Paretroplus menarambo Allgayer, 1996

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 : 90-97

Paretroplus menarambo Allgayer, 1996 Figures 35, 42–44; plate 1F; table 7

HOLOTYPE: MZUS ( MZS) 3653, 128.2 mm SL; northwestern Madagascar: Mahajanga Province: Potamasina: Sofia River basin: Lake Sarodrano ; J.-C. Nourissat and P. de Rham, 22-XI-1991. Not examined.

PARATYPES: MZUS ( MZS) 3654, 1 ex., 110.5 mm SL ; data as for holotype ; not examined. MNHN 1996-121 , 1 ex., 89.4 mm SL ; data as for holotype. MNHN 1996-122 , 1 ex., 92.8 mm SL ; data as for holotype .

ADDITIONAL MATERIAL EXAMINED: AMNH 97364 , 2 ex., 1 ex. C&S, 79.8– 124.0 mm SL ; northwestern Madagascar:

Mahajanga Province: Bemarivo River drainage (tributary of Sofia River): Lake Sarodrano ; P. V. Loiselle and local fishermen, 23- VI-1993. AMNH 226311 , 4 ex., 145–161 mm SL ; aquarium trade specimens from stock collected in northwestern Madagascar: Mahajanga Province: Bemarivo River drainage (tributary of Sofia River): Lake Sarodrano. MHNG 2537.49 , 4 ex., 95.0–185.4 mm SL ; northwestern Madagascar: Mahajanga Province: Bemarivo River drainage: 30 km north of Mampinkony: Lake Sarodrano ; P. de Rham and J.-C. Nourissat, 23-X-1992. UMMZ 233522 , 6 ex., 1 ex. S, 148.0– 160.00 mm SL ; northwestern Madagascar: Mahajanga Province: Bemarivo River drainage: near village of Sarodrano, approx. 30 km north of town of Mampikony : Lake Sarodrano ; aquarium specimens received from L. Demason, orig. collected by J. S. Sparks, K. J Riseng, and local fishermen. UMMZ 235013 , 1 ex., 84.5 mm SL ; northwestern Madagascar: Mahajanga Province: Bemarivo River drainage: near village of Sarodrano, approx. 30 km north of town of Mampikony: Lake Sarodrano : 15 ° 47 9 46.0 0 S, 47 ° 39 9 11.0 0 E ; JSS 96-23; J. S. Sparks, K. J Riseng, and local fishermen, 18-VII-1996. UMMZ 235014 , 28 ex., 2 ex. C&S, 72.0– 167.0 mm SL ; northwestern Madagascar: Mahajanga Province: Bemarivo River drainage: near village of Sarodrano, approx. 30 km north of town of Mampikony: Lake Sarodrano : 15 ° 47 9 46.0 0 S, 47 ° 39 9 10.0 0 E ; JSS 94-16; J. S. Sparks, K. J Riseng , and local fishermen, 19-VII-1994 .

DIAGNOSIS: A member of the deep-bodied clade of Paretroplus (Clade I) and distinguished from all congeners by the presence of a pinstriped lateral pigmentation pattern comprising numerous longitudinal series of prominent black spots on the flanks, owing to the presence of one large spot on each scale. Paretroplus menarambo is also unique among congeners in possessing a conspicuous black interorbital stripe and suborbital blotch.

DESCRIPTION: Morphometric and meristic data presented in table 7. Morphological characteristics and general pigmentation pattern in life and preservative can be observed in figs. 21D and 42–44. 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. maromandia , P. petiti , and P. polyactis ) (fig. 1). Head blunt and steeply sloping in lateral view. Predorsal profile

TABLE 7

Morphometric and meristic data for Paretroplus menarambo . For meristics, numerals in parentheses indicate number of specimens examined with that count. (P1) indicates counts corresponding to MNHN 1996-121 and

(P2) to MNHN 1996-122.

rounded and convex, particularly in larger individuals. Caudal peduncle short, deep, and laterally compressed. No sexually dimorphic characters apparent, although unpaired fins of males slightly more elongate and pointed distally than females of comparative standard length.

Total vertebral count 32–34 (mode 33), with formulae of: 15 + 17, 15 + 18, 15 + 19, and 16 + 18 precaudal and caudal vertebrae, respectively.

