Spawn predation and larval growth inhibition as mechanisms for niche separation in anurans

Summary We have investigated interspecific interactions between larvae of the 3 native British anurans, Rana temporaria, Bufo bufo and B. calamita in the field and in the laboratory. Over a 3 year period, predation of B. calamita spawn by the two competitively superior species accounted for only 0.3–3.0% of the total deposited in the dune pools of the study site. However, in ponds with marked niche overlap destruction of natterjack spawn was often 100%. Furthermore, growth rates of B. calamita larvae surviving to hatch were grossly inhibited when large numbers (>0.1 per l) of common frog or toad tadpoles were present. Overall survival to metamorphosis by the competitively inferior species was always negligible in these situations. These results were confirmed by experiments using caged larvae, and by movement of R. temporaria spawn between pools. Growth could be reduced to <25% of uninhibited controls, and time taken to achieve metamorphosis extended by at least 40% in the presence of larger larvae. Laboratory experiments confirmed that inhibitory effects of both R. temporaria and B. bufo larvae on growth by B. calamita tadpoles was demonstrable at tadpole densities comparable with those found in the field.     

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.     

Factors determining tadpole vulnerability to predators: can prior experience compensate for a suboptimal shape?

We investigated the role of constitutive morphology and previous experience in predator avoidance in two anuran species associated with different larval habitats. In Rana temporaria, deeper tails and larger body size conferred selective advantage against dragonfly predation. Previous experience with predators had a positive influence on the survival of R. temporaria tadpoles equivalent to predator selection. By contrast, survival in Bufo bufo seems unrelated to tail shape or experience. This suggests that B. bufo lacks constitutive morphological defenses against insect predators, and that morphological and behavioral defenses could result more effective than chemical deterrents for these insect predators. A key novelty of this study is the observation that Rana tadpoles having prior experience with predators have an enhanced success in further encounters, and this occurs before the morphological induced defense has been established. This induced modification for R. temporaria, and its lack of for B. bufo, may be an important determinant of larval survival.     

The introduced fish, rotan (Perccottus glenii), depresses populations of aquatic animals (macroinvertebrates, amphibians, and a fish)

The fish rotan (Perccottus gleniiDybowski) was accidentally introduced into European Russia from the Amur River basin. Rotan is capable of colonising small waterbodies – favourable breeding sites of native amphibians. To reveal its influence on the native aquatic fauna, monitoring of small waterbodies has been carried out since 1994 in the region of Lake Glubokoe Reserve (Moscow Province, Russia). The fish's diet includes a wide range of animal species of all trophic levels. Rotan considerably decreases the species richness of aquatic macroinvertebrates and larval amphibians. As a rule, most amphibian species (Triturus cristatus, T. vulgaris, Rana temporaria, R. arvalis, R. lessonae) and the fish Carassius carassius failed to breed successfully in ponds inhabited by rotan. In contrast, the toad Bufo bufo bred successfully in such sites because its larvae are distasteful to rotan. Rotan–amphibian interactions are discussed.     

Suspension feeding of anuran larvae at low concentrations of Chlorella algae (Amphibia,Anura)

Summary Ingestion and filtering rates in larval Xenopus laevis, Bufo calamita, Rana temporia and Bufo bufo fed suspensions of Chlorella fusca were investigated. Concentrations were measured with a Coulter Counter. (1) For all species, filtration occurred at concentrations far below those reported by other authors for Rana sylvatica feeding on Chlorella pyrenoidosa. For Bufo bufo, only larvae near metamorphosis showed ingestion at low particle concentrations. Since buccopharyngeal ventilation continues even in the absence of food particles, this threshold feeding behaviour in the younger larvae must be due to different mechanisms to those found in Daphnia and Calanus studied by other authors: probably reduction of the buccal pumping rate and the mucus production of the filter apparatus. (2) For B. calamita, R. temporaria and X. laevis the highest suspension feeding efficiency was at early tadpole stages, corresponding with the high growth rate of these stages. (3) The life histories of the species provide the basis for understanding their different retention efficiencies and functional responses.Supported by the Deutsche Forschungsgemeinschaft-DFG     

Spectrochemical and histochemical analyses of tissues of grass frog and gray toad tadpoles developing under simulation of pollution by plumbum and ferrum

Visual, spectrochemical, and histochemical analyses of tadpoles of two tailless amphibian species (Rana temporaria L. and Bufo bufo L) developed under simulation of water pollution by plumbum and ferruginous alloys demonstrated an alimentary way of the delivery of these metal ions in the organism. Accumulation of these metal ions occurs in intestine and liver tissues and increases with the development.     

