taxonID	type	description	language	source
170987F4C5375137A367FF33C4AFF9CF.taxon	description	As redefined herein, Crenicichla comprises 43 valid species divided into three subgenera: Crenicichla (one species), Batrachops (nine species), and Lacustria (33 species); see Table 1. The genus Crenicichla shows great morphological and ecological diversity, which challenges the formulation of a good diagnosis for the group. Thus, we make use of a combination of several characteristics to distinguish the species of Crenicichla from the other pike cichlids, organized by morphological groups. Diagnosis: All species of Crenicichla, except C. macrophthalma, can be distinguished from other pike cichlids by sexually dimorphic characters that include orange or reddish marking on the lateral abdomen of mature females. Apart from C. macrophthalma, which is known to lack such dimorphism, other species for which there is no information on the coloration of mature females probably show this feature (see the distribution of states of character 89). In most species of Crenicichla, female marking on the lateral abdomen is accompanied by one or more blotches on the dorsal fin; in some taxa, these blotches are eventually modified into a horizontal dark bar. While this characteristic is shared with some species of Wallaciia, it is absent in the remaining species of pike cichlids. All species of Crenicichla are distinguished from most species of Saxatilia and Lugubria by the absence of a broad reddish or purplish pigmentation on the ventral part of the abdomen of gravid females. All species of Crenicichla are additionally distinguished from Saxatilia by the absence of a humeral blotch and of sexual dimorphism consisting of scattered light spots on mature males — Crenicichla (Batrachops) jegui has light dots scattered on the flanks and head but their presence is not sexually dimorphic. All species of Crenicichla are distinguished from Lugubria by having fewer than 79 scales in the E 1 row, except Crenicichla (Lacustria) vittata with 79 – 93 scales (vs. 79 – 123 scales in the species of Lugubria). All species of Crenicichla have all post-lachrymal infraorbitals autogenous vs. infraorbitals 4 and 5 fused, forming a median pore in Teleocichla and Hemeraia. Exceptions are the unique specimen of C. (La.) igara examined herein, which shows infraorbitals 4 and 5 partially fused, and C. (La.) jupiaensis, with only four instead of five infraorbitals but without signal of co-ossification (see: Varella et al. 2018). Species of Crenicichla are additionally distinguished from Hemeraia by having most of the flank scales ctenoid [i. e. a combination of patterns 1 and B 2 in all Crenicichla except C. (C.) macrophthalma, which shows a combination of patterns 0 and B 1] vs. most of the flank scales cycloid (i. e. combination of pattern 3 and B 3). Species of Crenicichla are additionally distinguished from Teleocichla by having rounded pelvic fin with the second ray longest (vs. pointed pelvic fin with the third ray longest), by having a stick-like pharyngobranchial 1 (vs. globular pharyngobranchial 1), by showing the configuration of the urohyal similar to other pike cichlids, i. e. lateral wings wider than the depth of the medial crest (vs. lateral wings wide but medial crest rudimentary or absent), and by having the symmetrical medial processes of the basipterygia diverging anteriorly (vs. running very close, not diverging anteriorly). All species of Crenicichla are also distinguished from all species of Wallaciia, except W. heckeli, by the absence of serrations on the posterior margin of the supracleithrum. Additionally, Crenicichla comprises medium-sized species (max. SL 95 – 294 mm) with relatively small eyes (orbital diameter 4.6 – 10.9 % of SL, with minimum between 4.6 – 7.5 %) and clearly showing a negative allometry of eye size (eyes decreasing proportionally in size with ontogeny), with the exception of C. macrophthalma, a medium-sized species (max. SL 200 mm) with large eyes in small to large specimens (orbital diameter 9.4 – 10.6 % of SL), with less marked negative allometry in eye size. Thus, Crenicichla can be distinguished from Wallaciia, which comprises only small-sized species (max. SL 52 – 85 mm) with large eyes (orbital diameter 7.8 – 12.6 % of SL, with minimum between 7.8 – 8.6 %).	en	Varella, Henrique R, Kullander, Sven O, Menezes, Naércio A, Oliveira, Claudio, López-Fernández, Hernán (2023): Revision of the generic classification of pike cichlids using an integrative phylogenetic approach (Cichlidae: tribe Geophagini: subtribe Crenicichlina). Zoological Journal of the Linnean Society 198 (4): 982-1034, DOI: 10.1093/zoolinnean/zlad021, URL: https://academic.oup.com/zoolinnean/article/198/4/982/7204395
170987F4C5375137A367FF33C4AFF9CF.taxon	materials_examined	Type species: Crenicichla macrophthalma Heckel, 1840. Nominal species: Crenicichla macrophthalma and C. santaremensis (synonym of C. macrophthalma). The subgenus is recovered in all analyses performed herein with high support in ML and BI trees (BS 100 % and PP 100 %) and moderate support in the parsimony analysis of EW / DiscreteMatrixTE (ABS 5 / RBS 63 %; Table 6). The subgenus is diagnosed by 50 molecular transformations and 19 morphological synapomorphies (Table 10). The same morphological synapomorphies are found from the optimization on the parsimonious tree from EW / DiscreteMatrixTE, except for characters 98 and 168 (only in the ML optimization). Apart from some characters related to external morphology, such as squamation (characters 23, 24, 25, 26, 27, and 40) and colour patterns (51, 71, and 80), autapomorphies come from highly homoplastic characters inside the subtribe Crenicichlina and are of little value in formulating a diagnosis for the species. Character 98 (state 1: eyes visible in ventral view) was optimized as one of the apomorphic conditions of the subgenus, with convergences also optimized as synapomorphies of Saxatilia and Wallaciia. It was difficult to decisively infer whether the visibility of the eyes (in ventral view) resulted from two variables instead of one: size of the eye and its relative position in the head. In Saxatilia, the eyes are situated more laterally in the head (wide interorbital space). Crenicichla (C.) macrophthalma and Wallaciia are recognized to have larger eyes, which occupy a large portion of the upper lateral portion of the head because of their size and because the interorbital space is narrow. Further evaluation of this character based on ontogenetic information or even a reinterpretation may be important for subsequent analyses. Diagnosis: Crenicichla (Crenicichla) macrophthalma is a medium-sized species of pike cichlid (max. SL 200 mm) that can be distinguished from all other pike cichlids by the following combination of characters: large eyes (orbital diameter 9.4 – 10.6 % of SL); body almost entirely covered by ctenoid instead of cycloid scales (scales ctenoid on cheek, flank squamation following patterns 