Jaws isognathous. Single row of spatulate unicuspid teeth in both upper and lower jaws (fig. 3B). Teeth laterally expanded, flattened at crown, and procumbently implanted. In upper jaw, tooth on either side of premaxil- lary symphysis greatly enlarged, other teeth graded in size laterally (fig. 5A). 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 six to eight on each side, and total 14–15. Teeth in lower jaw number three to five on each side, and total 6–10. (Note: a single individual examined with three teeth on each side in lower jaw, otherwise four or five teeth on each side for a total of nine or 10.) Upper- and lower-jaw teeth widely set and more or less evenly spaced.

Upper and lower pharyngeal tooth plates well developed and dentition robust. Denti- tion 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. LPJ well sutured. Suture extensive with numerous finger-shaped interdigitating sutures on posteroventral margin. Five to seven robust tooth plates cover majority of dorsal surface of fourth ceratobranchial bones. Tooth plates not confluent with outer-row (5 lateral) gill rakers of fourth ceratobranchial elements. Bases of some tooth plates becoming confluent, 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 molariform posteromedially, hooked and bicuspid laterally and anteromedially. Dentition on second pharyngobranchial tooth plates hooked and bicuspid, and arrayed in three or four rows.

Nine or 10 (mode 9) elongate triangular gill rakers arrayed along lower limb of first gill arch. Rakers weakly denticulate dorsomedially. All other lower-limb rakers (i.e., those on gill arches 2–4) short, robust, more or less spherical in shape, and strongly denticulate dorsally. Teeth on rakers of gill arches 2–4 elongate, thin, conical, and curved near tip. Epibranchial rakers on first gill arch elongate and comparatively robust, numbering eight or nine.

Body covered with large, regularly imbricate, cycloid scales. Well-developed ridges of scales (5 scale sheathing) present along dorsal- and anal-fin bases (fig. 22A). Scale ridges free from both spiny and soft portions of dorsal and anal fins. Pelvic axillary scale present and well developed. Terminal scale of interpelvic series robust, elongate, and round- ed, or blunt and squared off distally. Lateralline scales number 34–38 (mode 37). Chest scales only slightly smaller than lateral body scales and weakly embedded (fig. 23B). Belly scales quite small compared to other body scales and strongly embedded, those along ventral midline smallest. Six or seven rows of scales on cheek. Opercle, subopercle, and interopercle scaled. Preopercle scaled except anteroventrally, and shaft fully scaled. Snout, lacrimal, and anterior portion of interorbital region asquamate. Scales on caudal fin reduced in size and extending posteriorly 2/3 to well over 3/4 length of fin on dorsal and ventral lobes, and 1/4 to 1/3 length of fin medially.

Dorsal with XV–XVII spines, 19–21 soft rays. Anal with VII–IX spines, 15–17 soft rays. Origin of dorsal fin ranging from about level of, to slightly posterior of, vertical through pectoral-fin insertion. Caudal fin deep-bodied, strongly emarginate and crescent shaped, trailing margins of upper and lower lobes quite produced, particularly in larger individuals. Pectoral fin broad and rounded at distal margin. Distal margins of soft dorsal and anal fins produced and pointed in larger specimens. Pelvic fin extending just beyond anal-fin origin when adducted.

MISCELLANEOUS OSTEOLOGY AND ANAT- OMY: Large, well-developed exoccipital foramina present (fig. 8B). 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 (fig. 8B). 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 ) (fig. 3B). An additional, fully ossified, anal- and dorsal-fin pterygiophore, not associated with any fin rays, present terminally in both fins. Modified ‘‘prezygopophyses’’ present on anterior neural arches (fig. 18B).

