Publications
My published papers, book chapters etc. Click on the title of each section to download the publication PDF.
The fossil record of true seals (Family Phocidae) is mostly made up of isolated bones, some of which are type specimens. Previous studies have sought to increase referral of non-overlapping and unrelated fossils to these taxa using the ‘Ecomorphotype Hypothesis’, which stipulates that certain differences in morphology between taxa represent adaptations to differing ecology. On this basis, bulk fossil material could be lumped to a specific ecomorphotype, and then referred to species in that ecomorphotype, even if they are different bones. This qualitative and subjective method has been used often to expand the taxonomy of fossil phocids, but has never been quantitatively tested. We test the proposed ecomorphotypes using morphometric analysis of fossil and extant northern true seal limb bones, specifically principal components analysis and discriminant function analysis. A large amount of morphological overlap between ecomorphotypes, and poor discrimination between them, suggests that the ‘Ecomorphotype Hypothesis’ is not a valid approach. Further, the analysis failed to assign fossils to ecomorphotypes designated in previous studies, with some fossils from the same taxa being designated as different ecomorphotypes. The failure of this approach suggests that all fossils referred using this method should be considered to have unknown taxonomic status. In light of this, and previous findings that phocid limb bones have limited utility as type specimens, we revise the status of named fossil phocid species. We conclude that the majority of named fossil phocid taxa should be considered nomina dubia.
Pinnipeds (seals, sea lions, walruses, and their fossil relatives) are one of the most successful mammalian clades to live in the oceans. Despite a well-resolved molecular phylogeny and a global fossil record, a complete understanding of their macroevolutionary dynamics remains hampered by a lack of formal analyses that combine these two rich sources of information. We used a meta-analytic approach to infer the most densely sampled pinniped phylogeny to-date (36 recent and 93 fossil taxa) and used phylogenetic paleobiological methods to study their diversification dynamics and biogeographic history. Pinnipeds mostly diversified at constant rates. Walruses however experienced rapid turnover in which extinction rates ultimately exceeded speciation rates from 12-6 Ma, possibly due to changing sea-levels and/or competition with otariids (eared seals). Historical biogeographic analyses including fossil data allowed us to confidently identify the North Pacific and the North Atlantic (plus or minus Paratethys) as the ancestral ranges of Otarioidea (eared seals + walrus) and crown phocids (earless seals), respectively. Yet, despite the novel addition of stem pan-pinniped taxa, the region of origin for Pan-Pinnipedia remained ambiguous. These results suggest further avenues of study in pinnipeds and provide a framework for investigating other groups with substantial extinct and extant diversity.
Toothed whales (odontocetes) emit high-frequency underwater sounds (echolocate)—an extreme and unique innovation allowing them to sense their prey and environment. Their highly specialized mandible (lower jaw) allows high-frequency sounds to be transmitted back to the inner ear. Echolocation is evident in the earliest toothed whales, but little research has focused on the evolution of mandibular form regarding this unique adaptation. Here, we use a high-density, three-dimensional geometric morphometric analysis of 100 living and extinct cetacean species spanning their 50-million-year evolutionary history. Our analyses demonstrate that most shape variation is found in the relative length of the jaw and the mandibular symphysis. The greatest morphological diversity was obtained during two periods of rapid evolution: the initial evolution of archaeocetes (stem whales) in the early to mid-Eocene as they adapted to an aquatic lifestyle, representing one of the most extreme adaptive transitions known, and later on in the mid-Oligocene odontocetes as they became increasingly specialized for a range of diets facilitated by increasingly refined echolocation. Low disparity in the posterior mandible suggests the shape of the acoustic window, which receives sound, has remained conservative since the advent of directional hearing in the aquatic archaeocetes, even as the earliest odontocetes began to receive sounds from echolocation. Diet, echolocation, feeding method, and dentition type strongly influence mandible shape. Unlike in the toothed whale cranium, we found no significant asymmetry in the mandible. We suggest that a combination of refined echolocation and associated dietary specializations have driven morphology and disparity in the toothed whale mandible.
