The evolution of the amphibian lateral line system and its bearing on amphibian phylogeny

In modern amphibians that are aquatic the lateral line system is organized, by order, as follows: caecilians have electroreceptive ampullary organs and single rows of mechanoreceptive neuromast organs; generalized anurans have single rows of neuromasts that divide in a transverse plane to form secondary neuromasts or stitches, they do not have ampullary organs; generalized urodeles have ampullary organs, transverse stitches, and double or triple rows of neuromasts. Fossil evidence indicates that early amphibians had both ampullary organs and single rows of neuromasts embedded in bone. With time, receptors became epidermal in all three orders. Modern caecilians have retained the primitive receptor arrangement. I propose that the common ancestor of anurans and urodeles had transverse stitches, and that this character allies these two groups. Subsequent to the anuranurodele split, anurans lost their ampullary organs, perhaps concomitant with developing specializations for herbivory. Urodeles developed orthogonal neuromast couplets und triplets. In modern anurans und urodeles, transverse stitches are correlated with pond dwelling, while ampullary organs are correlated with carnivory, suggesting that the anuran-urodele ancestor(s) was a (were) pond-dwelling carnivore(s).     

Ganglioside variations in amphibian brain tissue

Variation in the content and pattern of brain gangliosides from three anurans and two urodeles was analysed by thin-layer chromatography. The pattern for all three anurans was basically similar, though quantitative differences among species were detected. Ganglioside patterns from the two salamanders were distinctively different from that of the anurans and from one another, and more closely resembled the patterns seen in other tetrapods. Ganglioside variations in amphibians appear to reflect phylogenetic relationships better than life history or ecological factors.     

Stages in follicle cell/oocyte interface during vitellogenesis in caecilians <Emphasis Type="Italic">Ichthyophis tricolor</Emphasis> and <Emphasis Type="Italic">Gegeneophis ramaswamii</Emphasis>: a transmission electron-microscopic study

We describe the ultrastructural organization of the vitellogenic follicle stages in two caecilian species. Monthly samples of slices of ovary of Ichthyophis tricolor and Gegeneophis ramaswamii from the Western Ghats of India were subjected to transmission electron-microscopic analysis, with special attention to the follicle cell/oocyte interface. In order to maintain uniformity of the stages among the amphibians, all the stages in the caecilian follicles were assigned to stages I–VI, the vitellogenic and post-vitellogenic follicles being assigned to stages III–VI. Stage III commences with the appearance of precursors of vitelline envelope material in the perivitelline space. Stages IV and V have been assigned appropriate substages. During the transition of stage III to stage VI oocytes, a sequential change occurs in the manifestations of follicle cells, perivitelline space, vitelline envelope and oocyte cortex. The vitelline envelope becomes a tough coat through the tunnels of which the macrovilli pass to interdigitate between the microvilli. The oocyte surface forms pinocytic vesicles that develop into coated pits and, later, coated vesicles. Contributions of the oocyte cortex to the vitelline envelope and of the follicle cells to yolk material via synthesis within them are indicated. The follicle cell/oocyte interface of vitellogenic follicles of these two caecilians resembles that in anurans and urodeles, with certain features being unique to caecilians. Thus, this paper throws light on the possible relationships of caecilians to anurans and urodeles with special reference to ovarian follicles. This research was supported by funds from the Kerala State Council for Science, Technology and Environment (KSCSTE), through the SARD facility, and by the FIST scheme of Department of Science and Technology, Government of India, New Delhi, to the Department of Zoology, University of Kerala, Thiruvananthapuram, and to the Department of Animal Science, Bharathidasan University, Thiruchirapalli (SR/FST/LSI-233/2002).     

