Paretroplus lamenabe, Sparks, 2008

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 : 129-135

Paretroplus lamenabe , new species Figures 48, 58–59; plate 1L; table 13

Paretroplus aff. tsimoly Mahajamba : de Rham and Nourissat, 2004: 127–129.

Paretroplus sp. ‘‘Mahajamba’’: Sparks and Smith, 2004: fig. 1.

HOLOTYPE: MHNG 2676.30 (ex. MHNG 2640.039 ), holotype, 168.5 mm SL; northwestern Madagascar: Majunga (5 Mahajanga) Province: Mahajamba River at Androka (ex. MHNG 2640.039 , spec. 4); J.- C. Nourissat, 23-X-2001 to 06-XI-2001.

PARATYPES: AMNH 238557 , 2 ex., 165.4–167.6 mm SL ; data as for holotype (ex. MHNG 2640.039 , spec. 2 and 3). AMNH 238565 , 2 ex., 1 ex. C&S, tissue voucher specimens, 161.2–170.5 mm SL ; northwestern Madagascar: Mahajanga Province: Mahajamba River, behind Ankarafant- sika National Park ; JSS 39-2003; J.-C. Nourissat, XI-2003. AMNH 238562 , 2 ex., 178.3–181.3 mm SL ; northwestern Madagascar: Majunga Province: Mahajamba River ; JSS 36-2003; J.-C. Nourissat. AMNH 238566 , 1 ex., 184.3 mm SL ; northwestern Madagascar: Majunga Province: Mahajamba River ; JSS 41-2003; J.-C. Nourissat. MHNG 2640.039 , 10 ex., 161.4–183.9 mm SL ; northwestern Madagascar: Majunga Province :

TABLE 13

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

Mahajamba River at Androka; J.-C. Nourissat, 23-X-2001 to 06-XI-2001.

DIAGNOSIS: A shallow-bodied, elongate Paretroplus diagnosed from all congeners except P. nourissati and P. tsimoly by the presence of two wide and convergent (below the lateral midline) dark brown to black midlateral bands, representing the second and third or third and fourth bars in series. Paretroplus lamenabe is distinguished from P. nourissati and P. tsimoly by a deeper body (47.0%–54.3% vs. 38.1%–43.5% and 41.1%– 46.8% SL in P. nourissati and P. tsimoly , respectively), by the presence of pelvic fins that extend beyond origin of the anal fin when adducted, and by a larger adult size (regularly exceeding 180 mm SL vs. less than 160 mm SL in P. nourissati and P. tsimoly ).

DESCRIPTION: Morphometric and meristic data presented in table 13. Morphological characteristics and general pigmentation pattern in life and preservative can be observed in figs. 58–59. A shallow-bodied and elongate Paretroplus belonging to Clade F, which also includes P. nourissati and P. tsimoly (fig. 1). Paretroplus lamenabe is recovered as the sister taxon to a clade comprising P. nourissati and P. tsimoly , and is the largest member of this group, growing to nearly 200 mm SL. Head pointed, snout straight, and predorsal profile moderately curved. Dorsal body outline moderately curved, ventral outline mostly straight, except posteriorly. Lips normal in development and not hypertrophied or lobed. Caudal peduncle short, deep, and laterally compressed. No readily discernable sexually dimorphic characters apparent (but see below).

Total vertebral count 30 to 32 (mode 31), with formulae of: 14 + 16, 14 + 17, 15 + 16, and 14 + 18, precaudal and caudal vertebrae, respectively.

Jaws isognathous. Lips not notably fleshy, and covered with fine papillae. Upper and lower lips not extended into median lobes or flaps (versus condition in P. tsimoly ). Single row of spatulate unicuspid teeth in both upper and lower jaws. Teeth laterally expanded, flattened at crown, and implanted procumbently. 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 seven to nine on each side, and total 15–18. Teeth in lower jaw number three to five on each side, and total 7–10. Upper-jaw teeth relatively closely set for members of Paretroplus . Lower-jaw teeth irregularly and widely 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, becoming progressively enlarged medially; robust molariform teeth present posteromedially. LPJ weakly sutured, with few weak (5 shallow) interdigitations on the ventral suture, located posteroventrally. A narrow gap present between left and right fifth ceratobranchial elements posterior to ventral suture. Indentation present on ventral margin of posterior horns of LPJ. All members of Clade E exhibit both reduced number of (weak) interdigitations on ventral suture and distinct indentation on posterior horns of LPJ ( Stiassny et al., 2001: 37–38, fig. 9). Six or 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 ceratobran- chial 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 rows.

Twelve to 14 triangular and elongate gill rakers arrayed along lower limb of first gill arch. Rakers denticulate dorsally and medially. All other lower-limb rakers (i.e., those on gill arches 2–4) triangular, comparatively elongate as in other members of Clade F, and strongly denticulate dorsally. On gill arches 2–4, teeth much shorter than gill-raker bases. Epibranchial rakers on first gill arch slender and elongate, and number 12 or 13. Rakers on ventral half of first epibranchial denticulate dorsomedially; those on dorsal half edentate.