Adaptive landscapes are central to evolutionary theory, forming a conceptual bridge between micro- and macroevolution. Evolution by natural selection across an adaptive landscape should drive lineages toward fitness peaks, shaping the distribution of phenotypic variation within and among clades over evolutionary timescales. The location and breadth of these peaks in phenotypic space can also evolve, but whether phylogenetic comparative methods can detect such patterns has largely remained unexplored. Here, we characterize the global and local adaptive landscape for total body length in cetaceans (whales, dolphins, and relatives), a trait that spans 5 orders of magnitude, across their ~53 million year evolutionary history. Using phylogenetic comparative methods, we analyze shifts in long-term mean body length and directional changes in average trait values for 345 living and fossil cetacean taxa. Remarkably, we find that the global macroevolutionary adaptive landscape of cetacean body length is relatively flat, with very few peak shifts occurring after cetaceans entered the oceans. Local peaks aremore numerous and manifest as trends along branches linked to specific adaptations. These results contrast with previous studies using only extant taxa, highlighting the vital role of fossil data for understanding macroevolution. Our results indicate that adaptive peaks are dynamic and are associated with subzones of local adaptations, creating moving targets for species adaptation. In addition, we identify limits in our ability to detect some evolutionary patterns and processes and suggest that multiple approaches are required to characterize complex hierarchical patterns of adaptation in deep time.
Mysticetes (baleen whales) include the largest animals on Earth and are renowned for their songs and long-distance communication. Even so, the scope and origins of their hearing abilities remain poorly understood. Recent work on their sister clade, the toothed whales (odontocetes), has revealed notably convergent trends in the evolution of their inner ear. Here, we test whether the same applies to baleen whales via SURFACE, a phylogenetic method that fits Ornstein-Uhlenbeck models with stepwise Akaike Information Criterion to identify instances of convergent evolution. We identify a single convergent regime, including minke (Balaenoptera acutorostrata) and Bryde’s (Balaenoptera edeni) whales, which, however, is not statistically significant. We discuss potential reasons for the overall absence of convergence and suggest improvements for future work.
Within delphinoid cetaceans, snout shape is significantly correlated to diet, with long-snouted raptorial-feeding predators preying on smaller and more agile prey than shorter-snouted species. Although there have been several studies into longirostry from a functional perspective there have been no quantitative analyses of spatial variation in skull shape or how the pattern in skull shape morphospace occupation varies between assemblages. Here we analyse the cranial morphological variation of Delphinoidea assemblages. Firstly, we calculate mean and Gi* hotspot statistics of skull shape across the world’s oceans. We find that tropical and subtropical assemblages exhibit higher average measures of longirostry. This pattern is likely caused by differences in the availability of certain prey types in warmer and cooler environments. Secondly, we calculate mean pairwise distance as well as mean nearest taxon distance in functional traits between the members of 119 unique delphinoid assemblages. There was a trend for low latitude assemblages to exhibit greater overdispersion in PC1 (snout length) compared those from high latitudes. Our results suggest that ocean temperature is influential in determining the diversity, range limits and assemblage structure of delphinoid cetaceans.
Teeth are the primary tool used by most mammals to capture and process food. Over the lifetime of an individual, they progressively wear through contact with each other (attrition) and with food (abrasion), creating distinctive patterns that reflect function and diet. Unlike their terrestrial cousins, many marine mammals capture prey via suction, which so far has not been associated with a specific wear pattern. Here, we describe two new types of tooth wear across 18 species of modern marine mammal (beaked whales, belugas, killer whales, globicephalines, and various seals) that likely stem from this behaviour: “glossowear”, which primarily affects the lingual side of the crown and plausibly records piston-like tongue movements during suction feeding; and “hydrowear”, which wraps around the sides of the crown and occurs as water is expelled from the mouth. Both wear types differ from attrition and biting-related abrasion in their surface characteristics and location on the crown. Horizontal scratches suggest a physical wear process, rather than dental erosion (acid corrosion) and tooth abfraction (microfracture). Since suction specifically exploits the liquid properties of water, physical evidence of this behaviour may help to elucidate marine mammal feeding ecology and evolution. For example, glossowear is found in the toothed ancestors of baleen whales (mammalodontids, at least one aetiocetid, and likely Mystacodon), where it suggests an important role for suction in the emergence of filter feeding. By contrast, it is absent in most long-snouted toothed whales and dolphins, indicating that these animals mostly bite, rather than suck in, their prey.