COLORATION IN LIFE: Body ground coloration ranging from light gray to iridescent grayish green. Prominent black spots on flanks arranged in numerous longitudinal series, and alternating with light gray to grayish-green stripes, lend fish a pinstriped appearance, hence its common name in the aquarist literature, pinstripe damba (see de Rham and Nourissat, 2004: 98–100). Series of six or seven (anteriormost bar[s] frequently very faint in larger specimens) broad, vertical, dark charcoal to blackish bars on flanks, extending from anterior region of trunk to caudal peduncle. Lateral barring pattern very obvious in juveniles, becoming faint and less conspicuous, and frequently barely discernable, in large adults. Black interorbital stripe present as well as prominent black blotch on cheek below orbit. Head gray to iridescent grayish green, and generally somewhat darker dorsally. Snout and lacrimal gray. Unpaired fins gray, dark gray, or charcoal. Margins of dorsal, anal, and caudal fins vivid red, giving fish its Malagasy and scientific name menarambo , meaning ‘‘red tail’’ (see photos in de Rham and Nourissat, 2004: 100 and 133). Pectoral fins gray to grayish-green. Pelvic fins gray to charcoal. Breeding individuals exhibit intensification of dark vertical bars, but otherwise do not differ in coloration from sexually quiescent individuals.

COLORATION IN PRESERVATIVE: Body ground coloration grayish brown to dark brown. Paretroplus menarambo notably dark- er in preservation than in life. Alternating dark (dark brown) and light (lighter brown) horizontal-striping pattern evident on flanks. Dark brown horizontal stripes due to presence of dark spots on scales arranged in numerous longitudinal series, lending fish a pinstriped pigmentation pattern (figs. 21D, and 42–43). Pinstriped flank pattern obvious, but much less pronounced than in live fish. Dark spotting on scales also much less prominent than in life. Head generally uniform dark brown. Subopercle and (to lesser degree) interopercle golden in some specimens, otherwise dark brown as on rest of head. Series of six or seven broad, vertical, dark charcoal to blackish bars on flanks, extending from anterior region of trunk to caudal peduncle. Lateral barring pattern very obvious in juveniles (to about 85 mm SL), becoming faint and frequently barely discernable in large adults. Juveniles lighter overall in coloration (golden brown, not dark brown) and with seven easily visible lateral bars, extending from caudal margin of opercle to caudal peduncle (fig. 44).

DISTRIBUTION AND HABITATS: Paretroplus menarambo was until very recently considered extinct in the wild; no speci- mens had been collected since 1997 (de Rham and Nourissat, 2004). In mid- November of 2006, however, I received a message from Olaf Pronk in Madagascar stating that a friend of his (Jurgen Spannring) had collected what he considered to be P. menarambo from a single small lake about 30 km to the east of Port Berge´. Images of freshly caught specimens were included and the identity of P. menarambo could be easily confirmed. Prior to this, the species was only known from the type locality, Lake Sarodrano, although it was suspected that it might be present in similar floodplain lakes in the region, which remained poorly surveyed (fig. 35). Lake Sarodrano lies within the Bemarivo River floodplain between the towns of Mampinkony and Port Bergé in northwestern Madagascar. Lake Sarodrano is a typical floodplain lake characteristic of northwestern Madagascar. The lake is very shallow, extremely turbid, and lined and infested with large clumps of reeds. The Bemarivo River is a tributary of the extensive Sofia River drainage system, but the species is not reported from the Sofia. According to de Rham and Nourissat (2004: 99), who first discovered the species in November of 1991, local fishermen reported that P. menarambo once occurred in all of the Bemarivo River floodplain lakes, of which there are many, but that they had not collected specimens in recent years. Although P. menarambo apparently once occurred in lakes throughout the Bemarivo River drainage basin, up until November of 2006 the species had been collected by researchers only in Lake Sarodrano, locat ed just north of the town of Mampinkony. According to de Rham and Nourissat (2004), local fishermen confirmed the recent disappearance of P. menarambo from all of the Bemarivo floodplain lakes. Fortunately, the species still persists in similar habitats within the region, although it appears to have been extirpated from the type locality, Lake Sarodrano. In addition, colleagues and myself have exported specimens to the United States and Europe, where the species has done quite well in captivity and can now be obtained fairly easily from breeders.

LOCAL NAME: Menarambo.

ETYMOLOGY: The specific name menarambo is a compound Malagasy word that translates as mena (5 ‘‘red’’) and rambo (5 ‘‘tail’’) in reference to the vivid red caudalfin margin of mature specimens.

