Crenicichla Heckel

Synonymy

Crenicichla Heckel, 1840. Annln wien. Mus. Natges. 2, p. 416 (type by subsequent designation by Eigenmann & Bray (1894, p. 620), C. macrophthalma Heckel). — Feminine.
Batrachops Heckel, 1840. Annln wien. Mus. Natges. 2, p. 432 (type by subsequent designation by Eigenmann & Bray (1894, p. 620), B. reticulatus Heckel). — Masculine.
Boggiania Perugia, 1897. Ann. Mus. civ. Stor. nat. Genova (2) 18, p. 148 (type by original designation B. ocellata Perugia). — Feminine.

Crenicichla rosemariae.
Photo: S.O. Kullander.

Diagnosis

South American cichlids with attenuate body shape, reflected in a high number of vertebrae (lowest number 32, in the Crenicichla lepidota group (Kullander 1982)), small, numerous scales (33 to about 130), and high dorsal fin-ray count (rarely as few as 16, usually about 20-24 spines; usually about 15 rays). Crenicichla is similar to Teleocichla but lack the specializations for rheophily characteristic of that genus, and may be polyphyletic for want of proposed synapomorphy to distinguish from Teleocichla.

Crenicichla includes more than 80 species, many of them presently undescribed. Morphological variation, especially in snout and jaw shape is considerable, although still not extensively documented.

The following derived characters may justify recognition of Crenicichla until a thorough cladistic analysis has been performed. As in Teleocichla the supraoccipital crest is absent, but in contrast to Teleocichla there is a spinous caudal projection in Crenicichla which may be an autapomorphy of that genus. In Teleocichla there is only a slight strengthening or short bilateral processes to serve as attachment for the supracarinalis tendon. Crenicichla species usually have five separate infraorbitals, each separately innervated; in other cichlids, including Teleocichla the second and third infraobital bones are usually co-ossified terminally, retaining the opposed terminal foramina of the separate bones as a mesial foramen. Crenicichla hemera, however, has co-ossified second and third infraorbitals. The vertebral column in Crenicichla has more abdominal than caudal vertebrae, unlike most cichlids which have equal numbers or more caudal than abdominal vertebrae. Stiassny (1982) hypothesized that a higher ratio of abdominal to caudal vertebrae would be a synapomorphy of Cichla and Crenicichla.

Identification assistance

Among South American cichlids Crenicichla is a priority group for generic revision. Revisions by Pellegrin (1904) and Regan (1905, 1913) and most recently by Ploeg (1991), contain diagnoses and keys to species groups as well as to all species known to each of those authors. Later or simultaneously published papers (Kullander, 1990a, 1990b, 1991; Lucena & Kullander, 1992) suggested greater complexity than that put forward by Ploeg. Lucena and Kullander (1992) show that the 'lacustris' group discussed by Kullander (1981a, b), Casciotta (1987) and Ploeg (1991) is not diagnosable.

Using scale counts, Kullander (1991) distinguished two major groups among Crenicichla species; the large-scaled species with E1 scale counts 70 or less, and the small-scaled with E1 scale counts 80 or more. Crenicichla jegui (E1 scales 76-82) apparently extends the range for small-scaled species, but still there is no overlap bridging the 70-80 gap.

The Crenicichla lugubris species group, recognized for the small-scaled Crenicichla species can be separated into two groups.

The first is characterized by a blunt snout and generally more than 110 scales in the E1 row: C. lugubris, C. adspersa, C. lenticulata, C. strigata, C. cincta, C. johanna (with 90-110 E1 scales), C. rosemariae and C. marmorata. It corresponds to species 16-22 in Regan's (1913) key. As far as is known, these species undergo a more or less drastic ontogenetic change in colour pattern (Kullander, 1986; Stawikowski & Werner, 1989, p. 262ff.) and most species have a spotted head when young (C. adspersa, C. lenticulata, C. marmorata, C. strigata, C. johanna).

The species with pointed snout and 80-110 scales were contained by Ploeg (1991) in the lugubris group but they may be distinguished by a dorsoventrally compressed, pointed snout and mostly fewer than 110 E1 scales (Kullander, 1991): C. acutirostris (E1 scales 103-111), C. jegui (E1 scales 76-82), C. multispinosa (E1 scales 89-97), C. percna (E1 scales 95-104), C. phaiospilus (E1 scales 94-110), C. ternetzi (E1 scales 80-91), and C. vittata (E1 scales 80-90). These do not undergo drastic ontogenetic change of colour pattern and young apparently do not have a dark spotted head.

