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Node 110 = Stromateidae Stromateus brasiliensis, Pampus cinereus, Peprilus triacanthus and Peprilus paru. Unambiguous synapomorphies: Character 6 (24– 26> 34–39): number of anal-fin rays increased to 34–39; character 14 (60.1–60.6%> 62.1–63.3%): dorsal-fin base length increased to 62.1–63.3% of SL; character 18 (34.8–37.6%> 52.2%): anal-fin base length increased to 52.2% of SL; character 39 (0> 1): additional basibranchial cartilage located posterior to the fourth basibranchial present; character 56 (0> 1): pelvic fins absent in adults; character 68 (0> 1): number of supernumerary dorsal-fin spines associated with the first dorsal-fin pterygiophore decreased to one; character 107 (0> 1): levator operculi originating from both neurocranium and pectoral girdle; character 166 (0> 1): tooth patches on ventral surface of pharyngeal sac present; character 167 (0> 1): sphincter oesophagi fibres connecting the rear of the dorsal and ventral hemispheres of the pharyngeal sac; character 174 (0> 1): contralateral ligamentous association between basipterygia absent; character 204 (0> 1): fleshy groove at the ventral profile of the body restricted to area around urogenital papilla. Support: Relative Bremer = 100%. Remarks: The family Stromateidae (node 110; Fig. 70) is resolved herein as monophyletic based on 11 synapomorphies and a relative Bremer support of 100% (Fig. 69). Previous cladistic studies (Horn, 1984; Doiuchi et al., 2004; Doiuchi & Nakabo, 2006; Miya et al., 2013; Campbell et al., 2018; Friedman et al., 2019) have always recovered a monophyletic Stromateidae with robust support. Haedrich (1967) referred to Stromateidae as the current ‘zenith’ of stromateiform evolution, in allusion to the number of derived characteristics present in these fishes. He diagnosed the family based on a combination of characteristics that includes, among others, a lack of pelvic fins in adults and pharyngealsac papillae rakers not in bands or folds (Fig. 62A, B), with stellate bases (Figs 58C, 60C, D, 61D) and distributed in both the upper and lower halves of the sac (Fig. 58D). Horn’s (1984) stromateiform phylogeny was the first to test the monophyly of the family and recovered a monophyletic Stromateidae based on four synapomorphies: pharyngeal-sac papillae rakers with a distinct vertical axis, papillae rakers with a stellate base, lachrymals reduced and pelvic fins absent in adults (Horn, 1984: characters 7, 8, 11 and 19, respectively). The phylogeny of Doiuchi et al. (2004) also retrieved a monophyletic Stromateidae, this time based on 11 synapomorphies (Doiuchi et al., 2004: fig. 12). Despite the effort to include all the characters described by Horn’s (1984) and Doiuchi et al. (2004), many of them showed overlapping states when examined on a broader outgroup sampling and had to be excluded from our matrix. Nevertheless, three of the four stromateid synapomorphies listed by Horn (1984; our characters 56, 162 and 164) and four of the 11 listed by Doiuchi et al. (2004; our characters 39, 42, 56 and 66) could be included unambiguously in our dataset. The only character that is common among all three analyses is also the most obvious, i.e. pelvic fin absent in adult stromateids (our character 56). The other common synapomorphy between our study and previous morphological phylogenies is the presence of an extra basibranchial cartilage lying between the proximal portions of ceratobranchials 4 and 5 (Fig. 37), coded herein on character 39, which is modified from Doiuchi et al. (2004: character 19). The characters related to the pharyngeal sac of Stromateidae used by Horn (1984), i.e. pharyngeal-sac papillae rakers elongated and with stellate bases, were recovered herein as synapomorphies for a larger clade (i.e. node 94: Figs 69, 70). Likewise, the synapomorphies of Doiuchi et al. (2004), i.e. toothless palatine and continuous anterior and posterior dorsal fins, are optimized herein as plesiomorphic for the family.

Pastana, Murilo N L, Johnson, G David, Datovo, Aléssio (2022): Comprehensive phenotypic phylogenetic analysis supports the monophyly of stromateiform fishes (Teleostei: Percomorphacea). Zoological Journal of the Linnean Society 195 (3): 841-963, DOI: 10.1093/zoolinnean/zlab058, URL: https://academic.oup.com/zoolinnean/article/195/3/841/6372991

Tetragonuridae Tetragonurus cuvieri. Autapomorphies: Character 0 (11–12> 19): number of dorsal-fin spines increased to 19; character 1 (14> 11): number of dorsal-fin soft rays decreased to 14; character 3 (17–19> 14): number of branched pectoral-fin rays decreased to 14; character 5 (3> 1): number of anal-fin spines decreased to one; character 6 (15> 11): number of anal-fin soft rays decreased to 11; character 9 (33–36> 53): number of vertebrae increased to 53; character 10 (9> 11): number of dorsal procurrent rays increased to 11; character 13 (36.3–36.8%> 41.2%): predorsal length increased to 41.2%; character 14 (60.1–60.6%> 34.8%): dorsal-fin base length decreased to 34.8%; character 15 (24.9– 26%> 13.6%): pectoral-fin length decreased to 13.6%; character 16 (36.1–36.8%> 30.7%): prepelvic length decreased to 30.7%; character 18 (31.3–37.4%> 10.7%): anal-fin base length decreased to 10.7%; character 23 (0> 1): sclerotic bone absent; character 30 (1> 0): medial bony suture between anterior and posterior ceratohyals absent; character 38 (1> 0): interarcual cartilage absent; character 41 (1> 0): vomerine teeth present; character 42 (1> 0): palatine teeth present; character 45 (1> 0): basihyal/hypobranchial/ basibranchial teeth present; character 84 (0> 1): supraneurals absent; character 92 (0> 1): adductor mandibulae pars malaris reaching the dorsolateral edge of the preopercle; character 98 (1> 0): adductor mandibulae partes rictalis and