Paretroplus nourissati ( Allgayer, 1998 )

Sparks, J. S., 2008, Phylogeny Of The Cichlid Subfamily Etroplinae And Taxonomic Revision Of The Malagasy Cichlid Genus Paretroplus (Teleostei: Cichlidae), Bulletin of the American Museum of Natural History 2008 (314), pp. 1-151 : 97-104

Paretroplus nourissati ( Allgayer, 1998) Figures 45–48; plate 1G; table 8

HOLOTYPE: MNHN 1997-4172 , 98.2 mm SL; northwestern Madagascar: Sofia River drainage basin: near town of Mandritsara: Mangarahara River; J.-C. Nourissat, X- 1996.

PARATYPES: MNHN 1997-4173 , 4 ex., 84.1–96.6 mm SL ; data as for holotype .

ADDITIONAL MATERIAL EXAMINED: AMNH 229554 , 2 ex., 1 ex. C&S, 87.0– 106.0 mm SL ; northwestern Madagascar: Mahajanga Province: Sofia River drainage basin: near town of Mandritsara: Amboaboa River near confluence with Mangarahara River : 15 ° 50 9 1.0 0 S, 48 ° 42 9 52.0 0 E ; JSS 94-18; J. S. Sparks, K. J. Riseng, and local Malagasy guides, 24-VII-1994. AMNH 229555 , 3 ex., 1 ex. C&S, 72.0–78.0 mm SL ; northwestern Madagascar: Mahajanga Prov- ince: Sofia River drainage basin: near town of Mandritsara: Amboaboa River : 15 ° 50 9 1.0 0 S, 48 ° 42 9 51.0 0 E ; JSS 96-25: altitude 210.3 m a.s.l.; J. S. Sparks, K. J. Riseng, and local Malagasy guides, 10 and 11-VII-1996. AMNH 229562 , 1 ex., 64.0 mm SL ; northwestern Madagascar: Mahajanga Province: Sofia River drainage basin: west of Mandritsara: Amboaboa River ; PVL-93-VI-22; P. V. Loiselle, R. Morris, B. Vesta , and local villagers, 22-VI-1993. AMNH 236158 , 4 ex., 40.8–107.0 mm SL ; northwestern Madagascar: Mahajanga Province: Sofia River drainage basin: Amboaboa River at Andrahamamy village ; PVL-04-09; P. V. Loiselle. UMMZ 235205 , 76 ex., 4 ex. C&S, 1 ex. S ; 49.0–124.0 mm SL; data as for AMNH 229555. UMMZ 235206 , 40 ex., 3 ex. C&S, 7.0–160.0 mm SL ; data as for AMNH 229554. UMMZ 239530 , 6 ex., 76.0– 127.0 mm SL ; northwestern Madagascar: Mahajanga Province: Sofia River drainage basin: Mangarahara River just downstream from confluence with Amboaboa River , 15 ° 50 9 1.0 0 S, 48 ° 42 9 51.0 0 E: altitude 210.3 m a.s.l .; JSS 96-26; J. S. Sparks, K. J. Riseng, and local Malagasy guides, 11-VIII-1996. UMMZ 239545 , 3 ex., 76.0–90.0 m SL ; northwestern Madagascar: Mahajanga Province: Sofia River drainage basin: near town of Mandritsara: Amboaboa River near confluence with Mangarahara River : 15 ° 50 9 2.0 0 S, 48 ° 42 9 52.0 0 E ; JSS 94-53; J. S. Sparks, K. J. Riseng, and local Malagasy guides, 14-XI-1994. UMMZ 240363 , 1 ex. S, 62.5 mm SL ; northwestern Madagascar: Mahajanga Province: Sofia River drainage basin: near town of Mandritsara: Amboaboa River near confluence with Mangarahara River ; J. S. Sparks, K. J. Riseng, and local Malagasy guides, 10 and 11-VII-1996. UMMZ 243679 , 1 ex. S, 70.7 mm SL ; data as for UMMZ 240363 .

