Extant Vertebrates Research Articles

The oldest ionoscopiform from China sheds new light on the early evolution of halecomorph fishes

The Halecomorphi are a major subdivision of the ray-finned fishes. Although living halecomorphs are represented solely by the freshwater bowfin, Amia calva, this clade has a rich fossil history, and the resolution of interrelationships among extinct members is central to the problem of understanding the origin of the Teleostei, the largest clade of extant vertebrates. The Ionoscopiformes are extinct marine halecomorphs that were inferred to have originated in the Late Jurassic of Europe, and subsequently dispersed to the Early Cretaceous of the New World. Here, we report the discovery of a new ionoscopiform, Robustichthys luopingensis gen. et sp. nov., based on eight well-preserved specimens from the Anisian (242-247 Ma), Middle Triassic marine deposits of Luoping, eastern Yunnan Province, China. The new species documents the oldest known ionoscopiform, extending the stratigraphic range of this group by approximately 90 Ma, and the geographical distribution of this group into the Middle Triassic of South China, a part of eastern Palaeotethys Ocean. These new data provide a minimum estimate for the split of Ionoscopiformes from its sister clade Amiiformes and shed new light on the origin of ionoscopiform fishes.

The anatomy of the bifurcated neural spine and its occurrence within Tetrapoda.

Vertebral neural spine bifurcation has been historically treated as largely restrictive to sauropodomorph dinosaurs; wherein it is inferred to be an adaptation in response to the increasing weight from the horizontally extended cervical column. Because no extant terrestrial vertebrates have massive, horizontally extended necks, extant forms with large cranial masses were examined for the presence of neural spine bifurcation. Here, I report for the first time on the soft tissue surrounding neural spine bifurcation in a terrestrial quadruped through the dissection of three Ankole-Watusi cattle. With horns weighing up to a combined 90 kg, the Ankole-Watusi is unlike any other breed of cattle in terms of cranial weight and presence of neural spine bifurcation. Using the Ankole-Watusi as a model, it appears that neural spine bifurcation plays a critical role in supporting a large mobile weight adjacent to the girdles. In addition to neural spine bifurcation being recognized within nonavian dinosaurs, this vertebral feature is also documented within many members of temnospondyls, captorhinids, seymouriamorphs, diadectomorphs, Aves, marsupials, artiodactyls, perissodactyls, and Primates, amongst others. This phylogenetic distribution indicates that spine bifurcation is more common than previously thought, and that this vertebral adaptation has contributed throughout the evolutionary history of tetrapods. Neural spine bifurcation should now be recognized as an anatomical component adapted by some vertebrates to deal with massive, horizontal, mobile weights adjacent the girdles.

A unique mode of tissue oxygenation and the adaptive radiation of teleost fishes

Teleost fishes constitute 95% of extant aquatic vertebrates, and we suggest that this is related in part to their unique mode of tissue oxygenation. We propose the following sequence of events in the evolution of their oxygen delivery system. First, loss of plasma-accessible carbonic anhydrase (CA) in the gill and venous circulations slowed the Jacobs-Stewart cycle and the transfer of acid between the plasma and the red blood cells (RBCs). This ameliorated the effects of a generalised acidosis (associated with an increased capacity for burst swimming) on haemoglobin (Hb)-O2 binding. Because RBC pH was uncoupled from plasma pH, the importance of Hb as a buffer was reduced. The decrease in buffering was mediated by a reduction in the number of histidine residues on the Hb molecule and resulted in enhanced coupling of O2 and CO2 transfer through the RBCs. In the absence of plasma CA, nearly all plasma bicarbonate ultimately dehydrated to CO2 occurred via the RBCs, and chloride/bicarbonate exchange was the rate-limiting step in CO2 excretion. This pattern of CO2 excretion across the gills resulted in disequilibrium states for CO2 hydration/dehydration reactions and thus elevated arterial and venous plasma bicarbonate levels. Plasma-accessible CA embedded in arterial endothelia was retained, which eliminated the localized bicarbonate disequilibrium forming CO2 that then moved into the RBCs. Consequently, RBC pH decreased which, in conjunction with pH-sensitive Bohr/Root Hbs, elevated arterial oxygen tensions and thus enhanced tissue oxygenation. Counter-current arrangement of capillaries (retia) at the eye and later the swim bladder evolved along with the gas gland at the swim bladder. Both arrangements enhanced and magnified CO2 and acid production and, therefore, oxygen secretion to those specialised tissues. The evolution of β-adrenergically stimulated RBC Na(+)/H(+) exchange protected gill O2 uptake during stress and further augmented plasma disequilibrium states for CO2 hydration/dehydration. Finally, RBC organophosphates (e.g. NTP) could be reduced during hypoxia to further increase Hb-O2 affinity without compromising tissue O2 delivery because high-affinity Hbs could still adequately deliver O2 to the tissues via Bohr/Root shifts. We suggest that the evolution of this unique mode of tissue O2 transfer evolved in the Triassic/Jurassic Period, when O2 levels were low, ultimately giving rise to the most extensive adaptive radiation of extant vertebrates, the teleost fishes.

