Ptychochromis, STIASSNY & SPARKS, 2006

STIASSNY, MELANIE L. J & SPARKS, JOHN S, 2006, Phylogeny and Taxonomic Revision of the Endemic Malagasy Genus Ptychochromis (Teleostei: Cichlidae), with the Description of Five New Species and a Diagnosis for Katria, New Genus, American Museum Novitates 3535 (1), pp. 1-56 : 15-37

Ptychochromis sp. ‘‘sud-ouest Riv. Onilahy’’: de

Rham and Nourissat, 2002. Ptychochromis n. sp. ‘‘onilahy’’: Sparks, 2003. Ptychochromis nov. sp. ‘‘kotro/onilahy’’: Sparks and Stiassny, 2003: table 9.1.

HOLOTYPE: MNHN 1962-0201, 84.3 mm SL; southwestern Madagascar, Province of Tulear , Onilahy River, A. Kiener.

PARATYPES: AMNH 237130, 1 ex. (C&S in part), 76.8 mm SL; data as for holotype . MNHN 2006-0780, 3 ex., 66.4–85.8 mm SL; data as for holotype .

DIAGNOSIS: A Ptychochromis exhibiting the western-type palatine morphology (fig. 12A) and distinguished from all congeners by the presence of a lachrymal plate that extends

TABLE 2 Loadings of Morphometric Variables in Sheared Principal Components Analysis for Species of Ptychochromis Exhibiting the Western Type Palatine Configuration (figs. 6F, 9)

ventrally to cover a portion of the upper lip. It is further distinguished from all congeners except P. oligacanthus by the presence of cycloid scales above the upper lateral line on the flanks, and only two inner rows of teeth in both upper and lower jaws. Ptychochromis onilahy is distinguished from P. oligacanthus by a higher number of lateral line scales (33– 34 vs. 29–31 in P. oligacanthus ), a shorter head length (32.8–34.7 vs. 35.9–40.0% SL in P. oligacanthus ). Additionally, P. onilahy lacks all trace of the black blotch or bar covering the dorsoposterior margin of the opercle, diagnostic of P. oligacanthus .

DESCRIPTION: Morphometric and meristic data presented in table 4. Morphological characteristics and general pigmentation pattern in preservation can be observed in figure 11 and plate 2A. Body deep and laterally compressed. Lateral snout outline straight. Predorsal head profile convex from midorbit to dorsal-fin origin. Dorsal body profile from dorsal-fin origin smoothly convex. Ventral body profile more or less straight. Caudal peduncle short, deep, and laterally compressed. Dorsal-fin origin located anterior to vertical through pectoral-fin insertion. Pelvic-fin origin located well posterior to vertical through pectoral-fin insertion.

Total vertebral count 26 or 27, with a formula of 13 + 13 or 13 + 14 precaudal and caudal vertebrae, respectively.

Jaws isognathous. Oral dentition bilaterally symmetrical and bicuspid, with crowns somewhat expanded distally and slightly recurved. Outer row teeth of both jaws enlarged and graded in size laterally. Rostrally, outer row teeth procumbently implanted in lower jaw but only slightly so in upper jaw. Cusps well developed, particularly rostrally. Rostrally, upper and lower jaws with two inner rows of smaller teeth of the same morphology as those of outer rows. Total of three rows of teeth along rostral margin of both upper and lower jaws, tapering to a single row of teeth posteriorly. Dentition covers anterior half of dentary and nearly entire surface of premaxillary arcade. Uniquely with lachrymal bone on snout extending ventrally to cover a portion of upper lip, whereas in congeners the upper lip is fully exposed below lachrymal plate.

Lower pharyngeal jaw (LPJ) with a weakly interdigitating suture on posteroventral margin. Dentition on LPJ and upper pharyngeal toothplates (UPJ) composed of numerous, closely set, hooked, and bicuspid teeth. Medially on UPJ and posteromedially on LPJ, dentition becoming robust and molariform (fig. 12B). Four rows of hooked and bicuspid teeth on second pharyngobranchial toothplate. Three rows of teeth medially on ‘‘free’’ second epibranchial toothplate.

Lower limb rakers of first gill arch denticulate dorsomedially. Nine or ten (modally 10) triangular and somewhat elongate gill rakers (fig. 12C) arrayed along lower limb of first arch. Seven to nine epibranchial gill rakers. Rakers on remaining arches short, laterally expanded, and strongly denticulate. Teeth elongate and conical. Robust toothplates present on dorsal surface of fourth ceratobranchial bones; toothplates confluent with gill rakers of outer row of these elements. Teeth on fourth ceratobranchial toothplates unicuspid and conical laterally to hooked and bicuspid medially.

Body covered with large, regularly imbricate, weakly ctenoid scales from about level of midorbit to proximal portion of caudal fin. Scales above upper lateral line to about level of origin of soft dorsal fin cycloid. Scales on flanks becoming increasingly ctenoid posteriorly, but at most only weakly so. Scales on nape and head region cycloid. Four or five rows of cycloid scales on cheek. Scales on opercle and subopercle missing in all specimens. Anterior chest scales reduced in size and embedded. Scales on chest weakly ctenoid, except anteriorly where they are cycloid. Scales on belly cycloid. Snout, lachrymal, and anterior portion of interorbital region asquamate. Scales on caudal fin reduced in size. Lateral line scales 33 or 34. Five scales in diagonal from lateral line to dorsal-fin origin. Four or five scale rows between bases of pectoral and pelvic fins. No scale rows present along dorsal- and anal-fin bases.

Dorsal fin with XIII spines and 12 soft rays. Anal fin with III spines and eight or nine soft rays. First anal spine very short, second and third spines elongate and of similar length. All fins damaged, making determination of orig-

TABLE 3 Loadings of Morphometric Variables in Sheared Principal Components Analysis for Species of Ptychochromis Exhibiting the Eastern Type Palatine Configuration (figs. 6G, 10)

inal fin shape impossible. Pelvic fins short, extending at most to origin of anal fin.

MISCELLANEOUS OSTEOLOGY AND ANA- TOMY: Well-developed exoccipital foramina present on posterior of neurocranium. Paired anterior gas bladder extensions well developed but feeble, in contact with exoccipital region of neurocranium via connective tissue, but do not extend into exoccipital foramina. Infraorbital series composed of seven elements: lachrymal with four pores (fig. 12D), IO2 excluded from orbit by ventrally displaced IO3. Uncinate process and anterior arm of first epibranchial bone short and robust (fig. 12C). Well-developed process and deep indentation present on inner face of Cb4.