RELATIONSHIPS AND DISCUSSION: Paretroplus menarambo is a member of Clade I, comprising all of the deep-bodied and more or less disk-shaped members of Paretroplus (also including P. dambabe , P. maculatus , P. maromandia , 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 similar.

In the combined analysis of morphological features and nucleotide characters, P. menarambo is recovered in an unresolved polytomy 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 light and dark horizontal stripes. 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. menarambo is recovered as the sister taxon to P. maromandia (fig. 2), a result also obtained by the analysis of nucleotide characters only ( Sparks, 2004a; Sparks and Smith, 2004: fig. 1).

In addition to substantial differences in pigmentation pattern (pinstriped in P. menarambo vs. prominently laterally barred in P. maromandia ) and coloration (see respective descriptions in life and preservative), P. menarambo is distinguished from P. maro- mandia by number of scales in the lateral line (34–38 in P. menarambo vs. 39–41 in P. maromandia ), and generally by the number of gill rakers on the lower limb of the first arch (9 in P. menarambo [2 of 20 specimens had a count of 10] vs. 10 in P. maromandia ). Moreover, in P. menarambo the caudal fin is considerably more lunate and concave, with elongate trailing dorsal and ventral filaments in large specimens.

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

As discussed by Sparks (2002a), the holotype and only known specimen of P. petiti (fig. 34) is very similar in overall body shape and pigmentation pattern (in preservative) to P. menarambo Allgayer, 1996 (figs. 21D, and 42–43), a species historically restricted to lakes of the Bemarivo River floodplain within the Sofia River drainage system in northwestern Madagascar, but not occurring in the Sofia River itself (fig. 35). Despite being collected in the 1920s, the holotype of P. petiti is remarkably well preserved, particularly in terms of pigmentation pattern and coloration. In preservative, both P. petiti and P. menarambo are dark brown in overall coloration, with a lateral pattern characterized by numerous, thin, alternating light and dark horizontal stripes (cf. figs. 34 and 42). In preservative, no additional distinctive lateral markings are present in either of these species. In life P. menarambo is characterized by both a pinstriped lateral pigmentation pattern and broad red terminal bands in the unpaired fins. This unique pinstriped pattern is still obvious in many preserved specimens, whereas others appear more or less solid brown or grayish brown. Coloration in life for P. petiti is not known, and comparisons to P. menarambo cannot be made.

Pellegrin (1929) commented on the horizontal striping pattern mentioned above, but he did not note the presence of a series of broad vertical bars on the flanks of P. petiti . Pellegrin (1929) similarly made no reference to any additional distinctive markings in P. petiti . Although other members of the deepbodied clade of Paretroplus restricted to western basins (Clade J) possess this conspicuous alternating light and dark horizontal striping pattern, no species other than P. petiti and P. menarambo are dark brown with no additional lateral markings. As mentioned above, the vertical barring pattern on the flanks of P. menarambo is generally rather faint and difficult to discern in preservative, particularly in large adults. Vertical bars, if present at all, are very faint in the holotype of P. petiti , and although not mentioned in the original description may have faded considerably over time in preservative.

Specimens now assigned to P. dambabe Sparks, 2002a were historically (since at least the early 1960s) referred to P. petiti (e.g., Kiener, 1963; Kiener and Thérézien, 1963; Kiener and Mauge´, 1966). The type specimen of P. petiti , a juvenile of 82.8 mm SL, is much deeper bodied than any individual of P. dambabe of similar standard length, and is in fact deeper bodied than any individual of P. dambabe examined (57.9% in P. petiti vs. max. 57.1% SL in P. dambabe of nearly 200 mm SL). Members of Paretroplus generally become deeper bodied with increasing standard length. Nevertheless, the holotype of P. petiti is very similar in body depth to specimens of P. menarambo of similar standard length ( Sparks, 2002a: fig. 8). Sparks (2002a) discussed the possibility that P. petiti and P. menarambo are conspecific, but cautioned that further study is needed, which depends on acquiring additional material from the region of the type locality of P. petiti , which itself remains uncertain (see Sparks, 2002a). This is a difficult region of northwestern Madagascar to access, and no additional specimens have been collected to date (also see discussions above for both P. dambabe and P. maculatus ).