Klaas Post 1, Olivier Lambert 2 & Giovanni Bianucci 3 1 Natuurhistorisch Museum Rotterdam 2 Institut royal des Sciences naturelles de Belgique 3 Dipartimento di Scienze della Terra, Università di Pisa First record of Tusciziphius crispus (, Ziphiidae) from the Neogene of the US east coast

Post, K., Lambert, O. & Bianucci, G., 2008 - First record of Tusciziphius crispus (Cetacea, Ziphiidae) from the Neogene of the US east coast - DEINSEA 12: 1 - 10 [ISSN 0923-9308]. Published online 10 January 2008.

The genus Tusciziphius is represented by a single , which was previously only known from the holotype skull from the Early Pliocene of Tuscany, Italy. This article reports the second specimen of this species from Miocene/Pliocene strata of South Carolina (USA, east coast), kept in the Natuurhistorisch Museum Rotterdam. The specimen provides information on features not preserved on the holotype skull and reveals that the species was not endemic to the Mediterranean but must also have roamed the North Atlantic realm.

Correspondence: Klaas Post (corresponding author), Natuurhistorisch Museum Rotterdam, P.O. Box 23452, 3001 KL Rotterdam, the Netherlands, e-mail [email protected]; Olivier Lambert, Institut royal des Sciences naturelles de Belgique, Département de Paléontologie, Rue Vautier, 29, B-1000 Brussels, Belgium; e-mail [email protected]; Giovanni Bianucci, Dipartimento di Scienze della Terra, Università di Pisa, Via S. Maria, 53, I-56126 Pisa, Italy; e-mail [email protected]

Keywords: Tusciziphius crispus, Cetacea, Ziphiidae, Neogene, USA

INTRODUCTION thus saving it from oblivion. The cranium is In his revision of the fossil beaked whales of described here, assigned to Tusciziphius cris- the Italian Pliocene, Bianucci (1997) realised pus, and compared to other Neogene ziphiids. the unique features of a fossil neurocranium in the collection of the University of Florence METHODS and redescribed it as a new genus and species named Tusciziphius crispus. The holotype cra- Terminology and measurements nium had already been found during the 19th Anatomical terms are taken from Heyning century and was identified by Capellini (1885) (1989) and Bianucci et al. (2007). as Choneziphius planirostris (CUVIER, 1823). Measurements were taken following directions Since 1885, neither in Italy nor in the rest of of Ross (1984), Bianucci (1997), Lambert the world, another fossil of Tusciziphius was (2005) and Bianucci et al. (2007). found or recognised. In 2002 a fossil ziphiid skull from South Carolina was publicly sold by Institutional abbreviations the relatives of an American collector. The first IGF: Museo di Geologia e Paleontologia author realised the importance of the specimen dell’Università di Firenze, Florence, Italy. and could arrange that the fossil was donated NMB: Natuurhistorisch Museum Boekenberg, to the Natuurhistorisch Museum Rotterdam, Antwerp, Belgium. NMR: Natuurhistorisch

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Figure 1 Skull of Tusciziphius crispus (NMR 9991-3020): A dorsal view; B corresponding line drawing.

2 POST et al.: Tusciziphius crispus from the US east coast

Museum Rotterdam, Rotterdam, The Supraoccipital, posterior to the vertex, mod- Netherlands. NNM: Naturalis Nationaal erately constricted between the two maxillae. Natuurhistorisch Museum, Leiden, The Large temporal fossa and relatively elevated Netherlands. lambdoidal crest.

