Paretroplus maromandia Sparks and Reinthal, 1999

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 : 104-117

Paretroplus maromandia Sparks and Reinthal, 1999 Figures 35, 49–50; plate 1H; table 9

Paretroplus aff. maromandia from Lake Andrapongy and the Anjingo-Ankofia River: de Rham and Nourissat, 2004: 109–110.

Paretroplus aff. maromandia from the Maevarano River: de Rham and Nourissat, 2004: 107–108.

HOLOTYPE: UMMZ 234790 , holotype, adult, likely male (sex indeterminate due to poor internal preservation), 126.9 mm SL;

TABLE 9

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

northwestern Madagascar: Province of Mahajanga: region of Ambanja: immediately south of village of Maromandia : Maintsomalaza River , near confluence of Adranomalaza and Manongarivo rivers, downstream from where rivers unite: 14 ° 12 9 S, 48 ° 04 9 E ; JSS 96-13; obtained from fisherpersons; J. S. Sparks, K. J. Riseng, and Richard Randriamampionina, 7-VII-1996.

PARATYPE: AMNH 227336 , 1 ex., juvenile, presumably female (gonads reasonably well developed with abundant lipids, although no oocytes detected), 113.0 mm SL ; data as for holotype .

ADDITIONAL NONTYPE MATERIAL EXAM- INED: MHNG 2537.60 , 2 ex. (1 dried), 93.7– 130.0 (approx./dried) mm SL ; north- western Madagascar: region of Antsohihy: Ankofia River drainage: Lake Andrapongy ; P. de Rham and J.-C. Nourissat, 17-X-1992. MHNG 2623.080 , 3 ex., 1 ex. C&S, 119.8– 121.8 mm SL ; northwestern Madagascar: Region of Ambanja : near town of Maromandia: Manongarivo River , region of mouth of Manandtritzara [sic] River ; P. de Rham and J.-C. Nourissat, 22-X-1999. MHNG 2640.041 , 1 ex., 129.7 mm SL ; northwestern Madagascar: region of Ambanja: near town of Maromandia : Andranomalaza River ; P. de Rham and J.-C. Nourissat, X-1999.

DIAGNOSIS: Paretroplus maromandia is a member of the deep-bodied clade of Paretroplus (Clade I) and the only member of the genus, apart from P. polyactis , in which the entire lateral series of bars is prominent, including in adults. It is the only species of Paretroplus occurring in western drainages with strong lateral barring. Paretroplus maromandia is distinguished from P. polyactis by the presence of lateral bars that extend more or less over the entire flank, from the dorsalfin base (and sometimes extending onto the fin membrane itself) to the anal-fin base (vs. bars that terminate approximately under the lateral midline and do not extend to the analfin base in P. polyactis ), by a greater lateralline scale count (39–41 vs. 31–36 in P. polyactis ), fewer gill rakers on the lower limb of the first arch (10 vs. 11–13 in P. polyactis ), a higher dorsal-fin ray count (20–23 vs. 15–18 in P. polyactis ), and by bright yellow to greenishyellow body ground coloration interrupted by seven prominent, solid black vertical bars on the flanks (vs. bars and intervening regions highly speckled/spotted and resembling a chain-link fence, owing to pigmentation pattern in which scale margins are markedly darker than centers in P. polyactis ). In life, P. maromandia is further distinguished from P. polyactis by a golden iris (vs. red), and the presence of vivid red pigmentation on the flanks, below the lateral midline.

DESCRIPTION: Morphometric and meristic data presented in table 9. Morphological characteristics and general pigmentation pattern in life and preservative can be observed in figs. 49–50. A deep-bodied, laterally compressed Paretroplus belonging to Clade I, which comprises all deep-bodied and essentially disk-shaped members of Paretroplus (also including P. dambabe , P. maculatus , P. menarambo , P. petiti , and P. polyactis ) (fig. 1). Body laterally compressed and deep. Head blunt and steeply sloping in lateral view. Predorsal head profile moderately to strongly curved. Interorbital protuberance producing distinct concavity rostral to orbit. Lateral snout outline more or less straight, forming angle of approximately 50 ° to horizontal. Caudal peduncle short, deep, and laterally compressed. No sexually dimorphic features readily apparent.

Total vertebral count 32–34 (mode 33), with formulae of: 14 + 18, 15 + 18, and 16 + 18 precaudal and caudal vertebrae, respectively.