stegalis completely separated; character 100 (0> 1): adductor mandibulae segmentum mandibularis notched anteriorly; character101(1>0):adductormandibulaeparscoronalis not extending past the posterodorsal border of the lower jaw; character 102 (1> 0): adductor mandibulae pars mentalis not differentiated into prementalis and postmentalis sections; character 108 (1> 0): RLA-OP medial to the levator operculi; character 110 (1> 0): dilatator operculi and levator arcus palatini separated only in part; character 131 (0> 1): rectus ventralis I present; character 133 (1> 0): pharyngoclavicularis externus lateral to sternohyoideus; character 135 (0> 1): pharyngoclavicularis internus subdivided; character 139 (1> 0): pharyngoclavicularis internus anteriorly positioned in relationship to pharyngoclavicularis externus; character 146 (1> 0): anterior border of the anterolateral section of the epaxialis not reaching the vertical through the middle of the orbit; character 149 (1> 0): anterior border of the anteromedial section of the epaxialis not reaching the vertical line through the middle of the orbit; character 182 (1> 0): intradermal canal plexus absent; character 199 (1> 2): scales spinoid; character 203 (1> 0): fleshy groove at the ventral profile of the body absent. Support: –. Remarks: The family Tetragonuridae comprises one genus and three valid species, Tetragonurus atlanticus Lowe, 1839, Tetragonurus pacificus Abe, 1953 and Tetragonurus cuvieri (Haedrich & Horn, 1972; Fricke et al., 2020). The family groups the most modified species of Stromateiformes, as evidenced by its long branch containing 35 autapomorphies (Fig. 70). These include numerous features unique among stromateiforms, such as the highest number of dorsalfin spines (character 1), the lowest number of analfin spines (character 8), small dorsal- (character 17) and anal-fin base lengths (character 21), absence of supraneurals (character 101), absence of an intradermal canal system on the body (character 187) and presence of spinoid scales (character 206). These fishes additionally exhibit unique characteristics such as a highly modified lower jaw, equipped with teeth that resemble the edge of a saw (Fig. 48), a poorly ossified cranial skeleton (Fig. 53) and an imbricate scale pattern along the body (Fig. 71A) that, according to some authors, forms a water-filled channel system analogous to the intradermal canal plexus present on the remaining stromateiforms (Bone & Brook, 1973). Nevertheless, these characteristics were not included in the present analysis owing to their autapomorphic occurrence in our representative of Tetragonuridae (Tetragonurus cuvieri). Tetragonurids also differ from other stromateoids in their primary association with gelatinous invertebrates. Juveniles and young adults of Tetragonurus are always observed associated with salps and pyrosomes, whereas most stromateiforms are symbiotic with jellyfish and ctenophores. According to the behavioural study by Janssen & Harbison (1981), juvenile tetragonurids spend most of their life confined inside salps and pyrosomes, feeding on their guts and gonads. This highly specific symbiotic association with tunicates might be correlated with some of their aberrant morphological modifications, such as the specialized jaw and teeth (Fig. 48) and the elongate and shallow body (Fig. 71A). The symbiosis of Tetragonurus with tunicates, rather than jellyfishes, might also explain the absence of an intradermal canal plexus on their bodies, because one of the possible functions of this system is to protect against cnidarian stings. Given the number of morphological modifications present in tetragonurids, it is not surprising that the family has been hypothesized to belong to different lineages of stromateiforms. A Tetragonuridae + Ariommatidae sister-group relationship, as supported herein (and by Horn, 1984), is based mostly on shared pharyngeal-sac specializations (see Remarks on node 92). Alternative Tetragonuridae affinities include a sister-group relationship with stromateids (sensu Doiuchi et al., 2004; Doiuchi & Nakabo, 2006). Morphologically, this grouping was based on three shared features (anterior and posterior ceratohyals medially separated by a strip of cartilage, pharyngobranchial 3 articulation site with epibranchial 3 shifted anteriorly, and epurals reduced to two; Doiuchi et al., 2004: characters 13, 16 and 25, respectively). Molecular analyses with broader sampling of outgroup taxa offer more disparate hypotheses for the tetragonurid interrelationships. In the studies by Miya et al. (2013) and Friedman et al. (2019), Tetragonuridae is sister to the nonstromateiform Chiasmodontidae, and in the study by Campbell et al. (2018) it is grouped with Amarsipidae in a clade that is closer to scombrids (Scombriformes) than to the remaining stromateiforms., Published as part of Pastana, Murilo N L, Johnson, G David & Datovo, Aléssio, 2022, Comprehensive phenotypic phylogenetic analysis supports the monophyly of stromateiform fishes (Teleostei: Percomorphacea), pp. 841-963 in Zoological Journal of the Linnean Society 195 (3) on pages 949-951, DOI: 10.1093/zoolinnean/zlab058, http://zenodo.org/record/6758614, {"references":["Haedrich RL, Horn MH. 1972. A key to the stromateoid fishes (technical report WHOI- 72 - 15). Woods Hole, MA: Woods Hole Oceanographic Institution, 1 - 46.","Fricke R, Eschmeyer WN, Van der Laan R. 2020. Eschmeyer's catalog of fishes: genera, species, references. Available at: http: // researcharchive. calacademy. org / research / ichthyology / catalog / fishcatmain. asp","Bone Q, Brook CER. 1973. On Schedophilus medusophagus (Pisces: Stromateoidei). Journal of the Marine Biological Association of the United Kingdom 53: 753 - 761.","Janssen J, Harbison GR. 1981. Fish in salps: the association