Update of diagnoses , information on distribution , species , and key for identification of Laetacara species ( Teleostei , Cichlidae , Cichlasomatini )

The diagnoses of Laetacara species is updated herein based on either character states outlined in their original descriptions, redescriptions, or found in examined material (both type and non-type material) and additional observed data. Information on distribution of Laetacara species is herein updated. A key for species identification including all known species of Laetacara is provided. New information about the species of the genus is provided, based on examined material and compilation from literature.

About two decades later, Laetacara fulvipinnis staecK & schindler, 2007 was described based on six specimens collected in the upper Rio Negro, Casiquiare and Orinoco river basins (staecK & schindler, 2007).This species had been well known for some decades before its description from aquarium publications (see pricK, 1978;KOslOwsKi, 1985;schindler, 1991;röMer, 1992röMer, , 1994;;linKe & staecK, 1994;staecK, 2003).Just after the formal description of L. fulvipinnis, OttOni & cOsta (2009) re-described L. dorsigera based on Brazilian material, as well as described L. araguaiae OttOni & cOsta, 2009 from the Verde river drainage, part of the Araguaia river drainage, southern Amazonas river basin, Central Brazil.In the same year, OttOni et al. (2009) re-described L. curviceps based on both type material and specimens recently collected in the lower Amazonas river basin.More recently, L. flamannellus OttOni, Bragança, aMOriM & gaMa, 2012 was described based on several specimens from different coastal river basins of northern Brazil (OttOni et al., 2012).
The aim of this paper is to update the diagnoses of the valid species of Laetacara based on character states outlined in their original descriptions, re-descriptions, (re-) examination of voucher specimens (both type and nontype) and inclusion of additional observational data.Formal and explicit diagnoses were never proposed for both L. flavilabris and L. thayeri.Difference between species and some main characteres were just briefly cited in some sections of their original descriptions and redescriptions, not explicitly comparing them with all the valid congeners.What make a proposition of formal and explicit diagnoses for these species suitable for a better taxonomic understanding of the genus.In addition, the diagnoses of the other congeners have to be updated since a) new species have been described during the last decades, and b) additional data have been collected for all species of the genus.The final scope of this paper is to present an update of the comprehensive dichotomous key of OttOni L. thayeri: 14 specimens, and all specimens collected by the author.
Measurements and counts were made according to OttOni et al. (2011), OttOni et al. (2012) and OttOni & MattOs (2015).The smaller ray that occurs at the extremities of the dorsal, anal and pectoral fins may eventually be connected to the anterior ray.However, it is counted as a separate element.Some measurements and counts provided by OttOni et al. (2011) were erroneously and poorly defined, being herein redefined: Measurements: (1) standard length -from the upper jaw symphysis to posterior end of the caudal peduncle; and (4) preorbital depth -measured in an imaginary vertical line through the anteriormost eye margin.
Counts: (17) scales of caudal-peduncle depth is the number of scales (or series of scales) through a vertical line at approximately the middle of the caudal peduncle; The (18) caudal-fin rays are counted following the formula "number of dorsal unsegmented procurrent rays (Pc) + number of dorsal principal rays (segmented) + number of ventral principal rays (segmented) + number of ventral unsegmented procurrent rays (Pc)"; and The (21) scales of anal-fin origin series is the vertical count of scales from the anal-fin origin to the upper lateral line (not counting the lateral line scale) .
Counts of gill-rakers of the first branchial arch are made according OttOni & MattOs (2015), including rakers from both sides of each bone.The Prepelvic scales series (squ.prv) is a longitudinal count of scales, in a ventral view, from the base of pelvic fins (including the scales between these paired fins) to the last scale in of the ventral profile.The predorsal (squ.predorsal), is a longitudinal count of scales, in a dorsal view, from the first dorsal-fin spine, to the last scale in the dorsal profile (in the case of squamation pattern trisserial, this count begins on the pair of scales which are deeply in contact with the first dorsal-fin spine).

Diagnosis
Laetacara araguaiae differs from all its congeners by having one exclusive character state: longitudinal stripes on the mid-ventral portion of flank, below the lateral band, light brown on preserved specimens and yellowish brown to golden in live specimens (fig. 1) (vs.absent).In addition, L. araguaiae is distinguished from   1 C).However, when specimens were photographed some hours after collection, the redlines became inconspicuous, and absent in others.Therefore I consider here all these populations to belong to a single species.
Two specimens exhibiting abnormal higher vertebrae count (25) among the 67 examined specimens were recorded.In these specimens apparently there are an abnormal vertebrae, which are smaller than the others and have an uncommon shape, suggesting that one vertebra has been divided in two abnormal ones.As they apparently are a malformation and were rarely recorded, I excluded them from the species counts.Distribution.Laetacara curviceps is known from several localities of the lower Amazonas river basin in Brazil (fig.4).
Remarks.The exact type locality of Acara curviceps is unknown.This species was introduced in the German aquarium trade in 1909 as "Acara thayeri".Later it was recognized as a new species and described by ahl (1924), former curator of Herpetology in the ZMB, based on specimens obtained from the tropical aquarium-fish trade, collected in "Amazonenstrom" (Amazonas river) (OttOni et al. 2009).The examination of the type series did not help to conclude the exact type locality.
The major part of the gut content of seven specimens of UFRJ 4361 was composed of adult aquatic insects of Coleoptera and Hemiptera, and immature Chironomidae.In addition, few undetermined eggs, and adults of Amphipoda and Ostracoda were found.

