Memoirs of Museum Victoria 62(1): 67–89 (2005) ISSN 1447-2546 (Print) 1447-2554 (On-line) http://www.museum.vic.gov.au/memoirs/index.asp Pliocene marine mammals from the Whalers Bluff Formation of Portland, Victoria, Australia ERICHM.G. FITZGERALD School of Geosciences, Monash University, Vic. 3800, Australia and Museum Victoria, G.P.O. Box 666, Melbourne, Vic. 3001, Australia ([email protected]) Abstract Fitzgerald, E.M.G. Pliocene marine mammals from the Whalers Bluff Formation of Portland, Victoria, Australia. Memoirs of Museum Victoria62(1): 67–89. The most diverse and locally abundant Australian fossil marine mammal assemblages are those from late Neogene (Late Miocene through Late Pliocene) sediments in Victoria and Flinders Island, Tasmania. However, none of these assemblages have hitherto been described. The Pliocene (>2.5–4.8 Ma) Whalers Bluff Formation, exposed in beach cliff sections and offshore reefs, at Portland, western Victoria (38°19'S, 141°38'E) has yielded a small but moderately diverse assemblage of marine mammals represented by fragmentary material. Taxa present include: right whales (Balaenidae); rorqual whales (Balaenopteridae); a physeterid similar to the extant sperm whale (cf. Physetersp.); the first Australian fossil record of pygmy sperm whales (Kogiidae); at least three genera of dolphins (Delphinidae: cf. Tursiops sp.,Delphinussp. or Stenellasp., and an undetermined genus and species); and probable earless or true seals (Phocidae). This small assemblage represents the first Australian fossil marine mammal assemblage to be described in detail. The taxonomic composition of this Pliocene marine mammal assemblage is generally similar to the present day marine mammal assemblage in north-west Bass Strait. The occurrence of extant cetacean genera in the Portland Pliocene and Flinders Island Cameron Inlet Formation assemblages indicates that the marine mammal fauna off south-east Australia had acquired an essentially modern aspect by the Late Pliocene. Several of the cetacean genera recorded in the Portland Pliocene assemblage also occur in similar-aged assemblages in other ocean basins. This corroborates the hypothesis that many cetacean taxa that are widely distributed in the world’s oceans today were equally widespread during the Pliocene. Keywords Cetacea, Carnivora, Pinnipedia, Phocidae, Mysticeti, Odontoceti, Australia, Victoria, Portland, Whalers Bluff Formation, Pliocene Introduction especially the Palaeontology Collections of Museum Victoria, Melbourne. The Pliocene epoch (1.8–5.3 Ma) is generally considered to be The majority of Pliocene marine mammal fossils in these the time during which the modern marine mammal fauna collections are rather fragmentary with one partially complete evolved, with the extinction of archaic taxa (as well as some skull and associated skeleton known (NMV P179005, cf. taxa with novel adaptations), and widespread geographic Megaptera sp.). Despite this general lack of diagnostic skull distribution of extant families and genera (Barnes, 1977; material, some details of the SW Pacific Pliocene marine Fordyce, 1989; Fordyce and Barnes, 1994; Fordyce and mammal fauna may be filled in by the study of certain isolated Muizon, 2001; Fordyce et al., 2002; Deméré et al., 2003). skeletal elements such as periotics. As noted by Barnes (1977: Although the Pliocene marine mammals of the North Pacific 322), study of these isolated elements may provide data on the (e.g. Barnes, 1973a, 1977, 1998) and eastern tropical taxonomic diversity within an assemblage and wider fauna. Pacific (e.g. Muizon, 1981, 1984; Muizon and DeVries, 1985; Pliocene marine mammal assemblages have hitherto not been Muizon and Domning, 1985, 2002) have been described and described from Australia, so descriptions of even fragmentary discussed in some detail, the Pliocene marine mammals of material provide an initial basis for understanding marine the Southwest Pacific (Australia and New Zealand) remain mammal evolution off southern Australia during the Pliocene. poorly known (Fordyce et al., 2002: 53). This is despite the fact This description of marine mammals from the Pliocene Whalers that Australian late Neogene marine mammals (mostly Bluff Formation assemblage comprises a preliminary basis for cetaceans) are relatively abundant in museum collections, the late Neogene fossil record of marine mammals in Australia. 