PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 115(3):546-563. 2002.
Paretroplus dambabe, a new cichlid fish (Teleostei: Cichlidae) from northwestern Madagascar, with a discussion on the status of P. petiti
John S. Sparks
Division of Vertebrate Zoology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024-5192, U.S.A., e-mail: [email protected]
Abstract. —Paretroplus dambabe, n. sp., is described from Lake Kinkony, northwestern Madagascar. The new species is distinguished from congeners in Hfe and preservation by light yellowish-oUve body coloration in combination with a series of 6-7 vertical charcoal bars on the flanks, a blunt head profile, body depth not exceeding 57.1%SL, and uniform dark charcoal-gray or black fins. Paretroplus dambabe is further distinguished from P. petiti, a species to
which it has been mistakenly referred for decades, by overall pigmentation pattern (light yellow-olive vs. dark brown), the presence of bright red pigmen-
tation on the flanks in life, a prominent vertical barring pattern, and a shallower body.
Paretroplus Bleeker, 1868, the most spe- pelvic axillary scale, and well-developed ciose cichlid genus in Madagascar, com- ridges of scales ("scale sheathing" of Cich- prises nine described species, excluding the ocki, 1976) extending over, but not fused new taxon (Table 1). Paretroplus is endem- to, both the dorsal- and anal-fin bases. A ic to Madagascar, and members are distrib- number of additional features diagnostic of uted throughout the northwestern part of the both Etroplinae and Paretroplus, as well as island (8 species) and in eastern drainages a species-level phylogenetic analysis, are
(1 species), where the range of P. polyactis presented by Sparks (2001). All Paretro- extends nearly the length of the island (Fig. plus but a single described taxon, P. po-
1). Monophyly of Paretroplus, and that of lyactis, are restricted to northwestern Mad- a more inclusive clade encompassing Par- agascar; a number of species from that re- etroplus and Etroplus (endemic to southern gion still await description (J. Sparks, pers.
India and Sri Lanka), is well established obs.). based on numerous morphological and nu- Madagascar is generally considered to cleotide characters (Stiassny 1991, Sparks have a markedly depauperate freshwater & Reinthal 1999, Sparks 2001, Stiassny et ichthyofauna (Kiener & Richard- Vindard al. 2001). Sparks (2001) formally recog- 1972). Results of a recent summary study nized the latter clade, Etroplinae, and as- of ichthyofaunal diversity, however, indi- signed the taxon subfamilial rank. Major cate that Madagascar is comparable in modifications of the anterior swimbladder, terms of the number of native and endemic posterior neurocranium, palato-vomerine freshwater species to other landmasses of region, hyoid-mandibular and premaxillary- similar size (Sparks & Stiassny 2002a, maxillary linkages, and oral and pharyngeal Sparks & Stiassny 2002b). Moreover, the dentition characterize Paretroplus. Exter- number of extant freshwater species slightly nally, members of Etroplinae are easily rec- exceeds that expected for a landmass with ognized by an elevated number of anal-fin a surface area of just under 600.000 sq. km. spines (usually 7-10), a well-developed (Riseng 1997, Sparks & Stiassny 2002b). :
VOLUME 115, NUMBER 3 547
Table 1. —List of Malagasy and South Asian cich- morphological variation among the endem- lids according to current generic assignment, and sub- ic Malagasy and South Asian cichlids is familial designation (discussed in text). substantial (Reinthal & Stiassny 1997, Sparks 2001, Sparks & Reinthal 2001, Sparks 2002). Within this monophyletic as- Ptychochrominae semblage two major subfamilial lineages Ptychochromis oligacanthus (Bleeker) have been diagnosed based on both mor- Ptychochromis grandidieri Sauvage Ptychochromis inoniatiis Sparks phological and molecular characters; Ptych- Ptychochromoides betsileanus (Boulenger) ochrominae, comprising Ptychochromis, Ptychochromoides hatha Reinthal & Stiassny Ptychochromoides, and Oxylapia, and Etro- Ptychochromoides vondrozo Sparks & Reinthal plinae, comprising Paretroplus and Etro- Oxylapia polli Kiener & Mauge plus (Sparks 2001). Herein a new species Etroplinae: of Paretroplus from Lake Kinkony, a large Etroplus suratensis (Bloch) (oligotrophic) floodplain lake in northwest- Etroplus maculatus (Bloch) ern Madagascar and part of the Mahavavy Etroplus canarensis Day Paretroplus dami Bleeker du Sud basin, is described. Further, the Paretroplus polyactis Bleeker identity of P. petiti is clarified. For decades Paretroplus petiti Pellegrin members of the new species have been mis- Paretroplus kieneri Amoult takenly attributed to P. petiti, a taxon from Paretroplus maculatusJ^Qwer & Mauge which it is easily distinguished based on a Paretroplus menaramho Allgayer Paretroplus nourissati (Allgayer) number of morphological features. Paretroplus maromandia Sparks & Reinthal
Paretroplus tsimoly Stiassny et al. Materials and Methods Paratilapia: Paratilapia polleni Bleeker Counts and morphometric measurements
Paratilapia bleekeri Sauvage follow Barel et al. (1977), Kullander (1986) for upper-jaw length and pelvic-fin length, and Sparks & Reinthal (1999), unless noted Many new cichlid species endemic to Mad- otherwise. Measurements were recorded to agascar have recently been, or are in the the nearest 0. 1 mm using Sylvac digital cal- process of being, described (Allgayer 1996, ipers. Vertebral counts exclude the last hy- 1998; Reinthal & Stiassny 1997; Sparks & pural-bearing vertebra (i.e., last half cen- Reinthal 1999, 2001; Stiassny et al. 2001, trum) and were obtained from radiographs. Sparks 2002). Most of these recently dis- Members of Paretroplus frequently possess covered species are restricted to relatively abdominal ribs on the first caudal vertebra. remote regions of the island where limno- The first caudal vertebra is here defined as logical analyses indicate limited ecosystem the most anterior vertebra bearing a fully disturbance (Reinthal & Stiassny 1991, Ris- developed hemal spine, regardless of the eng 1997). Where there is a great deal of presence or absence of abdominal ribs. The ecosystem degradation, including a vast hemal spine of the first caudal vertebra ter-
majority of Madagascar's freshwater sys- minates distally either between the first and tems, primarily exotic species survive (Ben- second, or between the second and third stead et al. 1999, Sparks & Stiassny 2002b). anal-fin pterygiophores. All counts and Increased sediment load and water turbidi- measurements of fin elements were ob- ty, resulting from extensive deforestation tained from radiographs or cleared and and subsequent runoff, appear to be major stained preparations. Each of the terminal
factors leading to the rapid decline of en- dorsal- and anal-fin soft rays is counted as
demic species. two rays if this element is split completely
Despite relatively low species diversity. to the fin base (Barel et al. 1977), which is 548 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
P. polyactis
P. maromandia
P. menarambo
P. maculatus and P. kieneri
P. dambabe and P. kieneri
P. dami
P. nourissati and P. kieneri
P. petiti
P. tsimoly
Paretroplus n. sp. (Lac Andrapongy)
Fig. 1. Map of Madagascar illustrating cuiTent geographic ranges for members of Paretroplus. based on localities from which specimens have been collected in recent surveys (except P. petiti. which is only known from the holotype). These are only approximate distributions, as many remote areas of the island remain poorly surveyed. VOLUME 115, NUMBER 3 549
Fig. 2. Paretroplus dambabe, holotype, UMMZ 238724, adult male, 169.4 mm SL, northwestern Madagas- car, Province of Majunga, Lake Kinkony.
