Paretroplus dambabe Sparks, 2002

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 : 118-124

Paretroplus dambabe Sparks, 2002 Figures 35, 53–54; plate 1J; table 11

Paretroplus petiti: Kiener, 1963 (in part); Kiener and Thérézien, 1963; Kiener and Mauge´, 1966 (in part); Stiassny et al., 2001.

Paretroplus cf. petiti: de Rham and Nourissat, 2002 .

Paretroplus nov. sp. ‘‘dambabe’’: Sparks and Stiassny, 2003: table 9.1.

HOLOTYPE: UMMZ 238724 , adult male, 169.4 mm SL; northwestern Madagascar: Majunga (Mahajanga) Province: south of Mitsinjo: Mahavavy (du sud) drainage basin: Lake Kinkony : 16 ° 05 9 37.7 0 S, 45 ° 51 9 37.4 0 E; JSS 94-15; J. S. Sparks, K. J. Riseng, and local Malagasy guides, 14–16 July 1994.

PARATYPES: UMMZ 199406 , 3 ex., 67.0– 113.0 mm SL ; ex. MNHN 60579 ; Madagascar. AMNH 232398 , 10 ex., 61.0– 170.0 mm SL ; data as for holotype. UMMZ 235024 , 29 ex., 40.2–225.0 mm SL, 3 ex. C&S ; data as for holotype .

ADDITIONAL NONTYPE MATERIAL EXAM- INED: All from Madagascar, Majunga (Mahajanga) Province, Mahavavy (du sud) drainage basin, Lake Kinkony : AMNH 11707 , 2 ex., 137.5–153.5 mm SL ; Archbold Expedition: A. L. Rand and P. A. Dumont, III-1931. AMNH 97371 , 1 ex., 77.4 mm SL ; P. V. Loiselle and local fishermen, 27-VI- 1993. MHNG 2537.48 , 6 ex., 83.6–166.4 mm SL ; Mitsinjo; P. de Rham and J.-C. Nourissat, 23-X-1992. MHNG 2537.72 , 5 ex., 1 ex C&S ; 74.3–89.0 mm SL; Mitsinjo; P. de Rham and J.-C. Nourissat, XI-1991. MNHN 1960-0579 , 3 ex., 125.5–179.9 mm SL ; A. Kiener. MNHN 1962-0239 , 2 ex., 23.3– 24.6 mm SL ; A. Kiener. MNHN 1965-0316 , 2 ex., 64.2–129.3 mm SL ; G. Petit. MNHN 1996-123 , 1 ex., 105.2 mm SL ; J.-C. Nourissat and P. de Rham .

DIAGNOSIS: A member of the deepbodied clade of Paretroplus (Clade I) and distinguished from congeners, in preservative, by pale yellow to golden body coloration in combination with a series of six or seven faint to barely discernable (particularly in

adults) vertical, dark-olive bars on the flank, and in the possession of uniform gray or charcoal-gray unpaired fins. In life, P. dambabe is unique among congeners in the possession of a grayish to bluish-green base coloration, which is accompanied by a highly variable amount of bright red pigmentation on the flank, particularly below the upper branch of the lateral line. Paretroplus dambabe is further distinguished from P. petiti , a species with which it has been erroneously associated for decades, by overall coloration (light yellowish olive vs. dark brown), brownish spotting (reddish in life) on the flanks in preservative, particularly anteriorly and below the lateral midline, and a shallow- er body (mean 52.8% [ranges from 48.3%– 57.1%] vs. 57.9% SL).

DESCRIPTION: Morphometric and meristic data presented in table 11. Morphological characteristics and general pigmentation pattern in life and preservative can be observed in figs. 53–54. A deep-bodied, laterally compressed Paretroplus belonging to Clade I, discussed by Sparks and Reinthal (1999) and Sparks and Smith (2004), and comprising all deep-bodied and essentially disk-shaped members of Paretroplus ( P. dambabe , P. maculatus , P. maromandia , P. menarambo , P. petiti , and P. polyactis ) (fig. 1). Comparatively a large Paretroplus , frequently exceeding 200 mm SL, and reportedly attaining 400 mm TL (Kiener, 1963), although large specimens now rare (see below). Head blunt and steeply sloping in lateral view. Predorsal profile rounded and convex, especially pronounced in larger individuals. Caudal peduncle short, deep, and laterally compressed. Females smaller than males, but no additional sexually dimorphic characters apparent.

