Paretroplus kieneri Arnoult, 1960

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 : 77-90

Paretroplus kieneri Arnoult, 1960 Figures 36–38; plate 1D; table 5

Paretroplus cf. kieneri upper Kamoro River: de Rham and Nourissat, 2004: 111–112.

Paretroplus cf. kieneri Mahajamba River : de Rham and Nourissat, 2004: 111–112.

Paretroplus sp. of Lake Parinandrina: de Rham and Nourissat, 2004: 129–130.

HOLOTYPE: MNHN 1960-580 , adult female, 122.8 mm SL; northwestern Madagascar: Mahajanga (5 Majunga) Province: Mahavavy du Sud drainage basin: Lake Kinkony ; A. Kiener.

PARATYPE: MNHN 1960-581 , 1 ex., 117.7 mm SL ; data as for holotype .

ADDITIONAL MATERIAL EXAMINED: AMNH 97363 , 1 ex., 111.4 mm SL ; northwestern Madagascar: Mahajanga Province: Mahavavy du Sud drainage basin: Lake Kinkony ; P. V. Loiselle and local fishermen, 27-VI-1993. AMNH 229560 , 2 ex., 1 ex. C&S, 99.9–100.5 mm SL ; northwestern Madagascar: Mahajanga Province: Akalimilotrabe (5 Akalimilotra) River, north bank tributary of Betsiboka River, at Maevatanana : 16 ° 48 9 4.8 0 S, 47 ° 0 9 34.2 0 E ; P. V. Loiselle and local residents, 7-VI-1997. AMNH 238561 , ‘‘chocolat’’, 6 ex., 1 ex. C&S (in part), 109.0–134.0 mm SL ; northwestern Madagascar: Betsiboka River drainage basin: Kamoro River ; JSS 35-2003; J.-C. Nourissat, 2003. AMNH 238560 , ‘‘géant’’, 5 ex., 126.0– 160.0 mm SL ; northwestern Madagascar: Mahajanga Province: Mahajamba River ; JSS 32-2003; J.-C. Nourissat, 2003. AMNH 238567 , 1 ex. (dried), 94.0 mm SL ; northwestern Madagascar: Mahajanga Province: Betsiboka River drainage basin: Lake Parinandrina ; JSS 33-2003; J.-C. Nourissat, 2003. MHNG 2537.41 , 2 ex., 103.7– 114.2 mm SL ; Madagascar: northwest: P. de Rham and J.-C. Nourissat, X-1992. MHNG 2537.42 , 1 ex., 111.8 mm SL ; north- western Madagascar: Lac Sarodrano: 30 km north of Mampinkony : P. de Rham and J.-C. Nourissat, 22-X-1992. MHNG 2537.43 , 3 ex., 1 ex. C&S, 102.5–105.7 mm SL ; Madagascar: lac environment Boriziny ( Port Berger ): P. de Rham, X-II-1991. MNHN 1922- 0175 , 1 ex., 107.8 mm SL ; northwestern Madagascar: Ambatomainty. MNHN 1922- 0176 , 1 ex., 86.3 mm SL, northwestern Madagascar: Ambatomainty. MNHN 1922- 0177 , 1 ex., 82.0 mm SL, northwestern Madagascar: Ambatomainty. MNHN 1922- 0180 , 1 ex., 40.5 mm SL ; northwestern Madagascar: Ambatomainty. (Note: previous four specimens were misidentified as P. damii in the MNHN database.) MNHN 1960.581 , 3 ex. (preserved in lot with paratype), 131.1–142.2 mm SL ; northwestern Madagascar: Mahajanga Province: Mahavavy du Sud drainage basin: Lake Kinkony. MNHN 1966-1043 , 13 ex. ; northwestern Madagascar: Province of Majunga : Riviere Ankalamilotra. UMMZ 235018 , 5 ex., 1 ex. C&S, 70–108 mm SL ; northwestern Madagascar: Mahajanga Province: Betsiboka Riv- er drainage basin: near Ampijoroa Forestry station: Lake Ravelobe : 16 ° 18 9 31.0 0 S, 46 ° 48 9 59.0 0 E ; JSS 96-24; J. S. Sparks and K. J. Riseng, 14-VIII-1996. UMMZ 236592 , 4 ex., 1 ex. C&S, 84–102 mm SL ; northwest-

