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A New Species Of Ptychochromis From Northeastern Madagascar (Teleostei: Cichlidae), With An Updated Phylogeny And Revised Diagnosis For The Genus

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A new species belonging to the endemic Malagasy cichlid genus Ptychochromis is described. Ptychochromis ernestmagnusi is known only from the middle to lower reaches of the Mananara River in northeastern Madagascar. Based on a unique configuration of the palatine, the new species is placed among the species of Ptychochromis considered to have an eastern-type palatine morphology by Stiassny and Sparks (2006). The new species is further distinguished from members of the "western clade" of Ptychochromis by anterior displacement of the first supraneural such that it overlies the dorsoposterior margin of the supraoccipital. Among species of Ptychochromis with eastern-type palatine morphology, P. ernestmagnusi shares with P. makira the possession of supraneurals with a characteristically flattened dorsal profile. Although P. makira and P. ernestmagnusi both possess paired lateral barring pigmentation patterns, the lateral flank bars in P. makira are distinctively V-shaped, whereas in P. ernestmagnusi they are oriented vertically. Ptychochromis ernestmagnusi is further distinguished from P. makira by prominent iridescent spangling on the flank and dorsal fin near its base, dusky grayish-green base coloration (vs. whitish), four laterosensory foramina on the lachrymal (vs. three), and a series of seven (vs. six) infraorbital bones. In addition, an updated phylogeny and diagnosis for Ptychochromis is presented, including anatomical data for the new species and novel anatomical data for the genus.
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Accepted by J. Friel: 1 Dec. 2009; published: 22 Jan. 2010 33
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2010 · Magnolia Press
Zootaxa 2341: 3351 (2010)
www.mapress.com/zootaxa/Article
A new species of Ptychochromis from northeastern Madagascar (Teleostei:
Cichlidae), with an updated phylogeny and revised diagnosis for the genus
JOHN S. SPARKS & MELANIE L.J. STIASSNY
Department of Ichthyology, Division of Vertebrate Zoology, American Museum of Natural History, New York, NY 10024, USA.
E-mail: jsparks@amnh.org; mljs@amnh.org
Abstract
A new species belonging to the endemic Malagasy cichlid genus Ptychochromis is described. Ptychochromis
ernestmagnusi is known only from the middle to lower reaches of the Mananara River in northeastern Madagascar.
Based on a unique configuration of the palatine, the new species is placed among the species of Ptychochromis
considered to have an eastern-type palatine morphology by Stiassny and Sparks (2006). The new species is further
distinguished from members of the “western clade” of Ptychochromis by anterior displacement of the first supraneural
such that it overlies the dorsoposterior margin of the supraoccipital. Among species of Ptychochromis with eastern-type
palatine morphology, P. ernestmagnusi shares with P. makir a the possession of supraneurals with a characteristically
flattened dorsal profile. Although P. m ak ira and P. ernestmagnusi both possess paired lateral barring pigmentation
patterns, the lateral flank bars in P. m ak ira are distinctively V-shaped, whereas in P. ernestmagnusi they are oriented
vertically. Ptychochromis ernestmagnusi is further distinguished from P. makira by prominent iridescent spangling on
the flank and dorsal fin near its base, dusky grayish-green base coloration (vs. whitish), four laterosensory foramina on
the lachrymal (vs. three), and a series of seven (vs. six) infraorbital bones. In addition, an updated phylogeny and
diagnosis for Ptychochromis is presented, including anatomical data for the new species and novel anatomical data for
the genus.
Key words: Ptychochromis ernestmagnusi, Ptychochrominae
Introduction
Ptychochromis is endemic to fresh- and brackish-water habitats in northwestern, northern, and eastern
Madagascar and currently comprises eight valid species (Stiassny & Sparks, 2006). Historically, members of
the genus also occurred within the Onilahy River drainage basin in southwestern Madagascar (P. onilahy), as
well as other western basins, however, those populations are now presumed extinct (JSS, pers. obs.). Shortly
after the publication of a phylogenetic analysis and taxonomic revision of Ptychochromis (Stiassny & Sparks,
2006) additional specimens of an undescribed species collected from the middle to lower reaches of the
Mananara (du nord) River in northeastern Madagascar became available for study. The Mananara River is
located in a poorly sampled region of the island situated between the known geographic ranges of
Ptychochromis makira to the north and the widespread east coast species, P. grandidieri, to the south. The
presence there of a new species of Ptychochromis was not unanticipated given the inaccessibility and
consequent difficulty of conducting ichthyofaunal survey work in the region, which still contains intact tracts
of lowland rainforest (Rham & Nourissat, 2002; Stiassny & Sparks, 2006), and the new taxon is formally
described herein. Based on unique features of palatine morphology, the new species is placed among species
that have an eastern-type palatine (Stiassny & Sparks, 2006), which we refer to as the “eastern group” of
Ptychochromis, an assemblage comprised of Ptychochromis grandidieri Sauvage, 1882, P. m a kira Stiassny
and Sparks, 2006, P. loisellei Stiassny and Sparks, 2006, and P. curvidens Stiassny and Sparks, 2006.
Resolution of the intrarelationships of Ptychochromis is limited and no unambiguously derived anatomical
features have been identified to support monophyly of the “eastern group” of Stiassny and Sparks (2006: Fig.
SPARKS & STIASSNY34 · Zootaxa 2341 © 2010 Magnolia Press
2); however, the “western clade”, comprising P. oligacanthus (Bleeker, 1868), P. inornatus Sparks, 2002, P.
onilahy Stiassny and Sparks, 2006, and P. insolitus Stiassny and Sparks, 2006, was shown to be monophyletic
in that study. Here we present and analyze additional anatomical evidence unique to this “eastern group” of
five Ptychochromis species, generate an updated phylogeny for the genus with the inclusion of the new
species, and present a revised generic diagnosis.
Materials and methods
Counts and morphometric measurements follow Stiassny and Sparks (2006) and, for the new species, are
presented in Table 1. Specimens were cleared and stained (C&S) using a modified protocol based on Taylor
and Van Dyke (1985). Institutional abbreviations follow Leviton et al. (1985). Specimens examined in this
study are listed under Comparative materials at the end of this paper and are arranged alphabetically by
genus and species. Abbreviations used throughout the text are: C&S: cleared and stained preparations; ex.:
example(s); SL: standard length.
All methods of nucleotide sequence acquisition, sequence alignment, data treatment, and phylogeny
reconstruction (including search parameters under the optimality criterion of parsimony using direct
optimization) are as outlined in Sparks and Smith (2004) and Stiassny and Sparks (2006). The morphological
character matrix is presented in Table 2 and character state descriptions are listed in Appendix 1.