COLORATION IN LIFE: According to Kiener (1963) and Kiener and Maugé (1966), adults of this taxon were uniformly gray brown, paler ventrally and lacking any obvious body markings.

COLORATION IN PRESERVATIVE: Ground coloration brownish to golden-brown, lighter ventrally. All fins brownish, pectoral fins lightest and somewhat hyaline.

All five specimens appear quite faded in preservation, although a faint banding pattern is discernable in most specimens, whereas in two specimens a faded series of midlateral blotches remains.

DISTRIBUTION AND HABITAT: Ptychochromis onilahy is currently known only from the type series that was collected in the Onilahy River, a major westward flowing basin in southwestern Madagascar (fig. 1). Although detailed locality information is lacking, these specimens were collected by Kiener and almost certainly correspond to the ‘‘forme’’ of P. oligacanthus identified by Kiener and Maugé (1966) from the southwest of Madagascar. According to Kiener (1963) this taxon was particularly abundant in the Onilahy River to Ambohimahavelona (Lower Onilahy), and Kiener and Maugé (1966) indicate that it occurred, historically at least, from the Lower Onilahy River in the south, and northward in coastal basins to a little south of the Mangoky River.

CONSERVATION STATUS: Despite a number of collecting expeditions to southwestern Madagascar in recent years, no additional specimens referable to P. onilahy have been collected and the species may be extinct. However, many areas of southwestern Madagascar have not been thoroughly surveyed for freshwater fishes, and the Onilahy River basin is a large drainage system, thus, the possibility exists that remnant populations of P. onilahy persist in more remote areas within the region.

LOCAL NAME: According to Kiener (1963), in the Onilahy region P. onilahy was referred to as boramany, whereas fishermen around Lake Ihotry referred to the species as kotro.

ETYMOLOGY: The species is named after the basin in which the type, and only known,

TABLE 4 Morphometric and Meristic Data for Ptychochromis onilahy , new species Values in parentheses indicate number of specimens examined with that count. (H) indicates count corresponding to holotype.

specimens were reportedly collected. Specific epithet used as a noun in apposition.

DISCUSSION AND COMPARISONS: In sharing the western-type palatine morphology (fig. 6F, Character 14:1), Ptychochromis onilahy is recovered in a ‘‘western clade’’ of Ptychochromis (fig. 2), whose members are restricted to western drainages (fig. 1). In overall appearance it is most similar to P. oligacanthus from which it is readily distinguished by lateral line scale count (33 or 34 vs. 29–31 in P. oligacanthus ) and some proportional measurements (e.g., HL 32.8–34.7% SL vs. 35.9–40.0% SL in P. oligacanthus ). Unfortunately, no suitable tissue sample is available for inclusion in molecular phylogenetic studies and morphological character data gathered to date are not informative at this level of analysis; therefore, the sister group to P. onilahy is not known.

Ptychochromis makira , new species

Figures 13–14 , Plate 2B, Table 5

Ptychochromis sp. ‘‘Makira’’: Sparks and Smith, 2004: fig. 1, table 1.

HOLOTYPE: AMNH 237131, adult, 146.2 mm SL; northeastern Madagascar, Toamasina 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, Nov. 2003.

PARATYPE: AMNH 237132, 1 ex. (C&S in part), 112.5 mm SL; data as for holotype .

DIAGNOSIS: A Ptychochromis exhibiting the eastern-type palatine morphology (fig. 14A) and distinguished from all congeners by the presence of three (vs. four) laterosensory pore foramina on the lachrymal (the posteriormost [5 third] foramen forms part of the marginal canal between the lachrymal and second infraorbital bone; fig. 14D) and a unique pigmentation pattern consisting of four distinctive V-shaped black bars on the flanks superimposed on an overall whitish base. Ptychochromis makira is further distinguished from all congeners, except P. insolitus (with five), in the possession of a total of six (rather than seven) infraorbital bones.

DESCRIPTION: Morphometric and meristic data presented in table 5. Morphological characteristics and general pigmentation pattern can be observed in figure 13 and plate 2B. Extremely deep-bodied and laterally compressed. Dorsal body profile strongly curved, particularly in holotype. Ventral body profile moderately convex in paratype and relatively straight in holotype. Lateral snout outline straight to mildly curved. Predorsal head profile moderately convex from midorbit to dorsal-fin origin. Supraoccipital crest prominent and deep in larger specimen. Caudal peduncle short, deep, and laterally compressed. Dorsal-fin origin located slightly anterior to vertical through pectoral-fin insertion. Pelvic-fin origin located well posterior to vertical through pectoral-fin insertion.

Total vertebral count 27, with a formula of 13 + 14 precaudal and caudal vertebrae, respectively.

Jaws isognathous. Oral dentition bilaterally symmetrical and bicuspid, with moderately to well-developed distally expanded and slightly recurved cusps. Outer row teeth of both jaws enlarged and graded in size laterally. Rostrally, outer row teeth procumbently implanted in lower jaw but only slightly so in upper jaw. Rostrally, both upper and lower jaws with three inner rows of smaller teeth of the same morphology as those of outer rows. Total of four rows of teeth along rostral margin of both upper and lower jaw, tapering to a single row of teeth posteriorly. Dentition covers anterior 2/3 of dentary and nearly entire surface of premaxillary arcade.

LPJ robust with weakly interdigitating suture on posteroventral margin. Dentition on LPJ and UPJ comprised of numerous, closely set, hooked, and bicuspid teeth. Cusps on LPJ better developed posteriorly. Medially on both LPJ and third pharyngobranchial, dentition robust and molariform (fig. 14B). Five or six rows of hooked and bicuspid teeth on expansive second pharyngobranchial toothplates. Three rows of teeth present medially on ‘‘free’’ second epibranchial toothplate. Robust, laterally expanded, toothplates cover majority of dorsal surface of fourth ceratobranchial bones; toothplates confluent with outer row gill rakers of these elements. Dentition on fourth ceratobranchial toothplates unicuspid (and generally conical) or weakly hooked and bicuspid laterally, hooked and bicuspid medially (similar to lateral LPJ dentition).