Jaws isognathous and short, both upper and lower jaws bearing tiny, fleshy papillae. Single row of spatulate unicuspid teeth, generally widely and irregularly set, in both upper and lower jaws. Teeth wide and flattened at crown, and procumbently implanted. In upper jaw, dentition restricted to anterior portion of premaxillary arcade. Tooth on each side of premaxillary symphysis markedly enlarged; these symphyseal teeth closely set. Other upper-jaw teeth graded in size laterally. Lower-jaw teeth at symphysis not enlarged, but somewhat reduced in size compared to adjacent lateral teeth, presumably to accommodate enlarged upper symphyseal teeth. Including symphyseal teeth, teeth in upper jaw number five to seven on either side. Upper-jaw teeth laterally graded in size and widely set. Lower-jaw teeth, including those at symphysis, number seven to 10 in total. Lower-jaw teeth not uniform in size or spacing. Posteriormost two lower-jaw teeth reduced in size (i.e., anteromedial teeth not graded in size laterally). Dentition restricted to anterior portion of dentary.

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; robust molariform teeth present posteromedially. Hooked and bicuspid peripheral LPJ teeth numerous and closely set, whereas enlarged medial and molariform posteromedial teeth widely and/or irregularly set. LPJ well sutured, with numerous interdigitating sutures on posteroventral margin. Eight robust tooth plates cover majority of dorsal surface of fourth ceratobranchial elements; anteromedial tooth plates abut each other, whereas those on posterior half of fourth ceratobranchial not confluent. Tooth plates not confluent with outer-row (5 lateral) gill rakers of fourth ceratobranchial elements. 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 molariform posteromedially, hooked and bicuspid both laterally and anterome- dially. Dentition on second pharyngobranchial tooth plates hooked and bicuspid, closely set, and arrayed in four rows.

Ten robust, triangular, and rather stout gill rakers closely arrayed along lower limb of first gill arch, excluding raker located in angle of arch; anteriormost one or two rakers somewhat reduced in size. These ceratobranchial gill rakers denticulate dorsomedially. All other lower-limb rakers (i.e., those on gill arches 2–4) short, stout, and strongly denticulate dorsally. Epibranchial gill rakers of first arch elongate and slender, and number 10– 12.

Body covered with large, regularly imbricate, cycloid scales from level of orbit to base of caudal fin. Well-developed ridges of scales (5 scale sheathing) present along dorsal- and anal-fin bases. Scale ridges free from both spiny and soft portions of dorsal and anal fins (figs. 49–50). Pelvic axillary scale present and well developed. Interpelvic scale elongate and tapered distally. Lateral line scales number 39–41 (mode 40). Chest scales only slightly reduced in size compared to flank scales, and weakly embedded. Belly scales along ventral midline markedly reduced in size and embedded; much smaller than flank and chest scales. Cheek scales arrayed in oblique series, extending from ventral margin of midorbit to ventral margin of preopercle, and numbering six to seven rows. Snout, lacrimal, and anterior portion of interorbital region asquamate. Preopercle scaled anteriorly, asquamate ventrally and along posterior margin of shaft. Opercle and subopercle fully scaled; interopercle fully scaled except at rostral margin. Scales on caudal fin reduced in size and extending to about 3/4 length of fin on dorsal and ventral lobes, and 1/4 to 1/3 length of fin medially.

Dorsal with XV–XVI spines, 20–23 soft rays. Anal with VIII–IX spines, 14–16 soft rays. Origin of dorsal fin located somewhat posterior to vertical through pectoral-fin insertion (in closely related congeners, dorsal-fin origin at about level of pectoral-fin insertion). Origin of dorsal fin located slightly anterior to pelvic-fin insertion. Caudal margins of both the soft dorsal and anal fins notably rounded, extend only slightly beyond origin of caudal fin. Caudal fin emarginate and crescent shaped, trailing margins of upper and lower lobes slightly produced, lending forked appearance to fin. Pelvic fins extend just past origin of anal fin when adducted.

MISCELLANEOUS OSTEOLOGY AND ANAT- OMY: Large, well-developed exoccipital foramina present. Paired anterior gas bladder bullae with tough, thickened tunica externa and narrow tubular connections (5 diverticula) to main gas bladder chamber. Anteriormost chambers firmly lodged in exoccipital recesses. Expansive, wide and rounded excavation (5 supraoccipital notch of Stiassny et al., 2001) 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 Paretroplus within Cichlidae ). An additional, fully ossified, anal- and dorsal-fin pterygiophore, not associated with any fin rays, present terminally in both fins.