of squaretails (Tetragonurus spp.) with pelagic tunicates. Journal of the Marine Biological Association of the United Kingdom 61: 917 - 927.","Horn MH. 1984. Stromateoidei: development and relationships. In: Moser HG, Richards WJ, Cohen DM, Fahay MP, Kendall AW Jr, Richardson SL, eds. Ontogeny and systematics of fishes. Based on an International Symposium dedicated to the memory of Elbert Halvor Ahlstrom. New York: American Society of Ichthyologists and Herpetologists, 620 - 636.","Doiuchi R, Sato T, Nakabo T. 2004. Phylogenetic relationships of the stromateoid fishes (Perciformes). Ichthyological Research 51: 202 - 212.","Doiuchi R, Nakabo T. 2006. Molecular phylogeny of the stromateoid fishes (Teleostei: Perciformes) inferred from mitochondrial DNA sequences and compared with morphology-based hypotheses. Molecular Phylogenetics and Evolution 39: 111 - 123.","Miya M, Friedman M, Satoh TP, Takeshima H, Sado T, Iwasaki W, Yamanoue Y, Nakatani M, Mabuchi K, Inoue JG, Poulsen JY. 2013. Evolutionary origin of the Scombridae (tunas and mackerels): members of a Paleogene adaptive radiation with 14 other pelagic fish families. PLoS One 8: e 73535.","Friedman M, Feilich KL, Beckett HT, Alfaro ME, Faircloth BC, Cerny D, Miya M, Near TJ, Harrington RC. 2019. A phylogenomic framework for pelagiarian fishes (Acanthomorpha: Percomorpha) highlights mosaic radiation in the open ocean. Proceedings of the Royal Society B: Biological Sciences 286: 20191502.","Campbell MA, Sado T, Shinzato C, Koyanagi R, Okamoto M, Miya M. 2018. Multilocus phylogenetic analysis of the first molecular data from the rare and monotypic Amarsipidae places the family within the Pelagia and highlights limitations of existing data sets in resolving pelagian interrelationships. Molecular Phylogenetics and Evolution 124: 172 - 180."]}

Node 91 = Ariommatidae Ariomma indicum, Ariomma bondi and Ariomma melana. Unambiguous synapomorphies: Character 3 (17– 19> 20): number of branched pectoral-fin rays increased to 20; character 9 (33–36> 31): number of vertebrae decreased to 31; character 22 (1> 0): subocular shelf present; character 76 (1> 2): number of autogenous dorsal hypural plates decreased to one; character 78 (0> 1): ventral hypural plate fused to caudal centrum; character 117 (1> 0): adductor hyomandibulae insertion not advancing onto the anterior portion of the endopterygoid; character 125 (0> 1): retractor dorsalis vestigial; character 161 (0> 1): pharyngeal-sac papillae rakers on ventral portion of pharyngeal sac absent; character 169 (0> 1): dorsal portion of pharyngeal sac expanded anteriorly; character 170 (0> 1): accessory pharyngeal-sac muscular attachment to the cleithrum present; character 191 (1> 0): dorsal surface of the head scaled; character 194 (1> 0): dorsal-fin interradial membrane not scaled; character 197 (1> 0): anal-fin interradial membrane not scaled. Support: Relative Bremer = 78%. Remarks: The family Ariommatidae currently comprises the genus Ariomma, with seven valid species (Fricke et al., 2020). Monophyly of ariommatids is strongly supported herein by 13 synapomorphies (Fig. 70), including several characteristics unique to the family, such as a parhypural–hypural fusion (character 82; Fig. 46), pharyngeal-sac teeth absent on the ventral surface of the sac (character 161: Figs 36, 58B), anterodorsal expansion of the pharyngeal sac over the posteriormost gill-arch muscles and bones (character 141; Fig. 54A), an accessory muscular attachment of the sphincter oesophagi that connects the pharyngeal sac to the cleithrum (character 142) and a reduced/vestigial retractor dorsalis (character 148; Fig. 54A). The monophyly of Ariommatidae has been recovered by all previous analyses, those based on both morphological (Doiuchi et al., 2004) and molecular data (Doiuchi & Nakabo, 2006; Miya et al., 2013; Campbell et al., 2018; Friedman et al., 2019). Haedrich (1967) grouped ariommatids based on the presence of pelvic fins, separation of the dorsal fins, absence of teeth on the basihyal and basibranchials, six branchiostegals, fusion of theparhypuralandhypurals, andossifiedscleroticossicles (our characters 56, 66, 45, 12, 78 and 23, respectively). In addition, he listed the shape and distribution of the pharyngeal-sac papillae as diagnostic for the family (our characters 161 and 164). From these characters, only the parhypural–hypural fusion and the distribution of papillae on the pharyngeal sac were confirmed herein as valid ariommatid synapomorphies. Horn (1984) also used the shape of these papillae as diagnostic, together with the head squamation (scales on the opercle and preopercle) and number of hypurals. In our analysis, only one of Horn’s characters, the number of hypurals, is optimized as a valid ariommatid synapomorphy (coded herein in two independent characters: number of autogenous dorsal hypural plates, character 76; and hypural–caudal centrum fusion, character 78). The remaining characters are either plesiomorphies present in Stromateiformes (e.g. presence of pelvic fin, ossification of the sclerotic bones) or synapomorphies for larger clades including Ariommatidae (e.g. pharyngeal-sac papillae with round bases, shared with Tetragonuridae). The Ariommatidae clade of Doiuchi et al. (2004) was also represented as a long branch with ten autapomorphies. We have included five of the characters from Doiuchi et al. (2004) in our matrix (our characters 22, 36, 74, 117 and 192) and confirmed characters 22 and 117 as synapomorphies for Ariommatidae (presence of a subocular shelf,and adductor hyomandibulae not expanded onto the endopterygoid, respectively; Fig. 39). The remainder of the characters from Doiuchi et al. (2004) exhibited overlapping states when analysed over a broader taxon sampling and, consequently, were excluded from our dataset.