Do tracks yield reliable information on gaits? – Part 1: The case of horses

Abstract. During their lifetime animals leave many tracks and traces behind, which can provide insights into the animals' behaviour. Single footprints of extant vertebrates are frequently found in sediments all over the world, often arranged into trackways. The study of footprints and trackways lead to interpretations about the mode of locomotion of the trackmaker. Here we show an approach to identify gaits from tracks. A series of experiments with horses was performed to determine whether gaits could be identified on the basis of fossil trackways, e.g. those left behind by sauropod dinosaurs of the Mesozoic era or Tertiary mammals, to unveil their locomotor abilities. The generally valid rules for quadrupedal locomotion were taken into consideration. Symmetrical gaits result in very similar trackways; a further differentiation can be made by application of statistics on step lengths, excursion angles and overstepping. A clear difference exists between the trot and the pace. These rapid, symmetric gaits imply high ground reaction forces (GRF) because of their long phases of aerial suspension at higher speeds. The resulting GRF seem to be too high to be sustained by the limb bones of huge graviportal animals like sauropods. Unfortunately, most of these factors are rarely available in the case of fossil tracks. Likewise, the asymmetrical, springing gaits can be excluded for sauropods because of the enormous GRF. Provided that limb length as well as trunk length can be approximated, and left and right, as well as forefoot and hindfoot imprints can be discriminated, the symmetrical gaits (walk, amble, pace, trot) used when making a trackway can be discerned.

Evolutionarily conserved morphogenetic movements at the vertebrate head–trunk interface coordinate the transport and assembly of hypopharyngeal structures

The vertebrate head–trunk interface (occipital region) has been heavily remodelled during evolution, and its development is still poorly understood. In extant jawed vertebrates, this region provides muscle precursors for the throat and tongue (hypopharyngeal/hypobranchial/hypoglossal muscle precursors, HMP) that take a stereotype path rostrally along the pharynx and are thought to reach their target sites via active migration. Yet, this projection pattern emerged in jawless vertebrates before the evolution of migratory muscle precursors. This suggests that a so far elusive, more basic transport mechanism must have existed and may still be traceable today.Here we show for the first time that all occipital tissues participate in well-conserved cell movements. These cell movements are spearheaded by the occipital lateral mesoderm and ectoderm that split into two streams. The rostrally directed stream projects along the floor of the pharynx and reaches as far rostrally as the floor of the mandibular arch and outflow tract of the heart. Notably, this stream leads and engulfs the later emerging HMP, neural crest cells and hypoglossal nerve. When we (i) attempted to redirect hypobranchial/hypoglossal muscle precursors towards various attractants, (ii) placed non-migratory muscle precursors into the occipital environment or (iii) molecularly or (iv) genetically rendered muscle precursors non-migratory, they still followed the trajectory set by the occipital lateral mesoderm and ectoderm. Thus, we have discovered evolutionarily conserved morphogenetic movements, driven by the occipital lateral mesoderm and ectoderm, that ensure cell transport and organ assembly at the head–trunk interface.