COLORATION IN LIFE: Base body coloration bright yellow, greenish yellow, or olive. Seven prominent and broad black vertical bars present on flanks (anteriormost bar sometimes considerably weaker than rest, but always visible). Bars extend anteroposteriorly from caudal margin of opercle to caudal peduncle, and generally extend over entire flank, from dorsal-fin base (and sometimes extending onto fin membrane itself) to anal-fin base. Head and opercular region golden, golden brown, or greenish brown. Vivid red spotting on flanks, generally restricted ventral to lateral midline and concentrated anteriorly, posterior to pectoral-fin base. Red pigmentation may extend onto opercle and subopercle. Red flank pigmentation generally interrupted by dark lateral bars, but sometimes partially occluding anterior bars (see color photographs in de Rham and Nourissat, 2004: 103–109). In juveniles, red spotting covers entire flank (see photograph in de Rham and Nourissat, 2004: 104). Interorbital region gray. Snout golden, golden brown, greenish brown, or gray. Gular region, chest, belly and ventrum ranges from bright yellow, to gray, to nearly black. Grayish or black pigment may extend posteriorly along ventrum and dorsal to anal-fin base. Lips gray to dark gray. Dorsal and anal fins generally dark charcoal gray, both with thin red terminal band. Caudal fin dark gray proximally, becoming lighter at distal margin of fin, and fringed in red terminal band. Pelvic fins dark gray to blackish. Pectoral fins gray to dark charcoal. Solid black patch present in pectoral-fin axil, visible as black crescent when fin adducted. All fins, except pectorals, with slight reddish tinge in life.

COLORATION IN PRESERVATIVE: Base body coloration tan, with some goldenbrown hues, to golden brown. Red pigmentation on flanks lost in alcohol. Overall body pigmentation is somewhat darker dorsally, becoming lighter ventrally. Seven distinct, dark brown to black vertical bands present on flanks, extending from posterior margin of head to caudal peduncle (figs. 49–50). Bands most pronounced medially and dorsally on flanks, but extend over entire flank. Head and opercular region golden brown, dark brown, or grayish. Interorbital region, lacrimal, and snout dark brown to dark grayish brown. Lower lip, gular region, and lower portion of opercular series yellowish to golden brown. Upper lip yellowish to grayish brown. Chest and belly golden brown to dark grayish brown. Dorsal fin light grayish brown to dark brown. Anal fin a darker grayish brown to charcoal. Caudal fin tan, golden brown, or medium brown, and similar to flank ground coloration. Pelvic fins similar in coloration to the anal fin, dark grayish brown to charcoal overall, although leading edge and posterior rays tan or golden brown. Pectoral fins mostly hyaline, rays light tan to golden brown. Black patch present in pectoral-fin axil.

VISCERA AND DIET: Radiographs of holotype and paratype reveal that P. maromandia feeds primarily on small gastropods, as gut of each specimen was tightly packed with crushed shells. Presence of robust molariform pharyngeal dentition also consistent with this feeding behavior.

DISTRIBUTION AND HABITATS: The type series of P. maromandia was collected from the Maintsomalaza River, which is the local Malagasy name for the river just downstream from the confluence of the Manongarivo and Andranomalaza rivers, in the immediate vicinity of the town of Maromandia (14 ° 12 9 S, 48 ° 04 9 E) along the northwest coast of Madagascar (fig. 35). At the time of collection of the type series, corresponding to the dry season, this large, tidally influenced river was generally shallow, and the water was relatively clear, with a moderate current. The substrate comprised mostly sand and silt, although in places was extremely muddy. In addition to the Maintsomalaza River, local fishermen report catches of the new species from estuaries in the area and from the Andranomalaza River ( Nourissat, 1998).

Specimens that appear to be referable to P. maromandia have also been collected to the south of the Andranomalaza-Manongarivo drainage basin, from both the Maevarano River and Lake Andrapongy, a shallow floodplain lake located within the Ankofia River drainage basin. Although Sparks and Reinthal (1999) did not have material to examine from either locality at the time of their description, Sparks (personal obs.) did observe a dried/smoked specimen from Lake Andrapongy during a visit in 1994, which seemed to resemble P. maromandia in terms of body shape and the presence of a prominent vertical barring pattern. Subsequently, I have been able to examine two specimens (one dried/smoked) from that locality (MHNG 2537.60, 93.7–130.0 mm SL) collected by de Rham and Nourissat in 1991. Although the single specimen that is not dried is only a juvenile (93.7 mm SL), and much smaller than any available specimen of P. maromandia from the Andranomalaza- Manongarivo drainage basin, I an unable to find any morphological features to distinguish this individual from topotypic P. maromandia (apart from body depth, which is likely an artifact of its smaller size). Regrettably, de Rham and Nourissat (2004: 109) report that this population is now presumed extinct. No preserved material from the Maevarano River seems to have been deposited in museum collections (de Rham, personal commun.), although the color photographs presented in de Rham and Nourissat (2004: 107–108) closely resemble P. maromandia in the possession of a prominent vertical barring pattern and vivid red pigmentation on the flanks. In the absence of evidence to the contrary, the Maevarano River population is herein con- sidered to be conspecific with P. maromandia . In summary, the range of P. maromandia includes, from north to south, the Andranomalaza-Manongarivo drainage basin, the Maevarano River, and formerly, Lake Andrapongy, within the Ankofia drainage basin. The species has never been collected from the Ankofia or Anjingo rivers proper, and Lake Andrapongy presumably represented its southern range limit. Based on collection locality data, P. maromandia appears limited to the lower reaches of large tidally influenced rivers, estuaries, and shallow floodplain lakes (i.e., Lake Andrapongy) that experience periodic connections to the sea during the wet season ( Sparks and Reinthal, 1999; de Rham and Nourissat, 2004). Apart from the former Lake Andrapongy population, the species has not been collected upstream of tidally influenced sections of rivers within its range.

