Ptychochromis ernestmagnusi, Sparks, John S. & Stiassny, Melanie L. J., 2010
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publication ID |
https://doi.org/ 10.5281/zenodo.193312 |
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DOI |
https://doi.org/10.5281/zenodo.5624421 |
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persistent identifier |
https://treatment.plazi.org/id/03BC2F53-513C-FFE1-C1CA-F9485A9FFEF2 |
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treatment provided by |
Plazi |
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scientific name |
Ptychochromis ernestmagnusi |
| status |
sp. nov. |
Ptychochromis ernestmagnusi View in CoL , new species
Figures 1– 5 View FIGURE 1 View FIGURE 2 View FIGURE 3 View FIGURE 5 ; Table 1 View TABLE 1
Holotype: AMNH 249490, 146.6 mm SL, male, Mananara (du nord) River at Antanambaobe Village (16° 16′ 0 1″S, 49°40′ 0 1″E), 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′ 0 1″S, 49 ° 40′ 41″E), 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 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.
Diagnosis. A Ptychochromis exhibiting the eastern-type palatine morphology ( Fig. 2 View FIGURE 2 ) and further distinguished from all congeners except P. grandidieri , P. loisellei , P. m a k i r a and P. curvidens by an anterior displacement of the first supraneural such that it overlies the dorsoposterior margin of the supraoccipital ( Fig. 3 View FIGURE 3 A– C). Among eastern group Ptychochromis , P. m a k i r a 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. m a k i r a is the presence of strong (paired) lateral barring as a prominent component of pigmentation patterning; however, P. m a k i r a 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. m a k i r a 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 View TABLE 1 . External anatomical characteristics and general pigmentation pattern in life and preservation can be observed in Figures 1 View FIGURE 1 , 5 View FIGURE 5 and 7 View FIGURE 7 B. 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 View FIGURE 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.
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.
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)
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.
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.
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 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 View FIGURE 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. 3 View FIGURE 3 A). 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 View FIGURE 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. 3 View FIGURE 3 A) rostrally positioned, with first supraneural overlying dorsoposterior margin of supraoccipital crest. Both supraneurals characteristically shaped with flat dorsal margins ( Fig. 3 View FIGURE 3 A). Posterior supraneural often reduced in size or absent.
Coloration in life ( Figure 5 View 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 View 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 View 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 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).
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.
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 ethmoid process (i.e., “eastern type palatine”). On the cladogram presented by Stiassny and Sparks (2006: Fig. 2 View FIGURE 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 a k i r a, P. loisellei , and P. curvidens ) were represented as a polytomy ( Stiassny & Sparks, 2006: Fig. 2 View FIGURE 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. 3 View FIGURE 3 A–C).
To test this hypothesis, we reran our prior phylogenetic analysis ( Stiassny & Sparks, 2006: Fig. 2 View FIGURE 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. m a k i r a 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 View 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 View TABLE 2 . As the results indicate ( Fig. 7 View FIGURE 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. 3 View FIGURE 3 A–C) and “eastern type palatine” configuration ( Sparks & Stiassny, 2006: character 21; Fig. 2 View FIGURE 2 ) discussed above.
Characters
Among the species of Ptychochromis that exhibit an “eastern type palatine” configuration, P. m a k i r a shares with P. ernestmagnusi the possession of supraneurals with a characteristically flattened dorsal profile ( Fig. 3 View FIGURE 3 A, 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 View FIGURE 7 ; character 24). However, in P. m a k i r a these pairs of lateral flank bars are distinctively V-shaped ( Fig. 7 View FIGURE 7 A), whereas in P. ernestmagnusi the bars are vertically oriented ( Fig. 7 View FIGURE 7 B). Despite inconclusive results in our earlier phylogenetic analysis ( Sparks & Stiassny, 2006: Fig. 2 View FIGURE 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 a k i r a 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. m a k i r a by conspicuous iridescent spangling on the flank and dorsal fin membrane 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. Although only two specimens of P. m a k i r a 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. m a k i r a), has a larger eye (29.5–37.4 vs. 25.6–27.6% HL in P. m a k i r a), a longer head (34.2–37.1 vs. 32.0–32.2% SL in P. m a k i r a), a shorter preanal length (71.6–73.5 vs. 75.1% SL in P. m a k i r a), and a greater lateral line scale count (34–37 vs. 33 in P. makira ) than its sister taxon, P. m a k i r a.
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, 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- 0 780, 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° 58.49′S; 49° 57.81′E), 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° 38.49′S 49° 57.81′E), 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° 58.49′S 49° 57.81′E), 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° 57.30′S 49° 56.20′E), 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′ 1″S; 48° 42′ 51″E), 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′ 41.0″S, 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′ 20.4″S, 48º 14′ 52.2″E), 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′ 49.3″S, 48º 0 7′ 54.3″E), JSS 94-21. UMMZ 237067, paratypes, 7 ex., northwestern Madagascar, Antalaha Province, northeast of Antsohihy, Ankofia drainage, Anjingo River (14º 50′ 39.7″S, 48º 14′ 39.5″E), 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 ex., C&S, southeastern Madagascar, Mananjary, estuary of Mananjary River, 21º05′S 48º27′E. AMNH 88053, 2 ex., southeastern Madagascar, Mananjary, estuary of Mananjary River, 21º 0 5′S, 48º 27′E. AMNH 88076, 2 ex., eastern Madagascar, Vatomandry, 19º 20′S, 49º 0 0′E. AMNH 88090, 3 ex., eastern Madagascar, Mahanoro, 19º 55′S, 48º 50′E. 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 10′S, 49º 25′E. AMNH 88140, 1 ex., eastern Madagascar, 25 km north of Tamatave, Pangalanes canal, 18º 0 0′S, 49º 25′E. AMNH 88153, 11 ex., eastern Madagascar, between Fenerive and Tamatave, Pangalanes canal, 18º 0 0′S, 49º 25′E. 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 55′S, 48º50′E. 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′ 47″S, 47º 47′ 25″E. AMNH 231347, 1 ex., southeastern Madagascar, Ampataka village, Sahambavy River, 23º 21’ 04”S, 47º28′18″E. AMNH 231352, 4 ex., southeastern Madagascar, Manombo Special Reserve, Takoandra River, 23º 0 1′ 27″S, 47º 43′ 16″E. 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′ 47″S, 47º 22′ 0 5″E. 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 .
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.
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.
| Character | P. ernestmagnusi 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 |
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.
| 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 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 |
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