A Comparative Study of the Aminergic Structures in the Hypothalamus and the Hypophysis of Amphibian Larvae,with Special Reference to the Innervation of the Pars Distalis

The distribution and occurrence of the aminergic structures of the hypothalamus and the hypophysis of the investigated species are very similar, except for the pars distalis. In the tadpoles of the anuran species Rana temporaria, R. arvalis, R. dalmatina, R. escutenta, Bufo bufo and B. calamita, the pars distalis is aminergically innervated (the Falck-Hillarp technique), but these nerves disappear at the metamorphic climax. In the larvae of the urodelan species Triturus cristatus and T. vulgaris, the pars distalis is devoid of such nerves. Electron microscopical studies of R. temporaria, B. bufo and T. cristatus confirm these observations. A discussion of the functional significance of the pars distalis nerves and the phylogenetic role of the difference between anuran and urodelan species is given.     

Intra- und interspezifische Crowding-Effekte bei Kaulquappen der Kreuzkröte,Bufo calamita Laur.

Summary Tadpoles ofBufo calamita, a species that spawns in little temporary pools, were tested on intra- and interspecific crowding-effects. Water was conditioned by tadpoles of the following species:Bufo calamita (tadpoles of different age),Alytes obstetricans, Bombina variegata, Bufo bufo, Hyla arborea, Rana lessonae, R. ridibunda andR. temporaria. Tadpoles ofBufo calamita are most sensitive to water conditioned by older tadpoles of the own species during the first 2–3 weeks of development. They are also strongly inhibited by tadpoles ofRana temporaria andBufo bufo (that, as a rule, do not occur in the same habitat). Relative little effects are observed in water conditioned by tadpoles of the two species that occur very often in ecological sympatry withBufo calamita: Hyla arborea andBombina variegata. Young tadpoles ofBufo calamita are most tolerant of tadpoles of the same age of the own species. The crowding-effect increases with the number and weight of tadpoles/volume conditioning the water. In the age of 2–3 weeks the sensibility to intra- and interspecific crowding-effects fades. Thus inBufo calamita the age specific sensibility to crowding-effects in tadpoles may act as a density regulation mechanism in temporary pools.

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Mit Unterstützung des Schweizerischen Nationalfonds zur Förderung der wissenschaftlichen Forschung (Beitrag Nr. 3.256.69).     

Numbers,neighbors, and hungry predators: What makes chemically defended aposematic prey susceptible to predation?

Many chemically defended aposematic species are characterized by relatively low toxin levels, which enables predators to include them in their diets under certain circumstances. Knowledge of the conditions governing the survival of such prey animals—especially in the context of the co‐occurrence of similar but undefended prey, which may result in mimicry‐like interactions—is crucial for understanding the initial evolution of aposematism. In a one‐month outdoor experiment using fish (the common carp Cyprinus carpio) as predators, we examined the survival of moderately defended aposematic tadpole prey (the European common toad Bufo bufo) with varying absolute densities in single‐species prey systems or varying relative densities in two‐species prey systems containing morphologically similar but undefended prey (the European common frog Rana temporaria). The density effects were investigated in conjunction with the hunger levels of the predator, which were manipulated by means of the addition of alternative (nontadpole) food. The survival of the B. bufo tadpoles was promoted by increasing their absolute density in the single‐species prey systems, increasing their relative density in the two‐species prey systems, and providing ample alternative food for the predator. Hungry predators eliminated all R. temporaria individuals regardless of their proportion in the prey community; in treatments with ample alternative food, high relative B. bufo density supported R. temporaria survival. The results demonstrated that moderately defended prey did benefit from high population densities (both absolute and relative), even under long‐term predation pressure. However, the physiological state of the predator was a crucial factor in the survival of moderately defended prey. While the availability of alternative prey in general should promote the spread and maintenance of aposematism, the results indicated that the resemblance between the co‐occurring defended and undefended prey may impose mortality costs on the defended model species, even in the absence of actual mimicry.     

