Proceedings ofthe United StatesNational MuseumSMITHSONIAN INSTITUTION ? WASHINGTON, D.C.Volume 120 1967 Number 3555
REVIEW OF SOUTH AMERICAN FRESHWATERANGELFISHES?GENUS PTEROFHYLLUM
By Leonard P. SchultzSenior Scientist, Department of Vertebrate Zoology
The illustration of the new Plataxoides leopoldi Gosse (1963, Inst.Roy. Sci. Nat. Belgique Bull., vol. 39, no. 35, pp. 4-6, pi. 1, fig. 2)so closely resembled the illustration of Plataxoides dumerilii Castelnau(1855, Animaux nouveaux ou rares . . . d' Amerique du Sud, p. 21,pi. 11, fig. 3) that I decided to review the taxonomic status of all thenominal species of Pterophyllum. I shall attempt (1) to clarify thenomenclatural validity of the generic names Pterophyllum and Pla-taxoides, (2) to review the species complexes of Pterophyllum scalar
e
(=P. eimekei) and Plataxoides dumerilii (=P. leopoldi,), and (3) todiscuss the relationship between Pterophyllum altum PeUegrin andP. scalare (Lichtenstein).I have studied all nonaquarium specimens of Pterop>hyllum availablein the following museums: British Museum of Natural History(BMNH); California Academy of Sciences (CAS); Chicago NaturalHistory Museum (CNHM); Museum of Comparative Zoology,Harvard (MCZ); Museum National Histoire Naturelle, Paris(MNHN); Institut Royal des Sciences NatureUes de Belgique,Bruxelles (IRSNB); Stanford University (SU); U.S. National Museum(USNM). To the authorities of the above institutions, I wish toexpress my gratitude for their help and other courtesies. 1221-518?67 2
2 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120In the "Tropical Fish Hobbyist" (1953, vol. 1, no. 5, pp. 5-7, 20) Iattempted to distinguish three species of aquariiim-bred Pterophyllum.Since then I have studied specimens collected from their naturalhabitat, the syntypes of Pterophyllum altum Pellegrin, a syntype ofP. eimekei Ahl, 27 paratypes of Plataxoides leopoldi Gosse, and theholotype of P. dumerilii Castelnau.In my 1953 study I recognized Pterophyllum scalare, P. eimekei,and P. altum as valid species; however, with additional material myconclusions have changed and in this review I now recognize as validP. scalare, P. altum, and P. dumerilii. P. eimekei is a synonym ofP. scalare.In the synonomy for each species I have recorded only the mostpertinent references, omitting those related to aquaria.I have attempted to record each locahty from which Pterophyllumhas been reported by latitude and longitude as found in "The TimesAtlas of the World," 1957, vol. 5 ("The Americas"); "U.S. Board ofGeographic Names Gazetteer," no. 56 ("Venezuela") and no. 71("Brazil"); Eigenmann's "The American Characidae," 1917, pt. 1,pi. 1 ; and Eigenmann and AUen's "Fishes of Western South America,"1942, pp. 67-81. Since I was unable to find a few locaHties, specimensfrom such places are not included in the distributional map for thespecies (pi. 1). Some of the other records are so unspecific in locality,e.g., "Rio Orinoco," that these cannot be plotted on the map. I havestudied the routes followed by the collectors; the latitude and longitudeused represents the best estimate for the information available.The specific localities from which specimens of Pterophyllum havebeen collected are as foUows
:
locality
ANGELFISHES?SCHTJLTZ
locality longitude latitudeNazareth, Brazil 1?13' N 67?51' W [probably]Obidos, Brazil 1?52' S 55?30' WPacaya, Rio [see Yarinacocha]Pevas [=Pebas] Cafio, PeruPorto Alegre, BrazilPorto do Moz, BrazilPorto Negro [probably near Manaus], BrazilPurus, Rio, BrazilRockstone, British GuianaRupununi River, British GuianaSantar^m, BrazilSolimoes, Rio, at Manacapuru, BrazilTabatinga, near Leticia, BrazilTefe [=Teffe], Rio, BrazilTonantins, BrazilUeranduba, BrazilUrubu Rio, BrazilMUa Bella [Rio Ramos], BrazilXingu, Rio [Porto Alegre], BrazilYarinacocha [Rio Pacaya], PeruPterophyllum MeckelPterophyllum Heckel, 1840, Ann. Wien. Mus., vol. 2, no. 12, p. 334 [type species:P. scalare B.eckel= Platax scalaris Cuvier and Valenciennes].?Ahl, 1928,Zool. Anz., vol. 76, p. 252 [revision of 3 species].PZatoxoicZes Castelnau, 1855, Animaux nouveaux ou rares . . . de TAmerique duSud . . . , p. 21, pi. 11, fig. 3 [type species: P. dumerilii Castelnau].
