Interaction of pH,density, and priority effects on the survivorship and growth of two species of hylid tadpoles

Summary We examined the interactions of an abiotic factor (pH) and a biotic factor (density) on the survival and growth of two species of anuran larvae (Hyla gratiosa and Hyla femoralis) in outdoor tanks. Three levels of pH (4.3, 4.6, or 6.0) and three levels of density (0, 30 or 60 embryos) were arranged in a blocked design and replicated three times for Hyla gratiosa. At the end of this experiment the effects of pH (4.3, 4.6, or 6.0), density of H. femoralis (30 or 60), and prior use by H. gratiosa (at 0, 30, or 60 larvae per tank) on the survival and growth of H. femoralis, were examined. Higher density increased larval period and decreased size at metamorphosis of H. gratiosa. Lower pH decreased survival rate and also decreased size at metamorphosis. Body sodium concentrations were lowest at the low pH values. Lower pH increased the susceptibility of H. gratiosa tadpoles to the adverse effects of higher densities. For H. femoralis higher density decreased survival, increased larval period and decreased size at metamorphosis. Hyla femoralis also had lower survivorship at low pH and exhibited decreased size at metamorphosis. However, unlike the results with Hyla gratiosa, there were no interactive effects between pH and density for any of the life-history traits studied. The effect of previous colonization by H. gratiosa on H. femoralis survival was facilitative. Body sodium concentrations of H. femoralis were lowest at the highest pH value. Metamorphs of the same size had much lower levels of sodium in H. femoralis than H. gratiosa. In general, H. femoralis was less affected by pH variation than H. gratiosa. These results demonstrate that abiotic factors can interact strongly with biotic effects such as density and they suggest that interspecific interactions can be strongly modulated by the background abiotic environment.     

ADVERTISEMENT CALLS OF THE TREE FROGS,HYLA ARBOREA AND HYLA SAVIGNYI (ANURA: HYLIDAE) IN TURKEY

ABSTRACT

The systematic classification of tree frogs (Hyla) inhabiting different regions of the country of Turkey is unclear. Recordings of natural advertisement calls of individual male tree frogs in different locations in Turkey were analyzed to determine variation in acoustic features that may be related to taxonomic status. Multivariate analysis of covariance showed that call duration, intercali interval and number of pulses per call varied significantly between frogs in different locales. Call duration, intercall interval, and number of pulses per call were related to air temperature. Dominant frequency differed significantly between different groups of frogs, and was temperature-independent. These data are consistent with the hypothesis that tree frogs in Turkey represent two distinct species, Hyla arborea and Hyla savignyi.     

Ultrastructure of Spermiogenesis in Hyla japonica (Anura,Amphibia)

Spermiogenesis in Hyla japonica was examined using transmission electron microscopy. Features which have not been reported previously during spermiogenesis of anurans include: (1) the neck of pericentriolar material is well developed, (2) the perforatorium contains many microtubules, (3) the tail is composed of axoneme and axial rod, but has no undulating membrane. Such a structure has not been described in amphibians, although it occurs in fish and mammals, and (4) the modified microtubules have one to two long projections. So far they have not been observed in any species except Gerris and Aphyosemion. With the exception of the above characteristics, the spermatozoa of Hyla japonica are remarkably similar to those of Bufo sp.     

Osteological development of Cope's Gray Treefrog,Hyla chrysoscelis

Hyla chrysoscelis, Cope's Gray Treefrog, is a generalized treefrog found throughout much of east‐central North America. Although it is a model for many behavioural and ecological studies, little is known of its skeletal morphology or development. Herein, we describe the postembryonic skeletal development and adult osteology of H. chrysoscelis. The adult skull is well ossified with slight dermal ornamentation, the postcranial and tadpole skeletons are fairly non‐distinct with no obvious novel morphologies, and the Gosner stage by which bony elements first appear varies. We compare the rank order sequence of ossification to that of its sibling species Hyla versicolor and use examples from this study to demonstrate current complications with conducting ossification sequence meta‐analyses.     

