Paretroplus damii Bleeker, 1868

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 : 56-63

Paretroplus damii Bleeker, 1868 Figures 26–29; plate 1A; table 2

Paretroplus vandami: Sauvage, 1891 : pl. 46 (misnamed in caption; see footnote p. 446).

Paretroplus dami: Boulenger, 1899: 142 (does not mention or justify spelling change for species name).

Paretroplus dami: Pellegrin, 1904: 322 ; Pellegrin, 1933:115; Kiener and Mauge´,1966; Stiassny et al., 2001; Sparks, 2002a: table 1; Sparks, 2004a: fig. 3.

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

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

Paretroplus sp. Mahanara River: de Rham and Nourissat, 2004: 117–118.

SYNTYPES: RMNH 3939 , 1 ex., 69.2 mm SL ; northwestern Madagascar: island of Nosy Be: Lake Pambilao [‘‘Nossibe (Lacus Pambilao)’’] ; Van Dam and F. Pollen. RMNH 4478 , 1 ex., 58.2 mm SL ; northwestern Madagascar: island of Nosy Be : Lake Pambilao ; Van Dam and F. Pollen.

NOTES ON TYPE LOCALITY: Bleeker (1868: 311–314) originally described this species based on two specimens from ‘‘Nossibe

(Lacus Pambilao)’’, which most certainly refers to Lac Ampombilava on the satellite island of Nosy Be, located off of the northwest coast of Madagascar. This type locality has been questioned, given that no specimens referable to P. damii , other than those described by Bleeker in the type series, have subsequently been collected from the island of Nosy Be, despite substantial effort. De Rham and Nourissat (2004: 113–114) report that their Malagasy guide recently collected specimens of P. damii from Lake Djabala (an adjacent crater lake to the presumed type locality) on the island of Nosy

TABLE 2 Morphometric and meristic data for Paretroplus damii . For meristics, numerals in parentheses indicate number of specimens examined with that count. (S1) indicates counts corresponding to syntype RMNH 4478 and (S2) to syntype RMNH 3939. Note: In this and all subsequent data tables type material (if present) is included in number of specimens examined, and in range, mean, and standard deviation calculations.

Be, but neglected to preserve any material, concluding they were similar to specimens collected on the mainland.

NOTES ON TYPE SERIES: Although Eschmeyer (2006) lists two syntypes of Paretroplus damii under RMNH 3939, I was able to locate only a single specimen (69.2 mm SL) corresponding to this catalog number in the RMNH collections. The second syntype described by Bleeker (1868) is most likely a specimen cataloged under RMNH 4478 (the included label states ‘‘type’’ under the species name of ‘‘ Paretroplus dami ’’). Eschmeyer (2006), however, includes this individual within the syntypic series of Paretroplus polyactis ( Bleeker, 1878) , a taxon for which he lists two syntypes. An examination of this specimen, a juvenile, shows it to be clearly referable to P. damii , not P. polyactis (cf. tables 2 and 3). The handwritten labels included with both specimens are in identical handwriting, both stating ‘‘type’’ under the species name of ‘‘ Paretroplus dami ’’ (misspelling probably following Boulenger, 1899). Both specimens also agree in total length with that reported by Bleeker (1868) in the original description of the species (80’’’ and 95’’’), assuming that Bleeker reported total lengths (TL). RMNH 3939 measures 90.5 mm TL (caudal slightly damaged distally) and RMNH 4478 measures 76 mm TL (caudal also slightly damaged distally). In addition, fin spine/ray counts and lateral scale counts for both specimens also correspond well to those reported by Bleeker. It is unclear why Eschmeyer (2006) lists RMNH 4478 as a syntype of Paretroplus polyactis . The two specimens described as P. polyactis by Bleeker (1878) are reported as measuring 67’’’ and 89’’’ (presumably also referring to TL), neither measurement of which corresponds to that of RMNH 4478 (76 mm TL; 58.2 mm SL). The specimen catalogued under RMNH 4478 and measuring 58.2 mm SL is herein considered to be a syntype of P. damii .