DIAGNOSIS: A shallow-bodied, elongate Paretroplus diagnosed from all congeners except P. lamenabe and P. tsimoly by the presence of two wide and convergent (below the lateral midline) dark brown to black midlateral bands, representing the second and third or third and four bars in series. Paretroplus nourissati is diagnosed in life by a pigmentation pattern of broad vertical black

TABLE 8 Morphometric and meristic data for Paretroplus nourissati . For meristics, numerals in parentheses indicate number of specimens examined with that count. (H) indicates count corresponding to holotype.

bars on the flanks over bright orange to reddish-orange ground coloration. It is further distinguished from both P. lamenabe and P. tsimoly , in life and preservative, by the absence of bluish-gray to bluish-black lips, lower jaw, lower cheek, gular region, and belly. Paretroplus nourissati is further distinguished from P. lamenabe by a shallower body (38.1%–43.5% vs. 47.0%–54.3% in P. lamenabe ), pelvic fins that do not extend to origin of the anal fin when adducted, and by a smaller adult size (up to 160 mm SL vs. regularly exceeding 180 mm SL in P. lamenabe ). Paretroplus nourissati is further distinguished from P. tsimoly by the absence of enlarged, lobed lips. Only P. lamenabe and P. nourissati exhibit a lateral pigmentation pattern in which the central two bars, which converge below the lateral midline, are by far the most prominent. By contrast, in life and preservation, P. tsimoly exhibits a pigmentation pattern in which the posteriormost five (or four, as the bar on the caudal peduncle is sometimes pale) bars on the flanks are about equally prominent.

DESCRIPTION: Morphometric and meristic data presented in table 8. Morphological characteristics and general pigmentation pattern in life and preservative can be observed in figs. 24C and 45–47. A shallow-bodied and elongate Paretroplus belonging to Clade F, which also includes P. lamenabe and P. tsimoly (fig. 1). Paretroplus nourissati is recovered as the sister taxon to P. tsimoly , and this clade is in turn the sister taxon to P. lamenabe . Paretroplus nourissati is comparable in adult size to P. tsimoly , rarely exceeding 150 mm SL. Head pointed, snout and predorsal profile moderately curved, lending species a beaked appearance in lateral view. Premaxillary pedicels prominent, distinct notch present posterior to distal margins of ascending processes. Dorsal body outline mildly curved, ventral outline mostly straight, except posteriorly. Caudal peduncle short, deep, and laterally compressed. No sexually dimorphic features apparent; however, in photographs of breeding pair in de Rham and Nourissat (2004: 122), male appears to exhibit brighter and more vivid yellowish to orange coloration.

Total vertebral count 29 to 31 (mode 31), with formulae of: 13 + 16, 13 + 17, 14 + 16, and 14 + 17, precaudal and caudal vertebrae, respectively.

Jaws isognathous. Lips fleshy and well developed, but not lobed (i.e., fleshy medial flaps lacking on both upper and lower lips). Single row of spatulate unicuspid teeth in both upper and lower jaws. Teeth laterally expanded, flattened at crown, and procumbently implanted. In upper jaw, tooth on either side of premaxillary symphysis greatly enlarged, remaining upper-jaw teeth smaller and graded in size laterally. Lower-jaw teeth at symphysis not enlarged, but reduced in size compared to adjacent lateral teeth, presumably to accommodate enlarged upper symphyseal teeth. Teeth in upper jaw usually number six to nine on each side, and total 13– 16. Teeth in lower jaw number six or seven on each side, and total 12–14. Upper-jaw teeth comparatively closely set for Paretroplus . Lower-jaw teeth more or less widely set and evenly spaced.

Upper and lower pharyngeal tooth plates well developed and dentition robust. Dentition on lower pharyngeal tooth plates (5 lower pharyngeal jaws [LPJ] or fifth ceratobranchial tooth plates) hooked and bicuspid both laterally and anteriorly, becoming progressively enlarged medially; molariform teeth present posteromedially, but these teeth not particularly robust. LPJ weakly sutured (i.e., gap present posteriorly between right and left fifth ceratobranchial elements in specimens up to about 100 mm SL), with few weak interdigitations on posteroventral margin. Seven robust tooth plates cover majority of dorsal surface of fourth ceratobranchial bones (fig. 12C). Tooth plates not confluent with outer-row gill rakers of fourth ceratobranchial elements. Bases of some tooth plates becoming confluent with each other, but majority separate. Dentition on fourth ceratobranchial tooth plates unicuspid or weakly hooked and bicuspid laterally, hooked and bicuspid medially (similar to lateral LPJ dentition). Dentition on third upper pharyngobranchial tooth plates robust and becoming nearly molariform posteromedially, hooked and bicuspid laterally and anteromedially. Anteromedial teeth becoming enlarged, but not molariform. Posteromedial teeth retain a weak apical cusp, even in larger specimens examined. Dentition on second pharyngobranchial tooth plates hooked and bicuspid and arrayed in two complete rows (occasionally, incomplete third row present).

