Metamorphosis and evolution of feeding behaviour in salamanders of the family Plethodontidae

Plethodontid salamanders capture prey with enhanced tongue protraction relative to other salamander taxa, yet metamorphosing plethodontids are hypothesized to be constrained relative to direct-developing plethodontids in their degree of tongue evolution (protraction length and velocity) by the presence of a larval stage in development. In this biphasic life history the hyobranchial apparatus serves the conflicting functions of larval suction feeding and adult tongue protraction. The deletion of the larval stage removes one of the conflicting functions and has thus permitted direct-developing plethodontids to circumvent this constraint and evolve extremely long tongues, which in some species can be projected to 80% of body length. To evaluate this constraint hypothesis and explore taxonomic diversity of feeding behaviours, we studied feeding in larvae, adults and metamorphosing individuals of seven species of metamorphosing plethodontids from the basal taxa Desmognathinae and Hemidactyliini using direct observations, high-speed videography and kinematic analysis. We found that larval plethodontids suction feed, but feeding is suspended entirely during metamorphosis, and aquatic adults do not suction feed. Adults have exapted the terrestrial modes of tongue and jaw prehension for aquatic prey capture. These findings substantiate the premise that suction feeding and tongue protraction are conflicting functions, and thus our results support the constraint hypothesis. Plethodontid adults have evolved their extreme tongue protraction ability at the expense of adult suction feeding. The rapid metamorphosis that characterizes plethodontids may be an adaptation that minimizes the non-feeding period imposed by the evolution of derived tongue protraction in adults. © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society , 2002, 134 , 375–400.     

Cranial shape in fruit,nectar, and exudate feeders: Implications for interpreting the fossil record

At least 29 species of fossil primates have been referred to fruit, nectar, and/or exudate feeding dietary niches. Many studies have detailed the morphological correlates of fruit feeding in comparison to insectivory and folivory. In contrast, few studies have sought to differentiate the morphological correlates of fruit feeding from those of nectar and exudate feeding. This study investigates the differences between fruit, nectar, and exudate feeders using 22 cranial and dentary shape variables representing 28 species of living marsupials, bats, and primates. Discriminant function analysis is used to investigate the differences between these dietary categories using both the complete data set and a reduced data set composed of variables that might reasonably be available from fragmentary fossil material. The success rates of post-hoc classifications are 94 and 88%, respectively. These results demonstrate that it is possible to discriminate among fruit, nectar, and exudate feeders among fossil taxa with a reasonable degree of certainty using the data and techniques outlined here. Nectar feeders exhibit a unique combination of features that are associated with reduced masticatory strength and their role as pollination agents. Exudate feeder skulls and dentaries exhibit a combination of features that reflect the high stresses encountered by the anterior dentition through bark gouging behavior. Fruit feeders are morphologically diverse, exhibiting cranial and mandibular shape values that overlap with both nectar and exudate feeders. It is suggested that this diversity reflects the variety of physical properties represented among fruits, and the tendency for individual frugivore species to specialize on particular fruits. Am J Phys Anthropol 102:187–202, 1997 © 1997 Wiley-Liss, Inc.     

Angular roughshark Oxynotus centrina (Squaliformes: Oxynotidae) in captivity feeding exclusively on elasmobranch eggs: an overlooked feeding niche or a matter of individual taste?

A specimen of angular roughshark Oxynotus centrina has been kept successfully in captivity for the first time. Over a period of 24 months, the specimen preyed exclusively on the contents of elasmobranch egg cases, suggesting a specialized trophic niche.     

The mixosaurid ichthyosaur Contectopalatus from the Middle Triassic of the German Basin

The skull of the mixosaurid species Contectopalatus atavus (Quenstedt, 1851-52) is the most bizarre of any known ichthyosaur. It possesses a very high sagittal crest formed by the nasal, frontal and parietal bones which grows higher during ontogeny. This skull structure - found to a lesser extent in the other mixosaurid genera Mixosaurus and Phalarodon - is a synapomorphy of the family Mixosauridae. It is here interpreted as correlated with a unique arrangement of the jaw adductor musculature among tetrapods, with the internal jaw adductors extending over most of the skull roof up to the external narial opening. This reconstruction would increase the biting force considerably and the hypothesis is supported by peculiarities of the dentition and jaws of Contectopalatus. Contectopalatus probably reached a length of about 5 meters. It is therefore the largest known mixosaurid and one of the largest Triassic ichthyosaurs. The general text-book picture of mixosaurs as small, rather unspecialized, primitive ichthyosaurs is incorrect. Mixosaurs were a highly specialized, uniquely adapted and very diverse ichthyosaur family, some members of which rank among the marine top predators of their time.     