Laetacara dorsigera (heCkel, 1840)
Figs. 5 and   Distribution.Laetacara dorsigera is known from the middle Paraná and Paraguay river basins, Guaporé, Beni and Mamoré river drainages of the upper Madeira river basin in Brazil, Argentina, Paraguay and Bolivia (Kullander, 2003;staecK & schindler, 2007;OttOni & cOsta, 2009;pers. Observ.), and more recently, recorded from the Uruguay river basin (lanés et al., 2010).The major part of the gut content of ten specimens of UFRJ 3708 contained adult aquatic Hemiptera, immature Chironomidae and adult crustaceans of Copepoda.In addition was recorded few undetermined eggs, fish eggs, adults crustaceans of Cladocera and immature aquatic insects of Coleoptera.

Remarks. The type locality of
One specimen possessing an apparently additional abnormal caudal vertebra (13) was detected among six specimens examined for this character.Two of the caudal vertebrae are smaller than the others with uncommon shape, suggesting that one normal vertebra was divided in two abnormal ones.As this apparently is a malformation, I excluded it from counts., 1982;linKe & staecK, 1984;KOslOwsKi,1985;stawiKOwsKi & werner, 1998;staecK, 2003;OttOni et al., 2009); and from L. curviceps by having a yellow stripe at dorsal-fin mid-portion, along the whole fin of live specimens (fig.7) (vs.restricted to the posterior portion of fin in specimens with about 34 mm SL or over; fig.3), absence of a red or purple region on the anal-fin base (fig.7) (vs.presence; Fig. 3), and absence of a red region on the caudal-fin base (fig.7) (vs.presence of red region the on caudal-fin base in specimens with about 34 mm SL or over; fig.3).
Distribution.Laetacara flamannellus is known from coastal river basins of Amapá state, northern Brazil and the border area between Brazil and French Guiana.The southernmost record is the córrego Areal in the municipal area of Manzagão, and the northernmost the Oiapoque river in the municipal area of Oiapoque (fig.4).
Remarks.Laetacara flamannellus originally was considered being conspecific with L. curviceps (stawiKOwsKi, 1991;le Bail et al., 2000;araújO, 2010;le Bail et al., 2012).However, Laetacara flamannellus exhibits a yellow stripe along the whole dorsal-fin mid-portion of adults and juveniles (fig.7), whereas in L. curviceps, its most closely related congener, this yellow stripe of the dorsal-fin mid-portion is incomplete, restricted to the posterior portion of the fin in adults (specimens with about 34 mm SL or more).This is the main character which distuinguishes these species from each other.

Laetacara flavilabris (COpe
Remarks.Laetacara flavilabris is the type species of the genus.In addition to the character states cited in the diagnosis, it is also easily distinguished from all its congeners by possessing 16 -17 dorsal-fin spines, instead of  In the present work we agreed with the previous synonymization of A. freniferus with L. flavilabris pro posed by Kullander (1986).Examining both holotype of L. flavilabris and lectotype of Acara freniferus, it was observed that both possess vertebrae coutns included in the same range.Additionally, it was not possible to observe any clear differences between these species, and both type localities are geographically close.
The material of L. flavilabris herein examined does not present a row of scales between the rays of dorsaland anal fins.However, Kullander (1986) reported the presence of a row of scales between the rays of that fins, containing one or two scales, in some specimens over 69.2 mm SL.The caudal fin dots of L. flavilabris are pale brown, more evident on the posterior portion of the fin, being more conspicuous in well preserved or live specimens.during his voyage which explored the Rio Negro and Uaupés, between 1848 and 1852 (see tOledO-piza, 2002;p.472 -473 and fig. 205).Before its formal description, several aquarium publications also recorded this species (pricK, 1978;KOslOwsKi, 1985;schindler, 1991;röMer, 1992;1994;linKe & staecK, 1994;staecK, 2003).However, just in 2007 the species was formally described, based on six specimens, collected in the upper Negro, Casiquiare and Orinoco river basins (staecK & schindler, 2007).Information on L. fulvipinnis presented in this study is based on 26 wild specimens, as well as, on information and photographs of the type series and original description.Laetacara fulvipinnis also posses a distinct yellow or orange colouration surrounding the flank spot in live specimens (Fig. 9), which is typical of this species.Distribution.Laetacara thayeri is known from several tributaries of the lower Amazon, Madeira, Solimões  Remarks.Laetacara thayeri is the most widely distributed species of Laetacara, occurring in several tributaries of the lower Amazonas, Madeira, Solimões and Negro river drainages within the Amazonas river basin.Nevertheless, its morphology, including colouration in life of different populations is quite constant.The unique difference recorded among populations is that some specimens from the upper Negro river basin have more anal-fin rays (9) [vs.always 7 -8 in other populations, including the data provided by Kullander (1986)].