Structural characteristics of genome organization in amphibians: Differential staining of chromosomes and DNA structure

Summary Differential staining patterns on amphibian chromosomes are in some respects distinct from those on mammalian chromosomes; C-bands are best obtained, whereas G- and Q-bands are either unobtainable (on anuran chromosomes) or coincide with C-bands (chromosomes of urodeles). In amphibians, rRNA genes are located at secondary constrictions, but in urodeles they are also found at other chromosome sites, the positions of these sites being strictly heritable. DNA content in amphibian cells is tens and hundreds times higher than in mammals. DNA contents in anurans and urodeles differ within certain limits: from 2 to 25 pg/N and from 30 to over 160 pg/N respectively. Species characterized by slow morphogenesis have larger genomes. Genome growth is normally due to an increase in the amount of repetitive DNA (mostly intermediate repetitive sequences), the amount of unique sequences being almost constant (11 pg/genome in urodeles, and 1.5 pg/genome in anurans). In anurans in general no satellite DNA was found, whereas such fractions were found in manyUrodela species. Nucleosome chromatin structure in amphibians is identical to that of other eukariotes. It is postulated that differences in chromosome banding between amphibians and mammals are due to differences in chromatin packing which in turn is related to the distinct organization of DNA repetitive sequences. It is likely that fish chromosomes have a similiar structure. A comparison of such properties as the chromosome banding patterns, variations in nuclear DNA content and some genome characteristics enable us to group fishes and amphibians together as regards chromosome structure, as distinct from amniotes - reptiles, birds and mammals. It is probable that in the ancient amphibians - ancestors of reptiles - chromatin packing underwent a radical transformation, following changes in the organization of DNA repetitive sequences.     

Hair Cell Polarization in the Inner Ear of a Caecilian,Ichthyophis glutinosus (Amphibia: Gymnophiona)

The inner ear of Ichthyophis glutinosus is described with emphasis on the position of the sensory organs and the polarization of the hair cells. The hair cell polarization patterns of the maculae, cristae and papilla basilaris is similar to previous observations in other tetrapods. The papilla amphibiorum shows a simpler bidirectional polarization than described in other amphibians. The papilla neglecta, a sensory organ in the utriculus shows a unidirectional posteriorly directed polarization. A neglecta has not been found in the utriculus of anurans and urodeles previously.     

The amphibian octavo-lateralis system and its regressive and progressive evolution

The phylogenetic and ontogenetic changes in the octavolateralis system of sarcopterygian fish and tetrapods, presumed to be important for the formation of an amphibian auditory system, are reviewed. The lateral line system shows rudimentation of lines and loss of ampullary electroreceptors in many amphibians; in some amphibians it never develops. The metamorphic changes of the lateral-line system show different patterns in the different amphibian lineages with metamorphic retention in most urodeles and metamorphic loss in most anurans. The multitude of both ontogenetic and phylogenetic changes of the lateral line system among amphibians do exclude any prediction as to how this system might have changed in ancestral amniotes. The most important auditory epithelium of the tetrapod inner ear, the basilar papilla, seems to be primitively present in all tetrapods and Latimeria. In two amphibian lineages there is a trend towards rudimentation and loss of the basilar papilla. Only in the third order, the anurans, a tympanic ear develops and the inner ear shows a progressive evolution of the auditory epithelia. Together with the known differences in the periotic labyrinth of amphibians and amniotes, this scenario suggests a parallel evolution of the amniotic and anuran auditory periphery. All mechanoreceptive hair cells of the lateral line system and the inner ear appear to receive a common and bilateral efferent innervation. Among amphibians this pattern is represented only in some urodeles, whereas anurans show a derived pattern with loss of a bilateral component and presumably also of a common neuromast/inner ear component. Changes in the rhombencephalic nuclei which receive octavo-lateralis afferent fibers show a trend towards development of auditory nuclei only in the anuran lineage. The phylogenetic appearance of an auditory nucleus in this lineage coincides with the complete absence of formation of ampullary electroreceptors. In contrast, the earlier claim of a correlation between a metamorphic loss of the lateral line system and the formation of an auditory nucleus is not supported by more recent data: an auditory nucleus develops in anurans already prior to metamorphosis and is present in all anurans even when they retain the neuromast system. In anurans with a metamorphic loss of the neuromasts, the second order neurons degenerate as well. This independence of the auditory and the second order lateral line nuclei is further substantiated by their separate projection to other brain areas, like the torus semicircularis of the midbrain, and their functional properties.(ABSTRACT TRUNCATED AT 400 WORDS)     