The evolution of cetaceans (whales and dolphins) represents one of the most extreme adaptive transitions known, from terrestrial mammals to a highly specialized aquatic radiation that includes the largest animals alive today. Many anatomical shifts in this transition involve the feeding, respiratory, and sensory structures of the cranium, which we quantified with a high-density, three-dimensional geometric morphometric analysis of 201 living and extinct cetacean species spanning the entirety of their ∼50-million-year evolutionary history. Our analyses demonstrate that cetacean suborders occupy distinct areas of cranial morphospace, with extinct, transitional taxa bridging the gap between archaeocetes (stem whales) and modern mysticetes (baleen whales) and odontocetes (toothed whales). This diversity was obtained through three key periods of rapid evolution: first, the initial evolution of archaeocetes in the early to mid-Eocene produced the highest evolutionary rates seen in cetaceans, concentrated in the maxilla, frontal, premaxilla, and nasal; second, the late Eocene divergence of the mysticetes and odontocetes drives a second peak in rates, with high rates and disparity sustained through the Oligocene; and third, the diversification of odontocetes, particularly sperm whales, in the Miocene (∼18–10 Mya) propels a final peak in the tempo of cetacean morphological evolution. Archaeocetes show the fastest evolutionary rates but the lowest disparity. Odontocetes exhibit the highest disparity, while mysticetes evolve at the slowest pace, particularly in the Neogene. Diet and echolocation have the strongest influence on cranial morphology, with habitat, size, dentition, and feeding method also significant factors impacting shape, disparity, and the pace of cetacean cranial evolution.
Otarioids (fur seals, sea lions and the walrus, Odobenus rosmarus) are an ancient group of marine mammals that have adapted to feeding in water in a variety of ways. Fur seals and sea lions (otariids) primarily feed on fish and cephalopods, but opportunistically target a wide range of prey types and sizes, including sharks, penguins, and even their own kind. Like their terrestrial carnivoran relatives, otariids primarily rely on their teeth to catch and process their food. Suction—the ability to lower the pressure inside the oral cavity to draw in water and prey—also plays an important role, however, especially when ingested items are small. Osteological adaptations for suction are seemingly absent, but the behavior is nonetheless facilitated by the shape of the soft tissues surrounding the mouth. Walruses are suction specialists, as reflected in their robust skull, muscular lips and strong throat muscles. They primarily feed on benthic bivalves, gastropods and annelids, but sometimes also target larger prey, including birds and other pinnipeds. Their foraging activities affect vast areas of the (sub)Arctic seafloor, affecting the structure of benthic communities and leading to major increases in nutrient flux. These large-scale effects, plus a voracious appetite, make walruses a major ecosystem engineer.
Cetaceans (whales and dolphins) have some of the largest and most complex brains in the animal kingdom. When and why this trait evolved remains controversial, with proposed drivers ranging from echolocation to foraging complexity and high-level sociality. This uncertainty partially reflects a lack of data on extinct baleen whales (mysticetes), which has obscured deep-time patterns of brain size evolution in non-echolocating cetaceans. Building on new measurements from mysticete fossils, we show that the evolution of large brains preceded that of echolocation, and subsequently followed a complex trajectory involving several independent increases (e.g. in rorquals and oceanic dolphins) and decreases (e.g. in right whales and ‘river dolphins’). Echolocating whales show a greater tendency towards large brain size, thus reaffirming cognitive demands associated with sound processing as a plausible driver of cetacean encephalization. Nevertheless, our results suggest that other factors such as sociality were also important.