MOLECULAR EVOLUTION OF GPCRS: Kisspeptin/kisspeptin receptors

Following the discovery of kisspeptin (Kiss) and its receptor (GPR54 or KissR) in mammals, phylogenetic studies revealed up to three Kiss and four KissR paralogous genes in other vertebrates. The multiplicity of Kiss and KissR types in vertebrates probably originated from the two rounds of whole-genome duplication (1R and 2R) that occurred in early vertebrates. This review examines compelling recent advances on molecular diversity and phylogenetic evolution of vertebrate Kiss and KissR. It also addresses, from an evolutionary point of view, the issues of the structure-activity relationships and interaction of Kiss with KissR and of their signaling pathways. Independent gene losses, during vertebrate evolution, have shaped the repertoire of Kiss and KissR in the extant vertebrate species. In particular, there is no conserved combination of a given Kiss type with a KissR type, across vertebrate evolution. The striking conservation of the biologically active ten-amino-acid C-terminal sequence of all vertebrate kisspeptins, probably allowed this evolutionary flexibility of Kiss/KissR pairs. KissR mutations, responsible for hypogonadotropic hypogonadism in humans, mostly occurred at highly conserved amino acid positions among vertebrate KissR. This further highlights the key role of these amino acids in KissR function. In contrast, less conserved KissR regions, notably in the intracellular C-terminal domain, may account for differential intracellular signaling pathways between vertebrate KissR. Cross talk between evolutionary and biomedical studies should contribute to further understanding of the Kiss/KissR structure-activity relationships and biological functions.

Allometries of Maximum Growth Rate versus Body Mass at Maximum Growth Indicate That Non-Avian Dinosaurs Had Growth Rates Typical of Fast Growing Ectothermic Sauropsids

We tested if growth rates of recent taxa are unequivocally separated between endotherms and ectotherms, and compared these to dinosaurian growth rates. We therefore performed linear regression analyses on the log-transformed maximum growth rate against log-transformed body mass at maximum growth for extant altricial birds, precocial birds, eutherians, marsupials, reptiles, fishes and dinosaurs. Regression models of precocial birds (and fishes) strongly differed from Case’s study (1978), which is often used to compare dinosaurian growth rates to those of extant vertebrates. For all taxonomic groups, the slope of 0.75 expected from the Metabolic Theory of Ecology was statistically supported. To compare growth rates between taxonomic groups we therefore used regressions with this fixed slope and group-specific intercepts. On average, maximum growth rates of ectotherms were about 10 (reptiles) to 20 (fishes) times (in comparison to mammals) or even 45 (reptiles) to 100 (fishes) times (in comparison to birds) lower than in endotherms. While on average all taxa were clearly separated from each other, individual growth rates overlapped between several taxa and even between endotherms and ectotherms. Dinosaurs had growth rates intermediate between similar sized/scaled-up reptiles and mammals, but a much lower rate than scaled-up birds. All dinosaurian growth rates were within the range of extant reptiles and mammals, and were lower than those of birds. Under the assumption that growth rate and metabolic rate are indeed linked, our results suggest two alternative interpretations. Compared to other sauropsids, the growth rates of studied dinosaurs clearly indicate that they had an ectothermic rather than an endothermic metabolic rate. Compared to other vertebrate growth rates, the overall high variability in growth rates of extant groups and the high overlap between individual growth rates of endothermic and ectothermic extant species make it impossible to rule out either of the two thermoregulation strategies for studied dinosaurs.

A primitive placoderm sheds light on the origin of the jawed vertebrate face

Extant vertebrates form two clades, the jawless Cyclostomata (lampreys and hagfishes) and the jawed Gnathostomata (all other vertebrates), with contrasting facial architectures. These arise during development from just a few key differences in the growth patterns of the cranial primordia: notably, the nasal sacs and hypophysis originate from a single placode in cyclostomes but from separate placodes in gnathostomes, and infraoptic ectomesenchyme migrates forward either side of the single placode in cyclostomes but between the placodes in gnathostomes. Fossil stem gnathostomes preserve cranial anatomies rich in landmarks that provide proxies for developmental processes and allow the transition from jawless to jawed vertebrates to be broken down into evolutionary steps. Here we use propagation phase contrast synchrotron microtomography to image the cranial anatomy of the primitive placoderm (jawed stem gnathostome) Romundina, and show that it combines jawed vertebrate architecture with cranial and cerebral proportions resembling those of cyclostomes and the galeaspid (jawless stem gnathostome) Shuyu. This combination seems to be primitive for jawed vertebrates, and suggests a decoupling between ectomesenchymal growth trajectory, ectomesenchymal proliferation, and cerebral shape change during the origin of gnathostomes.

Comparative analysis of pleurodiran and cryptodiran turtle embryos depicts the molecular ground pattern of the turtle carapacial ridge.