Body covered with large, regularly imbricate, cycloid scales. Posterior field of lateral body scales thin and not ossified. Welldeveloped scale ridges (5 scale sheathing) present along dorsal- and anal-fin bases. Scale ridges free from spiny dorsal and anal fins, but becoming fused to membranes of both soft dorsal and anal. On membranes between fin rays, multiple discrete rows of scales extend well beyond scale ridges and onto both soft dorsal and anal fins. Pelvic axillary scale present and well developed. Interpelvic scale elongate and rather pointed terminally. Lateral-line scales number 36–39 (mode 37). Chest scales noticeably reduced in size and embedded. Belly scales markedly reduced in size compared to chest scales, those along ventral midline smallest and very embedded. Multiple rows of scales, markedly reduced in size, extend on flanks from chest (dorsal to pelvic fins and anus) to about analfin origin. Four to six rows of scales on cheek. Preopercle asquamate both ventrally and along posterior margin, scaled only dorsally, and anteriorly to a varying degree, on dorsal shaft. Opercle and interopercle scaled. Snout, lacrimal, and anterior portion of interorbital region asquamate. Scales on caudal fin reduced in size and extending posteriorly to over 3/4 length of fin on dorsal and ventral lobes, and 1/3 to 1/2 length of fin medially.

Dorsal with XV–XVII spines, 11–16 soft rays. Anal with VII–IX spines, 10–12 soft rays. Origin of dorsal fin located somewhat posterior to vertical through pectoral-fin insertion. Distal margins of soft dorsal and anal fins weakly produced and more or less rounded, and extend well beyond caudal-fin origin. Caudal fin weakly emarginate, upper and lower lobes broad and more or less rounded to weakly pointed. Trailing margins of upper and lower lobes weakly produced. Pectoral fin broad and rounded at distal margin. Pelvic fin extends beyond origin of anal fin when adducted. In few specimens that could reliably be sexed, caudal margins of both soft dorsal and anal fins, and caudal fin, pointed and more produced in males.

MISCELLANEOUS OSTEOLOGY AND ANAT- OMY: Large, well-developed exoccipital foramina 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) lacking along posterior margin of supraoccipital (a single individual was found to exhibit a very shallow concavity on posterior margin of supraoccipital, but no excavation or notch). 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. Nasal bones elongate. Lower pharyngeal jaw (LPJ) with reduced number of interdigitations on ventral suture and indentation on ventral face of posterior horns. Interdigitations along ventral suture of LPJ shallow and weak. Posterior horns of LPJ narrow, particularly along ventrally projecting ridges, and gap present between left and right fifth ceratobranchial elements, posterior to ventral suture.

COLORATION IN LIFE: (Based on photos and color description in de Rham and Nourissat, 2004: 127–128). Pinkish brown, grey, or bluish grey (nonbreeding, sexually quiescent) to bright yellow or orangish (presumably sexually active individual) with two prominent vertical and generally converging (below the lateral midline) brownish, dark gray, or black midlateral bars. A total of six to seven vertical bars present on flank. Apart from two prominent midlateral bars, representing the second and third or third and fourth bars in series, remaining bars generally faint, although those on posterior flank readily visible. Lips and gular region gray to light bluish grey (nonbreeding, sexually quiescent) to dark grayish blue (presumably sexually active individual). Fins dark gray (nonbreeding, sexually quiescent) to orangish yellow (presumably sexually active individual). Margins of soft dorsal and anal fins, and caudal fin, bright orange (presumably sexually active individual). Dark grey to black triangular patch generally visible in pectoral-fin axil.

COLORATION IN PRESERVATIVE: Base body coloration gray, bluish gray, or grayish brown. Six or seven dark gray vertical bars present on flanks. Apart from two prominent midlateral bars, which generally converge below the lateral midline, remaining bars comparatively faint, although those on posterior flank readily visible (figs. 58–59). Body coloration somewhat darker overall dorsally. Dark gray to blackish triangular patch present in pectoral-fin axil. Snout, lacrimal, and interorbital region beige to dark gray. Lips pale yellow to gray. Bluish-grey to dark grayish-blue coloration on lips and gular region not evident in any preserved specimens. Unpaired fins beige, brownish gray, or charcoal gray. Posterior margins of unpaired fins light and somewhat translucent gray. Pectoral fins beige to light gray. Pelvic fins solid beige, or gray to charcoal gray with lighter gray leading edge.

DISTRIBUTION AND HABITATS: Paretroplus lamenabe is known only from the lower reaches of the Mahajamba River, near the town of Androka, in northwestern Madagascar (fig. 48). De Rham and Nourissat (2004: 128) note that the Mahajamba River in this region is generally turbid and the banks muddy, but that all of the individuals were collected from a rocky stretch with an accelerated current. De Rham and Nourissat (2004) discuss a similar, and possibly conspecific, fish that was collected from the Andranomiditra River, a tributary of the Mahajamba River, in Ankarafantsika National Park, but I have not been able to locate any preserved material for comparison. Too little is currently known regarding the distribution and abundance of P. lamenabe to accurately assess its conservation status.