Comparing bivariate reaction norms among species: time and size at metamorphosis in three species of Hyla (Anura: Hylidae)

Summary Univariate and bivariate methods for comparing norms of reaction among species are discussed and illustrated with an example using North American hylid treefrogs. Norms of reaction for size at metamorphosis (SM) and length of larval period (LP) were compared among treefrog species raised at different food levels (Hyla cinerea vs H. gratiosa) and at different temperatures (H. cinerea vs H. gratiosa vs H. squirella). Hyla cinerea and Hyla gratiosa show parallel norms of reaction across food levels and temperatures. Across temperatures, H. squirella shows a much smaller change in SM relative to change in LP than do H. cinerea and H. gratiosa. This difference in shape of reaction norms may reflect different histories of selection resulting from these species' use of different larval habitats.     

Karyotypes of two Australian hylids

Karyotypes of the Australian frogs Hyla caerulea (White) and Hyla phyllochroa Günther were prepared and analysed from colchicine-treated, primary cultures of adult heart, lung and kidney. The diploid chromosome number for both species is 26, which resembles that of most known karyotypes of Papuan hylids but differs from that of the genus Hyla in other regions. A statistical comparison, involving data from arm ratios, centromere indices and relative lengths of metaphase chromosomes of H. caerulea incubated at 31 and 37° C respectively, showed that the differences were non-significant at the 5% level. Similar treatment of data from cells of Hyla caerulea and Hyla phyllochroa incubated at 31°C failed to show a significant species difference. Absolute lengths of haploid chromosome complements of the two species at the same temperature were found to differ significantly. No evidence of sexual heteromorphism was found. The occurrence of a terminally situated, heterochromatic zone associated with the longer arm of the 11th pair of chromosomes is described and compared with the known condition in other hylids.     

CALL ANALYSIS OF TRIPLOID HYBRIDS RESULTING FROM DIPLOID-TETRAPLOID SPECIES CROSSES OF HYLID TREE FROGS

ABSTRACT

Calls produced by hybrids resulting from laboratory crosses of tetraploid Hyla versicolor females and either diploid Hyla chrysoscelis (type I) or Hyla arborea (type II) males were induced through manipulation of environmental conditions. Type I hybrids produced trilled calls similar in note repetition to H. versicolor, but more similar in dominant frequency to H. chrysoscelis. Mean duration was shorter than in both parent calls. Type II hybrids produced calls which were longer in duration and lower in note repetition rate than H. versicolor, but shorter in duration and higher in note repetition rate than H. arborea. Dominant frequency of type II hybrids was lower than in H. arborea but not significantly different than in H. versicolor. Hybrid calls were not strictly intermediate, and may provide information regarding parental relationships.     

SPECIATION BY POLYPLOIDY IN TREEFROGS: MULTIPLE ORIGINS OF THE TETRAPLOID,HYLA VERSICOLOR

Speciation by polyploidy is rare in animals, yet, in vertebrates, there is a disproportionate concentration of polyploid species in anuran amphibians. Sequences from the cytochrome b gene of the mitochondrial DNA (mtDNA) were used to determine phylogenetic relationships among 37 populations of the diploid-tetraploid species pair of gray treefrogs, Hyla chrysoscelis and Hyla versicolor. The diploid species, H. chrysoscelis, consists of an eastern and a western lineage that have 2.3% sequence divergence between them. The tetraploid species, H. versicolor, had at least three separate, independent origins. Two of the tetraploid lineages are more closely related to one or the other of the diploid lineages (0.18%–1.4% sequence divergence) than they are to each other (1.9%–3.4% sequence divergence). The maternal ancestor of the third tetraploid lineage is unknown. The phylogenetic relationships between the two species and among lineages within each species support the hypothesis of multiple origins of the tetraploid lineages.     

Proximate developmental causes of limb length variation between Hyla cinerea and Hyla gratiosa (Anura: Hylidae)

Hyla cinerea and H. gratiosa are closely related treefrogs that differ at metamorphosis in the length of the hind limb relative to snout vent length. A developmental series of larvae of the two species raised under uniform laboratory conditions reveals that H. cinerea have proportionately longer legs at metamorphosis than H. gratiosa for the following reasons: 1) Hyla cinerea initiate limb growth at a smaller body size than H. gratiosa, 2) H. cinerea have a faster rate of limb growth, and 3) body growth rates in H. cinerea and H. gratiosa are nearly identical until just before metamorphosis.     