ADDITIONAL MATERIAL EXAMINED: AMNH 231248 , 3 ex., 1 ex. C&S, 71.6– 139.0 mm SL ; Madagascar: Antsiranana Province: Mahanara River: main channel of Mahanara River at Antsirabe Nord, at bridge on Route N-5: 13 ° 38 9 29.4 0 S, 49 ° 57 9 48.6 0 E ; P. V. Loiselle, 9-X-2000. AMNH 231257 , 4 ex., 1 ex. C&S, 74.9– 88.5 mm SL ; Madagascar: Antsiranana Province: Mahanara River : main channel at Antsirabe-Nord, at bridge on route N-5: 13 ° 58 9 29.4 0 S, 49 ° 57 9 48.6 0 E ; P. V. Loiselle, 13-X-2000. AMNH 232428 , 2 ex., 48.0– 49.0 mm SL ; Madagascar: Antsiranana Province: Ifasy River at Ambodipont, Andraniaomby: adjacent oxbow lake: 13 ° 22 9 7.8 0 S, 48 ° 52 9 22.8 0 E ; P. V. Loiselle, 25-X-2001. AMNH 232438 , 3 ex., 77– 108 mm SL ; Madagascar: Antsiranana Province: Sambirano River at Ambodidimaka village : 13 ° 45 9 18.0 0 S, 48 ° 29 9 30.0 0 E ; P. V. Loiselle and villagers, 26-X-2001. AMNH 232445 , 3 ex., 20.0–25.0 mm SL ; Madagascar: Antsiranana Province: Ramena River below bridge of Ambanja-Bemanevika road: 13 ° 45 9 16.2 0 S, 48 ° 31 9 9.6 0 E ; P. V. Loiselle, J. Miandriaza, and local fishermen, 26-X-2001. AMNH 232457 , 4 ex., 129.5–149.0 mm SL ; Madagascar: Antsiranana Province: Mahanara River : ca. 4 km northwest of Antsirabe- Nord: 13 ° 57 9 18.0 0 S, 49 ° 56 9 12.0 0 E ; P. V. Loiselle and local fishermen, 31-X-2001. AMNH 232461 , 3 ex., 96.0–134.1 mm SL ; Madagascar: Antsiranana Province: Mahanara River : at Antsirabe-Nord just upstream of bridge over route N-5: 13 ° 58 9 29.4 0 S, 49 ° 57 9 48.6 0 E ; P. V. Loiselle and local fishermen, 1-XI-2001. AMNH 232473 , 2 ex., 41.5– 84.0 mm SL ; Madagascar: Antsiranana Province: Ramena River at Antseva village : 13 ° 42 9 42.4 0 S, 48 ° 34 9 9.4 0 E ; local fisherman and C. J. Raxworthy, 1-IV-2001. AMNH 236145 , 112.5 mm SL ; Madagascar: Antsiranana Province: Mahanara River , main channel at Antsirabe-Nord ; P. V. Loiselle. AMNH 236154 , 2 ex., 52.4–110.8 mm SL ; Madagascar: Mahajanga Province: Anjingo River : main channel below bridge on Antsohihy-Bealanala Road ; P. V. Loiselle, 2004. MHNG 2537.44 , 7 ex., 91.4–162.2 mm SL ; Madagascar: Mahajanga Province: Riviere Ankofia (5 Anjingo) or Lac Andrapongy, environment Antsohihy: P. de Rham and J.-C. Nourissat, 17–19-X-1992. MHNG 2537.45 , 6 ex., 98.6–154.3 mm SL ; Madagascar: Mahajanga Province: Lac Andrapongy, environment Antsohihy: P. de Rham and J.-C. Nourissat, 17-X-1992. MHNG 2640.040 , 1 ex., 141.0 mm SL ; northern Madagascar: Ansiranana Province (Diego Suarez): Ramena River, near mouth of Sambirano River ; P. de Rham and J.-C. Nourissat, X-1999. MNHN 1931-0225 , 1 ex., 69.0 mm SL ; Antsiranana Province: Sambirano: Coll. by Waterlot. UMMZ 233523 , 14 ex., 6 ex. S, 123–179 mm SL ; northwestern Madagascar: Mahajanga Province: probably Anjingo Riv- er. UMMZ 235021 , 37 ex., 1 ex. C&S, 31– 138 mm SL ; northwestern Madagascar: Mahajanga Province: east of Antsohihy: Anjingo River (upstream and downstream of bridge over RN31 ): 14 ° 50 9 41.0 0 S, 48 ° 14 9 38.0 0 E ; JSS94-19; J. Sparks and K. Riseng, 28-VII- 1994. UMMZ 235022 , 34 ex., 1 ex. C&S, 82– 170 mm SL ; northwestern Madagascar: Mahajanga Province: northeast of Antsohihy : Lac Andrapongy, north basin: 14 ° 41 9 49.0 0 S, 48 ° 07 9 54.0 0 E ; JSS94-21B; J. Sparks and K. Riseng, 1994. UMMZ 235023 , 51 ex., 3 ex. C&S, 9–182 mm SL ; northwestern Madagascar: Antsiranana Province: north of Ambanja: Andranomaloto River (tributary of Mananjeba River ): 13 ° 12 9 46.0 0 S, 49 ° 10 9 34.0 0 E ; JSS96-22; J. Sparks and K. Riseng, 15-VII-1996. UMMZ 239543 , 5 ex., 1 ex. C&S, 56–72 mm SL ; northwestern Madagascar: Mahajanga Province: northeast of Antsohihy: Anjingo River at RN31: 14 ° 50 9 39.0 0 S, 48 ° 1 9 39.0 0 E ; JSS94-54; J. Sparks and K. Riseng, 15-XI-1994. UMMZ 239544 , 5 ex., 73–158 mm SL ; northwestern Madagascar: Antsiranana Province: market purchase near Ambanja: J. Sparks and K. Riseng, VII-1996. UMMZ 240355 , 1 ex., S, 86 mm SL ; no locality data, aquarium specimen.

