Experimental morphology of the feeding mechanism in salamanders

The subarcualis rectus I muscle (SAR) in the feeding mechanism of four tiger salamanders (Ambystoma tigrinum) was removed early in ontogeny and these individuals were allowed to complete metamorphosis. This procedure resulted in postmetamorphic tiger salamanders which differed from control individuals in the size (and thus force generating capacity) of the SAR muscle. The experimental manipulation of muscle ontogeny allowed a test of previous hypotheses of SAR function in postmetamorphic individuals. Multivariate analysis of variance for kinematic variables measured from high-speed video records of feeding revealed that experimentally modified tiger salamanders did not protract the hyobranchial apparatus or project the tongue from the mouth during feeding. Removal of the SAR muscle resulted in significantly reduced hyobranchial elevation in the buccal cavity and reduced maximum tongue projection distance.     

Functional morphology of the feeding mechanism in aquatic ambystomatid salamanders

This study addresses four questions in vertebrate functional morphology through a study of aquatic prey capture in ambystomatid salamanders: (1) How does the feeding mechanism of aquatic salamanders function as a biomechanical system? (2) How similar are the biomechanics of suction feeding in aquatic salamanders and ray-finned fishes? (3) What quantitative relationship does information extracted from electromyograms of striated muscles bear to kinematic patterns and animal performance? and (4) What are the major structural and functional patterns in the evolution of the lower vertebrate skull? During prey capture, larval ambystomatid salamanders display a kinematic pattern similar to that of other lower vertebrates, with peak gape occurring prior to both peak hyoid depression and peak cranial elevation. The depressor mandibulae, rectus cervicis, epaxialis, hypaxialis, and branchiohyoideus muscles are all active for 40–60 msec during the strike and overlap considerably in activity. The two divisions of the adductor mandibulae are active in a continuous burst for 110–130 msec, and the intermandibularis posterior and coracomandibularis are active in a double burst pattern. The antagonistic depressor mandibulae and adductor mandibulae internus become active within 0.2 msec of each other, but the two muscles show very different spike and amplitude patterns during their respective activity periods. Coefficients of variation for kinematic and most electromyographic recordings reach a minimum within a 10 msec time period, just after the mouth starts to open. Pressure within the buccal cavity during the strike reaches a minimum of ?25 mmHg, and minimum pressure occurs synchronously with maximum gill bar adduction. The gill bars (bearing gill rakers that interlock with rakers of adjacent arches) clearly function as a resistance within the oral cavity and restrict posterior water influx during mouth opening, creating a unidirectional flow during feeding. Durations of electromyographic activity alone are poor predictors of kinematic patterns. Analyses of spike amplitude explain an additional fraction of the variance in jaw kinematics, whereas the product of spike number and amplitude is the best statistical predictor of kinematic response variables. Larval ambystomatid salamanders retain the two primitive biomechanical systems for opening and closing the mouth present in nontetrapod vertebrates: elevation of the head by the epaxialis and depression of the mandible by the hyoid apparatus.     

Functional morphology and evolution of tail autotomy in salamanders

Basal tail constriction occurs in about two-thirds of the species of plethodontid salamanders. The constriction, which marks the site of tail autotomy, is a result of a reduction in length and diameter of the first caudal segment. Gross and microscopic anatomical studies reveal that many structural specializations are associated with basal constriction, and these are considered in detail. Areas of weakness in the skin at the posterior end of the first caudal segment, at the attachment of the musculature to the intermyotomal septum at the anterior end of the same segment, and between the last caudosacral and first caudal vertebrae precisely define the route of tail breakage. During autotomy the entire tail is shed, and a cylinder of skin one segment long closes over the wound at the end of the body. It is suggested that specializations described in this paper have evolved independently in three different groups of salamanders. Experiments and field observations reveal that, contrary to expectations, frequency of tail breakage is less in species with apparent provisions for tail autotomy than in less specialized species. The tail is a very important, highly functional organ in salamanders and it is suggested that selection has been for behavioral and structural adaptations for control of tail loss, rather than for tail loss per se.     

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.     

Ontogenetic convergence and evolution of foot morphology in European cave salamanders (Family: Plethodontidae)

Background  

A major goal in evolutionary biology is to understand the evolution of phenotypic diversity. Both natural and sexual selection play a large role in generating phenotypic adaptations, with biomechanical requirements and developmental mechanisms mediating patterns of phenotypic evolution. For many traits, the relative importance of selective and developmental components remains understudied.     