Paretroplus maromandia exhibits a very distinct pigmentation pattern and coloration. The additional populations from the Maevarano River and Lake Andrapongy, which were not part of the type series, share these distinctive features. Tissue vouchers were not available from either of these populations, and they could not be included in molecular phylogenetic analyses. Regardless, specimens from these two populations cannot currently be distinguished from topotypic P. maromandia on the basis of apomorphic morphological features and, therefore, there is no evidential basis for considering them not to be conspecific with P. maromandia .

Given that P. maromandia has only recently been discovered and that the surrounding region is poorly surveyed for freshwater fishes, little is known regarding the current status of this species. If P. maromandia can tolerate estuarine conditions, as indicated by collection-locality data, including the type locality, this species may not be as critically endangered as other members of Paretroplus , which exhibit highly restricted inland distributions, frequently comprising but a single basin. It has been hypothesized that estuaries serve as refugia for native Malagasy species ( Reinthal and Stiassny, 1991). Apart from P. maromandia , the only additional species of Paretroplus occurring almost exclusively in the tidally influenced portions of coastal rivers, or in estuarine conditions, is the widespread and still relatively common P. polyactis , which ranges along nearly the entire eastern coast of Madagascar (fig. 33). Regardless, throughout its limited range, P. maromandia seems nowhere abundant or even common and few collections have been made (there are only a handful of specimens in museum collections available for study, all of which have been examined here).

The region surrounding Maromandia, both to the north and south, is in need of comprehensive ichthyological surveys. The Tsaratanana Massif, the highest point in Madagascar (Maromakotro, 2876 m), is located inland to the north of Maromandia, and still contains large sections of intact rainforest that are difficult to access. The massif also creates what has been referred to as the Sambirano (micro-) climate (extending westward to the satellite island of Nosy Be), which is notably wetter than the dry western climate immediately to the south.

LOCAL NAME: Damba or damba mena. Damba is the Malagasy name used generally to refer to the deep-bodied species of Paretroplus throughout the northwest of Madagascar, and mena is Malagasy for ‘‘red’’, in reference to the vivid red flank pigmentation characteristic of this species.

ETYMOLOGY: Named for the town and general region from which the species was first collected. In English, the Malagasy prefix maro - translates as ‘‘many’’, and the Malagasy suffix - mandia means ‘‘to tread on or to go on a way/journey’’. The epithet, maromandia , is used as a noun in apposition.

RELATIONSHIPS AND DISCUSSION: Paretroplus maromandia is a member of Clade I, comprising all the deep-bodied and more or less disk-shaped members of Paretroplus (also including P. dambabe , P. maculatus , P. menarambo , P. petiti , and P. polyactis ), which is supported by three unambiguously optimized morphological features, two of which are unique and unreversed (features discussed and presented above) (fig. 1). Apart from pigmentation pattern and coloration, all members of this clade are morphologically very similar.

In the combined analysis of morphological features and nucleotide characters, P. maromandia is recovered in an unresolved polyt- omy with all other deep-bodied species of Paretroplus that have distributions restricted to western basins (fig. 1: Clade J). Monophyly of Clade J, comprising P. dambabe , P. maculatus , P. maromandia , P. menarambo , and P. petiti , is supported by two unambiguously optimized morphological features, the second of which is unique and unreversed: the presence of a blunt snout with a steeply sloping profile in lateral view and a distinctive lateral pigmentation pattern comprising alternating horizontal light and dark stripes (fig. 50). A lack of resolution within Clade J is due to the fact the P. petiti (known only from the formalin-fixed holotype) is included on the basis of morphological features only, which are insufficient for resolving relationships within this anatomically conservative clade. In a combined analysis of morphological features and nucleotide characters that specifically excluded P. petiti , P. maromandia is recovered as the sister taxon to P. menarambo (fig. 2), a result also obtained by the analysis of nucleotide characters only ( Sparks, 2004a; Sparks and Smith, 2004).