Total vertebral count 31–33 (mode 32), with formulae of: 14 + 18, 14 + 19, 15 + 16, 15 + 17, and 15 + 18, precaudal and caudal vertebrae, respectively.

Jaws isognathous. 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, and other teeth 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 eight on each side, rarely five. Teeth in lower

TABLE 11 Morphometric and meristic data for Paretroplus dambabe . For meristics, numerals in parentheses indicate number of specimens examined with that count. (H) indicates count corresponding to holotype.

jaw number three to five on each side. Lowerjaw teeth irregularly spaced and sized.

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 (fig. 14B). LPJ well sutured, with numerous 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 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 laterally and anteromedially (fig. 11D). Dentition on second pharyngobranchial tooth plates hooked and bicuspid, and arrayed in three to four rows (fig. 11D).

Nine or 10 triangular, somewhat elongate, gill rakers arrayed along lower limb of first gill arch. Rakers edentate in small individuals (, 100 mm SL) and weakly denticulate dorsomedially in larger specimens. All other lower-limb rakers (i.e., those on gill arches 2– 4) short and strongly denticulate dorsally. Epibranchial rakers on first gill arch elongate, numbering 8–10.

Body covered with large, regularly imbricate, cycloid scales. 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. Pelvic axillary scale present and well developed. Lateral-line scales number 36–38 (mode 36). Chest scales only slightly smaller than lateral body scales and weakly embedded. Belly scales along ventral midline markedly reduced in size and embedded, much smaller than lateral body and chest scales. Four to six rows of scales on cheek. Opercle, subopercle, and interopercle scaled. Snout, lacrimal, and anterior portion of interorbital region asquamate. Preopercle scaled dorsally along shaft, and ventrally only along anterior margin. Scales on caudal fin reduced in size and extending posteriorly 2/3 to 3/4 length of fin on dorsal and ventral lobes, and 1/4 to 1/3 length of fin medially.

Dorsal with XVI–XVIII spines, 17–19 soft rays. Anal with VIII–X spines, 13–15 soft rays. Origin of dorsal fin slightly anterior of vertical through pectoral-fin insertion. Caudal fin emarginate and crescent shaped, trailing margins of upper and lower lobes 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 fins extending just to anal-fin origin in smaller specimens, and well beyond origin in larger individuals.

MISCELLANEOUS OSTEOLOGY AND ANAT- OMY: Large, well-developed exoccipital foramina present (fig. 10B). Paired anterior gas bladder bullae with tough and thickened tunica externa, and 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. 10B). Supraoccipital extending anteriorly over median frontal pores of neurocranium (nlf 0 of Barel et al., 1977) (fig. 10B). 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: Ground coloration grayish, grayish pink, or bluish green (see de Rham and Nourissat, 2004: 95, 97). Body generally darker dorsally, but sometimes uniform in coloration. Alternating light and dark horizontal striping pattern evident on flank. Six or seven broad, vertical dark olive bars extending from anterior region of trunk to caudal peduncle; bars very obvious in juveniles, becoming less conspicuous to nearly indistinguishable in large adults. A variable amount of vivid red pigmentation (5 spotting) present on flanks, particularly below upper branch of lateral line. Some individuals with each flank scale outlined in red, whereas in others, red pigmentation restricted to region below upper branch of lateral line. Nape and region below anterior spinous dorsal golden in juveniles. Black interorbital bar present in adults, and dark golden in juveniles. Dorsal and anal fins uniformly black proximal to base. Soft dorsal and anal fins with white distal margin. Caudal fin orangish anteriorly, black on median portion of fin, and with white terminal band. Pelvic fin black with white leading edge. Pectoral fin reddish orange proximal to base, white distally. Pigmentation pattern of young fish (, 30 mm SL) blotchy and mottled (see juvenile P. petiti [now P. dambabe ] illustrated by Kiener, 1963: Pl. 16).