0 and B 1, and scales ctenoid covering the caudal fin almost entirely); and the absence of dark markings below the eye (suborbital marking) and on caudal blotch. Among pike cichlids, only C. (La.) vittata shows a similar squamation pattern but is readily distinguished from C. (C.) macrophthalma by having more scales in the E 1 series (79 – 93 vs. 65 – 68) and by having a suborbital marking, a conspicuous dark midlateral band on flank, and a caudal blotch. A d d i t i o n a l c o m p a r i s o n s b e t w e e n g r o u p s: T h e monotypic subgenus Crenicichla is distinguished from almost all species of the subgenera Batrachops and Lacustria by the absence of sexual dimorphism expressed by mature females with orange or reddish pigmentation on the lateral abodomen and by dark blotches on the dorsal fin of mature females. The latter character also distinguishes the subgenus Crenicichla from most species of Wallaciia. Crenicichla (Crenicichla) macrophthalma is distinguished from most species of Saxatilia and Lugubria by the absence of a reddish or purplish broad pigmentation on the ventral abdomen (belly) of mature females. The subgenus Crenicichla also differs from Saxatilia by the absence (vs. presence) of a humeral blotch, from Lugubria by having fewer scales in the E 1 series (65 – 68 vs. 88 – 123), from Wallaciia (except W. heckeli) by the absence of serrations on the posterior margin of supracleithrum, and from Teleocichla and Hemeraia by having all post-lachrymal infraorbitals autogenous vs. infraorbitals 4 and 5 co-ossified, forming a median pore. The subgenus also differs from Hemeraia by having regularly serrated instead of smooth preopercle and from Teleocichla by having a pelvic fin with rounded margin and second ray longest instead of pointed with third ray longest. Distribution: Crenicichla macrophthalma is known from tributaries (Río Negro, Río Trombetas, Río Tapajós and Río Xingu) and the main channel of the Amazonas River.	en	Varella, Henrique R, Kullander, Sven O, Menezes, Naércio A, Oliveira, Claudio, López-Fernández, Hernán (2023): Revision of the generic classification of pike cichlids using an integrative phylogenetic approach (Cichlidae: tribe Geophagini: subtribe Crenicichlina). Zoological Journal of the Linnean Society 198 (4): 982-1034, DOI: 10.1093/zoolinnean/zlad021, URL: https://academic.oup.com/zoolinnean/article/198/4/982/7204395
170987F4C5375137A367FF33C4AFF9CF.taxon	description	None of them corresponds to characteristics used in previous papers to diagnose the C. reticulata group, except for character 150 state 1, which distinguishes Crenicichla and Batrachops sensu Heckel (1840). The lack of unambiguous apomorphies in Batrachops is probably due to the inclusion of Crenicichla (Batrachops) jegui, with a morphology that diverges from the remaining species of the subgenus in several aspects (see Remarks). Indeed, C. (B.) jegui was previously placed in the C. lugubris group by Ploeg (1991) and has been placed among the species of Character transformation Apomorphic condition Observations Char. 48: 0 => 1 Lateral portion of the pelvic fin with skin Convergences occur in other groups of pike thickening. cichlids. Condition apparently related to rheophilic behaviour. Char. 155: 0 => 1 Medial process (ascendent) of the proximal extrascapula about twice longer than the distal process. Char. 162: 0 => 2 Lateral crest of epioccipitals on the eppiocitals only. Char. 163: 0 => 1 Caudal opening of the posterior myodome ab- Convergent transformations optimized as sent. autapomorphies of several species. Char. 211: 1 => 0 Posterior margin of the glossohyal convex. Convergent transformation as synapomorphy of Teleocichla. Ambiguous Char. 14: 1 => 2 A Horizontal portion of the preopercle longer Optimized as a synapomorphy, but with a than the vertical portion. reversal on the node 150, which comprises four of the six terminal taxa of the group. Char. 129: 0 => 1 A Decreasing to two or three rows of teeth in Convergences as synapomorphies of the C. the symphyseal region of the premaxilla. (La.) scottii complex (node 179) and as autapomorphy of C. (La.) jupiaensis. Char. 150: 2 => 1 A Posterior margin of the preopercle with few weak serrations irregularly distributed.	en	Varella, Henrique R, Kullander, Sven O, Menezes, Naércio A, Oliveira, Claudio, López-Fernández, Hernán (2023): Revision of the generic classification of pike cichlids using an integrative phylogenetic approach (Cichlidae: tribe Geophagini: subtribe Crenicichlina). Zoological Journal of the Linnean Society 198 (4): 982-1034, DOI: 10.1093/zoolinnean/zlad021, URL: https://academic.oup.com/zoolinnean/article/198/4/982/7204395
170987F4C533510AA367F8E6C5D6F9A3.taxon	diagnosis	Diagnosis: Species in the subgenus Batrachops, except C. (B.) jegui, are distinguished from all remaining pike cichlids by the reticulate colour pattern on the flanks emerging from the dark pigmentation on the base of the scales. However, the conspicuosness of the resulting horizontal stripes forming this pattern can vary across species and through ontogeny. Species in the subgenus Batrachops, except C. (B.) jegui, are also distinguished from all remaining pike cichlids by expressing the dark blotch on the caudal fin displaced posteriorly and dorsally in relation to the base of the caudal fin and the posterior branch of the lateral line, and from all remaining species of the subtripe Crenicichlina, except those in the subgenus Lacustria, by the orange or reddish marking of the lateral abdomen in sexually mature females. All species of the subgenus Batrachops, except C. (B.) jegui and C. (B.) cyclostoma, are distinguished from all other pike cichlids, except some species of Lugubria and C. (La.) vittata, by having almost the entire caudal fin densely covered by scales (vs. caudal fin only partially covered by scales and scales arranged in single or two series along the inter-radial membranes). All species in the subgenus Batrachops are additionally distinguished from the subgenus Crenicichla by the dark vertical bars expressed as a series of blotches along the midlateral area in adults, by the presence (vs. absence) of a caudal blotch, and by the cycloid (vs. ctenoid) scales on the cheek. Batrachops is additionally distinguished from the subgenus Lacustria by the absence of a dark suborbital marking. Crenicichla (B.) jegui has a well-defined, uniformly pigmented suborbital stripe running obliquely from the ventral margin of the orbit to the corner of the preopercle, a unique condition of the species among the subgenus, presumably not homologous with the suborbital marking formed by dark dots of the subgenus Lacustria. All species of the subgenus Batrachops are additionally distinguished from Saxatilia by the absence of a humeral blotch, from Lugubria by 55 – 75 (vs. 88 – 123) scales in the E 1 series, and from Wallaciia by their larger body size (max. SL 96 – 216 mm vs. 52 – 85 mm in Wallaciia) and by the absence of serrations on the posterior