TABLE 5

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

ern Madagascar: Mahajanga Province: Mahavavy du Sud drainage basin: Lake Kinkony : 16 ° 05 9 38.0 0 E, 45 ° 51 9 37.0 0 E ; JSS 94-15; J. S. Sparks and K. J. Riseng, 14-VII-1994. UMMZ 239565 , 1 ex., 76 mm SL ; northwestern Madagascar: Mahajanga Province: Betsiboka River drainage basin: across from Ampijoroa Forestry station: Lake Ravelobe : 16 ° 18 9 34.0 0 S, 46 ° 48 9 59.0 0 E ; JSS 94-10; J. S. Sparks and K. J. Riseng, VII-1994. UMMZ 240353 , 2 ex. S, ca. 80 mm SL ; northwestern Madagascar; received from L. de Mason II- 1995.

DIAGNOSIS: A shallow-bodied Paretroplus distinguished from all congeners except P. gymnopreopercularis by the presence of a blotchy, mottled, and irregular (grayish, orange, olive, and brown) pigmentation pattern, the absence of vertical bars on the flanks, and by a fleshy snout that extends rostral to the lips and also ventrally to cover a portion of the upper lip. Paretroplus kieneri is distinguished from its sister taxon, P. gymnopreopercularis , by the presence of a more or less fully scaled preopercle, except along the ventral margin (vs. completely asquamate), a pointed and mildly convex predorsal profile (vs. blunt and strongly convex), and a second lacrimal plate that forms a part (albeit small) of the orbit margin (vs. second lacrimal plate excluded from orbit margin). The pigmentation pattern of P. kieneri is conspicuously blotchy, mottled, and irregular, whereas P. gymnopreopercularis is characterized by a weakly mottled and essentially uniform chain-link lateral pigmentation pattern, owing to darkly pigmented scale margins.

DESCRIPTION: Morphometric and meristic data presented in table 5. Morphological characteristics and general pigmentation pattern in life and preservative can be observed in figs. 36–37. A shallow bodied and comparatively small Paretroplus belonging to Clade H (fig. 1) that rarely exceeds 150 mm SL (e.g., largest wild caught specimens examined, from Mahajamba River and referred to as ‘‘géant’’ measured 160.0 mm SL). Head moderately pointed and snout curved (5 convex), lending a beaked appearance to the species. Predorsal profile moderately (Lac Kinkony population) to strongly (Mahajamba River population) curved. Dorsal body outline slightly curved, ventral outline mostly straight, except posteriorly. Caudal peduncle short, moderately deep, and laterally compressed. No sexually dimorphic characters apparent.

Total vertebral count 32 or 33 (mode 32), with formulae of: 15 + 17, 15 + 18, 16 + 16, and 16 + 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, 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 number four to seven on each side, and total 8–13. Teeth in lower jaw number four to six on each side, and total 8–11. Upper- and lower-jaw teeth widely set, except for central (i.e., symphyseal and adjacent) teeth in the lower jaw, which are closely set.

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, be- coming progressively enlarged medially; robust molariform teeth present posteromedially. LPJ well sutured, with numerous interdigitating sutures on posteroventral margin. Six or seven robust tooth plates cover majority of dorsal surface of fourth ceratobranchial bones. In some specimens, tooth plates on fourth ceratobranchial bones become fused. 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 laterally and anteromedially. Dentition on second pharyngobranchial tooth plates hooked and bicuspid, and arrayed in three to four rows.

Nine or 10 (mode 9) triangular, somewhat elongate, gill rakers arrayed along lower limb of first gill arch. Rakers weakly to moderately denticulate dorsomedially in larger individuals. All other lower-limb rakers (i.e., those on gill arches 2–4) short, strongly denticulate dorsally, and spherical in shape. Teeth on rakers of gill arches 2–4 as long or longer than raker itself. Epibranchial rakers on first gill arch elongate, numbering seven or eight.