The turtle shell is a wonderful example of a genuine morphological novelty, since it has no counterpart in any other extant vertebrate lineages. The evolutionary origin of the shell is a question that has fascinated evolutionary biologists for over two centuries and it still remains a mystery. One of the turtle innovations associated with the shell is the carapacial ridge (CR), a bulge that appears at both sides of the dorsal lateral trunk of the turtle embryo and that probably controls the formation of the carapace, the dorsal moiety of the shell. Although from the beginning of this century modern genetic techniques have been applied to resolve the evolutionary developmental origin of the CR, the use of different models with, in principle, dissimilar results has hampered the establishment of a common mechanism for the origin of the shell. Although modern turtles are divided into two major groups, Cryptodira (or hidden-necked turtles) and Pleurodira (or side-necked turtles), molecular developmental studies have been carried out mostly using cryptodiran models. In this study, we revisit the past data obtained from cryptodiran turtles in order to reconcile the different results. We also analyze the histological anatomy and the expression pattern of main CR factors in a pleurodiran turtle, the red-bellied short-necked turtle Emydura subglobosa. We suggest that the turtle shell probably originated concomitantly with the co-option of the canonical Wnt signaling pathway into the CR in the last common ancestor of the turtle.

Lampreys Have a Single Gene Cluster for the Fast Skeletal Myosin Heavy Chain Gene Family

Muscle tissues contain the most classic sarcomeric myosin, called myosin II, which consists of 2 heavy chains (MYHs) and 4 light chains. In the case of humans (tetrapod), a total of 6 fast skeletal-type MYH genes (MYHs) are clustered on a single chromosome. In contrast, torafugu (teleost) contains at least 13 fast skeletal MYHs, which are distributed in 5 genomic regions; the MYHs are clustered in 3 of these regions. In the present study, the evolutionary relationship among fast skeletal MYHs is elucidated by comparing the MYHs of teleosts and tetrapods with those of cyclostome lampreys, one of two groups of extant jawless vertebrates (agnathans). We found that lampreys contain at least 3 fast skeletal MYHs, which are clustered in a head-to-tail manner in a single genomic region. Although there was apparent synteny in the corresponding MYH cluster regions between lampreys and tetrapods, phylogenetic analysis indicated that lamprey and tetrapod MYHs have independently duplicated and diversified. Subsequent transgenic approaches showed that the 5′-flanking sequences of Japanese lamprey fast skeletal MYHs function as a regulatory sequence to drive specific reporter gene expression in the fast skeletal muscle of zebrafish embryos. Although zebrafish MYH promoters showed apparent activity to direct reporter gene expression in myogenic cells derived from mice, promoters from Japanese lamprey MYHs had no activity. These results suggest that the muscle-specific regulatory mechanisms are partially conserved between teleosts and tetrapods but not between cyclostomes and tetrapods, despite the conserved synteny.

The origin of developmental mechanisms underlying vertebral elements: implications from hagfish evo-devo

The origins of the vertebral elements and the underlying developmental mechanisms have so far remained unclear, largely due to the unusual axial skeletal morphology of hagfish, one of two extant jawless vertebrate clades. Hagfish axial supporting tissue is generally believed to consist of the notochord and cartilaginous fin rays only. However, careful investigations of whether vertebral elements are truly absent in hagfish are scarce, and it is also unclear whether the axial skeletal morphology of the hagfish is an ancestral or a derived condition. To address these questions, we re-examined the axial skeletal morphology of the Japanese inshore hagfish (Eptatretus burgeri). Based on a report published a century ago which implied the existence of vertebral elements in hagfish, we conducted anatomical and histological analyses of the hagfish axial skeletal systems and their development. Through this analysis, we demonstrate that hagfish possesses sclerotome-derived cartilaginous vertebral elements at the ventral aspect of the notochord. Based on (i) molecular phylogenetic evidence in support of the monophyly of cyclostomes (hagfish and lampreys) and jawed vertebrates (gnathostomes), and (ii) the morphology of the vertebral elements in extant gnathostomes and cyclostomes, we propose that the embryos of the common ancestor of all vertebrates would have possessed sclerotomal cells that formed the segmentally arranged vertebral elements attached to the notochord. We also conclude that the underlying developmental mechanisms are likely to have been conserved among extinct jawless vertebrates and modern gnathostomes.

