Different pathways are involved in the early development of the transient oral apparatus in anuran tadpoles (Anura: Leiuperidae)

The oral apparatus of anuran tadpoles is a unique structure composed of soft and keratinized parts surrounding the mouth. Among the many variations, a common oral apparatus involves a dorsal gap in the marginal papillae, keratinized jaw sheaths, and two upper and three lower rows of labial teeth. In Leiuperidae, besides this generalized morphology, four configurations are distinguished by the arrangement of the lower marginal papillae and the number of lower tooth rows. Study of the early oral ontogeny in 12 species representing these five configurations shows variations in the development of the lower marginal papillae and the third lower labial tooth row. Similar configurations may result from similar pathways (e.g. Physalaemus cuvieri group and Pseudopaludicola falcipes) or different pathways (e.g. generalized oral discs of Pleurodema and Physalaemus). Different oral configurations may result from overlapping trajectories ending at different stages (e.g. Physalaemus riograndensis and Ph. biligonigerus) or different trajectories (e.g. Ph. henselii and Ph. gracilis). Further studies are needed to interpret the role that heterochrony has played in evolutionary change within this family. The unsuspected variation occurring in this transient structure highlights its evolutionary potential and might be insightful in studies of anuran phylogenies that are largely based on adult characters. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 330–345.     

Ultrastructure of neuroendocrine cells in the lungs of three anuran species

Neuroendocrine cells in the lungs of three species of anurans, Bombina variegata, Bufo bufo and Bu. viridis, occur both as single cells and in the form of neuroepithelial bodies. Neuroendocrine cells are covered by ciliated cells or pneumocytes, which separate them from the lumen of the lung. Neuroepithelial bodies are dispersed in the apical part of the main septa of lungs of Bo. variegata and Bu. bufo or are situated on special protrusions of septa in the lungs of Bu. viridis. Neuroepithelial bodies are innervated by intraepithelial nerve endings of afferent and efferent types.     

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).     

Oral disc muscles of anuran tadpoles

The presence and configurations of the small, extrinsic muscle fibers of the M. mandibulolabialis superior in the upper labium and the M. inferior in the lower labium are described for 24 species in 9 families. Although several species possess only the M. inferior, or both the M. inferior and superior, none has only the M. superior. Some species without tooth rows possess the M. inferior, and microhylid tadpoles have a single-stranded muscle different than the M. mandibulolabialis in each oral flap. The extrinsic oral muscles insert near the bottom of the inter-row valleys behind each tooth ridge and the bases of marginal papillae lateral to the ends of tooth rows. Muscles do not extend to marginal papillae beyond the most distal row of teeth. Muscle contraction is presumed to rotate the teeth distally so that they contact the substrate at the proper working angle. This action probably occurs at full extension of the oral disc just before disc closure starts.     

Sequence of ontogenetic development and atrophy of the oral apparatus of six anuran tadpoles

The development of the oral structures of six species of anuran tadpoles with four different types of mouth parts and the metamorphic atrophy of these structures in two species with different mouth parts are described. The oral labia of typical tadpoles, oral flaps of microhylids, and lateral oral folds of Rhinophrynus are assumed to be homologous. We also suggest that the barbels of the tadpoles of Rhinophrynus are homologs of the marginal papillae of species with an oral disc. Developmental patterns and sequences of the oral structures of all tadpoles examined follow a common pattern: stomodeal invagination, oral pad development, jaw sheeth delimitation, tooth row ridge development, jaw sheath keratinization, and labial tooth keratinization. Developmental patterns remain constant, while interspecific differences are apparent because of truncations of ontogeny at specific stages. Metamorphic atrophy of oral structures occurs roughly in the reverse order of development, although the procedure is rapid and more haphazard than development.     

中国林蛙蝌蚪的口器发育

采用扫描电镜和组织学技术观察了中国林蛙(Rana chensinensis)蝌蚪发育过程中口器外部形态结构的变化,以及中国林蛙蝌蚪口器内部结构特征.结果表明,在口器发育的初期,角质颌最先出现,接着出现唇乳突以及唇齿;在变态高峰期(G4l～G42),口器结构如唇齿、角质颌和唇乳突则是按以下顺序消失的,即唇齿最先消失,其次...     