SYSTEMATIC PALAEONTOLOGY DESCRIPTION AND COMPARISON While the holotype (Fig. 5) is a fairly complete Class Mammalia LINNAEUS, 1758 neurocranium without rostrum, NMR 9991- Order Cetacea BRISSON, 1762 3020 includes the rostrum, but misses most of Suborder Odontoceti FLOWER, 1867 the basicranium. At first sight NMR 9991-3020 Family Ziphiidae GRAY, 1865 seems smaller and less robust than the holo- Genus Tusciziphius BIANUCCI, 1997 type but observations and measurements (Table 1) confirm that this impression is only caused Type and only included species by the lack of the massive basicranium. Tusciziphius crispus BIANUCCI, 1997 As in the holotype, the vertex is anteriorly curved and covers - in dorsal view - the bony Holotype IGF-1594 V, incomplete cranium nares. Premaxillary crests, nasals and the pre- from Val di Pugna, Tuscany, Italy. served parts of the frontals of NMR 9991-3020 show the same general morphology as the hol- Referred specimen NMR 9991-3020, from otype but are completely fused (even more so South Carolina, USA; incomplete cranium than the holotype) to the extent that sutures are with rostrum, vertex and facial structures, but hardly or not visible. This extent of complete without basicranium, supraoccipital and man- co-ossifying is not seen in other Ziphiidae and dible (Figs. 1-4). might represent an autapomorphy of the genus. The vertex is moderately elevated, somewhat Emended diagnosis Skull equal in size lesser than in fossil Caviziphius BIANUCCI to extant Ziphius cavirostris. Rostrum very & POST, 2005 and Choneziphius, and by no dense and relatively short as in Choneziphius means as elevated as in extant Ziphius. The planirostris. Premaxillae on rostrum elevated, premaxillary crests are very robust, the left forming a dome near the rostral base and com- crest is smaller than the right one. The right pletely fused medially, closing the mesorostral premaxillary crest shows a large, semi-circular, groove. Broad and massive base of rostrum. partly flat, dorsal surface with a highest point Palatine extended anterior to the mid-length towards the right nasal. The left premaxillary of the rostrum. Premaxillary crests and sac crest shows a long wavy-like dorsal surface. fossae very asymmetrical (right side larg- The extremely long and slender nasals are est). Ascending process of the premaxilla somewhat oriented to the left (anteroposteri- anteriorly curved near the vertex, causing the orly) and laterally bordered by the premaxil- vertex to overhang the bony nares (in dorsal lary crests along their entire length, as in the view). Vertex elevated as in Choneziphius but holotype. In lateral view the nasals are not less than in Ziphius. Extreme ossification and extending in front of the premaxillae. In fact fusion of vertex elements. Premaxillary crests, the nasals seem reduced and depressed at especially the right one, extremely transver- their anterior midpoint (a condition seen in sally expanded, but not posteriorly curved Hyperoodon, Mesoplodon and their fossil rela- as in Mesoplodon and Hyperoodon. Nasals tives), but it is difficult to confirm this obser- large, antero-posteriorly elongated and later- vation on the holotype because of a crack. ally shifted to the left side of the skull. Nasals The frontals are preserved on the vertex and not protruding anteriorly past the premaxillary morphology and size are as in the holotype. crests. Frontals hardly exposed on the vertex. The vertex is most constricted (in dorsal view)

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Figure 2 Skull of Tusciziphius crispus (NMR 9991-3020): A (left) lateral view; B corresponding line drawing; C (right) lateral view.