COLORATION IN PRESERVATIVE: Ground coloration pale yellowish olive or light tan. Six or seven weak to barely discernable vertical, dark-olive bars on flank (fig. 53). Barring more conspicuous in subadults (, 150 mm SL) than adults ( Sparks, 2002a: fig. 3), and becoming obscure to barely visible in large adults (fig. 53). Pigmentation pattern of juveniles blotchy and mottled overall (fig. 54). Reddish coloration on anterior flank faded and appearing brownish in preservative. Alternating light and dark horizontal striping pattern obvious on flank. Unpaired fins and pelvic fins uniform charcoal gray. Pectoral fins olive proximal to base and more or less hyaline distally. Anterior interorbital region and snout light gray.

VISCERA AND DIET: Contents of gut in all individuals examined comprised entirely of crushed gastropods.

DISTRIBUTION AND HABITATS: Although Paretroplus dambabe is reported to have a slightly wider geographic range within the Mahavavy du Sud basin, preserved specimens are only available from Lake Kinkony, a large, shallow floodplain lake located to the southwest of Majunga in the northwestern part of the island (fig. 35). The species is also reported to occur in two smaller satellite lakes near the village of Antongomena (de Rham and Nourissat, 2004), and in the Mahavavy du Sud River (P. Loiselle, personal commun.). I have collected and examined material only from Lake Kinkony proper, and cannot confirm these additional localities. Lake Kinkony is a large (second largest lake in Madagascar, ca. 14,000 hectares), extremely shallow and turbid, oligotrophic floodplain lake characteristic of basins in northwestern Madagascar (Kiener, 1963). Similar lakes in northwestern Madagascar include lakes Andrapongy and Sarodrano. Several euryhaline and invasive marine species, including anguillids, carangids, Scatophagus , and Chanos , also inhabit the lake. The Lake Kinkony basin is moderately to highly disturbed and degraded. A substantial portion of the basin has been converted for rice cultivation and for grazing of livestock, and little original riparian vegetation remains in the basin.

According to local fishermen questioned in the mid-1990s, the species had suffered a dramatic decline in abundance in the preceding decade. Lake Kinkony is subjected to severe fishing pressure, especially from seasonal migrant fishermen who supply markets in the capital, Antananarivo. Regrettably, de Rham and Nourissat (2004: 93–96) report that P. dambabe is most certainly extinct in Lake Kinkony, but that in 1997 populations of P. dambabe were also discovered in two smaller satellite lakes, Andranobe and Lac de la Digue, situated near the town of Antongomena. Although they were unable to collect specimens in Lake Andranobe, which was under heavy fishing pressure, Lac de la Digue, an artificial lake used to store water for rice irrigation, still harbored a population of P. dambabe (de Rham and Nourissat, 2004) . Another member of Paretroplus , P. kieneri , also occurred in Lake Kinkony, which is the type locality for this species, but is also now reported to be extinct in the basin. In the mid-1990s, P. kieneri was extremely rare in Lake Kinkony, whereas a few specimens of P. dambabe could still be collected, but with substantial effort.

LOCAL NAME: Kotso, Damba.

ETYMOLOGY: Dambabe (pronounced ‘‘dambah bay’’) is a compound Malagasy word that translates as ‘‘large’’ or ‘‘big’’ (5 be) damba, in reference to the large size attained by this species relative to most other members of the genus. Damba is the Malagasy word used to refer to a number of species of Paretroplus throughout northwestern Madagascar. According to de Rham and Nourissat (2004: 88), damba was the name originally used by the Merina fish traders.

RELATIONSHIPS AND DISCUSSION: Paretroplus dambabe is a member of Clade I, comprising all the deep-bodied and essentially disk-shaped members of Paretroplus ( P. dambabe , P. maculatus , P. maromandia , 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). All members of this clade are morphologically similar apart from pigmentation pattern and coloration.