3?10'
4 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120Soc. New South Wales, p. 68) were in error in considering thatPterophyllum Heckel is not available as a generic name with the typespecies Pterophyllum scalare Heckel (=Zeus scalaris Lichtenstein)
.
Myers (1940, Stanford Ichthy. Bull., vol. 2, no. 1, p. 36) pointed outthat Neave (1940, Nomenclator zoologicus, vol. 3, p. 1028) was inerror in the following statement: "Pteropyhllum (pro-lla, Kirby 1825),Harris 1833, in Hitchcock, Kept. Geol. Min. Bot. Zool. Massach.,582-Orth." I have checked Harris (1833, 1835, Rep. Geol. Mineral.Bot. Zool. Massachusetts) and can verify Myers' observation thatHarris consistently used Pterophylla, leaving Pterophyllum available.Pterophyllum dumerilii (Castelnau)Plate 2Plataxoides dumerilii Castelnau, 1855, Animaux nouveaux ou rares . . . deI'Am^rique du Sud . . . , p. 21, pi. 11, fig. 3 [Parii=Belem].Plataxoides leopoldi Gosse, 1963, Inst. Roy. Sci. Nat. Belgique Bull., vol. 39,no. 35, p. 4, pi. 1, fig. 2 [mouth of Rio Solimoes, about 90 km above Mana-capuru].Specimens studied (see page 2 for more precise locality) : BMNH1902-11-4-10-12, Maraj6, 3 (27 to 40 mm) : BMNH 1902-2-11-4-9,Tonantins, 1 (41 mm); MCZ (out of 14992), Tefe, 2 (33 and 37mm);CNHM(outof 15254); Santarem, 1 (40 mm); CNHM 53846 Rupu-nuni River, 2 (33 and 54 mm); CNHM (out of 54370), Santarem, 1(37 mm.); USNM 167772, Rupununi River, 1 (40 mm.); USNM198177, Rio Urubu, 1 (45 mm.) ; USNM 198178, Rio Purus, 2 (29 and31 mm.) ; IRSNB 460, Rio Solimoes. 90 km above Manacapuru, 27paratypes of P. leopoldi, (34 to 61 mm) ; MNHN A 254, Pard, Brazil,holotype of P. dumerilii (49 mm); MNHN 221-61-3-7, Cacaual, 1(37 mm); MNHN 221-61-3-6, Caldron, 2 (42 and 51 mm).I examined the holotype of Plataxoides dumerilii Castelnau at theMuseum National Histoire Naturelle, Paris, and found thatCastelnau's illustration (1855, pi. 11) is in error. In his illustration,a black spot is shown behind the promment black vertical bar thatextends from the middle of the dorsal fin across body to the middleof the anal fin, whereas in the holotype this spot now shows as atrace in front of that dark bar, not behind it. Also, the dark verticalbars anteriorly on the illustration are incorrectly portrayed. On theholotype a short bar extends from the spiny dorsal origin ventrallyto the lateral line, another short bar occurs halfway between the eyeand the spiny dorsal origin; there is also a dorsoventral bar throughthe eye and across the cheek. These bars are prominent in well-preserved specimens and their positions differ notably from thoseof Pterophyllum scalare and P. altum. On the latter two species,the black bar through the eye extends dorso-posteriorly to the origin
NO. 3555 ANGELFISHES?SCHULTZ 5
of the spiny dorsal fin, replacing the two middle predorsal verticalbars on P. dumerilii. Thus, on the basis of coloration P. dumeriliican always be distinguished from P. scalare and P. altum. Thenmnber of vertebrae averages slightly fewer in P. dumerilii than inthe other two species (table 1) and there are also slightly fewer softdorsal and soft anal rays and fewer oblique scale rows (from rearof head to midbase of caudal fin) (table 2) than in P. scalare andTable 2.
?
