Histology and growth pattern of the pachy-osteosclerotic premaxillae of the fossil beaked whale Aporotus recurvirostris (Mammalia, Cetacea, Odontoceti)

Beaked whales (Ziphiidae) often show highly specialized features, involving bone morphology or structure, in the rostral region of their skulls. Previous studies revealed an extremely derived and peculiar histological structure in the rostrum of the extant Mesoplodon densirostris. In order to assess if this structure is a general feature of ziphiids, the swollen premaxillae of Aporotus recurvirostris, a Miocene species from the North Sea, were studied histologically. These bones are pachyostotic and strongly osteosclerotic. However, their structural organization is entirely different from that of M. densirostris rostrum: they are basically made of a non-remodeled, laminar tissue that was cyclically deposited by the periosteum. As compared to the generalized structure of the premaxillae of toothed whales exemplified by the bottlenose dolphin, Tursiops truncatus, the pachyostotic condition of Aporotus premaxillae was obviously due to a particularly high and sustained growth-rate, occurring in a dorso-lateral direction. The osteosclerotic structure of these bones resulted from a complete lack of inner resorption activity. The histological features of Aporotus premaxillae indicate that these bones are not likely to have been hypermineralized, and thus, their physical properties must have differed from those of the M. densirostris rostrum. The possible functional involvements of rostral peculiarities in beaked whales are discussed with reference to the whole set of available comparative data.     

Structure and growth pattern of the bizarre hemispheric prominence on the rostrum of the fossil beaked whale Globicetus hiberus (Mammalia,Cetacea, Ziphiidae)

The rostrum of most ziphiids (beaked whales) displays bizarre swollen regions, accompanied with extreme hypermineralisation and an alteration of the collagenous mesh of the bone. The functional significance of this specialization remains obscure. With the voluminous and dense hemispheric excrescence protruding from the premaxillae, the recently described fossil ziphiid Globicetus hiberus is the most spectacular case. This study describes the histological structure and interprets the growth pattern of this unique feature. Histologically, the prominence in Globicetus is made up of an atypical fibro‐lamellar complex displaying an irregular laminar organization and extreme compactness (osteosclerosis). Its development is suggested to have resulted from a protraction of periosteal accretion over the premaxillae, long after the end of somatic growth. Complex shifts in the geometry of this tissue are likely to have occurred during its accretion and no indication of Haversian remodeling could be found. X‐ray diffraction and Raman spectroscopy indicate that the bone matrix in the premaxillary prominence of Globicetus closely resembles that of the rostrum of the extant beaked whale Mesoplodon densirostris: apatite crystals are of common size and strongly oriented, but the collagenous meshwork within bone matrix seems to be extremely sparse. These morphological and structural data are discussed in the light of functional interpretations proposed for the highly unusual and diverse ziphiid rostrum. J. Morphol. 277:1292–1308, 2016. © 2016 Wiley Periodicals, Inc.     

No deep diving: evidence of predation on epipelagic fish for a stem beaked whale from the Late Miocene of Peru

Although modern beaked whales (Ziphiidae) are known to be highly specialized toothed whales that predominantly feed at great depths upon benthic and benthopelagic prey, only limited palaeontological data document this major ecological shift. We report on a ziphiid–fish assemblage from the Late Miocene of Peru that we interpret as the first direct evidence of a predator–prey relationship between a ziphiid and epipelagic fish. Preserved in a dolomite concretion, a skeleton of the stem ziphiid Messapicetus gregarius was discovered together with numerous skeletons of a clupeiform fish closely related to the epipelagic extant Pacific sardine (Sardinops sagax). Based on the position of fish individuals along the head and chest regions of the ziphiid, the lack of digestion marks on fish remains and the homogeneous size of individuals, we propose that this assemblage results from the death of the whale (possibly via toxin poisoning) shortly after the capture of prey from a single school. Together with morphological data and the frequent discovery of fossil crown ziphiids in deep-sea deposits, this exceptional record supports the hypothesis that only more derived ziphiids were regular deep divers and that the extinction of epipelagic forms may coincide with the radiation of true dolphins.     