Comparative enzymological study of catalytic properties of liver monoamine oxidases in frogs

Comparative enzymological study of catalytical properties of monoamine oxidase (MAO) of liver of the marsh frog Rana ridibunda and common frog Rana temporaria has revealed certain features of similarity and differences between these enzymes. The MAOs from both studied biological sources show catalytic properties resembling those of the classical MAO of terrestrial vertebrates: they deaminate tyramine, tryptamine, serotonin, and benzylamine and do not deaminate histamine, have sensitivity to clorgyline, the specific inhibitor of the MAO A form, and deprenyl, the specific inhibitor of the MAO B form, and are not inhibited by 10⁻² M semicarbazide. Based on data of substrate-inhibitor analysis, a suggestion is put forward about the existence of two molecular forms of the enzyme in liver of the studied frog species. Quantitative interspecies differences have been revealed between liver MAO of Rana ridibunda and Rana temporaria in values of kinetic parameters of reactions of deamination of several substrates and in sensitivity to the inhibitors, deprenyl and clorgyline. In the species Rana temporaria the MAO activity in reaction of deamination of serotonin and benzylamine were virtually identical, whereas in the species Rana ridibunda these parameters for serotonin were almost one order higher than for benzylamine. In the species Rana ridibunda, selectivity of action of deprenyl was expressed many times weaker, while selectivity of the clorgyline—one order of magnitude stronger than in the species Rana temporaria. The catalytic activities towards all studied substrates of liver MAO of both studied amphibian species were several times lower as compared with the enzyme of rat liver.     

The oral apparatus of tadpoles of Rana dalmatina, Bombina variegata, Bufo bufo, and Bufo viridis (Anura)

The morphology and development of the larval oral apparatus of Rana dalmatina, Bombina variegata, Bufo bufo, and Bufo viridis are described and compared using scanning electron microscopy. The species show different arrangements of the mouthparts. The small oral apparatus of R. dalmatina larvae has three labial tooth rows on the upper labium, while there are four tooth rows on the lower labium with a medial gap in row proximal to the mouth. The margins of the oral apparatus are defined by papillae that encircle the lower labium. B. variegata tadpoles have two upper labial tooth rows and three lower labial tooth rows that are uninterrupted, unlike the ones of R. dalmatina. The mouth is encircled by papillae that are larger than those of R. dalmatina. The oral discs of tadpoles of both B. bufo and B. viridis are similar. They are defined by two upper labial tooth rows (the second of which is interrupted by a medial gap) and by three lower tooth rows that differ in lengths in the two Bufo species. Both species develop papillae on the mouth angles and in two rows on the upper labium. Some morphological differences among the oral discs of R. dalmatina, B. variegata, B. bufo, and B. viridis tadpoles can be attributed to phylogenetic differences, but most can be related to their varying feeding habits and/or to their dietary specializations.     

Discovery of <Emphasis Type="Italic">Ascomorpha ovalis</Emphasis> (Bergendal, 1892) (Rotifera,Gastropodidae), an indicator of oligotrophic conditions,in Glubokoe Lake (Moscow Region,Russia)

The rotifer Ascomorpha ovalis (Bergendal, 1892) was found for the first time in Glubokoe Lake (Moscow Region) in June 2010. Long-term observations of zooplankton dating back to 1897, including our own studies starting in 2004 and carried out on a yearly basis, had never recorded this species before 2010. The density of the population was 25 ind./L in the 0–10 m water layer. In 2011–2012, the species did not disappear but was naturalized in the lake. A. ovalis is an indicator of oligotrophic conditions. The tendency for ongoing oligotrophication of the pelagial of Glubokoe Lake is supported by the penetration and naturalization of Ascomorpha ovalis during recent years and the presence of the oligotrophic rotifer Gastropus hyptopus (Ehrenberg, 1838) detected in the lake earlier.     