Suborder Amarsipoidei nom. nov. (new name containing Amarsipidae) Amarsipus carlsbergi. Autapomorphies: Character 0 (8–9> 11): number of dorsal-fin spines increased to 11; character 9 (31–42> 47): number of vertebrae increased to 47; character 11 (10–12> 13): number of ventral procurrent rays increased to 13; character 12 (7> 6): number of branchiostegals decreased to six; character 14 (49.6– 55.1%> 47.6%): dorsal-fin base length decreased to 47.6% of SL; character 16 (27.6–36.6%> 21.8%): prepelvic length decreased to 21.8% of SL; character 17 (53.6–54.7%> 45.9%): preanal length decreased to 45.9% of SL; character 30 (1> 0): medial bony suture between anterior and posterior ceratohyals absent; character 66 (0> 1): dorsal-fin anterior and posterior portions separated; character 82 (0> 1): epineural 4 to last originating from the ribs; character 85 (0> 1): first supraneural located in the first interneural space; character 86 (1> 2): second supraneural located in the second interneural space; character 87 (2> 3): third supraneural located in the third interneural space; character 97 (1> 0): adductor mandibulae partes rictalis and malaris completely separated along their entire extent; character 154 (1> 0): adductor medialis absent; character 198 (1> 0): interradial membrane of the caudal fin not scaled. Support: –. Remarks: Amarsipidae is a monotypic family containing the sole species Amarsipus carlsbergi, recovered herein as the sister group of the remaining Stromateiformes in a long terminal branch supported by 16 autapomorphies (Fig. 70). Haedrich (1969) diagnosed Amarsipus carlsbergi from the remaining Stromateiformes based on the combination of anteriorly placed pelvic fins, toothed vomer, six hypurals (erroneous, see Remarks on node 90), two epurals and absence of a pharyngeal sac. These characters are comparable with our characters 16, 41, 76 and 77, 79 and 157, respectively. When interpreted phylogenetically, only character 16 (anteriorly placed pelvic fins) represents an autapomorphy for Amarsipidae, while the remaining characters are plesiomorphies retained by this taxon. Haedrich (1969) suggested that Amarsipidae might warrant subordinal status, but refrained from doing that because it could mask its relationship with the remaining Stromateiformes, which at that time were classified as a suborder within the nonmonophyletic Perciformes. However, given that current classifications have elevated Stromateiformes to an ordinal level (Wiley & Johnson, 2010), the formal ranking of Amarsipoidei as the sister group of the Stromateoidei (node 100) is an appropriate taxonomic adjustment implemented herein. Node 100 = Stromateoidei (new usage for non-Amarsipidae Stromateiformes) Icichthys lockingtoni, Tubbia tasmanica, Centrolophus niger, Hyperoglyphe perciformis, Psenopsis cyanea, Psenopsis anomala, Seriolella porosa, Schedophilus sp., Psenes sio, Psenes cyanophrys, Nomeus gronovii, Cubiceps whiteleggii, Cubiceps pauciradiatus, Tetragonurus cuvieri, Ariomma indicum, Ariomma bondi, Ariomma melana, Stromateus brasiliensis, Pampus cinereus, Peprilus triacanthus, Peprilus paru. Unambiguous synapomorphies: Character 3 (16> 18): number of branched pectoral-fin rays increased to 18; character 41 (0> 1): vomerine teeth present; character 49 (0> 1): fourth upper pharyngeal tooth plate considerably longer than wide; character 108 (0> 1): orbitopectoral branch of the ramus lateralis accessorius (RLA-OP) lateral to the levator operculi; character 117 (0> 1): adductor hyomandibulae insertion advancing onto anterior portion of the endopterygoid; character 124 (0> 1): sphincter oesophagi pars anterior absent; character 129 (1> 0): dorsalis portion of the rectus communis associated with hypobranchial 3; character 130 (0> 1): fibres of the rectus communis disposed between hypobranchial 3 and urohyal; character 136 (0> 1): medial contact between antimeres of pharyngoclavicularis internus present; character 146 (0> 1): anterior border of epaxialis anterolateral section reaching or trespassing the vertical line through the middle of the orbit; character 157 (0> 1): pharyngeal sac present; character 194 (0> 1): interradial membrane of the dorsal fin scaled; character 197 (0> 1) interradial membrane of the anal fin scaled. Support: Relative Bremer = 64%. Remarks: The suborder Stromateoidei is herein proposed to encompass all non-amarsipid Stromateiformes, namely the families Centrolophidae, Nomeidae, Tetragonuridae, Ariommatidae and Stromateidae. The monophyly of this clade is supported by 13 morphological synapomorphies and obtained under all searching parameters used in this study. Among the characters supporting node 100, the presence of the pharyngeal sac is obviously the most remarkable synapomorphy of Stromateoidei, because this complex organ is, to our knowledge, unparalleled among vertebrates (Figs 36–41, 52–58). Accordingly, a natural assemblage comprising the pharyngeal sacbearing stromateiform fishes has long been recognized by morphological studies (e.g. Regan, 1902; Gilchrist, 1922; Bühler, 1930; Barnard 1948; Isokawa et al., 1965; Haedrich, 1967; Horn, 1984; Datovo et al., 2014). Our study aside, a Stromateoidei clade has been recovered within an explicit phylogenetic context only by Horn (1984), wherein it was supported by two synapomorphies: the presence of the pharyngeal sac (our character 53) and the juvenile association with floating objects (our character 206). Of these two characters, only the presence of a pharyngeal sac is corroborated as a synapomorphy for the Stromateoidei. The association between juvenile fishes and gelatinous organisms is broader than reported by Horn (1984), and it has been reported for juvenile amarsipids (Janssen & Harbison, 1981; Harbison, 1993) and for some other non-stromateiform percomorphs (Fig. 65; Supporting Information, Supplementary File S1: Table S3). A character reconstruction of this behaviour (character 206) indicates that it could either be optimized as a synapomorphy for Stromateiformes (node 90; DelTran, i.e. delayed transformation), with parallel acquisitions in bramids, caristiids, carangids and icosteids) or be a synapomorphy for a larger clade (node 76; AccTran, i.e. accelerated transformation), with reversals occurring within carangiforms, scombriforms and other taxa lacking juvenile