The most basal beaked whale Ninoziphius platyrostris Muizon, 1983: clues on the evolutionary history of the family Ziphiidae (Cetacea: Odontoceti)

Ninoziphius platyrostris, from the late Neogene of Peru, is one of the best‐known fossil beaked whales (Odontoceti: Ziphiidae), with a holotype including the skull with ear bones, mandibles, teeth, and postcranial elements. Furthermore, based on several characters, including a complete functional upper and lower dentition, it is usually considered as one of the most archaic ziphiids. However, the poorly preserved dorsal portion of the holotype skull has led to unresolved phylogenetic relationships. With the addition of two newly prepared skulls from the same Peruvian locality we redescribed N. platyrostris. In the light of recent ziphiid discoveries, an emended diagnosis of the species is proposed here. In our cladistic analysis Ninoziphius is the most basal stem ziphiid. Newly observed or reassessed morphological traits allow functional and ecological considerations. The morphology of the oral apparatus suggests that Ninoziphius was less specialized for suction feeding than most extant ziphiids. Tooth wear in the holotype may indicate benthic feeding. Although the vertebral column of Ninoziphius corresponds to less developed locomotor abilities for deep dives, its cranial morphology does not provide definitive arguments for an echolocation system less efficient than in deep diving extant ziphiids. Finally, the phylogenetic tree produced was used to detail the evolutionary history of several major ziphiid features (dental reduction, development of mandibular tusks, and increased body size). © 2013 The Linnean Society of London     

From Problem Taxa to Problem Solver: A New Miocene Family,Tranatocetidae, Brings Perspective on Baleen Whale Evolution

Miocene baleen whales were highly diverse and included tens of genera. However, their taxonomy and phylogeny, as well as relationships with living whales, are still a subject of controversy. Here, “Mesocetus” argillarius, a poorly known specimen from Denmark, is redescribed with a focus on the cranial anatomy. It was found to represent not only a new genus, Tranatocetus gen. nov., but also a new family; Tranatocetidae. The whales of this family have the rostral bones either overriding or dividing the frontals; the rostral bones are contacting the parietals and nasals dividing the maxillae on the vertex; the occipital shield is dorsoventrally bent. The tympanic bulla is particularly characteristic of this family featuring a short, narrow anterior portion with a rounded or squared anterior end and a wider and higher posterior portion that is swollen in the posteroventral area. A phylogenetic analysis including 51 taxa supports a monophyletic group comprising most Neogene and modern whales, with Tranatocetidae being possibly closer related to Balaenopteridae (rorquals) than to Cetotheriidae. Tranatocetidae exhibit a charahteristic bulla shape. In fact, all Neogene and modern mysticete families examined have a unique shape of the tympanic bulla that is diagnostic at family-level. Inclusion of problematic taxa like Tranatocetus argillarius in phylogenies brings new understanding of the distribution and diagnostic value of character traits. This underlines the need for re-examination of earlier described specimens in the light of the wealth of new information published in later years.     

Comparative paleohistology in osteoderms of Pleistocene Panochthus sp. Burmeister, 1886 and Neuryurus sp. (Xenarthra,Glyptodontidae)

Xenarthran osteoderms are integumentary bones with high fossilization potential presenting a high degree of morphological and histological diversity. Here, new data on the osteoderms histology of two glyptodonts, Panochthus and Neuryurus are presented. The poor spatial organization of the mineralized fibers and a large trabecular area in the middle zone identified in Neuryurus indicate a different bone pattern than the one found in Panochthus, which is mainly characterized by a middle zone with less spongiosa. Through the Bone Profiler program, the degree of compactness of the specimens was obtained, with about 70% for Neuryurus sp. and approximately 90% for Panochthus sp., showing the difference in bone pattern. These values confirm the visible difference in the histological patterns of these taxa, especially in the middle zone. This work demonstrates the microstructural variation studied in osteoderms and shows the importance of paleohistology as a starting point for a better understanding of extinct taxa.     

Dorsal rib histology of dinosaurs and a crocodylomorph from western Portugal: Skeletochronological implications on age determination and life history traits

Bone histology is an important tool for uncovering life history traits of extinct animals, particularly those that lack modern analogs, such as the non-avian dinosaurs. In most studies, histological analyses preferentially focus on long bones for understanding growth rates and determining age. Here we show, by analyzing ornithischians (a stegosaur and an ornithopod), saurischians (a sauropod and a theropod), and a crocodile, rib histology is a suitable alternative. The estimated age for all sampled taxa ranges between 14 to 17 years for Lourinhanosaurus antunesi and 27 to 31 years estimated for Draconyx loureiroi. The theropod Baryonyx was skeletally mature around 23–25 years of age but showed unfused neurocentral sutures, a paedomorphic feature possibly related to aquatic locomotion. Our results show that ribs can contain a nearly complete growth record, and reveal important information about individual age, point of sexual maturity, and, in some cases, sex. Because ribs are more available than long bones, this method opens new possibilities for studying rare and incomplete fossils, including holotypes.     