Dynamic bipedal walking of a dinosaur-like robot with an extant vertebrate's nervous system

SUMMARYIn this study, we attempt to develop a biped dinosaur-like walking robot by focusing on its nervous system as well as its mechanism. We developed a robot ‘Dinobot’ on the basis of palaeontological knowledge on dinosaurs and extant animals. In addition, we employed typical biologically inspired walking gait generation and control methods derived from an extant vertebrate's nervous system. In particular, we utilized a central pattern generator (CPG), which is a locomotion rhythm generator in a vertebrate's spinal cord, to generate the robot's walking rhythm. Moreover, a reflex centre was placed below CPG and it produced joint torque of the legs in the swing and stance phases. Thus, we successfully achieved adaptive 3D dynamic walking generated by the interaction between the original mechanism of dinosaurs and the nervous system of extant animals. Our future goal is to find out a dinosaur's robust locomotive nervous system suitable for its mechanism.

Diversity and history of the long-chain acyl-CoA synthetase (Acsl) gene family in vertebrates

BackgroundFatty acids, a considerable fraction of lipid molecules, participate in fundamental physiological processes. They undergo activation into their corresponding CoA esters for oxidation or esterification into complex lipids (e.g. triglycerides, phospholipids and cholesterol esters), a process that is carried out by acyl-CoA synthases (ACS). Here we analyze the evolution of the gene family encoding for the long-chain acyl-CoA synthetases (Acsl) in vertebrates.ResultsBy means of phylogenetics and comparative genomics we show that genome duplications (2R) generated the diversity of Acsl genes in extant vertebrate lineages. In the vertebrate ancestor two separate genes originated the current Acsl1/5/6 and the Acsl3/4 gene families, and the extra gene duplicates in teleosts are a consequence of the teleost specific third round of genome duplication (3R). Moreover, the diversity of Acsl family members is broader than anticipated. Our strategy uncovered a novel uncharacterized Acsl-like gene found in teleosts, spotted gar, coelacanth and possibly lamprey, which we designate Acsl2. The detailed analysis of the Acsl2 teleost gene locus strongly supports the conclusion that it corresponds to a retained 2R paralogue, lost in tetrapods.ConclusionsWe provide here the first evolutionary analysis of the Acsl gene family in vertebrates, showing the specific contribution of 2R/3R to the diversity of this gene family. We find also that the division of ACSL enzymes into two groups predates at least the emergence of deuterostomes. Our study indicates that genome duplications significantly contributed to the elaboration of fatty acid activation metabolism in vertebrates.

Regulation of a putative corticosteroid, 17,21-dihydroxypregn-4-ene,3,20-one, in sea lamprey, Petromyzon marinus

In higher vertebrates, in response to stress, the hypothalamus produces corticotropin-releasing hormone (CRH), which stimulates cells in the anterior pituitary to produce adrenocorticotropic hormone (ACTH), which in turn stimulates production of either cortisol (F) or corticosterone (B) by the adrenal tissues. In lampreys, however, neither of these steroids is present. Instead, it has been proposed that the stress steroid is actually 17,21-dihydroxypregn-4-ene-3,20-dione (11-deoxycortisol; S). However, there have been no studies yet to determine its mechanism of regulation or site of production. Here we demonstrate that (1) intraperitoneal injections of lamprey-CRH increase plasma S in a dose dependent manner, (2) intraperitoneal injections of four lamprey-specific ACTH peptides at 100μg/kg, did not induce changes in plasma S concentrations in either males or females; (3) two lamprey-specific gonadotropin-releasing hormones (GnRH I and III) and arginine-vasotocin (AVT), all at single doses, stimulated S production as well as, or to an even greater extent than CRH; (4) sea lamprey mesonephric kidneys, in vitro, converted tritiated 17α-hydroxyprogesterone (17α-P) into a steroid that had the same chromatographic properties (on HPLC and TLC) as S; (5) kidney tissues released significantly more immunoassayable S into the incubation medium than gill, liver or gonad tissues. One interpretation of these results is that the corticosteroid production of the sea lamprey, one of the oldest extant vertebrates, is regulated through multiple pathways rather than the classical HPI-axis. However, the responsiveness of this steroid to the GnRH peptides means that a reproductive rather than a stress role for this steroid cannot yet be ruled out.