border of the suplacleithrum (except W. heckeli, with smooth supracleithrum). All species of the subgenus Batrachops are also distinguished from Teleocichla and Hemeraia by having autogenous post-lachrymal infraorbitals (vs. infraorbitals 4 and 5 coossified, forming a median pore); they are further distinguished from Teleocichla by the rounded pelvic fin with the second ray longest (vs. pelvic pointed with third ray longest), and by prognathous or isognathous jaws (vs. hypognathous jaws). Further distinguished from Hemeraia by having most scales on flank ctenoid (combination of patterns 1 and 3 on the dorsal portion and pattern B 2 on the ventral portion of the body) vs. having most scales on flank cycloid (combination of patterns 3 and B 3; see characters 25 and 26 and Supporting Information, Appendix S 2, Fig. S 2). Remarks: Most species of the subgenus Batrachops correspond to the previously proposed C. reticulata group, traditionally characterized by a cylindrical body, depressed head, short snout, and large gape, all of which have been correlated with the occupation of benthic habitats (e. g. Kullander 1986; Ploeg 1991). However, in rapids of the Río Tocantins, the sympatric species C. (B.) jegui and C. (B.) cyclostoma represent strong deviations from the typical Batrachops morphology. Crenicichla (B.) jegui is a reophilic, sedentary (bottom sitter) species with very depressed body and head, eyes dorsally displaced on the head, and distinctly prognathous jaws. The reticulate colour pattern widespread among Batrachops species is replaced by a coarse pattern of light blotches and vermiculations on the entire dorsal portion of the body, suggesting cryptic coloration in fast-flowing, turbulent waters. Contrastingly, C. (B.) cyclostoma lives among dark rocks in moderate- to fast-flowing waters of the Río Tocantins and has laterally compressed body and head with a darker overall coloration. The two species also differ in the feeding apparatus, suggesting distinctive diets and foraging strategies. All species of the subgenus Batrachops have oral jaw teeth in the outer row firmly fixed, while teeth in the inner rows are slightly movable, a character that helps to distinguish Batrachops from the subgenus Crenicichla, the genera Saxatilia, Lugubria, Wallaciia, and Hemeraia, and from most species of the subgenus Lacustria. However, while outer row teeth in most species of Batrachops are unicuspid and moderately robust, C. (B.) jegui has caniniform teeth (long and thin), and the teeth of C. (B.) cyclostoma are unusually large and thick, suggesting pronounced differences in diet among the three lineages. Similarly, the lower pharyngeal jaw (LPJ) differs, with the common condition in the subgenus being a robust and wide LPJ bearing papilliform teeth on its medioposterior portion, whereas in C. (B.) jegui the LPJ is longer and thinner, bearing only cuspidated teeth, and in C. (B.) cyclostoma the LPJ is stout and strongly sutured, bearing strong molariform teeth. All this suggests that C. (B.) jegui may be a specialized, probably ambush, predator and C. (B.) cyclostoma is probably durophagous, foraging on rocky substrates similarly to Teleocichla preta (Varella et al. 2016). As a result of these ecomorphologically unique attributes, finding universally diagnostic characters shared between C. (B.) jegui, C. (B.) cyclostoma, and the remaining species of the subgenus Batrachops remains challenging. Distribution: Amazon basin (main channel and tributaries), Río Orinoco, and Essequibo river basins. Crenicichla (B.) semifasciata is the only species found in the Río Paraguai and in the lower and middle portions of the Río Paraná. Recent reports from the upper Río Paraná basin probably result from anthropogenic introductions (Roa-Fuentes et al. 2015). Node 127 — Crenicichla (Lacustria), new subgenus of Crenicichla u r n: l s i d: z o o b a n k. o r g: a c t: D 8 B 7 4 0 B 7 - E F 5 6 - 437 E- 8 DCA- 530741 D 4 D 775 Type species: Cycla lacustris Castelnau, 1855. The subgenus Lacustria is recovered in all analyses performed herein with high support in the ML and BI trees (BS 100 % and PP 91.1 %), but low support in the parsimony analysis of EW / DiscreteMatrix (ABS 3 / RBS 20 %; Table 6). It is also recovered in the recent phylogenies based on molecular data exclusively (Piálek et al. 2012; Burress et al. 2017, 2018). Six morphological synapomorphies (four unambiguous and two ambiguous) and 37 molecular unambiguous transformations are optimized for the Node 127 in the ML tree (Table 12) — all of them coincident with the optimization on the consensus tree based on parsimony analysis of the EW / DiscreteMatrixTE. Diagnosis: Species of the subgenus Lacustria are distinguished from all species of pike cichlids by the presence of a dark suborbital pattern formed by a variable number of conspicuous dark spots (punctulations), more or less scattered over the cheek. Species of Lugubria, Saxatilia, and Hemeraia also have a suborbital marking, as well as C. (B.) jegui, but this is interpreted as another pattern, not formed by spots but uniformly pigmented (see characters 57 – 60). Species of the subgenus Lacustria are further distinguished from most species of Lugubria by the presence of 41 – 75 scales in the E 1 row (vs. 88 – 123), with the exception of C. (La.) vittata (79 – 93 scales), and by the absence of a dark blotch behind the pectoral fin (vs. blotch appearing in adults of most Lugubria species). From Saxatilia, all species of Lacustria are distinguished by the absence of a humeral blotch and by sexually dimorphic females with an orange or reddish marking laterally on the abdomen instead of a broad reddish or purplish pigmentation on the ventral portion of the abdomen of Saxatilia. Lacustria is distinguished from the subgenus Batrachops, except C. (B.) cyclostoma, by the laterally compressed body (vs. nearly cylindrical or depressed) and from Batrachops, except C. (B.) jegui, by the absence of a reticulate colour pattern on the flank resulting from the dark pigmentation on the base of each flank scale. Lacustria differs from the subgenus Crenicichla by the presence of cycloid (vs. ctenoid) scales on the cheek, dorsal head, and area anterior to the dorsal fin, and on the chest; by the presence of a caudal blotch and by sexually dimorphic females showing orange or reddish marking on the lateral abdomen and, occasionally, one or more dark blotches on dorsal fin (vs. absence of these sexually dimorphic features related to coloration). Species of Lacustria are additionally distinguished from Wallaciia by their size (max. SL 95 – 294 mm vs. 52 – 85 mm) and by the smooth posterior margin of the supracleithrum (vs. posterior margin with serrations in all Wallaciia, except W. heckeli). The subgenus is additionally distinguished from Teleocichla and Hemeraia by having all the post-lachrymal infraorbitals autogenous (vs. infraorbitals 4 and 5 co-ossified forming a median pore). Distribution: Atlantic coastal rivers in Brazil (from the Río Buranhem to the Laguna dos Patos system) and in the Río Paraná, Río Uruguay, and Río Paraguay basins (Río de La Plata superbasin). A recently discovered, undescribed species expands the distribution of the subgenus to the Río São Francisco basin in north-eastern Brazil (H. Varella, pers. obs.). Remarks: Our results agree with all previous studies based on molecular data, which place Crenicichla vittata among the species of the subgenus Lacustria, former C. lacustris group (Fig. 3). This is the only species of the subgenus widely distributed in the three major rivers of the La Plata superbasin and falls in different positions of the subgenus depending on the analysis performed. Before those recent studies, C. (La.) vittata was considered a member of the C. lugubris group (e. g. Kullander 1991, 1997; Ploeg 1991; Fig. 3 I), mainly by being a medium to large species with numerous small scales on the flanks. Less inclusive groups in the subgenus Lacustria: Node 153 corresponds with what has been called the C. missioneira group or complex since Lucena and Kullander 1992 or Uruguay River species flock (URSF, Burress et al. 2017, 2018; Fig. 3 D−F). This group is recovered in all analyses performed herein and is well-supported in the ML and BI topologies (BS 100 % and PP 100 %) but has low support in the parsimony analysis of EW / DiscreteMatrixTE (ABS 2 / RBS 20 %; Table 6). Besides the species included as terminal taxa in the analyses performed herein, the C. missioneira Character transformation Apomorphic condition Observations Char. 59: 0 => 1 Presence of dark suborbital punctulations. A unique transformation of the Node 127. Char. 85: 0 => 1 Sexually dimorphic females showing dark one or Convergent transformations optimized more dark blotches on the dorsal fin. as synapomorphy of a less inclusive group of Wallaciia (Node 186) and as autapomorphy of C. (B.) cyclostoma, and a reversal occurs in C. (La.) vittatta. Char. 124: 0 => 1 Posterior border of the alveolar process of the Convergent transformations optimized premaxilla curved, bulbous. as autapomorphies of several species of Teleocichla and Hemeraia chicha. Char. 202: 1 => 0 Microbranchiospines regularly distributed on A reversal to the plesiomorphic state in- the external (lateral) face of the second to side the subtribe Crenicichlina. fourth branchial arches. Ambiguous Char. 66: 0 => 1 A Post-temporal dark marking present in adults. This character is very homoplastic inside the subtribe Crenicichlina. Optimized also as synapomorphies of the Node 132 (Clade Hemeraia-Lugubria-Saxatilia) and of Teleocichla, and with two reversals inside the subgenus Crenicichla (Lacustria). Char. 177: 1 => 2 A Posterior foramen of the nasal unique or This is a very homoplastic character and divided, diplaced anterior and with the pos- the state 2 represents a secondary modi- terior portion of the canal ossified, resulting in fication of the synapomorphic condition the openings placed at the middle of the nasal of pike cichlids (state 1). canal. complex also includes Acharches niederleinii, considered a nomen dubium of the Río Uruguay (Río Chapecó) that could correspond to C. (La.) missioneira, C. (La.) minuano, or C. (La.) hadrostigma (Varella 2011; Steinhauser 2019). All species are endemic to the middle and upper Río Uruguay basin. The C. missioneira complex can be characterized by 10 morphological synapomorphies (seven of them unambiguous) and 91 unambiguous molecular transformations. Among the morphological synapomorphies, four are useful to diagnose the group among the species of the subgenus Lacustria and were used in the previous characterization of the group by Lucena and Kullander (1992): char. 12: 2 => 1: lachrymal as long as deep (approximately square) vs. lachrymal longer than deep; char. 60: 1 => 2 D: dark suborbital marking composed of one or few spots restricted to the posterior portion of the cheek; char. 71: 0 => 1: presence of a dark blotch on the pectoral axila; and char. 150: 2 => 0 A: posterior margin of the preopercle smooth, with no serrations. Additionally, most species of the C. (La.) missioneira complex show sexually dimorphic males with the dark blotch of the caudal fin fragmented, accompanying the pattern of irregular small blotches scattered on the caudal peduncle.	en	Varella, Henrique R, Kullander, Sven O, Menezes, Naércio A, Oliveira, Claudio, López-Fernández, Hernán (2023): Revision of the generic classification of pike cichlids using an integrative phylogenetic approach (Cichlidae: tribe Geophagini: subtribe Crenicichlina). Zoological Journal of the Linnean Society 198 (4): 982-1034, DOI: 10.1093/zoolinnean/zlad021, URL: https://academic.oup.com/zoolinnean/article/198/4/982/7204395
170987F4C533510AA367F8E6C5D6F9A3.taxon	description	According to our results, the occurrence of hypertrophied lips is not restricted to C. tendybaguassu and C. tuca, but also observed in two other morphologically defined, putative new taxa belonging to the subgenus Lacustria in the Río Iguaçu basin. Three pairs of big-lipped / typical taxa of pike cichlids in the Río Iguaçu basin, matching in relation to overall shape and coloration, were included in the combined phylogenetic analysis. Crenicichla iguassuensis BL and C. tesay BL are big-lipped forms corresponding to C. iguassuensis and C. tesay, respectively. The third pair is C. tapii (typical) and C. tuca (big lips). Crenicichla iguassuensis BL and C. tesay BL were represented in the combined datasets only with morphological data and the remaining were represented with both molecular and morphological data.	en	Varella, Henrique R, Kullander, Sven O, Menezes, Naércio A, Oliveira, Claudio, López-Fernández, Hernán (2023): Revision of the generic classification of pike cichlids using an integrative phylogenetic approach (Cichlidae: tribe Geophagini: subtribe Crenicichlina). Zoological Journal of the Linnean Society 198 (4): 982-1034, DOI: 10.1093/zoolinnean/zlad021, URL: https://academic.oup.com/zoolinnean/article/198/4/982/7204395
170987F4C533510AA367F8E6C5D6F9A3.taxon	description	Considering our comparisons between big-lipped and typical-lipped correspondents, features on the LPJ (directly correlated to food processing, e. g. Burress et al. 2017), are more conservative than those on the oral jaws (correlated to foraging). LPJ of big-lipped and typical-lipped correspondents are similar in shape and robustness (character 205) and type of teeth on the medioposterior portion (character 206), but the LPJ of big-lipped taxa shows denser dentition [as already pointed out by Lucena and Kullander (1992) for C. tendybaguassu in the context of the C. missioneira complex]. Even in C. iguassuensis vs. C. iguassuensis BL pair, with pronounced differences on morphology of oral jaws, the LPJ of the big-lipped form has denser dentition but is very similar in shape and types of teeth (only unicuspid and bicuspid teeth, no papilliform or molariform teeth, which suggests preference for durophagy). The only clearly specialized molluscivorous species in the Río Iguaçu basin is what we identify as C. yaha (sensu Varella 2011; but for incongruences on species delimitations between authors, see also: Piálek et al. 2015), with much enlarged medioposterior teeth on the LPJ, and this is the only species that does not have a big-lipped correspondent.	en	Varella, Henrique R, Kullander, Sven O, Menezes, Naércio A, Oliveira, Claudio, López-Fernández, Hernán (2023): Revision of the generic classification of pike cichlids using an integrative phylogenetic approach (Cichlidae: tribe Geophagini: subtribe Crenicichlina). Zoological Journal of the Linnean Society 198 (4): 982-1034, DOI: 10.1093/zoolinnean/zlad021, URL: https://academic.oup.com/zoolinnean/article/198/4/982/7204395