Lower limb rakers of first gill arch denticulate dorsomedially. Eleven somewhat elongate gill rakers arrayed along lower limb of first arch. Nine epibranchial gill rakers. Gill rakers on remaining arches short, laterally expanded, and strongly denticulate. Teeth elongate and conical, somewhat bulbous distally.

Body covered with large, regularly imbricate, weakly ctenoid scales. Ctenoid scales extend from just posterior to dorsal-fin origin above upper lateral line and slightly anterior to pectoral-fin base below upper lateral line, to proximal portion of caudal fin. Scales on nape

TABLE 5 Morphometric and Meristic Data for Ptychochromis makira , new species Values in parentheses indicate number of specimens examined with that count. (H) indicates count corresponding to holotype.

and head region cycloid. Scales on opercle and subopercle also cycloid. Cheek scales cycloid and comprising four rows. Anterior chest scales slightly reduced in size and embedded. Snout, lachrymal, and anterior portion of interorbital region to about level of midorbit asquamate. Scales on caudal fin reduced in size and ctenoid anteriorly, markedly smaller and cycloid posteriorly. Lateral line scales 33. Five scale rows between bases of pectoral and pelvic fins. Five scales in diagonal from lateral line to dorsal-fin origin. No scale rows present along dorsal- and anal-fin bases.

Dorsal fin with XIII spines and 12 soft rays. Anal fin with III spines and eight or nine soft rays. First anal spine very short, second and third spines elongate and similar in length. Caudal fin emarginate, trailing margins of upper and lower lobes slightly produced. Pectoral fin elongate and tapered distally. Distal margins of soft dorsal and anal fins produced and pointed in holotype (5 larger specimen). Trailing margin of soft dorsal fin elongate and filamentous in holotype. Pelvic fin not extending to anal-fin origin in smaller specimen (5 paratype), slightly beyond origin in holotype.

MISCELLANEOUS OSTEOLOGY AND ANA- TOMY: Well-developed exoccipital foramina present on posterior of neurocranium. Paired anterior gas bladder extensions in contact with exoccipital region of neurocranium via connective tissue, but do not extend into exoccipital foramina. Infraorbital series (fig. 14D) composed of six elements. Although infraorbital number is extremely variable within Cichlidae (see Stiassny, 1991; Kullander, 1998), a total of seven infraorbital elements is resolved as the plesiomorphic condition for Cichlidae and is most commonly encountered among ptychochromins. A reduction to six is interpreted as a derived condition within Ptychochromis (Character 16, fig. 2). Lachrymal deep, with only three pores (fig. 14D), IO2 excluded from orbit by ventrally displaced IO3. Uncinate process and anterior arm of first epibranchial bone short and robust (fig. 14C). Well-developed process and deep indentation present on inner face of Cb4.

COLORATION IN LIFE: Unknown. Coloration and pigmentation pattern notes were taken within a couple days of preservation (plate 2B) and live coloration is probably similar to that reported below for preserved state.

COLORATION IN PRESERVATIVE: Ground coloration pale creamy white to yellow. Four black V-shaped bands present on flanks, which extend ventrally to lateral midline. Body much lighter ventrally, and essentially uniform pale creamy yellow. Fins pale yellow to grayish; fin rays dark gray to black. Fins, excluding pectorals, blackish terminally. Pectoral fin olive proximally and mostly hyaline distally. Anterior interorbital region, snout, and lachrymal grayish-green. Lips light olive. Scales on chest and ventral flanks, primarily rostral of anal-fin origin, brown along exposed margins.

DISTRIBUTION AND HABITAT: Ptychochromis makira is known only from the type series, collected in the Antainambalana River just north of Maroansetra in northeastern Madagascar (fig. 1). The southern range limit of P. makira is not known with certainty, but it is possible that P. makira is conspecific with populations of similarly pigmented Ptychochromis reported by local fishermen to occur as far south as the coastal towns of Manompana and Mananara.

CONSERVATION STATUS: Although once common throughout the region, P. makira has suffered a severe decline in abundance in recent years, according to local fishermen. Little original riparian vegetation remains in the lower reaches of the Antainambalana River basin. Fishing pressure is high in the region surrounding Maroansetra and habitat degradation has been widespread throughout the Antainambalana basin. Local fishermen report that the species is now rare.

LOCAL NAME: Saroy is the Malagasy name used throughout eastern Madagascar to refer to species of Ptychochromis .

ETYMOLOGY: The species is named after the region in which the type specimens were collected. The specific epithet, makira , is used as a noun in apposition.

DISCUSSION AND COMPARISONS: In our phylogenetic analysis, P. makira is recovered in a polytomy and its immediate relationships remain unresolved (fig. 2). Although there is currently no unambiguous evidence for the monophyly of an assemblage comprising P. makira , P. grandidieri , P. loisellei , and P. curvidens , the shared possession of an easterntype palatine morphology is unique to these four species and may indicate a close relationship among them. Molecular data analyzed alone indicates that P. makira is the sister taxon to the remaining species of Ptychochromis ( Sparks and Smith, 2004) . Regardless, morphometrically P. makira is most similar to P. grandidieri from which it is readily distinguished by pigmentation pattern and coloration, infraorbital morphology, and also in the possession of a considerably smaller LPJ with fewer and less well-developed molariform teeth (compare fig. 14B and fig. 24B).

Ptychochromis loisellei , new species

Figures 15–16 , Plates 1C and 2C, Table 6

Ptychochromis sp. ‘‘nord-est’’: de Rham and Nourissat, 2002.

Ptychochromis n. sp. ‘‘green garaka’’: Sparks, 2003.

Ptychochromis nov. sp. ‘‘green garaka’’: Sparks and Stiassny, 2003.

Ptychochromis sp. ‘‘Garaka’’: Sparks and Smith, 2004: fig. 1, table 1.

HOLOTYPE: AMNH 232462, 99.5 mm SL, male; northeastern Madagascar, Antsiranana 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, 1 Nov., 2001.

PARATYPES: AMNH 231249, 1 ex., 108.5 mm SL; northeastern Madagascar, Antsiranana 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, 9 Oct., 2000. AMNH 231258, 3 ex., 1 ex. C&S, 81.3–106.5 mm SL; northeastern Madagascar, Antsiranana 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, 13 Oct., 2000. MNHN 2006-0781, 1 ex., 97.2 mm SL; data as for AMNH 231258; AMNH 232458, 1 ex., 118.6 mm SL; northeastern Madagascar, Antsiranana 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, 31 Oct., 2001. MHNG 2676.095, 1 ex., 109.2 mm SL; data as for AMNH 232458. AMNH 237135, 5 ex., 39.3–50.8 mm SL; data as for holotype .