Evolutionary trends in adrenal gland of anurans and urodeles

Morphological, histological, ultrastructural, and developmental research on the adrenal gland of several species of anurans and urodeles belonging to different families is presented. Urodeles show a large variability in adrenal glandular structure without a clear taxonomic pattern, although increased compactness of the gland and mingling of steroidogenic and chromaffin cells are found only in some neourodeles. In anurans the glandular pattern may be divided into two subtypes: one more medial and diffuse, which is observed in frogs of the more primitive families; the other more lateral and aggregated, as seen in the more advanced families. The adrenal gland therefore increases in its compactness and aggregation of chromaffin and steroidogenic tissues in the transition from primitive to advanced families, both in urodeles and anurans. Until the end of metamorphosis, morphogenesis of the gland is similar in all amphibians studied. This process is extended after metamorphosis in the advanced anurans, in order that the gland may reach its definitive position.     

Amphibian behavioral diversity offers insights into evolutionary neurobiology

Recent studies have served to emphasize the unique placement of amphibians, composed of more than 8000 species, in the evolution of the brain. We provide an overview of the three amphibian orders and their respective ecologies, behaviors, and brain anatomy. Studies have probed the origins of independently evolved parental care strategies in frogs and the biophysical principles driving species-specific differences in courtship vocalization patterns. Amphibians are also important models for studying the central control of movement, especially in the context of the vertebrate origin of limb-based locomotion. By highlighting the versatility of amphibians, we hope to see a further adoption of anurans, urodeles, and gymnophionans as model systems for the evolution and neural basis of behavior across vertebrates.     

Neuromast topography in anuran amphibians

Generalized anuran tadpoles across families exhibit a similar neuromast morphology on their heads, as follows: (1) all neuromast lines known for anurans are present; (2) within these lines total neuromast number ranges from about 250 to 320; (3) neuromasts form linear stitches composed of two to three, but sometimes up to five, neuromasts; (4) neuromast linear dimensions are ? 10 μm; and (5) neuromasts contain ? 15 hair cells. Compared with generalized forms, stream, arboreal, carnivorous, and desert-pond forms have fewer neuromasts but they contain more hair cells. They do not, however, form stitches. Obligate midwater suspension-feeding forms, including Xenopus (Pipidae), Rhinophrynus (Rhinophyrnidae), and Phrynomerus (Microhylidae), form stitches that contain > six, but potentially up to 18 or more, loosely aggregated neuromasts. Xenopus and Rhinophrynus have large neuromasts (up to 40 μm across). Chiasmocleis (Microhylidae) tadpoles form stiches that are linearly arranged with up to ten neuromasts. Whereas urodeles can have more than one neuromast row per line and may form both linear and transverse stitches, anurans have only one row of neuromasts per line and form only transverse stitches. Neuromasts in anurans tend to be smaller and more circular than in urodeles and positioned flush with the epidermal surface. A greater percentage of anurans form stitches, and anurans have greater intrafamilial variation in stitch formation than do urodeles.     

Anuran postnasal wall homology: An experimental extirpation study

The nasal placode was extirpated unilaterally in Gosner stage 18–20 embryos of Rana sylvatica, R. palustris and R. pipiens, in order to test alternative proposed schemes of homology for the ethmoidal attachment of the palatoquadrate in anurans and urodeles. Absence of the nasal sac has no pronounced effect on the formation of larval chondrocranial structures. In contrast, in metamorphosed animals the lamina orbitonasalis and inferior prenasal process are the only nasal capsule structures present on the operated side. The medial nasal branch of the deep ophthalmic nerve passes forward over the dorsal surface of the lamina orbitonasalis, rather than through an orbitonasal foramen. Comparison with previous experimental work on urodeles supports the traditional homology of the anuran lamina orbitonasalis with the antorbital process of urodeles and other vertebrates. J. Morphol. 238:343–353, 1998. © 1998 Wiley-Liss, Inc.     