Shaping intraspecific variation: Development,ecology and the evolution of morphology and life history variation in tiger salamanders

The tiger salamander,Ambystoma tigrinum, is a geographically widespread, morphologically variable, polytipic species. It is among the most variable species of salamanders in morphology and life history with two larval morphs (typical and cannibal) and three adult morphs (metamorphosed, typical branchiate, cannibal branchiate) that vary in frequency between subspecies and between populations within subspecies. We report morphometric evidence suggesting that branchiate cannibals arose through intraspecific change in the onset or timing of development resulting in the wider head and hypertrophied tooth-bearing skull bones characteristic of this phenotype. We also quantified bilateral symmetry of gill raker counts and abnormalities, then evaluated fluctuating asymmetry as a measure of the developmental stability of each morph. There was a significant interaction between fluctuating asymmetry of developmental abnormalities in cannibals and typicals and the locality where they were collected, suggesting that relative stability of each phenotype could vary among populations. While altered timing of developmental events appears to have a role in the evolution and maintenance of morphs, novel phenotypes persist only under favorable ecological conditions. Predictability of the aquatic habitat, genetic variation, kinship, body size, intraspecific competition and predation all affect expression and survival of the morphs inA. tigrinum. This taxon provides an excellent model for understanding the diversity and complexity of developmental and ecological variables controlling the evolution and maintenance of novel phenotypes.     

Form and functional morphology of Raetellops pulchella (Bivalvia: Mactridae): an example of convergent evolution with anomalodesmatans

Abstract. The bivalve Raetellops pulchella is a highly specialized, deposit-feeding member of the Mactridae. Studies of its form and function provide an example of how the bivalve body plan can be modified to facilitate the exploitation of mud as a food resource, and help in understanding how this lifestyle has evolved. Adaptations to this lifestyle include an overall reduction in ctenidial size and loss of the descending lamellae of both outer demibranchs. This reduction is associated with the enlargement of the labial palps to process inhaled sediment. In the mantle cavity, a waste canal below the posterior mantle flaps facilitates pseudofeces removal. The midgut is long and capacious, presumably to cope with the large amounts of ingested organic material. In addition, individuals of R. pulchella have unusually thin, brittle, and rostrate shells, with narrow siphonal gapes. They possess a shell buttress in each valve extending from the hinge plate to above the posterior adductor muscle. This buttress functions to prevent the brittle shell valves from fracturing when adduction occurs. A buttress is also seen in some representatives of the Anomalodesmata; in particular, the situation in R. pulchella is most like that seen in individuals of the similarly deposit-feeding species Offadesma angasi (Anomalodesmata: Periplomatidae). I interpret the similar shell form of these deposit-feeding clams as an example of convergent evolution.     

Habits, functional morphology and the evolution of pycnogonids

Recent work on functional morphology has revealed not only how a wide range of animals work, but shows the significance of their shapes in great detail. Also, sound evidence of evolution is provided. A new approach towards the understanding of Pycnogonida comes from an appreciation of the significance of their general habits and shapes and from the structure and mode of action of their legs. Recent fossil evidence shows that the arachnids had at least two terrestrial landings, occurring millions of years apart in time. At least two, but probably more, separate arachnid lines lived in the sea. It is concluded that pycnogonids evolved from one such aquatic group which never became terrestrial.     

The role of climate in the dynamics of a hybrid zone in Appalachian salamanders

I examined the potential influence of climate change on the dynamics of a previously studied hybrid zone between a pair of terrestrial salamanders at the Coweeta Hydrologic Laboratory, U.S. Forest Service, in the Nantahala Mountains of North Carolina, USA. A 16‐year study led by Nelson G. Hairston, Sr. revealed that Plethodon teyahalee and Plethodon shermani hybridized at intermediate elevations, forming a cline between ‘pure’ parental P. teyahalee at lower elevations and ‘pure’ parental P. shermani at higher elevations. From 1974 to 1990 the proportion of salamanders at the higher elevation scored as ‘pure’P. shermani declined significantly, indicating that the hybrid zone was spreading upward. To date there have been no rigorous tests of hypotheses for the movement of this hybrid zone. Using temperature and precipitation data from Coweeta, I re‐analyzed Hairston's data to examine whether the observed elevational shift was correlated with variation in either air temperature or precipitation from the same time period. For temperature, my analysis tracked the results of the original study: the proportion of ‘pure’P. shermani at the higher elevation declined significantly with increasing mean annual temperature, whereas the proportion of ‘pure’P. teyahalee at lower elevations did not. There was no discernable relationship between proportions of ‘pure’ individuals of either species with variation in precipitation. From 1974 to 1990, low‐elevation air temperatures at the Coweeta Laboratory ranged from annual means of 11.8 to 14.2 °C, compared with a 55‐year average (1936–1990) of 12.6 °C. My re‐analyses indicate that the upward spread of the hybrid zone is correlated with increasing air temperatures, but not precipitation, and provide an empirical test of a hypothesis for one factor that may have influenced this movement. My results aid in understanding the potential impact that climate change may have on the ecology and evolution of terrestrial salamanders in montane regions.     

