Description of Laetacara fulvipinnis sp . n . ( Teleostei : Perciformes : Cichlidae ) from the upper drainages of the rio Orinoco and rio Negro in Venezuela

Laetacara fulvipinnis sp. n. wird aus dem oberen Einzugsgebieten des Orinoko und des rio Negro in Venezuela beschrieben. Die neue Art unterscheidet sich von allen anderen bisher beschriebenen Laetacara-Arten durch die Kombination folgender diagnostischer Merkmale: Rückenfl osse meist XV.10, 23–25 Schuppen in der Längsreihe, Schwanzfl osse ohne deutliches Muster heller Pünktchen und maximale SL 74 Millimeter.

The cichlid species described below from the upper drainages of the río Orinoco and rio Negro has probably been known for almost 30 years in the aquarium hobby (PRICK, 1978).In the aquarium literature it was fi rst provisionally referred to as Laetacara spec."Orangefl ossen" (KOSLOWSKI, 1985;STAECK, 2003) or "Orange-fi nned Laetacara" (LINKE & STAECK, 1994).The purpose of the present paper is to give a formal description of this species.

Material and Methods
The three specimens from the type locality were fi xed in 75 % ethanol, the other paratypes were fi xed in formalin and later transferred into 75 % ethanol.The holotype and 4 paratypes are deposited in the fi sh collection of the Museum für Tierkunde Dresden (MTD F), one paratype is in the collection of ZMB.Comparative material is in MTD F, ZMB or in the personal collections of the authors (CIS).
The techniques for taking measurements and meristic data follow those described in KULLANDER (1986KULLANDER ( , 1990) ) and KULLANDER & NIJSSEN (1989).Measurements were made with an electronic digital caliper reading to the nearest 0.1 mm.Numbers in brackets after counts indicate the number of specimens exam-ined with that condition.Nomenclature for colour pattern follows KULLANDER & SILFVERGRIP (1991) and for bones follows ROJO (1991).
The comparative material was used for the statistical evaluations.Principle component analyses (PCA) were used to investigate patterns of morphological variation between the species of Laetacara.PCA scores were calculated on the basis of the standardized metric or transformed meristic data, respectively, using the programme PAST (HAMMER et al., 2004).PCA for the metric data is based on the covariance matrix and for the meristic data on the correlation matrix.As recommended by VAN VELZEN et al., (1992) all measurements are standardized by expressing them as proportions of standard length in order to correct the differences in size before the data analysis.The formula log(x') = log(SL) -log(x) is used for this purpose (BLACKITH & REYMANT, 1971).Meristic data were transformed by x' = x 0.5 as recommended by KULLANDER (1996).
Abbreviations: E1 = row of scales in the horizontal series directly above the longitudinal row including the lower lateral line; CIS = personal collection of the authors, MTD F = Staatliche Naturhistorische Sammlungen Dresden, Museum für Tierkunde, Fischsammlung; SL = standard length; TL = total length; ZMB = Museum für Naturkunde, Berlin.
Laetacara fulvipinnis sp.n. , Tables 1-2) Holotype.MTD F 30607,male,74.6  Diagnosis.Laetacara fulvipinnis differs from all the other described species in the genus in the combination of the following characters: (1) dorsal fi n usually with 15 spines and 10 rays, (2) 23-25 scales in E1 row, (3) dorsal fi n base only moderately scaly, (4) caudal fi n without a conspicuous pattern of cross series of tiny light dots and ( 5) size (maximum SL 74 mm).
Etymology.The species epithet fulvipinnis is a compound derived from the Latin fulvus (= dark yellow, Vertebrate Zoology ■ 57 (1) 2007 orange) and pinna (= fi n).It refers to the colouration of the caudal and anal fi n, a distinguishing feature of this species.
Description.Based on the holotype, with notes on the paratypes.See Figs. 1 & 3-5 for general shape and colour patterns.Body proportions are summarized in Table 1.
Body moderately deep and laterally compressed.Head short, with bluntly rounded snout.Predorsal contour strongly curved in males; in females dorsal and ventral contours of head more or less evenly arched, with slight concavity in front of orbit.Dorsal-fi n base almost straight.Prepelvic contour less curved than dorsal profi le; abdominal contour straight or slightly concave; anal-fi n base contour slightly convex.Caudal peduncle short, with straight dorsal and ventral edge.In frontal aspect outline of body elliptic with rounded nape and chest.
Orbit in dorsal and chiefl y in anterior half of head.Snout little produced.Mouth terminal, jaws equal anteriorly; maxilla reaching to vertical from anterior  margin of orbit; lower jaw articula tion not reaching to vertical from anterior margin of orbit.Lip folds interrupted and comparatively thick in larger males.Scales on body and nape ctenoid.Prepelvic scales ctenoid with the exception of a few anterior ones.Triserial predorsal pattern with 4 median anterior scales.Cheek scales in 2 series.Anal fi n, pelvic and pectoral fi ns naked.In larger males (SL > 45 mm) a few marginal smaller body scales extend over soft dorsal-fi n base.One larger male with a single minute ctenoid interradial scale.Caudal-fi n base densely scaled; between one third and about half of the fi n covered by scales; posterior margin of the scaled area about truncate.Scales in E1 row 23(4), 24(1), 25(1).Scales on upper lateral line 15(1), 16(3), 17(2), on lower lateral line 8(2), 9(4), including 2 on caudal fi n base.No accessory lateral line on caudal fi n.
Anal-fi n origin opposite last or next to the last dorsal-fi n spine.Soft portion of anal fi n pointed, reaching slightly beyond middle of caudal fi n in adult females and beyond posterior border of caudal fi n in males.Soft dorsal fi n pointed, reaching beyond posterior end of caudal fi n in males.Caudal fi n subtruncate or with rounded posterior margin; caudal fi n length about one third of SL.Pelvic fi ns pointed, fi rst ray extending to the anus.Pectoral fi n rounded.Dorsal fi n XIV.10(1), XV.9(1), XV.10(4).Anal fi n III.8(6).
On fi rst gill arch 0 or 1 minute gill raker on epibranchial, 1 in the angle and 5-6 externally on ceratobranchial.Dorsal margin of anterohyal (Fig. 6) more or less straight (without deep notch), only with a shallow grove where the hyoid artery runs.