DIAGNOSIS: A Ptychochromis exhibiting the eastern-type palatine morphology (fig. 16A) and distinguished from all congeners by a unique pigmentation pattern and coloration comprising an expansive (generally vertically oriented) black band or blotched region of black pigment below the lateral midline and just anterior to origin of (and often extending over) the anal fin, and by an overall dark grayish-green body coloration. Ptychochromis loisellei is further diagnosed by an apomorphic lateral expansion of the LPJ toothplate anteriorly, which obscures the underlying bone in dorsal view (fig. 16B).

DESCRIPTION: Morphometric and meristic data presented in table 6. Morphological characteristics and general pigmentation pattern can be observed in figure 15 and plates 1C and 2C. Moderately deep-bodied and laterally compressed. Snout markedly acute and pointed. Lateral snout outline straight to moderately curved. Dorsal body profile moderately curved posterior to dorsalfin origin. Ventral body profile straight. Predorsal head profile moderately convex from midorbit to dorsal-fin origin. Caudal peduncle short, deep, and laterally compressed. Dorsal-fin origin located anterior to vertical through pectoral-fin insertion. Pelvicfin origin located well posterior to vertical through pectoral-fin insertion.

Total vertebral count 27 or 28, with a formula of 14 + 13 or 14 + 14 precaudal and caudal vertebrae, respectively.

Jaws isognathous or with lower jaw slightly prognathous. Oral dentition bilaterally symmetrical and bicuspid, with well-developed distally expanded and slightly recurved cusps. Outer row teeth in both jaws enlarged and graded in size laterally. Rostrally, outer row teeth procumbently implanted in lower jaw but only slightly so in upper jaw. Two to four inner rows of teeth in upper jaw, and two to three inner rows in lower jaw. Inner row teeth smaller than those of outer row but of similar morphology. Total of three to five rows of teeth along rostral margin of upper jaw and three or four in lower jaw, tapering to a single row of teeth posteriorly. Dentition covers anterior half of dentary and nearly entire surface of premaxillary arcade.

LPJ well sutured on posteroventral margin. Dentition on LPJ and UPJ comprised of numerous, closely set, hooked, and bicuspid teeth. Medially on both LPJ and UPJ dentition robust and becoming strongly molariform. The LPJ toothplate is markedly laterally expanded rostrally, and obscures the underlying bone when viewed from above (fig. 16B). Three to six rows of hooked and bicuspid teeth on second pharyngobranchial toothplates. Three rows of teeth medially on ‘‘free’’ second epibranchial toothplate. Robust toothplates covering majority of dorsal surface of fourth ceratobranchial bones; toothplates confluent with outer row gill rakers of these elements. Teeth on fourth ceratobranchial toothplates unicuspid and conical laterally to hooked and bicuspid medially.

Lower limb rakers of first gill arch denticulate dorsomedially. Usually with 10 or 11 (modally 10; one individual with 8) triangular and somewhat elongate gill rakers arrayed along lower limb of first arch. Seven to nine epibranchial gill rakers. Gill rakers on remaining arches short, laterally expanded, and strongly denticulate. Teeth elongate and conical.

Body covered with large, regularly imbricate, ctenoid scales, including region above upper lateral line, from posterior of nape to

TABLE 6 Morphometric and Meristic Data for Ptychochromis loisellei , new species Values in parentheses indicate number of specimens examined with that count. (H) indicates count corresponding to holotype.

proximal portion of caudal fin. Scales on flanks becoming slightly more ctenoid posteriorly, but not strongly so. Most of scales on nape ctenoid. Scales on anterior of nape, most of head region, and cheek cycloid. Three or four rows of scales on cheek. Anterior chest and belly scales cycloid; posterior chest scales ctenoid. Scales on opercle and subopercle cycloid or weakly ctenoid. Anterior chest scales reduced in size and embedded. Snout, lachrymal, and anterior portion of interorbital region asquamate. Scales on caudal fin reduced in size; ctenoid anteriorly and much smaller and cycloid posteriorly. Lateral line scales 28–32. Five scales in diagonal from lateral line to dorsal-fin origin. Five to seven scales in oblique series between bases of pectoral and pelvic fins. No scale rows present along dorsal- and anal-fin bases.

Dorsal fin with XIII or XIV spines and 12 or 13 soft rays. Anal fin with III spines and eight or nine soft rays. First anal spine very short, second and third spines elongate and similar in length. Caudal fin emarginate, trailing margins of upper and lower lobes quite produced. Pectoral fin elongate, tapered distally. Distal margins of soft dorsal and anal fins produced and markedly pointed in larger specimens. Trailing margin of soft dorsal fin elongate and filamentous. Pelvic fin extending just past anal-fin origin in smaller specimens, well beyond origin in larger individuals.

MISCELLANEOUS OSTEOLOGY AND ANAT- OMY: Well-developed exoccipital foramina present on posterior of neurocranium. Paired anterior gas bladder extensions well developed, in contact with exoccipital region of neurocranium via connective tissue, but do not extend into exoccipital foramina. Infraorbital series (fig. 16D) composed of seven elements. Lachrymal deep, with four pores (fig. 16D). IO2 notably narrow and elongate, excluded from orbit by slightly ventrally displaced IO3. Uncinate process and anterior arm of first epibranchial bone short and robust (fig. 16C). Well-developed process and deep indentation present on inner face of Cb4.

COLORATION IN LIFE (plate 1C): Sexually active individuals exhibit an intense copperorange wash on the flanks and an orange spot in the center of each scale of the middle four or five rows around the lateral line. A dusky, wedge-shaped bar extends from a point immediately anterior to the origin of the anal fin dorsad to the base of the dorsal fin. As it approaches the dorsum, the bar tapers to a point. Soft dorsal, anal, lower lobe of the caudal and pelvic fins dusky black. Spinous dorsal fin edged in yellowish-white. Caudal with an intense red distal margin. Iris of the eye marked with orange-red.