68 Erich M.G. Fitzgerald 140° 144° 148° SOUTH AUSTRALIA NEW SOUTH WALES Adelaide 36° 36° VICTORIA Melbourne 200 km Bass Strait 40° 40° 140° 144° 148° y a W 20 km 2 km n o utt Glenelg D Ri ver Portland Nelson Anderson Point Whalers Point Nunns Beach Bay Portland Portland Portland Bay Southern Ocean Figure 1. Locality of Portland in Victoria, south-east Australia, and the Portland fossil marine vertebrate localities. Fossils have been collected as float along the beach and from adjacent cliffs between Dutton Way and Portland Harbour. Black shading indicates areas of cliff outcrop of the Whalers Bluff Formation. Pliocene marine mammals 69 All fossils were collected from coastal exposures of the Materials and methods Whalers Bluff Formation lining Portland Bay in western All fossil specimens were collected by Mr Sean Wright of Portland and coastal Victoria, southeast Australia (38°19'S, 141°38'E) are in the Palaeontology Collections of Museum Victoria. Anatomical (Fig. 1). Unfortunately, details of the geological context of vir- terminology for periotics and tympanics follows Evans (1993), tually all fossils are unknown apart from whether the fossils Fordyce (1994), Fordyce and others (2002) and Kasuya (1973) with were collected from the Whalers Bluff Formation or underlying minor modifications. All periotic and tympanic measurements follow limestone. Fitzgerald (2004a, 2004b) mentioned the Portland the methods and dimensions outlined by Kasuya (1973) and were made fossil marine mammals in previous publications. Bearlin (1987: using vernier callipers. Photographs were taken using a 35 mm Nikon 177) briefly noted the occurrence of cf. Balaena sp., and cf. Nikkormat EL SLR with a 105 mm macro-lens, and a Nikon D70 Balaenopterasp. in private collections in an unpublished Ph.D. digital SLR with a 60 mm macro-lens. Where indicated, specimens were coated with a sublimate of ammonium chloride to enhance thesis. The vertebrate faunal list for Portland given by contrast in black and white (denoted by AC in figure captions). Where Fitzgerald (2004b: 186) was completed prior to the recognition necessary, fragile specimens were consolidated with a hardener con- of two distinct, stratigraphically/temporally disjunct marine sisting of 3% solution of Paraloid B72 (ethyl methacrylate/methyl vertebrate assemblages at Portland. The majority of the verte- acrylate copolymer) in acetone. brates listed by Fitzgerald (2004b) were derived from the Institutional abbreviations. NMVC, Museum Victoria Comparative Pliocene Whalers Bluff Formation although some vertebrates Anatomy Collection, Melbourne; NMV P, Museum Victoria from the Portland Late Miocene assemblage were listed under Palaeontology Collection, Melbourne; CD, Phylum Chordata cata- the Whalers Bluff Formation assemblage. The Portland Late logue, New Zealand Geological Survey, Lower Hutt; USNM, National Miocene marine vertebrate assemblage, from the Port Museum of Natural History (formerly United States National Campbell Limestone, is considerably more diverse than the Museum), Smithsonian Institution, Washington, DC. For a complete list of specimens referred to in this study, see table 1. Pliocene Whalers Bluff Formation assemblage, and is to be described in a subsequent publication. Below is an emended list of the vertebrates recorded in the assemblage from the Whalers Geology and age of the Whalers Bluff Formation Bluff Formation at Portland. For about 4 km along cliffs north of Portland Harbour the Chondrichthyes Pliocene Whalers Bluff Formation is exposed (Singleton et al., Isurussp. 1976). This formation is about 7.6 m thick, being comprised of Carcharodon carchariasLinnaeus, 1758 horizontally bedded fossiliferous clay, oyster-rich beds and Carcharodon megalodonAgassiz, 1835 sandy limestones (Abele et al., 1988). These sediments uncon- Myliobatissp. formably overlie the Upper Miocene Port Campbell Limestone Ischyodus dolloiLeriche, 1902 and infill a karst topography developed in the top of the Mammalia Miocene limestone (Boutakoff and Sprigg, 1953; Dickinson et Marsupialia al., 2002). The Whalers Bluff Formation is unconformably ?Dasyuromorphia incertae sedis capped by basalts. Diprotodontoidea gen. et sp. undet. The age of the Whalers Bluff Formation is well constrained Diprotodontoidea gen. et sp. undet. C relative to some other Neogene marine mammal-bearing units Diprotodontidae gen. et sp. undet. A in the SWPacific (Fig. 2). However, the determination of the Zygomaturinae gen. et sp. undet. T younger age limit of the Whalers Bluff Formation has proved Palorchestessp. problematic. The Port Campbell Limestone which underlies the Vombatidae gen. et sp. undet. Whalers Bluff Formation is Late Miocene (indicated by pres- Sthenurussp. ence of Globorotalia miotumida; planktonic foraminiferal Protemnodonsp. zones N16-basal N17; Tortonian; 8–10.8 Ma) (Dickinson et al., Macropussp. 