Modern pinnipeds (true and eared seals) employ two radically different swimming styles, with true seals (phocids) propelling themselves primarily with their hindlimbs, whereas eared seals (otariids) rely on their wing-like foreflippers. Current explanations of this functional dichotomy invoke either pinniped diphyly or independent colonizations of the ocean by related but still largely terrestrial ancestors. Here, we show that pinniped swimming styles form an anatomical, functional, and behavioral continuum, within which adaptations for forelimb swimming can arise directly from a hindlimb-propelled bauplan. Within phocids, southern seals (monachines) show a convergent trend toward wing-like, hydrodynamically efficient forelimbs used for propulsion during slow swimming, turning, bursts of speed, or when initiating movement. This condition is most evident in leopard seals, which have well-integrated foreflippers with little digit mobility, reduced claws, and hydrodynamic characteristics comparable to those of forelimb-propelled otariids. Using monachines as a model, we suggest that the last common ancestor of modern seals may have been hindlimb-propelled and aquatically adapted, thus resolving the apparent contradiction at the root of pinniped evolution.
The inner ear of the two higher clades of modern cetaceans (Neoceti) is highly adapted for hearing infrasonic (mysticetes) or ultrasonic (odontocetes) frequencies. Within odontocetes, Platanistoidea comprises a single extant riverine representative, Platanista gangetica, and a diversity of mainly extinct marine species from the late Oligocene onward. Recent studies drawing on features including the disparate tympanoperiotic have not yet provided a consensus phylogenetic hypothesis for platanistoids. Further, cochlear morphology and evolutionary patterns have never been reported. Here, we describe for the first time the inner ear morphology of late Oligocene–early Miocene extinct marine platanistoids and their evolutionary patterns. We initially hypothesized that extinct marine platanistoids lacked a specialized inner ear like P. gangetica and thus, their morphology and inferred hearing abilitieswere more similar to those of pelagic odontocetes. Our results reveal there is no “typical” platanistoid cochlear type, as the group displays a disparate range of cochlear anatomies, but all are consistent with high-frequency hearing. Stem odontocete Prosqualodon australis and platanistoid Otekaikea huata present a tympanal recess in their cochlea, of yet uncertain function in the hearing mechanism in cetaceans. The more basal morphology of Aondelphis talen indicates it had lower high-frequency hearing than other platanistoids. Finally, Platanista has the most derived cochlear morphology, adding to evidence that it is an outlier within the group and consistent with a >9-Myr-long separation from its sister genus Zarhachis. The evolution of a singular sound production morphology within Platanistidae may have facilitated the survival of Platanista to the present day.
Unlike most mammals, toothed whale (Odontoceti) skulls lack symmetry in the nasal and facial (nasofacial) region. This asymmetry is hypothesised to relate to echolocation, which may have evolved in the earliest diverging odontocetes. Early cetaceans (whales, dolphins, and porpoises) such as archaeocetes, namely the protocetids and basilosaurids, have asymmetric rostra, but it is unclear when nasofacial asymmetry evolved during the transition from archaeocetes to modern whales. We used three-dimensional geometric morphometrics and phylogenetic comparative methods to reconstruct the evolution of asymmetry in the skulls of 162 living and extinct cetaceans over 50 million years. In archaeocetes, we found asymmetry is prevalent in the rostrum and also in the squamosal, jugal, and orbit, possibly reflecting preservational deformation. Asymmetry in odontocetes is predominant in the nasofacial region. Mysticetes (baleen whales) show symmetry similar to terrestrial artiodactyls such as bovines. The first significant shift in asymmetry occurred in the stem odontocete family Xenorophidae during the Early Oligocene. Further increases in asymmetry occur in the physeteroids in the Late Oligocene, Squalodelphinidae and Platanistidae in the Late Oligocene/Early Miocene, and in the Monodontidae in the Late Miocene/Early Pliocene. Additional episodes of rapid change in odontocete skull asymmetry were found in the Mid-Late Oligocene, a period of rapid evolution and diversification. No high-probability increases or jumps in asymmetry were found in mysticetes or archaeocetes. Unexpectedly, no increases in asymmetry were recovered within the highly asymmetric ziphiids, which may result from the extreme, asymmetric shape of premaxillary crests in these taxa not being captured by landmarks alone. Early ancestors of living whales had little cranial asymmetry and likely were not able to echolocate. Archaeocetes display high levels of asymmetry in the rostrum, potentially related to directional hearing, which is lost in early neocetes—the taxon including the most recent common ancestor of living cetaceans. Nasofacial asymmetry becomes a significant feature of Odontoceti skulls in the Early Oligocene, reaching its highest levels in extant taxa. Separate evolutionary regimes are reconstructed for odontocetes living in acoustically complex environments, suggesting that these niches impose strong selective pressure on echolocation ability and thus increased cranial asymmetry.
