The petrosal and inner ear of the Late Jurassic cladotherian mammal Dryolestes leiriensis and implications for ear evolution in therian mammals

Dryolestes leiriensis is a Late Jurassic fossil mammal of the dryolestoid superfamily in the cladotherian clade that includes the extant marsupials and placentals. We used high resolution micro‐computed tomography (µCT) scanning and digital reconstruction of the virtual endocast of the inner ear to show that its cochlear canal is coiled through 270°, and has a cribriform plate with the spiral cochlear nerve foramina between the internal acoustic meatus and the cochlear bony labyrinth. The cochlear canal has the primary bony lamina for the basilar membrane with a partially formed (or partially preserved) canal for the cochlear spiral ganglion. These structures, in their fully developed condition, form the modiolus (the bony spiral structure) of the fully coiled cochlea in extant marsupial and placental mammals. The CT data show that the secondary bony lamina is present, although less developed than in another dryolestoid Henkelotherium and in the prototribosphenidan Vincelestes. The presence of the primary bony lamina with spiral ganglion canal suggests a dense and finely distributed cochlear nerve innervation of the hair cells for improved resolution of sound frequencies. The primary, and very probably also the secondary, bony laminae are correlated with a more rigid support for the basilar membrane and a narrower width of this membrane, both of which are key soft‐tissue characteristics for more sensitive hearing for higher frequency sound. All these cochlear features originated prior to the full coiling of the therian mammal cochlea beyond one full turn, suggesting that the adaptation to hearing a wider range of sound frequencies, especially higher frequencies with refined resolution, has an ancient evolutionary origin no later than the Late Jurassic in therian evolution. The petrosal of Dryolestes has added several features that are not preserved in the petrosal of Henkelotherium. The petrosal characters of dryolestoid mammals are essentially the same as those of Vincelestes, helping to corroborate the synapomorphies of the cladotherian clade in neural, vascular, and other petrosal characteristics. The petrosal characteristics of Dryolestes and Henkelotherium together represent the ancestral morphotype of the cladotherian clade (Dryolestoidea + Vincelestes + extant Theria) from which the extant therian mammals evolved their ear region characteristics. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166 , 433–463.     

Digital Reconstruction of the Otic Region and Inner Ear of the Non-Mammalian Cynodont Brasilitherium riograndensis (Late Triassic, Brazil) and Its Relevance to the Evolution of the Mammalian Ear

The external anatomy of the petrosal, the bony labyrinth of the inner ear, and the stapes of Brasilitherium riograndensis (specimen UFRGS-PV-1043-T) were investigated by digital 3D reconstructions based on μCT scan images. Brasilitherium is the most basal taxon bearing a distinct promontorium, although less inflated than that of Morganucodon and still lacking a flat medial facet. A bony wall formed by the petrosal separates the cochlear canal and the vestibule from the brain cavity, with an internal acoustic meatus bearing distinct foramina for the facial nerve (VII) and vestibulocochlear nerve (VIII). The semicircular canals are irregular in shape, the anterior canal being the largest and the lateral one the smallest. Brasilitherium has an elongated but straight cochlear canal. The stapes resembles the morphology of derived non-mammaliaform cynodonts, such as Probainognathus and Pachygenelus, and differs from Thrinaxodon. By the allometric relationship of the cochlear canal and the estimated body mass, Brasilitherium can be grouped with Yunnanodon and Morganucodon in a regression line, which is below the line of mammals and above the line of non-avian reptiles. Brasilitherium fits in a sequence of gradual elongation of the cochlear canal associated with the enhancement in the capacity to hear higher frequencies. Among the constraints that might have triggered these transformations in small, insectivorous, and possibly nocturnal Mesozoic cynodont taxa is the improvement of detecting acoustically active insects.     