generally the case in Paretroplus. There is 169.4 mm SL, Madagascar, Majunga Prov- only one supporting (articulating) ptery- ince, south of Mitsinjo, Mahavavy (du sud) giophore for this terminal split ray in Par- drainage basin. Lake Kinkony (16°05' etroplus. Without the use of radiographs 37.7"S, 45°51'37.4"E), JSS 94-15, 14-16 this condition is difficult to detect, and it July 1994, J. S. Sparks, K. J. Riseng, and appears as though two distinct rays are pre- local Malagasy guides. sent. Gill-raker counts the (if exclude raker Paratypes.—\jyvyVL 199406 (3, 67.0- present) in the angle of the arch. 113.0 mm SL), ex. MNHN 60579, Mada- Comparative anatomical analyses includ- gascar; AMNH 232398 (10, 61.0-170.0 ed specimens preserved in 70% ethanol, mm SL), data as for holotype; UMMZ specimens cleared and double stained for 235024 (29, 40.2-225.0 mm SL, 3 ex. bone and cartilage using a modified proto- C&S), data as for holotype. col based on Taylor & Van Dyke (1985), Non-type material examined. —All from and dry skeletal preparations. A list of com- Madagascar, Majunga Province, Mahavavy parative material is presented at the end of (du sud) drainage basin. Lake Kinkony: this paper. Institutional abbreviations are as MNHN 1960-0579 (3, 125.5-179.9 mm Usted in Leviton et al. (1985). SL), Kiener; MNHN 1962-0239 (2, 23.3- 24.6 SL), Kiener; 1965-0316 Paretroplus dambabe, new species mm MNHN 64.2-129.3 SL), Petit; Figs. 2-3, Table 2 (2, mm MNHN 1996-123 (1, 105.2 mm SL), Nourissat and Paretroplus petiti. Kiener (in — 1963 de Rham. part).—Kiener & Therezien 1963.—Kie- Differential diagnosis. —A deep-bodied ner & Mauge 1966 (in part). Paretroplus distinguished from congeners, Holotype.—UMML 238724, adult male, in life and preservative, by pale yellow to 550 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 3. Paretroplus dambabe, paratype, UMMZ 235024, juvenile, 93.5 mm SL.
Table 2. —Morphometric and meristic data for holotype and some paratypes of Paretroplus dambabe n. sp. Measurements (mm) in percent standard length (SL) or percent head length (HL), unless noted otherwise. (H) indicates count corresponding to holotype.
P. dambabe n. sp.
N Holotype Range Mean SD
Standard length (mm) 20 169.4 40.7-186.8 115.9 Head width (max.) % SL 20 17.4 16.0-17.5 16.7 0.49 Head length % SL 20 30.4 27.8-33.2 30.7 1.51 Body depth % SL 20 57.0 48.3-57.1 52.8 2.92 Caudal peduncle length % SL 20 6.2 5.1-8.6 6.4 0.66 Caudal peduncle width % SL 20 5.9 2.7-5.9 4.5 0.82 Caudal peduncle depth % SL 20 17.7 15.1-18.2 16.7 0.88 Pectoral fin length % SL 20 22.3 20.1-23.4 21.6 1.05 Pelvic fin length % SL 20 23.6 22.1-26.3 23.5 0.96 Last dorsal spine length % SL 19 14.5 13.3-16.9 15.4 0.97 Snout length % HL 20 48.5 31.9^8.9 43.7 4.64 Orbit diameter % HL 20 26.4 26.4-39.3 31.3 3.74 Upper jaw length % HL 20 30.1 25.7-31.3 28.3 1.59 Lower jaw length % HL 20 35.7 29.9-36.3 34.1 1.79 Interorbital width % HL 20 36.5 28.2^1.7 35.5 3.07 Preorbital depth % HL 20 34.4 22.2-37.6 31.4 4.10 Caudal peduncle length/width 20 1.1 1.1-2.0 1.5 0.26
Scales in lateral line 20 36 (10), 37 (7) (H), 38 (3)
Scales: lateral line to dorsal fin 20 6 (3), 6.5 (1), 7 (16) (H) Gill rakers (lower limb) 20 9(13), 10 (7) (H) Vertebrae (pre-caudal + caudal) 30 14 + 18 = 32 (5), 14 + 19 = 33 (1). 15 + 16 = 31 (1). 15 = + 17 = ;32 (17) (H), 15 + :18 33 (6)
Dorsal fin 30 XVI 18 (3),. XVI 19 (1), XVII 17 (6), XVII 18 (15 ) (H),
XVII 19 (4), XVIII 17 (I)
Anal fin 30 VIII 14 (3), VIII 15 (2). IX 13 (9). IX 14 (11) (H),. IX 15 (2).