Right whales (Balaenidae) are the most distinctive family of extant baleen whales, thanks to their highly arched rostrum, tall lips and robust body shape. They are also the oldest, originating as much as 20 million years ago (Ma). Nevertheless, their fossil record is patchy and frequently understudied, obscuring their evolution. Here, we describe a new stem balaenid, Antwerpibalaena liberatlas, from northern Belgium, adding to the rich but historically problematic baleen whale assemblage of the Pliocene North Sea. Within right whales, Antwerpibalaena forms a clade with two previously described extinct genera, Balaenella and Balaenula. The holotype preserves much of the postcranial skeleton, and informs the emergence of typical balaenid traits like fused neck vertebrae and paddle-shaped flippers. Its size is intermediate between that of extant right whales and most of their extinct forebears revealing a more complex pattern of balaenid size evolution than previously thought.
In morphological traits, variation within species is generally considered to be lower than variation among species, although this assumption is rarely tested. This is particularly important in fields like palaeontology, where it is common to use a single individual as representative of a species due to the rarity of fossils. Here, we investigated intraspecific variation in the cochleae of harbour porpoises (Phocoena phocoena). Interspecific variation of cochlear morphology is well characterised among odontocetes (toothed whales) because of the importance of the structure in echolocation, but generally these studies use only a single cochlea to represent each species. In this study we compare variation within the cochleae of 18 specimens of P. phocoena with variations in cochlear morphology across 51 other odontocete species. Using both 3D landmark and linear measurement data, we performed Generalised Procrustes and principal component analyses to quantify shape variation. We then quantified intraspecific variation in our sample of P. phocoena by estimating disparity and the coefficient of variation for our 3D and linear data respectively. Finally, to determine whether intraspecific variation may confound the results of studies of interspecific variation, we used multivariate and univariate analyses of variance to test whether variation within the specimens of P. phocoena was significantly lower than that across odontocetes. We found low levels ofintraspecific variation in the cochleae of P. phocoena, and that cochlear shape within P. phocoena was significantly less variable than across odontocetes. Although future studies should attempt to use multiple cochleae for every species, our results suggest that using just one cochlea for each species should not strongly influence the conclusions of comparative studies if our results are consistent across Cetacea.
Sound plays a crucial role in marine mammal ecology, and has led numerous species to evolve a diverse vocal repertoire (Dudzinski & Gregg, 2017; Winn & Schneider, 1977). Nonvocal auditory behaviors like flipper slaps and breaches are also important, but more limited in scope and thought to occur mostly at the surface of the water (Würsig & Whitehead, 2017). Here, we report a novel type of percussive signaling in wild gray seals (Halichoerus grypus), which demonstrates that nonvocal auditory behaviors may also be produced entirely underwater.
Background: Odontocetes (toothed whales) are the most species-rich marine mammal lineage. The catalyst for their evolutionary success is echolocation - a form of biological sonar that uses high-frequency sound, produced in the forehead and ultimately detected by the cochlea. The ubiquity of echolocation in odontocetes across a wide range of physical and acoustic environments suggests that convergent evolution of cochlear shape is likely to have occurred. To test this, we used SURFACE; a method that fits Ornstein-Uhlenbeck (OU) models with stepwise AIC (Akaike Information Criterion) to identify convergent regimes on the odontocete phylogeny, and then tested whether convergence in these regimes was significantly greater than expected by chance.
