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Fossil sperm whales (Cetacea, Physeteridae) from Gunma and Ibaraki prefectures, Japan; with observations on the Miocene fossil sperm whale Scaldicetus shigensis Hirota and Barnes, 1995
Abstract
Fossil specimens of sperm whales (members of the odontocete cetacean family Physeteridae) are rare in Japan, and this is true also in other parts of the world. Two physeterid fossils have been previously reported from Middle Miocene age deposits of Gunma and Ibaraki prefectures, Japan, and these specimens are re-identified here as cf. Scaldicetus sp. The morphology of the teeth of these specimens is consistent with the traditional concept of the genus Scaldicetus. However, because the genus Scaldicetus is a form taxon, based solely on isolated teeth, it is not appropriate to assign more completely known fossil sperm whales, several of which have other characters that are distinctive at the generic level, to the genus Scaldicetus. Thus, the Japanese Middle Miocene sperm whale Scaldicetus shigensis Hirota and Barnes, 1995, is here assigned to the new genus Brygmophyseter. We include here additional observations and illustrations of the holotype skeleton of Brygmophyseter shigensis.
... The stem physeteroid genus Scaldicetus du Bus, 1867 is mainly based on isolated teeth, which is considered material of limited diagnostic value (Bianucci & Landini, 2006;Toscano et al., 2013;Marra et al., 2016;Lambert et al., 2017;Reumer et al., 2017;Bosselaers & Van Nieulande, 2018). However, studies on physeteroid dentition of the last three decades do show characterizing features in size and morphology for different physeteroid species (Hirota & Barnes, 1994;Kazár, 2002;Bianucci et al., 2004;Bianucci & Landini, 2006;Kimura et al., 2006;Bloodworth & Odell, 2008;Lambert, 2008;Boersma & Pyenson, 2015;Lambert et al., 2017;Collareta et al., 2019;Benites-Palomino et al., 2020;Lambert et al., 2020;Kimura & Hasegawa, 2022;Peri et al., 2022). The genus Scaldicetus was introduced by du Bus (1867) based on 45 large physeteroid teeth with rugose enamel-capped crowns from the Upper Miocene of Borgerhout, Antwerp, Belgium. ...
... In line with the conclusions of Bianucci & Landini (2006) and Lambert & Bianucci (2019), former Scaldicetus species outside of the type material of Scaldicetus caretti solely defined by large isolated teeth are referred to as Physeteroidea indet. The specimens described by Hasegawa et al. (2001), Estevens & Antunes (2004), Kimura et al. (2006) and Toscano et al. (2013) all have been assigned to Scaldicetus, because these authors consider Scaldicetus to be a wastebasket genus for large enamel-capped physeteroid teeth or other less complete physeteroid fossils such as dentaries. We would suggest to formally refer to these and future isolated dental specimens as Physeteroidea indet. ...
... We would suggest to formally refer to these and future isolated dental specimens as Physeteroidea indet. Kimura et al. (2006) wrote in their paper: 'The size and general shape of the teeth GMNH-PV-581 and INM-4-012885 are also similar to the teeth of the holotype of B. shigensis, and there are no substantial morphological differences between all of these specimens ' (p. 8). ...
Here we report a well-preserved isolated physeteroid tooth of Late Miocene age from Liessel, the Netherlands. The presence of several morphological features allows attribution to the macroraptorial physeteroids. Size and morphology are to some extent comparable to Zygophyseter and almost identical to the primarily tooth-based Tortonian taxon Scaldicetus caretti. However, the genus Scaldicetus was declared unutilizable, which is supported here with an overview of modern classifications of Scaldicetus species and specimens. Despite the restrictions, the type species S. caretti is still valid, although the name is to be restricted to the type material. Based on its morphological resemblance, the tooth is identified as Physeteroidea indet. cf. Scaldicetus caretti.
... Deep water interchange between the Caribbean Sea and the Pacific Ocean had already been interrupted during the Middle to early Late Miocene and only a shallow connection between both sides of the Isthmus remained [12,[29][30][31][32][33]. Such events not only triggered speciation events in the Caribbean and Pacific or migrations in the terrestrial biota [31,33], but also affected oceanic waters and marine currents across the globe [34,35]. ...