X 13 (2)., X 14 (1) VOLUME 1 15, NUMBER 3 551 olive body coloration in combination with jaw usually number 6-8 on each side, rarely a series of 6—7 vertical, dark-olive bars on 5. Teeth in lower jaw number 3-5 on each the flanks, a blunt and steep head profile, side. Lower jaw teeth irregularly spaced body depth not exceeding 57.1%SL, and and sized. the possession of dark, uniform black or Upper and lower pharyngeal dentition ro- charcoal-gray fins. Paretroplus dambabe is bust, tooth plates well developed. Dentition further distinguished from P. petiti, a spe- on lower pharyngeal toothplates [= lower cies with which it has been erroneously as- pharyngeal jaws (LPJ) or fifth ceratobran- sociated, by overall pigmentation pattern chials] hooked and bicuspid both laterally and coloration (light yellow-olive vs. dark and anteriorly, becoming progressively en- brown), the presence of a prominent verti- larged medially; robust molariform teeth cal barring pattern on the flanks (vs. no bar- present postero-medially (Fig. 4). LPJ well ring), bright red pigmentation on the flanks sutured, with numerous interdigitating su- in life, and a shallower body [mean 52.8% tures on postero-ventral margin. Robust (ranges from 48.3-57.1) vs. 57.9% SL]. toothplates covering majority of dorsal sur- Description. —Morphometric and meris- face of fourth ceratobranchials. Toothplates tic data presented in Table 2. Comparative not confluent with outer-row gill rakers of data of other closely related Paretroplus these elements (Fig. 4). Dentition on fourth species presented in Table 3. Morphological ceratobranchial toothplates unicuspid or characteristics and general pigmentation weakly hooked and bicuspid laterally, pattern can be observed in Figs. 2-3. A hooked and bicuspid medially (similar to deep-bodied, laterally compressed Paretro- lateral LPJ dentition). Dentition on third up- plus, belonging to 'deep-bodied' clade per pharyngobranchial toothplates molari- identified and discussed by Sparks & Rein- form medially, hooked and bicuspid periph- thai (1999) and Sparks (2001). Compara- erally. Dentition on second pharyngobran- tively a large Paretroplus, frequently ex- chial toothplates hooked and bicuspid (Fig. ceeding 200 mm SL, and reportedly attain- 5). ing 400 mm TL (Kiener 1963), although Nine or 10 triangular, somewhat elon- large specimens now rare. Head blunt and gate, gill rakers arrayed along lower limb steeply sloping in lateral view. Predorsal of first gill arch. Rakers edentate in small profile rounded and convex, especially in individuals (<100 mm SL) and weakly larger individuals. Caudal peduncle short, denticulate dorso-medially in larger speci- deep, and laterally compressed. Females mens. All other lower-limb rakers (i.e., gill smaller than males, but no additional sex- arches 2—4) short and strongly denticulate ually dimorphic characters apparent. dorsally. Epibranchial rakers elongate, Total vertebral count 31-33 (mode 32), numbering 8-10. Robust toothplates pre- with formulae of: 14 + 18, 14 + 19, 15 + sent on dorsal surface of fourth ceratobran- 16, 15 + 17, and 15 + 18, precaudal and chials. caudal vertebrae, respectively. Body covered with large, regularly im- Jaws isognathous. A single row of spat- bricate, cycloid scales. Well-developed ulate, unicuspid teeth in both upper and scale ridges present along dorsal- and anal- lower jaws. Teeth laterally expanded, flat- fin bases (into which dorsal and anal fins tened at crown, and implanted procumbent- may partially retract). Scale ridges free ly. In upper jaw, tooth on either side of pre- from spiny dorsal and anal fins but becom- maxillary symphysis greatly enlarged, other ing weakly attached to both soft dorsal and teeth graded in size laterally. Lower-jaw anal. Pelvic axillary scale present and well teeth at symphysis not enlarged (and fre- developed. Lateral-line scales 36-38. Chest quently reduced to accommodate enlarged scales somewhat smaller than other body upper symphyseal teeth). Teeth in upper scales, but not greatly reduced in size, and ^ ; ' — ^' 1 —'
552 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
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U c/3 Q < O > VOLUME 1 15, NUMBER 3 553 1 mm Fig. 4. Isolated 4th and 5th ceratobranchial elements, dorsal view, in Paretroplus dambabe, UMMZ 235024, paratype, juvenile, 64.0 mm SL. Abbreviation: cb4tp = 4th ceratobranchial toothplates. Scale bar = 1 mm. embedded. Those along ventral midline tion. Caudal fin emarginate, trailing mar- smallest. Four to 6 rows of scales on cheek. gins of upper and lower lobes produced (es- Opercle and interopercle scaled. Snout, lac- pecially in larger individuals). Pectoral fin rimal, and anterior portion of interorbital re- broad and rounded at distal margin. Distal gion naked. Scales on caudal fin reduced in margins of soft dorsal and anal fins pro- size and extending posteriorly % to % length duced and pointed in larger specimens. Pel- of fin on dorsal and ventral lobes, and Va to vic fin extending just to anal-fin origin in Vj, length of fin medially. smaller specimens, well beyond origin in Dorsal with XVI-XVIII spines, 17-19 larger individuals. soft rays. Anal with VIII-X spines, 13-15 Miscellaneous osteology and anatomy. — soft rays. Origin of dorsal fin slightly an- Large, well-developed exoccipital excava- terior of vertical through pectoral-fin inser- tions present. Anterior swimbladder cham- 554 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON uptp4 1 mm Fig. 5. Isolated right upper pharyngeal elements, ventral view, in Paretroplus dambabe, UMMZ 235024, paratype, juvenile. 