The Mahajamba River is the next major basin to the north of the extensive Betsiboka system (fig. 48). The Mahajamba is captured by the Kamoro River, a tributary of the Betsiboka, east of Tsaramandroso and near Morafeno, such that the upper 153 km of the Mahajamba flows almost entirely into the Betsiboka basin ( Aldegheri, 1972). Capture of the upper Mahajamba by the Kamoro occurs well upstream of the range of P. lamenabe . Moreover, in periods of low water all of the Mahajamba’s water goes to the Kamoro, and from this point to the sea, the only water the Mahajamba receives is from its small tributaries ( Aldegheri, 1972). Thus, the lower reaches of the Mahajamba are effectively isolated from the Kamoro and Betsiboka basins to which P. tsimoly is endemic (see Distribution and Habitats above for P. tsimoly ). The isolation of the lower Mahajamba from both the upper Kamoro and Betsiboka basins may help to explain the existence of closely related yet distinct species in adjacent basins.

LOCAL NAMES: Lamena, Tsimoly, Damba.

ETYMOLOGY: Lamena is a Malagasy word that translates as ‘‘red one’’, and be is a Malagasy word meaning ‘‘big’’, in reference to both the live coloration of this species and its large size relative to members of its sister clade, P. tsimoly and P. nourissati .

RELATIONSHIPS AND DISCUSSION: Based on the simultaneous analysis of morphological features and nucleotide characters, P. lamenabe is recovered as a member of Clade F and as the sister taxon to a clade comprising P. nourissati and P. tsimoly (fig. 1). Clade F is diagnosed by two unambiguously optimized morphological transformations, the first of which is unique and unreversed: the presence of two prominent and converging midlateral bars (several fainter bars are also present on the flanks) (figs. 58–59) and elongate nasal bones. Paretroplus damii is recovered as the sister taxon to Clade F. Clade E, comprising P. damii , P. tsimoly , P. nourissati , and P. lamenabe , is united by five unambiguously optimized morphological features, two of which are unique and unreversed (features discussed and presented above) (fig. 1).

Interestingly, P. nourissati , the first member of the ‘‘lamena’’ group (Clade F) to be discovered, was not collected until November of 1991 (de Rham and Nourissat, 2004). The other two members of this clade, P. tsimoly and P. lamenabe , were not discovered until 1996 and 2000–2001, respectively (de Rham and Nourissat, 2004). These species are not all restricted to remote regions of the island, nor are they necessarily rare within their respective geographic ranges. For example, P. tsimoly was collected from the Kamilotra (5 Ankalimilotrabe) River, a tributary of the Betsiboka River, just upstream from where it is crossed by the main highway between Majunga and the capital, Antananarivo ( Stiassny et al., 2001; de Rham and Nourissat, 2004).

The members of Clade F are very similar morphologically. They differ primarily in body shape, development of the lips, tooth count in the oral jaws, pigmentation pattern, and coloration. Fin shape and development is also useful for distinguishing the members of this clade. In addition to greater body depth, the presence of pelvic fins that extend beyond origin of the anal fin when adducted and a larger adult size, P. lamenabe can generally be distinguished from P. nourissati and P. tsimoly by tooth count in the upper jaw (seven to nine on each side for a total of 15– 18 vs. 13–16 and 12–14 in P. nourissati and P. tsimoly , respectively), and much more extensive squamation extending onto the soft dorsal and anal fins. No other species of Paretroplus has more than 16 total teeth in the upper jaw. Although Stiassny et al. (2001) report a range of 14–20 teeth in the upper jaw for P. nourissati , I have not been able to find a specimen with more than a total of 16 teeth.

Stiassny et al. (2001: 37–38, fig. 9) listed two features of the LPJ (5 fused fifth ceratobranchial bones) that united P. nourissati and P. tsimoly , a reduced number of interdigitations on the ventral suture (poster- oventral margin of LPJ) and an indentation on the ventral face of the posterior horns. Paretroplus lamenabe also shares these two LPJ features. In addition, in all three of the species comprising Clade F, the posterior horns of the LPJ are also characteristically very narrow, particularly along the ventrally projecting ridges. In addition to a reduction in number, the interdigitations along the ventral suture of the LPJ in members of Clade F are shallow and weak. All three species belonging to Clade F are also characterized by the presence of a narrow gap between the left and right fifth ceratobranchial elements, posterior to the ventral suture. This gap is most pronounced in P. nourissati and P. tsimoly , and present only at the posterior margin of the LPJ in P. lamenabe .

Members of Clade E, comprising Clade F and P. damii , also share a number of unique features related to scale morphology and squamation pattern. Uniquely in this clade, the posterior field of the lateral body scales is thin and not ossified, multiple discrete rows of scales extend onto both the soft dorsal and anal fins (figs. 58–59), the chest scales are markedly reduced in size and highly embedded, and multiple rows of scales that are markedly reduced in size extend laterally on the flank from the chest and continue posteriorly dorsal to the pelvic fin and anus, terminating at about anal-fin origin (figs. 58– 59). In members of Clade F, the chest and belly scales are comparatively more reduced in size and more highly embedded than in other members of the genus, including P. damii . In Clade F, belly scales are so greatly reduced in size and highly embedded that the ventral chest and belly appears asquamate.