Body covered with large, regularly imbricate, cycloid scales. Well-developed scale ridges (5 scale sheathing) present along dorsal- and anal-fin bases. Scale ridges free from spiny dorsal and anal fins, but becoming weakly attached to both soft dorsal and anal. Pelvic axillary scale present and well developed. Lateral-line scales number 33–40 (mode 38). Chest scales only slightly smaller than lateral body scales and somewhat embedded. Belly scales along ventral midline markedly reduced in size and much smaller than lateral body and chest scales. Four to six rows of scales on cheek. Opercle, subopercle, and interopercle scaled. Preopercle fully scaled except ventrally and along extreme posterior margin of shaft. Snout, lacrimal, and anterior portion of interorbital region asquamate. Scales on caudal fin reduced in size and extending posteriorly 2/3 to over 3/4 length of fin on dorsal and ventral lobes, and about 1/4 to 1/3 length of fin medially.

Dorsal with XVIII–XX spines, 13–15 soft rays. Anal with IX–XI spines, 11 or 12 soft rays. Origin of dorsal fin at about level of, or slightly posterior to, a vertical through pectoral-fin insertion. Caudal fin emarginate, trailing margins of upper and lower lobes slightly produced in larger individuals. Pectoral fin broad and rounded at distal margin. Distal margins of soft dorsal and anal fins weakly produced and pointed in larger specimens. Distal margins of soft dorsal and anal fins extend well past caudal-fin origin, even in smaller individuals. Pelvic fins extending just past anal-fin origin when adducted.

MISCELLANEOUS OSTEOLOGY AND ANAT- OMY: Large, well-developed exoccipital excavations present. Paired anterior gas bladder bullae with tough and thickened tunica externa, and anteriormost chambers firmly lodged in exoccipital recesses. Prominent 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 premaxillarymaxillary 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 a blotchy, mottled, and speckled yellowish orange, grey, and olive, to a speckled dark brown (usually a combination of these colors are present) (see de Rham and Nourissat, 2004: 112, for color photographs). The Malagasy name for this fish, kotsovato, refers to its resemblance to the coloration of rocks (5 vato in Malagasy). Coloration varies depending on drainage basin. Body pigmentation generally darker dorsally, but sometimes nearly uniformly pigmented. Head region and fins frequently gray, dark gray, or mottled dark brown and gray. No vertical bars present, even in juvenile specimens. Juveniles mottled golden and brown, and appearing camouflaged (under about 75 mm SL). Mottling in juveniles is in the form of large wavy blotches, as compared to much finer mottling and speckling in adults.

COLORATION IN PRESERVATIVE: Body ground coloration ranges from a blotchy, mottled, and speckled pale yellow, to golden brown or grayish brown (figs. 36–37). As for live specimens, coloration varies depending on drainage basin from which the specimen was collected. Body pigmentation generally darker dorsally. Fin coloration olive, light gray, or uniform dark gray. Portions of head and dorsal flanks frequently dark gray or dark grayish brown (e.g., fig. 37). No other distinctive markings present. Juveniles mottled golden and brown, and appearing camouflaged (and similar in pigmentation pattern to juvenile P. gymnopreopercularis ) due to presence of much larger and more well-defined blotches as compared to adults.

DISTRIBUTION AND HABITATS: Paretroplus kieneri exhibits a relatively widespread distribution in northwestern Madagascar, and it occurs in both rivers and the shallow and turbid floodplain lakes characteristic of this region (fig. 38). The species’ range from southwest to northeast includes the Mahavavy du Sud basin and Lake Kinkony (where it is presumed extinct), the lower Betsiboka basin, including Lake Ravelobe, the Akalimilotrabe (5 Kalamilotra) River, Lake Parinandrina (see below), the Ikopa basin lakes, the upper Kamoro River, the Mahajamba River, and the Bemarivo basin and Lake Sarodrano, which are within the Sofia River drainage system (de Rham and Nourissat, 2004). To the north and northeast of the Bemarivo drainage basin, P. kieneri is replaced by P. gymnopreopercularis , which appears to be endemic to the Amboaboa-Mangarahara river system, a tributary of the extensive Sofia River drainage basin (fig. 38). No members of Paretroplus are known to occur to the southwest of the Mahavavy du Sud drainage system.