Results: We identified three convergent regimes: (1) True’s (Mesoplodon mirus) and Cuvier’s (Ziphius cavirostris) beaked whales; (2) sperm whales (Physeter macrocephalus) and all other beaked whales sampled; and (3) pygmy (Kogia breviceps) and dwarf (Kogia sima) sperm whales and Dall’s porpoise (Phocoenoides dalli). Interestingly the ‘river dolphins’, a group notorious for their convergent morphologies and riverine ecologies, do not have convergent cochlear shapes. The first two regimes were significantly convergent, with habitat type and dive type significantly correlated with membership of the sperm whale + beaked whale regime.
Conclusions: The extreme acoustic environment of the deep ocean likely constrains cochlear shape, causing the cochlear morphology of sperm and beaked whales to converge. This study adds support for cochlear morphology being used to predict the ecology of extinct cetaceans.
Mesoplodont beaked whales are one of the most enigmatic mammalian genera. We document a pod of four beaked whales in the Bay of Biscay breaching and tail slapping alongside a large passenger ferry. Photographs of the animals were independently reviewed by experts, and identified as True's beaked whales (Mesoplodon mirus). This is the first conclusive live sighting of these animals in the north-east Atlantic, and adds information to previous sightings that are likely to have been M. mirus. Photographs of an adult male appears to show two supernumerary teeth posterior to the apical mandibular tusks. Whilst analysed museum specimens (nD8) did not show evidence of alveoli in this location, there is evidence of vestigial teeth and variable dentition in many beaked whale species. This is the first such record of supernumerary teeth in True's beaked whales.
Neobalaenines are an enigmatic group of baleen whales represented today by a single living species: the pygmy right whale, Caperea marginata, found only in the Southern Hemisphere. Molecular divergence estimates date the origin of pygmy right whales to 2226 Ma, yet so far there are only three confirmed fossil occurrences. Here, we describe an isolated periotic from the latest Miocene of Victoria (Australia). The new fossil shows all the hallmarks of Caperea, making it the second-oldest described neobalaenine, and the oldest record of the genus. Overall, the new specimen resembles C. marginata in its external morphology and details of the cochlea, but is more archaic in it having a hypertrophied suprameatal area and a greater number of cochlear turns. The presence of Caperea in Australian waters during the Late Miocene matches the distribution of the living species, and supports a southern origin for pygmy right whales.
Kienle et al. [1] suggest amendments to our framework for feeding in predatory aquatic mammals [2]. Below we reply to their suggestions and demonstrate that they are fundamentally flawed from both a mechanical (feeding cycle, strategies) and an evolutionary perspective. They do, however, inspire an important addition to the range and structuring of capture behaviours encoded in our framework.
Cetaceans (whales and dolphins) primarily use sound to communicate and hunt for prey. Their auditory anatomy is highly specialised, but much about its function remains unknown. In particular, a feature of the cochlea known as the tympanal recess present in some mysticetes (baleen whales) and odontocetes (toothed whales) has defied functional explanation. Here, we present and discuss several hypotheses that may clarify the function and evolution of the tympanal recess. One potential function in particular, the vibroacoustic duct mechanism, seems most plausible although further work is needed to test the hypothesis, which hints at the possibility of sperm whales and beaked whales being able to detect both high and low frequencies.
The pygmy right whale, Caperea marginata, is the least understood extant baleen whale (Cetacea, Mysticeti). Knowledge on its basic anatomy, ecology, and fossil record is limited, even though its singular position outside both balaenids (right whales) and balaenopteroids (rorquals1 + grey whales) gives Caperea a pivotal role in mysticete evolution. Recent investigations of the cetacean cochlea have provided new insights into sensory capabilities and phylogeny. Here, we extend this advance to Caperea by describing, for the first time, the inner ear of this enigmatic species. The cochlea is large and appears to be sensitive to low-frequency sounds, but its hearing limit is relatively high. The presence of a well-developed tympanal recess links Caperea with cetotheriids and balaenopteroids, rather than balaenids, contrary to the traditional morphological view of a close Caperea-balaenid relationship. Nevertheless, a broader sample of the cetotheriid Herpetocetus demonstrates that the presence of a tympanal recess can be variable at the specific and possibly even the intraspecific level.