The vertebrate taxonomy ontology: a framework for reasoning across model organism and species phenotypes.

BackgroundA hierarchical taxonomy of organisms is a prerequisite for semantic integration of biodiversity data. Ideally, there would be a single, expansive, authoritative taxonomy that includes extinct and extant taxa, information on synonyms and common names, and monophyletic supraspecific taxa that reflect our current understanding of phylogenetic relationships.DescriptionAs a step towards development of such a resource, and to enable large-scale integration of phenotypic data across vertebrates, we created the Vertebrate Taxonomy Ontology (VTO), a semantically defined taxonomic resource derived from the integration of existing taxonomic compilations, and freely distributed under a Creative Commons Zero (CC0) public domain waiver. The VTO includes both extant and extinct vertebrates and currently contains 106,947 taxonomic terms, 22 taxonomic ranks, 104,736 synonyms, and 162,400 cross-references to other taxonomic resources. Key challenges in constructing the VTO included (1) extracting and merging names, synonyms, and identifiers from heterogeneous sources; (2) structuring hierarchies of terms based on evolutionary relationships and the principle of monophyly; and (3) automating this process as much as possible to accommodate updates in source taxonomies.ConclusionsThe VTO is the primary source of taxonomic information used by the Phenoscape Knowledgebase (http://phenoscape.org/), which integrates genetic and evolutionary phenotype data across both model and non-model vertebrates. The VTO is useful for inferring phenotypic changes on the vertebrate tree of life, which enables queries for candidate genes for various episodes in vertebrate evolution.

Origin and evolution of adaptive immunity.

The evolutionary emergence of vertebrates was accompanied by major morphological and functional innovations, including the development of an adaptive immune system. Vertebrate adaptive immunity is based on the clonal expression of somatically diversifying antigen receptors on lymphocytes. This is a common feature of both the jawless and jawed vertebrates , although these two groups of extant vertebrates employ structurally different types of antigen receptors and principal mechanisms for their somatic diversification . These observations suggest that the common vertebrate ancestor must have already possessed a complex immune system, including B- and T-like lymphocyte lineages and primary lymphoid organs, such as the thymus, but possibly lacked the facilities for somatic diversification of antigen receptors. Interestingly, memory formation, previously considered to be a defining feature of adaptive immunity, also occurs in the context of innate immune responses and can even be observed in unicellular organisms, attesting to the convergent evolutionary history of distinct aspects of adaptive immunity.

Unique characteristics of the trachea of the juvenile leatherback turtle facilitate feeding, diving and endothermy

The adult leatherback turtle Dermochelys coriacea overlaps in body size (300–500kg) with many marine mammals, yet develops from a 50g hatchling. Adults can dive deeper than 1200m and have core body temperatures of 25°C; hatchlings are near-surface dwellers. Juvenile leatherbacks have rarely been studied; here we present anatomical information for the upper respiratory tract of 3 turtles (66.7–83.0cm straight carapace length; 33.2–53.4kg body mass) incidentally captured by long-line fisheries. Combined with existing information from adults and hatchlings, our data show that there is an ontogenic shift in tracheal structure, with cartilaginous rings becoming broader and eventually fusing anteriorly. This ontogenic shift during independent existence is unique among extant deep-diving air breathing vertebrates. Tract wall thickness is graded, becoming progressively thinner from larynx to bronchi. In addition, cross-sectional shape becomes increasingly dorsoventrally flattened (more elliptical) from anterior to posterior. These characteristics ensure that the tract will collapse from posterior to anterior during dives. This study contains the first report of a double (=internally bifurcated) posterior section of the trachea; it is suggested that this allows continuous food movement along the esophagus without tracheal collapse. The whole upper respiratory tract (from larynx to lungs) has a vascular lining (thicker anteriorly than posteriorly) that appears to be a simple analog of the complex turbinates of birds and mammals. Our study confirmed that the leatherback tracheal structure represents a distinctive way of dealing with the challenges of diving in deep, cold sea water.