Systematic Accounts
Ptychochromis Steindachner, 1880
Type species: Ptychochromis oligacanthus (Bleeker, 1868)
Generic diagnosis. A ptychochromin genus diagnosed by the presence of a free second epibranchial
toothplate (Stiassny & Sparks, 2006: Fig. 7A), exclusion of the second infraorbital bone (IO2) from the orbit
by IO3 (Stiassny & Sparks, 2006: Fig. 8F), a prominent palatine groove (Stiassny & Sparks, 2006: Fig. 6F–
G), ventrally displaced insertion of the palatine-palatine ligament compared to other ptychochromin genera
(Stiassny & Sparks: Fig. 6F–G), a horizontally oriented palatine bone with an elongate palatine prong and
weakly elevated lateral ethmoid process (Fig. 2; Stiassny & Sparks, 2006: Fig. 6G; modified in members of
the “western clade”), and anterior displacement of the supraneural bones such that the anterior element comes
to overlie the dorsoposterior margin of the supraoccipital crest (Fig. 3A–C; lacking in members of the
“western clade”).
Note that due to a typographic error in our prior diagnosis for Ptychochromis (Stiassny & Sparks, 2006:
14) we erroneously state that IO3 is excluded from the orbital margin by IO2, however, this feature is
correctly described above (i.e., IO2 excluded from orbital margin by IO3). The remainder of the text,
character description, cladogram, figures, and dataset are correct with respect to the configuration, coding,
and interpretation of this feature (Stiassny & Sparks, 2006: character 11).
Ptychochromis ernestmagnusi, new species
Figures 1– 5; Table 1
Holotype: AMNH 249490, 146.6 mm SL, male, Mananara (du nord) River at Antanambaobe Village (16° 16
01S, 49° 40 01E), Madagascar, 120 m a.s.l., MG 10-06, coll. P.V. Loiselle, 4 October 2006.
Paratypes: AMNH 249488, 9 ex., incl. 1 ex. C&S, 75.9–99.5 mm SL, Mananara (du nord) River at
Antanibaolina Village (16° 15 01S, 49 ° 40 41E), Madagascar, 108 m a.s.l., MG 09-06, coll. P.V. Loiselle,
4 October 2006. AMNH 249489, 8 ex., incl. 1 ex. C&S, 51.5–99.0 mm SL, data as for holotype. MNHN
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NEW PTYCHOCHROMIN CICHLID FROM MADAGASCAR
2009-1674, 2 ex., 79.4–98.0 mm SL, data as for holotype. UMMZ 248823, 2 ex., 83.0–86.4 mm SL, data as
for AMNH 249488.
Non-Type Material Examined: MNHN 1935-0007, 1 ex., 128.5 mm SL, Mananara (du nord) River,
Madagascar. No additional collection locality information available.
FIGURE 1. Ptychochromis ernestmagnusi, AMNH 249490, holotype, adult male, 146.6 mm SL (scale bar = 1cm).
Diagnosis. A Ptychochromis exhibiting the eastern-type palatine morphology (Fig. 2) and further
distinguished from all congeners except P. grandidieri, P. loisellei, P. m akir a and P. curvidens by an anterior
displacement of the first supraneural such that it overlies the dorsoposterior margin of the supraoccipital (Fig.
3A– C). Among eastern group Ptychochromis, P. m aki ra shares with P. ernestmagnusi the possession of
supraneurals with a characteristically flattened dorsal profile, which is interpreted here as a synapomorphy
uniting these two geographically proximate species. Also shared with P. makir a is the presence of strong
(paired) lateral barring as a prominent component of pigmentation patterning; however, P. m a kira can be
distinguished by distinctively V-shaped lateral flank bars, whereas in P. ernestmagnusi the bars are oriented
vertically. Ptychochromis ernestmagnusi is further distinguished from P. makira by conspicuous iridescent
spangling on the flank and dorsal fin near its base, dusky grayish-green base coloration (vs. whitish), four
lachrymal laterosensory foramina (vs. three), and a total of seven (vs. six) infraorbital bones.
Description. Morphometric and meristic data presented in Table 1. External anatomical characteristics
and general pigmentation pattern in life and preservation can be observed in Figures 1, 5 and 7B. Moderately
deep bodied and laterally compressed. Dorsal body profile convex, becoming significantly more so in larger
specimens (ca. 100 mm SL and larger). Ventral body profile weakly to moderately convex. Lateral snout
outline straight in smaller specimens, and becoming weakly curved (convex) in larger individuals. Predorsal
profile moderately to strongly convex from mid-orbit to dorsal-fin origin, becoming more pronounced in
larger specimens and creating weak “nuchal hump”. Supraoccipital crest prominent in lateral view and
conspicuously deep bodied in larger individuals (ca. 100 mm SL). Caudal peduncle short, deep, and laterally
compressed. Origin of dorsal fin located well anterior to vertical through pectoral-fin insertion. Origin of
pelvic fin located considerably posterior to vertical through pectoral-fin insertion.
Total vertebral count 27 or 28, with formulae of 13 + 14 (mode), 13 + 15, and 14 + 14 precaudal and
caudal vertebrae, respectively.
Oral jaws isognathous and small. Oral dentition bilaterally symmetrical and bicuspid, with moderately to
well-developed distally expanded and slightly recurved cusps (Fig. 2). Outer row teeth of both premaxilla and
dentary enlarged relative to teeth of inner rows and graded in size (i.e., becoming smaller) posterolaterally.
SPARKS & STIASSNY36 · Zootaxa 2341 © 2010 Magnolia Press
Outer row teeth procumbently implanted in rostral portion of lower jaw, and oriented vertically elsewhere.
Outer row teeth in upper jaw more or less vertically oriented. Upper jaw with three or four rows of teeth
anteriorly and tapering to single (i.e., outer) row posteriorly. Lower jaw with three rows of teeth rostrally, and
tapering to single (i.e., outer) row posteriorly. Although smaller, inner rows of teeth on both premaxilla and
dentary of same morphology (bilaterally symmetrical and bicuspid) as those of respective outer row. Dentition
covers about anterior 2/3 of dentary and nearly entire surface (>80%) of premaxillary arcade.
TABLE 1. Morphometric and meristic data for Ptychochromis ernestmagnusi, new species. Proportional measurements
(mm) in percent standard length (SL) or percent head length (HL), unless noted otherwise. Values in parentheses indicate
number of specimens examined with that count. (H) indicates count corresponding to holotype.
