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. 2019 Oct 9;286(1912):20191336.
doi: 10.1098/rspb.2019.1336. Epub 2019 Oct 2.

The early elasmobranch Phoebodus: phylogenetic relationships, ecomorphology and a new time-scale for shark evolution

Affiliations

The early elasmobranch Phoebodus: phylogenetic relationships, ecomorphology and a new time-scale for shark evolution

Linda Frey et al. Proc Biol Sci. .

Abstract

Anatomical knowledge of early chondrichthyans and estimates of their phylogeny are improving, but many taxa are still known only from microremains. The nearly cosmopolitan and regionally abundant Devonian genus Phoebodus has long been known solely from isolated teeth and fin spines. Here, we report the first skeletal remains of Phoebodus from the Famennian (Late Devonian) of the Maïder region of Morocco, revealing an anguilliform body, specialized braincase, hyoid arch, elongate jaws and rostrum, complementing its characteristic dentition and ctenacanth fin spines preceding both dorsal fins. Several of these features corroborate a likely close relationship with the Carboniferous species Thrinacodus gracia, and phylogenetic analysis places both taxa securely as members of the elasmobranch stem lineage. Identified as such, phoebodont teeth provide a plausible marker for range extension of the elasmobranchs into the Middle Devonian, thus providing a new minimum date for the origin of the chondrichthyan crown-group. Among pre-Carboniferous jawed vertebrates, the anguilliform body shape of Phoebodus is unprecedented, and its specialized anatomy is, in several respects, most easily compared with the modern frilled shark Chlamydoselachus. These results add greatly to the morphological, and by implication ecological, disparity of the earliest elasmobranchs.

Keywords: Devonian; Morocco; chondrichthyes; gnathostomes; neurocranium.

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Conflict of interest statement

The authors declare no competing interests.

Figures

Figure 1.
Figure 1.
Phoebodus saidselachus sp. nov., (ad) PIMUZ A/I 4712 and (e) PIMUZA/I 4656. (a) Ferruginous nodule containing cranial and postcranial remains; (b) drawing, scale bar, 200 mm; (c) detail of visceral skeleton, scale bar, 100 mm; (d) tooth, scale bar, 5 mm; (e) tooth in labial, aboral, baso-lateral, and linguo-basal views, scale bar, 10 mm. adbc, anterior dorsal basal cartilage; bh, basihyal; cb, ceratobranchial; ch, ceratohyal; col, cololite; fs, fin spine; mc, Meckel̀s cartilage; mpt, metapterygium; n, neurocranium; na, neural arches; pdbc, posterior dorsal basal cartilage; pq, palatoquadrate; rad, radials; sc, scapulacoracoid. (Online version in colour.)
Figure 2.
Figure 2.
Otic and occipital region of P. saidselachus sp. nov., PIMUZ A/I 4711, reconstructed on the basis of CT scans: (a) anterior, (b) ventral, (c) dorsal, (d) lateral, and (e) posterior view. Braincase and articulated branchial arches: (f) ventral view of braincase and ceratohyals, (g) anterior aspect of ceratohyal, (h) lateral view of ceratohyal, (i) oblique lateral view of ceratohyal, (j) dorsolateral view on hyomandibula-braincase articulation. Scale bars, 30 mm. chy, ceratohyal; dor, dorsal otic ridge; endf, endolymphatic foramen; esc, external semicircular canal; fm, foramen magnum; glc, glossopharyngeal canal; hl, hypotic lamina; hym, hyomandibula; lda, lateral dorsal aorta; lof, lateral otic fossa; nc, notochordal canal; oc cot, occipital condyle; occr, occipital crest; psc, posterior semicircular canal; sac, sacculum. (Online version in colour.)
Figure 3.
Figure 3.
Otic and occipital region of the endocast of P. saidselachus (PIMUZ A/I 4711): (a) dorsal, (b) ventral, (c) lateral, (d) anterior, and (e) posterior views. Scale bar, 30 mm. esc, external semicircular canal; med, medulla; pa, posterior ampulla; psc, posterior semicircular canal; sac, sacculum; socc, spino-occipital. (Online version in colour.)
Figure 4.
Figure 4.
Cladogram (strict consensus tree) showing the placement of Phoebodus and Thrinacodus within the elasmobranchs. Colour coding: black, stem group gnathostome (outgroup) and Osteichthyes (Entelognathus to Raynerius); orange, stem Chondrichthyes including Acanthodii (Culmacanthus to Doliodus); purple, Holocephali (crown Chondrichthyes; Cladoselache to Chondrenchelys); blue, Elasmobranchii (crown Chondrichthyes, Cladodoides to Tribodus). White circles: bootstrap support of knot greater than 50% and/or Bremer decay values greater than 1; black circles: bootstrap support greater than 75% and/or Bremer decay values greater than 3. (Online version in colour.)
Figure 5.
Figure 5.
Possible body reconstruction of (a) P. saidselachus sp. nov., Late Devonian, (b) T. gracia [11], Early Carboniferous, and (c) picture of C. anguineus [44], Recent. (Online version in colour.)

References

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