Colouration in life.
Based on observations immediately after capture and on specimens kept in aquari-um.Live colouration very variable, depending on age and mood.Adult specimens with beige to turquoise ground colour.Nape and dorsal region dark grey to brown.A dark brown blotch on nape in front of the dorsal fi n.On snout and interorbital region alternating dark and light stripes: broad dark stripe from eye to eye across forehead and narrow dark stripe from eye to eye across upper lip.Both separated by contrasting broad light stripe from orbit to orbit across snout tip.Two additional alternating oblique dark and light stripes on cheek.Iris dusky golden or red.
Scales on the anterior body sides usually with thin black posterior edges.Posterior fl anks with four vertical dark bars growing gradually narrower and separated by narrow light interspaces: one on caudal peduncle, one behind and two above anal fi n.In front of anus an additional bar with midlateral spot, extending ventrally from lateral line scales 6-10 to scales of E1 row.Dark brown horizontal mid-lateral band about two scales wide, extending from distal margin of orbit over dorsal edge of operculum to midlateral spot.Vertical bar carrying midlateral spot margined anteriorly and posteriorly by contrasting narrow vertical orange zones approx.one scale wide.
Dorsal fi n grey, with narrow light margin, maroon submarginal band and with several short cross series of minute light dots on last membranes of soft portion.Pectoral fi ns hyaline.Pelvic fi ns greyish, with thin dark anterior edges of fi rst rays.Anal fi n dark orange, with black margin and a few tiny light dots on last three membranes.Caudal fi n translucent, with dark yellow to orange lower portion.
During courtship both sexes of Laetacara fulvipinnis develops a very dark, almost black colouration on cheek, preoperculum and operculum.After spawning and during parental care it is replaced by a conspicuous ruby-to purple-red colouration, which spreads over cheek, gill cover and chest between upper lip and the pectorals.