Homoplasy in the ear region of Tethytheria and the systematic position of Embrithopoda (Mammalia,Afrotheria)

The ear region of mammals has long been considered as morphologically very conservative and accordingly, phylogenetically useful. In this study, the anatomy of the petrosal and bony labyrinth (osseous inner ear) of Numidotherium (Proboscidea) and Arsinoitherium (Embrithopoda) are investigated and compared in order to assess the evolution of ear region characters in proboscideans and embrithopods. Using a cladistic analysis across Paenungulata based on ear region characters only, we found that Arsinoitherium is surprisingly best placed as a crown proboscideans to the exclusion of Numidotherium and Phosphatherium, which results in the paraphyly of proboscidean. The clade Proboscidea is actually well supported by dental and post-cranial characters, and we propose that this result underlines the great amount of morphological convergences in the ear region of Embrithopoda and Proboscidea, possibly due to convergent evolution of capabilities toward infrasonic hearing.     

Petrosal and Bony Labyrinth Morphology Supports Paraphyly of Elephantulus Within Macroscelididae (Mammalia,Afrotheria)

Interest in the phylogeny of Macroscelididae (sengis or elephant shrews) has been prompted by molecular studies indicating that Elephantulus rozeti is best placed as the sister group of Petrodromus tetradactylus (this clade being in turn the sister taxon to Macroscelides proboscideus) than among other species of the genus Elephantulus. Until now, no discrete morphological characters have been proposed to support the grouping of E. rozeti, Petrodromus, and Macroscelides into this single so-called ‘Panelephantulus’ clade. Here, we employed μCT scanning in order to investigate the petrosal and bony labyrinth (bony capsule of the inner ear) morphology of most species of extant Macroscelididae. We performed a cladistic analysis on ear traits and found that despite some convergences (e.g., concerning the bony arterial canals in Macroscelides and Rhynchocyon) the middle and inner ear morphology furnishes significant support for the ‘Panelephantulus’ clade. In our analysis, this clade is unambigously supported by the presence of a fully ossified stapediofacial tube. Two additional characters (the presence of a bony septum at the mouth of the fenestra cochleae dividing the D3 sinus into two distinct cavities and the absence of an accessory lateral pneumatic fossa) could also support ‘Panelephantulus.’ These newly discovered morphological characters support the molecular phylogenies published and highlight the importance of coding hitherto difficult to sample morphologies within cladistic analyses using micro-CT techniques. Taxonomic implications are briefly discussed.     

Cranial Morphology of a Pantolestid Eutherian Mammal from the Eocene Bridger Formation,Wyoming, USA: Implications for Relationships and Habitat

Pantolestinae is a eutherian subfamily of mammals whose members are known from the middle early Paleocene through at least the beginning of the Oligocene of North America. They are also known from Europe, and possibly Africa. A lack of information on pantolestine skulls has prevented the use of cranial anatomy in evaluation of this group’s enigmatic higher-level phylogenetic relationships. Conversely, postcranial skeletons are well known and locomotor interpretations based on them are robust. The most complete known skull of a pantolestine, Pantolestes longicaudus (YPM 13525), is described here and compared to potential close fossil relatives and extant mammals. Semicircular canal morphology is used to test locomotor hypotheses. YPM 13525 lacks an ossified bulla. It has a mediolaterally broad basioccipital, a large entoglenoid process, and a deeply incised glaserian fissure of the squamosal, caudal and rostral tympanic processes on the petrosal, a foramen for an internal carotid artery (ICA) that entered the tympanic cavity from a posteromedial position, bony tubes enclosing the main stem and transpromontorial branch of the ICA, a large anterior carotid foramen formed within the basisphenoid, evidence of a stapedial artery ramus superior, a groove on the dorsal aspect of the basisphenoid leading to the piriform fenestra possibly for drainage of the cavernous sinus to an extracranial inferior petrosal sinus, a dorsum sellae with well-developed posterior clinoid processes, a foramen rotundum within the alisphenoid, and a sphenorbital fissure between the alisphenoid and orbitosphenoid. Overall, the morphology is not strikingly similar to any potential close relative and the phylogenetic position of Pantolestinae cannot be estimated without cladistic analysis of a character matrix that includes this new morphology and broadly samples extant and extinct eutherian taxa. Semicircular canal morphology differs from that of two likely terrestrial Paleocene mammals, Aphronorus (another pantolestid) and Eoryctes (a palaeoryctid), suggesting a different, possibly semi-aquatic, lifestyle for Pantolestes.     