While in Madagascar in late 2003, I received a single dried specimen from J.-C. Nourissat that he labeled, ‘‘ Paretroplus new species, Lac Amparimenidrino’’. I did not know at the time that this specimen was referred to as the ‘‘Red Damba’’ from Lake Parinandrina, which de Rham and Nourissat (2004: 129–130) hypothesized may represent a new species. We immediately obtained DNA-sequence data for this dried specimen, and included it in a family-level phylogenetic study of cichlid fishes, in which this individual was robustly recovered as a member of a clade comprising several populations of P. kieneri ( Sparks and Smith, 2004: fig. 1). It is interesting to note that de Rham, having only seen a picture of this fish, thought that it could be an emaciated P. kieneri (de Rham and Nourissat, 2004: 130) , a result we have been able to corroborate using nucleotide characters.

Although P. kieneri is comparatively widespread in distribution for species of Paretroplus , which generally exhibit remarkably restricted (allopatric) distributions, I have found it to be nowhere abundant, or even common for that matter. Interestingly, throughout much of its range, specimens we have collected over the past decade generally appear to be in poor physical condition, being both emaciated and heavily infested with parasites. De Rham and Nourissat (2004) report that the Lake Kinkony (topotypic) population of P. kieneri is extinct, with no specimens collected since the mid-1990s (I last collected a few specimens from Lake Kinkony in 1994). At that time, P. dambabe was still relatively common in Lake Kinkony (now also presumed extinct in the basin), although our team encountered only four specimens of P. kieneri in several full days of fishing. Fortunately, de Rham and Nourissat (2004) report that P. kieneri is still abundant in some habitats, and although they do not specify which populations, one can assume they are referring to the recently discovered upper Kamoro and Mahajamba river populations (both of which have been included in this study).

LOCAL NAMES: Kotsovato.

ETYMOLOGY: Named after A. Kiener, a French fisheries researcher who conducted numerous studies in Madagascar in the 1950s and 1960s, and who is credited with obtaining the type series.

RELATIONSHIPS AND DISCUSSION: Based on the simultaneous analysis of morphological features and nucleotide characters, P. kieneri is recovered as a member of Clade H and as the sister taxon to P. gymnopreopercularis (fig. 1). Clade H is supported by a single unambiguously optimized unique and unreversed morphological feature, the pres- ence of a blotchy, mottled, and speckled overall pigmentation pattern (figs. 36–37). This clade in turn is recovered as the sister taxon to Clade I, which comprises all deepbodied members of the genus (i.e., P. polyactis , P. maculatus , P. dambabe , P. petiti , P. menarambo , and P. maromandia ) (fig. 1). Prior studies based only on the analysis of nucleotide characters ( Sparks, 2004a; Sparks and Smith, 2004) recovered a clade comprising P. kieneri and P. gymnopreopercularis as the sister taxon to a clade comprising the deep-bodied members of Paretroplus ( Sparks and Smith, 2004: fig. 1), less P. polyactis . In these molecular phylogenetic studies, P. polyactis was recovered as the sister taxon to a clade comprising P. kieneri + P. gymnopreopercularis and the deep-bodied members of Paretroplus restricted to western basins (here Clade J).

Noting slight differences in body shape, coloration, and pigmentation pattern among the various populations spanning several drainage systems in northwestern Madagascar, some researchers have suggested that P. kieneri may represent a complex of species (e.g., de Rham and Nourissat, 2004). A recent molecular phylogenetic study, using both mitochondrial and nuclear nucleotide characters and which included representatives from four geographically distinct populations of P. kieneri distributed throughout northwestern Madagascar, was unable to resolve their intrarelationships ( Sparks and Smith, 2004: fig. 1). Results of another, less comprehensively sampled, recent phylogenetic study ( Sparks, 2004a) also based on nucleotide characters provided evidence for the existence of two species within this clade, P. kieneri and a closely related and previously undescribed taxon from the Amboaboa- Mangarahara River system, P. gymnopreopercularis , which is formally described in the current publication.