Colouration in alcohol.
Based on male holotype and paratypes.Dark markings well preserved, but general appearance of preserved specimens paler.Turquoise areas become white, red pigmentation becomes pale reddish brown.Cheek and opercle dark grey.Snout stripe reddish brown.Subadult specimens with dark dot both near the upper and lower edge of each scale in the midlateral portion of their body.
Posterior bars (bars 2-5) usually more distinct than anterior ones (bars 6-8).Bar 1 reduced to an indistinct narrow greyish midbasal caudal spot (only visible in larger specimens).Bar 2 distinct, ventral part connected with bar 3. Bar 3 between posterior rays of dorsal and anal fi ns, partly covering root of caudal peduncle.Bar 4 between soft dorsal and anal fi n.Bar 5 very broad, between posterior part of spinous dorsal fi n and spinous anal fi n; vertically split into two parts; split indistinct in larger specimens.Bar 6 with midlateral spot, ventrally fused with bar 7; ventral third indistinct.Bar 7 ventrally indistinct, dorsally fused with bar 8 and horizontally interrupted by light interspace above lateral stripe.Bar 8 reduced to a blotch on nape and connected with bar 7. Lateral stripe from posterior orbit to bar 6 or 5 well defi ned; interrupted by a light zone in front and behind midlateral spot.Posterior part of lateral stripe indistinct, running to dorsal half of caudal peduncle.Dorsal fi n plain, posterior soft part with cross series of light dots.Anal fi n plain with dark margin and cross series of light dots on posterior soft part.Caudal fi n plain, in large specimens with a few indistinct light dots in posterior part.Pectoral fi n colourless.Pelvic fi ns hyaline, with dark margin.

Geographical distribution.
Laetacara fulvipinnis is known from several localities in the drainage of the upper and middle río Orinoco, the rio Casiquiare and the upper and middle rio Negro (SCHINDLER, 1991;RÖMER, 1992;1994).Confi rmed collecting sites are situated between Pozo Azul, approx.10 km north of Puerto Ayacucho, (pers.observ.) in the north and the rio Salgado near Barcelos de rio Negro (RÖMER, 1992) in the south.

Ecological notes.
Field observations indicate that Laetacara fulvipinnis prefers typical blackwater habitats with clear, acid and very soft tea-coloured water.The fi sh were usually collected along the banks of brooks and small rivulets in zones of extremely shallow water, i.e. in a water depth between approx.10 and 50 cm, where they were found either in a layer of dead leaves covering the bottom of the bank side or among submerged terrestrial vegetation.
The following water data were collected in February in the rio Casiquiare at the village El Niñal close to the mouth of rio Pasimoni: pH 4,4; electrical conductivity 10 μS/cm; total and temporary hardness < l °dH; water temperature 26,3 °Celsius.At this site the associated fi sh fauna included Apistogramma uaupesi, Mesonauta insignis, Heros severus, Heros sp. and several characid and silurid species.
Reproductive behaviour.Observations under aquarium conditions showed that Laetacara fulvipinnis is a monogamous substrate spawner and that both sexes share in all the duties of brood care.The male, however, is usually the more active partner.A detailed description of the reproductive behaviour was published by RÖMER (1998).Like most other open brooders these cichlids deposit their eggs on a horizontal surface.The preferred spawning site is a stout leaf of a water plant.At 27 ° Celsius hatching occurs about three days postspawning, and the fry attempt swimming six days thereafter.