4 POST et al.: Tusciziphius crispus from the US east coast

at the point where nasals and frontals meet. In attachment for facial muscles (Lambert 2005), anterior view the degree of asymmetry of the or might have been caused by blood vessels. vertex of Tusciziphius is striking and expressed A remarkable asymmetry is shown by the by the very wide, high and robust right pre- positions of the two anterior most foramina. maxillary crest. The right foramen is located just 11 mm from The premaxillary sac fossae show the same the medial maxillary foramen near the border architecture and asymmetry as in the fossil of the right premaxillary sac fossa, while the genera Caviziphius and Choneziphius, however left-one is located at a c. 40 mm more anterior the right fossa of Tusciziphius is not excavated position near the rostrum base. Maxillary and in a similar way as in these last two genera premaxillary foramina cannot be observed on (and especially in Caviziphius) but posteriorly the holotype due to bad preservation. On the excavated as well as anteriorly and antero- supraoccipital process, marked maxillary crests medially thickened and elevated. Contrary to towards the base of the rostrum - such as seen Beneziphius LAMBERT, 2005, Messapicetus in Choneziphius and Ziphius - are not present. BIANUCCI, LANDINI & VAROLA, 1992 and The rostrum base is fairly wide (Table 1) and Ziphirostrum DU BUS, 1868, no prenarial basin shows, at the lateral borders, a slight constric- is present in front of the prenarial sac fossae. tion (obviously a remnant of the antorbital Even a shallow basin, as in Choneziphius, is notch - see Figure 1). The dorsal maxillary sur- absent. The premaxillary foramina are difficult face at the rostrum base is slightly convex and to observe, the left foramen seems retained in highest at the border. At this level the joined the anterior corner of the deep premaxillary premaxillae are elevated to a bulbous 35 mm sac fossa, while the right foramen is absent (as high and 70 mm long dome, which is show- seems the case in Eboroziphius LEIDY, 1876). ing the rough surface as described earlier. This The premaxillae are completed fused immedi- condition seems also present in Eboroziphius. ately anterior to the premaxillary sac fossae. A similar but non-homologous feature (of This condition is also noted in Caviziphius, the ossified vomer) is present in Mesoplodon Eboroziphius, unnamed Belgian specimens tumidirostris (Miyazaki & Hasegawa 1992) and in Pelycorhamphus COPE, 1895 (Lambert and is observed in undescribed Mesoplodon- 2005). The fusion of the premaxillae extends like fossil rostra from the South African coast over the entire length of the rostrum till just (pers. obs.). More anteriorly the dorsal surface a few centimetres in front of the apex and a of the maxilla becomes flat to, near the apex, medial suture between the premaxillae is not concave. The fused premaxillae keep their visible. Creation of a mesorostral tunnel by the elevated ridge-like shape towards the apex joined and fused premaxillae is also noted in and are firmly pachyostosed, a condition also Caviziphius and Choneziphius, but in the last present in Caviziphius and Choneziphius. In genus a suture remains visible. lateral view the rostrum shows a blunt out- The antorbital notch area is not well pre- line (again reminding the general shape of served (as in the holotype). Six maxillary Choneziphius rostra), while the sharp border foramina are present posteriorly to the ros- at the rostrum base gradually eases out into a trum base and are surrounded by ridge-like ridge towards the apex. Remnants of a non- structures. These excrescences are also visible functional and shallow alveolar groove run on other parts of the dorsal surface of NMR below this line towards the apex. 9991-3020, mostly on the median premaxil- It seems that the apex is slightly restored lary dome and the anterior margin of each by the use of some sort of consolidate. The premaxillary sac fossa, and seem of the same eventual original damage, however, must have nature as is often noticed at or near the base of been marginal because the restoration surface the rostrum of Beneziphius and Choneziphius. is minor. The incomplete apex shows - in ante- These surface structures may have served as rior view - the large circular opening of the

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Figure 3 Skull of Tusciziphius crispus (NMR 9991-3020): A anterior view; B corresponding line drawing.

mesorostral tunnel, and - ventrally - two small The Morgan River, which runs between the foramina. Whether or not the dorsal fusion mentioned coordinates, is also a rich source of both premaxillae might have continued till of fossils, collected by divers. The Charleston the original apex, or might have stopped just Museum collections contain a rostrum of before, is not clear. Eurhinodelphinidae and teeth of Carcharodon The ventral part of the rostrum base is dam- megalodon collected from the bottom of this aged and important observations cannot be river, indicating either sub cropping of Middle- noted. However, laterally the anterior border of and/or Late Miocene strata, or the presence of the palatine was extended to at least 130 mm reworked Miocene strata in the area (pers.com. anterior from the rostrum base. The sutures of A. E. Sanders). Sanders (2002) reported Goose the pterygoid attachment are not clearly detect- Creek Limestone adherent to the holotype of able, but leave the possibility open that ptery- Ceterhinops longifrons (one of the nominal goid (and palatine) covered the ventral walls of odontocete taxa from South Carolina described the rostrum to a more anterior position than in by Leidy in 1877). Caviziphius and Choneziphius. Taking into account the absence of a more exact locality of the described cranium, the STRATIGRAPHIC ASSIGNMENT The skull was found by the late Vito Bertucci during 1999 in the Morgan River, Beaufort County, South Carolina, USA, at a loca- tion between 32º26'50"N, 80º35'57"W and 32º27'09"N, 80º28'44"W. These coordinates and the phosphatised condition of the fos- sil seem to hint to the ‘locally abundant Phosphate Beds’ which were mentioned in ancient literature and considered to be of Late Miocene origin (Leidy 1877). Recent stud- ies note sub-cropping of a small patch of the Early-Middle Pliocene Goose Creek Limestone at the indicated coordinates, surrounded by Early Miocene and Oligocene strata (Weems & Lewis 2002). The Goose Creek Limestone consists of quartzose and phosphatic cal- carenite with a base often marked by a lag Figure 4 Skull of Tusciziphius crispus (NMR 9991-3020): of phosphate pebbles (Weems et al. 1982). detail of vertex.