Trematode larvae in snails of Lake Glubokoe

In the Augusts of 1984 and 1985, 6 species of snails in Lake Glubokoe were investigated: Viviparus viviparus (Linne, 1758), Lymnaea ovata (Draparnaud, 1805), L. stagnalis (Linne, 1758), L. corvus (Gmelin, 1791), Planorbis planorbis (Linne, 1758), and Planorbarius corneus (Linne, 1758), which were found to be infected by 9 species of trematode larvae. V. viviparus showed the highest rate of infection — 48.15% ± 6.8. The larvae of trematodes belonging to the families Echinostomatidae and Plagiorchidae predominate in the snails of the lake.     

Reproductive interference between Rana dalmatina and Rana temporaria affects reproductive success in natural populations

Experimental evidence suggests that reproductive interference between heterospecifics can seriously affect individual fitness; support from field studies for such an effect has, however, remained scarce. We studied reproductive interference in 25 natural breeding ponds in an area where two ranid frogs, Rana dalmatina and Rana temporaria, co-occur. The breeding seasons of the two species usually overlap and males of both species are often found in amplexus with heterospecific females, even though matings between heterospecifics produce no viable offspring. We estimated species abundance ratios based on the number of clutches laid and evaluated fertilization success. In ponds with low spatial complexity and a species abundance ratio biased towards R. temporaria, the average fertilization success of R. dalmatina eggs decreased, while this relationship was not detectable in spatially more complex ponds. Fertilization success of R. temporaria did not decrease with increasing relative numbers of heterospecifics. This asymmetry in fitness effects of reproductive interference may be attributed to R. temporaria males being more competitive in scramble competition for females than R. dalmatina males. Our study is among the first to demonstrate that in natural breeding populations of vertebrates interference among heterospecifics has the potential to substantially lower reproductive success at the population level, which may in turn affect population dynamics.     

Density dependent growth in adult brown frogs Rana arvalis and Rana temporaria – A field experiment

In species with complex life cycles, density regulation can operate on any of the stages. In frogs there are almost no studies of density effects on the performance of adult frogs in the terrestrial habitat. We therefore studied the effect of summer density on the growth rate of adult frogs during four years. Four 30 by 30 m plots in a moist meadow were used. In early summer, when settled after post-breeding migration, frogs (Rana arvalis and Rana temporaria that have a very similar ecology and potentially compete) were enclosed by erecting a fence around the plots. Frogs were captured, measured, marked and partly relocated to create two high density and two low density plots. In early autumn the frogs were again captured and their individual summer growth determined. Growth effects were evaluated in relation to two density measures: density by design (high/low manipulation), and actual (numerical) density. R. arvalis in plots with low density by design grew faster than those in high density plots. No such effect was found for R. temporaria. For none of the species was growth related to actual summer density, determined by the Lincoln index and including the density manipulation. The result suggests that R. arvalis initially settled according to an ideal free distribution and that density had a regulatory effect (mediated through growth). The fact that there were no density effects on R. temporaria (and a significant difference in its response to that of R. arvalis) suggests it is a superior competitor to R. arvalis during the terrestrial phase. There were no density effects on frog condition index, suggesting that the growth rate modifications may actually be an adaptive trait of R. arvalis. The study demonstrates that density regulation may be dependent on resources in frogs' summer habitat.     

Vasoactive intestinal polypeptide (VIP)-like immunoreactivity in anuran intestine

Summary Immunocytochemical and radioimmunological techniques with region specific antisera have been used to identify a vasoactive intestinal polypeptide-like material in the anuran intestine. Seven species of Anura were investigated: Bombina bombina, Alytes obstetricans, Rana temporaria, Rana esculenta, Hyla arborea, Hyla crepitans and Bufo bufo.In five of the species (A. obstetricans, R. temporaria, H. arborea, H. crepitans and B. bufo) vasoactive intestinal polypeptide-like immunoreactive mucosal endocrine cells and nerve fibres in all layers of the gut wall, were detected by both immunofluorescence and peroxidase-antiperoxidase methods. In the other two species, R. esculenta and B. bombina, no mucosal endocrine cells were detected although the vasoactive intestinal polypeptide-immunoreactive nerve fibres were plentiful.Radioimmunoassay showed the presence of significant amounts of vasoactive intestinal polypeptide-immunoreactivity in intestinal extracts from all species. The highest quantities were present in those anurans with both immunostained cells and nerves. Gel permeation chromatography showed that most of the vasoactive intestinal polypeptide-like peptide eluted in a position identical to that of natural mammalian (porcine) vasoactive intestinal polypeptide.The results indicate that a vasoactive intestinal polypeptide-like peptide is well represented in the Anura and that it is immunologically very similar to the mammalian peptide.Part of this work was presented at the European Society of Comparative Endocrinology, 1979; see Buchan et al. 1980a     