In addition to considerable differences in pigmentation pattern and coloration, P. maromandia is distinguished from P. menarambo by number of scales in the lateral line (39–41 vs. 34–38 in P. menarambo ), and usually by the number of gill rakers on the lower limb of the first arch (10 vs. 9 in P. menarambo ; 2 of 20 specimens had a count of 10 in P. menarambo ). Moreover, in P. menarambo the caudal fin is considerably more lunate, with elongate trailing dorsal and ventral filaments in large specimens, and a concave, rounded caudal margin. In P. maromandia , the caudal margins of both the soft dorsal and anal fins are deep bodied and notably rounded compared to congeners of similar standard length (i.e.,. 100 mm SL; figs. 49–50), a feature it shares only with P. maculatus . In similarly sized specimens of P. maculatus , however, the caudal margin of the anal fin is somewhat more pointed than in P. maromandia .

Paretroplus maromandia and P. menarambo are the only species of Paretroplus with a dorsal-fin ray count that exceeds 20 (range 20–23 in P. maromandia and 19–21 in P. menarambo ). Otherwise, only P. maculatus has a dorsal-fin ray count that reaches 20 (range 16–20).

Paretroplus tsimoly Stiassny, Chakrabarty, and Loiselle, 2001 Figures 48, 51–52; plate 1I; table 10

HOLOTYPE: AMNH 229558 , 140.3 mm SL, adult female; northwestern Madagascar: Majunga (5 Mahajanga) Province: Betsiboka River drainage basin: Akalimilotrabe River (5 Kalamilotra or Kamilotra River ) at village of same name, ca. 43 km northwest of Maevatanana : 16 ° 48 9 8.0 0 S, 47 ° 00 9 57.0 0 E: alt. 318 m a.s.l.; P. V. Loiselle , 7-VI-1997.

PARATYPES: AMNH 229559 , 2 ex., 1 ex. C&S, 99.3–138.1 mm SL, larger specimen adult female, smaller indeterminate sex ; data as for holotype. MNHG 2609.44 , 1 ex. 127.2 mm SL, adult female ; data as for holotype ; not examined. UMMZ 236893 , 1 ex., 123.6 mm SL, adult female ; data as for holotype .

ADDITIONAL NONTYPE MATERIAL EXAM- INED: AMNH 229556 , 2 ex., 110.1–129.5 mm SL, adult males ; northwestern Madagascar: Mahajanga Province: region of Maevatanana: Betsiboka drainage (but with no more specific locality data). AMNH 229557 , 1 ex. C&S, 52.0 mm SL, juvenile ; data as for holotype. MHNG 2640.038 , 2 ex., 133.6– 137.0 mm SL ; northwestern Madagascar: Mahajanga Province: Kamoro River at Antsalahina ; P. de Rham and J.-C. Nourissat, 20-X-2001. AMNH 238563 , 1 ex., 120.6 mm SL ; northwestern Madagascar: Mahajanga Province: Betsiboka River drainage basin: Kanero / Kamoro River ; JSS 37- 2003; J.-C. Nourissat, C. Toumy, and J. Andriamianamihaja, X-2003. AMNH 238564 , 1 ex., tissue voucher specimen, 89.7 mm SL ; northwestern Madagascar: Mahajanga Province: Betsiboka River drainage basin: Kamoro River ; JSS 38-2003, EE-1- 2003; J.-C. Nourissat, C. Toumy, and J. Andriamianamihaja , X-2003 .

DIAGNOSIS: Paretroplus tsimoly is distinguished from all congeners in life and preservation by the possession of markedly enlarged, lobed bluish-gray to bluish-black (in life) or dark bluish-gray (in preservation) lips in adults. In addition, P. tsimoly , is distinguished from all congeners except P. lamenabe and P. nourissati by the presence of two wide, dark brown to black midlateral bands that converge below the lateral mid- line, representing the second and third, or third and fourth, bars in series. Paretroplus tsimoly is further distinguished from both P. lamenabe and P. nourissati by a lateral pigmentation pattern in which the posteriormost five (or four, as the bar on the caudal peduncle is sometimes pale) bars on the flank are about equally prominent (vs. only central two bars strongly pigmented and prominent in P. lamenabe and P. nourissati ).

DESCRIPTION: Morphometric and meristic data presented in table 10. Morphological characteristics and general pigmentation pattern in life and preservative can be observed in figs. 51–52. A shallow-bodied and elongate Paretroplus belonging to Clade F, which also includes P. lamenabe and P. nourissati (fig. 1). Paretroplus tsimoly is recovered as the sister taxon to P. nourissati , and this clade is in turn the sister taxon to P. lamenabe .

TABLE 10

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

Preserved material of P. tsimoly available for study is more or less comparable in size to P. nourissati , reaching a maximum size of less than 150 mm SL, but P. tsimoly possesses a noticeably deeper and more robust body. Although the holotype (140.3 mm SL) is the largest preserved specimen, de Rham and Nourissat (2004: 124) state that P. tsimoly can be distinguished from P. nourissati by a larger adult size. In addition, Stiassny et al. (2001: 35) report that field observations reveal that P. tsimoly regularly exceeds 200 mm SL. Head pointed, snout and predorsal profile moderately curved, giving species a beaked appearance in lateral view. Premaxillary pedicels prominent, and notch present posterior to distal ends of ascending processes in some specimens. Dorsal body outline mildly curved, ventral outline mostly straight, except posteriorly. Caudal peduncle short, deep, and laterally compressed. No sexually dimorphic characters apparent in preserved material available for study.