水蛭捕食两栖类卵

许多卵生动物具有对付食卵者的精巧策略以降低死亡率,因为早期发育阶段的捕食压力能显著影响具复杂生活史动物的生存。两栖动物卵被水蛭等很多物种所捕食。我们查阅了水蛭捕食两栖类卵的文献,发现27篇论文。在所报道的水蛭捕食两栖类卵的例子中,无尾类(3.6%,n=591)明显多于有尾类(1.6%,n=255)。此外,我们第一次记录到水蛭捕食四趾螈(Salamandrina perspicillata)卵,这也是西欧有尾类的第一例报道。我们没有发现两个两栖动物类群间存在差异。然而,当我们分别分析来自北美、欧洲和亚洲三个地区的数据时,发现北美的有尾类和无尾类间存在显著差异。水蛭可捕食各生活史阶段的两栖动物,因此,水蛭应归为一些两栖类的天敌。     

The skin of Ichthyophis (Amphibia: Caecilia): an ultrastructural study

The skin of the adult Ichthyophis spp. has been investigated using light microscopy and transmission electron microscopy, and the various epidermal and dermal integral cellular components of this genus, of the order Caecilia, are compared and contrasted with those of members of the Anura and Urodela among the Amphibia. In general the epidermis of Ichthyophis is typically amphibian in appearance, though in terms of size the cells are large and of urodelan dimensions. Apart from the epithelial cells, in addition the epidermis includes a number of specifically different cells, some doubtless arising in situ , possibly from the stratum germinativum, others entering the epidermis from elsewhere.
The dermis of caecilians is unique among living amphibians in possessing scales located in pockets, and for completeness their gross structure and arrangement are described and existing information on their ultrastructure is summarized. The cellular composition and arrangement of the dermal glands are described and the glandular components in the Amphibia are compared.     

Mortality of vertebrates on a road crossing the Biebrza Valley (NE Poland)

Mortality of vertebrates was monitored on a local road running across Poland’s Biebrza River Valley during 2 years (August 2005–July 2006). On the basis of distance from the river and surrounding habitats, the road (of total length 2,510 m) was divided into three stretches. The road was monitored on foot by two people every month, over a few consecutive days. A total of 1,892 road kills representing at least 47 species were found. Of these, 90.7% were amphibians, 4.2% mammals, 3.1% birds and 2.0% reptiles. Most (70%) of the amphibians were anurans, with the common toad, common frog and moor frog among them together accounting for 82% of the total. Mortality among amphibians differed between months, most anurans dying in May and August, while a majority of Urodela are lost in October. The peaks in mortality were connected with the migration of adult amphibians in spring and juveniles in summer and autumn. The number of amphibians killed was greatest on the (wettest) stretch adjacent to the river and decreased with distance from it. Mortality among birds was highest in July—probably in association with the dispersal of young individuals. Among recorded mammalian road kills, there was a prevalence of small rodents (mainly voles) and insectivores (mainly shrews). Medium-sized mammals were found only accidentally. Mortality in general was conditioned by the number of anurans killed.     

Phylogeny and Differentiation of Reptilian and Amphibian Ranaviruses Detected in Europe