Cranial morphology of the early tertiary Phenacolemur and its bearing on primate phylogeny

An early Eocene skull of the paromomyid Phenacolemur, a plesiadapoid primate, is described with particular emphasis on the ear region. The auditory bulla is composed of the petrosal and of a large ectotympanic plate which is outside of the bulla. The preserved morphology of the middle ear is distinctly more primitive than that of the older Plesiadapis. It cannot be determined with certainty whether Phenacolemur had the carotid circulation enclosed in bony tubes or not. The auditory bulla of early primates and relevant living ones is discussed and it is suggested that an extrabullar ectotympanic, as seen in all non-lemuriform fossil and extant primates, was probably the primitive ordinal condition, rather than the intrabullar ring-like ectotympanic in the Lemuriformes. Aspects of the carotid circulation are discussed as they pertain to the relationship of early Tertiary primates, living Tarsiiformes, Lemuriformes and Lorisiformes.     

Systematics,phylogeny and evolutionary pattern of the hystricognath rodent Eumysops (Echimyidae) from the Plio–Pleistocene of southern South America

?Eumysops is a peculiar representative of the currently tropical family Echimyidae, which evolved in increasingly dry and cold Plio–Pleistocene environments of southern South America. The results of a systematic and stratigraphic review of the genus, and of phylogenetic analyses based on both morphology and a combined morphological–molecular dataset in the context of extant representatives, are presented here. Recognised diversity includes four previously described species plus a new one from the late Pliocene. These species form a well-supported monophyletic clade, sister to the late Miocene ?Pampamys and the extant Thrichomys. The position of ?Eumysops–?Pampamys–Thrichomys in a major clade including non-‘eumysopine’ echimyids constrains the traditional taxon Eumysopinae only to these three genera. Phylogeny and stratigraphic distribution of ?Eumysops species suggest an essentially cladogenetic evolutionary pattern. Beyond this, a gradual directional change, involving increase in size and in molar hypsodonty, is shown by ?Eumysops chapalmalensis as part of a late Pliocene faunal turnover interpreted as a local representation of the 2.5-Ma cooling global event. Distinctive skeletal and dental anatomy of ?Eumysops, including large orbits, shortened braincase, marked hypsodonty and postcranial specialisations, would be a result of its southern history related to a particular palaeoclimatic context.     

Petrosal and inner ear anatomy and allometry amongst specimens referred to Litopterna (Placentalia)

New isolated petrosals from the Itaboraí beds of Brazil (late Palaeocene or early Eocene) are here described and referred to the early diverging litoptern Miguelsoria parayirunhor, based on phylogenetic, size, and abundance arguments. Both the external and internal anatomy of these specimens were investigated, which for the first time document many details of the auditory region of a Palaeogene litoptern. Our cladistic analysis, which included our new observations, failed to recover a monophyletic Litopterna but did not exclude it. A constrained analysis for the monophyly of this order showed that several features such as a (sub)quadrangular and anteroposteriorly elongated tensor tympani fossa and a large notch in the vicinity of the external opening of the cochlear canaliculus may constitute synapomorphies for Litopterna. The evolution of several other auditory characters amongst Litopterna is discussed and the relative dimensions of the inner ear and surrounding petrosal in the group were also investigated. This allowed detection of negative allometry of the bony labyrinth within the petrosal, which was confirmed by measurements and regression analysis across a larger sample of placental mammals. This scaling effect probably has an important influence on several characters of the bony labyrinth and petrosal, amongst which are the length of the vestibular aqueduct and cochlear canaliculus. It demonstrates that many aspects of the morphological variation of the bony labyrinth need to be thoroughly investigated before being incorporated into phylogenetic analyses. © 2015 The Linnean Society of London     