... There are four post-symphyseal alveolar teeth on the right ramus and four preserved symphyseal teeth on the left one. As in Acrophyseter sp., the convex ventral region of the mandible is enhanced by the bow-like lateral aspect of it, thus contrasting the more linear profile observed in the mandible of Zygophyseter and Brygmophyseter [35]. The external surface of the left ramus displays many borings and marking that resemble those left by Osedax-like worms (electronic supplementary material, figure S2). ...
Fossil cetaceans are often found in Miocene marine outcrops across the globe. However, because this record is not homogeneous, the dissimilar increase in occurrences, along with the sampling bias has created regions with extensive records and others with great scarcity. Among these, the Caribbean has remained enigmatic due to the lack of well-preserved cetacean fossils. Here, we report new Caribbean fossil cetaceans from the Upper Miocene Chagres Formation exposed along Piña beach, Eastern Panama, including a scaphokogiine kogiid, an Acrophyseter-like physeteroid and the phocoenid Piscolithax. Along with previous records of the iniid Isthminia panamensis and the kogiine Nanokogia isthmia, the Chagres cetacean fauna shows some similarities with other Late Miocene cetacean communities such as the Californias in the North Pacific, although their closest affinities lie with the eastern South Pacific Pisco Formation, Peru. Such findings indicate that though deep and intermediate Caribbean-Pacific water interchange was reduced by the Middle Miocene due to the shallowing of the Central American Seaway, shallow waters marine connection that persisted until the Pliocene might have facilitated the dispersal of coastal species across both sides of the Isthmus.
... Furthermore, the teeth of these fossil sperm whales exhibit deep occlusal facets, which are sulci on the tooth surface produced by repeated tooth-totooth contacts (attritional wear), and fractures attributed to strong occlusion or to the contact with hard material (e.g., bone) (Bianucci and Landini 2006;Lambert et al. 2017;Lambert and Bianucci 2019;Peri et al. 2020). Taxonomically diagnostic fossil remains of macroraptorial physeteroids have been retrieved in several localities all around the world, both in the middle Miocene (Albicetus oxymycterus, California, USA; Brygmophyseter shigensis, Japan) and in the upper Miocene (Acrophyseter deinodon, Acrophyseter robustus and Livyatan melvillei, Peru; Zygophyseter varolai, Italy) (Bianucci and Landini 2006;Kimura et al. 2006;Lambert et al. 2008Lambert et al. , 2010Lambert et al. , 2017Boersma and Pyenson 2015). One of the best known macroraptorial physeteroids, Z. varolai from Tortonian strata of the Pietra leccese formation (Salento Peninsula, southern Italy), is characterised by having a dorsal concavity on the skull (i.e., supracranial basin) that is wide and hemispherical, an extremely elongated zygomatic process (probably supporting a developed masseter muscle), and several dental features associated to a strong occlusion and repetitive use of the bite (e.g. ...
... The results obtained in this study need to be framed in the context of the complex ecology of Miocene seas. Indeed, besides a number of genera and species that are based on taxonomically diagnostic skeletal materials, isolated teeth referable to macroraptorial physeteroids have been reported from middle and upper Miocene deposits of several localities of the Americas, Asia and Europe (e.g., Kimura et al. 2006;Hasegawa et al. 2006;Marra et al. 2016;Reumer et al. 2017;Piazza et al. 2018;Lambert and Bianucci 2019). As already mentioned, the chronostratigraphic and geographic distribution of the published remains of macroraptorial sperm whales suggests that this group impressively radiated during the middle and late Miocene (Lambert et al. 2017). ...