association with pelagic gelatinous invertebrates. Another synapomorphy for Stromateoidei is the shared pattern 10 of the RLA, in which its orbitopectoral branch overlies the levator arcus palatini, dilatator operculi and levator operculi muscles (Figs 27, 50, 51; see character 108). The sharing of this innervation pattern has previously been posited as evidence for a possible relationship between stromateoids and the Kyphosidae s.l. (including Scorpididae, Girellidae and Microcanthidae), Oplegnathidae, Kuhliidae, Arripidae, Terapontidae (Freihofer, 1963; Johnson & Fritzsche, 1989; Springer & Johnson, 2004) and Dichistiidae (Leis & Lingen, 1997). Yet, our analysis contrasts with these hypotheses by not supporting the monophyly of an RLA-10-pattern fish assemblage. Our topology indicates that this branching pattern has arisen at least four times independently, in: (1) Pomatomidae (Fig. 67, optimized as an autapomorphy); (2) Arripidae (Fig. 67, also optimized as an autapomorphy); (3) a clade encompassing Kyphosidae s.l., Kuhliidae, Terapontidae, Oplegnathidae and Haemulidae, with reversals in Scorpis chilensis and Orthopristis ruber (Fig. 67: node 114, AccTran); and (4) Stromateoidei (Fig. 69: node 100, with reversal in Tubbia, Psenes and Tetragonurus). Node 100 = Stromateoidei (new usage for non-Amarsipidae Stromateiformes) Icichthys lockingtoni, Tubbia tasmanica, Centrolophus niger, Hyperoglyphe perciformis, Psenopsis cyanea, Psenopsis anomala, Seriolella porosa, Schedophilus sp., Psenes sio, Psenes cyanophrys, Nomeus gronovii, Cubiceps whiteleggii, Cubiceps pauciradiatus, Tetragonurus cuvieri, Ariomma indicum, Ariomma bondi, Ariomma melana, Stromateus brasiliensis, Pampus cinereus, Peprilus triacanthus, Peprilus paru. Unambiguous synapomorphies: Character 3 (16> 18): number of branched pectoral-fin rays increased to 18; character 41 (0> 1): vomerine teeth present; character 49 (0> 1): fourth upper pharyngeal tooth plate considerably longer than wide; character 108 (0> 1): orbitopectoral branch of the ramus lateralis accessorius (RLA-OP) lateral to the levator operculi; character 117 (0> 1): adductor hyomandibulae insertion advancing onto anterior portion of the endopterygoid; character 124 (0> 1): sphincter oesophagi pars anterior absent; character 129 (1> 0): dorsalis portion of the rectus communis associated with hypobranchial 3; character 130 (0> 1): fibres of the rectus communis disposed between hypobranchial 3 and urohyal; character 136 (0> 1): medial contact between antimeres of pharyngoclavicularis internus present; character 146 (0> 1): anterior border of epaxialis anterolateral section reaching or trespassing the vertical line through the middle of the orbit; character 157 (0> 1): pharyngeal sac present; character 194 (0> 1): interradial membrane of the dorsal fin scaled; character 197 (0> 1) interradial membrane of the anal fin scaled. Support: Relative Bremer = 64%. Remarks: The suborder Stromateoidei is herein proposed to encompass all non-amarsipid Stromateiformes, namely the families Centrolophidae, Nomeidae, Tetragonuridae, Ariommatidae and Stromateidae. The monophyly of this clade is supported by 13 morphological synapomorphies and obtained under all searching parameters used in this study. Among the characters supporting node 100, the presence of the pharyngeal sac is obviously the most remarkable synapomorphy of Stromateoidei, because this complex organ is, to our knowledge, unparalleled among vertebrates (Figs 36–41, 52–58). Accordingly, a natural assemblage comprising the pharyngeal sacbearing stromateiform fishes has long been recognized by morphological studies (e.g. Regan, 1902; Gilchrist, 1922; Bühler, 1930; Barnard 1948; Isokawa et al., 1965; Haedrich, 1967; Horn, 1984; Datovo et al., 2014). Our study aside, a Stromateoidei clade has been recovered within an explicit phylogenetic context only by Horn (1984), wherein it was supported by two synapomorphies: the presence of the pharyngeal sac (our character 53) and the juvenile association with floating objects (our character 206). Of these two characters, only the presence of a pharyngeal sac is corroborated as a synapomorphy for the Stromateoidei. The association between juvenile fishes and gelatinous organisms is broader than reported by Horn (1984), and it has been reported for juvenile amarsipids (Janssen & Harbison, 1981; Harbison, 1993) and for some other non-stromateiform percomorphs (Fig. 65; Supporting Information, Supplementary File S1: Table S3). A character reconstruction of this behaviour (character 206) indicates that it could either be optimized as a synapomorphy for Stromateiformes (node 90; DelTran, i.e. delayed transformation), with parallel acquisitions in bramids, caristiids, carangids and icosteids) or be a synapomorphy for a larger clade (node 76; AccTran, i.e. accelerated transformation), with reversals occurring within carangiforms, scombriforms and other taxa lacking juvenile association with pelagic gelatinous invertebrates. Another synapomorphy for Stromateoidei is the shared pattern 10 of the RLA, in which its orbitopectoral branch overlies the levator arcus palatini, dilatator operculi and levator operculi muscles (Figs 27, 50, 51; see character 108). The sharing of this innervation pattern has previously been posited as evidence for a possible relationship between stromateoids and the Kyphosidae s.l. (including Scorpididae, Girellidae and Microcanthidae), Oplegnathidae, Kuhliidae, Arripidae, Terapontidae (Freihofer, 1963; Johnson & Fritzsche, 1989; Springer & Johnson, 2004) and Dichistiidae (Leis & Lingen, 1997). Yet, our analysis contrasts with these hypotheses by not supporting the monophyly of an RLA-10-pattern fish assemblage. Our topology indicates that this branching pattern has arisen at least four times independently, in: (1) Pomatomidae (Fig. 67, optimized as an autapomorphy); (2) Arripidae (Fig. 67, also optimized as an autapomorphy); (3) a clade encompassing Kyphosidae s.l., Kuhliidae, Terapontidae, Oplegnathidae and Haemulidae, with reversals in Scorpis chilensis and Orthopristis ruber (Fig. 67: node 114, AccTran); and (4) Stromateoidei (Fig. 69: node 100, with reversal in Tubbia, Psenes and Tetragonurus)., Published as part of Pastana, Murilo N L, Johnson, G David & Datovo, Aléssio, 2022, Comprehensive phenotypic phylogenetic