64.0 mm SL. Abbreviations: eb 1-4 = epibranchials 1-4; pb 2—3 = pharyngobranchials 2—3; uptp4 = 4th upper toothplate. Scale bar = I mm. bers thick-walled, anteriormost chambers tion pale yellow or pink to light olive. Body lodged in exoccipital recesses. Well-devel- usually darker dorsally, but sometimes uni- oped concavity (= supraoccipital notch of form. Six or seven broad dark olive bars Stiassny et al. 2001) present along posterior extending from anterior region of trunk to margin of supraoccipital. Supraoccipital ex- caudal peduncle; bars very obvious in ju- tending anteriorly over neurocranial median veniles, becoming less conspicuous in large frontal pores (NLFo of Barel et al. 1977). adults. Vivid reddish spots or patches pre- Two distinct and well-separated proximal sent on flank; degree and intensity of spot- premaxillary-maxillary ligaments present ting varying considerably. Some individu- (rostral ligament unique to Paretroplus als with each flank scale outlined in red; in within Cichlidae). An extra, fully ossified, others, red pigmentation restricted to region anal- and dorsal-fin pterygiophore, not as- below upper branch of lateral line. Nape sociated with any fin rays, present termi- and region below anterior spinous dorsal nally. golden in juveniles. Black interorbital bar Coloration in life. —Base body colora- present in adults, dark gold in juveniles. VOLUME 1 15, NUMBER 3 555 Dorsal and anal fins uniform black proxi- verted for rice cultivation and grazing of mally. Soft dorsal and anal fins with white livestock. Little original riparian vegetation distal margin. Caudal fin orangish anteri- remains in the basin. orly, becoming black distally, with white Relationships. —The new species is a terminal band. Pelvic fin black with white member of the 'deep-bodied' clade of Par- leading edge. Pectoral fin reddish-orange etroplus recognized by Sparks & Reinthal proximally, white distally. Pigmentation (1999) and Sparks (2001) on the basis of pattern of young fish (<30 mm SL) blotchy both morphological and molecular evi- and mottled [see juvenile P. petiti (now P. dence. This clade also includes P. macula- dambabe) illustrated by Kiener 1963: PL tus, P. maromandia, P. menarambo, P. pe- 16]. titi, and P. polyactis (Sparks 2001). All Coloration in preservative. —Ground members of this clade are morphologically coloration pale yellow-olive or light tan. very similar except for pigmentation pattern Six or seven broad brownish to charcoal and coloration (Figs. 2-3, 6-7). Morpho- bars on flank. Barring more conspicuous in logically, members of this clade are diag- younger individuals (<150 mm SL) (Fig. nosed by the presence of deep, disk-shaped 3), becoming somewhat obscure in larger bodies (body depth 48-60.5 %SL, regard- specimens (Fig. 2). Reddish coloration on less of standard length) in which body anterior flank lost in preservative. Unpaired depth exceeds 57 %SL in adults, and is not fins and pelvic fin near uniform charcoal less than 48 %SL in young. Body depth in gray. Pectoral fin olive proximally and all other Paretroplus ranges from 38-55 mostly hyaline distally. Anterior interorbital %SL, with body depth in young measuring region and snout light gray. less than 45 %SL. Viscera and diet. —Gut contents com- Paretroplus polyactis is in turn the sister prised entirely of macerated gastropods in taxon to a clade comprising P. dambabe, P. all individuals examined. Paretroplus dam- maculatus, P. maromandia, P. menarambo, babe appears to feed exclusively on gastro- and P. petiti. Morphologically, members of pods. this clade are diagnosed by the presence of Distribution and habitats. —Paretroplus conspicuous alternating, light and dark hor- dambabe has been collected only in Lake izontal stripes on the flank (Sparks 2001, ICinkony, but is also reported to occur in Figs. 2-3, 6-7). Based on the combined surrounding 'satellite' lakes and the Maha- analysis of nucleotide characters from two vavy du Sud River (P. V. Loiselle, pers. mitochondrial genes, a 548 bp region of the comm.). I have only collected and exam- large ribosomal subunit (16S) and a 649 bp ined material from Lake Kinkony proper, region of cytochrome c oxidase subunit I and cannot verify these additional localities. (COI), P. maculatus is the hypothesized sis- Lake Kinkony is a large (second largest ter taxon to the new species (Sparks 2001, lake in Madagascar, ca. 14,000 hectares), fig. 1.2). extremely shallow and turbid, oligotrophic Conservation status. —According to local floodplain lake characteristic of northwest- fishermen the new species has suffered a ern Madagascar (Kiener 1963). Similar severe decline in abundance in recent years. lakes in northwestern Madagascar include Lake Kinkony is subjected to severe fishing lakes Andrapongy and Sarodrano. Several pressure, especially from migrant fishermen euryhaline and marine species, including (catch is shipped to the capital, Antanana- anguillids, carangids, Scatophagus, and rivo). The shores of the lake are home to a Chanos, also inhabit the lake. The Kinkony number of villages that subsist on fishing basin is moderately to highly disturbed and alone. Another member of Paretroplus, P. degraded. Seasonally, fishing pressure is kieneri, is also found in the lake. This spe- very high. Much of the basin has been con- cies has become increasingly rare in recent 556 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Paretrophts petiti Pellegrin, holotype, MNHN 1928-282, juvenile, 82.8 mm