2002). Deposition of the Whalers Bluff Formation began Macropodidae gen. et sp. undet. C during planktonic foraminiferal zone N19 (indicated by the Ektopodontidae gen. et sp. undet. presence of Globorotalia puncticulata at the base of the for- Rodentia mation) and ensued into the base of planktonic foraminiferal Rodentia incertae sedis zone N21 (Fig. 2) (Singleton et al., 1976; Dickinson et al., Carnivora 2002). K-Ar dates of 2.51 Ma from basalts that cap the Whalers ?Phocidae gen. et sp. indet. Bluff Formation have been reported (Singleton et al., 1976). Cetacea Beu and Darragh (2001) have suggested that the Whalers Balaenidae gen. et sp. indet. Bluff Formation is latest Pliocene to earliest Pleistocene Balaenopteridae gen. et sp. indet. (>1.5–2.0 Ma) based on the presence of the pectinid bivalve cf.Physetersp. Pecten fumatus Reeve, 1852 in a section along the Glenelg Kogiidae gen. et sp. indet. River. However, Pecten fumatusdoes not occur in the Whalers Delphinoidea incertae sedis Bluff Formation at Portland (Darragh in Singleton et al., 1976; cf.Tursiopssp. T.A. Darragh, pers. comm.). Zenatiopsis ultima Darragh and Delphinussp. or Stenellasp. Kendrick, 1971 occurs in the Whalers Bluff Formation at Delphinidae gen. et sp. undet. A Portland but is a Pliocene species and never occurs with 70 Erich M.G. Fitzgerald Table 1. Specimens referred to in this study. For more detailed locality/stratigraphic data see Fitzgerald (2004b). Abbreviations: e=Early; m=Middle; l=Late; M=Miocene; P=Pliocene; Pt=Pleistocene; R=Recent; Vic. = Victoria; Tas. = Tasmania; NZ = New Zealand. Specimen Taxon Locality Formation Age NMVP221242 Isurussp. Portland, Vic. Whalers Bluff P NMVP218415 Carcharodon megalodon Portland, Vic. Whalers Bluff P NMVP218418 Carcharodon carcharias Portland, Vic. Whalers Bluff P NMVP218470 Myliobatissp. Portland, Vic. Whalers Bluff P NMVP218296 Ischyodus dolloi Portland, Vic. Whalers Bluff P NMVP221241 ?Dasyuromorphia gen. et sp. undet. Portland, Vic. Whalers Bluff P NMVP218500 Diprotodontoidea gen. et sp. undet. Portland, Vic. Whalers Bluff P NMVP218498 Diprotodontoidea gen. et sp. undet. C Portland, Vic. Whalers Bluff P NMVP218499 Diprotodontidae gen. et sp. undet. A Portland, Vic. Whalers Bluff P NMVP221230 Zygomaturinae gen. et sp. undet. T Portland, Vic. Whalers Bluff P NMVP221231 Palorchestessp. Portland, Vic. Whalers Bluff P NMVP221238 Vombatidae gen. et sp. undet. Portland, Vic. Whalers Bluff P NMVP221232 Protemnodonsp. Portland, Vic. Whalers Bluff P NMVP221235 Macropussp. Portland, Vic. Whalers Bluff P NMVP221227 ?Macropussp. Portland, Vic. Whalers Bluff P NMVP221237 Sthenurussp. Portland, Vic. Whalers Bluff P NMVP221229 Macropodidae gen. et sp. undet. C Portland, Vic. Whalers Bluff P NMVP197795 Ektopodontidae gen. et sp. undet. Portland, Vic. Whalers Bluff P NMVP221240 Rodentia indet. Portland, Vic. Whalers Bluff P NMVP218273 ?Phocidae gen. et sp. indet. Portland, Vic. Whalers Bluff P NMVP218465 ?Phocidae gen. et sp. indet. Portland, Vic. Whalers Bluff P NMVP218269 Balaenidae gen. et sp. indet. Portland, Vic. Whalers Bluff P NMVP218268 Balaenopteridae gen. et sp. indet. Portland, Vic. Whalers Bluff P NMVP218298 cf. Physetersp. Portland, Vic. Whalers Bluff P NMVP218407 Kogiidae gen. et sp. indet. Portland, Vic. Whalers Bluff P NMVP218283 Delphinoidea incertae sedis Portland, Vic. Whalers Bluff P NMVP218284 Delphinoidea incertae sedis Portland, Vic. Whalers Bluff P NMVP218286 Delphinoidea incertae sedis Portland, Vic. Whalers Bluff P NMVP218266 cf. Tursiopssp. Portland, Vic. Whalers Bluff P NMVP218265 DelphinusorStenellasp. Portland, Vic. Whalers Bluff P NMVP218264 Delphinidae gen. et sp. undet. A Portland, Vic. Whalers Bluff P NMVP16198 ?Phocidae gen. et sp. indet. Beaumaris, Vic. Black Rock Sandstone lM-eP NMVP41759 Phocidae gen. et sp. indet. Beaumaris, Vic. Black Rock Sandstone lM-eP NMVP42523 ?Phocidae gen. et sp. indet. Beaumaris, Vic. Black Rock Sandstone eP NMVP160399 Phocidae gen. et sp. undet. Beaumaris, Vic. Black Rock Sandstone eP NMVP160433 Phocidae gen. et sp. undet. Beaumaris, Vic. Black Rock Sandstone eP NMVP160441 Phocidae gen. et sp. undet. Grange Burn, Vic. Grange Burn eP NMVP215759 ?Phocidae gen. et sp. indet. Beaumaris, Vic. Black Rock Sandstone eP NMVP16195 Balaenidae gen. et sp. indet. Beaumaris, Vic. Black Rock Sandstone eP NMVP48865 Balaenidae gen. et sp. indet. Grange Burn, Vic. Grange Burn eP NMVP160438 Balaenidae gen. et sp. indet. Beaumaris, Vic. Black Rock Sandstone lM-eP NMVP197824 Balaenidae gen. et sp. indet. Beaumaris, Vic. Black Rock Sandstone lM-eP NMVP171503 Balaenoptera edeniorB. brydei Tidal River, Vic. N/A R NMVP179005 Megapterasp. Lakes Entrance, Vic. Jemmys Point eP NMVP23961 ?Mesoplodonsp. Cameron Inlet, Tas. Cameron Inlet P NMVP13033 “Steno”cudmorei Beaumaris, Vic. Black Rock Sandstone lM-eP NMVP204352 Delphinidae gen. et sp. undet. A Henley, England Red Crag P-Pt NMVP218481 Delphinidae gen. et sp. undet. A Henley, England Red Crag P-Pt NMVC24972 Kogia breviceps Cape Conran, Vic. N/A R NMVC24976 Kogia breviceps Shelley Beach, Vic. N/A R NMVC27879 Eubalaena australis Altona Bay, Vic. N/A R NMVC28892 Megaptera novaeangliae Venus Bay, Vic. N/A R CD 53 Delphinidae gen. et