analysis supports the monophyly of stromateiform fishes (Teleostei: Percomorphacea), pp. 841-963 in Zoological Journal of the Linnean Society 195 (3) on pages 941-942, DOI: 10.1093/zoolinnean/zlab058, http://zenodo.org/record/6758614, {"references":["Haedrich RL. 1969. A new family of aberrant stromateoid fishes from the equatorial Indo-Pacific. Dana Reports 76: 1 - 14.","Wiley EO, Johnson GD. 2010. A Teleost classification based on monophyletic groups. In: Nelson JS, Schultze H-P, Wilson MVH, eds. Origin and phylogenetic interrelationships of teleosts. Munich: Dr Friedrich Pfeil, 123 - 182.","Regan CT. 1902. A revision of the fishes of the family Stromateidae. Annals and Magazine of Natural History 10: 115 - 131.","Gilchrist JDF. 1922. On the oesophagal teeth of the Stromateidae. Annals and Magazine of Natural History 9: 249 - 255.","Buhler H. 1930. Die Verdauungsorgane der Stromateidae (Teleost.). Zoomorphology 19: 59 - 115.","Barnard KH. 1948. Further notes on South African marine fishes. Annals of the South African Museum 36: 341 - 406.","Isokawa S, Kubota K, Kosaikai T, Satomura I, Tsubouchi M, Sera A. 1965. Some contributions to study of esophageal sacs and teeth of fishes. Journal of Nihon University School of Dentistry 7: 103 - 111.","Haedrich RL. 1967. The stromateoid fishes: systematics and a classification. Bulletin of the Museum of Comparative Zoology 135: 310 - 139.","Horn MH. 1984. Stromateoidei: development and relationships. In: Moser HG, Richards WJ, Cohen DM, Fahay MP, Kendall AW Jr, Richardson SL, eds. Ontogeny and systematics of fishes. Based on an International Symposium dedicated to the memory of Elbert Halvor Ahlstrom. New York: American Society of Ichthyologists and Herpetologists, 620 - 636.","Datovo A, de Pinna MCC, Johnson GD. 2014. The infrabranchial musculature and its bearing on the phylogeny of percomorph fishes (Osteichthyes: Teleostei). PLoS One 9: e 110129.","Janssen J, Harbison GR. 1981. Fish in salps: the association of squaretails (Tetragonurus spp.) with pelagic tunicates. Journal of the Marine Biological Association of the United Kingdom 61: 917 - 927.","Harbison GR. 1993. The potential of fishes for the control of gelatinous zooplankton. International Council for the Exploration of the Sea (ICES) 74: 1 - 10.","Freihofer WC. 1963. Patterns of ramus lateralis accessorius and their systematic significance in teleostean fishes. Stanford Ichthyological Bulletin 8: 80 - 189.","Johnson GD, Fritzsche RA. 1989. Graus nigra, an omnivorous girellid, with a comparative osteology and comments on relationships of the Girellidae (Pisces: Perciformes). Proceedings of the Academy of Natural Sciences of Philadelphia 141: 1 - 27.","Springer VG, Johnson GD. 2004. Study of the dorsal gillarch musculature of Teleostome fishes, with special reference to the Actinopterygii. Bulletin of the Biological Society of Washington 11: 1 - 205.","Leis JM, Van der Lingen CD. 1997. Larval development and relationships of the perciform family Dichistiidae (= Coracinidae), the galjoen fishes. Bulletin of Marine Sciences 60: 100 - 116."]}

Datovo, Aléssio (2014): A new species of Ituglanis from the Rio Xingu basin, Brazil, and the evolution of pelvic fin loss in trichomycterid catfishes (Teleostei: Siluriformes: Trichomycteridae). Zootaxa 3790 (3): 466-476, DOI: http://dx.doi.org/10.11646/zootaxa.3790.3.5

Datovo, Aléssio, Vari, Richard P. (2014): The adductor mandibulae muscle complex in lower teleostean fishes (Osteichthyes: Actinopterygii): comparative anatomy, synonymy, and phylogenetic implications. Zoological Journal of the Linnean Society 171 (3): 554-622, DOI: 10.1111/zoj.12142, URL: http://dx.doi.org/10.1111/zoj.12142

Trichomycterus perkos, new species (Figs. 1–2, Table 1) Trichomycterus aff. itatiayae: Kantek et al. (2007: 797). Trichomycterus sp. aff. T. itatiayae: Torres et al. (2004: 124). Trichomycterus sp. aff. T. itatiyae [sic]: Sato et al. (2004: 45, 46, 47, Table 1, Table 2). Holotype. MCP 46679 (93.5 mm SL), XR; Brazil, Rio Grande do Sul State, Passo Fundo Municipality; Uruguai River basin, Passo Fundo River, under the bridge on BR– 285 road; approx. 28 ° 14 '07"S 52 ° 24 ' 53 "W; W. Bruschi Jr. & J. F. P. Silva; 0 3 January 2003. Paratypes. LIRP 8344, 4 (28.0–93.2 mm SL): 1 CS (28.0 mm SL), 1 MS (35.3 mm SL), 1 XR (28.3 mm SL); Brazil, Rio Grande do Sul State, Passo Fundo Municipality; Uruguai River basin, unnamed stream tributary of Passo Fundo River, about 1 km from BR– 285 road; approx. 28 ° 15 ' 14 "S 52 ° 21 ' 19 "W; W. Bruschi Jr. & J. F. P. Silva; 0 3 January 2003. MCP 31763, 10 (16.2–99.8 mm SL): 1 CS (52.4 mm SL), 1 XR (99.81 mm SL); same data as LIRP 8344. MCP 31764, 2 (17.9–33.1 mm SL; XR); same data as holotype. MCP 31774, 1 (61.2 mm SL); same data as LIRP 8344. MCP 31776, 3 (27.0–43.3 mm SL); Brazil, Rio Grande do Sul State, Passo Fundo Municipality; Uruguai River basin, Passo Fundo River; 28 ° 14 ' 44 "S 52 ° 20 ' 53 "W; W. Bruschi Jr. & J. F. P. Silva; 0 4 January 2003. MCP 46701, 1 (48.9 mm SL); Brazil, Rio Grande do Sul State, São Valentim Municipality; Uruguai River basin, Passo Fundo River basin, unnamed stream tributary of Erechim River; 27 ° 38 ' 29 "S 52 ° 35 ' 51 "W; J. F. P. Silva; 20 October 2011. MCP 46711, 1 (69.1 mm SL); Brazil, Rio Grande do Sul State, Campinas do Sul Municipality; Uruguai River basin, unnamed stream tributary of Sepultura Stream, an affluent of reservoir of the Passo Fundo Hydroelectric Power Plant; 27 ° 48 ' 56 "S 52 ° 37 ' 51 "W; J. F. P. Silva; 20 October 2011. MCP 46715, 1 (32.9 mm SL); Brazil, Rio Grande do Sul State, Campinas do Sul Municipality; Uruguai River basin, unnamed stream affluent of reservoir of the Passo Fundo Hydroelectric Power Plant; 27 ° 51 ' 18 "S 52 ° 35 ' 16 "W; J. F. P. Silva; 20 October 2011. MCP 46718, 1 (71.4 mm SL); Brazil, Rio Grande do Sul State, Campinas do Sul Municipality; Uruguai River basin, unnamed stream tributary of Cipó Stream, an affluent of reservoir of the Passo Fundo Hydroelectric Power Plant; 27 ° 52 ' 24 "S 52 ° 33 ' 35 "W; J. F. P. Silva; 20 October 2011. MCP 46721, 1 (50.2 mm SL); Brazil, Rio Grande do Sul State, Campinas do Sul Municipality; Uruguai River basin, Cipó Stream, an affluent of reservoir of the Passo Fundo Hydroelectric Power Plant; 27 ° 52 ' 27 "S 52 ° 33 ' 25 "W; J. F. P. Silva; 20 October 2011. MZUSP 82372, 1 (72.2 mm SL); Brazil, Paraná State, Castrolanda Municipality; Paranapanema River basin; Tibaji River sub-basin, Iapó River sub-basin, unnamed stream tributary of da Onça River; approx. 