A New Look at Ichthyosaur Long Bone Microanatomy and Histology: Implications for Their Adaptation to an Aquatic Life

Background

Ichthyosaurs are Mesozoic reptiles considered as active swimmers highly adapted to a fully open-marine life. They display a wide range of morphologies illustrating diverse ecological grades. Data concerning their bone microanatomical and histological features are rather limited and suggest that ichthyosaurs display a spongious, “osteoporotic-like” bone inner structure, like extant cetaceans. However, some taxa exhibit peculiar features, suggesting that the analysis of the microanatomical and histological characteristics of various ichthyosaur long bones should match the anatomical diversity and provide information about their diverse locomotor abilities and physiology.

Methodology/Principal Findings

The material analyzed for this study essentially consists of mid-diaphyseal transverse sections from stylopod bones of various ichthyosaurs and of a few microtomographic (both conventional and synchrotron) data. The present contribution discusses the histological and microanatomical variation observed within ichthyosaurs and the peculiarities of some taxa (Mixosaurus, Pessopteryx). Four microanatomical types are described. If Mixosaurus sections differ from those of the other taxa analyzed, the other microanatomical types, characterized by the relative proportion of compact and loose spongiosa of periosteal and endochondral origin respectively, seem to rather especially illustrate variation along the diaphysis in taxa with similar microanatomical features. Our analysis also reveals that primary bone in all the ichthyosaur taxa sampled (to the possible exception of Mixosaurus) is spongy in origin, that cyclical growth is a common pattern among ichthyosaurs, and confirms the previous assumptions of high growth rates in ichthyosaurs.

Conclusions/Significance

The occurrence of two types of remodelling patterns along the diaphysis, characterized by bone mass decrease and increase respectively is described for the first time. It raises questions about the definition of the osseous microanatomical specializations bone mass increase and osteoporosis, notably based on the processes involved, and reveals the difficulty in determining the true occurrence of these osseous specializations in ichthyosaurs.     

Phylogenetic signal in bone histology of amniotes revisited

Legendre, L, Le Roy, N, Martinez‐Maza, C, Montes, L, Laurin, M & Cubo, J. (2012). Phylogenetic signal in bone histology of amniotes revisited. —Zoologica Scripta, 42, 44–53. There is currently a debate about the presence of a phylogenetic signal in bone histological data, but very few rigorous tests have fuelled the discussions on this topic. Here, we performed new analyses using a larger set of seven histological traits and including 25 taxa (nine extinct and 16 extant taxa), using three methods: the phylogenetic eigenvector regression, the tree length distribution and the regressions on distance matrices. Our results clearly show that the phylogenetic signal in our sample of bone histological characters is strong, even after correcting for multiple testing. Most characters exhibit a significant phylogenetic signal according to at least one of our three tests, with the phylogeny often explaining 20–60% of the variation in the histological characters. Thus, we conclude that the phylogenetic comparative method should be systematically used in interspecific analyses of bone histodiversity to avoid problems of non‐independence among observations.     

Palaeobiology of Triassic procolophonids,inferred from bone microstructure

Procolophonoidea represent the most successful radiation of Parareptilia that lived during the Permo-Triassic. They are one of the few vertebrate groups that survived the end-Permian extinction and are thus important for studying the recovery of the post-extinction terrestrial ecosystem. Here, we investigate the palaeobiology of three Triassic procolophonid parareptiles, namely Sauropareion anoplus, Procolophon trigoniceps and Teratophon spinigenis, from the Karoo Basin of South Africa, inferred from histological analyses of their limb bones. Results reveal that all three taxa exhibit parallel-fibered bone tissue. Growth rings are absent in the Early Triassic Sauropareion and Procolophon whereas annuli are present in the Middle Triassic Teratophon, even during early ontogeny, suggesting a difference in life histories. Morphology and bone histology imply fossorial lifestyles for all three taxa, suggesting that burrowing may have played an important role in their survival during the harsh post-extinction Triassic environment.     