Thirteen or 14 elongate triangular gill rakers arrayed along lower limb of first gill arch. Rakers on lower limb of first gill arch edentate to weakly denticulate medially. All other lower-limb rakers (i.e., those on gill arches 2–4) triangular and moderately to strongly denticulate dorsally. These rakers elongate compared to short, spherical rakers in members of Clade G, comprising deepbodied Paretroplus (Clade I) and P. kieneri species complex (Clade H). Teeth on rakers of gill arches 2–4 thin, conical and curved distally near crown, but much shorter than body of raker. Epibranchial rakers on first gill arch elongate and thin, numbering 12.

Body covered with large, regularly imbricate, cycloid scales. Posterior field of lateral body scales thin and not ossified. Posterior field of head scales ossified. Well-developed scale ridges (5 scale sheathing) present along dorsal- and anal-fin bases. Scale ridges free from spiny dorsal and anal fins, but becoming fused to membranes of both soft dorsal and anal (fig. 22B). On membranes between fin rays, multiple discrete rows of scales extend well beyond scale ridges and onto both soft dorsal and anal fins. Pelvic axillary scale present and well developed. Interpelvic scale elongate and somewhat pointed terminally. Lateral-line scales number 35–39 (mode 37). Chest scales noticeably reduced in size and embedded (fig. 24C). Belly scales markedly reduced in size compared to chest scales, those along ventral midline smallest and very embedded (such that the belly appears asquamate). Multiple rows of scales, markedly reduced in size, extend on flanks from chest (and continue posteriorly dorsal to pelvic fins and anus) to about anal-fin origin (fig. 24C). Three to five (usually three or four) rows of scales on cheek. Opercle and subopercle scaled. Interopercle scaled only posteriorly or asquamate. Preopercle asquamate. Snout, lacrimal, and anterior portion of interorbital region asquamate. Scales on caudal fin reduced in size and extending posteriorly about 3/4 length of fin on dorsal and ventral lobes, and about 1/3 length of fin medially.

Dorsal with XIV–XVI spines, 12–15 soft rays. Anal with VII–VIII spines, 10–13 soft rays. Origin of dorsal fin at about level of vertical through anterior margin of pectoralfin insertion. Caudal fin weakly emarginate, margins of upper and lower lobes broad and rounded, and not produced. Pectoral fin broad and rounded at distal margin, rays slightly excised from fin terminally. Distal margins of soft dorsal and anal fins rounded, slightly produced in larger specimens, but with no trailing margins. Posterior margins of soft dorsal and anal fins terminate just anterior of caudal-fin origin in juveniles and extend slightly past origin in large adults (from about 120 mm SL). Pelvic fins terminate anterior to origin of anal fin when adducted.

MISCELLANEOUS OSTEOLOGY AND ANAT- OMY: Large, well-developed exoccipital foramina present. Paired anterior gas bladder bullae with tough and thickened tunica externa, and anteriormost chambers firmly lodged in exoccipital recesses (fig. 20C). Prominent excavation (5 supraoccipital notch of Stiassny et al., 2001) lacking along posterior margin of supraoccipital. Supraoccipital extending anteriorly over median frontal pores of neurocranium (nlf 0 of Barel et al., 1977). Two distinct and well-separated proximal premaxillary-maxillary ligaments present (rostral ligament unique to Paretro- plus within Cichlidae ). An additional, fully ossified, anal- and dorsal-fin pterygiophore, not associated with any fin rays, present terminally in both fins (structures appear less well developed in P. nourissati and P. tsimoly than in other species of Paretroplus ). Nasal bones elongate. Lower pharyngeal jaw (LPJ) with reduced number of interdigitations on ventral suture and indentation on ventral face of posterior horns. Interdigitations along ventral suture of LPJ shallow and weak. Posterior horns of LPJ narrow, particularly along ventrally projecting ridges, and gap present between left and right fifth ceratobranchial elements, posterior to ventral suture.