Extant aquatic mammals are a key component of aquatic ecosystems. Their morphology, ecological role and behaviour are, to a large extent, shaped by their feeding ecology. Nevertheless, the nature of this crucial aspect of their biology is often oversimplified and, consequently, misinterpreted. Here, we introduce a new framework that categorizes the feeding cycle of predatory aquatic mammals into four distinct functional stages (prey capture, manipulation and processing, water removal and swallowing), and details the feeding behaviours that can be employed at each stage. Based on this comprehensive scheme, we propose that the feeding strategies of living aquatic mammals form an evolutionary sequence that recalls the land-to-water transition of their ancestors. Our new conception helps to explain and predict the origin of particular feeding styles, such as baleen-assisted filter feeding in whales and raptorial ‘pierce’ feeding in pinnipeds, and informs the structure of present and past ecosystems.
Living baleen whales (mysticetes) produce and hear the lowest-frequency (infrasonic) sounds among mammals. There is currently debate over whether the ancestor of crown cetaceans (Neoceti) was able to detect low frequencies. However, the lack of information on the most archaic fossil mysticetes has prevented us from determining the earliest evolution of their extreme acoustic biology. Here, we report the first anatomical analyses and frequency range estimation of the inner ear in Oligocene (34–23 Ma) fossils of archaic toothed mysticetes from Australia and the USA. The cochlear anatomy of these small fossil mysticetes resembles basilosaurid archaeocetes, but is also similar to that of today’s baleen whales, indicating that even the earliest mysticetes detected low-frequency sounds, and lacked ultrasonic hearing and echolocation. This suggests that, in contrast to recent research, the plesiomorphic hearing condition for Neoceti was low frequency, which was retained by toothed mysticetes, and the high-frequency hearing of odontocetes is derived. Therefore, the low-frequency hearing of baleen whales has remained relatively unchanged over the last approximately 34 Myr, being present before the evolution of other signature mysticete traits, including filter feeding, baleen and giant body size.
The origin of baleen, the key adaptation of modern whales (Mysticeti), marks a profound yet poorly understood transition in vertebrate evolution, triggering the rise of the largest animals on Earth. Baleen is thought to have appeared in archaic tooth-bearing mysticetes during a transitional phase that combined raptorial feeding with incipient bulk filtering. Here we show that tooth wear in a new Late Oligocene mysticete belonging to the putatively transitional family Aetiocetidae is inconsistent with the presence of baleen, and instead indicative of suction feeding. Our findings suggest that baleen arose much closer to the origin of toothless mysticete whales than previously thought. In addition, they suggest an entirely new evolutionary scenario in which the transition from raptorial to baleen-assisted filter feeding was mediated by suction, thereby avoiding the problem of functional interference between teeth and the baleen rack.
Australia has a fossil record of penguins reaching back to the Eocene, yet today is inhabited by just one breeding species, the little penguin Eudyptula minor. The description of recently collected penguin fossils from the re-dated upper Miocene Port Campbell Limestone of Portland (Victoria), in addition to reanalysis of previously described material, has allowed the Cenozoic history of penguins in Australia to be placed into a global context for the first time. Australian pre-Quaternary fossil penguins represent stem taxa phylogenetically disparate from each other and E. minor, implying multiple dispersals and extinctions. Late Eocene penguins from Australia are closest to contemporaneous taxa in Antarctica, New Zealand and South America. Given current material, the Miocene Australian fossil penguin fauna is apparently unique in harbouring ‘giant penguins’ after they went extinct elsewhere; and including stem taxa until at least 6 Ma, by which time crown penguins dominated elsewhere in the southern hemisphere. Separation of Australia from Antarctica during the Palaeogene, and its subsequent drift north, appears to have been a major event in Australian penguin biogeography. Increasing isolation through the Cenozoic may have limited penguin dispersal to Australia from outside the Australasian region, until intensification of the eastwards-flowing Antarctic Circumpolar Current in the mid-Miocene established a potential new dispersal vector to Australia.
