Mauicetus’ Lophocephalus
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1 This is a postprint that has been peer reviewed and published in Zoological Journal of 2 the Linnean Society. The final, published version of this article is available online. 3 Please check the final publication record for the latest revisions to this article. 4 [Author's note: one figure, the only known tooth of an eomysticetid, is not present in 5 this postprint because it was added during final revisions. Consult the formally 6 published version of record.] 7 8 Boessenecker, R.W., and R.E. Fordyce. 2015. A new genus and species of 9 eomysticetid (Cetacea: Mysticeti) and a reinterpretation of ‘Mauicetus’ lophocephalus 10 Marples, 1956: transitional baleen whales from the upper Oligocene of New Zealand. 11 Zoological Journal of the Linnean Society 175:607-660. doi: 10.1111/zoj.12297 12 1 1 ABSTRACT 2 The early evolution of toothless baleen whales (Chaeomysticeti) remains elusive 3 despite a robust record of Eocene-Oligocene archaeocetes and toothed mysticetes. 4 Eomysticetids, a group of archaic longirostrine and putatively toothless baleen whales 5 fill in a crucial morphological gap between well-known toothed mysticetes and more 6 crownward Neogene Mysticeti. A historically important but perplexing cetacean is 7 “Mauicetus” lophocephalus (upper Oligocene South Island, New Zealand). The 8 discovery of new skulls and skeletons of eomysticetids from the Oligocene Kokoamu 9 Greensand and Otekaike Limestone permit a redescription and modern 10 reinterpretation of “Mauicetus” lophocephalus, and indicating that this species may 11 have retained adult teeth. A new genus and species, Tokarahia kauaeroa, is erected on 12 the basis of a well-preserved subadult to adult skull with mandibles, tympanoperiotics, 13 and cervical and thoracic vertebrae, ribs, sternum, and forelimbs from the Otekaike 14 Limestone (>25.2 Ma). “Mauicetus” lophocephalus is relatively similar and 15 recombined as Tokarahia lophocephalus. Phylogenetic analysis supports inclusion of 16 Tokarahia within the Eomysticetidae alongside Eomysticetus, Micromysticetus, 17 Yamatocetus, and Tohoraata, and strongly supports monophyly of Eomysticetidae. 18 Tokarahia lacked extreme rostral kinesis of extant Mysticeti and primitively retained 19 a delicate archaeocete-like posterior mandible and synovial temporomandibular joint, 20 suggesting that Tokarahia was capable of at most, limited lunge feeding in contrast to 21 extant Balaenopteridae, and utilized an alternative as-yet unspecified feeding strategy. 22 Baleen whales-Oligocene-Cetacea-Mysticeti-Eomysticetidae 23 24 INTRODUCTION 2 1 Sometime before 1937 an unassuming fragmentary skull was collected from a 2 limestone quarry near Milton in south Otago, South Island, New Zealand. It was 3 formally described by University of Otago zoology Professor W.B. Benham (1937) as 4 Lophocephalus parki, and thought to be a new archaeocete; it is now known to have 5 been collected from the latest Oligocene-earliest Miocene Milburn Limestone (Willett, 6 1946; Waitakian Stage, 25.2-21.7 Ma, Raine et al., 2012). Subsequent correspondence 7 led Benham to realize that the genus name was preoccupied and that Lophocephalus 8 parki was in fact an archaic baleen whale, which he later renamed Mauicetus parki 9 (Benham, 1942). Additional mysticetes collected by Brian J. Marples from North 10 Otago from the somewhat older Kokoamu Greensand (Duntroonian stage). In 1956 he 11 named these Mauicetus lophocephalus (based on a partial braincase, mandible, 12 tympanic bullae, and cervical and thoracic vertebrae), Mauicetus waitakiensis (based 13 on an occipital, tympanic bullae, and cervical vertebrae) and Mauicetus brevicollis 14 (based on a partial vertebral column and scapula). Recently collected material of 15 Mauicetus parki demonstrates that it is Although a reasonable referral at the time 16 owing to the incomplete knowledge of these Oligo-Miocene mysticetes, new 17 discoveries of fossil mysticetes from the South Island of New Zealand and other 18 continents in the past thirty years indicates that at least two of the species of 19 Mauicetus described by Marples (1956) represent fossil mysticetes dramatically 20 different from anything else known to twentieth century paleocetologists, and are not 21 closely related to Mauicetus parki (Fordyce, 2005; 2006; Boessenecker and Fordyce, 22 2014a). 