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Node 191—genus Teleocichla Kullander 1988

Teleocichla Kullander, 1988: 196 View in CoL (type species Teleocichla centrarchus Kullander 1988 View in CoL )

Crenicichla View in CoL non Heckel (1840) —Ploeg (1991: 122): synonym not followed by subsequent authors.

Teleocichla monophyly is always recovered with high support (ML-BS 100%; BI-PP 100%; parsimony- ABS> 72/RBS 100%; Table 6) and characterized by 39 unambiguous molecular transformations and 32 morphological synapomorphies (19 unambiguous and 13 ambiguous; Table 14). Especially useful to diagnose the genus are characters 19, 47, 48, 73, and 75 combined, 91, 105, 113, and 174. For simplicity, Table 14 only presents six of the 13 ambiguous synapomorphies optimized on the ML tree. These characters are not only diagnostic of the genus (char. 75) but represent repeated modifications apparently associated with rheophyly (chars. 13, 128, 159, and 206) and body size reduction (char. 169) in the context of pike cichlids.

Most synapomorphies of Teleocichla agree with the putative synapomorphies proposed or discussed as related to rheophyly by Kullander (1988) in his diagnosis and anatomical description of the genus. We offer a simplified diagnosis that follows the standard for other groups proposed herein.

Diagnosis: Teleocichla comprises small-sized species (max. SL 48–90 mm, with exception of T. preta with max. 121.3 mm SL) that are distinguished from all other pike cichlids, except species of Hemeraia and Crenicichla (La.) igara , by having infraorbitals 4 and 5 co-ossified, forming a median pore (vs. infraorbitals 4 and 5 autogenous). Teleocichla can be further distinguished from all pike cichlids by the combination of the following apomorphic conditions: pelvic fin long and pointed, with third ray longest (almost reaching the genital papilla) and with a skin thickening on the lateral portion; mouth hypognathous; cheek squamation restricted to its posterior portion; rostral pore of the nasal canal displaced posteriorly from the postlabial margin of the snout; and two or more caudal than precaudal vertebrae. Teleocichla also presents several apomorphic conditions of internal morphology that are useful to diagnose the genus ( Table 14).

Species of Teleocichla are additionally distinguished from all species of the sister-group Wallaciia , except W. heckeli , by the absence of serrations on the posterior margin of the supracleithrum. Teleocichla is further distinguished from Hemeraia , Saxatilia , subgenus Lacustria of Crenicichla , and many species of Lugubria by the absence of suborbital markings. Teleocichla also differs from Saxatilia by the absence of a humeral blotch, from Lugubria by having fewer scales in the E1 series (53–79 scales, except T. centisquama with 85–86 vs. 88–123 scales), from subgenus Batrachops of Crenicichla , except C. (B.) jegui , by the absence of a reticulate colour pattern on the flank (vs. reticulate colour pattern on the flank formed by the dark pigmentation on the base of individual scales), and from Crenicichla (Crenicichla) by having all cycloid (vs. ctenoid) scales on cheek, anterodorsal portion of the body, and on the chest.

Distribution: The nine species of Teleocichla are apparently restricted to the rapids of clear-water tributaries of the Amazon basin, with species distributed in the Río Tapajós, Río Xingu, Río Tocantins- Araguaia, Río Tocantins, and Río Jari basins.

Remarks: In this study, a clade including all species of Teleocichla and Wallaciia as sisters to each other is recovered in all analyses combining morphological and molecular data, with good support in the ML and BI trees (BS 99% and PP 100%), and moderate support in the parsimony analysis based on EW/ DiscreteMatrixTE (ABS 10/RBS 53%; Table 6). This grouping is also recovered in the ML analysis of the molecular data only, and by the parsimony analysis Character transformation Apomorphic condition Observations

Char. 8: 2=>3 Five or more vertebrae + halfcentrum Condition related to the elongation of caudal contained witin the caudal peduncle. peduncle, with several convergent transformations occuring inside the subtribe