Living in fast‐flowing water: Morphology of the gastromyzophorous tadpole of the bufonid Rhinella quechua (R. veraguensis group)

We describe the bufonid gastromyzophorous tadpoles of Rhinella quechua from montane forest streams in Bolivia. Specimens were cleared and stained, and the external morphology, buccopharyngeal structures, and the musculoskeletal system were studied. These tadpoles show a combination of some traits common in Rhinella larvae (e.g., emarginate oral disc with large ventral gap in the marginal papillae, labial tooth row formula 2/3, prenarial ridge, two infralabial papillae, quadratoorbital commissure present, larval otic process absent, mm. mandibulolabialis superior, interhyoideus posterior, and diaphragmatopraecordialis absent, m. subarcualis rectus I composed of three slips), some traits apparently exclusive for the described species of the R. veraguensis group (e.g., second anterior labial tooth row complete, lingual papillae absent, adrostral cartilages present), and some traits that are shared with other gastromyzophorous tadpoles (e.g., enlarged oral disc, short and wide articular process of the palatoquadrate, several muscles inserting on the abdominal sucker). In the context of the substantial taxonomic and nomenclatural changes that the former genus Bufo has undergone, and despite the conspicuous morphological differences related to the presence of an abdominal sucker, the larval morphology of R. quechua supports including it in the genus Rhinella and placing it close to species of the R. veraguensis assemblage. J. Morphol., 2009. © 2009 Wiley‐Liss, Inc.     

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.     

First report of a green toad (Bufo viridis sensu lato) in the Early Pleistocene of Spain: Palaeobiogeographical and palaeoecological implications

For the first time unequivocal fossil remains of a green toad (Bufo viridis s.l.) are described in the Iberian Peninsula. The fossils come from the Cueva Victoria site, a late Early Pleistocene (ca. 1.1–1.2 Ma) karstic filling in semi-arid southeastern Spain (Murcia region). By extension, other remains from two other Early Pleistocene Spanish localities, Barranco León D (ca. 1.3 Ma) and Almenara-Casablanca 3 (ca. 1.1 Ma), are cautiously attributed to the group B. viridis. The B. viridis group was previously reported with some uncertainty to the west of its current distribution area in Western Europe (Spain and France) in the Pliocene (Bufo cf. viridis) and less probably in the Early Miocene (Bufo aff. viridis). Since no osteological differences have been established between the recently described extant species of B. viridis s.l. (e.g. Bufo balearicus, Bufo siculus, Bufo boulengeri, B. viridis sensu stricto and Bufo variabilis) no precise palaeobiogeographical relationships can be drawn for the Spanish fossils. However, the occurrence of a third species of bufonid toad during the Pleistocene in the South of the Iberian Peninsula raises some interesting ecological questions in relation to the local disappearance of the green toad, which can be hypothetically linked to the intensification of the Pleistocene glacial/interglacial climate dynamic or to probable competition with another toad, Bufo calamita.     

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.     

How can ontogeny help us to understand the morphology of anuran pectoral girdle?

The ontogenetic development of the pectoral girdle in seven anuran species (Xenopus laevis, Discoglossus pictus, Bombina bombina, Bombina variegata, Pelobates fuscus, Bufo bufo and Rana dalmatina) was studied using cleared and stained specimens. The epicoracoid cartilage was found to develop in two different ways resulting in an arciferal or firmisternal type of the pectoral girdle. In the arciferal one, the epicoracoid originates from a medial prolongation of the procoracoid cartilage and broadly overlaps its counter part during further development. In the firmisternal pectoral girdle, the epicoracoids are formed by the widened cartilaginous medial edges of the coracoids that fuse together along the midline. Polarization of ontogenetic characters shows, that omosternum evolves inside Anura, and the type of sternum occurring in basal Anura seems to be an apomorphy of all Batrachia. The sternal elements have a single or paired rudiment, their development is connected with M. rectus abdominis or a zonal area and they remain cartilaginous or ossify during postmetamorphic development. An occurrence of omosternum in Barbourula busuangensis was described for the first time.     

