Morgan Churchill

A Reevaluation of the Morphology, Paleoecology, and Phylogenetic Relationships of the Enigmatic Walrus Pelagiarctos

Background A number of aberrant walruses (Odobenidae) have been described from the Neogene of the North Pacific, including specialized suction-feeding and generalist fish-eating taxa. At least one of these fossil walruses has been hypothesized to have been a specialized predator of other marine mammals, the middle Miocene walrus Pelagiarctos thomasi from the Sharktooth Hill Bonebed of California (16.1–14.5 Ma). Methodology/Principal Findings A new specimen of Pelagiarctos from the middle Miocene “Topanga” Formation of southern California (17.5–15 Ma) allows a reassessment of the morphology and feeding ecology of this extinct walrus. The mandibles of this new specimen are robust with large canines, bulbous premolars with prominent paraconid, metaconid, hypoconid cusps, crenulated lingual cingula with small talonid basins, M2 present, double-rooted P3–M1, single-rooted P1 and M2, and a P2 with a bilobate root. Because this specimen lacks a fused mandibular symphysis like Pelagiarctos thomasi, it is instead referred to Pelagiarctos sp. This specimen is more informative than the fragmentary holotype of Pelagiarctos thomasi, permitting Pelagiarctos to be included within a phylogenetic analysis for the first time. Analysis of a matrix composed of 90 cranial, dental, mandibular and postcranial characters indicates that Pelagiarctos is an early diverging walrus and sister to the late Miocene walrus Imagotaria downsi. We reevaluate the evidence for a macropredatory lifestyle for Pelagiarctos, and we find no evidence of specialization towards a macrophagous diet, suggesting that Pelagiarctos was a generalist feeder with the ability to feed on large prey. Conclusions/Significance This new specimen of Pelagiarctos adds to the knowledge of this problematic taxon. The phylogenetic analysis conclusively demonstrates that Pelagiarctos is an early diverging walrus. Pelagiarctos does not show morphological specializations associated with macrophagy, and was likely a generalist predator, feeding on fish, invertebrates, and the occasional warm-blooded prey item.

Cope's rule and the evolution of body size in Pinnipedimorpha (Mammalia: Carnivora)

Copes rule describes the evolutionary trend for animal lineages to increase in body size over time. In this study, we tested the validity of Copes rule for a marine mammal clade, the Pinnipedimorpha, which includes the extinct Desmatophocidae, and extant Phocidae (earless seals), Otariidae (fur seals and sea lions), and Odobenidae (walruses). We tested for the presence of Copes rule by compiling a large dataset of body size data for extant and fossil pinnipeds and then examined how body size evolved through time. We found that there was a positive relationship between geologic age and body size. However, this trend is the result of differences between early assemblages of small‐bodied pinnipeds (Oligocene to early Miocene) and later assemblages (middle Miocene to Pliocene) for which species exhibited greater size diversity. No significant differences were found between the number of increases or decreases in body size within Pinnipedimorpha or within specific pinniped clades. This suggests that the pinniped body size increase was driven by passive diversification into vacant niche space, with the common ancestor of Pinnipedimorpha occurring near the minimum adult body size possible for a marine mammal. Based upon the above results, the evolutionary history of pinnipeds does not follow Copes rule.

Functional Implications of Variation in Tooth Spacing and Crown Size in Pinnipedimorpha (Mammalia: Carnivora)

Pinnipeds (seals, sea lions, and walruses) show variation in tooth morphology that relates to ecology. However, crown size and spacing are two aspects of morphology that have not been quantified in prior studies. We measured these characters for nearly all extant pinnipeds and three fossil taxa and then determined the principal sources of variation in tooth size and spacing using principal components (PCAs) and hierarchical cluster analysis (HCA). PCA and HCA showed that species sorted into three groups: taxa with small crowns and large diastemata, taxa with large crowns and small diastemata, and taxa that fell between these two extremes. We then performed discriminant function analysis (DFA) to determine if tooth morphology correlated with foraging strategy or diet. DFA results indicated weak correlation with diet, and stronger correlation with prey capture strategies. Tooth size and spacing were most strongly correlated with the importance of teeth in prey acquisition, with tooth size decreasing and tooth spacing increasing as teeth become less necessary in capturing food items. Taxa which relied on teeth for filtering prey from the water column or processing larger or tougher food items generally had larger crowns and smaller tooth spacing then taxa which swallowed prey whole. We found the fossil taxa Desmatophoca and Enaliarctos were most similar in tooth morphology to extant otariids, suggesting that both taxa were generalist feeders. This study established the relationship between tooth size and feeding behavior, and provides a new tool to explore the paleoecology of fossil pinnipeds and other aquatic tetrapods. Anat Rec, 298:878–902, 2015.