23 In 2002, the strange new longirostrine toothless mysticete Eomysticetus 24 whitmorei was described from the upper Oligocene of South Carolina, U.S.A. 25 (Sanders and Barnes 2002b). Eomysticetus is characterized by a toothless palate, 3 1 kinetic rostrum, delicate frontals, a poorly “telescoped” skull, enormous temporal 2 fossae, elongate and cylindrical zygomatic processes, and basilosaurid-like 3 tympanoperiotics and postcrania (Sanders and Barnes 2002b). While these authors 4 recognized that Mauicetus lophocephalus was another archaic chaeomysticete and 5 similar in some regards to Eomysticetus, albeit not as archaic, they considered it to not 6 be “[included] within the clade containing Eomysticetus whitmorei and [Yamatocetus 7 canaliculatus]”. Their assertion that “Mauicetus” lophocephalus is not an 8 eomysticetid was not tested by cladistic analysis. Regardless, several features are 9 shared between “Mauicetus” lophocephalus and Eomysticetus, including enormous 10 temporal fossae, transversely narrow intertemporal region with a high sagittal crest, 11 elongate and subcylindrical zygomatic processes, all unique features amongst 12 Chaeomysticeti (although primitively present as well in Basilosauridae and certain 13 toothed mysticetes). In recognition of these similarities with Eomysticetus and 14 concomitant differences from Mauicetus parki, the fragmentary “Mauicetus” 15 waitakiensis was transferred to the newly described eomysticetid genus Tohoraata 16 (Boessenecker and Fordyce, 2014a). Unfortunately, the holotype skull and scapulae of 17 “Mauicetus” lophocephalus are missing (Fordyce, 1980:20), likely discarded by 18 university maintenance staff not long before 1962 (J.T. Darby, personal 19 communication to R.E. Fordyce, May 1978). However, the tympanoperiotics, 20 mandible, and much of the postcrania remain, permitting limited reassessment of this 21 critical but poorly understood fossil mysticete. 22 Newly discovered fossil material from the upper Oligocene Otekaike 23 Limestone of New Zealand (Fig. 1, 2) including a spectacularly preserved and nearly 24 complete skull with mandibles, tympanoperiotics, and partial postcranial skeleton 25 (cervical and thoracic vertebrae, ribs, sternum, scapulae, humeri, radius, ulnae) share 4 1 near identical tympanoperiotic morphology with “Mauicetus” lophocephalus, and 2 critically share similar skull morphology with published eomysticetids and 3 “Mauicetus” lophocephalus. The skull and skeleton of this new fossil mysticete is 4 remarkable in its transitional morphology between toothed Mysticeti and Neogene 5 crown Mysticeti (=Balaenomorpha), serving as an exemplar for comparison between 6 the two. Another partial skull and skeleton is tentatively referred to “Mauicetus” 7 lophocephalus, and notably includes an isolated tooth differing in morphology from 8 all other cetaceans and indicates that eomysticetids may have primitively retained 9 non-functional adult teeth (out of convention, “toothed mysticete” hereafter refers to 10 stem Mysticetes not including eomysticetids). The aim of this study is to report the 11 new morphological details preserved in this new genus and species of archaic 12 mysticete and other similar specimens (Fig. 3), provide a new description through 13 which “Mauicetus” lophocephalus may be reinterpreted in light of recent advances in 14 paleocetology, and establish a phylogenetic context for these distinctive fossil 15 cetaceans through cladistic analysis. 16 17 MATERIALS AND METHODS 18 Preparation, Anatomical Description, and Illustration: Fossil material in OU 19 collections was mechanically prepared with pneumatic air scribes. Fine preparation 20 was performed under a Zeiss binocular microscope. Anatomical terminology follows 21 Mead and Fordyce (2009), Oishi and Hasegawa (1995) and Ekdale et al. (2011). 22 Tympanoperiotic orientation follows Mead and Fordyce (2009) using anatomical 23 structures (e.g. anterior process, posterior process, lateral tuberosity) to dictate 24 orientation of tympanoperiotics when in isolation from the skull (in contrast to 25 orientation in situ) to facilitate comparisons between taxa. 5 1 Osteohistology: Histologic sections were taken from rib fragments of OU 22235 2 (Tokarahia kauaeroa holotype) and OU 22081 (Tokarahia sp., cf. T. lophocephalus). 3 Sections were embedded in epoxy and thin sections were prepared by University of 4 Otago Petrology Technician B. Pooley. Photomicrographs were captured under non- 5 polarized light. 6 Cladistic Methodology: A cladistic analysis was executed in order to assess the 7 phylogenetic relationships of Tokarahia spp. in addition to the previously described 8 Tohoraata raekohao. This analysis includes 74 terminal taxa (Archaeoceti, n=3; 9 Odontoceti, n=2, Mysticeti, n=69; 12 extant mysticetes and one extant odontocete) 10 and a total of 363 morphological characters including cranial (n=251; 104 characters 11 for the tympanoperiotic complex), mandibular (n=27), dental (n=15), postcranial 12 (n=46), and soft tissue characters (n=24). Character inclusion began with the matrix 13 of Marx (2011) and included 101 characters modified from Fitzgerald (2010), 142 14 characters modified from Fordyce and Marx (2013), and