Cerebellum and spatial cognition in goldfish

The cerebellum of mammals has recently been linked to spatial navigation, as indicated by the results of a number of studies performed in animal models with cerebellar abnormalities. However, nothing is known about the contribution of this structure to spatial cognition in other vertebrate groups such as teleost fish. To investigate the involvement of the teleostean cerebellum in navigation, sham-operated (Sh) and cerebellum-ablated (Cb) goldfish were trained in a “hole-board” task in which they had to locate the baited feeder within a 5×5 feeder matrix surrounded by visual cues. Cb goldfish were significantly impaired in the acquisition and performance of the task, as revealed by their low spatial accuracy, the number of errors committed, and the stereotyped searching pattern exhibited relative to Sh goldfish. Probe tests, performed during the final training sessions, showed that Cb animals could not integrate experimental cues into an internal representation of the environment (as an allocentric strategy would require) and they resorted to a guiding strategy to locate the goal. The results of this experiment demonstrated that the cerebellum might have a modulatory role in the declarative component of navigation by which an animal develops an internal spatial representation. Our results constitute the first evidence of the involvement of the fish cerebellum in spatial cognition. Our results also suggest that the cognitive functions of the cerebellum may have appeared early in vertebrate evolution and been conserved throughout the phylogenetic history of extant vertebrates.

The articulation of sauropod necks: methodology and mythology.

Sauropods are often imagined to have held their heads high atop necks that ascended in a sweeping curve that was formed either intrinsically because of the shape of their vertebrae, or behaviorally by lifting the head, or both. Their necks are also popularly depicted in life with poses suggesting avian flexibility. The grounds for such interpretations are examined in terms of vertebral osteology, inferences about missing soft tissues, intervertebral flexibility, and behavior. Osteologically, the pronounced opisthocoely and conformal central and zygapophyseal articular surfaces strongly constrain the reconstruction of the cervical vertebral column. The sauropod cervico-dorsal vertebral column is essentially straight, in contrast to the curvature exhibited in those extant vertebrates that naturally hold their heads above rising necks. Regarding flexibility, extant vertebrates with homologous articular geometries preserve a degree of zygapophyseal overlap at the limits of deflection, a constraint that is further restricted by soft tissues. Sauropod necks, if similarly constrained, were capable of sweeping out large feeding surfaces, yet much less capable of retracting the head to explore the enclosed volume in an avian manner. Behaviorally, modern vertebrates generally assume characteristic neck postures which are close to the intrinsic curvature of the undeflected neck. With the exception of some vertebrates that can retract their heads to balance above their shoulders at rest (e.g., felids, lagomorphs, and some ratites), the undeflected neck generally predicts the default head height at rest and during locomotion.

Bite marks on an ichthyodectiform fish from Australia: possible evidence of trophic interaction in an Early Cretaceous marine ecosystem

Wretman, L. & Kear, B.P., 2013. Bite marks on an ichthyodectiform fish from Australia: possible evidence of trophic interaction in an Early Cretaceous marine ecosystem. Alcheringa 38, 000–000. ISSN 0311-5518.A well-preserved fish skull from late Albian deposits of the Allaru Mudstone near Richmond in Queensland displays a conspicuous V-shaped pattern of indentations, punctures and depression fractures consistent with a vertebrate bite trace. This is the first direct evidence of trophic interaction between vertebrates within an Early Cretaceous marine ecosystem from Australia. The specimen is taxonomically referable to the large-bodied (ca 1 m snout–tail length) ichthyodectiform Cooyoo australis, but the size and spacing of the tooth marks is incompatible with attack by a conspecific individual. The lack of osseous growths concordant with healing also suggests that the bite occurred shortly before or after the animal’s death. Comparison with the dentitions of other coeval vertebrates indicates compatible tooth arrangements in longirostrine amniote predators such as polycotylid plesiosaurians, ornithocheiroid pterosaurs and especially the ichthyosaurian Platypterygius. The implications of this as a potential predator–prey association are that Early Cretaceous actinopterygians occupied middle-level trophic niches and were in turn consumed by higher-level amniote carnivores, similar to many extant marine vertebrate communities of today.Lovisa Wretman [lovisa.wretman@ebc.uu.se], Subdepartment of Evolution and Development, Department of Organismal Biology, Uppsala University, Norbyvägen 18A, SE-752 36 Uppsala, Sweden (address for correspondence); Benjamin P. Kear [benjamin.kear@geo.uu.se], Palaeobiology Programme, Department of Earth Sciences, Uppsala University, Villavägen 16, SE-752 36 Uppsala, Sweden. Received 8.8.2013, revised 17.9.2013, accepted 23.9.2013.