The phylogenetic analysis presented herein and based on the simultaneous analysis of morphological features and nucleotide characters further corroborates the hypothesis that P. kieneri and P. gymnopreopercularis are distinct species (fig. 1). Although the Amboaboa–Mangarahara River population, P. gymnopreopercularis , could be diagnosed on the basis of a single apomorphic morpho- logical feature, none of the other populations spanning the range of P. kieneri could be distinguished from topotypical P. kieneri by unique (5 apomorphic) morphological features. It is important to note that in the aforementioned molecular phylogenetic studies, the Bemarivo River (Lake Sarodrano) population could not be included due to lack of a tissue sample suitable for molecular studies. The Bemarivo River, like the Amboaboa-Mangarahara drainage to which P. gymnopreopercularis is endemic, is a tributary of the extensive Sofia River drainage basin. Although members of these two populations from within the Sofia River basin could be distinguished by squamation of the preopercle, it would be informative to see whether nucleotide characters also support this distinction. In conclusion, until the interrelationships of these populations can be sorted out, assuming they can be via either the use of additional, more quickly evolving nucleotide sequences or apomorphic morphological features, I believe that it is prudent to consider P. kieneri to represent a single, widespread species that spans several drainage basins in northwestern Madagascar.

In addition to differences in body shape (e.g., the body is noticeably deeper in the Lake Ravelobe individuals compared to all other populations examined), pigmentation pattern, and coloration among the various populations currently referred to P. kieneri , the degree of squamation on the preopercle is also to some extent variable among (and even within) the various geographic populations. Regardless, there is a distinct difference between the completely asquamate condition in P. gymnopreopercularis and all populations of P. kieneri , in which the preopercle is asquamate only along its ventral margin (e.g., in the Lake Kinkony and Lake Ravelobe populations). Both P. kieneri and P. gymnopreopercularis are characterized by a fleshy snout that extends rostrally anterior of the lips and also ventrally to slightly cover a portion of the upper lip.

Apart from degree of squamation on the preopercle (scaled in P. kieneri vs. asquamate in P. gymnopreopercularis ), other useful (but not diagnostic) features for distinguishing P. kieneri and P. gymnopreopercularis are overall coloration and shape of the caudal peduncle. Paretroplus kieneri generally exhibits dark gray coloration somewhere on the body (frequently on the head), whereas P. gymnopreopercularis is never dark gray, but orangish to yellow overall. The caudal peduncle in P. kieneri is notably shorter and deeper than in P. gymnopreopercularis (caudal peduncle length/depth 0.4%–0.6%SL [mean 0.5] in P. kieneri vs. 0.5%–0.8%SL [mean 0.7] in P. gymnopreopercularis ).

Paretroplus maculatus Kiener and Mauge´, 1966 Figures 35, 39–41; plate 1E; table 6

Paretroplus damii: Kiener, 1963 (in part): Pl. 15, illustration 2: Specimen is an amalgam of P. damii (body) and P. maculatus (only member of genus with large black lateral blotch).

Paretroplus cf. maculatus Lake Marajory : de Rham and Nourissat, 2004: 101–102, including photograph.

SYNTYPES: MNHN 1965-315 , 2 ex., 88.6– 124.5 mm SL ; northwestern Madagascar: Province of Majunga : Ambato-Boeny (Ambatoboemy [sic] in MNHN database) ; Kiener and Mauge´, IX-1964.

NOTES ON TYPE LOCALITY: In the original description of P. maculatus, Kiener and Maugé (1966: 69–70) state ‘‘cette espèce, originaire du nord-ouest de Madagascar, est couramment capturée dans la zone du lac Amparihibe-Sud , Tsaramandroso, and Kamoro’ ’; however, no specific type locality was provided for the two syntypes described. In the MNHN database, Ambato-Boeny (‘‘Ambatoboemy’’ [sic]) is listed as the collection locality. This most likely was because Ambato-Boeny is the nearest town of substantial size somewhat centrally located within this region of occurrence. Regardless, the specific basin in which the two syntypes were collected remains unknown .