DIAGNOSIS: A Paretroplus belonging to Clade E, comprising P. damii , P. nourissati , P. tsimoly , and P. lamenabe , and distinguished from all congeners by the presence of a triangular, black pectoral-axil patch in combination with the absence of a series of vertical bars on the flanks. In life, unstressed individuals can easily be distinguished from all congeners by the presence of a broad, vertical pale yellow band on the anterior half of the flanks. Young P. damii are unique among congeners in the possession of a dark blotch, surrounded by a hyaline ring, posteriorly on the soft dorsal fin near its base. In preservation, P damii is the only member of Paretroplus that exhibits a solid grayishbrown to brown base coloration and the absence of vertical barring.

DESCRIPTION: Morphometric and meristic data presented in table 2. Morphological characteristics and general pigmentation pattern in life and preservative can be observed in figures 26–28. A relatively shallow bodied Paretroplus belonging to Clade E: fig. 1), which also includes P. nourissati , P. tsimoly , and P. lamenabe . Paretroplus damii is the largest and most deep-bodied member of this clade, and specimens of nearly 40 cm TL have been collected (de Rham and Nourissat, 2004). Head moderately pointed, snout straight to slightly curved, and predorsal profile moderately curved. Dorsal body outline slightly curved, ventral outline mostly straight (except posteriorly). Caudal peduncle short, deep, and laterally compressed. No sexually dimorphic features apparent.

Total vertebral count 31 or 32, with formulae of: 14 + 17, 15 + 16, and 15 + 17, 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, 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 number four to seven on each side, and total 12–13 (a single specimen was examined from UMMZ 235023 [C&S] with a total of eight teeth in the upper jaw). Teeth in lower jaw number five to seven on each side, and total 11–13. Teeth in both upper and lower jaws frequently irregularly spaced and graded in size laterally.

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, but only posteromedially on LPJ. LPJ well sutured, with numerous interdigitating sutures on posteroventral margin. Seven to nine robust tooth plates cover majority of dorsal surface of fourth ceratobranchial bones. Fourth ceratobranchial tooth plates not confluent with outer-row gill rakers of these elements. Individual tooth plates separate at base in smaller individuals, and becoming fused in larger specimens. Dentition on fourth ceratobranchial tooth plates unicuspid or weakly hooked and bicuspid laterally, hooked and bicuspid medially (similar to lateral LPJ dentition) (fig. 13). 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 rows.

Eleven to 13 triangular, somewhat elongate, gill rakers arrayed along lower limb of first gill arch. Rakers edentate in small individuals, becoming weakly denticulate dorsomedially in larger specimens. All other lower-limb rakers (i.e., gill arches 2 through 4) short, triangular in shape (not spherical), and strongly denticulate dorsally. Epibranchial rakers on first gill arch elongate, numbering 9–11.