Total vertebral count 31 or 32 (mode 31), with formulae of: 14 + 17 and 14 + 18 precaudal and caudal vertebrae, respectively.

Jaws isognathous. Lips quite fleshy and well developed. Fleshy medial flaps present 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 implant- ed. In upper jaw, tooth on either side of premaxillary symphysis greatly enlarged, remaining 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 number 12–14 in total, and those in lower jaw number 12 or 13 in total (also see Stiassny et al., 2001). Upperjaw teeth generally closely set, but also irregularly set. Lower-jaw teeth generally widely set, but also irregularly set in places.

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. LPJ weakly sutured, with few weak interdigitations on posteroventral margin. As in P. lamenabe and P. nourissati , apart from weak interdigitations, gap present posteriorly between right and left fifth ceratobranchial elements. Seven or eight robust tooth plates cover majority of dorsal surface of fourth ceratobranchial bones. Bases of some tooth plates becoming confluent with each other, but majority separate. Tooth plates not confluent with outer-row gill rakers of fourth ceratobranchial elements. 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 becoming molariform posteromedially, although most posteromedial teeth not molariform and retain an apical cusp. Dentition on third upper pharyngobranchial tooth plates hooked and bicuspid laterally and anteromedially. Anteromedial teeth enlarged, but not molariform. Dentition on second pharyngobranchial tooth plates hooked and bicuspid, and arrayed in two or three (two complete) rows.

Twelve to 14 triangular and somewhat elongate gill rakers arrayed along lower limb of first gill arch. Rakers on lower limb of first gill arch denticulate dorsomedially. 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 deep- bodied Paretroplus (Clade I) and P. kieneri 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 markedly elongate and slender, numbering nine or 10 (also see Stiassny et al., 2001).

Body covered with large, regularly imbricate, cycloid scales. Posterior field of lateral body scales thin and not ossified (fig. 23C). Posterior field of head scales ossified. Welldeveloped 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. 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 (fig. 23C). Interpelvic scale series present, terminal scale somewhat pointed distally. Lateral-line scales number 36–39 (mode 38). Chest scales noticeably reduced in size and embedded. 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. 23C). These scales of reduced size comprise a deep patch of several rows that is easy to visualize in lateral view. Four or five rows (sometimes also a partial sixth row, e.g., holotype with a single sixth-row scale) of scales on cheek. Opercle and subopercle scaled. Interopercle fully scaled posteriorly, asquamate anteriorly. Preopercle asquamate. Snout, lacrimal, and anterior portion of interorbital region asquamate. Scales on caudal fin reduced in size and extending posteriorly 2/3 to 3/4 length of fin on dorsal and ventral lobes, and about 1/3 length of fin medially.

Dorsal with XV–XVII spines, 13 or 14 soft rays. Anal with VII–VIII spines, 11 or 12 soft rays. Origin of dorsal fin at about level of, or slightly posterior to, vertical through dorsal margin of pectoral-fin insertion. Caudal fin emarginate, upper and lower lobes rounded. Pectoral fin broad and rounded at distal margin. Distal margins of soft dorsal and anal fins slightly produced and rounded. Lobes extend well beyond origin of caudal fin. Adducted pelvic fins extend to, or terminate just anterior to, anal-fin origin.

MISCELLANEOUS OSTEOLOGY AND ANAT- OMY: Large, well-developed exoccipital foramina present. Paired anterior gas bladder bullae present, with tough and thickened tunica externa, and anteriormost chambers firmly lodged in exoccipital recesses. 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 Paretroplus 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: Stiassny et al. (2001: 38) present a description of live coloration for both sexually quiescent (nonbreeding) and parental (breeding) adults. In addition, de Rham and Nourissat (2004: 125, 127) present several photographs of live aquarium specimens. Body ground coloration pale beige to orangish or reddish brown (nonbreeding, sexually quiescent individuals) to bright yellow, yellowish orange, or bright red (breeding, sexually active individuals), with series of six or seven vertical dark gray to blackish bars extending from the dorsoposterior margin of opercle to caudal peduncle and continuous from dorsum to ventrum. Posteriormost five (or four, as bar on caudal peduncle sometimes pale) bars on the flank are strongly pigmented and more or less equally prominent ( Stiassny et al., 2001: fig. 10; de Rham and Nourissat, 2004: 125). Lips, lower cheek, gular region, and belly dark grayish blue, dark purplish gray, or bluish black. Unpaired fins and pelvic fins grayish brown to reddish brown with red margins (nonbreeding), or charcoal with vivid red margins to more or less uniform bright red (breeding). Pelvic fins with white leading edge. Pectoral fins reddish gray, golden brown, or bright red. Grayish to dark gray triangular patch generally visible in pectoral-fin axil.