Plant-feeding and non-plant feeding phytoseiids: differences in behavior and cheliceral morphology

In previous studies plant feeding behavior of plant- and non-plant feeding phytoseiids was never examined directly. Moreover, in these studies the cheliceral morphology of phytoseiids was not associated with their ability to feed on plants. In the present study, we monitored the plant-feeding behavior of Euseius scutalis and Amblyseius swirskii. Only E. scutalis was observed penetrating the leaf surface with the movable digit and feeding. Second, using a dye and coloring the gut as an indicator for feeding, we found that E. scutalis pierced an artificial membrane and fed whereas A. swirskii did not. Finally, to identify morphological characteristics typical of plant feeders versus non-plant feeders, we used scanning electron microscopy to examine the adaxial (inner) profile of the chelicerae in 13 phytoseiid species. The only parameter that distinguished between plant- and non-plant feeders was the ratio of the dorsal perimeter length of the fixed digit to the ventral perimeter length of the movable digit. Plant-feeders were characterized by ratio values greater than one whereas the values for non plant-feeders were lower than one. We suggest that a shorter and less curved movable digit, expressed by a high ratio, will facilitate the penetration of the leaf surface. Cheliceral traits proposed here as typical of plant feeders, were observed for five genera, indicating that plant-feeding may be more common in the Phytoseiidae than previously reported. We propose that the ability to feed on plants be added as a cross type trait of phytoseiid life-style types.     

Diet,feeding behavior,and jaw architecture of Taï monkeys: Congruence and chaos in the realm of functional morphology

We review feeding and mandibular anatomy in a community of West African monkeys. We use field observations, food material property data, and skeletal specimens from the Ivory Coast's Taï Forest to explore the factors that shape mandibular architecture in colobines and cercopithecines. Despite excellent geographic control across our sample, the fit between bone form (as conventionally described) and functional activity (as we perceive it) is not spectacular. We present a thought experiment to assess how well we could reconstruct diet in the Taï monkeys if we only had skeletons and teeth to study. This exercise indicated that we would be correct about half the time. Our analyses reinforce the notion that diet is anything but a monolithic variable and that better success at relating mandibular form to food must incorporate information on ingestive and processing behavior, geometric and material properties of foods, and both material and structural data on jaws themselves.     

Speciation, phylogeography and evolution of life history and morphology in plethodontid salamanders of the Eurycea multiplicata complex

Understanding the complex interactions among environment, genotype and ontogeny in determining organismal phenotypes is cental to many biological disciplines. The Eurycea multiplicata complex, endemic to the Interior Highlands (Ozark Plateau and Ouachita Mountains) of eastern North America, comprises a diverse radiation of paedomorphic surface-dwelling (E. tynerensis), metamorphic surface-dwelling (E. multiplicata multiplicata and E. m. griseogaster) and metamorphic subterranean (Typhlotriton spelaeus) hemidactyliine plethodontid salamanders. Portions of two mitochondrial genes, cytochrome-b and NADH dehydrogenase-4, totalling 1818 base pairs (bp) were sequenced for 70 ingroup individuals plus numerous outgroup taxa, to examine the biogeography and relationships among these morphologically disparate species. Results show the E. multiplicata complex to be monophyletic, with its two most divergent clades corresponding to geography, not morphology or life history. Transforming surface-dwelling populations from the Ouachitas (E. m. multiplicata) are sister to the Ozark taxa, including paedomorphic surface-dwelling (E. tynerensis), subterranean (T. spelaeus) and transforming surface-dwelling salamanders assigned to the 'subspecies'E. m. griseogaster. Among Ozark taxa T. spelaeus (deeply nested within Eurycea) is sister to a clade that includes E. m. griseogaster and E. tynerensis. Current taxonomy suggests that paedomorphic populations (E. tynerensis) from the western Ozarks are distinct from nearby transforming populations (E. m. griseogaster). However, paedomorphic and transforming salamanders do not form reciprocally monophyletic groups and many populations share almost identical haplotypes. Ancestral state reconstruction of life history traits shows that paedomorphosis arose independently from three to nine times. Most populations are either completely paedomorphic or completely transforming. This suggests that local habitat parameters strongly influence life history mode in this complex, either facultatively or by selection for particular genotypes.     