170987F4C50E5108A0CDF998C653FB07.taxon	materials_examined	Type species: Crenicichla wallacii Regan, 1905. The genus Wallaciia is recovered in the ML and BI analyses, as well as in the parsimony analysis, under extended implied weighting (XIW / DiscreteMatrixTE and XIW / ContinuousMatrixTE). However, it is not recovered in the parsimony analyses under equal weighiting (EW / DiscreteMatrix and EW / ContinuousMatrixTE). We hypothesize that the discordance is a result of long-branch attraction caused by convergence in characters associated with specializations to rheophily (see Remarks). Such convergence clustered W. heckeli and W. urosema, two species that inabit rapids and were represented by morphological characters only, with Teleocichla. Although the node corresponding to the genus Wallaciia was moderately supported in the ML tree (BS 71 %; Table 6), it shows high support in the BI tree (PP 100 %) and is recovered as monophyletic in all previous molecular studies (Kullander et al. 2010; Piálek et al. 2012; Burress et al. 2017, 2018; Fig. 3). Moreover, three of the five unambiguous morphological synapomorphies optimized for the group (Table 13) are congruent with characters used to diagnose the C. wallacii group in previous studies and also to diagnose the genus Wallaciia as proposed herein. Diagnosis: Wallaciia includes small-sized species (max. SL 52 – 85 mm) that differ from all pike cichlids by the following combination of traits: large eyes (orbital diameter 7.8 – 12.6 % of SL); presence of serrations on the posterior margins of the supracleithrum; posterior margin of preopercle with prominent, spine-like serrations; pectoral fin with 13 – 14 rays (vs. 15 – 18; in modal values); reduced predorsal squamation (not extending on to the NFL 0; see char. 28); and pelvic fin rounded with second ray longest and all post-lachrymal infraorbitals autogenous (vs. infraorbitals 4 and 5 co-ossified, forming a median opening). Wallaciia heckeli deviates from the typical Wallaciia morphology by having some characters convergently shared with Teleocichla, another small-bodied genus. For example, W. heckeli has smooth preopercle and suprachleitrum, jaws not prognathous, and the pattern of fixation of oral teeth and configuration of the lower pharyngeal jaw is different from the typical Wallaciia species. Also, it can be distinguished from other Wallaciia species by having smaller eyes, infraorbitals co-ossified forming a median opening (vs. 4 and 5 autogenous), and pelvic fin pointed, with third ray longest (vs. rounded). Besides, Wallaciia differs from Teleocichla by lacking several other osteological modifications associated to life in rapids and diagnostic of the later (see below). Species of Wallaciia are additionally distinguished from Saxatilia by the absence of a humeral blotch and suborbital marking, from Lugubria by having fewer scales in the E 1 series (52 – 64 vs. 88 – 123 scales) and fewer vertebrae (31 – 34 vs. 38 – 42), from Hemeraia by having most scales on the flanks ctenoid (combination of patterns 1 and 2 on dorsal portion and pattern B 2 on ventral portion of the flank) vs. most flank scales cycloid (combination of patterns 3 and B 3). Wallaciia are additionally distinguished from the subgenus Crenicichla of Crenicichla, by cycloid (vs. ctenoid) scales on the cheek and on the chest. Wallaciia differs from the subgenus Batrachops of Crenicichla by the absence of a reticulate colour pattern on the flank (vs. reticulate pattern on flanks formed with the dark pigmentation on the base of the scales), and from the subgenus Lacustria of Crenicichla by the absence of a suborbital marking (vs. presence of dark puntulations more or less scattered on the cheek forming different patterns of suborbitals markings). Distribution: As proposed here, Wallaciia includes eight species that are distributed in the Amazon basin, Río Orinoco, and Essequibo river basins. Species of Wallaciia are present in almost all major rivers of the Amazon basin from the Río Madeira to the Río Tocantins, but are not found in the Western portion of the Amazon basin, i. e. tributaries and main channel of the Amazon basin west of Santo Antonônio do Içá. Remarks: This is the first phylogenetic study of pike cichlids that includes Wallaciia heckeli and W. urosema, two rheophilic species that inhabit rapids of the Río Trombetas and Río Tapajós basins, respectively. In the two parsimony analyses of the combined datasets under the equal weighting scheme, these species cluster with Teleocichla, as successive sister-groups. This is probaby a result of long-branch attraction of some convergences shared by W. heckeli and Teleocichla, resulting in synapomorphies of an expanded Teleocichla (including W. heckeli and W. urosema) or of the group Teleocichla + W. heckeli in the aforementioned parsimony trees. Many of these convergent characters are correlated with specializations for rheophilic habitats and specialized benthic diets (e. g. Kullander 1988; Varella et al. 2016), including reduced cheek squamation, differences in number of precaudal and caudal vertebrae, acute mouth with narrow gape, ridged vomer (linked with a downturned mouth), and modifications on the format and teeth of the lower pharyngeal jaw (char. 19: 0 => 2; char. 113: 0 => 1; char. 128: 0 => 1; char. 159: 1 => 2 A; char. 205: 2 => 3; char. 206: 0 => 1). Interestingly, these characters are shared between Teleocichla and three species of Wallaciia instead of two — the third being W. compresiceps, also a rheophilic species that inhabits rapids in the Río Tocantins. Further study is needed to help clarify the relationships of rheophilic Wallaciia species, because W. compressiceps is the only taxon in our matrix represented by both molecular and morphological data, whereas the others are represented by morphology only, which is probaby the reason why W. compressiceps grouped with Wallaciia instead of Teleocichla, as did the two species with only morphological data.	en	Varella, Henrique R, Kullander, Sven O, Menezes, Naércio A, Oliveira, Claudio, López-Fernández, Hernán (2023): Revision of the generic classification of pike cichlids using an integrative phylogenetic approach (Cichlidae: tribe Geophagini: subtribe Crenicichlina). Zoological Journal of the Linnean Society 198 (4): 982-1034, DOI: 10.1093/zoolinnean/zlad021, URL: https://academic.oup.com/zoolinnean/article/198/4/982/7204395