Ranaviruses in amphibians and fish are considered emerging pathogens and several isolates have been extensively characterized in different studies. Ranaviruses have also been detected in reptiles with increasing frequency, but the role of reptilian hosts is still unclear and only limited sequence data has been provided. In this study, we characterized a number of ranaviruses detected in wild and captive animals in Europe based on sequence data from six genomic regions (major capsid protein (MCP), DNA polymerase (DNApol), ribonucleoside diphosphate reductase alpha and beta subunit-like proteins (RNR-α and -β), viral homolog of the alpha subunit of eukaryotic initiation factor 2, eIF-2α (vIF-2α) genes and microsatellite region). A total of ten different isolates from reptiles (tortoises, lizards, and a snake) and four ranaviruses from amphibians (anurans, urodeles) were included in the study. Furthermore, the complete genome sequences of three reptilian isolates were determined and a new PCR for rapid classification of the different variants of the genomic arrangement was developed. All ranaviruses showed slight variations on the partial nucleotide sequences from the different genomic regions (92.6–100%). Some very similar isolates could be distinguished by the size of the band from the microsatellite region. Three of the lizard isolates had a truncated vIF-2α gene; the other ranaviruses had full-length genes. In the phylogenetic analyses of concatenated sequences from different genes (3223 nt/10287 aa), the reptilian ranaviruses were often more closely related to amphibian ranaviruses than to each other, and most clustered together with previously detected ranaviruses from the same geographic region of origin. Comparative analyses show that among the closely related amphibian-like ranaviruses (ALRVs) described to date, three recently split and independently evolving distinct genetic groups can be distinguished. These findings underline the wide host range of ranaviruses and the emergence of pathogen pollution via animal trade of ectothermic vertebrates.     

Taste organ growth and development in the direct developing frog Eleutherodactylus coqui (Lissamphibia: Eleutherodactylidae)

Previous research on amphibian taste organs concerned amphibians with a biphasic life history, that is, with larval period and metamorphosis. Direct developing frog species, such as Eleutherodactylus coqui, undergo a cryptic metamorphosis before hatching, and many larval‐specific features are vestigial or have been lost entirely from their ontogeny. Taste buds are present in larval stages of biphasically developing anurans and are replaced by taste discs during metamorphosis. One goal of this study was to characterize the ontogeny of taste buds and/or discs in E. coqui. The other goal was to examine correlations between body size and taste organ density and size in different regions of oral epithelium. The research reveals the presence of only one type of taste organ, characteristic of metamorphs of biphasic amphibians, namely taste disc. In addition, taste disc density and the area of the taste disc sensory zone change dramatically during growth.     

An identification of invariants in life history traits of amphibians and reptiles

While many morphological, physiological, and ecological characteristics of organisms scale with body size, some do not change under size transformation. They are called invariant. A recent study recommended five criteria for identifying invariant traits. These are based on that a trait exhibits a unimodal central tendency and varies over a limited range with body mass (type I), or that it does not vary systematically with body mass (type II). We methodologically improved these criteria and then applied them to life history traits of amphibians, Anura, Caudata (eleven traits), and reptiles (eight traits). The numbers of invariant traits identified by criteria differed across amphibian orders and between amphibians and reptiles. Reproductive output (maximum number of reproductive events per year), incubation time, length of larval period, and metamorphosis size were type I and II invariant across amphibians. In both amphibian orders, reproductive output and metamorphosis size were type I and II invariant. In Anura, incubation time and length of larval period and in Caudata, incubation time were further type II invariant. In reptiles, however, only number of clutches per year was invariant (type II). All these differences could reflect that in reptiles body size and in amphibians, Anura, and Caudata metamorphosis (neotenic species go not through it) and the trend toward independence of egg and larval development from water additionally constrained life history evolution. We further demonstrate that all invariance criteria worked for amphibian and reptilian life history traits, although we corroborated some known and identified new limitations to their application.     

Evolution of predetermined germ cells in vertebrate embryos: implications for macroevolution