COLORATION IN PRESERVATIVE: Base body coloration gray to grayish brown. Generally with an overall speckled or blotchy appearance (figs. 51–52). Scale margins generally light gray on flanks, except ventrally where margins sometimes edged in dark brown. Six or seven darker gray to blackish vertical bars present on flanks. In specimens with six lateral bars, first bar at posterior margin of head and next bar in series are coalesced. Two midlaterally located bars, which represent the second and third or third and fourth bars in series, and those on posterior flank relatively strongly pigmented and about equally prominent (fig. 51). Bars on anterior flank quite faint. Body somewhat darker overall dorsally. Dark gray to blackish triangular patch present in pectoral-fin axil (faint in some specimens). Lateral line canals lined with melanophores in some specimens. Head, chest, and belly brownish gray. Snout, lacrimal, and interorbital region brownish gray to dark purplish gray. Lips, lower cheek, gular region, and belly dark charcoal gray to dark bluish gray or purplish gray in some specimens and light grayish brown to gray in others. Unpaired fins gray, grayish black, or black proximal to body, and light gray distally. Dorsal and anal fins sometimes black along margins. Pectoral fins medium gray proximal to body and light gray distally, or uniform gray. Pelvic fins gray proximal to body and charcoal gray distally, or uniform dark gray. Leading edge of pelvic fins whitish to light gray. Lateral barring more prominent in juveniles than adults. In juveniles the entire series of lateral bars is strongly pigmented.

DISTRIBUTION AND HABITATS: The type series of P. tsimoly was collected from the Ankalimilotrabe (5 Kalamilotra or Kamilo- tra) River, a tributary of the Betsiboka River, at the town of the same name and just upstream from where it is crossed by the main highway between Majunga (5 Mahajanga) and the capital, Antananarivo ( Stiassny et al., 2001; de Rham and Nourissat, 2004), approximately 43 km to the northwest of Maevatanana (fig. 48). The Akalimilotrabe River is connected to the Betsiboka River by a series of two floodplain lakes ( Stiassny et al., 2001). The substrate at the type locality is rocky, interspersed with patches of gravel and small stones, with numerous rocky outcrops creating a series of pools (also see Stiassny et al., 2001: 39; de Rham and Nourissat, 2004: 124–127). Given that the type locality is at the crossing of the major highway to the north of Madagascar from the capital, Antananarivo, it is surprising that the species was not discovered until the late 1990s. In the original description of this species, Stiassny et al. (2001: 39) note that although no specimens were collected, P. tsimoly was also observed in another north bank tributary of the Betsiboka River, the Boinakely River, about 33 km to the northwest of Maevatanana, where it is crossed by the major north-south highway, RN-4. De Rham and Nourissat (2004: 124) mention another population of P. tsimoly located in a lake (no name provided) three hours walk to the east of the Kalamilotra River. As far as I can determine, no specimens from this population were preserved.

Subsequent to the description of P. tsimoly, de Rham and Nourissat (2004) collect- ed very similar specimens in the upper reaches of the Kamoro River, a tributary of the extensive Betsiboka basin, near the village of Antsalahina, and which they refer to as Tsimoly (fig. 48). The upper Kamoro specimens share the characteristic enlarged and lobed lips and dark grayish-blue to bluishblack lips, lower cheek, belly, and gular region with topotypic P. tsimoly . Although de Rham and Nourissat (2004: 126) report that, in life, specimens from the upper Kamoro differ slightly in pigmentation pattern and coloration from topotypic P. tsimoly , in preservative specimens from these two populations are indistinguishable. Moreover, I am unable to find any anatomical differences to suggest that the Kamoro River population is a distinct species from topotypic P. tsimoly .

Interestingly, there appears to be hydrological evidence to support the hypothesis that the Kalamilotra and Kamoro populations are conspecific, as well as to explain species boundaries within Clade F. Paretroplus lamenabe , the sister species to the clade comprising P. nourissati and P. tsimoly , is known only from the lower reaches of the Mahajamba River, near the town of Androka, a locality well downstream from any connection with the Kamoro River (fig. 48). The Mahajamba is the next major basin to the north of the Betsiboka. As noted by Aldegheri (1972), the Mahajamba River is captured by the Kamoro River, a tributary of the Betsiboka, east of Tsaramandroso and near Morafeno, such that the upper 153 km of the Mahajamba flows almost entirely into the Betsiboka basin, whereas the distance from the capture zone to the mouth of the Mahajamba is 145 km. Capture of the upper Mahajamba by the Kamoro occurs well upstream of the range of P. lamenabe . Aldegheri (1972: 276) further notes that in periods of low water all of the upper Mahajamba’s water goes to the Kamoro, and from this point to the sea, the only water the lower Mahajamba receives is from its small tributaries. Thus, the lower reaches of the Mahajamba are effectively isolated from the Kamoro and Betsiboka basins.