Body covered with large, regularly imbricate, cycloid scales. Posterior field (5 caudal margin) of flank scales thin, lacking circuli, and unossified, forming a flexible ‘‘flap’’. Well-developed ridges of scales (5 scale sheathing) present along dorsal- and analfin bases. Scale ridges free from spiny dorsal and anal fins, but becoming weakly attached to both soft dorsal and anal fins. Pelvic axillary scale present and well developed. Lateral-line scales number 35–39 (mode 37). Chest scales markedly reduced in size and embedded. Scales along ventral midline smallest. Four to five rows of scales on cheek. Opercle, subopercle, and interopercle scaled. Snout, lacrimal, and anterior portion of interorbital region asquamate. Preopercle asquamate ventrally and only scaled on dorsal portion of shaft. Scales on caudal fin reduced in size and extending posteriorly about 1/2 to 2/3 length of fin on dorsal and ventral lobes, and 1/4 to 1/3 length of fin medially.

Dorsal with XVIII–XX spines, 11–16 soft rays. Anal with IX–XI spines, 9–12 soft rays. Origin of dorsal fin at about level of vertical through pectoral-fin insertion. Caudal fin weakly emarginate, and upper and lower lobes broad and more-or-less rounded. Pectoral fin broad and rounded at distal margin. Distal margins of soft dorsal and anal fins at most weakly produced and pointed in larger specimens. Pelvic fin extending to about level of anal-fin origin.

MISCELLANEOUS OSTEOLOGY AND ANAT- OMY: Large, well-developed exoccipital foramina present (figs. 7C, 8A, and 9B). Paired anterior gas bladder bullae with tough and thickened tunica externa, anteriormost chambers firmly lodged in exoccipital recesses. Characteristic excavation (5 supraoccipital notch of Stiassny et al., 2001) along posterior margin of supraoccipital either poorly developed or lacking (i.e., notch may be lacking entirely or, if present, ranges from shallow, narrow, and poorly developed in riverine populations to moderately pronounced in deeper-bodied, lacustrine populations). 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 ranges from golden brown, to grayish, to dark brownish olive (de Rham and Nourissat, 2004: 114–117). Triangular black patch present in the pectoral-fin axil, and visible in lateral view dorsal to pectoral-fin base. Fin coloration ranges from gray, to grayish brown, to dark brownish olive. Juveniles less than about 40 mm SL mottled (appear camouflaged) with characteristic large dark predorsal saddle and concentrated black blotch on posterior margin of soft dorsal fin (fig. 28). Unstressed individuals are dark gray to grayish brown and exhibit a wide, pale yellow band on the anterior flank. The head, dorsal opercle, cheek, interorbital region, and snout are also pale yellow in unstressed specimens. This pale yellow coloration quickly disappears if the fish is disturbed (see de Rham and Nourissat [2004: 116] for a photograph). In breeding individuals from west coast populations, this yellow region becomes brick red (P. Loiselle, personal commun.).

COLORATION IN PRESERVATIVE: Large adults dark grayish brown to dark brown, and smaller specimens golden brown to gray; body somewhat darker dorsally, particularly in smaller individuals (fig. 27). Prominent triangular black patch present in pectoral-fin axil and extending dorsal to pectoral-fin base (fig. 25A). Wide, pale (yellowish) lateral band present in some large adults. Juveniles less than about 40 mm SL mottled gray, grayish brown, or golden brown, with concentrated black blotch present on posterior margin of soft dorsal fin (fig. 28).

DISTRIBUTION AND HABITATS: Widespread in northwestern and far northern Madagascar (fig. 29). Along the northwestern versant of the island, the known range of P. damii extends in the south from the Anjingo-Ankofia River basin and Lake Andrapongy northward to the rivers (Sahinana and Sampiana) draining the western slopes of the Montagne d’Ambre massif in far northern Madagascar (de Rham and Nourissat, 2004). The species has also recently been collected by Paul Loiselle (NY Aquarium/WCS) from the middle reaches of the eastward draining Mahanara River to the north of Sambava in northeastern Madagascar (fig. 29, gray circle). The geographic range of P. damii is noteworthy in that it is the only member of the genus that occurs in both eastward and westward draining basins.

LOCAL NAMES: Damba, damba mena, lamena, dridrimena, ventitry. De Rham and Nourissat (2004) also list filaopisaka and loakapisaka as local common names.