The germ line is established in animal embryos with the formation of primordial germ cells (PGCs), which give rise to gametes. Therefore, the need to form PGCs can act as a developmental constraint by inhibiting the evolution of embryonic patterning mechanisms that compromise their development. Conversely, events that stabilize the PGCs may liberate these constraints. Two modes of germ cell determination exist in animal embryos: (a) either PGCs are predetermined by the inheritance of germ cell determinants (germ plasm) or (b) PGCs are formed by inducing signals secreted by embryonic tissues (i.e., regulative determination). Surprisingly, among the major extant amphibian lineages, one mechanism is found in urodeles and the other in anurans. In anuran amphibians PGCs are predetermined by germ plasm; in urodele amphibians PGCs are formed by inducing signals. To determine which mechanism is ancestral to the tetrapod lineage and to understand the pattern of inheritance in higher vertebrates, we used a phylogenetic approach to analyze basic morphological processes in both groups and correlated these with mechanisms of germ cell determination. Our results indicate that regulative germ cell determination is a property of embryos retaining ancestral embryological processes, whereas predetermined germ cells are found in embryos with derived morphological traits. These correlations suggest that regulative germ cell formation is an important developmental constraint in vertebrate embryos, acting before the highly conserved pharyngula stage. Moreover, our analysis suggests that germ plasm has evolved independently in several lineages of vertebrate embryos.     

Helminth community structure of sympatric eastern American toad, Bufo americanus americanus, northern leopard frog, Rana pipiens, and blue-spotted salamander, Ambystoma laterale, from southeastern Wisconsin

One-hundred twelve amphibians, including 51 blue-spotted salamanders, Ambystoma laterale, 30 eastern American toads, Bufo americanus americanus, and 31 northern leopard frogs, Rana pipiens, were collected during April-October 1996 from Waukesha County, Wisconsin and examined for helminth parasites. The helminth compound community of this amphibian assemblage consisted of at least 10 species: 9 in American toads, 8 in leopard frogs, and 3 in blue-spotted salamanders. American toads shared 7 species with leopard frogs, and 2 species occurred in all 3 host species. Although there was a high degree of helminth species overlap among these sympatric amphibians, statistically significant differences were found among host species and percent of indirect or direct-life cycle parasites of amphibian species individual component communities (chi2 = 1,015, P < 0.001). American toads had a higher relative abundance of nematodes, 59%, than larval cestodes, 31%, and larval and adult trematodes, 10%, whereas leopard frogs had a higher relative abundance of larval cestodes, 71.3%, and larval and adult trematodes, 25.3%, than nematodes 3.4%. This is related to ecological differences in habitat and dietary preferences between these 2 anuran species. Helminth communities of blue-spotted salamanders were depauperate and were dominated by larval trematodes, 94%, and few nematodes, 6%. Low helminth species richness in this host species is related to this salamander's relatively small host body size, smaller gape size, lower vagility, and more fossorial habitat preference than the other 2 anuran species. Adult leopard frogs and toads had significantly higher mean helminth species richness than metamorphs, but there was no significant difference in mean helminth species richness among adult and metamorph blue-spotted salamanders. Considering adult helminths, the low species richness and low vagility of caudatans as compared with anurans suggest that local factors may be more important in structuring caudatan helminth communities of salamanders than of anuran hosts. Helminth species infecting salamanders may be more clumped in their geographic distribution as compared with anurans, and the role of other hosts and their parasites at the compound community level may be important in structuring helminth communities of salamanders.     

Ready and Waiting: Delayed Hatching and Extended Incubation of Anamniotic Vertebrate Terrestrial Eggs

Some anamniotic aquatic vertebrates lay eggs in a terrestrialhabitat that is hostile to the survival of hatchings or larvae.These terrestrial eggs are ready and able to hatch at a particulardevelopmental time, but do not hatch until presented with suitableconditions for aquatic larval survival. Beyond this time, hatchingis possible whenever aquatic conditions occur. The durationof extended terrestrial incubation is dependent on the availabilityof energy for metabolism from the yolk. Extended incubationis useful for anamniotic eggs laid in terrestrial habitats whereconditions suitable for larval survival arrive with unpredictableor variable timing. Examples of anamniotes with delayed hatchingand extended terrestrial incubation can be found among teleostfishes, anurans, and caudate amphibians. This paper characterizesthe embryonic period, compares this mode with other forms ofdevelopmental plasticity in anamniotes, evaluates the constraintsand advantages of this life history mode, and examines how somefishes and amphibians are able to obtain the benefits of terrestrialityfor their eggs when the timing of the return to aquatic conditionsis not entirely predictable.