SL, northwestern Mad- agascar, Province of Majunga, Maintimaso River. Fig. 7. Fiirciropliis meiuinunln). paralypc, MNHN 1996-122. juvenile, 92.9 mm SL. northwestern Mada- gascar, Potamasina, Lake Sarodrano. VOLUME 1 15, NUMBER 3 557 years according to local fishermen. All near the Betsiboka River, 35 km south of specimens of P. kieneri that were collected Majunga (NW of Maintimaso). Further, by our group, which were few, were in very while examining the type specimen at the poor condition and heavily infested with MNHN in Paris, I noticed an old tag in the both external and internal parasites. jar listing the species name and also includ- Local name. —Kotso. ing the word "Ambanja," a town located Etymology. —Dambabe (pronounced several hundred km to the north of Main- dambah bay), a Malagasy word that trans- timaso and Mahajunga. I have been unable lates as large or big damba, in reference to to obtain any additional information regard- the comparatively large size attained by this ing either the Ambila or the Ambanja lo- species relative to other members of the ge- cality references, and believe that the most nus. Damba is the Malagasy name that is prudent course of action is to accept the lo- used to refer to a number of species of Par- cality information presented by Pellegrin etroplus in northwestern Madagascar, and (1929). As with many other fish species de- be translates as big or large in Malagasy. scribed from Madagascar in the latter part Specific epithet used as a noun in apposi- of the 19th century through the first half of tion. the 20th century, collection localities are frequently ambiguous and questionable Discussion and Comparisons (Sparks 2002). Whether this is the case for P. petiti remains to be determined, as the Since at least the early 1960's individuals region of the putative type locality has been of the new species collected from Lac Kin- poorly surveyed to date. kony have mistakenly been referred to P. According to locality data presented in petiti [Kiener 1963 (in part); Kiener & Pellegrin's original description, it appears Therezien 1963; Kiener & Mauge 1966 (in likely that the type specimen of P. petiti part)] (Fig. 6). Based on the results of this was collected from within the Betsiboka ba- investigation, all specimens previously as- sin, one of Madagascar's largest drainage signed to P. petiti that were collected in Lac systems. Regardless of the exact original Kinkony are determined to be members of collection locality, it is clear from the in- the new species, and P. petiti is only known formation that is available that the type from the holotype described by Pellegrin specimen of P. petiti was not collected from (1929). the Mahavavy (du Sud) drainage basin, to There is some confusion regarding the which the new species is endemic (Fig. 1). type locality of P. petiti, Pellegrin 1929, The holotype and only specimen of P. which is listed in the original description as petiti is very similar in overall body shape the Maintimaso River in the province of and pigmentation pattern (in preservation) Mahajunga (northwestern Madagascar). As to P. menarambo, Allgayer 1996, a species far as I have been able to determine, the restricted to floodplain lakes of the Sofia type specimen of P. petiti (MNHN 1928- basin, northwestern Madagascar (Figs. 1, 282) was collected from a tributary (Main- 6-7). Despite being collected in the 1920's, timaso River) of the westward flowing Bet- this specimen is remarkably well preserved, siboka River near the town of Maintimaso especially in terms of coloration and pig- (16°10'60"S, 46°43'00"E), located approxi- mentation pattern. Both P. petiti and P. mately 37 km southeast of Majunga (= Ma- menarambo are dark brown in overall col- hajunga). In the MNHN database, however, oration, with a lateral pattern of numerous, the collection locality for P. petiti is listed thin, alternating light and dark horizontal as "Ambila." No town of this exact spell- stripes (Figs. 6-7). In preservation, no other ing appears to be in the vicinity of Majun- distinctive lateral markings are present in ga, although the town of Ambilo is located either of these species. In life, P. menar- 558 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 20 40 60 80 100 120 140 160 180 200 Standard Length (mm) Fig. 8. Plot of body depth (BD) as a percentage of standard length (SL) verses SL for P. dambabe (D), P. menarambo (M), and holotype of P. petiti (P). Letters following specific name correspond to symbols shown on plot. ambo is characterized by red terminal bands dark brown with no additional lateral mark- in the unpaired fins. Unfortunately, colora- ings. Although the type specimen of P. pe- tion in life for P. petiti is not known, and titi, a juvenile, is much deeper bodied than comparisons to P. menarambo cannot be any individual of P. dambabe of similar made at this time. standard length, and in fact is deeper bodied In his original description of P. petiti, than any individual of P. dambabe exam- Pellegrin (1929) likewise made no mention ined, it is quite similar in body depth to regarding any additional distinctive mark- specimens of P. menarambo of similar stan- ings. Although Pellegrin (1929) commented dard length (Fig. 8). The possibility exists on the horizontal striping pattern mentioned that P. petiti and P. menarambo are con- above, he did not note the presence of bars specific; however, further studies are war- on the flanks of P. petiti. Other members of ranted and depend on the availability of ad- the 'deep-bodied' clade of Paretroplus (ex- ditional specimens from the region of the cept P. polyactis) possess this conspicuous type locality of P. petiti. horizontal striping pattern; however, none It is clear that P. petiti is not conspecific other than P. petiti and P. nieiuiraiiibo are with P. dambabe from Lac Kinkony, al- VOLUME 115, NUMBER 3 559 though members of the latter species have ex. C&S); UMMZ 235046 (2, 1 ex. C&S). been assigned to P. petiti for decades (Kie- All from Madagascar, Tamatave Province, ner 1963, Kiener & Therezien 1963, Kiener Mangoro drainage, village of Marolambo, & Mauge 1966). Paretroplus petiti is cur- Nosivolo River. rently known only from the holotype. As Paratilapia polleni: RMNH 3.934 (2), discussed, the new species is readily distin- syntypes, island of Nosy be, Madagascar; guished from P. petiti by overall pigmen- RMNH 4.483 (1), syntype, Madagascar; tation pattern and coloration (light yellow- RMNH 6.690 (2), syntypes, "Ambassuana, ish-olive vs. dark brown), the presence of Madagascar septentrionalis, in fluvis"; prominent broad vertical bars on the flanks MNHN A.4195 (3), "marais et rizieres de (except very large individuals in which the rimerina, pres d' Antananarivo"; RMNH bars are less conspicuous), and a shallower 6.692 (1), "Madagascar orientalis, in flu- body (mean 52.8% vs. 57.9% SL). Paretro- vis"; UMMZ 235043 (2 ex. C&S), Lac An- plus petiti is further distinguished from the javibe, island of Nosy be, Madagascar; new species by a much deeper supraoccip- UMMZ uncat., Lac de Deux Soures, island ital crest that is markedly pointed (vs. of Nosy be, Madagascar; UMMZ 235044, rounded) at its dorso-caudal margin. Vahifito River, near Vevembe, southeastern Madagascar; UMMZ 235045 (2 ex. C&S), Comparative Material Sahapindra River, near Vevembe, south- eastern Madagascar; UMMZ uncat. (2 ex. Materials are arranged alphabetically by C&S), rivers near Vevembe, southeastern genus and species, with type specimens list- Madagascar; UMMZ uncat., Manombo ed first followed alphabetically by museum Special Reserve, south of Farafangana, acronym (C&S = cleared and stained prep- southeastern Madagascar; UMMZ uncat. aration, S = dry skeletal preparation; values (1), Ampijoroa, Lake Ravelobe, northwest- in parentheses indicate number of speci- ern Madagascar. mens examined, and do not necessarily cor- Paretroplus dami: RMNH 3.939 (1), respond to the total number of specimens syntype, "Nossibe (Lacus Pambilao)"; in the lot): RMNH 4.478 (1), syntype, "Nossibe (La- Etroplus canarensis: yiCZ 4316 (1), pos- cus Pambilao)"; UMMZ 233523 (S), north- sible syntype, Canara, Cannanore on Mal- western Madagascar; UMMZ 235021 (3 ex. abar Coast, Kerala, India; AMNH 217754 C&S), Anjingo River, east of Antsohihy, (1), River Kumaradhara at Kalikai, southern northwestern Madagascar; UMMZ 235022 Canara, India; RMNH 1103 (1), Canara, In- (C&S), Lac Andrapongy, north basin, dia. northeast of Antsohihy, northwestern Mad- Etroplus maculatus: FMNH 58986 (2), agascar; UMMZ 235023 (1 ex. C&S), An- Colombo, Sri Lanka; FMNH 76016 (37, 2 dranomaloto River (tributary of Mananjeba ex. C&S), India; MCZ 4311 (1), Kerala, River), north of Ambanja, northwestern Laccadive Sea, Canara, Cannanore on Mal- Madagascar; UMMZ uncat. (C&S, S), abar Coast, India; USNM 301168 (6, 2 ex. northwestern Madagascar. C&S), near Negombo Point, Sri Lanka. Paretroplus kieneri: MNHN 1960-580, Etroplus suratensis: FMNH 58987 (8), holotype, Lac Kinkony, northwestern Mad- Colombo, Sri Lanka; MCZ 4306 (2), Ker- agascar; MNHN 1960-581, paratype, Lac ala, Laccadive Sea, Canara, Cannanore on Kinkony, northwestern Madagascar; MNHN Malabar Coast, India; USNM 301178 (6, 2 1966-1043 (13), Riviere Ankalamilotra, ex. C&S), near Negombo Point, Sri Lanka. province of Majunga, northwestern Mada- Oxylapia polli: MNHN 1965-317 (1), gascar; UMMZ 235018 (5, 1 ex. C&S), Lac syntype; AMNH 97098 (1); AMNH 97111 Ravelobe, near Ampijoroa Forestry station, (10, 1 ex. C&S); MNHN 1966-1034 (2, 1 northwestern Madagascar; UMMZ 236592 560 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (4, 1 ex. CS), Lac Kinkony, near Majunga, UMMZ 235015 (10), Mananjary market northwestern Madagascar; UMMZ uncat. (2 and port, southeastern Madagascar; UMMZ ex. C&S, S), Amboaboa and Mangarahara 235016 (15, 3 ex. C&S, S), Farafangana Rivers, near Mandritsara, northern Mada- market, southeastern Madagascar; UMMZ gascar. 235017 (C&S), Manombo Special Reserve, Paretroplus maculatus: MNHN 1965- southeastern Madagascar; UMMZ 236593 315 (2), syntypes, Ambato-boeny, province (1), mangrove swamp near Lalona river, of Majunga, northwestern Madagascar; west coast of Masoala Peninsula, north- UMMZ 235019 (2 ex. C&S), Lac Ravelo- eastern Madagascar; UMMZ uncat. (4), be, near Ampijoroa Forestry Station, north- Maroansetra market, northeastern Madagas- western Madagascar; UMMZ 235020 (2 ex. car. C&S), Lac Ravelobe, near Ampijoroa For- Paretroplus tsimoly: AMNH 229558, ho- estry Station, northwestern Madagascar; lotype, AMNH 229559 (1 ex. C&S), para- UMMZ uncat. (2 ex. S), Lac Ravelobe and type, UMMZ 236893 (1), paratype. Speci- vicinity, northwestern Madagascar. mens from Akalimilotrabe River, Betsiboka Paretroplus maromandia: UMMZ Basin, northwestern Madagascar. 234790, holotype; AMNH 227336, para- Paretroplus cf. maromandia: MHNG type. Specimens from Antalaha province, 2537.60 (2), Lac Andrapongy, near Antso- region of Maromandia, Maintsomalaza Riv- hihy, northwestern Madagascar. er, immediately south of Maromandia, Ptychochromis oligacanthus: RMNH northwestern Madagascar. 