170987F4C50C510DA0CDFB3CC4DFFA90.taxon	description	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 E 1 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	en	Varella, Henrique R, Kullander, Sven O, Menezes, Naércio A, Oliveira, Claudio, López-Fernández, Hernán (2023): Revision of the generic classification of pike cichlids using an integrative phylogenetic approach (Cichlidae: tribe Geophagini: subtribe Crenicichlina). Zoological Journal of the Linnean Society 198 (4): 982-1034, DOI: 10.1093/zoolinnean/zlad021, URL: https://academic.oup.com/zoolinnean/article/198/4/982/7204395
170987F4C5095103A367FAD0C5DEFBA3.taxon	materials_examined	Type species: Crenicichla hemera Kullander, 1991 Hemeraia, comprising only H. hemera and H. chicha, is recovered in all analyses performed herein, although in different positions within the subtribe Crenicichlina (see Remarks): as sister-group of the clade Saxatilia (ML and BI trees); in a trichotomy with Saxatilia and Lugubria (Parsimony / EW / DiscreteMatrixTE); as sister-group of the the clade Saxatilia + Lugubria (Parsimony / XIW / DiscreteMatrixTE); and as sister-group of all remaining pike cichlids (parsimony analysis of ContinuousMatrixTE under EW and XIW schemes). As this is the first phylogenetic study that included both H. hemera and H. chicha, there are no previously published phylogenetic hypotheses for comparisons. Nevertheless, monophyly of the genus is well supported in the ML and BI trees (BS 98 % and PP 100 %), and has moderate support in the parsimony tree based on EW / DiscreteMatrixTE (ABS 5 / RBS 63 %; Table 6). The two known species in the genus were represented only by morphological data, but nonetheless the clade was diagnosable by six unambiguous and one ambiguous synapomorphies (Table 15). Synapomorphic conditions of characters 25, 26, and 105 help to diagnose the group among pike cichlids and coincide with diagnostic characters in previous taxonomic papers that included the two species (Varella et al. 2012, 2018). Diagnosis: Hemeraia is distinguished from other pike cichlids by the combination of: reduced distribution of ctenoid scales (vs. cycloid) on flanks (pattern 3 and B 3; see characters 25, 26 and Supporting Information, Appendix S 2, Fig. S 2); a smooth preopercle (vs. serrated); infraorbitals 4 and 5 co-ossified, forming a median pore (vs. autogenous); and presence of a suborbital marking. Hemeraia is further distinguished from Lugubria by having fewer scales in the E 1 series (58 – 75 vs. 89 – 123). Hemera is further distinguished from Saxatilia by the absence of a humeral blotch that appears in early ontogenetic stages (only adults of H. hemera shows a dark blotch just posterior to the pectoral fin that resembles a humeral blotch but appears only late in ontogeny). Hemeraia is further distinguished from all species of Teleocichla, which also have infraorbitals 4 and 5 co-ossified and (most of them) a smooth preopercle, by having typical pike-cichlid morphology without the various characters related to rheophyly found in Teleocichla, including the pelvic pointed with third ray longest, almost reaching the genital papilla, mouth downturned, reduced cheek squation, and some degree of molarization of LPJ teeth. Distribution: Hemeraia hemera is distributed in the Río Aripuanã basin and H. chicha in the Río Juruena, Río Tapajós basin. Remarks: The position of species of Hemeraia species within the subtribe Crenicichlina is controversial in both previous taxonomic studies and in our present work. Ploeg (1991) allocated Crenicichla guentheri (junior synonym of C. hemera) among the species of the C. saxatilis group, which herein corresponds to Saxatilia. Varella et al. (2012), in the description of Crenicichla chicha and based mainly on external morphology, considered Hemeraia hemera and H. chicha as sister-taxa by sharing features such as reduced ctenoid squamation on flanks, smooth preopercle, and infraorbitals 4 and 5 co-ossified. However, their placement in any of the major groups of pike cichlids remained unclear because these species share some characters (mostly related to meristics) with the C. saxatilis (herein Saxatilia) group and others with the C. lugubris (herein Lugubria) group (related to ontogenetic change of coloration). The discordance of the position of Hemeraia between our different analyses may be partially due to the lack of molecular data, together with the use of the first 16 characters as continuous data. The clade gathering Hemeraia, Saxatilia, and Lugubria, i. e. node 131 in ML tree (Fig. 2 left) and node 129 in the parsimony tree based on EW / DiscreteMatrixTE (Fig. 2 right), is supported by five and six unambiguous synapomorphies, respectively. Among these, characters 4 (number of dorsal-fin rays) and 5 (number of anal-fin rays) are multistate quantitative characters treated as continuous in the ContinuousMatrixTE. As a side-effect of treating additive characters as continuous, these characters are downweighted in the analysis and the resulting shifts are different from the discretizations of the states performed a priori. This side-effect would have been of little impact if molecular data of these species were available to be included in the analyses.	en	Varella, Henrique R, Kullander, Sven O, Menezes, Naércio A, Oliveira, Claudio, López-Fernández, Hernán (2023): Revision of the generic classification of pike cichlids using an integrative phylogenetic approach (Cichlidae: tribe Geophagini: subtribe Crenicichlina). Zoological Journal of the Linnean Society 198 (4): 982-1034, DOI: 10.1093/zoolinnean/zlad021, URL: https://academic.oup.com/zoolinnean/article/198/4/982/7204395
170987F4C5075103A0FFFB98C22CF9CA.taxon	materials_examined	Type species: Perca saxatilis Linnaeus, 1758 Saxatilia is recovered as monophyletic with high support in the ML and BI trees (BS 100 %, PP 100 %), and moderate to low support in the parsimony analysis based on EW / DiscreteMatrixTE (ABS 3 / RBS 43 %; Table 6). This genus comprises 23 valid species and corresponds almost entirely to the C. saxatilis group, considered monophyletic in all previous studies since Ploeg (1991; see Fig. 3). Thirty-one unambiguous molecular transformations and 11 morphological synapomorphies (eight of them unambiguous) are optimized for the group (Table 16). Diagnosis: Species of Saxatilia can be diagnosed from all other pike cichlids by the presence of humeral blotch, which is evident from juvenile stages. Most species of Lugubria also have a dark blotch posterior to the pectoral fin that resembles a humeral blotch but appears only later in ontogeny. Species of Saxatilia are additionally distinguished from all pike cichlids by the combination of 31 – 71 scales in the E 1 series, suborbital marking present, serrated preopercle, slender lachrymal, and sexual dimorphism