P. ernestmagnusi
Character N Holotype Range Mean SD
Standard length (mm) 18 146.6 51.5–146.6 83.5
Percentage of SL
Head length 18 35.7 34.2–37.1 35.3 0.92
Body depth 18 46.1 41.7–46.1 43.9 1.40
Predorsal length 18 42.3 40.4–44.6 42.3 1.10
Preanal length 18 71.6 71.6–73.5 72.7 0.66
Prepelvic length 18 41.3 39.8–48.7 42.1 1.97
Head width (max.) 18 18.6 15.9–18.6 17.3 0.74
Caudal peduncle length 18 15.0 12.4–15.9 14.4 0.82
Caudal peduncle width 16 6.2 4.9–6.7 5.9 0.60
Caudal peduncle depth 18 16.5 13.3–16.5 15.2 0.85
Pectoral-fin length 18 31.0 29.8–34.1 32.1 1.24
Pelvic-fin length 18 30.9 24.3–31.2 28.7 1.76
Percentage of HL
Snout length 18 46.3 35.3–46.3 39.3 2.71
Orbit diameter 18 29.5 29.5–37.4 34.2 2.13
Upper-jaw length 18 37.7 26.3–37.7 30.1 2.69
Lower-jaw length 18 39.6 35.6–41.2 38.4 1.38
Interorbital width 18 32.1 25.6–32.1 28.3 1.55
Preorbital depth 18 28.7 20.0–28.7 24.6 2.24
Caudal peduncle length/depth 18 0.9 0.8–1.1 1.0 0.07
Caudal peduncle length/width 16 2.4 2.1–3.3 2.5 0.33
Scales in lateral line 16 34 (8), 35 (5, H), 36 (2), 37 (1)
Scales: lateral line to dorsal fin 18 4 (8), 4.5 (2), 5 (7, H), 6 (1)
Scales: pectoral to pelvic bases 18 4 (4), 5 (14, H)
Gill rakers (lower limb 1st arch) 18 11 (15, H), 12 (3)
Vertebrae (pre-caudal + caudal) 18 13+14 = 27 (15), 13+15 = 28 (1), 14+14 = 28 (2, H)
Dorsal fin 18 XII 13 (1), XIII 11 (1), XIII 12 (15, H), XIII 13 (1)
Anal fin 18 III 7 (2), III 8 (13, H), III 9 (3)
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NEW PTYCHOCHROMIN CICHLID FROM MADAGASCAR
FIGURE 2. Left lateral view of the oral jaws and anterior region of the suspensorium of P. ernestmagnusi (AMNH
249489, paratype, 87.5 mm SL, female, C&S) illustrating the “eastern type palatine” (shaded in gray) morphology.
Lower pharyngeal jaw (LPJ = fused 5th ceratobranchial elements) robust with interdigitating suture on
posteroventral margin. Dentition on LPJ and upper pharyngeal jaw (UPJ) comprised of numerous, closely set,
strongly hooked and bicuspid teeth. Cusps on LPJ teeth better developed posteriorly. Posteromedially on both
LPJ and third pharyngobranchial toothplate, dentition becoming robust and molariform (Fig. 4A); other teeth
of these elements hooked and bicuspid. Expansive second pharyngobranchial toothplate bearing five or six
rows of well-developed, hooked and bicuspid teeth. Two or three rows of hooked and bicuspid teeth present
on “free” second epibranchial toothplate. Fourth upper toothplate covered with numerous, closely-set rows of
smaller hooked and bicuspid teeth; teeth becoming progressively smaller and much less well developed
posteriorly. Strong concavity and associated sickle-like prong present on caudomedial margin of fourth upper
toothplate. Dorsal surface of fourth ceratobranchial elements bearing numerous robust, laterally expanded,
toothplates. Fourth ceratobranchial toothplates confluent with outer row gill rakers of these elements.
Dentition on fourth ceratobranchial toothplates unicuspid and more or less conical to weakly hooked and
bicuspid laterally, and becoming progressively more strongly hooked and bicuspid medially (similar to pattern
observed for lateral LPJ dentition). Contralateral fourth ceratobranchial elements bearing strong concavity
and associated prong (= hook) on medial margin.
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FIGURE 3. Left lateral view of the dorsoposterior margin of the neurocranium illustrating morphology of the
supraneural bones (in black) and their relationship to the supraoccipital bone. A) Ptychochromis ernestmagnusi, B) P.
makira, C) P. c urv ide ns, and D) P. oligacanthus. Arrows indicate degree of overlap of supraoccipital crest by anterior
supraneural.
Eleven or 12 relatively elongate gill rakers arrayed along lower limb of first arch, excluding raker in angle
of arch (Fig. 4B). Lower limb rakers of first gill arch denticulate dorsomedially, bearing numerous conical to
weakly hooked and bicuspid teeth. Nine weakly developed and triangular epibranchial gill rakers. Remaining
gill arches bearing short, robust, and strongly laterally expanded (particularly, distally near crown) rakers.
These rakers strongly denticulate dorsally, bearing numerous elongate and conical (becoming bulbous
apically) to weakly hooked and bicuspid teeth.
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NEW PTYCHOCHROMIN CICHLID FROM MADAGASCAR
FIGURE 4. Ptychochromis ernestmagnusi. (A) Dorsal view of the lower pharyngeal jaw (fused 5th ceratobranchial
elements) illustrating molariform caudomedial tooth morphology. Left lateral view of the: (B) lower limb of the first gill
arch (ceratobranchial 1 and hypobranchial 1) with gill rakers shaded in gray, and (C) lachrymal (pores shaded in gray)
and additional bones of the infraorbital series.
Flank squamation comprised of large, regularly imbricate, weakly ctenoid scales. Scale margins becoming
progressively more ctenoid posteriorly on flank. Ctenoid scales extend from about dorsal-fin origin (i.e.,
ranging from slightly anterior to somewhat posterior of fin insertion) dorsal to upper branch of lateral line and
somewhat posterior to pectoral-fin base below upper lateral line, to proximal portion of caudal fin. Scales on
SPARKS & STIASSNY40 · Zootaxa 2341 © 2010 Magnolia Press
anterior portion of nape and throughout head region cycloid. Scales on opercle and subopercle cycloid. Cheek
scales cycloid and comprising four rows; fourth (ventral) row frequently poorly developed and comprising
few scales. Snout, lachrymal, and anterior portion of interorbital region to about level of mid-orbit asquamate.
Anterior chest scales somewhat reduced in size and embedded. Scales extending onto caudal fin reduced in
size and ctenoid anteriorly, markedly smaller and cycloid posteriorly. Pored scales of lower branch of lateral
line frequently extending onto caudal fin (i.e., beyond hypural flexure) for one or two rows. Lateral-line scales
with well-developed canals, and numbering 34 to 37 (mode 34). Four or five (mode) scale rows between bases
of pectoral and pelvic fins. Four (mode) to six scales in diagonal from upper branch of lateral line to dorsal-fin
origin. No scale rows extending onto dorsal- and anal-fin membranes proximal to base of fins.