Predictive equations for the estimation of body size in seals and sea lions (Carnivora: Pinnipedia).

Body size plays an important role in pinniped ecology and life history. However, body size data is often absent for historical, archaeological, and fossil specimens. To estimate the body size of pinnipeds (seals, sea lions, and walruses) for today and the past, we used 14 commonly preserved cranial measurements to develop sets of single variable and multivariate predictive equations for pinniped body mass and total length. Principal components analysis (PCA) was used to test whether separate family specific regressions were more appropriate than single predictive equations for Pinnipedia. The influence of phylogeny was tested with phylogenetic independent contrasts (PIC). The accuracy of these regressions was then assessed using a combination of coefficient of determination, percent prediction error, and standard error of estimation. Three different methods of multivariate analysis were examined: bidirectional stepwise model selection using Akaike information criteria; all‐subsets model selection using Bayesian information criteria (BIC); and partial least squares regression. The PCA showed clear discrimination between Otariidae (fur seals and sea lions) and Phocidae (earless seals) for the 14 measurements, indicating the need for family‐specific regression equations. The PIC analysis found that phylogeny had a minor influence on relationship between morphological variables and body size. The regressions for total length were more accurate than those for body mass, and equations specific to Otariidae were more accurate than those for Phocidae. Of the three multivariate methods, the all‐subsets approach required the fewest number of variables to estimate body size accurately. We then used the single variable predictive equations and the all‐subsets approach to estimate the body size of two recently extinct pinniped taxa, the Caribbean monk seal (Monachus tropicalis) and the Japanese sea lion (Zalophus japonicus). Body size estimates using single variable regressions generally under or over‐estimated body size; however, the all‐subset regression produced body size estimates that were close to historically recorded body length for these two species. This indicates that the all‐subset regression equations developed in this study can estimate body size accurately.

The oldest known fur seal

The poorly known fossil record of fur seals and sea lions (Otariidae) does not reflect their current diversity and widespread abundance. This limited fossil record contrasts with the more complete fossil records of other pinnipeds such as walruses (Odobenidae). The oldest known otariids appear 5–6 Ma after the earliest odobenids, and the remarkably derived craniodental morphology of otariids offers few clues to their early evolutionary history and phylogenetic affinities among pinnipeds. We report a new otariid, Eotaria crypta, from the lower middle Miocene ‘Topanga’ Formation (15–17.1 Ma) of southern California, represented by a partial mandible with well-preserved dentition. Eotaria crypta is geochronologically intermediate between ‘enaliarctine’ stem pinnipedimorphs (16.6–27 Ma) and previously described otariid fossils (7.3–12.5 Ma), as well as morphologically intermediate by retaining an M2 and a reduced M1 metaconid cusp and lacking P2–4 metaconid cusps. Eotaria crypta eliminates the otariid ghost lineage and confirms that otariids evolved from an ‘enaliarctine’-like ancestor.