3.936 (2), syntypes, "Madagascar, in flu- Paretroplus menarambo: MNHN 1996- mine Samberano; (var. nossibeensis) Nos- 121 and 1996-122 (2), paratypes, Lac Sar- sibe; in lacu Pambilao"; AMNH 215522 odrano, Sofia basin, Potomasina, Madagas- (4), Lac Bemapaza, island of Nosy be, car; UMMZ 233522 (6, 1 ex. S), Lac Sar- northwestern Madagascar; AMNH 215523 odrano, approx. 30 km north of Mampi- (15), Lacs Djabala and Ampombilava, is- kony, northwestern Madagascar; UMMZ land of Nosy be, northwestern Madagascar; 235013 (1), Lac Sarodrano, approximately MNHN 1962-322 (1), Sambirano River, 30 km north of Mampikony, northwestern northwestern Madagascar; UMMZ 236591 Madagascar; UMMZ 235014 (2 ex. C&S), (26, 4 ex. C&S), Lake Ampombilava, is- Lac Sarodrano, near village of Sarodrano, land of Nosy be, northwestern Madagascar; approx. 30 km north of Mampikony, north- UMMZ 237498 (22, 2 ex. C&S), Lake Dja- western Madagascar; UMMZ uncat., Lac bala, island of Nosy be, northwestern Mad- Sarodrano, northwestern Madagascar. agascar; UMMZ 237493 (3), Lac de Deux Paretroplus nourissati: MNHN 1997- Soures, island of Nosy be, northwestern 4172, holotype, Amboaboa River, Sofia ba- Madagascar; UMMZ 237494 (1), Lac Am- sin, Madagascar; MNHN 1997-4173 (4), parihibe, island of Nosy be, northwestern paratypes, Amboaboa River, Sofia basin, Madagascar; UMMZ 237496 (6), Lac Bem- Madagascar; UMMZ 235205 (2 ex. C&S, pazava, island of Nosy be, northwestern S), Amboaboa River, near Mandritsara, Madagascar; UMMZ 237497 (8), Lac An- northern Madagascar; UMMZ 235206 (3 javibe, island of Nosy be, northwestern ex. C&S, S), Amboaboa River near conflu- Madagascar; UMMZ 237499 (11, 1 ex. ence with Mangarahara River, near Man- C&S), Mananjeba drainage, Andranomalo- dritsara, northern Madagascar. to River, northeast of Ambanja. northwest- Paretroplus petiti: MNHN 1928-282, ho- ern Madagascar. lotype, Maintimaso River, Province of Ma- Ptychochromis graiididieri: MNHN junga, Ambila, northwestern Madagascar. A.310 and A.4147 (2). syntypes. Madagas- Paretroplus polyactis: UMMZ 199407 car, region of high forests. (3), Madagascar, formerly MNHN 60-222; Ptychochromis cf. grandidieri: MNHN VOLUME 1 15, NUMBER 3 561 A. 7896 (2), originally designated as syn- Ptychochromoides katria: AMNH 217739, types of P. madagascariensis, Lac Itasy, holotype; AMNH 217740 (8), paratypes; central Madagascar to the east of Antana- AMNH 93701 (10, 2 ex. C&S); UMMZ un- narivo; MNHN 1901-0020 (1), Tamatave, cat (5, 1 ex. C&S). All from Tamatave eastern Madagascar; MNHN 1901-0021 Province, Mangoro drainage, Nosivolo Riv- (1), Tamatave, eastern Madagascar; MNHN er, Marolambo, Madagascar. 1901-0486 (3), Ankobo, Manambolo River, Ptychochromoides vondrozo: UMMZ Madagascar; MNHN 1901-0487 (3), An- 235297, holotype, Fianarantsoa Province, kobo, Manambolo River, Madagascar; Region of Vondrozo, near village of Vev- MNHN 1932-0014 (1), Taraouy River, embe, Mananara drainage basin, Ramanara Madagascar; MNHN 1932-0082 (1), Man- River, tributary of Sahampindra River, ompana, eastern Madagascar; MNHN southeastern Madagascar; AMNH 228488 1932-0083 (13), Manompana, eastcoast, (2), data as for holotype; UAZ 2000-1-1 Madagascar; MNHN 1935-0007 (1), Man- (1), data as for holotype; UMMZ 235293 anara, eastern Madagascar; UMMZ 233524 (1), Sahampindra River, southeastern Mad- (17, S), eastcoast, Madagascar; UMMZ agascar; UMMZ 235294 (3, 1 ex. C&S), 237311 (9), Mananjary, southeastern Mad- Sahampindra River, approx. 10 km up- agascar; UMMZ 237312 (3, 3 ex. C&S), stream from Vevembe Camp, southeastern Manombo Special Reserve, southeastern Madagascar; UMMZ 235295 (1), Voatavo- Madagascar; UMMZ uncat. (C&S), south- be River, tributary of Vahafito River; eastern Madagascar; UMMZ 237495 (1), UMMZ 235296 (3, 1 ex. C&S), data as for Karianga, Rianila drainage, Andriambondro holotype. River, southeastern Madagascar. Ptychochromis sp.: MNHN 1962-0201 Acknowledgments (6), Onilahy River, southwestern Madagas- car. Thanks to Karen Riseng and Roger Ran- Ptychochromis sp.: UMMZ 237066 (1), driamampionina for considerable assistance Amboaboa River, Sofia drainage, northeast- in the field. Thanks to Peter Reinthal for his ern Madagascar. generous support, which allowed me to Ptychochromoides betsileanus: BMNH conduct fieldwork in Madagascar. I am 1882.2.25:69, lectotype, Betsileo, Madagas- grateful to Melanie Stiassny (AMNH) for car; BMNH 1882.2.25:70 (1), paralectotype, the loan of specimens, for support to visit Betsileo, Madagascar; BMNH 1909.7.27:53 AMNH, and for generously sharing her (1), ex. Paris Museum, no locality informa- knowledge of cichlid fishes. For the loan of tion; AMNH 217753 (1), Ilanana River, near material I am also thankful to Barrel Siebert Ranohira, Onilahy drainage, Madagascar; (BMNH), and Susan Jewett (USNM). Col- AMNH 217763 (1), Manantanana River, lecting efforts and visits to Madagascar headwaters near laritsena, Ambalavao Re- were facilitated by Benjamin Andriamahaja gion, Mangoky drainage, Madagascar; and the MICET (Institute for the Conser- MNHN 1907-0104 (2), Madagascar; MNHN vation of Tropical Environments, Madagas- 1919-1 1 (1 ex. C&S), Lac Itasy, central Mad- car) staff, to whom I am indebted. For help- agascar; MNHN 1960-255 (1), Madagascar; ful comments on this manuscript I thank M. MNHN 1965-314 (1), Ambalavao, south- Stiassny and P. Reinthal. This work was central Madagascar; UMMZ 199409 (3), funded by grants from the National Science Madagascar; UMMZ 238114 (3), Ilanana Foundation (DEB-9300996), and the Unit- River, south of Isalo National Park, south- ed States Agency for International Devel- central Madagascar; UMMZ 238115 (5 ex. opment (University Development Linkage S), Ilanana River, south of Isalo National Program, US AID Cooperative Agreement, Park, south-central Madagascar. PCE-5063-A-00-3035-00) to P Reinthal, 562 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON and by the Carl L. and Laura C. Hubbs Re- ology: Part I. Standard symbolic codes for in- search Fellowship and support from the stitutional resource collections in herpetology and ichthyology.