Crenicichlina . Char. 19: 0=>2 External area corresponding to the adductor This condition is generally related to mandibulae muscles covered by scales only rheophyly and convergent transform-

on a small dorsoposterior portion. ations are abundant inside the subtribe

Crenicichlina , mainly in the clade

Teleocichla + Wallaciia and in the genus

Crenicichla . Char. 47: 2=>1 Pelvic pointed medially, with third ray Exclusive synapomorphy without reversal.

longest. Char. 48: 0=>1 Presence of skin thickening on the lateral Condition generally related to rheophyly,

portion of the pelvic fin. with convergent transformations occurring as synapomorphies of Crenicichla

(Batrachops) and less inclusive groups of Crenicichla (Lacustria) , as well as autapomorphy of Lugubria phaiospilus . Char. 73: 0=>1 Dark vertical vertical bars expressed as a Convergences occur as synapomorphy of series of blotches along the midlateral area a less inclusive group in Crenicichla in adults. ( Batrachops ) and as synapomorphies of two small groups and autapomorphies of some species of Crenicichla (Lacustria) . Char. 91: 1=>2 Lower jaw hypognathous—upper jaw The transformation from a distinctly prog-

extending more anteriorly than the nathous lower jaw to a hypognathous lower jaw. lower jaw (1=>2) occurs only as synapomorphy of Teleocichla . However, transformations from isognathous or slightly prognathous to a hypognathous lower jaw

(0=>2) occur three times in Crenicichla

(Lacustria). Char. 105: 0=>1 Infraorbitals 4 and 5 fused, forming a median pore. Convergent transformations occurring as synapomorphy of Hemeraia and as autapomorphy of C. (La.) igara . Char. 113: 0=>1 Two to six more caudal vertebrae than Inside the subtribe Crenicichlina , convergent precaudal vertebrae (vs. more precaudal transformations occur as autapomorphies vertebrae, equal numbers or only one more of W. heckeli , C. (La.) hadrostigma and C.

caudal vertebra). (La.) jupiaensis . Char. 131: 0=>2 Heterogeneous fixation of teeth onto This character is already mentioned in other oral jaws: teeth of the outer row firmly groups, since it is also optimized as syn-

fixed and teeth of inner rows slightly apomorphies of Crenicichla (Batrachops)

mobile. and of C. ( Lac .) scottii complex. Other convergences occur inside the subgenus

Crenicichla (Lacustria) . Char. 138: 1=>0 Dorsal process of the anguloarticular Inside the subtribe Crenicichlina , other three taller than the posterior border of the convergent transformations occur in the alveolar arm or approximately equal in genus Crenicichla .

height. Char. 142: 0=>1 Maxillary process of the palatine ap- Reversals occurs in Teleocichla centisquama ,

proximately cylindrical. T. proselytus and T. gephyrogramma , and convergences occur repeatedly inside

Crenicichla (Lacustria) . of the ContinuousMatrix (i.e. morphology only) under the equal weighting scheme. This grouping agrees with the phylogenetic inferences made by Ploeg (1991; Fig. 3I), based exclusively on morphological data, and was also recovered in some of the topologies presented by Burress et al. 2017 using UCEs ( Fig. 3E). Other analyses in Burress et al. (2017; Fig. 3D), and other previous studies ( Burress et al. 2018; Ilves et al. 2018; Fig. 3C, F) based on phylogenomic datasets, disagree with our hypotheses and find Teleocichla View in CoL as sister-group of the remaing groups of clade A.

On the ML tree, 14 morphological transformations are optimized as synapomorphies of this group, seven of them unambiguous. Many of these synapomorphies are modifications related to reduction in body size, including char. 10: 1=>0: maximum body size <100 mm SL, with a reversal only in T. preta that reaches 125 mm SL. The others are the reduction of the squamation on the anterodorsal portion of the body, or predorsal area (char. 28: 0=>1); a reduction of the supraoccipital medial crest and of the paired lateral crests of the neurocranium (char. 160: 2=>3 A; char. 162: 0=>2 A); opening of the NFL3 along the canal connecting NLF2 and NLF4, lacking an accessory canal (char. 172: 0=>1); and pharyngobranchial 1 diminute, globular (char. 180: 0=>3).