sp. undet. A Chatham Island, NZ Unnamed ?P USNM 22953 Orycterocetus crocodilinus Calvert County, USA Calvert mM USNM 183007 Physeteridae gen. et sp. undet. Lee Creek Mine, USA Yorktown eP USNM 452993 Scaphokogia cochlearis Aguada de Lomas, Peru Pisco lM USNM 183008 Kogiidae gen. et sp. undet. Lee Creek Mine, USA Yorktown eP USNM 251118 Kogiidae gen. et sp. undet. Lee Creek Mine, USA Yorktown eP Pliocene marine mammals 71 Ma Chronostratigraphy PSltaFanonkrdataomrndisc ReSgEi oAnuasl tSrataliganes StPraotrigtlraanpdhy SE Australia SE Pacific NE Pacific NW Atlantic -MAanrtianrec tPiclaain 2543 PLIOC. EarlyLate PZiaacnecnlezaiann NN1N1928-120 WKearliikmonoaiann BWasBaFlt FCoarImmnleaeBrttoiRonnS SSAAOS SUDAFF FYoorrmktaotwionn 6 Messinian Cheltenhamian GBF Sørsdal 7 E e N17 MATGML LAF Formation 98 EN Lat Tortonian N16 Mitchellian Port Campbell ELJ 10 C Limestone CLB 11 O N15 N14 Bairnsdalian 12 MI N13 111543 Middle SeLrarnagvhailaliann NNNNN11198120 BBaatelcsofomrbdiiaann BL MRooSuunilnttadin FoCramlvaetirotn 16 Figure 2. Stratigraphic correlation of the Portland fossil marine mammal-bearing formations with selected major late Neogene marine mammal- bearing units. Stratigraphy and geochronology are from Barnes (1973, 1977, 1984, 1998), Muizon and DeVries (1985), Muizon and Bellon (1986), Gottfried et al. (1994), Whitmore (1994), Prothero (1998), Fordyce (2002a), Fordyce et al. (2002), Fitzgerald (2004b), Muizon et al. (2004), Barnes et al. (2005) and Gradstein et al. (2004). Abbreviations: AGL, Pisco Formation, Aguada de Lomas level; BL, Batesford Limestone; BRS, Black Rock Sandstone; CLB, Pisco Formation, Cerro la Bruja; ELJ, Pisco Formation, El Jahuay level; GBF, Grange Burn Formation; LAF, Lower Member, Almejas Formation; MTM, Pisco Formation, Montemar level; SAO, Pisco Formation, Sacaco level; SAS, Pisco Formation, Sud- Sacaco level; SDF, San Diego Formation; UAF, Upper Member, Almejas Formation; WBF, Whalers Bluff Formation. P. fumatus, the latter first appearing at the base of the Description. P218269 is highly polished and abraded. The Pleistocene (T.A. Darragh, pers. comm.). anterior process is blunt and globose, being indistinct from the These data indicate that the Whalers Bluff Formation at superior process. There is marked lateral exostosis of the super- Portland is Early to Late Pliocene (Zanclean to Piacenzian; ior process lateral to the epitympanic recess. The lateral aspect >2.5–4.8 Ma). Further study of terrestrial mammals from the of the anterior process is rugose and pitted. Posteriorly, this pit- Whalers Bluff Formation may help refine the geological age of ting decreases in density. Only the lateralmost region of the this unit. Unfortunately, the exact bed from which fossil verte- pars cochlearis is preserved. In medial view, the most notable brates were collected within the Whalers Bluff Formation is feature is the sulcus for the facial nerve (cr. VII), the course of unknown. Thus, a finer age resolution than that given above for cr. VII being preserved from its entry into the body of the the assemblage is currently unavailable. The age range of the periotic at the aperure of the internal facial foramen, to its Whalers Bluff Formation presented herein is considered to ventral exit into the epitympanic cavity via the ventral facial be the best estimate based on available data and the reader must foramen. All other features of the pars cochlearis and epitym- be cautioned that future work may yield a younger limit on the panic recess have been obliterated. Posterior to the broad and minimum age. shallow hiatus epitympanicus is a remnant of the base of the The Whalers Bluff Formation (as well as, in part, laterally posterior process (which is directed posterolaterally and equivalent Victorian marine mammal-bearing units such as the somewhat ventrally). Grange Burn Formation, Black Rock Sandstone and Jemmys Point Formation) has been interpreted as representing a Discussion. Miller (1924: 8–9) listed the following features that prograding quartz-carbonate barrier system with the clastic- distinguish the periotics of Balaenidae from those of dominated units listed above representing an initial marine Balaenopteridae (and other baleen-bearing Mysticeti): (1) axis incursion (Dickinson et al., 2002: 290). of anterior process of periotic parallel with axis of internal acoustic meatus; (2) [longitudinal] axes of anterior and posteri- Systematics of marine mammals or processes converge at an acute angle; and (3) pars cochlearis small relative to rest of periotic. In addition to the preceding OrderCetaceaBrisson, 1762 features, the possession of massive lateral exostosis of the SuborderMysticetiFlower, 1864 anterior process and anterolateral superior process, such that the anterior process appears swollen (as noted by Fordyce, FamilyBalaenidaeGray, 1821 1982: 48), seems to be a feature shared by all extant and late Neogene balaenid periotics. It is largely on the basis of the Genus and species indeterminate latter character and the phenetic similarity of P218269 to a periotic (P16195) from the Lower Pliocene Black Rock Referred specimen. NMVP218269, incomplete right periotic; anterior and superior processes virtually complete, but pars cochlearis almost Sandstone of Beaumaris identified as belonging to cf. entirely worn off, and only anteriormost base of posterior process “Balaena” (Gill, 1957) that P218269 is referred to Balaenidae, preserved (Fig. 3A). genus and species indeterminate. 