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Comparative Osteology and Phylogenetic Relationships Of bs_bs_banner Zoological Journal of the Linnean Society, 2012, 166, 876–911. With 22 figures Comparative osteology and phylogenetic relationships of Miocaperea pulchra, the first fossil pygmy right whale genus and species (Cetacea, Downloaded from https://academic.oup.com/zoolinnean/article-abstract/166/4/876/2627067 by guest on 12 September 2019 Mysticeti, Neobalaenidae) MICHELANGELO BISCONTI* Museo di Storia Naturale del Mediterraneo, Via Roma 234, 57125, Livorno, Italy Received 18 August 2011; revised 6 July 2012; accepted for publication 26 July 2012 A fossil pygmy right whale (Cetacea, Mysticeti, Neobalaenidae) with exquisitely preserved baleen is described for the first time in the history of cetacean palaeontology, providing a wealth of information about the evolu- tionary history and palaeobiogeography of Neobalaenidae. This exquisitely preserved specimen is assigned to a new genus and species, Miocaperea pulchra gen. et sp. nov., and differs from Caperea marginata Gray, 1846, the only living taxon currently assigned to Neobalaenidae, in details of the temporal fossa and basicranium. A thorough comparative analysis of the skeleton of M. pulchra gen. et sp. nov. and C. marginata is also provided, and forms the basis of an extensive osteology-based phylogenetic analysis, confirming the placement of M. pulchra gen. et sp. nov. within Neobalaenidae as well as the monophyly of Neobalaenidae and Balaenidae; the phylogenetic results support the validity of the superfamily Balaenoidea. No relation- ship with Balaenopteroidea was found by the present study, and thus the balaenopterid-like morphological features observed in C. marginata must have resulted from parallel evolution. The presence of M. pulchra gen. et sp. nov. around 2000 km north from the northernmost sightings of C. marginata suggests that dif- ferent ecological conditions were able to support pygmy right whale populations in what is now Peru, and that subsequent environmental change caused a southern shift in the distribution of the living neobalaenid whales. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166, 876–911. doi: 10.1111/j.1096-3642.2012.00862.x ADDITIONAL KEYWORDS: Balaenoidea – Caperea marginata – Miocene – Peru – phylogeny. INTRODUCTION Neobalaenidae (Baker, 1985). In fact, most mor- phological analyses support a close relationship of In the last few decades the evolutionary history of Neobalaenidae and Balaenidae (right and bowhead baleen whales (Mammalia, Cetacea, Mysticeti) has whales; Bisconti, 2005; Deméré, Berta & McGowen, been investigated by morphological and molecular 2005), but the molecular and few morphological analyses, but only a limited consensus has emerged studies performed have almost invariably indica- from this effort. One of the major points of disagree- ted a close relationship of Neobalaenidae and the ment concerns the phylogenetic relationships of the Eschrichtiidae–Balaenopteridae clade (gray, rorqual, pygmy right whale Caperea marginata Gray, 1846, and humpback whales; Sasaki et al., 2005; Nikaido the only species currently assigned to the family et al., 2006; Deméré et al., 2008). Neobalaenids are characterized by balaenid-like features, such as: arched rostrum; high number of *E-mail: [email protected] long baleen plates; fused cervical vertebrae; low and 876 © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166, 876–911 FIRST FOSSIL NEOBALAENID WHALE 877 wide tympanic bulla, with low tympanic cavity; anteriorly thrusted supraoccipital that is superim- posed on the parietal, preventing the parietal from appearing at the skull vertex; and alimentary behaviour based on continuous ram feeding to exploit calanoid copepods (Beddard, 1901; Baker, 1985). Balaenopterid- and eschrichtiid-like features include: presence of ventral throat grooves in some individu- als; squamosal cleft; a dorsal fin, radius, and ulna Downloaded from https://academic.oup.com/zoolinnean/article-abstract/166/4/876/2627067 by guest on 12 September 2019 longer than humerus; and flat supraorbital process of the frontal (Baker, 1985; Marx, 2010). Given that C. marginata possesses a mix of balaenid and bal- aenopterid characters, it is difficult to understand which features are the result of convergence and which are those representing the proof of true phylo- genetic relationships. Up to present times, the fossil record could not help reconstruct the ancestral mor- phological conditions of Neobalaenidae because it was considered to be non-existent (Fordyce & De Muizon, 2001; Fordyce, 2009). A recent find from New Zealand suggested that neobalaenid whales were existent in the southern hemisphere around 6.2–5.4 Mya (Fitzgerald, 2012), but the find consisted