Phylogenetic relationships of Neotropical honeycreepers and the evolution of feeding morphology

Phylogenetic relationships among Neotropical honeycreepers were studied by using cytochrome b sequence data. Although honeycreepers were once placed in their own family (Coerebidae), these species did not form a monophyletic group in any of the phylogenies in this study. Thus, our results indicate that nectar-feeding evolved independently multiple times among these birds. The Neotropical honeycreepers are best considered disparate members of a larger radiation of tanagers and finches, in which bill size and shape, and associated feeding behaviors have changed frequently to fill a variety of niches. Our phylogenies also provide specific taxonomic recommendations on the placement of each honeycreeper genus. The use of the taxon Coerebidae is no longer warranted given the well-supported relationships between different honeycreeper species and a variety of tanagers and finches.     

Mammalian feeding and primate evolution: An overview

Most of the papers included in this volume are derived from presentations in a symposium on Mammalian Feeding at the 65th Annual Meetings of the American Association of Physical Anthropologists in North Carolina in 1996. The aims of this symposium were to gather together the preeminent researchers on mammalian mastication and document the state of research in that field. The symposium emphasized in vivo studies of mammalian feeding because of a paucity of recent reviews of this field, but included morphometric and modeling papers as well. Subsequently the papers were revised, and were submitted in spring 1998 for publication, pending the outcome of peer review. Copyright 2000 Wiley-Liss, Inc.     

Molecular mechanisms underlying sensorimotor cortex pyramidal neuron involvement in the feeding behavior of rabbits

Response in pyramidal neurons belonging to the sensorimotor cortex (37% of total nerve cells investigated in this zone) which were identified by stimulating the bulbar pyramids, were investigated during experiments on unrestrained rabbits. Pyramidal neurons having connections with the lateral hypothalamus were activated during operation of feeding behavior, while activity was inhibited in those unconnected with the lateral hypothalamus. Microiontophoretic application of a protein synthesis inhibitor to pyramidal neurons caused their ability to respond to ascending activating influences from the lateral hypothalamus to disappear. When pentagastrin was applied to these neurons following the protein synthesis blocker, they recovered their ability to participate in hypothalamic feeding reaction. It is suggested that synthesis and release of a gastrin-like peptide into the perineuronal space is required for sensorimotor cortex pyramidal neurons to participate in the organization of feeding behavior.P. K. Anokhin Institute of Normal Physiology, Academy of Medical Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 19, No. 5, 601–606, September–October, 1987.     

Leptin and the central control of feeding behavior

The discovery of leptin by Friedman and coll. in 1995 was a major step forward in our comprehensive view of energy homeostasis. Since the original paper, a tremendous amount of work has been performed in laboratories all over the world. Many recent reviews have described this work in details. In the present review, we focus on the role of leptin on food intake. It is accepted by most authors working in this field that the control of food intake can be divided in two closely-related system: the homeostatic system and the hedonic system. Leptin has been shown to act on both systems.     

Temporal variation in feeding morphology and size-structured population dynamics in fishes

Considerable variation in morphology associated with resource use is a classic example of local adaptation to the environment. We demonstrate that a temporal change in feeding morphology might occur within a population. In a 5-year natural field experiment, we estimated gill raker morphology, resource density and population dynamics of the roach and its competitor, the perch. Despite a variation in density of zooplankton resources and perch across years, no change in roach density was observed. However, gill raker morphology in roach covaried with size structure of the zooplankton resource across years. A laboratory experiment confirmed that gill raker morphology has a great effect on roach foraging efficiency on zooplankton and that there is a functional trade-off with regard to zooplankton foraging. We suggest that the response in gill raker structure is an adaptation to deal with the rapid population dynamics of zooplankton, which are in turn mediated by changes in the size structure of the competing perch.