ADDITIONAL MATERIAL EXAMINED: AMNH 97362 , 1 ex., 114.3 mm SL ; northwestern Madagascar: Mahajanga Province: Lake Ravelobe: Station Forestiere d’Ampijoroa ; P. V. Loiselle, R. Morris, and station staff, 25-VI-1993. AMNH 238559 , 2 ex., 117.0–128.5 mm SL ; northwestern Madagascar: Province of Mahajanga: Betsiboka drainage basin: near Ambato-Boeny: Lake Marajory ; JSS 31-2003; J. Andriamianami- haja and J.-C. Nourissat, X-2003 to XI-2003. AMNH 238568 , 2 ex., 106.3–112.7 mm SL ; northwestern Madagascar: Province of Mahajanga: Betsiboka drainage basin: near Ambato-Boeny: Lake Marajory ; JSS 43- 2003; J. Andriamianamihaja and J.-C. Nourissat, 2002. MHNG 2537.47 , 6 ex., 1 ex. C&S, 90.1–143.2 mm SL ; northwestern Madagascar: Province of Mahajanga: Ampijoroa Forestry Station: Lake Ravelobe ; P. de Rham and J.-C. Nourissat, 24-X-1992. UMMZ 235019 , 31 ex., 2 ex. C&S, 2 ex. S, 68–149 mm SL ; northwestern Madagascar: near Ampijoroa Forestry Station : Lac Ravelobe: 16 ° 18 9 31.0 0 S, 46 ° 48 9 59.0 0 E ; JSS 96-24; J. S. Sparks and K. J. Riseng, 14-VIII-1996. UMMZ 235020 , 31 ex., 2 ex. C&S, 48.4– 145.0 mm SL ; northwestern Madagascar: Province of Mahajanga: across highway from Ampijoroa Forestry Station: Lac Ravelobe : 16 ° 18 9 34.0 0 S, 46 ° 48 9 59.0 0 E ; JSS 94- 10; J. S. Sparks and K. J. Riseng, VII-1994. UMMZ 240354 , 1 ex. S, 85 mm SL ; northwestern Madagascar: Lake Ravelobe ; Laif Demason, 1995. UMMZ 240364 , 1 ex. S, 76 mm SL ; northwestern Madagascar: Lake Ravelobe ; J. S. Sparks and K. J. Riseng. UMMZ 242081 , 3 ex. S, 58–70 mm SL ; northwestern Madagascar: near Ampijoroa Forestry Station: Lake Ravelobe ; J. S. Sparks and K. J. Riseng , from aquarium shipment, 1996 .

DIAGNOSIS: A member of the deep-bodied clade of Paretroplus (Clade I) and distinguished from all congeners by the presence of a large and essentially square to rounded solid black pectoral (5 humeral) patch, located above and slightly posterior to the pectoral-fin base, and generally not extending dorsal to the upper branch of the lateral line.

DESCRIPTION: Morphometric and meristic data presented in table 6. Morphological characteristics and general pigmentation pattern in life and preservative can be observed in figs. 24B and 39–41. A deep-bodied, laterally compressed Paretroplus belonging to Clade I, which comprises 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). Head blunt and steeply sloping in lateral view. Predorsal profile rounded and markedly convex, particularly in larger individuals. Caudal peduncle short, deep, and laterally compressed. Apart from unpaired fins of males slightly more elongate

TABLE 6 Morphometric and meristic data for Paretroplus maculatus . For meristics, numerals in parentheses indicate number of specimens examined with that count. (S) indicates counts corresponding to syntypes (n = 2).

and pointed distally than females of comparative standard length, no additional sexually dimorphic features readily apparent.

Total vertebral count 32 or 33 (mode 33), with formulae of: 14 + 18, 14 + 19, 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 accom- modate enlarged upper symphyseal teeth. Teeth in upper jaw usually number six to eight on each side, and total 13–14. Teeth in lower jaw number five or six on each side, and total 10–11. Upper-jaw teeth widely set and evenly spaced, lower-jaw teeth widely set and irregularly spaced.

Upper and lower pharyngeal tooth plates well developed and dentition robust. Dentition on lower pharyngeal tooth plates (5 lower pharyngeal jaws [LPJ] or fused fifth ceratobranchial tooth plates) hooked and bicuspid both laterally and anteriorly, becoming progressively enlarged medially; robust molariform teeth present posteromedially. LPJ well sutured, with numerous interdigitating sutures on posteroventral margin. 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. 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 to 11 (mode 10) elongate triangular gill rakers arrayed along lower limb of first gill arch. Ceratobranchial rakers on first gill arch edentate in smaller specimens, becoming denticulate dorsomedially in larger individuals. All other lower-limb rakers (i.e., those on gill arches 2–4) short, stout, and strongly denticulate dorsally. Epibranchial rakers on first gill arch somewhat elongate and number 10 or 11.