COLORATION IN PRESERVATIVE: Ground coloration dark grayish-green or green. Large irregular black band, bands, or blotched region just anterior to and extending above anal-fin base (i.e., posterior region of flank). Black pigment mainly concentrated below lateral midline, but may extend dorsally onto dorsal-fin membrane. Some individuals with irregular black blotching on flanks. Some specimens with blue-gray spangling on flanks, particularly below lateral midline. Chest, belly, opercular region, and ventral flanks golden in some individuals, dark blackish-gray in others. Base body coloration slightly lighter ventrally. Upper lip medium to dark gray, lower lip pale yellow to golden. Dorsal and anal fins dark gray to blackish, anal fin lighter distally. Pectoral fins light gray and hyaline. Gular region blackish. Snout, lachrymal, and interorbital region dark gray.

DISTRIBUTION AND HABITAT: Ptychochromis loisellei is currently known only from the Mahanara River and its tributaries in northeastern coastal Madagascar near the town of Antsirabe-Nord (fig. 1).

CONSERVATION STATUS: According to local fishermen, P. loisellei is still relatively common throughout its limited range within the Mahanara River basin.

LOCAL NAME: Garaka is the local Malagasy name used to refer to the species of Ptychochromis of this region.

ETYMOLOGY: Named for our colleague Paul V. Loiselle, who collected the type series, in recognition of his many contributions to the understanding and conservation of Madagascar’s freshwater fishes.

DISCUSSION AND COMPARISONS: In our phylogenetic analysis, P. loisellei is recovered in a polytomy and its immediate relationships remain unresolved (fig. 2). Although there is currently no unambiguous evidence for the monophyly of an assemblage comprising P. loisellei , P. grandidieri , P. makira , and P. curvidens , the shared possession of an easterntype palatine morphology is unique to these four species and may indicate a close relationship among them. An indepenedent analysis of nucleotide characters also was unable to resolve the intrageneric placement of P. loisellei ( Sparks and Smith, 2004: fig. 1). Morphometrically P. loisellei is most similar to P. grandidieri from which it is readily distinguished by pigmentation pattern and coloration, infraorbital morphology, and also in the possession of a considerably smaller LPJ with fewer and less well-developed molariform teeth (compare fig. 16B with fig. 24B).

Ptychochromis curvidens , new species

Figures 17–18 , Plates 1B and 2D, Table 7

Ptychochromis sp. ‘‘vert-ouest de la Montagne d’Ambre’’: de Rham and Nourissat, 2002.

Ptychochromis n. sp. ‘‘montagne d’ambre’’ Sparks, 2003.

HOLOTYPE: MHNG 2623.82, 127.6 mm SL; northern Madagascar, Antsiranana (Diego Suarez) Province, Andranfanjava, Andranofanjava-Sandriapiana River system, P. de Rham and J.-C. Nourissat, 9 Oct., 1999.

PARATYPES: MHNG 2676.096, 2 ex., 92.8– 146.8 mm SL, data as for holotype. AMNH 237133, 2 ex., 1 ex. C&S, 92.3–105.1 mm SL, data as for holotype. MHNG 2623.84, 1 ex., 90.2 mm SL; northern Madagascar, Antsiranana (Diego Suarez) Province, Mirosolava, P. de Rham and J.-C. Nourissat, 10 Oct., 1999 .

DIAGNOSIS: A Ptychochromis exhibiting the eastern-type palatine morphology (fig. 18A) and distinguished from all congeners by the presence of a characteristically robust and strongly recurved oral dentition (fig. 18E). Ptychochromis curvidens is further distinguished by the possession of a feeble LPJ with only weakly molariform posteromedian dentition (fig. 18B), and a characteristic iridescent golden-green coloration in life.

DESCRIPTION: Morphometric and meristic data presented in table 7. Morphological characteristics and general pigmentation pattern can be observed in figure 17 and plates 1B and 2D. An extremely deep-bodied and robust species, rather less laterally compressed than other members of the genus. Head rather small (HL 30.4–31.7% SL), snout rounded, and ventral body outline convex and rounded. Lateral snout outline straight. Predorsal head profile straight from midorbit to dorsal-fin origin. Dorsal body profile relatively straight posterior to dorsal-fin origin, and strongly curved caudally. Ventral body profile moderately to strongly curved caudally. Caudal peduncle short, deep, and laterally compressed. Dorsal-fin origin located at about level of, to slightly posterior of, vertical through pectoral-fin insertion. Pelvicfin origin located well posterior to vertical through pectoral-fin insertion.

Total vertebral count 27 or 28, with a formula of 13 + 14 or 13 + 15 precaudal and caudal vertebrae, respectively.

Jaws isognathous. Oral dentition bilaterally symmetrical and bicuspid. Teeth extremely robust particularly rostrally, with notably recurved cusps (fig. 18E) that are L-shaped in lateral view. Outer row teeth enlarged and graded in size laterally. Rostrally, outer row teeth procumbently implanted in lower jaw. Rostrally, upper jaw with three to four and lower jaw with two to three inner rows of smaller teeth of similar shape to those of outer rows although with less markedly recurved cusps. Total of three to five rows of teeth along rostral margin of upper jaw and three or four in lower jaw, tapering to a single row of teeth posteriorly in both. Dentition covers anterior 2/3–3/4 of dentary and nearly entire surface of premaxillary arcade.

LPJ with a few weakly interdigitating sutures on posteroventral margin. Lower pharyngeal toothplates markedly smaller and less robust compared to those of similarly sized congeners. Dentition on LPJ and UPJ comprised of numerous, closely set, weakly hooked, and bicuspid teeth. Medially on UPJ and posteromedially on LPJ dentition becoming progressively robust, but only weakly so, and teeth are enlarged but rarely molariform. Three rows of hooked and bicuspid teeth on second pharyngobranchial toothplates. Two rows of teeth on ‘‘free’’ toothplate associated with second epibranchial bone. Robust, laterally expanded toothplates cover majority of dorsal surface of fourth ceratobranchial bones. Toothplates confluent with outer-row gill rakers of these elements. Dentition on fourth ceratobranchial toothplates unicuspid laterally, weakly to moderately hooked and bicuspid medially.

Lower limb rakers of first gill arch denticulate dorsomedially. Ten or eleven (modally 11) triangular and somewhat elongate gill rakers arrayed along lower limb of first arch. Eight or nine epibranchial gill rakers. Gill rakers on remaining arches laterally expanded, and strongly denticulate. Gill raker teeth on remaining arches elongate and conical.