Dorsal fin with XII or XIII spines and 11 to 13 soft rays. Anal fin with III spines and seven to nine soft
rays. First anal-fin spine conspicuously short, whereas second and third spines elongate and more or less
similar in length. Distal margins of soft dorsal and anal fins becoming produced and tapered in larger
specimens; posteriorly margins reaching to about caudal-fin origin in smaller individuals (<80 mm SL) and
extending well beyond origin in larger specimens (Fig. 1). Pectoral fin elongate, deep bodied and paddle-like;
becoming tapered distally (i.e., dorsal rays much longer than ventral). Adpressed pelvic fin terminating well
before anal-fin origin in smaller specimens, and extending to about anal-fin origin in larger individuals (>90
mm SL). Caudal fin emarginate, trailing margins of upper and lower lobes becoming at most weakly produced
in larger individuals (>80 mm SL).
Miscellaneous osteology and anatomy. Exoccipital foramina on posterior of neurocranium poorly
developed; simple and lacking complex interior chamber (see Stiassny, 1991, and Sparks, 2008, for a
discussion of exoccipital foramen development in Malagasy-South Asian cichlids). Paired, anterior gas
bladder diverticula well developed, elongate, and tube-like; however, rather feeble in structure (i.e., not rigid
and thick walled) and similar to main gas bladder chamber. Diverticula in contact with exoccipital region of
neurocranium via connective tissue but not penetrating into exoccipital foramina (Sparks, 2008; Fig. 3A).
Infraorbital series composed of seven distinct elements (Fig. 4C). Lachrymal (= first infraorbital or IO1) with
four neurosensory pores; fourth pore communicating with anterior pore of second infraorbital (IO2). Second
infraorbital excluded from orbital margin by IO3. Cephalic laterosensory canals well developed with enlarged
pores (e.g., including those on preopercle and dentary; Figs. 2 & 4C). Uncinate process and anterior arm of
first epibranchial element short and robust (Fig. 4B). Well-developed process (= prong/hook) and deep
indentation (= excavation) present on medial face of fourth ceratobranchial element. Supraneurals (Fig. 3A)
rostrally positioned, with first supraneural overlying dorsoposterior margin of supraoccipital crest. Both
supraneurals characteristically shaped with flat dorsal margins (Fig. 3A). Posterior supraneural often reduced
in size or absent.
Coloration in life (Figure 5). Overall uniform greenish base coloration with a dusky gray overlay, not
notably darker dorsally than ventrally. Many scales bearing a small, conspicuous iridescent spot along
posterior scale margin. Pigmentation pattern composed of five or six (mode, holotype) prominent midlateral
blotches intersected by six to eight less strongly pigmented vertical bars. Nape dark gray, snout and cheek
dusky grey, and gular region black. Fins uniformly dark blackish-gray, with some small iridescent spots
proximally in soft dorsal. Pectoral fin hyaline.
Coloration in preservation (Figure 1). Ground coloration reddish-brown, slightly paler ventrally than
dorsally. Traces of iridescent spangling present, particularly in larger specimens, and most evident ventrally
on flank. Pigmentation pattern consisting of five or six (mode, holotype) prominent midlateral blotches with
significantly paler intersecting vertical bars retained in preservation. Most posterior blotch, located on caudal
peduncle (i.e., sixth blotch in series if present), significantly paler than others. Fins pale reddish-brown,
trailing margins of soft anal and dorsal fins blackish. Pectoral fin hyaline. Pelvic fin pale reddish-brown, and
becoming charcoal to blackish distally. Anterior interorbital region, snout, and lachrymal dark gray. Lower lip
creamy brown. Gular region dark grayish-black.
Distribution and habitat (Figure 6). Currently known only from the type localities, which are located in
the middle to lower reaches of the Mananara (du nord) River, northeastern Madagascar. This region is
characterized by humid, lowland rainforest (UNESCO - MAB Biosphere Reserves Directory, 2009), and
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NEW PTYCHOCHROMIN CICHLID FROM MADAGASCAR
remains poorly surveyed for freshwater fishes. It is unknown whether the new species is more widely
distributed within the region, and also whether it occurs in smaller tributaries of the Mananara (du nord) River
or other adjacent drainage basins to the north and south. Substantial forest cover remains in the region and it is
likely the new species has a significantly more widespread geographic distribution than current collection data
would indicate. The Mananara (du nord) River is a relatively large basin that extends from its headwaters
through regions of intact forest and low population density. It is a typical eastern drainage, with a steep overall
profile, and a generally rocky to sandy substrate. It is likely that the new species of Ptychochromis is restricted
to the middle to lower reaches of the river and its tributaries, given that suitable habitat and adequate trophic
resources are most likely lacking at higher elevations, as is typical for other eastern basins (Sparks, 2005a, b).
FIGURE 5. Ptychochromis ernestmagnusi, AMNH 249490, holotype, 146.6 mm SL, adult male. Freshly captured
specimen illustrating adult coloration in life. Photo by Paul Loiselle.
Conservation status. Although forested portions of the coastal region to the south of the Mananara (du
nord) River are encompassed by the Mananara-Nord Biosphere Reserve (designated in 1990), which extends
from 16° 09' to 16° 36'S and 49° 31' to 49° 53'E, and covers 140,000 hectares, the Mananara River and its
south bank tributaries are not included within the terrestrial component of the protected area (which also
includes a smaller marine component) (UNESCO - MAB Biosphere Reserves Directory, 2009). We anticipate
that the new species is not only more widespread in the region due to similar available habitats, but that it also
receives some degree of protection from habitat degradation, deforestation, and overfishing within the
biosphere reserve, which protects some of the last remaining remnants of lowland rainforest in eastern
Madagascar. In general, species of Ptychochromis seem able to tolerate a moderate degree of habitat
degradation (in particular P. grandidieri, which remains relatively common and widespread along the highly
developed and densely populated eastern coast of the island). Nevertheless, members of the genus are rapidly
extirpated from areas where habitats have become severely disturbed and water quality is negatively impacted
(e.g., severe deforestation and development resulting in highly turbid drainage basins) (Sparks & Stiassny,
2003, 2008).
Etymology. Named in honor of Mr. Ernest Magnus of Berlin, Germany, and New York City, at the request
of the family of Dr. Rudolph G. Arndt, whose support of ichthyological exploration and research at the
American Museum of Natural History is gratefully acknowledged.
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FIGURE 6. Relative geographic distributions of species of Ptychochromis exhibiting the “eastern type palatine”
configuration, including the geographically proximate sister species, P. mak ir a and P. ernestmagnusi, new species. Note:
Arrow for P. grandidieri indicates that distribution for species extends southward along the eastern coast of Madagascar.
Discussion and comparisons. Stiassny and Sparks (2006) corroborated the monophyly of, and resolved
intergeneric relationships within, the endemic Malagasy cichlid subfamily Ptychochrominae based on the
analysis of a combination of nucleotide sequence data and anatomical features, but resolution of relationships
within Ptychochromis was problematic in that study. Based primarily on features of the palatine bone of the
suspensorium, Stiassny and Sparks (2006) recognized two groups of Ptychochromis: a “western clade”
characterized by a compact palatine head with an upright orientation and marked elevation of the lateral
ethmoid process (i.e., “western type palatine”), and an “eastern group” in which the palatine exhibits a
markedly horizontal orientation, an elongate palatine prong, and less prominent (weakly elevated) lateral
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NEW PTYCHOCHROMIN CICHLID FROM MADAGASCAR
ethmoid process (i.e., “eastern type palatine”). On the cladogram presented by Stiassny and Sparks (2006: Fig.