in which males usually have light, small botches scattered on flanks and females show a round belly with a broad reddish or purplish pigmentation. Saxatilia also differs from Hemeraia by having most scales on the flank ctenoid (combination of patterns 1 or 2 on the dorsal area and pattern B 2 on the ventral area; see characters 25 – 26 and Supporting Information, Appendix S 2, Fig. S 2) vs. cycloid (combination of patterns 3 and B 3; see characters 25 – 26 and Supporting Information, Appendix S 2, Fig. S 2). From Lugubria, Saxatilia differs by the number of vertebrae (31 – 36 vs. 38 – 42 in Lugubria), from Wallaciia by the absence (vs. presence) of serrations on the posterior margin of supracleithrum, and from Teleocichla by all post-lachrymal infraorbitals autogenous (vs. infraorbitals 4 and 5 co-ossified), pelvic fin short and rounded with second ray longest (vs. pointed with third ray longest almost reaching genital papilla), and by the absence of osteological modifications related to rheophyly. Saxatilia is distinguished from the subgenus Crenicichla by the presence (vs. absence) of a dark blotch on the caudal fin and by cycloid (vs. ctenoid) scales on cheek and chest. From all species of the subgenus Batrachops of Crenicichla, except C. (B.) jegui, Saxatilia differs by the absence of a reticulate colour pattern on the flank (vs. having a reticulate pattern formed by the pigmentation on the base of each scale); and from the subgenus Lacustria of Crenicichla by a suborbital marking uniformly pigmented instead of formed by dark punctulations more or less scattered on the suborbital region. Distribution: Saxatilia is the most widespread genus of pike cichlids, occurring in all major rivers of South America, including the Río Orinoco basin, the Amazon basin, all major drainages of the La Plata basin, the coastal drainages of Trinidad and Tobago, the Guianas, and Atlantic coastal drainages from north-eastern Brazil to the Lagoa dos Patos and Lagoa Mirim systems in southern Brazil and Uruguay. Recent collections from areas influenced by hydroelectric dams in coastal drainages in south-eastern Brazil (e. g. Río Paraíba do Sul) have shown the presence of Saxatilia (preliminarily identified as S. britskii), but it is unclear if these records represent native distribution or recent human introductions.	en	Varella, Henrique R, Kullander, Sven O, Menezes, Naércio A, Oliveira, Claudio, López-Fernández, Hernán (2023): Revision of the generic classification of pike cichlids using an integrative phylogenetic approach (Cichlidae: tribe Geophagini: subtribe Crenicichlina). Zoological Journal of the Linnean Society 198 (4): 982-1034, DOI: 10.1093/zoolinnean/zlad021, URL: https://academic.oup.com/zoolinnean/article/198/4/982/7204395
170987F4C5075107A367F961C4B2F8E4.taxon	materials_examined	Type species: Crenicichla lugubris Heckel, 1840.	en	Varella, Henrique R, Kullander, Sven O, Menezes, Naércio A, Oliveira, Claudio, López-Fernández, Hernán (2023): Revision of the generic classification of pike cichlids using an integrative phylogenetic approach (Cichlidae: tribe Geophagini: subtribe Crenicichlina). Zoological Journal of the Linnean Society 198 (4): 982-1034, DOI: 10.1093/zoolinnean/zlad021, URL: https://academic.oup.com/zoolinnean/article/198/4/982/7204395
170987F4C5075107A367F961C4B2F8E4.taxon	diagnosis	Diagnosis: Lugubria includes species with relatively large body (max. SL 225 – 300 mm) distinguished from all other pike cichlids by the combination of reduced size of the scales in the whole body reflected in the high number in the E 1 row (88 – 123 scales; see Remarks), the high count of soft rays in the dorsal and anal fins (13 – 18 and 9 – 13, respectively), and the high number of vertebrae (39 – 44 vertebrae). Crenicichla (Lacustria) vittata is similar to species of Lugubria by having high counts of vertebrae and scales in the E 1 row but differs from all species of that genus by females showing orange or reddish pigmentation on the lateral portion of the abdomen, whereas in species of Lugubria this pattern is not observed — in most species, sexually dimorphic females have a broad reddish or purplish broad pigmentation on the ventral portion of the abdomen (belly). Crenicichla (Lacustria) vittata also shows a suborbital marking composed of distinct dark spots that together form a stripe, which is characteristic of other species of the subgenus Lacustria of Crenicichla, whereas species of Lugubria do not show suborbital marking or have it uniformly pigmented. Finally, most species of Lugubria show a dark blotch posterior to the pectoral fin that appears only late during ontogeny and is absent in C. (La.) vittata. Species of Lugubria are further distinguished from Saxatilia by the absence (vs. presence) of a humeral blotch that appears early in ontogeny and remains in adults; from Hemeraia and Teleocichla by having all post-lachrymal infraorbitals autogenous (vs. infraorbitals 4 and 5 co-ossified) and posterior margin of preopercle strongly serrated (vs. preopercle smooth or with weak serrations irregularly distributed along the posterior margin); and from Wallaciia by the absence (vs. presence) of serrations on the posterior margin of the supracleithrum. Species of Lugubria are additionally distinguished from the subgenus Crenicichla by having all scales on head cycloid (vs. scales ctenoid on cheek and predorsal area) and by the presence (vs. absence) of a dark blotch on the caudal fin, with the exception of Lugubria johanna and Lu. monicae, which lack the caudal fin blotch. Species of Lugubria are futher distinguished from all species of the subgenus Batrachops of crenicichla, except C. (B.) jegui, by the absence of a reticulate pattern of coloration on the flank (vs. presence of a reticulate pattern resulting from the dark pigmentation on the base of each scale on the flank). Distribution: Lugubria is distributed in all major tributaries of the Amazon basin, in the Río Orinoco basin, and coastal drainages of the Guianas. There are no records of Lugubria in the Atlantic coastal drainages or the La Plata basin. Remarks: Since Kullander (1991, 1997), species of the C. lugubris group have been subdivided into the C. acutirostris group and C. lugubris group s. s. (e. g. Montaña et al. 2008; Kullander and Varella 2015). Crenicichla (Lacustria) vittata and C. (Batrachops) jegui were tentatively allocated in the C. acutirostris group, which has recently been contested (Piálek et al. 2012; Burress et al. 2017, 2018; this paper; see Fig. 3). The distinction between these subgroups was based on a series of characters in combination, including a depressed head and long, pointed snout in the C. acutirostris group, with nostril halfway between the postlabial skin fold and the anterior margin of orbit vs. blunt snout in the C. lugubris group s. s., with nostril closer to postlabial skin fold, as well as some details of coloration related to ontogeny. All analyses performed herein recover a Lu. lugubris group s. s. (Node 137) as monophyletic. Among the synapomorphies for this group, some are related to the blunt snout (char. 13: 1 => 0 and char. 96: 0 => 1) and the changes on the coloration during ontogeny, i. e. the presence of dark spots scattered on the anterior portion of the body (in juveniles) and the presence (in adults) of a dark blotch posterior to the pectoral fin and another on the pectoral axilla (chars. 