Flanks covered with large, regularly imbricate, weakly ctenoid scales from posterior of nape to base of caudal fin. Scales on flanks becoming increasingly ctenoid posteriorly, but never strongly so. Scales on head, anterior portion of nape, and cheek cycloid. Three or four rows of scales on cheek. Scales on opercle and subopercle cycloid. Chest scales weakly ctenoid, except anteriorly. Scales on belly and anterior of chest cycloid, somewhat reduced in size and embedded. Snout, lachrymal, and anterior portion of interorbital region to about midorbit asquamate. Scales on caudal fin reduced in size, increasingly so posteriorly; ctenoid anteriorly and cycloid posteriorly.

TABLE 7 Morphometric and Meristic Data for Ptychochromis curvidens , new species Values in parentheses indicate number of specimens examined with that count. (H) indicates count corresponding to holotype.

Lateral line scales 30–32. Five-and-a-half to 6 1/2 scales in diagonal from lateral line to dorsal-fin origin. Five or six scales in oblique row between bases of pectoral and pelvic fins. No scale rows present along dorsal- and analfin bases.

Trailing margin of soft dorsal fin elongate and filamentous, particularly in larger specimens; trailing margin of anal fin elongate and filamentous, but to lesser degree than dorsal filament. Dorsal fin tall, with XIV spines and 11 or 12 soft rays. Anal fin with III spines and nine soft rays. First anal spine short, second and third spines elongate, although second spine noticeably shorter than third. Caudal fin very large and fanlike, weakly emarginate. Pectoral fin somewhat elongate, tapered distally. Pelvic fin extending to about level of anal-fin origin or slightly beyond when adducted.

MISCELLANEOUS OSTEOLOGY AND ANA- TOMY: Well-developed exoccipital foramina present on posterior of neurocranium. Paired anterior gas bladder extensions in contact with exoccipital region of neurocranium via connective tissue, but do not extend into exoccipital foramina. Infraorbital series (fig. 18D) composed of seven elements. Lachrymal rath- er shallow, with four pores (fig. 18D). IO2 broad and platelike, excluded from orbit by ventrally displaced IO3. Uncinate process of first epibranchial roughly twice diameter of anterior arm (fig. 18C). Well-developed process and deep indentation present on inner face of Cb4. Preopercular and mandibular canals markedly enlarged and pores expanded.

COLORATION IN LIFE (plate 1B): Base body coloration iridescent golden-green. Body dark golden-green dorsal of midline, and bright, iridescent golden below. Opercular region bright golden, except dorsally which is greenish. No distinctive lateral markings. Snout, lachrymal, interorbital region, gular region, and lips dusky charcoal gray. Belly charcoal gray. Unpaired fins dusky charcoal gray, membranes grayish to hyaline. Pelvic fins hyaline proximal to base, blackish distally. Pectoral fins bright yellowish-orange.

COLORATION IN PRESERVATIVE: Ground coloration dark brownish-green. Body darker dorsally. Smaller specimens yellowish to golden ventrally. No distinct lateral markings. Larger specimens with some light grayish spangling/spotting posteriorly on flanks, for the most part restricted ventral of lateral midline. Opercular and preopercular region golden. Gular region, lower lip, belly, and chest yellowish to golden in smaller individuals. Gular region and lower lip in larger specimens dark brownish-gray. Interorbital region, lachrymal, snout, and upper lip dark brownish-gray.

DISTRIBUTION AND HABITAT: Currently known only from a few westward flowing rivers draining the western flank of Montagne d’Ambre in extreme northern Madagascar . Most specimens are from the Andranofanjava-Sandriapiana River system, located southwest of the town of Antsiranana, but a single specimen purportedly collected from the Mirosolava River (Sahinana-Sampiena system) is included here in the type series. However, we note that although this single specimen (MHNG 2623.84) is labelled as coming from ‘‘northern Madagascar ; Antsiranana (Diego Suarez) Province, Mirosolava, P. de Rham and J.-C. Nourissat,’’ Patrick de Rham (in litt.), does not recall preserving any specimen of this species from Mirosolava, and, therefore, some doubt as to the exact provenance of this specimen persists. De Rham (in litt.) has expressed reservations as to whether the Andranofanjava and Sahinana river populations are conspecific, nevertheless our examination supports the presence of a single species in the far north of the island .

CONSERVATION STATUS: According to Patrick de Rham (in litt.), the conservation status of P. curvidens is difficult to assess. The species is not abundant anywhere within its range and exotic fishes, especially Oreochromis sp. , are present in the region. However, several other native species persist in reasonable numbers and exotic species are not abundant, suggesting that the ecosystem remains relatively healthy. This is further indicated by a diverse aquatic flora recorded at many sample sites, where endemic plant species such as Aponogeton cf. bovinianus remain plentiful. Human populations in these remote and poorly accessible regions remain small, and fishing pressure is consequently also low.

LOCAL NAME: Ptychochromis curvidens is referred to locally as garaka, a name that is used throughout much of northeastern and northern Madagascar to refer to species of Ptychochromis .

ETYMOLOGY: From the Latin, curvus, a ‘‘curve’’, and dens, a ‘‘tooth’’, in reference to the apomorphically recurved oral jaw dentition characteristic of this species.

DISCUSSION AND COMPARISONS: Although P. curvidens occurs in westward flowing basins in far northern Madagascar, the species exhibits the eastern-type palatine morphology (fig. 18A) otherwise found only in members of the genus restricted to eastern basins: P. grandidieri , P. makira , and P. loisellei . As such, P. curvidens represents the only species of Ptychochromis occurring in a western drainage with the eastern-type palatine configuration. Regrettably, no tissue sample suitable for molecular studies was available from specimens of P. curvidens , and the species could not be included in molecular phylogenetic analyses ( Sparks and Smith, 2004). In our phylogenetic analysis, P. curvidens is recovered in a polytomy and its intrageneric relationships remain unresolved (fig. 2).

Ptychochromis insolitus , new species

Figures 19–20 , Plates 1D and 2E, Table 8

Ptychochromis oligacanthus , race géographique ‘‘region de Mandritsara’’: Kiener and Mauge´, 1966.

Ptychochromis sp. ‘‘Mangarahara ou Mandritsara’’: de Rham and Nourissat, 2002.

Ptychochromis n. sp. ‘‘sofia’’: Sparks, 2003.

