jan 2007.
Stigmatochromis pleurospilus. Holo-
type: BMNH 1935.6.14.1475, 40.3 mm
SL, Tanzania, Lupembe Sandbank,
Christy, 1925–1926.
Stigmatochromis pholidophorus. Holo-
type: BMNH 1935.6.14.1544, 85.7 mm
SL, Malaw
ˆi, Lake Malaw
ˆi, Vua, Christy,
1925–1926.
Key to the Species of Stigmatochromis
1a. Head length less than 35 SL . . . . . 2
1b. Head length greater than 35 SL . . . . 3
2a. Snout length less than 30%HL . . .
..................... S. pleurospilus
2b. Snout length greater than 30%HL . . .
..................... S. pholidophorus
3a. Snout to pelvic-fin origin less than
41%SL ....................... 4
3b. Snout to pelvic-fin origin greater
than 41%SL ................... 5
4a. Anterior dorsal to pelvic-fin origin less
than 31%SL ...... S. macrorhynchos
4b. Anterior dorsal to pelvic-fin origin great-
er than 31%SL ..... S. woodi (in part)
5a. Posterior dorsal fin to anterior anal
fin greater than 28.5%SL ....... 6
5b. Posterior dorsal fin to anterior anal fin
less than 28.5%SL ..... S. modestus
6a. Horizontal eye diameter 25.2–26.9%
HL ................. S. melanchros
6b. Horizontal eye diameter 27.0–32.6%
HL ............. S. woodi (in part)
Literature Cited
Bookstein, F., B. Chernoff, R. Elder, J. Humphries,
G. Smith, & R. Strauss. 1985. Morphometrics
in evolutionary biology.—Academy of Natu-
ral Sciences, Philadelphia, Special Publication
15:1–277.
Fig. 4. Second sheared principal components (morphometric data) plotted against first principal
components (meristic data) of Stigmatochromis spp.
VOLUME 124, NUMBER 3 221
Cleaver, R. M., A. F. Konings, & J. R. Stauffer, Jr.
2009. Two new cave-dwelling cichlids of Lake
Malaw
ˆi, Africa.—Ichthyological Explora-
tions of Freshwaters 20(2):163–178.
Eccles, D. H., & E. Trewavas. 1989. Malaw
ˆian
cichlid fishes: the classification of some
haplochromine genera. Lake Fish Movies,
Herten, Germany,
Humphries, J. M., F. L. Bookstein, B. Chernoff, G.
R. Smith, R. L. Elder, & S. G. Poss. 1981.
Multivariate discrimination by shape in rela-
tion to size.—Systematic Zoology 30(30):291–
308.
Leviton,A.E.,R.H.Gibbs,E.Heal,&C.E.Dawson.
1985. Standards in herpetology and ichthyolo-
gy: part I. Standard symbolic codes for
institutional resource collections in herpetology
and ichthyology.—Copeia 1985:802–832.
Stauffer, J. R., Jr. 1991. Description of a facultative
cleanerfish (Teleostei: Cichlidae) from Lake
Malaw
ˆi, Africa.—Copeia 1991(1):141–147.
———, & E. Hert. 1992. Pseudotropheus callainos,
a new species of mbuna (Cichlidae), with
analyses of changes associated with two intra-
lacustrine transplantations in Lake Malaw
ˆi,
Africa.—Ichthyological Explorations of
Freshwaters 3(3):253–264.
———, & A. F. Konings. 2006. Review of
Copadichromis (Telostei: Cichlidae) with the
description of a new genus and six new
species.—Ichthyological Explorations of
Freshwaters 17(1):9–42.
Associate Editor: Jeffrey T. Williams.
222 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
ResearchGate has not been able to resolve any citations for this publication.
Article
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A new species of the cichlid fish genus Pseudotropheus from Lake Malawi, Africa, is described. The new species superficially resembles Pseudotropheus lanisticola and P. livingstonii, but is clearly distinguished by head shape. The behavior pattern of cleaning ectoparasites from other cichlids is unique among all known species of Pseudotropheus.