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Figure 5 Tusciziphius crispus (holotype, IGF 1594 V): A anterior view; B corresponding line drawing. presence of Miocene and Pliocene strata in the extreme development of the right premaxillary area, and the fact that advanced ziphiids are crest is seen in the first two genera, but lacks only known from the Late Miocene onward, in Choneziphius and Ziphius. it seems prudent to assume a Late Miocene / Extant Ziphius - on the other hand - shows Middle Pliocene background for NMR 9991- essential differences from the three fossil 3020. The holotype (IGF-1594 V) originates forms in the architecture of the rostrum (ossi- from Early Pliocene strata (Bianucci et al. fied vomer as in several Mesoplodon species), 2001). the more elevated vertex, and a vertical and rectilinear outline of the lateral margins of DISCUSSION AND PHYLOGENETIC the ascending processes of the premaxilla in OBSERVATIONS anterior view (the premaxillary crest is not dis- Tusciziphius crispus is closely related to tinctly wider than the ascending processes of the North Atlantic fossil ziphiid species the premaxilla). Caviziphius altirostris and Choneziphius NMR 9991-3020 is - at present - the best planirostris by several characters such as the preserved fossil ziphiid specimen from the East short and blunt rostrum, the rostral fusion of Coast of North America. The many nominal the premaxillae (closing dorsally the mesoros- ziphiid species from South Carolina described tral groove), the extreme rostral pachyostosis by E.D. Cope in 1869 (Cope 1869a,b) and without mesorostral ossification of vomer and/ J. Leidy in 1877 (Leidy 1877), are based on or mesethmoid, the architecture and extreme fragmentary specimens and do not resemble, or asymmetry of the premaxillary sac fossae, and cannot be compared to Tuscizphius. However the moderately high vertex. Some features of Eboroziphius coelops LEIDY, 1877 and Caviziphius and Tusciziphius such as extreme Pelycorhamphus pertortus COPE, 1895 deserve lateral development of the premaxillary crests, special attention. Both species are only known complete fusion of the vertex, elevation of the by their holotype: the former is a rostral frag- anterior margin of the premaxillary sac fossa, ment known from the Phosphate Beds of South and complete fusion of premaxillae on dorsal Carolina and the latter is a very fragmented surface of the rostrum, seem more derived than specimen from the Miocene of the Chesapeake in Choneziphius. Group (East Coast USA). The preserved parts Tusciziphius and Caviziphius also share of these rostral fragments clearly show simi- an anteriorly projecting vertex overhanging larities with Caviziphius and Tusciziphius at the bony nares, which is also noted in extant the level of the premaxillary sac fossae and Ziphius but lacks in Choneziphius. Moreover fused premaxillae lacking a prenarial basin

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Table 1 Measurements (in mm) of the two known fossil skulls of Tusciziphius crispus: IGF 1594 V (holotype) and NMR 9991-3020; e = estimation ______