Adaptive evolution of tight junction protein claudin-14 in echolocating whales

Toothed whales and bats have independently evolved specialized ultrasonic hearing for echolocation. Recent findings have suggested that several genes including Prestin, Tmc1, Pjvk and KCNQ4 appear to have undergone molecular adaptations associated with the evolution of this ultrasonic hearing in mammals. Here we studied the hearing gene Cldn14, which encodes the claudin-14 protein and is a member of tight junction proteins that functions in the organ of Corti in the inner ear to maintain a cationic gradient between endolymph and perilymph. Particular mutations in human claudin-14 give rise to non-syndromic deafness, suggesting an essential role in hearing. Our results uncovered two bursts of positive selection, one in the ancestral branch of all toothed whales and a second in the branch leading to the delphinid, phocoenid and ziphiid whales. These two branches are the same as those previously reported to show positive selection in the Prestin gene. Furthermore, as with Prestin, the estimated hearing frequencies of whales significantly correlate with numbers of branch-wise non-synonymous substitutions in Cldn14, but not with synonymous changes. However, in contrast to Prestin, we found no evidence of positive selection in bats. Our findings from Cldn14, and comparisons with Prestin, strongly implicate multiple loci in the acquisition of echolocation in cetaceans, but also highlight possible differences in the evolutionary route to echolocation taken by whales and bats.     

Bone microstructure of Vegavis iaai (Aves,Anseriformes) from the Upper Cretaceous of Vega Island,Antarctic Peninsula

Vegavis iaai is a neornithine bird coming from the Late Cretaceous Sandwich Bluff Member of the López de Bertodano Formation (Maastrichtian), Antarctic Peninsula. Vegavis constitutes the only unquestionable Cretaceous neornithine bird, and is known by the holotype and specimen MACN-PV 19.748. The goal of this paper is to present a detailed osteohistological analysis of V. iaai. Vegavis shows a highly vascularized fibrolamellar matrix lacking lines of arrested growths, features widespread among modern birds. This is consistent with previous hypotheses indicating that modern birds were dominant at high latitudes. This is probably related to high-metabolic rates shared by modern birds, whereas archaic taxa as Enantiornithes are absent or form a minority part of High-Latitude bird assemblages. Vegavis was a diver, characterised by a certain degree of limb osteosclerosis, with an increase of bone inner compactness, and inhibition of secondary remodelling, with no effect on the external dimensions of the bone, a combination of characters related to diving lifestyle. Based on Relative Bone Thickness it is possible to infer that Vegavis was a foot-propelled diving bird, similar to some extant anseriforms. Occurrence of osteosclerotic limb bones in Vegavis and Polarornis may constitute a derived shared feature, sustaining the hypothesis that both taxa are phylogenetically related.     

Unexpected microanatomical variation among Eocene Antarctic stem penguins (Aves: Sphenisciformes)

The microanatomical and histological structure of Eocene Antarctic stem penguin tarsometatarsi is examined in order to characterise the bone microstructure. Eight adult tarsometatarsi belonging to eight fossil species (Palaeeudyptes gunnari, Palaeeudyptes klekowskii, Anthropornis grandis, Anthropornis nordenskjoeldi, Archaeospheniscus wimani, Marambiornis exilis, Delphinornis arctowskii and Delphinornis larseni) collected from the Antarctic A. nordenskjoeldi Biozone (La Meseta Formation, ～34.2 Ma) were examined. The thin sections revealed a distinctive microanatomical variation among taxa. Whereas Anthropornis spp., A. wimani and P. gunnari possess massive, clearly osteosclerotic bones (medullary cavities absent or strongly reduced), the bones of Delphinornis spp., P. klekowski and M. exilis exhibit well-developed medullary cavities. The cortical bone in all the specimens consists of primary, well-vascularised fibro-lamellar bone and variable amounts of secondary bone. Medullary cavities are coated by a thick layer of lamellar bone tissue and coarse compacted cancellous bone. Although several causes can explain the striking microanatomical variation (e.g. ontogeny), we interpret that such variation is related to differential adaptations to the aquatic life, for which taxa with more massive bones were possibly adapted to deeper and more prolonged diving excursions.     