Ptychochromis nov. sp. ‘‘sofia’’: Sparks and Stiassny, 2003: table 9.1.

Ptychochromis sp. ‘‘Sofia’’: Sparks and Smith, 2004: fig. 1, table 1.

HOLOTYPE: UMMZ 237066, 54.8 mm SL, juvenile; northeastern Madagascar, Mahajanga Province, near town of Mandritsara, Sofia drainage basin, Amboaboa (5 Ambomboa) River (15 ° 50910S; 48 ° 429510E), J. S. Sparks and K. J. Riseng, 10 July 1996.

DIAGNOSIS: A Ptychochromis exhibiting the western-type palatine morphology (fig. 20A) and distinguished from congeners by a reduced number of precaudal vertebrae (12 vs. 13–14 in congeners) and a reduced number of infraorbital elements (five vs. six or seven in congeners). Ptychochromis insolitus is further distinguished from congeners by the presence (most obvious in preservation) of a faint midlateral stripe, beginning just posterior to the dorsocaudal margin of the opercle and extending to caudal-fin origin. Ptychochromis insolitus and P. inornatus are distinguished from the remaining species of Ptychochromis by the presence of strongly ctenoid scales, with the entire caudal scale margin bearing well-developed cteni (vs. weakly ctenoid scales, with only the central portion of the caudal margin bearing weak cteni).

DESCRIPTION: Morphometric and meristic data for the holotype presented in table 8. Morphological characteristics and general pigmentation pattern can be observed in figure 19 and plates 1D and 2E. The following description is based on the holotype, a wild caught specimen from the region of Mandritsara within the Sofia River drainage. However, a number of field photographs of additional specimens from the region have been made available to us and these are incorporated into the color description for the species. Additionally, a number of aquarium-raised specimens (purportedly F2) are available for examination; however, some morphological anomalies were noted in these individuals (see discussion below) and for that reason they have not been formally included within the type series.

A comparatively shallow bodied and laterally compressed Ptychochromis . Lateral snout outline straight to somewhat convex. Predorsal head profile mildly convex from midorbit to dorsal-fin origin. Dorsal body profile relatively straight posterior of dorsalfin origin. Ventral body profile relatively straight. Caudal peduncle comparatively elongate and laterally compressed. Dorsal-fin origin located slightly anterior to vertical through pectoral-fin insertion. Pelvic-fin origin located well posterior to vertical through pectoral-fin insertion.

Total vertebral count 27, with a formula of 12 + 15 precaudal and caudal vertebrae, respectively.

Jaws isognathous. Oral dentition bilaterally symmetrical and bicuspid, cusps expanded distally. Outer row teeth in both jaws enlarged and graded in size laterally. Rostrally, outer row teeth somewhat recurved. Teeth procumbently implanted in lower jaw, only slightly so in upper jaw. Rostrally, upper and lower jaws with three inner rows of smaller teeth of the same morphology as in outer rows; inner teeth tapering to a single row of teeth posteriorly in both jaws. Dentition covers anterior 2/3 of dentary and nearly entire surface of premaxillary arcade.

LPJ well sutured, with few weakly interdigitating sutures on posteroventral margin. Dentition on LPJ and UPJ comprised of numerous, closely set, hooked, and bicuspid teeth. Medially on both LPJ and UPJ dentition robust, but not molariform (fig. 20B). Three rows of hooked and bicuspid teeth on second pharyngobranchial toothplates. Two rows of teeth on ‘‘free’’ toothplate associated with second epibranchial bone. Robust, laterally expanded, toothplates cover most of dorsal surface of fourth ceratobranchial bones. Dentition on fourth ceratobranchial toothplates unicuspid or weakly hooked and bicuspid laterally, hooked and bicuspid medially.

Lower limb rakers of first gill arch denticulate dorsomedially. Nine triangular and somewhat elongate gill rakers arrayed along lower limb of first arch. Eight or nine epibranchial gill rakers. Gill rakers on remaining arches laterally expanded and strongly denticulate; teeth elongate and conical.

Body covered with large, regularly imbricate, strongly ctenoid scales from nape to proximal portion of caudal fin. Entire caudal margin of most scales with well-developed cteni. Scales on flanks becoming increasingly

TABLE 8 Morphometric and Meristic Data for Holotype of Ptychochromis insolitus , new species

Standard length (mm) 54.8

Percentage of SL

Head length 33.4 Body depth 38.7 Predorsal length 41.2 Preanal length 70.4 Prepelvic length 42.2 Head width (max.) 17.5 Caudal peduncle length 17.2 Caudal peduncle width 6.6 Caudal peduncle depth 15.3 Pectoral-fin length 29.6 Pelvic-fin length 30.1

Percentage of HL

Snout length 28.4 Orbit diameter 37.2 Upper jaw length 32.2 Lower jaw length 42.1 Interorbital width 27.9 Preorbital depth 17.5

Caudal peduncle length/depth 1.1 Caudal peduncle length/width 2.6

Scales in lateral line 34 Scales: lateral line to dorsal fin 6 Scales: pectoral to pelvic bases 5 Gill rakers (lower limb) 9 Vertebrae (precaudal + caudal) 12 + 15 5 27 Dorsal fin XIII 11 Anal fin III 8

ctenoid posteriorly. Scales on head and anterior portion of nape cycloid; posterior nape scales ctenoid. Scales on opercle and subopercle cycloid. Belly scales cycloid and chest scales weakly to moderately ctenoid, both somewhat reduced in size and embedded. Four rows of cycloid scales on cheek. Snout, lachrymal, and anterior portion of interorbital region asquamate. Scales on caudal fin reduced in size; ctenoid anteriorly and cycloid posteriorly. Lateral line with 34 scales. Seven scales in diagonal from lateral line to dorsalfin origin. Five scale rows between bases of pectoral and pelvic fins. No scale rows present along dorsal- and anal-fin bases.

Trailing margin of soft dorsal fin somewhat elongate. Dorsal fin with XIII spines and 11 soft rays. Anal fin with III spines and eight soft rays. First anal spine very short, second and third spines elongate and similar in length. Caudal fin emarginate. Pectoral fin elongate, tapered distally. Pelvic fin extending well beyond anal-fin origin when adducted.