A SKULL OF THE GIANT BONY‐TOOTHED BIRD DASORNIS (AVES: PELAGORNITHIDAE) FROM THE LOWER EOCENE OF THE ISLE OF SHEPPEY

Abstract: The first substantial skull of a very large Paleogene bony‐toothed bird (Pelagornithidae) is described from the Lower Eocene London Clay of the Isle of Sheppey in England. The specimen is assigned to Dasornis emuinus (Bowerbank), based on a taxonomic revision of the large London Clay Pelagornithidae. Very large bony‐toothed birds from the London Clay were known previously from fragmentary remains of non‐comparable skeletal elements only, and Dasornis londinensis Owen, Argillornis emuinus (Bowerbank), A. longipennis Owen, and Neptuniavis miranda Harrison and Walker are considered junior synonyms of D. emuinus. The new specimen allows a definitive assignment of Dasornis to the Pelagornithidae and documents that this taxon closely resembles other bony‐toothed birds in cranial morphology. It is hypothesized that giant size (i.e. a wingspan above 4 m) evolved only once within Pelagornithidae and that Dasornis emuinus is the sister taxon of the giant Neogene bony‐toothed birds, which share a derived wing morphology.     

The femur of extinct bunodont otters in Africa (Carnivora,Mustelidae, Lutrinae)

This study compares fossil femora attributed to extinct African bunodont lutrines with extant mustelids and ursids to reconstruct locomotor behavior. Due to the immense size differences among taxa, shape data were used to compare morphology. Based on morphological differences, the fossil femora are suggested to belong to different taxa with different locomotor abilities and habitat preferences. The Langebaanweg femur is the oldest and has a typical mustelid morphology suggesting that it was a locomotor generalist like most mustelids. The West Turkana form is more like extant nonbunodont otters, but much larger, and may have belonged to a semiaquatic taxon. The enormous Omo femur shares some features with truly aquatic taxa (e.g., Enhydra) and is the most likely to have been fully aquatic. The same may hold true for the Hadar species as it is most similar to that from the Omo. If these femora truly belong to bunodont lutrines, then they are more diverse in postcranial morphology than in dental morphology.     

The inner ear of <Emphasis Type="Italic">Protungulatum</Emphasis> (Pan-Euungulata,Mammalia)

We present new anatomical details about the bony labyrinth of Protungulatum based on micro CT-scan investigation of an isolated petrosal bone retrieved at the Puercan locality of Bug Creek Anthills and referred to Protungulatum sp. The exceptional state of preservation of the specimen allowed us to reconstruct the very fine details of the inside of the petrosal bone, including the bony labyrinth, the innervation of the vestibule and the innervation and vasculature of the cochlea. Estimation of the auditory capability of Protungulatum based on cochlear morphology indicate that Protungulatum was specialized for high-frequency hearing, with estimated low frequency limits above 1 KHz. Comparisons with Late Cretaceous non-placental eutherians and with early Tertiary pan-euungulates indicate that the bony labyrinth of Protungulatum is closer in general morphology to Mesozoic forms (low coiling and low aspect ratio of the cochlea, posterior orientation of the common crus, dorsal outpocketing of the cochlear fossula), and shares only a few characters with pan-euungulate and euungulate taxa. Interestingly, the bony labyrinth of Protungulatum also shares some morphological features with South American notoungulates and litopterns recently described from Itaboraí, Brazil. These new observations provide new morphological features of potential phylogenetic interest.     

STRUCTURE AND FUNCTION IN WHALE EARS

ABSTRACT

Ultrasonic echolocation abilities are well documented in several dolphin species, but hearing characteristics are unknown for most whales. Vocalization data suggest whale hearing spans infra- to ultrasonic ranges. This paper presents an overview of whale ear anatomy and analyzes 1) how whale ears are adapted for underwater hearing and 2) how inner ear differences relate to different hearing capacities among whales.

Whales have adaptations for rapid, deep diving and long submersion; e.g., broad- bore Eustachian tubes, no pinnae, and no air-filled external canals, that impact sound reception. In odontocetes, two soft tissue channels conduct sound to the ear. In mysticetes, bone and soft tissue conduction are likely. The middle ear is air-filled but has an extensible mucosa. Cochlear structures are hypertrophied and vestibular components are reduced. Auditory ganglion cell densities are double land mammal averages (2000–4000/mm). Basilar membrane lengths range 20–70 mm; gradients are larger than in terrestrial mammals. Odontocetes have 20–60% bony membrane support and basal ratios >0.6, consistent with hearing >150 kHz. Mysticetes have apical ratios <0.002 and no bony lateral support, implying acute infrasonic hearing. Cochlear hypertrophy may be adaptive for high background noise. Vestibular loss is consistent with cervical fusion. Exceptionally high auditory fiber counts suggest both mysticetes and odontocetes have ears “wired” for more complex signal processing mechanisms than most land mammals.     