24 ° 49 ' 30 "S 49 ° 54 ' 28 "W; A. L. A. Alves, R. Devid, A. Ferreira, & Y. R. Arvex; 20 March 2002; UFRGS 5922, 1 (41.1 mm SL), XR; Brazil, Rio Grande do Sul State, Condor Municipality; Uruguai River basin, Ijuí River sub-basin, Alegre River sub-basin, Palmeiras River; approx. 28 ° 14 '09"S 53 ° 28 '01"W; J. A. Anza & J. F. P. Silva; 0 2 November 2002. Diagnosis. Large-sized specimens (more than 65.8 mm SL), presumably adults, of Trichomycterus perkos are distinguished from all their congeners in having a distinctive coloration with melanophores arranged in two distinct skin layers, forming (1) a freckled pattern, with minute light brown spots scattered on the superficial tegumentar layer of the dorsum and caudal peduncle, and (2) a striped pattern, with three – sagittal, midlateral, and ventrolateral – wide, irregularly bordered, and sometimes interrupted dark brown stripes running along an inner skin layer of the trunk and caudal peduncle (Fig. 1; vs. melanophores in only one skin layer in most other Trichomycterus or melanophores occurring in two layers, but with the inner layer forming blotches in T. brasiliensis Lütken, T. castroi de Pinna, T. crassicaudatus Wosiacki and de Pinna, T. diabolus Bockmann, Casatti, and de Pinna, T. igobi Wosiacki and de Pinna, T. maracaya Bockmann and Sazima, T. mimonha Costa, T. stawiarski (P. Miranda Ribeiro), and T. tropeiro Ferrer and Malabarba, or forming solely a midlateral stripe in T. giganteus Lima and Costa; pers. obs.; Bockmann et al., 2004; Bockmann & Sazima, 2004; de Pinna, 1992 b; Lima & Costa, 2004; Wosiacki & de Pinna, 2008 a, 2008 b). Smaller individuals (less than 43.3 mm SL), presumably juveniles, of Trichomycterus perkos lack the superficial freckled pattern, but already exhibit the three wide dark stripes found in the adults (Fig. 2). This juvenile color pattern, although not identical, may resemble that of some T. duellmani Arratia and Menu-Marque, T. itatiayae A. Miranda Ribeiro, T. nigroauratus Barbosa and Costa, T. pauciradiatus Alencar and Costa, T. reinhardti (Eigenmann), and T. taenia Kner. Nevertheless, both juveniles and adults of T. perkos unequivocally differ from these congeners in a number of morphological traits: (1) from T. duellmani in having modally seven pectoral-fin rays (vs. modally eight), two pores in the lateral line (vs. four), distal margin of the adpressed pelvic fin not reaching the anus (vs. distal margin extending posteriorly beyond the anus), modally ten branchiostegal rays (vs. six or seven), and modally 41 post-Weberian vertebrae (vs. 33–36) (Arratia & Menu-Marque, 1984); (2) from T. itatiayae by the first pectoral-fin ray not filamentous (vs. filamentous), distal margin of the adpressed pelvic fin not reaching the anus (vs. distal margin extending posteriorly beyond the anus), modally ten branchiostegal rays (vs. eight), and modally 41 post-Weberian vertebrae (vs. 35–37) (Barbosa & Costa, 2008; Caramaschi & Caramaschi, 1991; A. Miranda Ribeiro, 1906); (3) from T. nigroauratus by the first pectoral-fin ray not filamentous (vs. filamentous), modally seven pectoral-fin rays (vs. modally eight), modally ten branchiostegal rays (vs. seven or eight), and modally 41 post-Weberian vertebrae (vs. 35–36) (Barbosa & Costa, 2008); (4) from T. pauciradiatus in having the first pectoral-fin ray not filamentous (vs. filamentous), modally seven pectoral-fin rays (vs. modally six), five pelvic-fin rays (vs. four), distal margin of the adpressed pelvic fin not reaching the anus (vs. distal margin extending posteriorly beyond the anus), pores i 1 and i 3 of the infraorbital laterosensory canal lacking (vs. i 1 and i 3 present), modally ten branchiostegal rays (vs. eight), and modally 41 post-Weberian vertebrae (vs. 36–38) (Alencar & Costa, 2006); (5) from T. reinhardti by the first pectoral-fin ray not prolonged as a filament (vs. ray filamentous), distal margin of the adpressed pelvic fin not reaching the anus (vs. distal margin extending posteriorly beyond the anus), absence of the pores i 1 and i 3 of the infraorbital laterosensory canal (vs. i 1 and i 3 present), modally ten branchiostegal rays (vs. seven or eight), and modally 41 post-Weberian vertebrae (vs. 38) (Eigenmann, 1917, 1918; Morris et al., 2006); (6) from T. taenia by the first pectoral-fin ray not filamentous (vs. filamentous), and distal margin of the adpressed pelvic fin not reaching the anus (vs. distal margin reaching the anus) (Eigenmann, 1918). Description. Morphometric data for type series given in Table 1. Refer to Figs. 1–2 for general external aspects. Body elongate. Dorsal and ventral profiles of trunk ranging from straight to slightly convex; dorsal and ventral profiles of caudal peduncle from straight to slightly concave. Cross section of trunk nearly oval, becoming gradually more compressed posterior to pectoral girdle. Head wide, depressed, trapezoidal in dorsal view. Larger specimens usually with region lateral to eyes swollen by hypertrophied adductor mandibulae A 2 A 3 muscles. Dorsal profile of head straight; ventral profile ranging from straight to convex. Eyes on dorsolateral region of head, progressively migrating towards more dorsal position with specimen growth. Orbital rim not free. Thin and translucent skin covering eye, not adhered to eyeball surface and forming anteroposteriorly elongated ocular capsule. Anterior nostril surrounded by tubular flap continuous with nasal barbel base; posterior nostril opening slightly smaller than anterior one and with crescent thin flap on anterior border. Mouth subterminal and slightly curved. Lower lip with lateral fleshy lobes posteromedial to rictal-barbel base. Nasal barbel emerging from lateral region of anterior nostril and reaching center of neurocranium. Maxillary and rictal barbels about same size and usually reaching anterior portion of interopercular patch of odontodes. Branchial membranes thick, united to isthmus only anteriorly and forming small free fold across isthmus. Branchiostegal rays modally ten (one among five CS, MS, and XR specimens with nine); medial most rays hardly visualized through skin. Opercular patch of odontodes