Polymorphism in the location of the 18S and 28S rRNA genes on the chromosomes of the diploid-tetraploid treefrogs Hyla chrysoscelis and H. versicolor

The chromosomes of the diploid Hyla chrysoscelis and its tetraploid sibling species H. versicolor were studied with AgNO₃ staining and in situ hybridization to determine the chromosome location of the 18S and 28S ribosomal RNA (rRNA) genes. A total of 236 Hyla chrysoscelis from 34 localities in 15 U.S. states and 100 H. versicolor from 15 localities in 12 states were examined. The rRNA gene sites were extremely variable in H. chrysoscelis, and also variable, but to a lesser extent, in H. versicolor. The most common rRNA gene site in both H. chrysoscelis and H. versicolor was on the short arm of chromosome 6. All of the rRNA gene locations seen in H. versicolor were also seen in H. chrysoscelis, supporting the hypothesis that the tetraploid H. versicolor arose from H. chrysoscelis. Although polymorphic rRNA gene sites in H. versicolor may reflect the positions of the rRNA genes in H. chrysoscelis ancestors, the origin of the extreme variability of such sites in H. chrysoscelis seems more obscure. Possible explanations include inversions, translocations, mobile genetic elements or a combination of some or all of these.     

Configurations of the rectus abdominis muscle of anuran tadpoles

Configurations of the rectus abdominis in tadpoles of 60 anuran species in 13 families were examined. This muscle is present by Gosner stage 25 and does not change until late in metamorphosis. The anterior terminus of the r. abdominis usually is a straight, transverse front or fan-shaped array only loosely associated with the rectus cervicis. In some suspension feeders (Rhinophrynus, Xenopus, but not microhylids), macrophagous suction feeders (Lepidobatrachus) and bromeliad inhabitants (Hyla bromeliacia) the r. abdominis is contigous with the r. cervicis which continues anterolaterally and dorsally from the wall of the spiracular cavity. Suctorial forms (Scutiger), those that live in confined spaces (burrowers, Centrolenella; bromeliad inhabitants, Hyla bromeliacia), and the taxa that have the r. cervicis and r. abdominis contiguous all have closed myosepta; the myosepta of other taxa; except for ones with large tadpoles (e.g., Rana catesbeiana), have large gaps between at least anterior myotomes. These initial data suggest that the configurations of the r. abdominis have responded to selection based on ecomorphological function and convergence among lineages are noted. © 1992 Wiley-Liss, Inc.     

Diet composition of the endangered giant water bug Lethocerus deyrolli (Hemiptera: Belostomatidae) in the rice fields of Japan: Which is the most important prey item among frogs,fish, and aquatic insects?

The food habits of the endangered giant water bug, Lethocerus deyrolli, were studied in the rice fields of Nose, in the north of Osaka Prefecture, Japan. Field observations revealed that frogs were the most important prey item. Frogs represented 86.4% and 78.6% in the diet of L. deyrolli in spring and summer, respectively. Among seven species of three families (Hylidae, Rhacophoridae, and Ranidae) exploited by L. deyrolli, the most important food item was adult Hyla japonica in spring and juvenile Rana nigromaculta in summer. Fish and aquatic arthropods were not considered important foods for L. deyrolli. The frog‐dependent food habits indicate that the recovery and conservation of frogs should be prioritized to protect L. deyrolli from extinction.     

Comparative skeletogenesis of the Oriental Tree Frog Hyla orientalis (Anura: Hylidae)

The ontogeny of skeletal development of the Oriental Tree Frog, Hyla orientalis (Bedriaga, 1890), is followed from tadpole to froglet and compared with four other hylid species i.e., Hypsiboas lanciformis, Hypsiboas pulchellus, Phyllomedusa vaillanti and Pseudis platensis. Our analysis and comparison is based on cleared and double-stained specimens. The parasphenoid is the first skeletal element that ossified in Hyla orientalis is followed by the exoocipital, prootic, axial and appendicular skeletal elements, frontoparietal, nasal, upper jaw, squamosal, pterygoid, lower jaw, vomer and quadratojugal. Ossification initiated at early Stage 33 in Hyla orientalis whereas in other hylids it begins after Stage 37. Major transformations of the larval jaw were visible after Stage 42. Cranial ossifications were completed after appendicular and axial ossifications of Hyla orientalis. All skeletal elements ossified before the end of the metamorphosis, a unique feature of Hyla orientalis in comparison to the other hylids studied here.     