Bone Inner Structure Suggests Increasing Aquatic Adaptations in Desmostylia (Mammalia,Afrotheria)

Background

The paleoecology of desmostylians has been discussed controversially with a general consensus that desmostylians were aquatic or semi-aquatic to some extent. Bone microanatomy can be used as a powerful tool to infer habitat preference of extinct animals. However, bone microanatomical studies of desmostylians are extremely scarce.

Methodology/Principal Findings

We analyzed the histology and microanatomy of several desmostylians using thin-sections and CT scans of ribs, humeri, femora and vertebrae. Comparisons with extant mammals allowed us to better understand the mode of life and evolutionary history of these taxa. Desmostylian ribs and long bones generally lack a medullary cavity. This trait has been interpreted as an aquatic adaptation among amniotes. Behemotops and Paleoparadoxia show osteosclerosis (i.e. increase in bone compactness), and Ashoroa pachyosteosclerosis (i.e. combined increase in bone volume and compactness). Conversely, Desmostylus differs from these desmostylians in displaying an osteoporotic-like pattern.

Conclusions/Significance

In living taxa, bone mass increase provides hydrostatic buoyancy and body trim control suitable for poorly efficient swimmers, while wholly spongy bones are associated with hydrodynamic buoyancy control in active swimmers. Our study suggests that all desmostylians had achieved an essentially, if not exclusively, aquatic lifestyle. Behemotops, Paleoparadoxia and Ashoroa are interpreted as shallow water swimmers, either hovering slowly at a preferred depth, or walking on the bottom, and Desmostylus as a more active swimmer with a peculiar habitat and feeding strategy within Desmostylia. Therefore, desmostylians are, with cetaceans, the second mammal group showing a shift from bone mass increase to a spongy inner organization of bones in their evolutionary history.     

Impact of increased character sampling on the phylogeny of Cetartiodactyla (Mammalia): combined analysis including fossils

The phylogenetic position of Cetacea (whales, dolphins and porpoises) is an important exemplar problem for combined data parsimony analyses because the clade is ancient and includes many well‐known and relatively complete fossil species. We combined data for 71 terminal taxa (43 extinct/28 extant) to test where Cetacea fits within Cetartiodactyla, and where various fossil hoofed mammals (e.g., ?entelodonts, “?anthracotheriids” and ?mesonychians) are positioned. We scored 635 phenotypic characters (osteology, dentition, soft tissue, behavior), approximately three times the number of characters in the last major analysis of this clade, and combined these with > 40 000 molecular characters, including new data from 10 genes. The analysis supported a topology consistent with the majority of recently published molecular studies. Cetacea was the extant sister taxon of Hippopotamidae, followed successively by Ruminantia, Suina and Camelidae. Several extinct taxa were phylogenetically unstable, upsetting resolution of the strict consensus and limiting branch support, but the positions of several key fossils were consistently resolved. The wholly extinct ?Mesonychia was more closely related to Cetacea than was any “artiodactylan.”“?Anthracotheriids” were paraphyletic, and, with the exception of one species, were more closely related to Hippopotamidae than to any other living taxon. The total evidence analysis overturned a highly nested position for Moschus supported by molecular data alone. The character partition that could be scored for the fossil taxa (osteological and dental characters) included more informative characters than most molecular partitions in our analysis, and had the fewest missing data. The osteological–dental data alone, however, did not support inclusion of cetaceans within crown “Artiodactyla.” Recently discovered ankle bones from fossil whales reinforced the monophyly of Cetartiodactyla but provided no particular evidence of derived similarities between hippopotamids and fossil cetaceans that were not shared with other “artiodactylans”. © The Willi Hennig Society 2007.     

Relationships of Cetacea (Artiodactyla) Among Mammals: Increased Taxon Sampling Alters Interpretations of Key Fossils and Character Evolution

Background

Integration of diverse data (molecules, fossils) provides the most robust test of the phylogeny of cetaceans. Positioning key fossils is critical for reconstructing the character change from life on land to life in the water.