2), unambiguous optimization of the “western type palatine” configuration (Sparks & Stiassny, 2006:
character 14, node D) was interpreted as evidence for monophyly of a “western clade”; however, the “eastern
type palatine” morphology described above was not unambiguously optimized on that phylogenetic
reconstruction. As a result, the relationships of the four eastern species included in our 2006 analysis (viz.,
Ptychochromis grandidieri, P. m aki ra, P. loisellei, and P. curvidens) were represented as a polytomy (Stiassny
& Sparks, 2006: Fig. 2).
Here we have identified an additional putatively homologous anatomical feature unique to the “eastern
group”, rostral displacement of the supraneural elements such that the anterior supraneural comes to overlie
the dorsoposterior margin of the supraoccipital of the neurocranium. This supraneural configuration is lacking
in all other members of Ptychochrominae, and, therefore, is hypothesized to represent an apomorphic feature
diagnostic of an “eastern clade” of Ptychochromis (e.g., Fig. 3A–C).
To test this hypothesis, we reran our prior phylogenetic analysis (Stiassny & Sparks, 2006: Fig. 2) using
the same methodological approach, search parameters, and combined morphological and molecular dataset,
with the inclusion of the new species and including coding for the unique “eastern group” supraneural
configuration described above and the two features (discussed below) hypothesized to unite P. maki ra and the
new species (for a total of 24 morphological transformations and 2053 total characters; characters 1–21 are
described in detail in Sparks & Stiassny, 2006: Table 1 and Results). A complete list of the morphological
character descriptions is presented in the Appendix and the corresponding data matrix is presented in Table 2.
As the results indicate (Fig. 7), we again failed to recover a monophyletic “eastern group” despite the fact that
these five species possess both the novel supraneural feature (character 22, Fig. 3A–C) and “eastern type
palatine” configuration (Sparks & Stiassny, 2006: character 21; Fig. 2) discussed above.
TABLE 2. Morphological character matrix for species of Ptychochromis and outgroups included in phylogenetic
analysis. Inapplicable character assignments are designated by (-). “A” = character states 0 & 1. Character state
descriptions presented in Appendix 1.
Characters
12345678910 11 12 13 14 15 16 17 18 19 20 21 22 23 24
Paretroplus damii 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0
Retroculus 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 1 1 0 0 0 0 0
Heterochromis 0 0 0 0 0 0 1 0 0 0 1 1 0 0 0 1 0 1 0 - 0 0 0 0
Paratilapia polleni 0 0 0 0 0 0 0 0 0 0 A 0 0 0 0 0 1 1 A - 0 0 0 0
Oxylapia polli 1 A 1 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 A 0 0 0 0 0
Ptychochromoides betsileanus 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 1 1 0 0 1 0 0 0 0
Ptychochromoides itasy 1 1 1 1 1 0 0 0 0 0 0 0 0 0 1 1 1 0 0 1 0 0 0 0
Ptychochromoides vondrozo 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 1 1 0 0 1 0 0 0 0
Katria katria 1 1 1 1 1 1 1 1 1 0 0 0 0 0 1 1 0 1 1 1 0 0 0 0
Ptychochromis oligacanthus 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 0 0 0 0
Ptychochromis insolitus 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 0 0 0 0
Ptychochromis onilahy 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 0 0 0 0
Ptychochromis inornatus 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0
Ptychochromis grandidieri 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 1 1 1 1 1 0 0
Ptychochromis makira 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 1 1 1 1 1 1 1 1
Ptychochromis loisellei 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 1 1 1 1 1 0 0
Ptychochromis curvidens 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 1 1 1 1 1 0 0
Ptychochromis ernestmagnusi 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 1 1 1 1 1 1 1
SPARKS & STIASSNY44 · Zootaxa 2341 © 2010 Magnolia Press
FIGURE 7. Lateral pigmentation pattern in preservation for: A) Ptychochromis makira (AMNH 237131, holotype,
151.0 mm SL, adult male), and B) P. ernestmagnusi (AMNH 249488, paratype, 99.5 mm SL, adult). Dashed lines
indicate relative adult lateral barring pattern and orientation.
Among the species of Ptychochromis that exhibit an “eastern type palatine” configuration, P. ma kira
shares with P. ernestmagnusi the possession of supraneurals with a characteristically flattened dorsal profile
(Fig. 3A, B, character 23). Also shared with P. makira is the presence of a characteristic lateral barring pattern,
in which each prominent midlateral blotch is intersected by a pair of narrow and less strongly pigmented
vertical or oblique bars (Fig. 7; character 24). However, in P. ma k ira these pairs of lateral flank bars are
distinctively V-shaped (Fig. 7A), whereas in P. ernestmagnusi the bars are vertically oriented (Fig. 7B).
Despite inconclusive results in our earlier phylogenetic analysis (Sparks & Stiassny, 2006: Fig. 2), in which P.
makira was recovered in a polytomy with other “eastern group” species of Ptychochromis, our new analysis
recovered a sister group relationship between P. m aki ra and P. ernestmagnusi supported by the two
unambiguously optimized anatomical features (characters 23 & 24) discussed above.
Although similar in many respects, P. ernestmagnusi is readily distinguished from P. maki ra by
conspicuous iridescent spangling on the flank and dorsal fin membrane near its base, dusky grayish-green
Zootaxa 2341 © 2010 Magnolia Press · 45
NEW PTYCHOCHROMIN CICHLID FROM MADAGASCAR
base coloration (vs. whitish), four lachrymal laterosensory foramina (vs. three), and a total of seven (vs. six)
infraorbital bones. Although only two specimens of P. makira have been collected to date and admittedly
represent a rather limited size range, the new species is also more shallow bodied (41.7–46.1 vs. 48.1–48.9%
SL in P. makir a), has a larger eye (29.5–37.4 vs. 25.6–27.6% HL in P. ma kira ), a longer head (34.2–37.1 vs.
32.0–32.2% SL in P. maki ra), a shorter preanal length (71.6–73.5 vs. 75.1% SL in P. mak ira), and a greater
lateral line scale count (34–37 vs. 33 in P. makira) than its sister taxon, P. m akira.
FIGURE 8. Strict consensus cladogram of nine optimal topologies recovered (tree length = 1018, CI = 0.67, RI = 0.61)
based on the simultaneous analysis of the nucleotide dataset of Stiassny and Sparks (2006) and 24 morphological
transformations. Unambiguously optimized morphological features supporting recovered nodes are designated by solid
(unique feature) and open (homoplasious feature) circles. Character numbers (above braches) and states (below
branches) correspond to the morphological transformations listed in Table 2 and Appendix 1. Indices in bold at nodes
indicate jackknife support. Character state descriptions and statistics are listed in Appendix 1. The morphological
character matrix is presented in Table 2.