Osteology Supports a Stem-Galliform Affinity for the Giant Extinct Flightless Bird Sylviornis neocaledoniae (Sylviornithidae,Galloanseres)

The giant flightless bird Sylviornis neocaledoniae (Aves: Sylviornithidae) existed on La Grande Terre and Ile des Pins, New Caledonia, until the late Holocene when it went extinct shortly after human arrival on these islands. The species was generally considered to be a megapode (Megapodiidae) until the family Sylviornithidae was erected for it in 2005 to reflect multiple cranial autapomorphies. However, despite thousands of bones having been reported for this unique and enigmatic taxon, the postcranial anatomy has remained largely unknown. We rectify this deficiency and describe the postcranial skeleton of S. neocaledoniae based on ~600 fossils and use data from this and its cranial anatomy to make a comprehensive assessment of its phylogenetic affinities. Sylviornis neocaledoniae is found to be a stem galliform, distant from megapodiids, and the sister taxon to the extinct flightless Megavitiornis altirostris from Fiji, which we transfer to the family Sylviornithidae. These two species form the sister group to extant crown-group galliforms. Several other fossil galloanseres also included in the phylogenetic analysis reveal novel hypotheses of their relationships as follows: Dromornis planei (Dromornithidae) is recovered as a stem galliform rather than a stem anseriform; Presbyornis pervetus (Presbyornithidae) is the sister group to Anseranatidae, not to Anatidae; Vegavis iaai is a crown anseriform but remains unresolved relative to Presbyornis pervetus, Anseranatidae and Anatidae. Sylviornis neocaledoniae was reconstructed herein to be 0.8 m tall in a resting stance and weigh 27–34 kg. The postcranial anatomy of S. neocaledoniae shows no indication of the specialised adaptation to digging seen in megapodiids, with for example, its ungual morphology differing little from that of chicken Gallus gallus. These observations and its phylogenetic placement as stem galliforms makes it improbable that this species employed ectothermic incubation or was a mound-builder. Sylviornis neocaledoniae can therefore be excluded as the constructor of tumuli in New Caledonia.     

Bony labyrinth morphometry indicates locomotor adaptations in the squirrel-related clade (Rodentia,Mammalia)

The semicircular canals (SCs) of the inner ear detect angular acceleration and are located in the bony labyrinth of the petrosal bone. Based on high-resolution computed tomography, we created a size-independent database of the bony labyrinth of 50 mammalian species especially rodents of the squirrel-related clade comprising taxa with fossorial, arboreal and gliding adaptations. Our sampling also includes gliding marsupials, actively flying bats, the arboreal tree shrew and subterranean species. The morphometric anatomy of the SCs was correlated to the locomotion mode. Even if the phylogenetic signal cannot entirely be excluded, the main significance for functional morphological studies has been found in the diameter of the SCs, whereas the radius of curvature is of minor interest. Additionally, we found clear differences in the bias angle of the canals between subterranean and gliding taxa, but also between sciurids and glirids. The sensitivity of the inner ear correlates with the locomotion mode, with a higher sensitivity of the SCs in fossorial species than in flying taxa. We conclude that the inner ear of flying and gliding mammals is less sensitive due to the large information flow into this sense organ during locomotion.     

Functional morphological adaptations of the bony labyrinth in marsupials (Mammalia,Theria)