Predator-prey relationships among larval dragonflies,salamanders, and frogs

Summary Tadpoles of the barking tree frog, Hyla gratiosa, are abundant in spring and summer in some ponds and Carolina bays on the Savannah River Plant near Aiken, South Carolina. To determine how these tadpoles survive in the presence of predaceous salamander larvae, Ambystoma talpoideum, and larvae of an aeshnid dragonfly, Anax junius, we determined fields densities and sizes of the predators and the prey and conducted predation experiments in the laboratory. Tadpoles rapidly grow to a size not captured by Ambystoma, although Anax larvae can capture slightly larger tadpoles. Differing habitat preferences among the tadpoles and the two predator species probably aid in reducing predation pressure. Preliminary work indicates that the tadpoles may have an immobility response to an attack by a predator. In addition, the smallest, most vulnerable tadpoles have a distinctive color pattern which may function to disrupt the body outline and make them indiscernable to predators.     

A case of Batesian mimicry between a myrmecophilous staphylinid beetle, Pella comes, and its host ant, Lasius (Dendrolasius) spathepus: an experiment using the Japanese treefrog, Hyla japonica as a real predator

Some myrmecophilous animals show myrmecomorphy, however, its adaptive significance is still controversial. We investigated a possible benefit of Batesianmimicry between a myrmecophilous staphylinid beetle, Pella comes, and its host ant, Lasius (Dendrolasius) spathepus, by using a common ant predator, the Japanese treefrog, Hyla japonica. In the field, H. japonica were found to feed on numerous ants and other insects, but in laboratory experiments they refused feeding on L. spathepus. L. spathepus was highly repellent to these frogs, while P. comes was potentially palatable. After repeated contacts with L. spathepus which led to its avoidance the treefrogs started to reject P. comes as well . This suggests that myrmecomorphy is beneficial to P. comes, reducing the risk of predation, and that it , may represent a case of Batesian mimicry. may represent a case of Batesian mimicry. Received 15 February 2005; revised 12 April 2005; accepted 18 April 2005.     

Laich-Fressen durch Kaulquappen als mögliche Ursache spezifischer Biotoppräferenzen und kurzer Laichzeiten bei europäischen Froschlurchen (Amphibia,Anura)

Zusammenfassung In einem Kunstweiher aufgezogene Kaulquappen von Rana temporaria fressen in der Versuchsanordnung von Abb. I den Laich folgender sympatrischer Anuren-Arten: Rana temporaria (Kannibalismus), Rana esculenta, Rana ridibunda, Bufo bufo, Bufo calamita, Bombina variegata und Hyla arborea (Laichräubern). — Daß die Kaulquappen der früh im Jahr laichenden Rana temporaria später gelegten arteigenen und artfremden Laich fressen, wirkt als Selektionsdruck in folgende Richtungen: Rana temporaria züchtet sich selbst eine kurze Laichzeit. Spät in seichtem Wasser laichende Arten werden im Biotop von Rana temporaria unterdrückt (Bombina variegata, Bufo calamita, Hyla arborea). Rana esculenta, Rana ridibunda und Bufo bufo sind dem Laichräubern durch Kaulquappen wenig ausgesetzt, weil sie in größeren Wassertiefen laichen. Zudem ist Bufo bufo ebenfalls ein Frühlaicher. Alytes obstetricans steht außer Konkurrenz, weil er die Eier nicht ins Wasser setzt. — Untersuchungen über die Biotoppräferenzen und Laichzeiten der betroffenen Arten bestätigen diese Interpretation.

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Summary Tadpoles of Rana temporaria which have been raised in an artificial pond prey in the experimental situation shown in Fig. 1 upon the spawn of the following sympatric European Anura species: Rana temporaria (cannibalism), Rana esculenta, Rana ridibunda, Bufo bufo, Bufo calamita, Bombina variegata and Hyla arborea (predation). — Rana temporaria is an early breeder. Thus selection pressure acts in the following ways: Rana temporaria establishes for itself an explosive breeding pattern. Species which breed later in the year in shallow water are suppressed in the biotope of Rana temporaria and develop other biotope preferences (Bombina variegata, Bufo calamita, Hyla arborea). Rana esculenta, Rana ridibunda and Bufo bufo spawn in deeper water. Therefore they are scarcely exposed to the tadpoles of Rana temporaria. Moreover Bufo bufo, too, is an early breeder. Alytes obstetricans laying no spawn in the water is not endangered. — An analysis of biotope preferences and breeding times of the species involved confirms these interpretations.