Comparative materials. Values following catalog numbers indicate count of specimens examined, and
do not necessarily correspond to the total number of individuals in that particular lot:
Katria katria: AMNH 217739, holotype, eastern Madagascar, Tamatave Province, River Nosivolo, below
Zule’s Village, large side-pool off mainstream. AMNH 93701, 20 ex., 10 ex. C&S, eastern Madagascar,
SPARKS & STIASSNY46 · Zootaxa 2341 © 2010 Magnolia Press
Tamatave Province, River Nosivolo below Ampasimaniona Village, 26 km east-northeast of Marolambo.
UMMZ 240358, 1 ex. C&S, eastern Madagascar, Marolambo.
Oxylapia polli: AMNH 97111, 10 ex., 1 ex. C&S, eastern Madagascar, Tamatave Province, Mangoro
drainage, village of Marolambo, Nosivolo River. AMNH 97150, 4 ex., 1 ex. C&S, eastern Madagascar,
Tamatave Province, River Nosivolo by Ambarimasina Village, ca. 16 km east northeast of Marolambo.
UMMZ 235046, 1 ex. C&S, eastern Madagascar, Tamatave Province, Nosivolo River, near village of
Marolambo, Mangoro drainage.
Paratilapia polleni: AMNH 216068, 25 ex., eastern Madagascar, large baylake, behind dunes 1 km south
of turnoff from Marolambo-Mananjary road, ca. 100 meters from sea. UMMZ 235043, 2 ex. C&S,
northeastern Madagascar, Lac Anjavibe, Nosy be. UMMZ 235045, 2 ex. C&S, southeastern Madagascar,
Sahapindra River, near Vevembe.
Ptychochromis onilahy: MNHN 1962-0201, holotype, southwestern Madagascar, Province of Tulear,
Onilahy River, A. Kiener. AMNH 237130, paratype, 1 ex. (C&S in part), data as for holotype. MNHN 2006-
0780, paratypes, 3 ex., data as for holotype.
Ptychochromis makira: AMNH 237131, holotype, northeastern Madagascar, Antalaha Province, north of
Maroansetra, near town of Marovonona, Antainambalana River, purchased from local fishermen by Augustin
Sarovy, J. S. Sparks, W. L. Smith, and K. L. Tang. AMNH 237132, paratype, 1 ex. (C&S in part), data as for
holotype.
Ptychochromis loisellei: AMNH 232462, holotype, male, northeastern Madagascar, Antalaha Province,
north of Sambava, Mahanara River at Antsirabe-Nord, just upstream of bridge over route N-5 (13° 58.49S;
49° 57.81E), PVL-01-29, P.V. Loiselle and local fishermen. AMNH 231249, paratype, 1 ex., northeastern
Madagascar, Antalaha Province, main channel of the Mahanara River at Antsirabe-Nord, at bridge on Route
N-5 (13° 38.49S 49° 57.81E), PVL-00-07, P.V. Loiselle. AMNH 231258, paratypes, 3 ex., 1 ex. C&S,
northeastern Madagascar, Antalaha Province, main channel of the Mahanara River at Antsirsabe-Nord, at
bridge on Route N-5 (13° 58.49S 49° 57.81E), PVL-00-12, P.V. Loiselle. MNHN 2006-0781, paratype, 1 ex.,
data as for AMNH 231258. AMNH 232458, paratype, 1 ex., northeastern Madagascar, Antalaha Province,
Mahanara River, ca. 4 km northwest of Antsirabe-Nord (13° 57.30S 49° 56.20E), PVL-01-27, P.V. Loiselle
and local fishermen. MHNG 2676.095, paratype, 1 ex., data as for AMNH 232458. AMNH 237135,
paratypes, 5 ex., data as for holotype.
Ptychochromis curvidens: MHNG 2623.82, holotype, northern Madagascar, Antsiranana (Diego Suarez)
Province, Andranofanjava, Andranofanjava-Sandriapiana River system, P. de Rham and J.-C. Nourissat.
MHNG 2676.096, paratypes, 2 ex., data as for holotype. AMNH 237133, paratypes, 2 ex., 1 ex. C&S, data as
for holotype. MHNG 2623.84, paratype, 1 ex., northern Madagascar, Antsiranana (Diego Suarez) Province,
Mirosolava, P. de Rham and J.-C. Nourissat.
Ptychochromis insolitus: UMMZ 237066, holotype, juvenile; northeastern Madagascar, Antalaha
Province, near town of Mandritsara, Sofia drainage basin, Amboaboa (= Ambomboa) River (15° 50 1S; 48°
42 51E), J. S. Sparks and K. J. Riseng.
Ptychochromis inornatus: UMMZ 237492, holotype, adult female, northwestern Madagascar, Antalaha
Province, northeast of Antsohihy, Ankofia drainage, Anjingo River (14º 50 41.0S, 48º 14 38.3E), JSS 94-
19, J. S. Sparks, K. J. Riseng, and local Malagasy guides. UMMZ 237063, paratypes, 5 ex., 1 ex. C&S, data as
for holotype. AMNH 230746, paratypes, 2 ex., data as for holotype. UMMZ 237064, paratypes, 5 ex., 2 ex.
C&S, northwestern Madagascar, Antalaha Province, northeast of Antsohihy, Ankofia drainage, Bora Special
Reserve, Bemahavony River (tributary of Anjingo River) (14º 52 20.4S, 48º 14 52.2E), JSS 94-20. AMNH
230747, paratypes, 2 ex., data as for UMMZ 237064. UMMZ 237065, paratype, 1 ex., northwestern
Madagascar, Antalaha Province, northeast of Antsohihy, Ankofia drainage, Lake Andrapongy (14º 41 49.3S,
48º 07 54.3E), JSS 94-21. UMMZ 237067, paratypes, 7 ex., northwestern Madagascar, Antalaha Province,
northeast of Antsohihy, Ankofia drainage, Anjingo River (14º 50 39.7S, 48º 14 39.5E), JSS 94-54.