Differing from the extant physeteroids, macroraptorial sperm whales are currently regarded as apex predators of the Miocene seas based on several morphofunctional observations. Here, we estimate the bite force of Zygophyseter varolai, a macroraptorial physeteroid from lower upper Miocene strata of the Pietra leccese formation (Apulia, Italy) using the finite element analysis (FEA). To explore multiple bite scenarios, we set four different load cases on a 3D model of the cranium obtained via digital photogram-metry, considering the temporalis and masseter muscles as jaw adductors. Our FEA simulations indicate that Z. varolai exerted an anterior bite force of more than 4000 N and a posterior bite force of more than 10000 N. These values are similar to those estimated for other marine predators known for their powerful bite. This suggests that Z. varolai might have fed upon medium-sized marine vertebrates like other odontocetes. Considering the significant difference observed between the anterior and posterior bite forces, Z. varolai likely fed via 'grip-and-shear' feeding, snapping the food items with an anterior bite and then cutting them with a powerful posterior bite. Other macroraptorial sperm whales such as the roughly coeval Acrophyseter from Peru likely employed the same feeding technique. ARTICLE HISTORY
... Tooth morphology has been traditionally used to diagnose extinct sperm whales and some species or genera have been defined based on one or more isolated teeth (e.g., Hoplocetus Gervais, 1848; Scaldicetus Du Bus, 1867) (Piazza et al., 2018;Paolucci et al., 2021a,b). However, the diagnostic potential of dental characters in the taxonomy of extinct odontocete taxa has been questioned by several researchers (e.g., Kellogg, 1925;Bianucci and Landini, 2006;Hampe, 2006;Lambert et al., 2017;Reumer et al., 2017) and there is a broad consensus that dental morphology is not sufficient to identify and differentiate taxa neither at the genus nor the species level (Kimura and Hasegawa, 2006;Paolucci et al., 2021a,b). This is related to the difficulty to evaluate homodont dentition in odontocetes, but also because there is scarce information on ontogenetic and variation in their teeth (Paolucci et al., 2021a,b). ...
Remains of cetaceans, particularly odontocetes, are scarce in the Cuban fossil record. Here we describe the second
odontocete (toothed whale) record from the Neogene of Cuba and extends the temporal distribution of this
group in the Insular Caribbean into the Middle Miocene. This account is based on two isolated teeth found in
limestones of the Güines Formation that crop out in a quarry near the town of Guanábana, Matanzas City, northwestern
Cuba. The specimens are referred to the superfamily Physeteroidea incertae sedis. These findings, along
with other fossils recovered from other Early and Middle Miocene localities in the region support the hypothesis
of a complex, diverse marine vertebrate fauna on the formerly called “Habana-Matanzas channel” and circum-
Cuban waters during the Neogene.
Los restos fosilizados de ballenas y delfines son escasos en el registro fósil cubano.
Aquí reportamos el primer
fósil Odontoceto del Mioceno Medio de Cuba. El reporte se basa en restos dentales desarticulados encontrados en
calizas de la Formación Güines que afloran en una cantera cercana al poblado de Guanábana, sur de la ciudad de
Matanzas, al noroeste de Cuba. Los especímenes se clasifican como especies no identificadas dentro de la familia
de ballenas dentadas Physeteroidea incertae sedis. Estos hallazgos apoyan la hipótesis de una fauna de vertebrados
marinos diversa y compleja en el llamado “canal Habana-Matanzas” y aguas circum-cubanas-caribeñas durante
el Neógeno
... of Odontocetes from Mysticetes around 30 Mya (Thewissen & Williams, 2002), many of the larger predators, such as Otodus megalodon (Shimada et al., 2016;Pimiento et al., 2017;Cooper et al., 2022) and a large physeteroid (sperm whale) (Kimura et al., 2006;Lambert et al., 2014;Peri et al., 2022) disappeared around the end of the Pliocene, a period marked by climatic variability and sea-level fluctuations (Pimiento et al., 2017). The earliest known fossil of a killer whale, O. citonensis, dates back to the Pliocene Epoch (5.3 million to 2.6 Mya) and was only about 4 m in length, similar in size to a typical dolphin (Galatius et al., 2020). ...
Unlabelled:
A widespread pattern in vertebrate life-history evolution is for species to evolve towards either fast or slow life histories; however, the underlying causes of this pattern remain unclear. Toothed whales (Odontoceti) are a diverse group with a range of body sizes and life histories, making them an ideal model to investigate potential drivers of this dichotomy. Using ancestral reconstruction, we identified that certain groups of odontocetes evolved more-streamlined, presumably faster, body shapes around the same time that killer whales (Orcinus orca) evolved into whale predators approximately 1 Mya during the Pleistocene. This suggests that the evolution of a streamlined body shape may have been an adaptation to escape killer whale predation, leading to longer life-history events. To test this hypothesis, we performed a cluster analysis of odontocete whales and confirmed the dual pattern of life-history traits, with one group referred to as 'reproducers' characterized by early age of maturity, short gestation, short interbirth interval, and short lifespan, and the other group referred to as 'bet-hedgers' exhibiting the opposite pattern. However, we found that life history grouping was relatively unrelated to whale shape (i.e., more streamlined or less streamlined). Therefore, we incorporated principal component results into mixed effects models, and the model results indicated that body shape was positively related to neonate length (a measure of investment in progeny), but not significantly related to the temporal life-history traits. Thus, whale body shape is not a sufficient explanation for the evolution of fast-slow life histories in odontocete whales.