Modulation in Feeding Kinematics and Motor Pattern of the Nurse Shark <Emphasis Type="Italic">Ginglymostoma cirratum</Emphasis>

Synopsis Studies of feeding in bony fishes have almost universally demonstrated the ability of individuals to modulate their method of capture in response to differing stimuli. Preliminary evidence indicates that morphologically specialized inertial suction feeding sharks are the most likely fishes to lack inherent modulatory ability. We examined the ability of the nurse shark, Ginglymostoma cirratum, to modulate its feeding behavior based on different food types and sizes. G. cirratum is an inertial suction feeding fish that is apparently stereotyped in its food capture behavior. Electromyography showed no statistical difference between feeding motor patterns based on food type (squid or fish) or size (gape width or twice gape width), although there were slight inter-individual differences in the onset of muscle firing for some muscles. Kinematic analysis showed a statistical difference in variables associated with durations for different food types, with the durations for all variables being faster for squid bites than fish bites, but no difference based on the size of the food item. This apparent lack of modulation may be associated with specialization of the morphology and behavior of G. cirratum for obligate suction prey capture. This functional specialization constrains the method in which G. cirratum captures prey but does not appear to result in dietary specialization. An unusual post capture spit-suck manipulation allows this shark to handle and ingest large prey.     

Prey capture behavior and kinematics of the Atlantic cownose ray, Rhinoptera bonasus

The structurally reinforced jaws of the cownose ray, Rhinoptera bonasus testify to this species' durophagous diet of mollusks, but seem ill-suited to the behaviors necessary for excavating such prey. This study explores this discordance by investigating the prey excavation and capture kinematics of R. bonasus. Based on the basal suction feeding mechanism in this group of fishes, we hypothesized a hydraulic method of excavation. As expected, prey capture kinematics of R. bonasus show marked differences relative to other elasmobranchs, relating to prey excavation and use of the cephalic lobes (modified anterior pectoral fin extensions unique to derived myliobatiform rays). Prey are excavated by repeated opening and closing of the jaws to fluidize surrounding sand. The food item is then enclosed laterally by the depressed cephalic lobes, which transport it toward the mouth for ingestion by inertial suction. Unlike in most sharks, upper jaw protrusion and mandibular depression are simultaneous. During food capture, the ray's spiracle, mouth, and gill slit movements are timed such that water enters only the mouth (e.g., the spiracle closes prior to prey capture and reopens immediately following). Indigestible parts are then hydraulically winnowed from edible prey portions, by mouth movements similar to those used in excavation, and ejected through the mouth. The unique sensory/manipulatory capabilities of the cephalic lobes, as well as the cownose ray's hydraulic excavation/winnowing behaviors and suction feeding, make this species an effective benthic predator, despite its epibenthic lifestyle.     

Models in palaeontological functional analysis

Models are a principal tool of modern science. By definition, and in practice, models are not literal representations of reality but provide simplifications or substitutes of the events, scenarios or behaviours that are being studied or predicted. All models make assumptions, and palaeontological models in particular require additional assumptions to study unobservable events in deep time. In the case of functional analysis, the degree of missing data associated with reconstructing musculoskeletal anatomy and neuronal control in extinct organisms has, in the eyes of some scientists, rendered detailed functional analysis of fossils intractable. Such a prognosis may indeed be realized if palaeontologists attempt to recreate elaborate biomechanical models based on missing data and loosely justified assumptions. Yet multiple enabling methodologies and techniques now exist: tools for bracketing boundaries of reality; more rigorous consideration of soft tissues and missing data and methods drawing on physical principles that all organisms must adhere to. As with many aspects of science, the utility of such biomechanical models depends on the questions they seek to address, and the accuracy and validity of the models themselves.