The evolution of biosonar (production of high-frequency sound and reception of its echo) was a key innovation of toothed whales and dolphins (Odontoceti) that facilitated phylogenetic diversification and rise to ecological predominance. Yet exactly when high-frequency hearing first evolved in odontocete history remains a fundamental question in cetacean biology. Here, we show that archaic odontocetes had a cochlea specialized for sensing high-frequency sound, as exemplified by an Oligocene xenorophid, one of the earliest diverging stem groups. This specialization is not as extreme as that seen in the crown clade. Paired with anatomical correlates for high-frequency signal production in Xenorophidae, this is strong evidence that the most archaic toothed whales possessed a functional biosonar system, and that this signature adaptation of odontocetes was acquired at or soon after their origin.
Pseudaptenodytes macraei is redescribed in detail using current avian anatomical terminology. The age of the species is constrained to the late Miocene, and two humeral autapomorphies are identified for the species: a flattened elliptical ventral portion of the fossa tricipitalis and a curved margo cranialis lacking a preaxial angle. Pseudaptenodytes macraei is a distinct taxon based on a diagnostic-type specimen, but it is recommended that the nominal congener Pseudaptenodytes minor be designated a nomen dubium pending the discovery of more complete material.
Australian fossil penguins (Sphenisciformes) are reviewed as a basis for future primary research. The five named species are based on type specimens of Eocene, Miocene—Pliocene and Holocene age collected from South Australia, Victoria and Tasmania. The phylogenetic affinities of these taxa remain unresolved. Only one type specimen is represented by clearly associated elements of a skeleton; the rest are single bones (isolated partial humeri and a pelvis). Further research is required to establish the taxonomic status of Pachydyptes simpsoni, Anthropodyptes gilli, Pseudaptenodyes macraei, ?Pseudaptenodytes minor and Tasidyptes hunteri. Additional described specimens include isolated postcranial elements from the Late Oligocene of South Australia and Late Miocene—Early Pliocene of Victoria. Other Miocene and Pliocene specimens are housed in Museum Victoria. These specimens have the potential to shed light on the Neogene palaeobiogeography and diversification of crown group penguins.
An incomplete tarsometatarsus identified as an indeterminate species of Dromornithidae is described from the upper Miocene–lower Pliocene shallow marine Black Rock Sandstone at Beaumaris, Victoria, Australia. This isolated specimen represents one of the few pre-Pleistocene dromornithids with a well-constrained geologic age. Additionally, it is one of the few pre-Quaternary dromornithid fossils recorded from southeast Australia. Comparisons with known dromornithid taxa suggest that the Beaumaris dromornithid is distinct from previously established species. This hitherto unknown species of dromornithid in the late Neogene of southeastern Australia cautions against deriving evolutionary patterns solely on the basis of fossils from northern Australia.
The Pelagornithidae, or ‘giant bony-toothed birds,’ are enigmatic extinct seabirds with a long history spanning the late Paleocene–late Pliocene (Harrison, 1985; Mourer-Chauvire and Geraads, 2008, 2010; Bourdon, 2011). In parallel with their extensive chronostratigraphic distribution, pelagornithids have thus far been recorded from all continents with the notable exception of Australia (Mayr, 2011; contra Boessenecker and Smith, 2011). Here we document the first evidence of Pelagornithidae in Australia: a diagnostic Pelagornis tibiotarsus and non-associated fragmentary appendicular elements from the Mio–Pliocene of Beaumaris, Victoria (Fig. 1). This discovery confirms the distribution of pelagornithids on every continent and the global distribution of the genus Pelagornis during the late Neogene.