Temperature dependence of oocyte cleavage in three thermophilically different frogs of the genus Rana

The duration of the mitotic cycle (τ₀) at different temperatures at the time of synchronous cleavage division of oocytes of frogs Rana temporaria L., Rana arvalis Nilss. (= Rana terrestris Andr.) and Rana ridibunda Pall. has been studied. The first species is the least, and the last the most, thermophilic. In the semilogarithmic scale, the dependence of τ₀ on temperature is linear in the middle of the temperature scale, and is limited on both sides by inflexion points. This zone comprises the temperatures at which the larvae develop normally. For R. temporaria these temperatures are between 11–23°C. for R. arvalis 13–24°C and for R. ridibunda 16–27°C. At the midpoints of each zone (17.5, 18.5 and 21.5°C respectively) there is no difference in τ₀ between the species. At the same temperature, however, the mitotic cycle is the shortest for R. temporaria, is somewhat longer for R. arvalis and is the longest for R. ridibunda. The difference in the optimal temperature zones agree with different thermal stabilities of the cells and proteins of these species.     

A comparative study of physiological and structural changes at the myoneural junction in two species of frog after transection of the motor nerve

Summary Structural and functional behaviour of motor end-plates after transection of the motor nerve has been studied in two species of frog: Rana esculenta and Rana temporaria. The physiological results show that in both species there is a transient cessation of spontaneous activity followed by a resumption of miniature end-plate potentials (min. e.p.p.s.) after denervation. The characteristics of these potentials (frequency, distribution of amplitudes, time-course) are similar in the two species. However, some differences have been observed: Firstly, the period of silence lasts for 2–4 days in the case of Rana temporaria whereas it is prolonged to about 15 days in Rana esculenta. Secondly, the resumption of min. e.p.p.s. is gradual and after the 10th day of denervation remains constant in Rana temporaria. It is inconstant independent of the period of denervation in Rana esculenta. The morphological results show that the Schwann cell is constantly in contact with the post-synaptic membrane after about 6 days of denervation in both species. It is suggested that either the Schwann cell is capable of functioning for a limited period of time in Rana esculenta or is activated to produce min. e.p.p.s. only in certain cases.     

The use of artificial spawning substrates as media to support the reproduction of Eurasian perch in Lake Piediluco

Lake Piediluco is a naturally regulated lake used for hydroelectric purposes. Management of the lake and the eutrophic conditions of its waters have had a negative impact on fish populations, particularly that of Eurasian perch (Perca fluviatilis L.). In order to increase the natural reproduction of this species, an experimental program was undertaken from 1997 to 2003 to facilitate reproduction by providing artificial spawning substrates. In Lake Piediluco perch spawn from March to May. Egg ribbons ranged in length from 30 to 180 cm and contained from 12,740 to 205,466 eggs. The fraction of degenerated eggs ranged from 0.20% to 63.30% (average 10.20%). The results demonstrate the effectiveness of artificial substrates as suitable substrates for the reproduction of perch. Guest editors: R. L. Welcomme & G. Marmulla Hydropower, Flood Control and Water Abstraction: Implications for Fish and Fisheries     

Toxicity of an Anionic Detergent to the Spawn and Larvae of Anurans (Amphibia)

Several experiments were performed to test the toxicity of FIT, a washing-up and general cleaning liquid containing about 10 % sodiumalkanemonosulfonate (industrial name E 30) to spawn and larvae of anura. The LC₅₀ values for spawn were between 1.62 × 10⁻³ % (Rana dalmatina) and 4.97 × 10⁻³ % (Rana temporaria). The sensitivity of anuran larvae depends on the concentration and duration of exposure to the detergent as well as on the developmental and differentiation stage of the organisms. The toxicity of FIT is not due solely to the alkanesulfonate it contains, but results from the joint action of the surfactant, polyphosphates and solvents. The results indicate that detergents may be harmful to the amphibian fauna of Central Europe.