Ptychochromis grandidieri: MNHN A.4147, holotype, Madagascar, region of high forests, Humblot and
Grandidier. See discussion in Sparks (2003) regarding the locality of the holotype. Additional Non-Type
Material Examined: MNHN A.310, 1 ex., rivers that cross the eastern slope, Lantz. AMNH 88018, 56 ex., 1
Zootaxa 2341 © 2010 Magnolia Press · 47
NEW PTYCHOCHROMIN CICHLID FROM MADAGASCAR
ex., C&S, southeastern Madagascar, Mananjary, estuary of Mananjary River, 21º05S 48º27E. AMNH 88053,
2 ex., southeastern Madagascar, Mananjary, estuary of Mananjary River, 21º 05S, 48º 27E. AMNH 88076, 2
ex., eastern Madagascar, Vatomandry, 19º 20S, 49º 00E. AMNH 88090, 3 ex., eastern Madagascar,
Mahanoro, 19º 55S, 48º 50E. AMNH 88092, 17 ex., eastern Madagascar, Mahanoro, Pangalanes canal north
of Mangoro River. AMNH 88102, 36 ex., 14 ex. C&S, eastern Madagascar, Baylake behind dunes, ca. 100 m
from sea. AMNH 88117, 18 ex., eastern Madagascar, Tamatave market, 18º 10S, 49º 25E. AMNH 88140, 1
ex., eastern Madagascar, 25 km north of Tamatave, Pangalanes canal, 18º 00S, 49º 25E. AMNH 88153, 11
ex., eastern Madagascar, between Fenerive and Tamatave, Pangalanes canal, 18º 00S, 49º 25E. AMNH
96999, 52 ex., 2 ex. C&S, eastern Madagascar, Tamatave Province, Mangoro River near mouth. AMNH
97008, 185 ex., eastern Madagascar, Salehy village, 1 km south of turnoff from Marolambo-Mananjary road,
19º 55S, 48º50E. AMNH 97012, 12 ex., eastern Madagascar, Mahanoro market. AMNH 97028, 7 ex., 3 ex.
C&S, eastern Madagascar, Tamatave Province, Bay Lake behind first dune ca. 100 m from sea, east of road by
Sahey Village, 1 km south of turnoff from Marolambo-Manajary Road. AMNH 97057, 1 ex., eastern
Madagascar, Ambodisovoka village, Savalany River. AMNH 228067, 3 ex., southeastern Madagascar,
Lopary, Mananizo River. AMNH 228072, 6 ex., southeastern Madagascar, Ampataka village, Sahambavy
River. AMNH 228074, 3 ex., southeastern Madagascar, 12 km north of Farafangana, Manampatrona River,
22º 43 47S, 47º 47 25E. AMNH 231347, 1 ex., southeastern Madagascar, Ampataka village, Sahambavy
River, 23º 21’ 04”S, 47º2818E. AMNH 231352, 4 ex., southeastern Madagascar, Manombo Special
Reserve, Takoandra River, 23º 01 27S, 47º 43 16E. MNHN A.7896, 2 ex., central Madagascar to the west
of Antananarivo, Lac Itasy(?). MNHN 1901-0020, 1 ex., eastern Madagascar, Tamatave. MNHN 1901-0021,
1 ex., eastern Madagascar, Tamatave. MNHN 1932-0082, 1 ex., eastern Madagascar, Manompana. MNHN
1932-0083, 13 ex., eastern Madagascar, Manompana. UMMZ 233524, 17 ex., 2 ex. C&S, Madagascar,
southeastern coastal region. UMMZ 237311, 22 ex., southeastern Madagascar, Mananjary. UMMZ 237312, 3
ex., 1 ex. C&S, southeastern Madagascar, Manombo Special Reserve. UMMZ 237495, 5 ex., southeastern
Madagascar, 6 km north of Karianga at Mahavelo, Rienana drainage, Andriambondro River, 22º 21 47S, 47º
22 05E. UMMZ 238453, 2 ex., southeastern Madagascar, near Manombo Special Reserve. UMMZ 238471,
1 ex., southeastern Madagascar, Mananjary port. UMMZ 238472, 11 ex., 2 ex. C&S, southeastern
Madagascar, Farafangana market. UMMZ 238476, 7 ex., Madagascar, southeastern coastal region.
Ptychochromis oligacanthus: RMNH 3.936, lectotype, “Madagascar, in flumine Samberano, Nossibé, in
lacu Pambilao”, Pollen and van Dam. Additional Non-Type Material Examined: AMNH 18841, 1 ex.,
Madagascar (probably mainland, Sambirano region). AMNH 58491, 9 ex., northwestern Madagascar, Lake
Amparihibe, at the mouth of the inflowing small stream, Lake Antsidihy, Nosy Be. AMNH 215522, 4 ex.,
northwestern Madagascar, Lake Bemapaza, Nosy Be. AMNH 215523, 15 ex., northwestern Madagascar,
Lakes Djabala and Ampombilava, Nosy Be. AMNH 230699, 3 ex., northwestern Madagascar, Lake
Andjavibe, Nosy Be. AMNH 232399, 2 ex., northwestern Madagascar, Lake Ampombilava, Nosy Be. AMNH
232415, 3 ex., northwestern Madagascar, Lake Djabala, Nosy Be. MNHN 1962-322, 1 ex., northwestern
Madagascar, Sambirano River. UMMZ 236591, 26 ex., 4 ex. C&S, northwestern Madagascar, Lake
Ampombilava, Nosy Be. UMMZ 237498, 22 ex., 2 ex. C&S, northwestern Madagascar, Lake Djabala, Nosy
Be. UMMZ 237493, 3 ex., northwestern Madagascar, Lac de Deux Soeurs, Nosy Be. UMMZ 237494, 1 ex.,
northwestern Madagascar, Lake Amparihibe, Nosy Be. UMMZ 237496, 6 ex., northwestern Madagascar,
Lake Bempazava, Nosy Be. UMMZ 237497, 8 ex., northwestern Madagascar, Lake Anjavibe, island of Nosy
Be. UMMZ 237499, 11 ex., 1 ex. C&S, northwestern Madagascar, Mananjeba drainage, Andranomaloto
River, northeast of town of Ambanja.
Ptychochromoides betsileanus: BMNH 1882.2.25:69, lectotype, Betsileo, Madagascar. BMNH
1882.2.25:70, paralectotype, Betsileo, Madagascar. AMNH 217753, 1 ex., southwestern Madagascar, Ilanana
River, near Ranohira, Onilahy drainage. AMNH 217763, 1 ex., south-central Madagascar, Manantanana
River, headwaters near Iaritsena, Ambalavao Region, Mangoky drainage. UMMZ 238115, 5 ex., dry
skeletons, south-central Madagascar Ilanana River, south of Isalo National Park.
Ptychochromoides itasy: AMNH 2336643, paratype, 1 ex., Madagascar. MNHN 1919-11, paratype, 1 ex.
C&S, Madagascar, central highlands, Region of Antananarivo, Lake Itasy.
SPARKS & STIASSNY48 · Zootaxa 2341 © 2010 Magnolia Press
Ptychochromoides vondrozo: AMNH 228488, paratypes, 2 ex., southeastern Madagascar, Fianarantsoa
Province, Region of Vondrozo, near Village of Vevembe, Mananara (du sud) drainage. UMMZ 235294,
paratypes, 3 ex., 1 ex. C&S, southeastern Madagascar, Fianarantsoa Province, Region of Vondrozo, near
Village of Vevembe, Mananara (du sud) drainage.