Discussion
Although all Laetacara species have the same general appearance and a similar colour pattern, Laetacara fulvipinnis is easily distinguished by a set of divergent characteristic features and the specifi c colouration of its translucent caudal fi n, which in live specimens has a dark yellow or orange lower portion and, in contrast to all the congeners, lacks a conspicuous pattern of minute light dots.Subadult preserved specimens of Laetatacara fulvipinnis can be distinguished from all the congeners by a dark dot both near the upper and lower edge of each scale in the midlateral portion of their body.
In addition Laetacara fulvipinnis differs from L. dorsigera and L. curviceps by its size (SL >70 mm versus SL<50 mm), potentially more scales in E1 row (23-25 versus <24) and the lack of a large dark blotch in the dorsal fi n above bar 6 (present in both sexes in L. dorsigera and at least in females of L. curviceps).Furthermore it can be distinguished from L. dorsigera by the shape of bar 4, which is vertically not split (versus split in L. dorsigera), and from L. curviceps by a higher number of dorsal-fi n rays (9-10 versus < 9).
Laetacara fulvipinnis differs from the two larger species L. fl avilabris (SL up to 110 mm) and L. thayeri (SL up to 65 mm) by the absence of a squamatation of the dorsal-fi n base in smaller specimens (SL <45 mm) or a weak squamatation in larger specimens in contrast to a densely scaly dorsal-fi n base in L. thayeri at all sizes and in larger specimens (SL >50 mm) of L. fl avilabris.
Laetacara fulvipinnis is distinguished from L. fl avilabris by well-defi ned snout stripes (less intense in L. fl avilabris), a dark blotch on the nape (versus absence of such blotch), the lack of an extension of the upper part of the midlateral spot (versus dorso-caudally extended midlateral spot at least in specimens up to 60 mm SL), less dorsal-fi n spines (usually XV versus XVI), less scales in E1 row (modally 23 versus 24), no or only one minute gill raker on fi rst epibranchial (versus 1 or 2 well developed ones) and the more steeply arched predorsal contour.Breeding males and females of Laetacara fulvipinnis develops a characteristic conspicuous red colouration on their cheeks, gill covers and chest.Sexually active pairs of Laetacara dorsigera have a similar colouration, which, however, spreads over the whole abdomen.
KULLANDER (1986) documented a deep notch in the dorsal margin of the anterohyal of Laetacara fl avilabris.CASCIOTTA (1998), however, found only a shallow groove in L. dorsigera, an observation confi rmed by our examination of the anterohyal of a L. dorsigera specimen.In Laetacara fulvipinnis the dorsal margin of the anterohyal is also more or less straight, lacks a deep notch (fi g. 6) and is slightly more elongated than in the remaining species.Thus, the shape of the anterohyal of L. fulvipinnis is a further sign of the specifi c differentiation.
Principal component analyses (PCA) were based on the pooled samples of 34 specimens of all described species: Laetacara fulvipinnis (n=6), L. dorsigera (n=13), L. fl avilabris (n=5), L. thayeri (n=3) and L. curviceps (n=7).In the PCA of 12 standardized measurements (Fig. 7a) the PC1 (35% of contribution) shows a high correlation with SL (r = 0.91).This correlation is probably a refl ection of the diagnostic difference in the size between the smaller and larger species.The PC1 is mainly infl uenced by the preorbital depth and caudal peduncle length (highest scores, see Table 2) and, therefore, indicates shape determinants which possibly get the strongest modifi cation during ontogeny.PC2 accounts for 27 % of the variance (no signifi cant correlation with SL, r = -0.21).The loadings of this axis (Table 2) indicate that PC2 is mainly affected by snout length, orbital diameter and pectoralfi n length.The difference between larger specimens of L. fulvipinnis and the remaining species is most apparent in the PC2 axis and the plots of these specimens (L.fulvipinnis SL > 45 mm) are clearly separated in the PC1-PC2 diagram (Fig. 7a).
In the PCA of the meristic data (Table 2) L. fulvipinnis is well separated from the remaining species (Fig. 7b).The difference is mainly caused by the high-  er number of dorsal-fi n rays and scales in E1 row.The difference between L. fulvipinnis and L. fl avilabris on PC2-axis is mainly based on the number of dorsal-fi n spines.The remaining taxa are also signifi cantly different and clearly separated from the cluster of the specimens of L. fulvipinnis.
Despite the comparatively limited number of samples the differences refl ected in the multidimensional analysis are convincing evidence of the distinctness of L. fulvipinnis.

Fig. 2 .
Fig. 2. Collecting site of Laetacara fulvipinnis sp.n. at the río Casiquiare close to the village El Niñal and the mouth of Río Pasimoni.
Tab. 2. Character loadings on PC1 and PC2 for standardized morphometric and transformed meristic data.