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Mit Unterstützung des Schweizerischen Nationalfonds zur Förderung der wissenschaftlichen Forschung.     

Embryonic development of the tree frog, Hyla arborea

A staged series with time data and original photographs of the embryonic development of the tree frog, Hyla arborea are presented, following Gosner’s generalized table. To contribute to species specific embryological studies of anurans, the external features of development of this species were described. Duration of embryonic development, from fertilization to 25^th stage, is about 9 days at 20 ± 1 °C. Cleavage is holoblastic and unequal. Unlike the typical anuran development, neurular rotation was not observed in stage 15 or in any other stages. Other developmental stages resulted in a normal-looking. Embryos of H. arborea hatched at stage 20 or 21, which varied individually. Hatching occurred in the 4^th or 5^th day after fertilization. The mean total length of hatchlings was calculated as 6.38 mm. The mean size of ovum was estimated 1.4 mm in diameter.     

The separate and combined effects of harmonic structure, phase, and FM on female preferences in the barking treefrog (Hyla gratiosa)

In natural advertisement calls of the barking treefrog, Hyla gratiosa, a small amount of incoherent frequency modulation (FM) is present. Incoherency in the FM of a call creates inharmonicity and phase changes between its frequency components. In this study, the combined and separate effects of the harmonic structure, phase spectrum, and FM of an advertisement call on female choice were tested. The harmonic structure of a call can have a direct effect on female preference; females showed a significant preference for static-inharmonic calls over static-harmonic calls. Neither differences in phase or FM alone conferred a preference in two choice tests. However, when FM is present in both calls it does influence female preference for harmonic structure -namely harmonic calls become preferable to inharmonic calls. This reversal of female preference for inharmonicity in a call by the presence of FM suggests that call parameters may interact, and thereby effect mate choice.Abbreviations AP amphibian papilla - BP basilar papilla - FM frequency modulation - PM phase modulation - HS harmonic structure - GB Gaussian Band     

Opalinidae (Sarcomastigophora) in North American Amphibia. GenusOpalina Purkinje & Valentin, 1835

During 1992, the cloaca of 83 specimens of North American Amphibia from Algonquin Park and Kingston, Ontario, Canada, and from El Paso County, Texas, USA, comprising 14 species in seven families, were checked for opalinids. Opalina martinonae n. sp. was found in five of nine Rana sylvatica (Ranidae); O. obtrigonoidea Metcalf, 1923 in six of 11 Bufo americanus (Bufonidae) and two of nine Pseudacris crucifer (Hylidae); O. pickeringi Metcalf, 1923 in two of nine Pseudacris crucifer and in one of six Rana pipiens; and O. virguloidea Metcalf, 1923 in two of six Hyla versicolor (Hylidae). Opalina flava Stokes, 1884 is proposed for Cepedea flava (Stokes, 1884) Metcalf, 1923. The occurrence, evolution and biogeography of North American Opalina are discussed.     

FORMS OF CALL OVERLAP AND THEIR IMPACT ON ADVERTISEMENT CALL ATTRACTIVENESS TO FEMALES OF THE GRAY TREEFROG,HYLA VERSICOLOR

ABSTRACT

Male gray Lreefrogs, Hyla versicolor, advertise for females in choruses in which calls often overlap. Previous research has demonstrated that such acoustic interference among conspecifics can impair the ability of a male to attract a mate. However, the stimulus set used in those studies was quite limited. Here we performed single-speaker and two-speaker tests of phonotaxis with female Hyla versicolor to explore the impact of different degrees and forms of call overlap on the relative attractiveness of male signals. We found that: (1) overlapped calls are recognized as acceptable signals by females, although they are less attractive than calls that are not overlapped; (2) the relative phase of the overlapping calls matters. When calls were time-shifted by 50% of their pulse-period (180 degree offset), attractiveness of overlapped calls was reduced relative to non-overlapped calls as much as it would be by a reduction in intensity of about 12 dB SPL or a drop in call duration of between one third to one half. A 90 degree phase offset was equivalent to a 3 dB drop in SPL; (3) partial call overlap may not impair call attractiveness if one third or less of the call is overlapped; (4) if the ending portion of a call is overlapped it may impair attractiveness less than if previous segments of a call are obscured; (5) if most (three fourths or more) of a male's calls are free of interference, overlap of his remaining calls does not significantly impair his attractiveness.