Diprotodontia represents the largest and ecologically most distinct order of marsupials occurring in Australasian being highly divers in size, locomotion, habitat preferences, feeding, and activity pattern. The spatial orientation in the habitat and therefore the three‐dimensional space is detected by the vestibular system of the inner ear, more precisely by the three semicircular canals. In this study, we investigated the bony labyrinth of diprotodontian and selected non‐diprotodontian marsupial mammals of almost all genera with noninvasive micro‐CT scanning and 3D‐reconstructions. In principal component analyses, the subterranean taxon can be separated from gliding and saltatorial taxa, whereas arboreal species can be separated from saltatorial specimens. The highest PCA loadings of this functional distinction are clearly found in the diameter of the semicircular canals, whereas the overall shape (height, width, length) of the semicircular canals is less important. Additionally, the investigated arboreal and fossorial species of South America are nested in the morphospace of the Australasian taxa. Even if a phylogenetic signal in the anatomy of the bony labyrinth cannot be excluded entirely, the main functional morphological signal of the vestibular system is found in the diameter of the semicircular canals. With the large dataset of extant marsupial mammals analysed here, the locomotion mode of extinct taxa can be inferred in future studies independent of any evidence of postcranial material.     

Additional postcranial elements of Teilhardina belgica: The oldest European primate

Teilhardina belgica is one of the earliest fossil primates ever recovered and the oldest fossil primate from Europe. As such, this taxon has often been hypothesized as a basal tarsiiform on the basis of its primitive dental formula with four premolars and a simplified molar cusp pattern. Until recently [see Rose et al.: Am J Phys Anthropol 146 (2011) 281–305; Gebo et al.: J Hum Evol 63 (2012) 205–218], little was known concerning its postcranial anatomy with the exception of its well‐known tarsals. In this article, we describe additional postcranial elements for T. belgica and compare these with other tarsiiforms and with primitive adapiforms. The forelimb of T. belgica indicates an arboreal primate with prominent forearm musculature, good elbow rotational mobility, and a horizontal, rather than a vertical body posture. The lateral hand positions imply grasps adaptive for relatively large diameter supports given its small body size. The hand is long with very long fingers, especially the middle phalanges. The hindlimb indicates foot inversion capabilities, frequent leaping, arboreal quadrupedalism, climbing, and grasping. The long and well‐muscled hallux can be coupled with long lateral phalanges to reconstruct a foot with long grasping digits. Our phyletic analysis indicates that we can identify several postcranial characteristics shared in common for stem primates as well as note several derived postcranial characters for Tarsiiformes. Am J Phys Anthropol 156:388–406, 2015. © 2014 Wiley Periodicals, Inc.     

The sculpture and morphology of postcranial dermal armor plates and associated bones in gasterosteiforms and syngnathiforms inhabiting Estonian coastal waters

Lees, J., Märss, T., Wilson, M. V. H., Saat, T. and ?pilev, H. 2011. The sculpture and morphology of postcranial dermal armor plates and associated bones in gasterosteiforms and syngnathiforms inhabiting Estonian coastal waters. ―Acta Zoologica (Stockholm) 93 : 422–435. Five fish species inhabiting Estonian coastal waters (Gasterosteus aculeatus L., Pungitius pungitius (L.), and Spinachia spinachia (L.) of the order Gasterosteiformes and Syngnathus typhle L. and Nerophis ophidion (L.) of the order Syngnathiformes) are described on the basis of the sculpture and morphology of their postcranial dermal armor plates, as revealed and illustrated by SEM images. This study shows that the shapes of these superficial skeletal elements vary by species as well as by their position on the body, whereas the sculpture on the bones is taxon specific. The detailed features allow the identification of isolated fossil and subfossil remains and show promise for future systematics studies.     

A hominoid distal tibia from the Miocene of Pakistan

A distal tibia, YGSP 1656, from the early Late Miocene portion of the Chinji Formation in Pakistan is described. The fossil is 11.4 million years old and is one of only six postcranial elements now assigned to Sivapithecus indicus. Aspects of the articular surface are cercopithecoid-like, suggesting some pronograde locomotor activities. However, YGSP 1656 possesses an anteroposteriorly compressed metaphysis and a mediolaterally thick medial malleolus, ape-like features functionally related to orthograde body postures and vertical climbing. YGSP 1656 lacks specializations found in the ankle of terrestrial cercopithecoids and thus Sivapithecus may have been primarily arboreal. Nevertheless, the morphology of this tibia is unique, consistent with other interpretations of Sivapithecus postcranial functional morphology that suggest the locomotion of this ape lacks a modern analog. Based on the limited postcranial remains from S. indicus, we hypothesize that this taxon exhibited substantial body size dimorphism.