Supplementary information:
The online version contains supplementary material available at 10.1007/s11692-023-09605-4.
... Tooth morphology has been traditionally used to diagnose extinct sperm whales and some species or genera have been defined based on one or more isolated teeth (e.g., Hoplocetus Gervais, 1848; Scaldicetus Du Bus, 1867) (Piazza et al., 2018;Paolucci et al., 2021a,b). However, the diagnostic potential of dental characters in the taxonomy of extinct odontocete taxa has been questioned by several researchers (e.g., Kellogg, 1925;Bianucci and Landini, 2006;Hampe, 2006;Lambert et al., 2017;Reumer et al., 2017) and there is a broad consensus that dental morphology is not sufficient to identify and differentiate taxa neither at the genus nor the species level (Kimura and Hasegawa, 2006;Paolucci et al., 2021a,b). This is related to the difficulty to evaluate homodont dentition in odontocetes, but also because there is scarce information on ontogenetic and variation in their teeth (Paolucci et al., 2021a,b). ...
... The marine deposits of the Tomioka-Annaka groups yield abundant and well-preserved marine vertebrate fossils (e. g., Takakuwa et al, 2001;Matsuoka et al., 2002;Kimura et al., 2003;Takakuwa and Hasegawa, 2004;Hasegawa et al., 2005;Hasegawa and Kimura, 2008). Even only from the Haraichi Formation, many sharks (Goto et al., 1978(Goto et al., , 1983Takakuwa, 1999Takakuwa, , 2005Takakuwa et al., 2003), Allodesminae pinniped (Hasegawa et al., 2000), desmostylian mammal Paleoparadoxia (Hasegawa and Kimura, 2008), whales and dolphins including originally established Joumocetus shimizui Kimura and Hasegawa, 2010, Kentriodon nakajimai Kimura and Hasegawa, 2019, and Norisdelphis annakaensis Kimura and Hasegawa, 2020, 2002Kimura et al., 2006;Kimura and Hasegawa, 2010a, 2010b, 2019, 2020, and a bird "Annaka Short-winged Swan" (Matsuoka et al., 2001(Matsuoka et al., , 2004 have been reported. Invertebrate fossils reported from the Tomioka-Annaka groups include: fossil decapod crustaceans (Kato, 2001), molluscs (Kurihara, 2000(Kurihara, , 2002, barnacles (Nomura and Takakuwa, 2009), and crinoids and so on. ...
Abstract: A new genus and two new species of flightless fossil Cygnini (Aves: Anatidae), Annakacygna hajimei and A.
yoshiiensis are described from the horizon of about 11.5 Ma of the Miocene Haraichi Formation, Annaka Group of Gunma,
central Japan. The type specimen of A. hajimei (GMNH-PV-678) is the associated skeleton of an adult bird. Osteological
features of A. hajimei indicate that this large anatid bird belongs to tribe Cygnini as it possesses the diagnostic features of the
tribe such as the long neck and long pelvis with dorsally swelling ala postacetabularis illi. Annakacygna were a group of
flightless birds characterized by the distally small wing elements, large body, and pachyostotic bones. In addition, the
autapomorphies of Annakacygna indicate that this bird was an animal with the body plan that specialized in breeding behavior,
or sex appeal in a broad sense, and uniquely developed the filter feeding with its large head. In a sense, Annakacygna acquired
remarkable evolutionary forms for two essential biological activities, eating and reproduction, is the “ultimate bird” ever exist.
A. yoshiiensis, known only from the distal end of tibiotarsus (GMNH-PV-1685), was a gigantic species and showed the
diversification within this unique genus in the sea of Miocene Japan.
... Traditionally, teeth displaying this particular morphology were identified as Scaldicetus (e.g., Barnes, 1976). However, it is now known that there are multiple taxa of stem physeteroids that share a similar morphology with the teeth from the Monterey Formation (e.g., Brygmophyseter- Kimura et al., 2006;Acrophyseter-Lambert et al., 2017a). Crown physeteroids are absent in the Monterey Formation in Southern California, whereas they are represented in the middle Miocene Sharktooth Hill bone bed by Aulophyseter morricei and Idiophyseter merriami from beds mapped as Monterey Formation further north (Kellogg, 1925b(Kellogg, , 1927Dibblee and Minch, 2006;Velez-Juarbe, 2018). ...
