Stromateidae

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), pp. 841-963 : 952

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.

Figures

Figure 37. Gill arches of Peprilus sp. (SML 34163). A, dorsal gill arches from left side in dorsal view. B, dorsal gill arches from left side in dorsal view.C, ventral view. Gill rakers omitted. Photographs taken by Miguel Montalvo. Scale bars: 0.5 mm.

Figure 58. Micro-computed tomography scans of pharyngeal-sac skeletons in left lateral view, whole (A, C) and sagittal (B, D) view. A, B, the ariommatid Ariomma bondi (MZUSP 86717). C, D, the stromateid Peprilus triacanthus (MZUSP 123240). Scale bar: 2 mm.

Figure 60. Stromateiform pharyngeal-sac rakers in dorsal (A, C, E) and lateral (B, D, F) view.A, B, the ariommatid Ariomma bondi (MZUSP 86717), showing round base and elongate axis. C, D, the stromateid Peprilus triacanthus (MZUSP 123240), showing stellate base and elongate axis. E, F, the nomeid Psenes cyanophrys (MZUSP 106392), showing stellate base and less elongate axis. Scale bars: 0.5 mm.

Figure 61. The muscular attachment of the pharyngeal-sac raker in Stromateiformes.Raker bases are visible after removal of the external most layer of the sphincter oesophagi. A, in the ariommatid Ariomma melana (MZUSP 123246), rakers are large, round and with partly overlapping bases, forming a scale-like pattern. B, in the tetragonurid Tetragonurus cuvieri (MZUSP 123241), rakers are small, round, poorly ossified and with non-overlapping bases. C, D, in both the nomeid Nomeus gronovii (MZUSP 67590; C) and the stromateid Stromateus brasiliensis (MZUSP 51279; D), the raker bases are large, well ossified and stellate. Scale bars: 0.5 mm.

Figure 62. Diagrammatic cross-sections of the pharyngeal sac of the ariommatid Ariomma indicum (A), the stromateid Stromateus fiatola (B), the centrolophid Centrolophus niger (C) and the nomeid Cubiceps capensis (D). The pointy projections emerging from the internal walls of the pharyngeal sac (A–D) represent the sac papillae, and arrows (in C, D) indicate the greatly enlarged, posteriorly directed, toothed gill rakers. In ariommatids (A) and stromateids (B), the internal lining of the sac is classified as papillary (or polypoid), whereas the presence of enlarged rakers entering the pharyngeal sac of centrolophids (C) and nomeids (D) gives the sac a folded (or wrinkled) aspect. The obliquely shaded oval structures (C, D) represent the elongated fourth upper pharyngeal tooth plates. They are not shown in ariommatids (A) and stromateids (B) because they extend only into the anterior half of the sac. Figure modified from Barnard (1948: fig. 10).

Figure 69. Reference tree obtained under implied-weights parsimony (k = 13.56554), with sensitivity analysis of alternative weighting schemes, highlighting the interrelationships of Stromateiformes (node 90). Squares representing each value of k are coloured according to their respective element-based scores. Numbers below sensitivity plots indicate the average element-based comparison score for the clade. Node numbers referenced in the text displayed above each clade.

Figure 70. Unambiguous phenotypic synapomorphies for Stromateiformes superimposed on the reference tree of the order obtained under implied-weights parsimony (k = 13.56554).