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. Interpelvic scale elongate and pointed. Lateral-line scales number 34–39 (mode 38). 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. Five or 6 rows of scales on cheek. Opercle, subopercle, and interopercle scaled. Preopercle ranges from asquamate only ventrally and along posterior margin of shaft to more or less completely 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 1/4 to 1/3 length of fin medially.

Dorsal with XV–XVIII spines, 16–20 soft rays. Anal with VIII–X spines, 13–16 soft rays. Origin of dorsal fin slightly anterior of vertical through pectoral-fin insertion. Cau- dal fin deep-bodied, strongly 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, particularly males. Pelvic fins extending slightly beyond anal-fin origin when adducted.

MISCELLANEOUS OSTEOLOGY AND ANAT- OMY: Large, well-developed exoccipital excavations 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. Prominent 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) and nearly to anterior margin of frontals (fig. 6C). Two distinct and wellseparated 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: Conspicuous alternating light (pale golden brown, yellowy green, or grayish green) and dark (dark olive to dark reddish brown) horizontal stripes present on flanks (de Rham and Nourissat, 2004: 90–91). Presence of large, essentially square to rounded, solid black pectoral (5 humeral) patch, usually four scale rows deep, located above and slightly posterior to pectoral-fin base. Black patch generally not extending dorsal to upper branch of lateral line (but see photograph in de Rham and Nourissat, 2004: 91). Flanks usually somewhat darker dorsally, but frequently uniform. Six or seven (bar on caudal peduncle is very faint to essentially lacking in larger specimens) broad, vertical, dark olive bars on flanks, extending from anterior region of trunk to caudal peduncle. Bars more prominent dorsally and midlaterally on flank, yet always faint; bars very obvious in juveniles, becoming less conspicuous and barely discernable in large adults. Patch of red spots on flanks reported in some specimens (de Rham and Nourissat, 2004: 89–90). Nape, postorbital region, and region below anterior spinous dorsal ranges from grayish brown to reddish. Opercular region and cheek golden brown, golden green, or reddish. Black interorbital bar present in adults. Interorbital region grayish to pale golden brown anterior to black bar. Gular region ranges from light gray or grayish blue to blackish. Ventral chest and belly grayish brown to black. Dorsal and anal fins olive or gray proximal to base, and reddish distally. Caudal fin charcoal anteriorly and medially, and reddish distally. Spines of dorsal fin red distally. Soft dorsal and anal fins, and caudal fin with vivid red terminal band. Pectoral fins dark gray to black. Pigmentation pattern of young fish (to about 30 mm SL) characterized by large goldenbrown blotches and mottling, such that fish appears camouflaged (e.g., see juvenile P. petiti [now P. dambabe ] illustrated by Kiener [1963: Pl. 16], which exhibits a similar pigmentation pattern).

COLORATION IN PRESERVATIVE: Similar to live coloration, except that reddish coloration on flanks, head, and fins is more or less lost in preservative. Conspicuous alternating light (pale golden brown or light olive) and dark (dark olive, dark grayish brown, or dark brown) horizontal stripes present on flanks. Large, more or less square to rounded, black pectoral (5 humeral) patch, present on anterior flank, above and slightly posterior to pectoral-fin base (figs. 24B and 39–41). Six or seven broad and faint dark olive bars on flanks; bars more conspicuous in juveniles and smaller individuals. Interorbital region, lacrimal, and snout grayish. Lips goldenbrown to grayish. Gular region, anterior chest, belly, interopercle, and ventral cheek dark brown to black. Pelvic fins black. Pectoral fins pale golden-brown to olive. Unpaired fins golden brown, brownish gray, or near uniform dark gray. Anal fin with black terminal band; sometimes anal fin almost uniform black. Dorsal fin usually with weak black terminal band. Base of caudal fin golden to dark grayish-brown. Young fish (to about 30 mm SL) characterized by large golden-brown blotches and mottling, such that fish appears camouflaged.

VISCERA AND DIET: Gut contents comprised entirely of macerated gastropods in all individuals examined. Paretroplus maculatus appears to feed exclusively on gastropods.