This study provides new stratigraphic data and identifications for fossil marine mammals from the Monterey Formation in the Capistrano syncline, Orange County, California, showing that there are two distinct marine mammal assemblages. Until now, marine mammals from the Monterey Formation of Orange County have been considered to represent a single assemblage that is 13.0-10.0 Ma in age. By combining data from diatoms with the geographic positions of sites, faunal analysis, and data from the literature, we can assign 59 sites to three main levels: the lower part (ca. 16-13 Ma), the middle part (ca. 13-10 Ma), and the upper part (ca. 10-8 Ma). We assigned 308 marine mammal specimens to 38 taxa, resulting in 97 occurrences (unique record of a taxon for a given site). Of the 38 taxa we identified within the study area, 15 taxa are restricted to the lower part of the Monterey Formation, 15 are restricted to the upper part of the Monterey Formation, eight were found in both, and none has yet been reported from the middle (possibly condensed) section. Six of the eight taxa that occur in both the lower and upper parts of the Monterey Formation are higher-level taxa, which accounts for their broad temporal range. The recognition of two distinct marine mammal assemblages in the Monterey Formation of Orange County is an important step toward a better-calibrated sequence of faunal evolution in the region while improving the utility of marine mammals for regional biostratigraphy.
... Historically, the physeteroid skull has been considered as the main source to diagnose taxa and resolve phylogenetic relationships among species. However, the diagnostic value of teeth and tympano-periotic bones has been questioned by some authors (Kellogg 1925(Kellogg , 1927Caviglia and Jorge 1980;Hirota and Barnes 1994;Luo and Marsh 1996;Bianucci and Landini 2006;Kimura et al. 2006;Hampe 2006;Reumer et al. 2017), especially in the case of isolated materials or incomplete skulls. ...
Sperm whales (Physeteroidea) include today only two genera of morphologically disparate odontocetes: the largest
toothed whale known (Physeter macrocephalus) and small sized forms (Kogia spp.). In contrast, their fossil record indicates
a high diversity for the group during the Miocene, with over 20 species recognized. Miocene marine sediments from
Patagonia (Argentina) record this diversity, including at least five species. Among them, Preaulophyseter gualichensis,
from the Miocene of Gran Bajo del Gualicho Formation, has been one of the most enigmatic. Despite the fragmentary
nature of the type and referred materials (isolated teeth and periotics), which casts some doubts on its validity, this species
has not been revised since its original description. In this contribution, we re-describe the materials referred to P. gualichensis,
revise the taxonomic status of the species and evaluate the phylogenetic signal of ear bones among Physeteroidea.
Our results indicate that the physeteroid tympano-periotic complex morphology is poorly diagnostic at the species level.
Intraspecific variation (including ontogeny and sexual dimorphism) and/or taphonomic processes cannot be ruled out as
the causes of the minor differences observed among specimens. We suggest that sperm whale tympano-periotics
retain
many plesiomorphic characters and are diagnostic only between kogiids and non-kogiid physeteroids. Based on the fragmentary
and isolated state of the studied specimens, and the lack of diagnostic characters in both teeth and periotics, we
consider P. gualichensis as nomen dubium and we re-assign the referred specimens as Physeteroidea indet. A conservative
morphology of the tympano-periotic and, to a lesser extent, the nasal complex in sperm whales, might result from the
morpho-functional constraints imposed by a highly specialized but successful echolocation system.
We describe a new specimen of physeteroid from the lower Miocene (Burdigalian) of Japan. This specimen was recovered from the Toyohama Formation, Chita County, Aichi Prefecture, Japan in 1984 and includes a finely preserved cranium with detached teeth and ear bones (periotic, tympanic bulla, and malleus). Here we refer this specimen to a new genus and species of the Physeteroidea, Miophyseter chitaensis gen. et sp. nov. Our phylogenetic analysis suggests that Miophyseter is a physeteroid more closely related to the crown Physeteroidea (Physeteridae and Kogiidae) than the macroraptorial physeteroids that flourished in middle and late
Miocene times. A deep and large excavation on the ventral surface of the palatine and pterygoid in Miophyster suggests an adaptation for deep dives and/or the development of robust pterygoid muscles for active biting.
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