Counts recorded for Pterophyllum dumerilii
Localities
6 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120Pterophyllum altiim PellegrinPlate 3Pterophyllum altum Pellegrin, 1903a, Bull. Mus. Hist. Nat. Paris, vol. 9, p. 125[Rio Atabapo, Orinoco]; 1903b, Mem. Soc. Zool. France, vol. 16, p. 252,pi. 4, fig. 4 [Atabapo].?Regan, 1905, Ann. Mag. Nat. Hist., ser. 7, vol. 16,p. 442 [Rio Orinoco].?Eigenmann, 1910, Rep. Princeton Univ. Exped.Patagonia 1896-1899, vol. 3 (Zool.), pt. 4, p. 479 [Orinoco].?Ahl, 1928, Zool.Anz., vol. 76, p. 255 [Orinoco].?Schultz, 1949, Proc. U.S. Nat. Mus., vol. 99,p. 167 [Rio Atabapo]; 1953, Tropical Fish Hobbyist, vol. 1, no. 5, pp. 5-7,20 [key to aquarium-bred angelfish].Specimens studied (see page 2 for more precise locality) : BMNH1904-6-28-2-3, Rio Orinoco, 2 (59 and 60 mm); MNHN 221-61-1-3,Chaffanjon (Rio Atabapo), 5 sjmtypes of P. altum (42 to 62 mm);MNHN 221-61-1-1, Chaffanjon, 4 syntypes of P. altum (63 to 75mm); MNHN 221-61-1-2, Chaffanjon, 5 syntypes of P. altum(59 to 65 mm); USNM 163204, Venezuela, 1 (79 mm); USNM196007, Colombia, 1 (67 mm); CAS [no number], Rio Casiquiare,1 (53 mm).Ptewphyllum altum Pellegrin has a color pattern identical withthat of P. scalare; however, no black spot occurs on the upper midsideas in P. dumerilii. The black bar through the eye curves dorso-posteriorly to the origin of the spiny dorsal and is not broken intothree separate bars as in P. dumerilii.This species is best recognized from P, scalare by the greateraverage number of median fin rays, oblique scale rows, and vertebrae(tables 1-4). Undoubtedly P. altum represents the P. scalare typeof angelfish in the upper Orinoco, and in having a higher averagenumber of dorsal, anal, oblique scale rows and vertebrae than P.scalare, it might be considered to represent only a subspecies ofP. scalare; however, since P. altum has been taken so far only in theupper Orinoco basin, I prefer tentatively to recognize it as a distinctspecies. Pterophyllum scalare (Lichtenstein)Plate 4Zeus scalaris Lichtenstein, 1823, Verz. Doubl. Zool. Univ. Berlin, p. 114 [Brazil].Platax scalaris Cuvier and Valenciennes, 1831, Histoire naturelle des poissons,vol. 7, p. 237 [Brazil, in "Collection de Bloch, au Musee de Berlin"].Pterophyllum scalaris Heckel, 1840, Ann. Wien. Mus., vol. 2, p. 335, pi. 30,figs. 5, 6, 7-7a, 8-8b [Reo Negro].Pterophyllum scalare Glinther, 1862, Catalogue of the fishes in the British Museum,vol. 4, p. 316 [Rio Cupai; Brazil].?Kner, 1862, Sitz. Akad. Wiss. Wien.,vol. 46, p. 295, pi. 1, figs. 1-la.?Cope, 1872, Proc. Acad. Nat. Sci. Phila-delphia, p. 250 [Rio Ambyiacu].?Steindachner, 1875, Sitz. Akad. Wiss.Wien., vol. 71, p. 76 [Amazon at Santarem, Monte Alegre, Villa Bella,Obidos, Coary, Ueranduba, Tonantins, Tabatinga, Rio Jutahy, Xingu, LagoManacapuru, Lago Maximo, Para, Rio Ambyiacvi; ?Barra do Rio Negro].
?
Eigenmann and Eigenmann, 1891, Proc. U.S. Nat. Mus., vol. 14, p. 71
PROC. U.S. NAT. MUS. VOL. 120 SCHULTZ?PLATE 1
.J^V
i^
PROC. U.S. NAT. MUS. VOL. 120 SCHULTZ?PLATE 2
Pterophyllum dumerilii (Castelnau).
PROC U.S. NAT. MUS. VOL. 120 SCHULTZ?PLATE 3
Pterophyllum altum Pellegrin.