In the combined analysis of morphological features and nucleotide characters, P. dambabe is recovered in an unresolved polytomy with all other deep-bodied species of Paretroplus that have distributions confined 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. 53). 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 relation- ships within this morphologically conservative clade. In a combined analysis of morphological features and nucleotide characters that specifically excluded P. petiti , P. dambabe is recovered as the sister taxon to P. maculatus (fig. 2), a result also obtained on the basis of nucleotide characters only ( Sparks, 2004a; Sparks and Smith, 2004).

Although P. dambabe was only recently described ( Sparks, 2002a), since at least the early 1960s individuals of this species collect- ed from Lac Kinkony have erroneously been referred to P. petiti (e.g., Kiener, 1963; Kiener and Thérézien, 1963; Kiener and Mauge´, 1966). Based on results presented by Sparks (2002a), all specimens previously assigned to P. petiti that were collected in Lac Kinkony are determined instead to be members of P. dambabe . Paretroplus dambabe is readily distinguished from P. petiti in preservative by overall coloration (light yellowish olive vs. dark brown), brownish spotting (reddish in life) on the flanks, particularly anteriorly and below the lateral midline, and a shallower body (mean 52.8% [ranges from 48.3%–57.1%] vs. 57.9% SL). Paretroplus petiti is, therefore, currently only known from the holotype described by Pellegrin (1929).

As discussed by Sparks (2002a), there has been some confusion and misunderstanding regarding the type locality of P. petiti Pellegrin, 1929 , which in the original description is listed as the Maintimaso River in the province of Mahajunga, northwestern Madagascar. As far as I can determine, the type specimen of P. petiti (MNHN 1928-282) was collected from the Maintimaso River, a tributary of the westward flowing Betsiboka River, near the town of Maintimaso (16 ° 10 9 60 0 S, 46 ° 43 9 00 0 E), which is situated about 37 km southeast of Majunga (5 Mahajunga) (see fig. 35). The MNHN database, however, lists the collection locality of this specimen as ‘‘Ambila’’. No town with this spelling could be located in the vicinity of Majunga, although the town of Ambilo is located near the Betsiboka River, 35 km south of Majunga and to the northwest of Maintimaso. In addition, in the jar with the type specimen at the MNHN (Paris) was an old tag with the species name and also including the word ‘‘Ambanja,’’ a town located several hundred km to the north of Maintimaso and Mahajunga. No additional information with regard to either the Ambila or Ambanja localities could be located. As with many other fish species described from Madagascar in the latter part of the 19th century through the first half of the 20th century, collection localities are frequently ambiguous and questionable ( Sparks, 2002a). Nevertheless, it is clear from the information that is available that the type specimen of P. petiti was not collected from the Mahavavy du Sud drainage basin, to which P. dambabe is endemic, but instead was obtained from a tributary of the Betsiboka River, one of Madagascar’s largest drainage basins (fig. 35).

As noted by Sparks (2002a), the holotype of P. petiti , a juvenile, is considerably deeper bodied than P. dambabe of similar standard length, and is deeper bodied than any individual of P. dambabe examined, whereas it is similar in body depth to specimens of P. menarambo of similar standard length ( Sparks, 2002a: fig. 8). The holotype and only known specimen of P. petiti is remarkably well preserved and similar in body shape and pigmentation pattern, in preservation, to P. menarambo , a species restricted to floodplain lakes of the Sofia basin, northwestern Madagascar (cf. figs. 34 and 42–43). Both species are dark brown, with a pattern of alternating horizontal light and dark stripes on the flank. Although other deep-bodied members of Paretroplus (Clade J) possess this characteristic horizontal striping pattern, none other than P. petiti and P. menarambo are dark brown with no additional lateral markings apart from a faint vertical barring pattern in P. menarambo . Pellegrin (1929) commented on the horizontal striping pattern in P. petiti , but he did not mention any additional distinctive markings. Paretroplus menarambo is characterized in life by bright red terminal margins to the unpaired fins, whereas coloration in life for P. petiti is not known. Based on these observations, Sparks (2002a) hypothesized that P. petiti and P. menarambo might be conspecific, but deferred judgment until more material from the region of the putative type locality of P. petiti became available.

Paretroplus gymnopreopercularis , new species