—Copeia 1985:802-832. Rackham School of Graduate Studies of the Pellegrin, J. 1929. Cichlides de Madagascar recuellis University of Michigan to the author. par M. Georges Petit. Description d'une espece nouvelle.—Bulletin de la Societe Zoologique de Literature Cited France 54:252-255. Reinthal, R N., & M. L. J. Stiassny. 1991. The fresh- Allgayer, R. 1996. Description espece nouvelle d'une water fishes of Madagascar: a study of an en- Paretroplus Blacker (Teleostei: Cich- du genre dangered fauna with recommendations for a lidae) de Madagascar. Revue Fran^aise des — conservation strategy.—Conservation Biology Cichlidophiles 159:6-20. 5:231-243. . 1998. Descriptions de Lamena nourissati sp. , & M. L. J. Stiassny. 1997. Revision of the n. genre et espece nouveaux, endemiques de Madagascan genus Ptychochromoides (Teleos- Madagascar (Teleostei: Etroplinae). Revue — tei: Cichlidae), with description of a new spe- Fran^aise des Cichlidophiles 179:7-17. cies.—Ichthyological Exploration of Freshwa- Barel C. D. N., M. J. R Van Oijen, F Witte, & E. ters 7:353-368. Witte-Maas. 1977. An introduction to the tax- Riseng, K. J. 1997. The distribution of fishes and the onomy and morphology of the haplochromine conservation of aquatic resources in Madagas- Cichlidae from Lake Victoria. —Netherlands car. Unpublished M.S. thesis. University of Journal of Zoology 27:333-389. Michigan, Ann Arbor, 149 pp. Benstead, J. R, M. L. J. Stiassny, R V. Loiselle, K. J. Sparks, J. S. 2001. Phylogeny and biogeography of the Riseng, & N. Raminosoa. 2000. River conser- Malagasy and South Asian cichhd fishes (Te- vation in Madagascar. Pp. 205-231 in P. J. leostei: Perciformes: Cichlidae), including a Boon, B. R. Davies, & G. E. Petts, eds.. Global survey of the freshwater fishes of Madagascar. perspectives on river conservation: science, pol- Unpublished Ph.D. dissertation. University of icy, and practice. John Wiley and Sons, New Michigan, Ann Arbor, 518 pp. York, 548 pp. . 2002. Ptychochromis inornatiis, a new cichlid Bleeker, P. 1868. Description de trois especes inedites (Teleostei: Cichlidae) from northwestern Mad- de Chromidoides de Madagascar.—Verslagen agascar, with a discussion of intrageneric vari- en Medeedelingen der Koninklijke Akademie ation in Ptychochromis. —Copeia 2002:120- van Wetenschappen Amsterdam, Afdeeling Na- 130. turukunde 2:307-314. , & P. N. Reinthal. 1999. Paretroplus maro- Cichocki, F. P. 1976. Cladistic history of cichlid fishes mandia, a new cichlid fish from the northwest and reproductive strategies of the American of Madagascar.—Occasional Papers of the Mu- genera Acarichthys, Biotodoma, and Geopha- seum of Zoology, The University of Michigan gus. Vol. 1, Unpublished Ph.D. dissertation. 727:1-18. University of Michigan, Ann Arbor, 356 pp. , & P. N. Reinthal. 2001. A new species of Kiener, A. 1963. Poissons, peche et pisciculture a Ptychochromoides from southeastern Madagas- Madagascar.—Publication du Centre Technique car (Teleostei: Cichlidae), with comments on Forestier Tropical 24:1-244. monophyly and relationships of the ptychoch- , & M. Mauge. 1966. Contribution a I'etude romine cichlids. Ichthyological Exploration of systematique et ecologique des poissons Cich- — Freshwaters 12:115-132. lidae endemiques de Madagascar—Memoires , L. J. Stiassny. 2002a. Madagascar's du Musee National d'Histoire Naturelle (Serie & M. A), Zoologique, Paris 40:51-99. freshwater fishes: an imperiled treasure. In M. Thieme, R. Abell, M. L. J. Stiassny. D. Olsen, , & G. Richard- Vindard. 1972. Fishes of the continental waters of Madagascar. Pp. 477-499 E. Dinerstein. J. D'Amico, and E. Underwood, in R. Battistini & G. Richard- Vindard, eds., eds.. Freshwater ecoregions of Africa. A con- Biogeography and ecology in Madagascar. Dr. servation assessment. Island Press, Washington, W. Junk. The Hague, 765 pp. D.C. (in press). . Introduction to , & Y. Therezien. 1963. Principaux poissons du & M. L. J. Stiassny. 2002b. lac Kinkony.—Bulletin de Madagascar 204: Madagascar's freshwater fishes. /;; S. M. Good- 395-415. man and J. P. Benstead. eds.. The natural history KuUander, S. O. 1986. Cichlid fishes of the Amazon of Madagascar. The University of Chicago River drainage of Peru. Swedish Mu.seum of Press, Chicago (in press). Natural History, Stockholm, 431 pp. Stiassny, M. L. J. 1991. Phylogenetic intrarelationships Leviton, A. E., R. H. Gibbs Jr., E. Heal, & C. E. Daw- of the family Cichlidae: an overview. Pp. 1-35 son. 1985. Standards in herpetology and ichthy- //( M. H. A. Keenleyside. ed., Cichlid fishes: VOLUME 115, NUMBER 3 563 behaviour, ecology, and evolution. Chapman of northwestern Madagascar.—Ichthyological and Hall, London, 378 pp. Exploration of Freshwaters 12:29-40. , P. Chakrabarty, & P. V. Loiselle. 2001. Rela- Taylor, W. R., & G. C. Van Dyke. 1985. Revised pro- tionships of the Madagascan cichlid genus Par- cedures for staining and clearing small fishes etroplus Bleeker 1868, with description of a and other vertebrates for bone and cartilage new species from the Betsiboka River drainage study. —Cybium 9:107-119.