measured unit IGF 1594 V NMR 9991-3020 ______

Greatest postorbital width 415 >372 Width of premaxillae at base of rostrum - 86 Greatest width between zygomatic processes of squamosals 450 - Distance from base of rostrum to anterior tip of palatine - e130 Height of skull between pterygoids and vertex 339 - Length of temporal fossa 159 - Height of temporal fossa 88 - Width of cranium between posterior borders of temporal fossa 214 - Width of condyles 128 - Width of foramen magnum 46 - Height of right condyle 69 - Width of right condyle 46 - Minimal posterior distance between maxillae on the vertex - 84 Transverse width of premaxillary crests 192 e184 Minimal distance between premaxillary crests 52 e39 Width of bony nares - 69 Width of premaxillary sac fossae - 173 Width of right premaxillary sac fossa - 108 Width of left premaxillary sac fossa - 51 Length of medial suture between nasals 70 68 Greatest width of nasals 64 - Greatest width of right nasal - e27 Length of right nasal 90 82 Width of right premaxillary crest - >101 Width of left premaxillary crest - 51 Minimal width of right premaxillary dorsal to the premaxillary sac fossa - 68 ______

(see description and comparison). However the aim and reach of this article. The study on the deep right premaxillary sac fossa observed eight newly described genera in these specimens link them to Caviziphius from South Africa (Bianucci et al. 2007), and rather than to Tusciziphius. Unfortunately, more future works on newly discovered, well pre- in-depth observations are hindered by the fact served ziphiids from Peru by same the same that Eboroziphius coelops and Pelycorhamphus authors, will review the present status of fossil pertortus are based on too fragmentary speci- ziphiids in general, and might unravel the com- mens with few diagnostic features, and - in fact plicated phylogeny of this diversified toothed - must be considered nomina nuda. whale family. Based on dimensions and morphology of the rostrum, the unnamed isolated rostrum NMB CONCLUSIONS 002 from the North Sea - which was compared Tusciziphius crispus is a Late Miocene- to Eboroziphius by Lambert (2005) - might be Middle Pliocene beaked whale and its dis congeneric with Caviziphius. tribution is noted on both sides of the North All observations do confirm a close relation- Atlantic realm (the Mediterranean and the ship between Caviziphius and Tusciziphius. east coast of the USA). Tusciziphius shares A detailed phylogenetic analysis of above- this transatlantic distribution with Choneziphius mentioned partly (or seemingly) contradicting (Lambert 2005) and Messapicetus (Fuller & observations and a comparison of characters of Godfrey 2005, 2007). the known fossil and extant ziphiids is beyond  NMR 9991-3020 adds new data to the mor-

8 POST et al.al.:: Tusciziphiusmute swan crispusbiometrics from the US east coast

phology of the genus: size and architecture of tion on essential morphological features (apex the rostrum, median fusion of the premaxillae, of rostrum, prominental-antorbital notch, median fusion and elevation of the premax- petro-tympanic complex, post-crania) and illae, extension of the palatine to the ventral geographic distribution. side of the rostrum, position of the maxillary foramina, ossification and fusion of the vertex ACKNOWLEDGEMENTS elements. Henry van der Es (NMR) and John de Vos  Tusciziphius seems closely related to two (NNM) are thanked for access to fossils stored other Late Miocene-Early Pliocene ziphiids under their care. Jonathan H. Geisler (Georgia from the North Atlantic (Choneziphius and Southern Museum) and Albert E. Sanders especially Caviziphius), and is - in some (Charleston Museum) kindly provided litera- characters - more derived than both of them. ture and comments on the geology of Tertiary However their essential phylogenetic relation deposits from South Carolina. Michelangelo and the relationships to other ziphiids (fossil Bisconti (Natural History Museum of Livorno) and extant) are still not clear. assisted in many ways and Mark Broch helped The species is known from two fossils only; in textual matters. new fossils might provide additional informa-