PROC. U.S. NAT. MUS. VOL. 120 SCHULTZ?PLATE 4
NO. 3555 ANGELFISHES?SCHULTZ 7[Amazon].?Eigenmann and Bray, 1894, Ann. New York Acad. Sci., vol. 7p. 624.?Pellegrin, 1903, Mem. Soc. Zool. France, vol. 16, p. 251 [high Peru;upper Amazon, Tefe; Pard].?Regan, 1905a, Ann. Mag. Nat. Hist., ser. 7,vol. 16, p. 441 [Rio Cupai; Tabatinga; Manaus; Tonantins; Marajo Islandin Rio Amazon]; 1905b, Proc. Zool. Soc. London, pt. 1, p. 190 [Rio Negro,Brazil].?Ihering, 1907, Rev. INIus. Paulista, vol. 7, p. 335 [Amazon atTabatinga and tributaries].?Eigenmann, 1910, Rep. Princeton Univ.Exped. Patagonia, 1896-1899, vol. 3 (Zool.), pt. 4, p. 479 [Amazon].?Haseman, 1911, Ann. Carnegie Mus., vol. 7, p. 372 [Santarem, Manaus].
?
Eigenmann, 1912, Mem. Carnegie Mus., vol. 5, p. 521 [Rupununi River;Rockstone; Crab Falls].?Ribeiro, 1915, Arch. Mus. Nac. Rio de Janeiro,vol. 17, p. 56 [Amazon and tributaries].?Ahl, 1928, Zool. Anz., vol. 76,p. 254 [Amazon].?Eigenmann and Allen, 1942, Fishes of western SouthAmerica, p. 406 [Iquitos, Yarinacocha].?^Fowler, 1945, Los peces delPeru, Mus. Hist. Nat. Javier Prado, Univ. Nac. Mayor de San Marcos, p.253, fig. 88 [Amazon, Peru].?Schultz, 1949, Proc. U.S. Nat. Mus., vol. 99,p. 167 [Venezuela]; 1953, Tropical Fish Hobbyist, vol. 1, pp. 5-7, 20 [keyto aquarium-bred angelfish].Pterophyllum eimekei Ahl, 1928, Zool. Anz., vol. 76, p. 252, fig. 1 [Rio Negro inthe Amazon].Specimens studied (see page 2 for more precise locality),?Thefollo^^dng have USNM catalog numbers: 163210, probably BritishGuiana, 20 (28 to 63 mm); 198180, Belem, 5 (72 to 78 mm); 198179,Rio Puriis, 4 (46 to 67 mm) ; 179565 and 179566, Rio Urubu, 54 (33to 80 mm) ; 163210, probably lower Amazon, 6 (72 to 81 mm) ; 191591,upper Rio Araguaia, 1 (17.5 mm); 167754, Yarinacocha, 1 (54 mm);167755, Iquitos, 2 (41 and 63 mm); 26664, Tabatinga, 1 (42 mm);174942, British Guiana, 3 (71 to 81 mm); 175933, Pevas, 3 (27 to 31mm). The following have BMNH numbers: 1926-3-3-3, Rio Negro,1 (60 mm); 1925-10-28-452-457, Manacapuru, 6 (44 to 99 mm);1926-10-27-488, Monte Alegre, 1 (50 mm) ; 53-3-19-42, Rio Cupai[probably Rio Cupari], 1 [not measured]; 81-5-13-129, Lago Cashi-boya, 1 (43 mm); 85-1-14-71, Tabatinga, 1 (55 mm). The followinghave MNHN numbers: 221-61-2-1, Rio Negro, 1 syntype of P.eimekei (55 mm): 221-61-3-10, French Guiana, 3 (63 to 71 mm);221-61-3-2, Tefe, 2 (90 and 101 mm); 221-61-3-3, Peru (Amazon),5 (41 to 51 mm); 221-61-3-4, Peru (Amazon), 3 (50 to 82 mm);221-61-3-5, Tefe, 5 (30 to 44 mm); 221-61-3-6, Caldron, 2 (65 and79 mm); 221-61-3-9, Brazil, 3 (49 to 52 mm). The following haveCAS numbers: [no number], Rio Negro, 4 (72 to 84 mm); [no num-ber], Santarem, 11 (34 to 50 mm); lU 17780, Yarinacocha, 1 (47 mm)
;
lU 2096, Tabatinga, 1 (43 mm) ; lU 5106, Lower Amazon, 1 (51 mm)lU 12502, Rupununi River, 1 (34 mm) ; lU 12504, Essequibo River,2 (48 and 50 mm); lU 13368, Manaus, 1 (84 mm); [no number]
"Porto Negro," 4 (46 to 61 mm) ; 5100, Marajo, 1 (57 mm) ; lU 12503,Essequibo River, 2 (43 and 44 mm); lU 15988, Iquitos, 4 (42 to 69mm). The following have MCZ numbers: 14989, Coary, 4 (84 to
8 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 12094 mm); 14993, Manacapuru, 2 (59 and 60 mm); 14984, Porto doMoz, 2 (47 and 51 mm); 14988, Tonantins, 3 (59 to 94 mm); 14997,Tabatinga, 2 (92 and 99 mm) ; 14985, Porto do Moz, 20 (37 to 48 mm)