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The status of the genus Copadichromis is discussed. Copadichromis prostoma and C. boadzulu are placed in Nyassachromis. Copadichromis cyclicos, C. thinos, C. conophoros, C. fl avimanus, C. inornatus, and C. eucinostomus are placed in Mchenga, new genus, diagnosed principally on morphology, bower construction, and habitat preference. In the remaining Copadichromis, three species groups are identifi ed, the C. quadrimaculatus group, the C. mbenjii group, and the C. virginalis group. Six new species are described in the C. mbenjii group: C. melas, C. chizumuluensis, C. diplostigma, C. insularis, C. cyanocephalus, and C. parvus.
Article
Full-text available
Six different populations of the Lake Malawi cave-dweller Otopharynx lithobates, each of them distinguished by differences in the male breeding coloration, were examined. We found that morphometric and meristic data of all populations overlap, and that no significant differences could be found to distinguish them. Morphological data of Otopharynx walteri were completely within the range of O. lithobates and we regard them as conspecific. Two other cave-dwelling cichlids are described: Otopharynx spelaeotes, new species, and Otopharynx antron, new species. No geographical variation in the males’ breeding colors of these species was observed. Otopharynx antron is described from a 20 km stretch of the eastern shore of the lake. It is sympatric with O. spelaeotes, but seems to have a more predatory feeding regime. Stigmatochromis modestus, another cave-dwelling species, and Stigmatochromis pleurospilus are compared to the three cave-dwelling members of Otopharynx
1 mm SL; MalawîMalawî: Lake MalawîMalawî: Nametumbwe, 13u38.2909S, 34u51.3349E
  • Stigmatochromis Modestus
  • Lake Holotype
  • Williams Malawîmalawî
Stigmatochromis modestus. Holotype: BMNH 1893 1.17.5, 120.0 mm SL, Lake MalawîMalawî, Williams, 1891; PSU 4466, 6, 67.3–124.1 mm SL; MalawîMalawî: Lake MalawîMalawî: Nametumbwe, 13u38.2909S, 34u51.3349E, Stauffer & Konings, 25 Jan 2007. Stigmatochromis pleurospilus. Holotype: BMNH 1935.6.14.1475, 40.3 mm SL, Tanzania, Lupembe Sandbank, Christy, 1925–1926. Stigmatochromis pholidophorus. Holotype: BMNH 1935.6.14.1544, 85.7 mm SL, MalawîMalawî, Lake MalawîMalawî, Vua, Christy, 1925–1926.
  • Mazinzi Reef
  • Lake Malawîmalawî
  • J Stauffer
Mar 1995; PSU 4946, 2, 147.3–162.6 mm SL, Mazinzi Reef, Lake MalawîMalawî, J. Stauffer, 5 Mar 1996; PSU 4948, 2, 142.5–177.4 mm SL, Mazinzi Reef, Lake MalawîMalawî, J. Stauffer, 5 Mar 2002.
1 mm SL; Paralectotypes: BMNH 1921
  • Material Stigmatochromis Woodi
  • Lectotype
Material Stigmatochromis woodi. Lectotype: BMNH 1921.9.6.139–144, 173.1 mm SL; Paralectotypes: BMNH 1921.9.6.139–144, 5, 103.99–170.2 mm SL; PSU 4610, 2, 171.7–174.0 mm SL, Mazinzi Reef, Lake MalawîMalawî, MalawîMalawî, Africa, J. Stauffer, 7
Lectotype: BMNH 1921.9.6.139-144, 173.1 mm SL
  • Stigmatochromis
Stigmatochromis woodi. Lectotype: BMNH 1921.9.6.139-144, 173.1 mm SL;
3-162.6 mm SL, Mazinzi Reef, Lake MalawîMalawî
Mar 1995; PSU 4946, 2, 147.3-162.6 mm SL, Mazinzi Reef, Lake MalawîMalawî, J. Stauffer, 5 Mar 1996; PSU 4948, 2, 142.5-177.4 mm SL, Mazinzi Reef, Lake MalawîMalawî, J. Stauffer, 5 Mar 2002.