72 Erich M.G. Fitzgerald The fossil record of Balaenidae begins in the Late Oligocene what attenuated anteriorly. The dorsal surface of the anterior (c. 28 Ma: Fordyce, 2002b), although the record only becomes process is smooth, with only slight rugosity, as seen in the reasonably well known from the Mio-Pliocene boundary periotics of extant Balaenopteridae. An oblique groove on the onwards (McLeod et al., 1993; Bisconti, 2003). Morenocetus dorsolateral surface of the anterior process near its preserved parvus Cabrera, 1926 is the geologically oldest named apex is interpreted as a trace of a vascular sulcus. The latter balaenid, from the early Early Miocene (Aquitanian) of feature has previously been considered a sulcus for the cap- Patagonia. From the end Aquitanian to early Tortonian of the suloparietal emissary vein (Geisler and Luo, 1998; Geisler and Miocene the evolutionary history of Balaenidae is virtually Sanders, 2003) or as a sulcus marking the path of an artery, unknown. The extant balaenids include Balaena mysticetus specifically part of the middle meningeal artery (Fordyce, Linnaeus, 1758, Eubalaena australis Desmoulins, 1822, 1994). Fordyce (1994) and Watson and Fordyce (1994) E. glacialis Müller, 1776, and E. japonica Lacépède, 1818 described this feature as the anteroexternal sulcus whereas (e.g., Cummings, 1985; Reeves and Leatherwood, 1985; Geisler and Sanders (2003) treated the anteroexternal sulcus Bannister, 2002). Note that Rice (1998) included all extant and sulcus for the capsuloparietal emissary vein as separate fea- balaenids in the genus Balaena and recognised only two tures. Further work is required to better establish the species,B. mysticetusandB. glacialis. The taxonomic scheme venous/arterial correlate of this osteological feature which in of Bannister (2002) is used herein. Balaenais known from the this study is referred to as the sulcus for the capsuloparietal Early Pliocene of the North Atlantic (McLeod et al., 1993; emissary vein. In extant balaenopterids, this sulcus usually Westgate and Whitmore, 2002). There are very few confirmed courses posteriorly to a point level with the position of the pre-Quaternary fossil records of Eubalaena. Bisconti (2003, mallear fossa. However, in P218268 any more posterior con- 2005) referred the Pliocene Balaena belgica Abel, 1941 to tinuation of the sulcus for the capsuloparietal emissary vein, if Eubalaena belgica. McLeod and others (1993: 63) suggested formerly present, no longer occurs due to abrasion. that a balaenid periotic from the Early Pliocene of South The ventral presentation of the periotic exhibits several Australia (originally recorded by Howchin: 1919) could repre- features. As occurs in extant BalaenopteraandMegaptera, the sentEubalaena(as opposed to its original referral to Balaena). lateralmost eminence of the ventrolateral ridge of the superior Dixon (1990) described an incomplete Recent Eubalaena process (sensu Geisler and Luo, 1996) is situated at the same australisskeleton from Altona Bay, near Melbourne, Victoria. level as the anterior margin of the pars cochlearis. The mallear The latter specimen (C27879) includes tympanics and perio- fossa is poorly differentiated from the rest of the epitympanic tics. The extinct genera Balaenula and Balaenotus have been recess. recorded from the Late Miocene through Pliocene of the N The preserved posterolateral margin of the periotic is Pacific (Barnes, 1977; McLeod et al., 1993) and N Atlantic formed by the hiatus epitympanicus. The course of the facial (McLeod et al., 1993; Bisconti, 2003 and references therein). nerve on the ventral surface of the periotic is marked by the Recently, Bisconti (2005) described a new genus and species of facial sulcus which is bounded anteriorly by the aperture of the relatively small balaenid, Balaenella brachyrhynus, from the ventral facial foramen. In ventral view, a distinct bridge of Early Pliocene of Belgium. bone at the anterolateral corner of the preserved pars cochlearis The incompleteness of P218269 and lack of information on represents the ventral roof of the ventral facial foramen. The the extent of intraspecific and ontogenetic variation