ETYMOLOGY: Named in honor of the Dutch naturalist Van Dam, who along with F. Pollen, collected the type material.

RELATIONSHIPS AND DISCUSSION: Paretroplus damii is the sister taxon to a clade (Clade F) comprising P. tsimoly , P. nourissati , and P. lamenabe (fig. 1). Clade F is diagnosed by the presence of two prominent and converging midlateral bands (several fainter bands are also present on the flanks) and elongate nasal bones, features that are both absent in P. damii (cf. figs. 24C and 25–27). 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 present- ed above) (fig. 1).

It has been suggested that specimens historically assigned to P. damii , as well as populations of morphologically similar fishes recently discovered in far northern and northeastern Madagascar, represent more than one valid species, and possibly as many as three (P. Loiselle, personal commun.). These allegedly unique species have been referred to as Paretroplus sp. ‘‘dridrimena’’ (far northern Madagascar) and P. sp. ‘‘ventitry’’ (Mahanara drainage, northeastern Madagascar) (de Rham and Nourissat, 2002, 2004). Paretroplus sp. ‘‘dridrimena’’ is reported from the western slope of the Massif d’Ambre, and the Mananjeba, Mahavavy du Nord, and Ifasy rivers, all western drainages in northwestern Madagascar. Paretroplus sp. ‘‘ventitry’’ has to date been collected only in the Mahanara River drainage in northeastern Madagascar, north of the Masoala Peninsula.

The type locality for P. damii is a crater lake (Lake Ampombilava; recorded by Bleeker, 1868, as ‘‘Nossibe (Lacus Pambilao)’’) located on the island of Nosy Be, which lies only a few kilometers offshore from mainland Madagascar between the outlets of the Sambirano and Mananjeba rivers and their mainland tributaries (comprising a portion of the putative range of P. sp. ‘‘ dridrimena ’’, which extends northward to rivers draining the western slopes of Montagne d’Ambre) along the northwestern coast of Madagascar. During the course of this study, specimens from populations that represent both P. sp. ‘‘dridrimena’’ and P. sp. ‘‘ventitry’’ have been examined. Numerous lots of P. damii were also examined from throughout its range. No apomorphic features were identified to differentiate these allegedly distinct northern populations from P. damii . Given the close proximity to the type locality of P. damii and lack of apomorphic diagnostic features, specimens that have been referred to P. sp. ‘‘dridrimena’’ are instead concluded to be conspecific with P. damii . Likewise, the Mahanara River population, P. sp. ‘‘ventitry’’, located on the northeastern coast and opposite to the Mananjeba River and its tributaries on the western coast, appears to be indistinguishable from P. damii on the basis of apomorphic morphological features. Therefore, it is concluded herein that the populations from northern Madagascar that have been referred to as ‘‘ventitry’’ and ‘‘dridrimena’’ do not warrant recognition as distinct species.

This statement should not be taken to indicate that additional study is not likely to reveal features useful for distinguishing among these geographic populations, and in particular between the single known eastcoast (‘‘ventitry’’) and various western populations (including ‘‘dridrimena’’). For example, despite some overlap in all of the following characteristics, in overall appearance, members of the east-coast ‘‘ventitry’’ population are in general more shallow bodied, the caudal peduncle is usually more elongate, and they are generally more darkly pigmented as adults than P. damii confined to western basins. In addition, the east-coast ‘‘ventitry’’ population differs in breeding coloration from all known west-coast populations of P. damii . Sexually active ‘‘ventitry’’ individuals are bright golden yellow and black, versus brick red and black in western populations of P. damii .

Nucleotide characters may prove useful in differentiating the various populations currently included in P. damii ; however, preliminary results to date indicate that nucleotide sequence divergence among these populations is quite low (W. L. Smith and J. S. Sparks, unpubl. data). In addition, based on DNA sequence data collected to date for other species shared between the mainland (i.e., occurring in the Mananjeba River and its tributaries) and Nosy Be (e.g., Ptychochromis oligacanthus and Paratilapia polleni [Sparks, 2003; Stiassny and Sparks, 2006]), landmasses which have repeatedly been connected during periods of low sea level resulting from Recent and Pleistocene glaciations, differences between mainland and Nosy Be populations of P. damii , should the latter be extant (e.g., see Loiselle, 2005), are anticipated to be minimal.