Methodology/Principal Findings

We reexamine relationships of critical extinct taxa that impact our understanding of the origin of Cetacea. We do this in the context of the largest total evidence analysis of morphological and molecular information for Artiodactyla (661 phenotypic characters and 46,587 molecular characters, coded for 33 extant and 48 extinct taxa). We score morphological data for Carnivoramorpha, †Creodonta, Lipotyphla, and the †raoellid artiodactylan †Indohyus and concentrate on determining which fossils are positioned along stem lineages to major artiodactylan crown clades. Shortest trees place Cetacea within Artiodactyla and close to †Indohyus, with †Mesonychia outside of Artiodactyla. The relationships of †Mesonychia and †Indohyus are highly unstable, however - in trees only two steps longer than minimum length, †Mesonychia falls inside Artiodactyla and displaces †Indohyus from a position close to Cetacea. Trees based only on data that fossilize continue to show the classic arrangement of relationships within Artiodactyla with Cetacea grouping outside the clade, a signal incongruent with the molecular data that dominate the total evidence result.

Conclusions/Significance

Integration of new fossil material of †Indohyus impacts placement of another extinct clade †Mesonychia, pushing it much farther down the tree. The phylogenetic position of †Indohyus suggests that the cetacean stem lineage included herbivorous and carnivorous aquatic species. We also conclude that extinct members of Cetancodonta (whales + hippopotamids) shared a derived ability to hear underwater sounds, even though several cetancodontans lack a pachyostotic auditory bulla. We revise the taxonomy of living and extinct artiodactylans and propose explicit node and stem-based definitions for the ingroup.     

On the Middle Miocene avifauna of Maboko Island,Kenya

The Middle Miocene sediments of Maboko Island (Lake Victoria) in western Kenya yielded numerous avian bones, which remained, however, little studied. The significance of this material is shown by the recent identification of an opisthocomiform bird. In the present study, further avian remains from Maboko Island are described. Most of the specimens belong to aquatic or semi-aquatic groups, of which some are closely related to taxa known from Early and Middle Miocene European avifaunas, that is, Nectornis cormorants (N. africanus nov. sp.) and Laricola-like Laromorphae. The fossil material also includes Ciconiidae (cf. Ciconia), Pelecanidae, Phoenicopteridae (Leakeyornis aethiopicus), Musophagidae, and a species of Ardeidae, which closely resembles the taxon Pikaihao from the Early Miocene of New Zealand. Some avian remains from Maboko Island belong to higher-level taxa unknown from the Middle Miocene of Europe. The occurrence of a giant Jacanidae (?Nupharanassa mabokoensis nov. sp.) is of particular interest, because these are globally absent in extant avifaunas and were previously only known from the Late Eocene/Early Oligocene of Egypt. Further unknown from contemporaneous European sites are small representatives of Jacanidae, Bucerotidae, and Alcedinidae, with the fossils of the latter two taxa being among the earliest published records of their respective groups. Several of the taxa that are common in contemporaneous European avifaunas have not been found in Maboko, and in spite of less pronounced climatic differences, Middle Miocene Afrotropical avifaunas already appear to have been distinct from contemporaneous European ones.     

New skull of Schizodelphis sulcatus Gervais, 1861 (Mammalia, Odontoceti, Eurhinodelphinidae) from the Lower Miocene of Pignan (Hérault, France) and its implications for systematics of Eurhinodelphinidae

Schizodelphis sulcatus is the type species of the Eurhinodelphinid genus Schizodelphis and was until now only known from a rather fragmentary specimen, lacking some important parts of the skull (vertex, ear bones and apex of rostrum). The study of a well-preserved skull of S. sulcatus Gervais, discovered in the lower Miocene of Hérault (France), helps to improve the diagnosis of this enigmatic species. Diagnostic features include the low height at the base of the rostrum, the lower position of the paroccipital process compared to the height of the postglenoid process in lateral view, the ventrally deflected postglenoid process and the ovoid-shaped nasals. It also confirms that the type species of Schizodelphis is indeed distinct. For the first time in Schizodelphis, the vertex is preserved and the ear bones are found in anatomic connection with the basicranium. A cladistic analysis of cranial characters of Eurhinodelphinidae, including ear bones, is performed and reveals a close phylogenetic relationship between S. sulcatus and the North American species Schizodelphis barnesi. This analysis also highlights some apomorphies of the genus as the smoothly laterally deflected anterior spine of the bulla in ventral view and the reestablishment of the contact between the falciform process of the squamosal and the lateral lamina of the pterygoid. Conversely, the attribution of Schizodelphis morckhoviensis to the genus Schizodelphis is questioned by this analysis.