in balaenid endocranial aspect of the pars cochlearis preserves the aperture periotics, hampers comparisons with described extant and of the internal facial foramen. Anterior to this aperture is a deep fossil balaenid taxa. Furthermore, there are as yet no published excavation in the medial surface of the periotic at the base of criteria for discriminating between the periotics of Balaenaand the anterior process. This region (composed of cancellous bone Eubalaena. Despite these problems, it may be noted that in extant balaenopterids) marks the site of ankylosis between P218269 is similar in overall size to several isolated balaenid the anterior process and the body of the periotic. As in other periotics from the uppermost Miocene to Lower Pliocene Black Balaenopteridae, the pars cochlearis appears to have been Rock Sandstone and Grange Burn Formation of Victoria (e.g. elongated towards the cranial cavity. P16195, P48865, P160438, and P197824). The discovery of a Discussion. That P218268 is a balaenopterid periotic is evident more complete periotic (including the pars cochlearis) is neces- by the possession of: (1) elongated, triangular and anteriorly sary before any further comparisons between the Portland attenuated anterior process; (2) a triangular lateral eminence of Pliocene balaenid and the other Victorian specimens listed the ventrolateral ridge; and (3) a relatively large pars cochlearis above can be made. elongated towards the cranial cavity. Because P218268 is rep- resented only by an incomplete periotic, it is not possible for it FamilyBalaenopteridaeGray, 1864 to be identified below family level. The size of P218268 is Genus and species indeterminate comparable to that of periotics of subadult Balaenoptera edeni Anderson, 1879 or B. brydeiOlsen, 1913 (P171503) and juve- Referred specimen.NMVP218268, incomplete right periotic; lacking nile Megaptera novaeangliae Borowski, 1781 (C28892). medial three-quarters of pars cochlearis and posterior process However, the periotics of extant Megaptera novaeangliae and (Fig. 3B). an undescribed species of Megaptera from the Early Pliocene Description. P218268 is polished, rolled and may be secondar- of Victoria (P179005) possess the following features which ily phosphatised. The anterior process is elongated and some- differentiate them from P218268: (1) the anterior process is Pliocene marine mammals 73 AA C BB Figure 3. Mysticeti and Physeteridae from the Pliocene Whalers Bluff Formation, Portland. A, Balaenidae gen. et sp. indet., incomplete right peri- otic, NMVP218269, in ventrolateral view (AC);B, Balaenopteridae gen. et sp. indet., incomplete right periotic, NMVP218268, in ventral view (AC);C, cf. Physetersp., apical crown of tooth, NMVP218298, in side view (AC). Scale bars equal 10 mm. 74 Erich M.G. Fitzgerald relatively shorter; (2) anterior process is more dorsoventrally Discussion. P218407 is referred to the Kogiidae on the basis of compressed; (3) in endocranial view, there is an anteroposter- the following features shared with extant and fossil kogiids: (1) iorly thickened region of cancellous bone in the pars cochlearis small overall size [as Kasuya (1973: 25) noted among extant anterior to the internal facial foramen; and (4) no deep excava- Odontoceti only Pontoporia blainvilleiGervais and d’Orbigny, tion in the endocranial surface of the periotic anterior to the 1844 has a smaller tympanic, and P218407 does not resemble pars cochlearis. Many of these features may be related to onto- the tympanic of that taxon]; (2) distinct embayment in genetic variation. At least, the lack of these four features in the medial edge of the involucrum between the inner anterior P218268 may indicate that this periotic is not referrable to and posterior prominences; and (3) squared-off anterior edge Megaptera and may belong to an indeterminate species in the of the involucrum and outer lip. It should be noted that the genusBalaenoptera. second feature is also seen in the tympanic bullae of Physeter macrocephalusLinnaeus, 1758, Orycterocetus croco- SuborderOdontocetiFlower, 1867 dilinus Cope, 1868 (USNM 22953) (Kellogg, 1965) (Fig. 5C–D), and an Early Pliocene physeterid (USNM FamilyPhyseteridaeGray, 1821 183007) (Fig. 5A, B). Despite this similarity between P218407 cf.Physetersp. Linnaeus, 1758 and the tympanics of Physeteridae, the relatively large size of physeterid tympanics precludes P218407 from being con- Referred specimen. NMV P218298, isolated, worn-down apical sidered as a physeterid. Furthermore, the inner posterior region of tooth crown (Fig. 3C). prominence in physeterid tympanics is generally more pointed Description. P218298 has an ovoid cross-section at its base in outline than the rounded outer posterior prominences in which becomes more ellipsoid towards the preserved apex of kogiid tympanics. the