Laetacara fulvipinnis
In addition to the character states cited in the diagnosis, L. thayeri can be distinguished from its congeners by the presence of a row of scales between the rays of dorsal and anal fins; contaning a maximum of five scales on each row between rays of dorsal fin, and one or two scales on each row between rays of anal fin (Kullander, 1986;fig. 154).The unique exception is L. flavilabris, for which Kullander (1986) has reported the presence of a row of scales between the rays of that fins, containing one or two scales, in some specimens over 69.2 mm SL.However, the material of L. flavilabris herein examined does not present these rows of scales between the rays of dorsal and anal fins.

Diagnosis.
Laetacara flavilabris differs from all its congeners by having two exclusive character states: presence of 26 or 27 total vertebrae (vs.23 -25) and more radial proximal on the dorsal-fin base (25 vs. 20 -24).In addition, L. flavilabris is distinguished from L. curviceps, L. dorsigera and L. flamannellus by absence of brown dorsal-fin base spot in all specimens examined (fig.8) (vs.presence; figs.3, and 5 -7); from L. araguaiae by missing of longitudinal stripes on the mid-ventral portion of flank, below lateral band, in all specimens (fig.8) (vs.presence of longitudinal stripes on the mid-ventral portion of flank, below lateral band, light brown on preserved, yellowish brown to golden in live specimens; fig.1); from L. thayeri by having flank spot approximately rounded, without intense dorsal extension to the dorsalfin base (fig.8) (vs.flank spot with intense dorsal extension to the dorsal-fin base; fig.10), and absence of brown suborbital bar in adult specimens (fig.8) (vs.presence; fig.10); and from L. fulvipinnis by the presence of brown dots on the caudal fin, more conspicuous in live specimens (fig.8) (vs.absence; fig.9).

Fig. 8 .
Fig. 8. Laetacara flavilabris: not preserved from (A) drainage of the Napo river, in the vicinity of the town Cocain in Ecuador; (B) Drainage of the Juruá, near Cruzeiro do Sul in Brazil.Photographed by W. Staeck.

Diagnosis.
Laetacara thayeri differs from all its congeners by two exclusive character states in both preserved and live specimens: flank spot with intense dorsal extension to the dorsal-fin base (fig.10)(vs.flank spot approximately rounded, without intense dorsal extension to the dorsal-fin base), and presence of brown suborbital bar (fig.10) (vs.absence).In addition, L. thayeri is distinguished from L. flavilabris and L. fulvipinnis by possessing 24 total vertebrae (vs. 25 -27); from L. curviceps, L. dorsigera and L. flamannellus by absence of brown dorsal-fin base spot in both preserved and live specimens (fig.10) (vs.presence; figs.3, and 5 -7); and from L. ful vipinnis by presence of brown dots on the caudal fin of preserved specimens and pale brown of live specimens (fig.10C and D) (vs.absence; fig.9); and from L. ara guaiae by the absence of longitudinal stripes on the midventral portion of flank, below lateral band, in all specimens (fig.10) (vs.presence of longitudinal stripes on the mid-ventral portion of flank, below lateral band, light brown on preserved specimens and yellowish brown to golden in live specimens; fig.1).
6; table2 Diagnosis.Laetacara dorsigera differs from all its congeners (except L. curviceps and L. flamannellus) by a brown dorsal-fin base spot, just above trunk bar 5 in preserved and live specimens (figs.5 and 6) (vs.absent).In addition, L. dorsigera is distinguished from L. fla vilabris and L. fulvipinnis by absence of brown suborbital bar in adults of preserved and live specimens (figs.5 and 6) (vs.presence; fig.10); and from L. fulvipinnis by brown dots on the caudal fin of preserved, and bluish brown on that of live specimens (figs.5 and 6) (vs.absence; fig.9).Lae ta cara dorsigera differs from L. curviceps and L. flamannellus by having red pigmentation of the caudal fin along the entire poste- Tenente de Dragões Antônio Pinto Rego e Carvalho.In 1874, Vila-Maria do Paraguai was elevated to the category of city, receiving the name of São Luiz de Cáceres (http://www.mteseusmunicipios.com.br/NG/conteudo.php?sid=127&cid=442 14 October 2013).
A. dorsigera is "Sümpfe in der Nähe des Paraguay-Flusses bei Villa Maria" meaning "marshes near the Paraguay river basin at Villa Maria".Villa Maria, more correctly Vila-Maria do Paraguai, was a village located at the Mato Grosso state, founded in 1778 by the