COLORATION IN LIFE: Pinkish-brown, grey, or reddish-brown (nonbreeding, sexually quiescent individuals) to bright yellowish-orange, orangish-red, or bright red (breeding, sexually active individuals) ground coloration, with two prominent vertical and generally converging (below the lateral midline) dark gray to jet black midlateral bars (de Rham and Nourissat, 2004: 122). Total of six or seven vertical bars present on flanks. Apart from two prominent midlateral bars, representing the second and third or third and fourth bars in series, remaining bars generally faint, although those on posterior region of flank easily visible. Fins gray, pinkish brown, or reddish brown and similar to base body coloration in nonbreeding individuals, and solid yellowish orange, orangish red, or bright red in breeding individuals. Lips, lower cheek, gular region, and belly very pale blue or grey in sexually active individuals (see photographs in de Rham and Nourissat, 2004: 122).

COLORATION IN PRESERVATIVE: Body ground coloration reddish brown, golden brown, or dark grayish brown. Six or seven dark brown or blackish vertical bars present on flanks. Two midlaterally located bars most prominent and usually converging below lateral midline (figs. 24C, and 45–46). Apart from two prominent midlateral bars, remaining bars comparatively faint, although those on posterior flank readily visible. Body somewhat darker overall dorsally. Dark gray, golden brown, or brownish triangular patch easily visible in pectoral-fin axil in most specimens (faint in others). Unpaired fins olive, brown, or grayish. Pectoral fins olive or gray. Pelvic fins tan, brown, or light grey. Lower lip, lower cheek, and gular region ranges from pale yellow, to olive, to light brown. Upper lip ranges from pale yellow to dark grayish brown. Juvenile specimens strongly barred (to about 70 mm SL), with series of lateral bars prominent over entire flank (vs. only two midlateral bars strongly pigmented in adults) (fig. 47).

DISTRIBUTION AND HABITATS: The known range of P. nourissati is restricted, encompassing only the Amboaboa and Mangarahara rivers near their confluence, which is in the vicinity of the town of Mandritsara in northeastern Madagascar (fig. 48). The species is also reported to occur in neighboring lakes, located a few kilometers from the confluence of the Mangarahara and Amboaboa rivers (de Rham and Nourissat, 2004: 121–122). The Amboaboa River is a tributary of the Mangarahara River, which itself is a moderately sized left-bank tributary of the extensive westward flowing Sofia drainage. Paretroplus nourissati co-occurs in these rivers with P. gymnopreopercularis ; neither species has been collected to date from the Sofia River proper. Ichthyological surveys in this remote region of Madagascar have not been extensive and P. nourissati could potentially be more widespread in distribution.

The Amboaboa and Mangarahara rivers near the type locality are shallow, clear (low in turbidity), and the current is swift, with many areas of small cascades and riffles. These rivers flow over large areas of exposed bedrock, and the substrate is generally rocky, with many exposed boulders, and interspersed with areas of sand. Paretroplus nourissati frequents shallow, rocky stretches with a swift current and riffles. The species is frequently collected in swift-flowing water only a few inches deep, where it hides under rocks.

Although not widespread in distribution, in the mid-1990s P. nourissati was still relatively common within its limited range in the Mangarahara and Amboaboa rivers. Nevertheless, in consideration of its very restricted distribution, P. nourissati would seem to be vulnerable to increased fishing pressure and continued habitat degradation. Unfortunately, de Rham and Nourissat (2004) report that on their most recent visit to the region (in 1999) they found the Mangarahara River upstream from its confluence with the Amboaboa River to be completely dry due to the combined effects of a prolonged drought in the region and water diversion to irrigate rice fields yearround via an upstream dam (for additional information see Distribution and Habitats for P. gymnopreopercularis below). Subsequent ichthyological surveys of the Amboaboa River found P. nourissati to be abundant in 2004, but greatly reduced in numbers in 2006, possibly as a consequence of the many years of severe drought that the region has experienced (P. Loiselle, personal commun.).

LOCAL NAME: Lamena , a Malagasy word, which translates literally as ‘‘the red one’’.

ETYMOLOGY: The species was named in honor of the French aquarist, naturalist, and intrepid traveler Jean-Claude Nourissat, who with Patrick de Rham discovered the species (in 1991) and collected the type specimens.

RELATIONSHIPS AND DISCUSSION: Based on the simultaneous analysis of morphological features and nucleotide characters, P. nourissati is recovered as a member of Clade F and as the sister taxon to P. tsimoly (fig. 1). Clade F is diagnosed by two unambiguously optimized morphological transformations, the first of which is unique and unreversed, the presence of two prominent and converging midlateral bars (several fainter bars are also present on the flanks) (figs. 24C, and 45– 46) and elongate nasal bones. Paretroplus damii is recovered as the sister taxon to Clade F. Clade E, comprising P. damii , P. tsimoly , P. nourissati , and P. lamenabe , is united by five unambiguously optimized morphological features, two of which are unique and unreversed (features discussed and presented above) (fig. 1).