DISTRIBUTION AND HABITATS: Paretroplus maculatus is endemic to turbid, shallow floodplain lakes in the lower reaches of the Betsiboka and Ikopa drainage basins in northwestern Madagascar (fig. 35). The range of P. maculatus extends in the north from Lake Ravelobe and surrounding small rivers in the vicinity of Ampijoroa Forestry Station, located to the southeast of Mahajunga (5 Majunga), southward to a number of small lakes in the vicinity of Maevatanana (de Rham and Nourissat, 2002, 2004), including lakes in the region of Ambato-Boeni (Kiener and Mauge´, 1966: ‘‘dans la zone du lac Amparihibe-Sud, Tsaramandroso, and Kamoro’’; de Rham and Nourissat, 2004), which is more or less centrally located between Ampijoroa and Maevatanana. Lake Andimaka, located to the southwest of the town of Mahazoma, which itself is located to the southwest of Maevatanana, is the most southerly collection locality known for P. maculatus . De Rham and Nourissat (2004) hypothesize that the rapids of the middle courses of the Betsiboka and Ikopa rivers act as barriers to the upstream dispersal of P. maculatus , restricting it to the lakes (and small rivers, personal obs.) of the lower reaches of these drainage basins.

Based on collections I have made and those of colleagues spanning several years, P. maculatus appears to have suffered a severe decline in abundance in recent years throughout its range. Severe fishing pressure, habitat degradation, and the introduction of a number of exotic species, most notably the Asian snakehead, Channa sp. , are probably the largest factors accounting for the rapid attrition of P. maculatus . Another member of Paretroplus , P. kieneri , is also found throughout the range of P. maculatus . This species has become increasingly rare in recent years according to local fishermen, and is considered to be extinct in its type locality, Lake Kinkony.

LOCAL NAME: Damba.

ETYMOLOGY: The specific name maculatus (from the Latin maculae, meaning ‘‘spot’’) refers to the large black humeral patch, which is diagnostic of this taxon.

RELATIONSHIPS AND DISCUSSION: Paretroplus maculatus is a member of Clade I, comprising all of the deep-bodied and essentially disk-shaped members of Paretroplus (also including P. dambabe , 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). 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. maculatus 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 horizontal light and dark 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 morphologically conservative clade. In a combined analysis of morphological features and nucleotide characters that specifically excluded P. petiti , P. maculatus is recovered as the sister taxon to P. dambabe (fig. 2), a result also obtained on the basis of nucleotide characters only ( Sparks, 2004a; Sparks and Smith, 2004).

In a paper describing a new species of Paretroplus from Lake Kinkony ( Sparks, 2002a), I reported that P. petiti is currently known only from the holotype (MNHN 1928.282), a juvenile specimen. After communicating my findings regarding the status of P. petiti Pellegrin, 1929 , which I consider to be very closely related to, and possibly conspecific with, the fish later described as P. menarambo Allgayer, 1996 (see Sparks, 2002), J.-C. Nourissat collected and provided me with specimens (AMNH 238559 and AMNH 238568) from Lake Marajory in the Betsiboka-Kamoro drainage basin, which were labeled P. cf. maculatus . These specimens exhibit the large black humeral patch diagnostic of P. maculatus . They are also reported to possess red spotting on the flanks (de Rham and Nourissat, 2004), a feature lacking in other populations of P. maculatus . De Rham and Nourissat (2004: 100–101) refer to these specimens as ‘‘ Paretroplus sp. aff. menarambo Betsiboka-Kamoro’’ and discuss whether they may be referable to P. petiti . After examining the specimens, I disagree with this assessment, and consider the specimens to be clearly referable to P. maculatus , the only species with a black humeral blotch. (In an addendum to de Rham and Nourissat (2004) [dated August 2004], de Rham likewise considers these specimens to be conspecific with P. maculatus .) The holotype of P. petiti (MNHN 1928.282) is quite well preserved, particularly in terms of pigmentation pattern, and there is no trace of a humeral blotch (fig. 34). In the original description of P. maculatus, Kiener and Maugé (1966) list its range to include lake habitats in the regions of Amparihibe- Sud, Tsaramandroso, and Kamoro. The specimens in question from Lake Marajory were collected from within the Betsiboka- Kamoro drainage basin, and within the range attributed to this species in the original description.