Acknowledgments
We are extremely grateful to P. Loiselle (WCS) who collected a majority of the material upon which this
description is based. Thanks to P. Loiselle, P. de Rham, and the late J.-C. Nourissat for freely sharing their
collections, field data, and knowledge of Malagasy cichlids. We are grateful to R. Schelly (AMNH) and W.L.
Smith (FMNH) for contributing to various aspects of this study. For the loan or gift of specimens in their care
we are grateful to W. Fink, G. Smith, and D. Nelson (UMMZ), D. Siebert (BMNH), G. Duhamel, P. Pruvost,
and R. Causse (MNHN), C. Weber and S. Fisch-Muller (MHNG), and M. van Oijen (RMNH). For curatorial
assistance with materials at the AMNH and for assistance with radiographs, we are grateful to B. Brown, R.
Arindell, and R. Schelly. Collecting efforts in Madagascar were facilitated by the efforts of B. Andriamahaja
and the MICET (Institute for the Conservation of Tropical Environments, Madagascar) staff, and we are
grateful for their continued support of ichthyological research. Collection permits were obtained from the
Direction des Eaux et Forêts and the Association National pour la Gestion des Aires Protégées (ANGAP),
Madagascar. This study was supported by a generous gift from Dr. Rudolph G. Arndt and by an NSF award to
JSS (IOS-0749943). Finally, our thanks to two anonymous reviewers for helpful comments on the manuscript.
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SPARKS & STIASSNY50 · Zootaxa 2341 © 2010 Magnolia Press
APPENDIX 1. Character numbers and states are as listed in Table 2. Values in brackets correspond to
consistency index (CI) and retention index (RI) in the format [CI, RI].
Character 1. Median frontal neurocranial laterosensory foramina (NLF0): [1.00, 1.00]
State 0 = pores broadly coalesced in midline.
State 1 = pores widely separated.
Character 2. Oral jaw dentition: [1.00, 1.00]
State 0 = unicuspid, tricuspid, or asymmetrically bicuspid.
State 1 = bilaterally symmetrical and bicuspid.
Character 3. Anteriormost dorsal-fin pterygiophore: [1.00, 1.00]
State 0 = lacking procurrent spur.
State 1 = procurrent spur present.
Character 4. Supraneural shape: [1.00, 1.00]
State 0 = anterior supraneural straight with at most weak dorsoanterior projection or hook.
State 1 = anterior supraneural in shape of an inverted “L”.
Character 5. Anal-fin spine configuration: [0.50, 0.75]
State 0 = graded in size, becoming progressively longer posteriorly.
State 1 = extremely short first anal-fin spine followed by two elongate spines of more or less equal length.
Character 6. Inner margin of fourth ceratobranchial bone: [1.00, 1.00]
State 0 = smooth.
State 1 = bearing an elongate process/prong and deeply rounded indentation.
Character 7. Cephalic laterosensory canals and pores: [0.50, 0.83]
State 0 = not enlarged.
State 1 = markedly enlarged and inflated.
Character 8. Supraneural pattern: [1.00, 1.00]
State 0 = first and second supraneural separated by first neural spine.
State 1 = both first and second supraneurals located anterior to first neural spine.
Character 9. Lateral ethmoid process of palatine: [1.00, 1.00]
State 0 = process absent or at most weakly developed.
State 1 = process well developed, expansive, and elevated in lateral view.
Character 10. “Free” second epibranchial toothplate: [1.00, 1.00]
State 0 = toothplate absent.
State 1 = separate (“free”) toothplate present.
Character 11. Second infraorbital (IO2) configuration: [0.50, 0.80]
State 0 = IO2 included in orbit margin.
State 1 = IO2 excluded from orbital margin by IO3.
Character 12. Palatine groove: [0.50, 0.85]
State 0 = groove lacking on palatine head.
State 1 = deep groove present on palatine head.
Character 13. Insertion of palato-palatine ligament: [1.00, 1.00]
State 0 = palato-palatine ligament inserts directly or via a small dorsally-elevated process onto dorsal face of
contralateral palatine elements.
State 1 = insertion of palato-palatine ligament ventrally displaced onto small nub-like process on dorsolateral face of
contralateral palatine elements.
Zootaxa 2341 © 2010 Magnolia Press · 51
NEW PTYCHOCHROMIN CICHLID FROM MADAGASCAR
Character 14. Palatine with compact head and elevated lateral ethmoid process, both with an upright, vertical orientation
(“western type palatine” configuration): [1.00, 1.00]
State 0 = absent.
State 1 = present.
Character 15. Scale ctenoidy: [0.33, 0.50]
State 0 = flank scales cycloid or weakly ctenoid; only central portion of each scale margin bearing small cteni.
State 1 = flank scales strongly ctenoid, with entire caudal scale margin bearing well-developed cteni.
Character 16. Number of infraorbital elements in series (including lachrymal): [0.40, 0]
State 0 = 6 infraorbital bones.
State 1 = 7 infraorbital bones.
State 2 = 5 infraorbital bones.
Character 17. Anterior gas bladder morphology: [0.33, 0.33]
State 0 = anterior gas bladder horns absent or weakly developed and not approaching exoccipital region of
neurocranium.
State 1 = paired anterior gas bladder horns (= diverticula) elongate and contacting exoccipital region of
neurocranium.
Characer 18. Palatine splint: [0.50, 0.75]
State 0 = palatine splint lacking.
State 1 = palatine splint present.
Character 19. Dentition on second pharyngobranchial (Pb2) toothplate: [0.50, 0.66]
State 0 = single row of teeth on Pb2.
State 1 = two or more rows of teeth on Pb2.
Character 20. Configuration of fourth ceratobranchial (Cb4) toothplates: [0.50, 0.50]
State 0 = Cb4 toothplates separate from gill rakers of outer row.
State 1 = Cb4 toothplates confluent (= fused) with gill rakers of outer row.
Character 21. Palatine with horizontally oriented head, elongate anterior prong, weakly elevated lateral ethmoid process,
and posterior palatine lamina more or less horizontally aligned (“eastern type palatine” configuration): [0.50, 0.75]
State 0 = absent.
State 1 = present.
Character 22. Supraneural elements rostrally displaced or shifted, such that anterior supraneural overlies dorsoposterior
margin of supraoccipital crest: [0.50, 0.75]
State 0 = absent.
State 1 = present.
Character 23. Dorsal surface of supraneural elements: [1.00, 1.00]
State 0 = curved and convex in lateral view.
State 1 = dorsal profile characteristically flattened in lateral view.
Character 24. Characteristic lateral barring pattern on flank, such that pair of narrow and considerably less strongly
pigmented bars intersect each prominent midlateral blotch: [1.00, 1.00]
State 0 = absent.
State 1 = present.
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