Given the isolation of the lower Mahajamba from both the upper Kamoro and Betsiboka basins, it is not hard to imagine that populations of Paretroplus tsimoly in the upper Kamoro and the Kalamilotra (5 Ankalimilotrabe) rivers, both tributaries of the Betsiboka River, would be conspecific, whereas a distinct species, P. lamenabe , is present in the more or less isolated lower Mahajamba River. Somewhat surprising, however, is the fact that P. tsimoly is recovered as the sister taxon to P. nourissati , endemic to the Sofia basin located to the north of the Mahajamba, instead of P. lamenabe , which occurs in the adjacent basin to the north. Nevertheless, during exceptionally rainy years, the floodplains of the Mahavavy du Sud, Betsiboka, Kamoro, Mahajamba, and Sofia rivers are often contiguous (P. Loiselle in Courtney et al., 2004), which in part may explain the close relationship between Betsiboka and Sofia endemics.

The conservation status of P. tsimoly appears to be stable. De Rham and Nourissat (2004: 126) report that at least in the upper Kamoro River, P. tsimoly was relatively abundant when they visited the region in 2001 (and J.-C. Nourissat in 2003). However, judging from the small number of specimens collected from the type locality since the species was discovered in 1996, it would seem that the species is not abundant in the Kalamilotra River. I had briefly fished the Kalamilotra near the type locality in 1994 and did not collect any native cichlids.

LOCAL NAME: Tsimoly, which is pronounced ‘‘see-MOOL’’.

ETYMOLOGY: The Malagasy name of this species in the local Sakalava dialect is tsimoly .

RELATIONSHIPS AND DISCUSSION: Based on the simultaneous analysis of morphological features and nucleotide characters, P. tsimoly is recovered as a member of Clade F and as the sister taxon to P. nourissati (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. 51–52) 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).

In addition to the characteristically pigmented and hypertrophied, lobed bluish-gray to bluish-black (in life) or dark bluish-gray (in preservation) lips, a number of additional features serve to distinguish P. tsimoly from congeners. Paretroplus tsimoly is further distinguished from other members of Clade F, P. lamenabe and P. nourissati , by a lateral pigmentation pattern in which the posteriormost four or five bars on the flanks are strongly pigmented and about equally prominent (vs. only central two bars, representing the second and third or third and fourth bars in series, strongly pigmented in P. lamenabe and P. nourissati ), and by an overall speckled or blotchy lateral pigmentation pattern (vs. uniform base coloration).

Additionally, P. tsimoly can be distinguished from P. lamenabe by a shallower body (41.1%–46.8% vs. 47.0%–54.3% in P. lamenabe ) and by shorter pelvic fins that do not extend to anal-fin origin when adducted, and from P. nourissati by posterior extensions of the soft dorsal and anal fins that extend well beyond caudal-fin origin even in the smallest specimen examined (89.7 mm SL) in which the fins were not damaged (vs. posterior fin margins either not reaching, or, in some large specimens, extending to or rarely slightly beyond caudal-fin origin in P. nourissati ). Paretroplus tsimoly and P. nourissati can generally be distinguished from P. lamenabe by a lower tooth count in the upper jaw (12–14 in P. tsimoly and 13–16 in P. nourissati and respectively, vs. 15–18 in P. lamenabe ), and by less extensive squamation extending onto the membranes of the soft dorsal and anal fins.

As in other members of Clade F, P. tsimoly exhibits both a reduced number of interdigitations (which are shallow and weak) on the ventral suture and an indentation on the ventral face of the posterior horns of the LPJ. In addition, members of Clade F are united by narrow posterior horns of the LPJ, especially along the ventral ridges, and, posterior to the ventral suture, by a narrow gap between left and right fifth ceratobranchial elements.

Paretroplus damii and its sister taxon, Clade F, comprise Clade E. Members of Clade E are unique among Paretroplus in a number of features of scale morphology and squamation pattern, including: the posterior field of the flank scales is unossified and thin (fig. 23C), multiple rows of scales extend onto the membranes of both the soft dorsal and anal fins, the chest scales are markedly reduced in size and highly embedded, and multiple rows of scales of reduced size extend from the chest, dorsal to the pelvic fins and anus, to approximately anal-fin origin (fig. 23C). Members of Clade F can be distinguished from congeners, including P. damii , by exceedingly reduced (in size) and strongly embedded chest and belly scales, such that the ventral chest and belly appear asquamate.