70: 0 => 1; char. 71: 0 => 1; and 81: 0 => 1). The Lu. acutirostris group sensu Kullander (1991, 1997), including all long-snout species of Lugubria, is recovered only in the parsimony analyses under extended implied weighting but not recovered in our main hypothesis (ML tree), in the BI tree and in the parsimony analyses under equal weighting. CHARACTER CONGRUENCE AND INSIGHTS ON PHENOTYPIC DIVERSIFICATION Discussions on macroevolution of the clade of pike cichlids (subtribe Crenicichlina) have been focused on the subgenus Lacustria of Crenicichla, which is restricted to the rivers of southern South America (Piálek et al. 2012, 2019 a; Burress et al. 2017, 2018). These studies suggest the existence of adaptive radiations in the C. (La.) missioneira complex in the Río Uruguay and of the C. (La.) mandelburgeri complex in the Río Paraná. Except for their occurrence in complex riverine (instead of lacustrine) habitats, these radiations were considered similar to the species flocks in the African Great Lakes and Middle America (Piálek et al. 2012), i. e. monophyletic assemblages of closely related species (shallow genetic divergence) that coexist in the same area with a high level of endemicity and ecomorphological specialization. Our results agree with these previous studies in demonstrating the occurrence of parallel evolution of ecomorphological traits in geographically isolated habitats between the C. missioneira and C. mandelburgeri complexes, most of them related to feeding and microhabitat occupation (e. g. Burress et al. 2018; Piálek et al. 2019 a). Several characters present in these clades correlated to rheophyly and durophagy were highly homoplastic among pike cichlids. These characters were clearly homoplastic and phylogenetically misleading when morphological characters were analysed alone, resulting in lack of resolution and long-branch attraction. More generally, ecomorphological convergence is rampant beyond the well-known cases within the subgenus Lacustria, with clear examples among multiple pike cichlid taxa coexisting in complex environments such as rapids. Examples are the assemblages comprising C. (B.) jegui, C. (B.) cametana, and C. (B.) cyclostoma occurring in sympatry in rapids of the Río Tocantins basin and the diverse assemblages of Teleocichla and Lugubria in rapids of the Río Xingu basin. Varella et al. (2018) discussed characteristics related to microhabitat occupation and feeding within the assemblage of Teleocichla from the Río Xingu, most of them reanalysed herein in a phylogenetic framework and confirmed as convergent. Likewise, Varella and Ito (2018) explored the large diversity of the C. lugubris group (= Lugubria) in the Río Xingu basin and highlighted the convergent dark coloration between Lugubria dandara, Teleocichla preta, and Crenicichla (La.) hu, which is comparable to that of Crenicichla (B.) cyclostoma and C. (B.) cametana. Dark coloration in these species might be understood as a convergent adaptation for inhabiting swift, clear-water rapids over beds of dark rocks widespread in the Brazilian Shield. On the other hand, the diversity of the genera Lugubria, Saxatilia, and Wallaciia and of the subgenus Batrachops of Crenicichla suggest widespread allopatric divergence resulting in ecomorphologically similar species across river basins. Most species or populations within their widespread species (see: Říčan et al. 2021 a) are distributed following hydrography or geological history. In these groups, there is widespread variation in sexually selected traits (sexual dimorphism and dichromatism), but traits associated with feeding or habitat occupation vary much less, with ecomorphological variation mostly restricted to pairs or small groups of sympatric species living in fast current environments. From our study, it remains unclear if sympatric assemblages diversified in situ (like those of Lacustria radiations in the Uruguay and Paraná rivers), because our taxonomic sampling, character coding, and analyses focused on assessing monophyly clarifying relationships among the major groups of pike cichlids, sacrificing resolution of less-inclusive groups. The taxonomic sampling of recent phylogenomic analyses (Piálek et al. 2012, 2019 a; Burress et al. 2017, 2018) also prevents inference of relationships within these less studied sympatric assemblages. Nevertheless, Říčan et al. ’ s (2021 a) recent study using a large taxonomic sampling but only two mtDNA markers suggests monophyly of the assemblage of Lugubria in the Río Xingu basin and of Crenicichla (Batrachops) in the Río Tocantins basin, but not of the large sympatric assemblage of Teleocichla in the Río Xingu basin. Říčan et al. (2021 a, b) studies also suggest the existence of smaller sympatric groupings of Crenicichla (Lacustria) that evolved parallel ecomorphs in different rivers basins of the Río Paraná basin. In that context, resource partitioning apparently plays an important role in the origin or maintenance of the phenotypic diversity among rheophilic pike cichlids in different river basins or, more generally, among coexisting but phylogenetically distant species that occupy more or less specialized ecological niches. Moreover, resource partitioning apparently plays a role even within the already specialized rheophilic assemblage of Teleocichla in the Río Xingu (Zuanon 1999; Varella et al. 2016). From this perspective, the role of ecological speciation may be a fruitful avenue for subsequent studies on the evolution of pike cichlids beyond the C. (La.) missioneira and C. (La.) mandelburgeri complexes. Recent studies with broader taxonomic scope (e. g. Arbour and López-Fernández 2014; Astudillo-Clavijo et al. 2015) place the phenotypic and funcional diversification of pike cichlids in the broader context of the tribe Geophagini and Neotropical cichlids, highlighting that pike cichlids occupy a vast and largely unique functional morphospace among their South American relatives. However, because few representatives of pike cichlids were included in these analyses, much about the evolution within the clade remains unexplored and will be the subject of future studies.	en	Varella, Henrique R, Kullander, Sven O, Menezes, Naércio A, Oliveira, Claudio, López-Fernández, Hernán (2023): Revision of the generic classification of pike cichlids using an integrative phylogenetic approach (Cichlidae: tribe Geophagini: subtribe Crenicichlina). Zoological Journal of the Linnean Society 198 (4): 982-1034, DOI: 10.1093/zoolinnean/zlad021, URL: https://academic.oup.com/zoolinnean/article/198/4/982/7204395