of a single posterior process of a periotic that is not diagnostic enough to provide information about taxonomy and phylogenetic relationships. In this article, the anatomy of an exquisitely preserved neobalaenid skull is reported and its phylogenetic impli- cations are discussed. The new specimen is the holotype of Miocaperea pulchra gen. et sp. nov., found from the upper Miocene Pisco Formation at Aguada de Loma, Peru (Fig. 1), and is now permanently housed in the Staatliches Museum für Naturkunde, Stuttgart, Germany, as specimen no. 46978 of the palaeonto- logical collection. The specimen was excavated by Jakob Siber in 1985, and was legally exported by the Siber+Siber Aathal/Zürich company (E.P.J. Heizmann, pers. comm.). Mr Siber confirmed the legal status of the specimen before the Society of Friends of the Natural History Museum Stuttgart bought it (E.P.J. Heizmann, pers. comm.). The legal documenta- tion can be provided by Siber+Siber Aathal/Zürich. The specimen came to the Stuttgart collection as a present from the Society of Friends of the State Museum of Natural History (SMNS), together with further Peru- vian material (e.g. the skeleton of Balaenoptera siberi Pilleri, 1989, which is exhibited in the Schloss Rosen- stein building of the SMNS). Detailed comparisons with the living pygmy right whale C. marginata are provided to form a solid Figure 1. Locality of the discovery of Miocaperea basis for a new and comprehensive cladistic analysis pulchra gen. et sp. nov. A, South America; B, Peruvian of Mysticeti, directed at discovering the phylogenetic territory with Aguada de Lomas indicated by a line; C, relationships of Neobalaenidae. An in-depth analy- close-up view of Aguada de Lomas in Peru. sis of the basicranium, earbones, and postcranial skeleton of the extant C. marginata are included to supplement the section dealing with the compara- © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166, 876–911 878 M. BISCONTI tive anatomy of M. pulchra gen. et sp. nov., so as to foramen; ofh, humerus facet for olecranon process of provide the first detailed morphological descriptions ulna; oft, open foramen transversarium; opu, olecra- of some anatomical parts of the neobalaenid skel- non process of ulna; oul, outer lip; ow, oval window; eton, and to provide data about individual variation p, parietal; patb, posterior site for attach with peri- and the growth trajectory of C. marginata. otic; pch, pars cochlearis of periotic; pe, posterior In this article, the anatomical terminology is taken end of the pelvis; per, proximal epiphysis of radius; from Mead and Fordyce (2010) and, for limited parts, pg, promontorial groove; pgp, postglenoid process of from Nickel et al. (1999), Schaller (1999), and Struth- squamosal; plc, posterolateral corner; plf, perilym- ers (1895). phatic duct; pmc, posteromedial corner; pmx, pre- Downloaded from https://academic.oup.com/zoolinnean/article-abstract/166/4/876/2627067 by guest on 12 September 2019 Anatomical abbreviations: aar, area of acetabulum maxilla; pop, postorbital process of supraorbital in the pelvis; ab-pch, anterior border of pars cochle- process of the frontal; ppp, posterior process of peri- aris; ae, anterior end of pelvis; afa, atlas articular otic; pppb-br, broken base of posterior process of facet for occipital condyle; afi, (incus) articular facet periotic; p-sq, parietal–squamosal suture; pt, ptery- for malleus; alc, anterolateral corner; amc, antero- goid; ptg, groove for insertion of pterygoid muscle; medial corner; ang, angular process of dentary; ap, ptf, pterygoid fossa; rfh, radial facet of humerus; rw, acromion process of scapula; aplf, area of posterior round window; sc, sagittal crest; sip, sigmoid ligament of femur in the pelvis; app, anterior process; smf, stylomastoid fossa; soc, supraoccipital; process of periotic; b, baleen; bC5 and bC6, body sop, supraorbital process of frontal; spf, supras- of cervical vertebrae 5 and 6; bdp, basioccipital pinous fossa of scapula; sq, squamosal; ss, scapular descending process; bg, groove for vasculature spine; ssf, subscapular fossa; st, stapes; stfo, stape- of the baleen-bearing epithelium; boc, basioccipi- dial foramen (stapes); ST(T1), scala tympani, first tal; bst, base of stapes; c, coana; C2–C7, cervical cochlear turn; SV(T1), scala vestibuli, first cochlear vertebrae 2–7; cb, crus breve (incus); cdp, caudal turn; T, thoracic vertebra; tc, temporal crest; T2, process of periotic; cop, conical process; cp, coracoid second cochlear turn; td, trapezoid; tf, temporal process of scapula; cs, cranial surface of periotic; cu, fossa; tyc, tympanic cavity; typ, tympanic plate; ufh, cuneiform; dc, dorsal crest of periotic; eam external ulnar facet of humerus; um, umbo (stapes); un,
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