REFERENCES Cope, E.D., 1869b - On two new genera of extinct Bianucci, G., 1997 - The odontoceti (Mammalia Cetacea) Cetacea - The American Naturalist 3(8): 444-445 from Italian Pliocene. The Ziphiidae - Palaeonto- Cuvier, G., 1823 - Recherches sur les ossements fossiles, graphia Italica 83: 163-192 5 (1ère partie) - G. Dufour et E. D’Ocagne, Paris, Bianucci, G., Lambert, O. & Post, K., 2007 - A high pp 405 diversity in fossil beaked whales (Odontoceti, du Bus, B.A.L., 1868 - Sur différents Ziphiides nouveaux Ziphiidae) recovered by trawling from the sea floor du Crag d’Anvers - Bulletin de l’Académie Royale off South Africa - Geodiversitas 29(4): 5-62 des Sciences de Belgique 25: 621-630 Bianucci, G., Landini, W. & Varola A., 1992 - Fuller, A.J. & Godfrey, S.J., 2005 - Miocene ziphiids Messapicetus longirostris, a new genus and spe (Cetacea: Odontoceti) from the Chesapeake Group cies of Ziphiidae (Cetacea) from the Late Miocene of of Calvert Cliffs, Maryland, USA - Evolution of ‘Pietro Leccese’ (Apulia, Italy) - Bolletino della aquatic tetrapods. Fourth Triannual Convention Società Paleontologica Italiana 31(2): 261-264 Abstracts 1: 26 Bianucci, G., Mazza, P., Merola, D., Sarti, G. & Cascella Fuller, A.J. & Godfrey, S.J., 2007 - A Late Miocene A., 2001 - The Early Pliocene assemblage ziphiid (Messapicetus sp.: Odontoceti: Cetacea) from of Val di Pugna (Tuscany, Italy) in the light of calca the St. Marys Formation of Calvert Cliffs, Maryland reous plankton biostratigraphical data and paleoeco- - Journal of Vertebrate Paleontology 27(2): 535-540 logical observations - Rivista Italiana di Paleonto- Heyning, J.E., 1989 - Comparative facial anatomy of logica e Stratigrafia 107(3): 425-438 beaked whales (Ziphiidae) and a systematic revision Biannuci, G. & Post, K., 2005 - Caviziphius altirostris, a among the families of extant Odontoceti - Contribu- new beaked whale from the Miocene southern North tions in Science, Nat. Mus. LA County 405: 1-63 Sea basin - Deinsea 11: 1-6 Lambert, O., 2005 - Systematics and phylogeny of the Capellini, G., 1885 - Del Zifioide fossile (Choneziphius fossil beaked whales Ziphirostrum du Bus, 1868 and planirostris) scoperto nelle sabbie plioceniche di Choneziphius Duvernoy, 1851 (Cetacea, Odontoceti), Fangonero presso Siena - Atti della Reale Accademia from the Neogene of Antwerp (North of Belgium) - dei Lincei. Memorie 8: 205-210 Geodiversitas 27(3): 443-497 Cope, E.D., 1869a - On two extinct Mammalia from Leidy, J., 1876 - Remarks on fossils from the Ashley the United States - Proceedings of the American Phosphate Beds - Proceedings of the Academy of Philosophical Society 11(82): 188-190 Natural Sciences of Philadelphia 1876: 80-81, 86-87

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Leidy, J., 1877 - Description of vertebrate remains, chie- and Georgia - American Philosophical Society, 152 p. fly from the Phosphate Beds of South Carolina. Weems, R.E., Lemon, E.M., Jr., McCartan, L., Bybell, Journal of the Academy of Natural Sciences of L.M. & Sanders, A.E., 1982 - Recognition and for Philadelphia 2:(8): 209-261 malization of the Pliocene “Goose Creek phase” in Miyazaki, N. & Hasegawa, Y., 1992 - A new species of the Charleston, South Carolina area, Stratigraphic fossil beaked whale, Mesoplodon tumidirostris sp. notes, 1980-1982 - United States Geological Survey nov. (Cetacea, Ziphiidae) from the Central North Bulletin, 1529-H: H137-H148 Pacific - Bulletin of the National Science Museum, Weems, R.E. & Lewis, W.C., 2002 - Structural and Tokyo, Ser. A, 18:(4) 167-174 tectonic stetting of Charleston, South Carolina, Ross, G.J.B., 1984 - The smaller cetaceans of the south region: Evidence from the tertiary stratigraphic record east coast of southern Africa - Annals of the Cape - Bulletin of the Geological Society of America Provincial Museums of Natural History 15:(2) 173- 104(1): 24-42 410 Sanders, A.E., 2002 - Additions to the Pleistocene Received 2 June 2007 mammal faunas of South Carolina, North Carolina Accepted 3 September 2007

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