The evolution of aquatic feeding in seals: insights from Enaliarctos (Carnivora: Pinnipedimorpha), the oldest known seal

The development of pierce‐feeding and loss of oral processing represented major adaptations for underwater feeding in marine mammals. We examined the evolution of pierce‐feeding and its association with changes in tooth spacing and tooth size to determine whether pierce‐feeding was practiced by the earliest known pinnipeds. Data on crown size and spacing in postcanine dentition were collected and 1) analysed by principal components analysis (PCA) to determine the tooth morphospace of arctoid carnivores, 2) analysed by least squares (LS) regression and phylogenetic independent contrasts (PIC) to determine what morphological variables were associated with increases in tooth spacing, and 3) used to reconstruct the evolution of feeding related traits within a phylogenetic context. The PCA analysis revealed that within arctoid carnivores, the greatest differences in morphospace were associated with pierce‐feeding, and the early‐diverging seal Enaliarctos was placed within the pinniped morphospace. Increased tooth spacing within Pinnipedia is a result of decreased postcanine crown size. When the evolution of dental characters is reconstructed, ‘enaliarctines’ were found to represent an intermediate stage in evolution between ‘fissiped’ and pinniped carnivores. They retained the limited tooth spacing of terrestrial carnivores, possessed postcanine crown lengths intermediate in size between pinnipeds and fissipeds, and possessed reduced heterodonty characteristic of crown pinnipeds. Our study indicated that pierce‐feeding evolved early within pinnipeds. This suggested either that pierce‐feeding evolved prior to the loss of mastication, or that pierce‐feeding evolved at the same time as loss of mastication, and well before simplification of the dentition was completed.

Unique Biochemical and Mineral Composition of Whale Ear Bones

Cetaceans are obligate aquatic mammals derived from terrestrial artiodactyls. The defining characteristic of cetaceans is a thick and dense lip (pachyosteosclerotic involucrum) of an ear bone (the tympanic). This unique feature is absent in modern terrestrial artiodactyls and is suggested to be important in underwater hearing. Here, we investigate the mineralogical and biochemical properties of the involucrum, as these may hold clues to the aquatic adaptations of cetaceans. We compared bioapatites (enamel, dentine, cementum, and skeletal bone) of cetaceans with those of terrestrial artiodactyls and pachyosteosclerotic ribs of manatees (Sirenia). We investigated organic, carbonate, and mineral composition as well as crystal size and crystallinity index. In all studied variables, bioapatites of the cetacean involucrum were intermediate in composition and structure between those of tooth enamel on the one hand and those of dentine, cementum, and skeletal bone on the other. We also studied the amino acid composition of the cetacean involucrum relative to that of other skeletal bone. The central involucrum had low glycine and hydroxyproline concentrations but high concentrations of nonessential amino acids, unlike most bone samples but similar to the tympanic of hippos and the (pachyosteosclerotic) ribs of manatees. These amino acid results are evidence of rapid bone development. We hypothesize that the mineralogical and amino acid composition of cetacean bullae differs from that of other bone because of (1) functional modifications for underwater sound reception and (2) structural adaptations related to rapid ossification.

Taxonomy and biogeography of the Pleistocene New Zealand sea lion Neophoca palatina (Carnivora: Otariidae)

Abstract. The Otariidae (fur seals and sea lions) are an important and highly visible component of Southern Hemisphere marine mammal faunas. However, fossil material of Southern Hemisphere otariids is comparatively rare and often fragmentary. One exception is the Pleistocene sea lion Neophoca palatina King, 1983a, which is known from a nearly complete skull from the North Island of New Zealand. However, the phylogenetic affinities of this taxon are poorly known, and comparisons with other taxa have been limited. We provide an extensive redescription of Neophoca palatina and diagnose this taxon using a morphometric approach. Twenty measurements of the skull were collected for N. palatina, as well as for all extant Australasian otariids and several fossil Neophoca cinerea Perón, 1816. Using principal component analysis, we were able to segregate taxa by genus, and N. palatina was found to cluster with Neophoca according to overall size of the skull as well as increased width of the intertemporal constriction and interorbital region. N. palatina can be distinguished from all other Australasian otariids by its unusually broad basisphenoid. Discriminant function analysis supported referral of Neophoca palatina to Neophoca with very high posterior probability. These results confirm the treatment of Neophoca palatina as a distinct species of Neophoca and highlight the former broad distribution and greater tolerance for colder temperatures of this genus. These results also suggest that New Zealand may have played a pivotal role in the diversification of Southern Hemisphere otariid seals.