crown. In occlusal view, the apex of the tooth crown is Among fossil Kogiidae, the tympanics of Praekogia incised by a deep anteroposterior cleft which is attributed to cedrosensis(Barnes, 1973b) have not been described. Askull of tooth wear. Shallow longitudinal grooves in the surface occur Scaphokogia cochlearis Muizon, 1988 (USNM 452993) on all faces of the crown. The basal end of the tooth is broken includes an associated incomplete left tympanic (Fig. 4E, F). to reveal a cross-section through the crown. The only notable The tympanic of S. cochlearis is similar to the tympanics of feature of this aspect of the tooth is the presence of thick layers extantKogiain its relatively small size and possession of a dis- of cementum. tinct embayment in the medial side of the involucrum between the inner anterior and posterior prominences. The most notable Discussion. The large size of this tooth and the thick layers difference between the tympanic of S. cochlearis and the of cementum suggest that P218298 represents a physeterid. Portland kogiid tympanic lies in the more marked inflation of Little more can be said about the systematics of this speci- the inner posterior prominence of P218407. S. cochlearispos- men although its size and similarity to teeth of large adult sesses a less expanded inner posterior prominence, such that Physeter macrocephalus suggest affinities with the extant the embayment in the medial face of the involucrum is not as sperm whale. deep as in P218407. In this respect, Scaphokogia cochlearisis similar to two undescribed Early Pliocene kogiids from the Lee FamilyKogiidae Gill, 1871 Creek Mine, North Carolina (USNM 183008, Fig. 4G–H; Genus and species indeterminate USNM 251118, Fig. 4C, D) and the extant Kogia breviceps Blainville, 1838 (Fig. 4I, J). Referred specimen. NMV P218407, incomplete left tympanic bulla The features which Kasuya (1973) used to differentiate (Fig. 4A-B). between Recent Kogia brevicepsandK. simaOwen, 1866 are Description. The most striking aspect of the morphology of not preserved in P218407. However, comparison between fig- P218407 is its small size. The tympanic is polished with rolled ures of the tympanic of K. sima(Kasuya, 1973: plate VIII), and edges being rounded off. The preserved portion includes only actual specimens of K. breviceps (C24972, C24976), indicate the medial half of the bulla with very little of the outer lip pres- that P218407 differs from both extant Kogia species in: (1) ent. The base of the posterior process of the tympanic has been embayment in medial edge of involucrum between the inner worn off. In dorsal and ventral view, there is a distinct furrow prominences is markedly deeper; and (2) the involucrum is less in the medial edge of the involucrum between the inner poster- dorsoventrally and mediolaterally inflated. Despite these dif- ior prominence and the inner anterior prominence. In ventral ferences, P218407 is almost identical in size to the tympanics view, the furrow between the prominences of the involucrum ofKogia breviceps. Given that the currently available evidence forms an obtuse angle. There is no preserved ventral keel and is meagre, P218407 is not identified below family level. Table the median furrow is very shallow such that it is poorly differ- 2 compares some dimensions of the tympanics of kogiids and entiated from the surrounding ventral surface of the tympanic. physeterids discussed above. The interprominential notch is relatively broad. The anterior P218407 is the first fossil record of Kogiidae from edge of the involucrum and outer lip is squared-off. In dor- Australia. Fossil kogiids have previously been reported in the sal view, the involucrum has a consistent width along its SW Pacific region, from the ?Late Miocene of the Chatham length, but is expanded at the level of the inner posterior Rise, east of New Zealand (Fordyce, 1984a) but that record has prominence. a poorly constrained age (Fordyce, 1989, 1991b). Pliocene marine mammals 75 Table 2. Measurements of tympanics of Physeteridae and Kogiidae. Measurements follow methodology of Kasuya (1973). All measurements are in mm. NMV NMV USNM USNM USNM USNM USNM C24972 P218407 22953 183007 452993 183008 251118 Standard length of tympanic bulla, distance from anterior tip to posterior end of outer posterior prominence 26.0 – 31.0 42 – 29.0 31 Distance from anterior tip to posterior end of inner posterior prominence 19.9 21.6 27.0 38 28 31.0 28 Distance from posteroventral tip of outer posterior prominence to tip of sigmoid process 21.28 – – – – – – Distance from posteroventral tip of outer posterior prominence to tip of conical process 15.3 – – – – – – Width of tympanic bulla at the level of the sigmoid process 26.0 – – – – – – Height of tympanic bulla, from tip of sigmoid process to ventral keel 26.5 – – – – – – Width across inner and outer posterior prominences 19.0 – 20.2 24 – 22.5 +20+ SuperfamilyDelphinoideaGray, 1821 Perrin, 1994; LeDuc et al., 1999) and that some of the pre- sumed ancestors of Delphinidae, the Kentriodontidae (Barnes, Incertae sedis 1978, 2002; LeDuc, 2002), possessed Steno-like crown orna- Referred specimens.NMVP218283, P218284 and P218286, all isolat- mentation (e.g., Kellogg, 1966), the anastomosing striae on the ed teeth (not figured). crown of P13033 (and P218283, P218284, P218286) are of dubious use in assessing the phylogenetic affinities of isolated Description. P21283, P218284 and P218286 all represent small teeth. Furthermore, non-delphinid small odontocetes such as odontocete teeth possessing conical enamel-covered crowns Lipotes vexillifer Miller, 1918 possess anastomosing wrinkling that bear fine wrinkling ornamentation, and curve lingually on tooth crown enamel (Miller, 1918; Brownell and Herald, towards the crown apex. As in kentriodontids, there is a lingual 1972; Barnes, 1985) which casts doubt on any perceived bulge at the base of the crown but this feature is not as promi- taxonomic or phylogenetic signal present in these teeth. nent in the teeth from Portland. None possesses an open pulp The isolated teeth from Portland are assigned to cavity suggesting that all were derived from adult individuals. Delphinoidea incertae sedis. None of the Portland Pliocene P218283 is an incomplete tooth, its preserved maximum teeth share demonstrably close affinities with the holotype length and maximum width of the crown being 16 mm and 6 tooth of “Steno”cudmorei. mm respectively. Due to the incomplete nature of this tooth it does not warrant further description. P218284 is the most highly polished and the most complete. FamilyDelphinidaeGray, 1821 It differs from the others in having a mediolaterally compressed cf.Tursiopssp. Gervais, 1855 sp. root with a more prominent mesial-distal bulge at its midpoint. The preserved apex of the root curves posteriorly. Referred specimen. NMV P218266, virtually complete right periotic The most notable feature of P218286 distinguishing it from (Fig. 6). the other teeth is its bulbous root, which contrasts with the Description. P218266 is typically delphinid in possessing: (1) transversely flattened morphology of P218284. posterior process of the periotic projects laterally and postero- Discussion. Only one delphinoid odontocete has previously laterally; (2) longitudinal grooves on the articular facet of the been described from the Tertiary of Australia, the latest posterior process of the tympanic (this feature also occurs in Miocene-earliest Pliocene “Steno” cudmorei Chapman (1917) Monodontidae); (3) relatively low crista transversa; (4) internal from the Black Rock Sandstone of Beaumaris, Victoria facial foramen opens at the same level as the tractus spiralis (Fitzgerald, 2004b). Fordyce (1982) questioned the taxonomic foraminosus in the internal acoustic meatus; (5) a short, blunt, identity of “S.”cudmorei(the holotype, P13033, being an iso- rectangular anterior process of the periotic which in anterior, lated tooth) and Fitzgerald (2004b) referred “S.” cudmorei to dorsal and ventral views appears laterally compressed; (6) a Delphinidae, genus and species indeterminate. Chapman large fovea epitubaria for the accessory ossicle eliminates the (1917) assigned P13033 to Steno on the basis of the resem- anterior bullar facet on the anterior process; (7) prominent blance of its crown ornamentation to that seen in the teeth of parabullary ridge; (8) inflated pars cochlearis; and (9) rela- the extant Steno bredanensis Cuvier in Lesson, 1828 (e.g., tively shallow internal acoustic meatus (Kasuya, 1973; Fordyce Miyazaki and Perrin, 1994). Given that Stenois probably in a et al., 2002). The first character is usually only seen in basal position in the phylogeny of Delphinidae (Miyazaki and Delphinidae but several taxa in the extinct delphinoid grade- 76 Erich M.G. Fitzgerald A B C D E F G H I J Figure 4. Miocene to Recent Kogiidae tympanics. A-B, Kogiidae gen. et sp. indet. (Pliocene Whalers Bluff Formation, Portland, Victoria, Australia), incomplete left tympanic, NMVP218407 (AC).C-D, Kogiidae gen. et sp. undet. (Lower Pliocene Yorktown Formation, Lee Creek Mine, North Carolina, U.S.A.), incomplete left tympanic, USNM 251118. E-F,Scaphokogia cochlearis(Upper Miocene Pisco Formation, Aguada de Lomas level, Arequipa Department, Peru), incomplete left tympanic, USNM 452993. G-H, Kogiidae gen. et sp. undet. (Lower Pliocene Yorktown Formation, Lee Creek Mine, North Carolina, U.S.A.), incomplete right tympanic, USNM 183008. I-J,Kogia breviceps(Recent, Shelley Beach, Victoria, Australia), incomplete left tympanic, NMVC24976. A,C,E,G,I, all in dorsal view. B,D,F,H,J,all in ventral view. Scale bars equal 10 mm.