;
14994, Santarem, 10 (46 to 54 mm); 14998, Obidos, 4 (47 to 71 mm);14991, Jutahy, 5 (43 to 73 mm); 14992, Jutahy, 3 (39 to 41 mm);14987, Porto do Moz, 5 (40 to 44 mm); 14977 to 14980, Tefe, 10 (51to 101 nrni) ; 14977 to 14980, 14982 and 14983, Tefe, 6 (74 to 100 mm)14996, Santarem, 3 (35 to 46 mm) ; 14990, Villa Bella, 6 (36 to 41mm) ; 14999, Para, 1 (54 mm) ; 14995, Santarem, 1 (44 mm) ; 14976,Monte Alegre, 6 (33 to 39 mm) ; 14981, Tefe, 10 (63 to 98 mm). Thefollowing have CNHM nmnbers: 15254, Santarem, 1 (38 mm) ; 54371,Manaus, 4 (81 to 90 mm) ; 54233, Manaus, 1 (49 mm) ; 53847, Esse-quibo River, 2 (41 and 43 mm) ; 54370, Santarem, 3 (40 to 56 mm)53848, Essequibo River, 4 (44 to 47 mm); 15562, Peru, 1 (53 mm).The following have SU numbers: 36799, Pevas, 1 (62 mm); 36660,near Pevas, 1 (28 mm); 2205, Lower Amazon, 1 (59 mm); 36661,near Pevas, 2 (23 and 26 mm) ; 36797, near Pevas, 1 (37 mm) ; 54245,Santarem, 3 (45 to 57 mm); 60504, Pevas, 1 (75 mm); 36798, nearP^vas, 3 (22 to 36 mm) ; 36659, near Pevas, 2 (37 and 46 mm) ; 36796,near Pevas, 3 (47 to 74 mm)
.
P. scalare (Lichtenstein) is the common angelfish of aquarists.Since 1928, when Ahl described P. eimekei from the mouth of theRio Negro in the Amazon basin considerable confusion has occurredas to the vahdity of that species. P. eimekei was thought to averagefewer dorsal and anal rays and fewer obhque rows of scales (Schultz,1953) and as indicated by Ahl (1928) and Ladiges (1949, DeutscheAquar. Terr., year 2, no. 3, pp. 50-52); however, most of theircounts were based on aquarium specimens, undoubtedly inbred strainsin aquaria, which most probably did not represent the species com-plex of natural populations found in the Amazon basin. The coimtsrecorded in tables 3 and 4 are based on specimens from definitenatural locaHties and not on aquarium-reared material.The counts for the syntype of P. eimekei with XIII, 24 dorsal,VI, 26 anal, 10 pectoral fin rays, and 36 scale rows on the side fallnearly in the middle of the frequency distributions for Amazon speci-mens of P. scalare (tables 3 and 4). Thus, I conclude that P. eimekeiis a synonym of P. scalare.The differences between the color patterns of P. scalare and P.dumerilii, discussed under the latter species and shown in plates 1and 3, should make identification easy; however, P. scalare andP. altum cannot be distinguished on the basis of coloration. Thechief differences are the greater number of soft dorsal and soft analfin rays and the number of scales in P. altum (tables 3 and 4). P.altum has 27 to 31 soft dorsal rays whereas P. scalare rarely hasmore than 26.
ANGELFISHES?SCHULTZ 9
00 C^t^ Ci CO t^ ?0 CD^ O 00 Oii-H -^-^05 i-l CO ?) M ^(N
CO 0.-1010 00 (M(MCOC<5
.-I CO i-IIM
O IMOO OS -^ lOGO
t^ fH CC 00 TtH r-H ^M ?0 POrH Oq -( I-H
iH O .-I O -H 03 CO r^ ?o
t^ CD CO CS O
o
"3 (N ,-1CI -H rt (M
1^CO
10 PROCEEDINGS OF THE NATIONAL MUSEUMAlthough proportional measurements were made on all three speciesof Pterophyllum, the great variabihty even at nearly equal sizessuggests that little rehance can be placed on measurements for iden-tification purposes.Table 4.
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Character index: Total number of rarjs in dorsal and anal fins added tonumber of scale rows for Pterophyllum
Species and localities