The Crenicichla saxatilis group includes species with fairly large scales, less than 70, and of fairly small size, less than 30 cm, and usually only 15-20 cm SL. Practically all species have a distinct humeral blotch, usually with a light rim around it, situated across or below upper lateral line scales. A homologous marking is always present, but in some species may be little differentiated from the anterior portion of the lateral band. There is usually a narrow black stripe below they eye. Females have ocellated spots in the dorsal fin, or a contrasting black marginal and white submarginal band. These species are found all over the range of the genus except along the south Brazilian coast between the Rio Jacui and the Rio São Francisco, both in lowlands and at a few hundred meters elevation: Crenicichla albopunctata, C. alta, C. anthurus, C. brasiliensis, C. britskii, C. coppenamensis, C. frenata, C. hummelincki, C. inpa, C. labrina, C. lepidota, C. lucius, C. menezesi, C. nickeriensis, C. proteus, C. pydanielae, C. santosi, C. saxatilis, C. semicincta, C. sipaliwini, C. sveni, C. vaillanti.

The Crenicichla reticulata group is often separated as a distinct genus, Batrachops. It includes a range of species, often with fewer, more firmly implanted teeth than other Crenicichla, and always with a colour pattern consisting of vertical bars from juvenile to adult. Scale counts range generally between 50 and 70. There is a dark spot at the margin of each scale. The caudal ocellus as a rule is well developed. Sizes vary from 10 to 30 cm. Some of these species, viz. C. cametana, C. cyclostoma, C: geayi, C. sedentaria, occur in fast running water, but C. reticulata, C. cyanonotus, C. stocki and C. semifasciata are lowland quiet water fishes.

The Crenicichla wallacii group includes small species, less than 80 mm SL, with fairly large eyes and plain sides, the caudal spot either small and ocellated or appearing as a large midbasal blotch. Sex dimorphism is marked in some, with females having large ocellated spots in the dorsal fin, but sexes may also be isochromatic, at least in preservative. In distinction to other Crenicichla the supracleithrum is serrated in most species. All species are found in Amazonian, Guianan and Orinocoan lowlands, and in the Tocantins, predominantly in black or clear waters. The group was reviewed independently by Ploeg (1991) and Kullander (1990a), from slightly different angles. Crenicichla notophthalmus, C. regani, C. urosema, C. wallacii, probably also C. heckelii, C. compressiceps and C. virgatula.

The Crenicichla scottii group consists of three species, viz. C. scottii, C. gaucho and C. prenda, restricted to the Uruguay and nearby Argentina. Sizes range between 85 and ca 300 mm SL. Lucena & Kullander (1992) characterised this group by wide interorbital space, maxilla extending well beyond a vertical from the anterior margin of the orbit, lachrymal bone wider than deep (as in saxatilis group species), and a wide lateral band, continuous from head to caudal fin.

The Crenicichla missioneira group consists of the Uruguaian endemic species C. missioneira, C. minuano, C. tendybaguassu, C. igara and C. jurubi. They grow to about 20 cm SL. Scale counts range between 52 and 68. Lucena & Kullander (1992) characterised the group by smooth preopercular margin, suborbital stripe reduced to one or a few spots just below eye, caudal spot situated immediately at base of caudal fin instead of slightly removed from it, and posttemporal blotch well developed and frequently ocellated.

The Crenicichla lacustris group is currently under work by Kullander & Lucena. Those species, including C. lacustris, C. dorsocellata and C. punctata may be recognized by extensively spotted sides and fins. They reach about 30 cm SL and have about 60-70 scales in a lateral row.

The type species, C. macrophthalma, cannot be assigned to any particular group. It with extremely large eyes and the only species with ctenoid predorsal scales. Lateral scales number about 60-70. It reaches about 20 cm SL and is restricted to the central Amazon basin.

Etymology

Crenicichla — crena, Latin noun meaning notch; and Cichla, name of another cichlid genus, from the Greek kichle, a labrid fish known to the Ancient Greek. The name refers to the serrated preopercle.
Batrachops — batrachos, Greek noun meaning frog, and ops, Greek noun meaning face or eye. The name alludes to the short broad snout and large mouth of the originally included species.
Boggiania — for the collector of the type-species, Guido Boggiani.

Geographical distribution

Found over most of cis-Andean South America, including the island of Trinidad, W.I., the Tuy and Paria basins of Venezuela, the Orinoco, Amazon, Paraguay-Paraná, most coastal rivers between the Río de La Plata and the Amazon, as well as all of the Guianas. The southernmost locality was reported to be Puerto Madryn, Chubut Province, Argentina (Kullander, 1981a), but as turns out that collection was mislabeled (S. Muller, in litt.), instead the C. scottii reported by Mac Donagh (1937) (as Crenicichla lacustris), and Casciotta (1987) from the Río Negro in northern Patagonia most likely represents the most southern locality of South American cichlids.