Paretroplus nourissati was originally described as the type species of a new genus, Lamena Allgayer, 1998 , and represented the first member of the ‘‘lamena’’ group (Clade F) to be discovered. The results of a number of systematic studies that followed soon thereafter indicated that Lamena nourissati was nested within Paretroplus , and it was argued that Lamena be synonymized with Paretroplus Bleeker, 1868 ( Sparks and Reinthal, 1999; Sparks, 2001; Stiassny et al., 2001; Sparks, 2004a; Sparks and Smith, 2004). Paretroplus damii Bleeker, 1868 is the type species of the genus, and is the sister taxon to Clade F, which includes P. nourissati . Synonymy of Lamena with Paretroplus reinstated Paretroplus as a monophyletic assemblage.

Although members of Clade F are very similar morphologically, sexually active (breeding) individuals of Paretroplus nourissati exhibit a diagnostic pigmentation pattern consisting of broad vertical black bars on the flanks over bright orange to reddish-orange ground coloration. Paretroplus nourissati is further distinguished from the other members of Clade F, P. lamenabe and P. tsimoly , in life and preservative, by the absence of dark bluish-gray to bluish-black lips, lower jaw, lower cheek, gular region, and belly. Although this region may be pale blue or very pale gray in breeding P. nourissati (see photographs of aquarium specimens in de Rham and Nourissat, 2004: 122), it is never darkly pigmented. Additionally, P. nourissati can be distinguished from P. lamenabe by a shallower body (38.1%–43.5% vs. 47.0%– 54.3% in P. lamenabe ), shorter pelvic fins that do not extend to anal-fin origin when adducted, and by a smaller adult size (, 160 mm SL vs. regularly. 180 mm SL in P. lamenabe ). Paretroplus nourissati is further distinguished from P. tsimoly by the absence of markedly enlarged and lobed lips. Paretroplus nourissati and P. lamenabe are unique in sharing a lateral pigmentation pattern, in life and preservation, in which the central two bars, which converge below the lateral midline, are the most strongly pigmented (vs. the posteriormost five [or four, as the bar on the caudal peduncle is sometimes pale] bars on the posterior flanks about equally prominent in P. tsimoly ). Paretroplus nourissati and P. tsimoly can generally be distinguished from P. lamenabe by tooth count in the upper jaw (13–16 and 12–14 in P. nourissati and P. tsimoly , respectively, vs. 15–18 in P. lamenabe ), and by a much lesser degree of squamation extending onto the soft dorsal and anal fins. Stiassny et al. (2001) report a range of 14–20 teeth in the upper jaw for P. nourissati ; however, I have not been able to find a specimen with more than a total of 16 upper-jaw teeth.

All members of Clade F share two features of the lower pharyngeal jaw (LPJ) that were noted by Stiassny et al. (2001: 37–38, fig. 9), a reduced number of interdigitations on the ventral suture (posteroventral margin of LPJ) and an indentation on the ventral face of the posterior horns. In addition, the interdigitations along the ventral suture of the LPJ in members of Clade F are shallow and weak. Members of Clade F are also united by the presence of posterior horns on the LPJ that are very narrow, particularly along the ventrally projecting ridges, and by the presence of a narrow gap between left and right fifth ceratobranchial elements, posterior to the ventral suture.

Members of Clade E, which includes Clade F and P. damii , are unique among Paretroplus in sharing a number of apomorphic features related to scale morphology and squamation pattern. Uniquely in members of Clade E, the posterior field of the lateral body scales is unossified and feeble (fig. 22B), multiple discrete rows of scales extend onto the membranes of both the soft dorsal and anal fins (fig. 24C), the chest scales are distinctly reduced in size and quite embedded, and multiple rows of scales of reduced size extend posteriorly from the chest (dorsal to the pelvic fins and anus) to about anal-fin origin (figs. 45–46). Within Paretroplus , members of Clade F exhibit a squamation pattern on the chest and belly in which the scales are both more reduced in size and more highly embedded than in other members of the genus, including P. damii , lending the ventral chest and belly an asquamate appearance.