MISCELLANEOUS OSTEOLOGY AND ANA- TOMY: Well-developed exoccipital foramina present on posterior of neurocranium. Paired anterior gas bladder extensions in contact with exoccipital region of neurocranium via connective tissue, but do not extend into exoccipital foramina. Infraorbital series (fig. 20D) composed of five elements. As noted previously, although infraorbital number is extremely variable within Cichlidae , a total of seven infraorbital elements is resolved as the plesiomorphic number for Cichlidae and is the number most often encountered among ptychochromins. A reduction to five is interpreted as a derived condition within Ptychochromis (Character 16(2), fig. 2). Lachrymal shallow, with four pores (fig. 20D). IO2 only marginally excluded from orbit by ventrally displaced IO3. Uncinate process and anterior arm of first epibranchial bone short and robust (fig. 20C). Well-developed process and deep indentation present on inner face of Cb4.

COLORATION IN LIFE (plate 1D): Based on photos of freshly captured specimens in de Rham and Nourissat (2002) and from the photo collection of de Rham. Base body coloration silvery to silvery-golden, particularly below lateral midline. Cheek and opercular series silvery. Body golden with blackish speckles and blotches above midline, including nape, top of head, and infraorbital series. Snout, interorbital region, and area dorsal to orbit gray to dark gray-green. Unpaired fins reddish, particularly distally, or dark gray. Caudal fin dark gray to black proximal to base, reddish distally. Thin, golden midlateral stripe; caudally stripe along lower lateral line particularly prominent in juveniles. Another thin, dark brown to blackish stripe paralleling dorsal-fin base. Pectoral fins pale and reddish. Pelvic fins pale reddish and may exhibit some black pigmentation along caudal margin. Some photographed, freshly captured specimens exhibit faint vertical barring on dorsal flanks.

COLORATION IN PRESERVATIVE: Ground coloration pale yellowish-olive. Faint dark stripe present along lateral midline, more pro-

Plate 1. Ptychochromis live coloration: (A) Ptychochromis inornatus , northwestern Madagascar, Anterano River. Photo by Patrick de Rham. (B) Ptychochromis curvidens , new species, northern Madagascar, Andranofanjava-Sandriapiana River system. Photo by Patrick de Rham. (C) Ptychochromis loisellei , new species, northeastern Madagascar, Mahanara River. Photo by Paul Loiselle. (D) Ptychochromis insolitus , new species, northeastern Madagascar, Amboaboa River. Photo by Patrick de Rham. (E) Ptychochromis grandidieri , male, eastern Madagascar, Lake Salehy. Photo by Paul Loiselle. (F) Ptychochromis grandidieri , female, eastern Madagascar, Ambila Lemaitso. Photo by Paul Loiselle. (G) Ptychochromis oligacanthus , male, northwestern Madagascar, Nosy Be, Lake Bemapaza. Photo by Paul Loiselle. (H) Katria katria , eastern Madagascar, Nosivolo River. Photo by M. Stiassny,

nounced posteriorly. No additional lateral markings evident. Body slightly lighter posterolaterally. Fins hyaline to pale yellowish-olive. Black pigment distally on dorsal fin. Caudal and anal fins dusky gray distally. Small, brownish spot present near base of anterior portion of soft dorsal fin, near margin of spinous and soft dorsal.

Plate 2. Ptychochromis in preservative: (A) Ptychochromis onilahy , new species, holotype, MNHN 1962- 0201, 84.3 mm SL. (B) Ptychochromis makira new species, holotype, AMNH 237131, 151.0 mm SL. (C) Ptychochromis loisellei , new species, holotype, AMNH 232462, 99.5 mm SL. (D) Ptychochromis curvidens , new species, holotype, MHNG 2623.82, 127.6 mm SL. (E) Ptychochromis insolitus , new species, holotype, UMMZ 237066, 54.8 mm SL. (F) Katria katria , holotype, AMNH 217739, 105.2 mm SL.

DISTRIBUTION AND HABITAT: Ptychochromis insolitus is known only from tributaries of the Sofia River (viz., Mangarahara and Amboaboa rivers near the town of Mandritsara), a westward flowing basin in northeastern Madagascar (fig. 1). Habitat at type locality mostly sand and rock substrate. Deeper pools present in sheltered areas. Water clear, shallow and current swift .

CONSERVATION STATUS: Ptychochromis insolitus appears to be rare throughout its limited range, and few specimens have been collected despite sizeable collections of other species having been made in this region, including other cichlids (viz., Paretroplus nourissati and P. cf. kieneri ).

LOCAL NAME: Joba or Juba is the Malagasy name used throughout much of northwestern

Figures

Fig. 13. Ptychochromis makira, new species, holotype, AMNH 237131, adult, 146.2 mm SL, northeastern Madagascar, Toamasina Province, north of Maroansetra, near town of Marovonona, Antainambalana River. Scale bar 5 1 cm. Drawing by Ian Hart.

Fig. 14. Ptychochromis makira, new species, paratype, AMNH 237132: (A) palatine, (B) LPJ, (C) first gill arch, and (D) infraorbital series.

Fig. 15. Ptychochromis loisellei, new species, holotype, AMNH 232462, 99.5 mm SL, northeastern Madagascar, Antalaha Province, north of Sambava, Mahanara River at Antsirabe-Nord. Scale bar 5 1 cm. Drawing by Ian Hart.

Fig. 16. Ptychochromis loisellei, new species, paratype, AMNH 231258: (A) palatine, (B) LPJ, (C) first gill arch, and (D) infraorbital series.

Fig. 17. Ptychochromis curvidens, new species, holotype, MHNG 2623.82, 127.6 mm SL, northern Madagascar; Diego Suarez Province, Andranofanjava, Andranofanjava-Sandriapiana River system. Scale bar 5 1 cm. Drawing by Ian Hart.

Fig. 18. Ptychochromis curvidens, new species, paratype, AMNH 237133: (A) palatine, (B) LPJ, (C) first gill arch, (D) infraorbital series, and (E) premaxilla.

Fig. 19. Ptychochromis insolitus, new species, holotype, UMMZ 237066, juvenile, 54.8 mm SL, northeastern Madagascar, Antalaha Province, near town of Mandritsara, Sofia drainage basin, Amboaboa River. Scale bar 5 1 cm. Drawing by Ian Hart.

Fig. 20. Ptychochromis insolitus, new species, holotype, UMMZ 237066: (A) palatine, (B) LPJ, (C) first gill arch, and (D) infraorbital series.