Included species

Crenicichla acutirostris Günther, 1862 Crenicichla adspersa Heckel, 1840 Crenicichla funebris Heckel, 1840 Crenicichla albopunctata Pellegrin, 1904 Crenicichla alta Eigenmann, 1912 Crenicichla pterogramma Fowler, 1914 ?Crenicichla cardiostigma Ploeg, 1991 Crenicichla anthurus Cope, 1872 Crenicichla brasiliensis (Bloch, 1792) Sparus Nhoquunda Lacépède, 1802 Crenicichla britskii Kullander, 1982 Crenicichla cametana Steindachner, 1911 Crenicichla astroblepa Ploeg, 1986 Crenicichla celidochilus Casciotta, 1987 Crenicichla cincta Regan, 1905 Crenicichla brasiliensis var. fasciata Pellegrin, 1904 Crenicichla compressiceps Ploeg, 1986 Crenicichla coppenamensis Ploeg, 1987 Crenicichla cyanonotus Cope, 1870 Crenicichla cyclostoma Ploeg, 1986 Crenicichla frenata Gill, 1858 Crenicichla dorsocellata Haseman 1911 Crenicichla biocellata R. von Ihering, 1914 Crenicichla gaucho Lucena & Kullander, 1992 Crenicichla Geayi Pellegrin, 1903 Crenicichla haroldoi Luengo & Britski, 1974 Crenicichla heckeli Ploeg, 1989 Crenicichla hemera Kullander, 1990 Crenicichla guentheri Ploeg, 1991 Crenicichla hummelincki Ploeg, 1991 Crenicichla iguassuensis Haseman, 1911 Crenicichla inpa Ploeg, 1991 Crenicichla igara Lucena & Kullander, 1992 Crenicichla isbrueckeri Ploeg, 1991 Crenicichla jaguarensis Haseman, 1911 Crenicichla jegui Ploeg, 1986 Crenicichla johanna Heckel, 1840 Cychla fasciata Jardine, 1843 Crenicichla obtusirostris Günther, 1862 Crenicichla Johanna; var. carsevennensis Pellegrin, 1905 Crenicichla jupiaensis Britski & Luengo, 1968 Crenicichla jurubi Lucena & Kullander, 1992 Cychla labrina (Agassiz, 1831) Cycla lacustris (Castelnau, 1855) Crenicichla lenticulata Heckel, 1840 Crenicichla ornata Regan, 1905 Crenicichla lepidota Heckel, 1840 Crenicichla edithae Ploeg, 1991 Crenicichla lucius Cope, 1870 Crenicichla lugubris Heckel, 1840 Cychla? rutilans Jardine, 1843 Crenicichla macrophthalma Heckel, 1840 Crenicichla santaremensis Haseman, 1911 Crenicichla marmorata Pellegrin, 1904 Crenicichla menezesi Ploeg, 1991

Crenicichla minuano Lucena & Kullander, 1992 Crenicichla missioneira Lucena & Kullander, 1992 Crenicichla mucuryna von Ihering, 1914 Crenicichla multispinosa Pellegrin, 1903 Crenicichla nickeriensis Ploeg, 1987 Crenicichla niederleinii (Holmberg, 1891) Crenicichla notophthalmus Regan, 1913 Crenicichla pellegrini Ploeg, 1991 Crenicichla percna Kullander, 1991 Crenicichla phaiospilus Kullander, 1991 Crenicichla prenda Lucena & Kullander, 1992 Crenicichla punctata Hensel, 1870 Crenicichla polysticta Hensel, 1870 Crenicichla proteus Cope, 1872 Batrachops nemopterus Fowler, 1940 ? Crenicichla nijsseni Ploeg, 1991 Crenicichla pydanielae Ploeg, 1991 Crenicichla regani Ploeg, 1989 Crenicichla reticulata (Heckel, 1840) Crenicichla elegans Steindachner, 1881 Batrachops punctulatus Regan, 1905 Crenicichla rosemariae Kullander, 1997 Crenicichla santosi Ploeg, 1991 Crenicichla saxatilis (Linnaeus, 1758) Sparus biocellatus Walbaum, 1792 Sparus Pavo Lacépède, 1802 Scarus pavoninus Gray, 1854 Crenicichla scottii (Eigenmann, 1907) Crenicichla multidens Steindachner, 1915 Labrus amarus Larrañaga, 1923 Crenicichla lacustris semifasciata Devincenzi,1939 Crenicichla sedentaria Kullander, 1986 Crenicichla semicincta (Steindachner, 1892) Crenicichla clancularia Ploeg, 1991 Crenicichla semifasciata (Heckel, 1840) Acharnes chacoensis Holmberg, 1891 Boggiania ocellata Perugia, 1897 Crenicichla simoni Haseman, 1911 Crenicichla sipaliwini Ploeg, 1987 Crenicichla strigata Günther, 1862 Crenicichla stocki Ploeg, 1991 Crenicichla sveni Ploeg, 1991 Crenicichla tendybaguassu Lucena & Kullander, 1992 Crenicichla ternetzi Norman, 1926 Crenicichla tigrina Ploeg, Jégu & Ferreira, 1991 Crenicichla urosema Kullander, 1990 Crenicichla vaillanti Pellegrin, 1903 Crenicichla virgatula Ploeg, 1991 Crenicichla vittata Heckel, 1840 Crenicichla wallacii Regan, 1905 Crenicichla nanus Regan, 1913 Nomina inquirendae Cycla conibos Castelnau, 1855 Cycla multifasciata Castelnau, 1855

References

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