rounded and dorsolaterally placed on head; 8–14 conical odontodes. Interopercular patch of odontodes narrow, posteriorly curved, and placed anterior to opercular patch; 11–17 conical odontodes. Odontodes progressively larger and more curved towards posterior region of both opercular and interopercular patches. Pectoral-fin rays modally I+ 6 (four among 27 specimens with I+ 5); first ray not prolonged as a pectoral filament. Pectoral-fin posterior margin convex. Anterior portion of pectoral-fin base covered by branchial membrane. Axillary pore present. Pelvic-fin rays I+ 4, thin pelvic splint parallel to first pelvic ray. Pelvic-fin origin anterior to origin of dorsal fin; posterior margin convex. Bases of pelvic fins contacting to each other. Distal margin of adpressed pelvic fin not reaching urogenital and anal openings. Dorsal-fin rays i–ii+II+ 7; eight basal radials distributed between neural spines of 20 th– 21 st and 25 th– 26 th post- Weberian vertebrae. Dorsal fin located on posterior half of trunk; origin approximately at vertical through posterior margin of adpressed pelvic fin; posterior margin convex. Anal-fin rays i–iii+II+ 5; six basal radials distributed between haemal spines of 22 nd– 23 rd and 26 th– 27 th post- Weberian vertebrae. Anal-fin origin slightly posterior to vertical through dorsal-fin origin; posterior margin convex. Caudal-fin rays xiv–xvii+I+ 5 on dorsal lobe and xii–xiv+I+ 6 on ventral lobe. Caudal-fin posterior margin slightly convex with rounded dorsal and ventral corners. Two upper hypural plates, presumably hypural 3 (ventral) and compound hypural 4 + 5 (dorsal); single lower hypural plate (compound hypural 1 + 2) fused to parhypural (de Pinna & Ng, 2004; Lundberg & Baskin, 1969). First complete haemal arch on seventh–ninth post-Weberian vertebrae; first complete haemal spine on 15 th– 16 th post-Weberian vertebrae. Post-Weberian vertebrae modally 41 (one specimen with 39 and another with 40 among eight CS and XR specimens). Ribs 12–14. Laterosensory cephalic canals with simple (non-dendritic) tubes ending in single pores. Supraorbital canal mostly into frontal bone with pores s 1, s 3 and s 6. Infraorbital canal mostly into soft tissue ventroposterior to eyeball and with branches and pores i 10 and i 11. Otic canal without pores and running through sphenotic-prootic-pterosphenoid. Postotic canal traversing pterotic and posttemporo-supracleithrum, with pores po 1 (= preoperculomandibular of Schaefer & Aquino, 2000) and po 2 (= pterotic of Schaefer & Aquino, 2000) located anterodorsal to opercular patch of odontodes. Short lateral line canal with pores ll 1 and ll 2 positioned dorsoposterior to pectoral-fin base. Coloration in alcohol. Unpigmented body background pale yellow (Figs. 1, 2). Dark brown melanophores located on inner skin layer and forming three wide body stripes: sagittal, midlateral, and ventrolateral. Sagittal stripe unpaired; midlateral and ventrolateral stripes bilaterally paired. All stripes with notched borders and usually interrupted at some points, thereby occasionally forming few isolated irregular blotches. Stripes borders gradually more notched in larger specimens. Sagittal and midlateral stripes running along entire lengths of trunk and caudal peduncle; sagittal stripe about two times wider than midlateral one. Ventrolateral stripe thinnest and running along posterior part of trunk and caudal peduncle; ventrolateral stripe most often interrupted, sometimes taking form of row of irregular blotches. Dark brown pigmentation from inner tegumentar layer forming highly irregular marks on dorsolateral region of head and small spots on barbel bases. Smallest individuals lacking dark pigmentation on all fins. Median fins and dorsal surface of pectoral fin progressively acquiring increasing number of small spots with specimen growth. Spots on fins also dark brown and located on inner tegumentar layer. Larger individuals (more than 65.8 mm SL), presumably adults, with caudal peduncle and dorsum covered with tiny light brown spots forming a freckled pattern (Fig. 1). Freckled pattern of larger individuals located on outer skin layer and covering part of dark stripes and blotches of inner tegumentar layer. Ventral surface of head and trunk devoid of dark pigmentation (Figs. 1, 2). Etymology. From the Greek perkos (περκς), meaning “spotted or streaked with black marks (Valpy, 1826: 227), in allusion to the color pattern of the new species formed by either dark stripes (small-sized specimens) or dark stripes combined with small spots (larger individuals). An adjective. Distribution. A single paratype (MZUSP 82372) is from the Tibaji River basin, a tributary of the Paranapanema River (Fig. 3; see Discussion below). All remaining type series is from two sub-basins of the Uruguai River system, the Ijuí and Passo Fundo drainages (Fig. 3). The Paranapanema is an affluent of the Paraná River, which joins the Uruguai to form the La Plata River/Estuary. Habitat notes. Ecological data from all localities is unavailable. Most types were collected in clear water streams with 1 to 3 m wide, with bottoms composed of stone, gravel, and sand (Fig. 4). Streamlets were usually surrounded by modified riparian vegetation.

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No. of bitstreams: 0 Previous issue date: 2016-11-23 Villanova Univ, Dept Biol, Villanova, PA 19085 USA Museu Nacl Hist Nat & Ciencia, Lisbon, Portugal Univ Brasilia, Dept Zool, Inst Ciencias Biol, BR-70910900 Brasilia, DF, Brazil Smithsonian Inst, Natl Museum Nat Hist, Washington, DC 20560 USA Sorbonne Univ, Museum Natl Hist Nat, ISYEB, Paris, France Univ Nacl Tucuman, San Miguel De Tucuman, Argentina King Saud Univ, Riyadh, Saudi Arabia Cornell Univ, Ithaca, NY USA Univ Fed Sao Paulo, Diadema, Brazil Upper Austrian Museum, Ctr Biol, Linz, Austria Villanova Univ, Villanova, PA 19085 USA Univ Salzburg, Salzburg, Austria Plazi, Bern, Switzerland Univ Brasilia, Brasilia, DF, Brazil Univ Sao Paulo, Ribeirao Preto, Brazil Zool Forsch Museum A Koenig, Bonn, Germany Museu Paraense Emilio Goeldi, Belem, Para, Brazil Russian Acad Sci, St Petersburg, Russia Pontificia Univ Catolica Rio Grande do Sul, Porto Alegre, RS, Brazil Museo Reg Sci Nat, Turin, Italy Univ Fed Rio Grande do Sul, Porto 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