浙江丽水中华大蟾蜍和黑眶蟾蜍蝌蚪对水位变化的表型响应

全球气候变暖引发栖息地干涸将对生活在水中的无尾类幼体提出了挑战。通过浙江丽水中华大蟾蜍(Bufo gargarizans)和黑眶蟾蜍(Duttaphrynus melanosticus)蝌蚪在实验条件下对不同水位变化的表型响应,检测表型可塑性的遗传性和环境近因性影响。结果表明,水位变化对中华大蟾蜍蝌蚪早期发育历期、头宽和体重影响不显著,对体长影响显著,其中逐减水位最大、恒低水位最小,慢波、恒高与快波、逐增水位依次减少;水位变化对黑眶蟾蜍蝌蚪早期发育历期、体长、头宽和体重影响均显著;发育历期以恒高水位最大,恒低水位最小;体长以逐减水位最大,恒低、快波和慢波水位显著偏小,逐增和快波水位居中;头宽以恒低水位最小,逐增水位居中,其余较大;体重以恒低水位最小、恒高水位最大,其余居中。水位变化对中华大蟾蜍蝌蚪的变态时间、体长、头宽和体重影响均不显著;水位变化对黑眶蟾蜍蝌蚪的变态时间、体长和体重影响均显著,对头宽影响不显著;恒低水位的变态时间最长,恒高水位的变态时间最短,其他水位变化之间差异不显著;恒高水位的体长最大,恒低和快波水位最小,其他居中;逐增和快波水位的体重最大,恒低水位最小。研究结果表明,繁殖季节不同的中华大蟾蜍和黑眶蟾蜍蝌蚪响应水位变化的表型可塑性差异显著,长期在容易发生干旱和水位变化的冬季繁殖的中华大蟾蜍蝌蚪的表型可塑性低,在雨水充沛的春季繁殖的黑眶蟾蜍蝌蚪的表型可塑性高,表现出表型可塑性的种间差异和遗传性;在早期发育过程中,两种蝌蚪体长的共同的表型变异与缺乏遗传基础的环境近因性影响有关;黑眶蟾蜍蝌蚪对低水位或水位下降作出减速分化的消极响应,响应程度与环境信号的强弱直接相关。     

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.     

Growth and behaviour of six species of amphibian larvae in a winter pond in Israel

The effect of different ecological niches on growth and behaviour of larvae of four frogs (Bufo viridis, Hyla arborea, Pelobates syriacus, Rana ridibunda), and two salamanders (Salamandra salamandra and Triturus vittatus) found in a winter pond was studied. S. salamandra, T. vittatus, R. ridibunda and B. viridis were found most of the time on the bottom of the pond. However, H. arborea tadpoles were found throughout the pond and were usually sedentary, as compared with P. syriacus which moved up and down constantly. S. salamandra, T. vittatus, R. ridibunda and R viridis tadpoles from the bottom of the pond grew faster than the tadpoles from the surface of the pond. However, tadpoles of H. arborea and P. syriacus growing at the bottom or on the surface developed at similar rates.The invertebrate biomass increases during the summer and was higher at the bottom of the pond than at the surface. However the amount of chlorophyll a was about the same at the surface and at the bottom of the pond. S. salamandra and T. vittatus tadpoles feed on various types of of invertebrates, R. ridibunda and H. arborea and B. viridis tadpoles feed on vascular plants and algae, and P. syriacus tadpoles feed on both invertebrates and plants.     

On the biology of Amphibia in the ecosystem of Lake Glubokoe

In Lake Glubokoe, the Amphibia are represented by five species. Two numerous species, Rana temporaria L. and Bufo bufo L., spawn in the lake and appear to be a significant component of its ecosystem. Thus, in autumn R. temporaria is a common food for predatory fish (pike, perch). During their wintering R. temporaria participate in the consumption of oxygen from the water. They may be a source of dangerous infections. Their tadpoles and those of B. bufo feed in the lake and annually remove substances from it. Adult R. temporaria in autumn, and the young of the year of R. temporaria and of B. bufo influence the population numbers of small littoral invertebrates. Several dozen pairs of Rana terrestris Andrzejw are observed to spawn regularly. Rana esculenta L. and Rana ridibunda Pallas are scarce. The influence of introducing a nature preserve on the numbers of Amphibia in Lake Glubokoe is discussed.     

凹耳蛙的分类地位及蛙亚科一新属

对凹耳蛙模式标本产地的蛙类及蝌蚪进行了调查,并对蝌蚪进行了人工饲养和生长发育到完成变态的观察,结果发现一种没有腹吸盘的小型蝌蚪是凹耳蛙的蝌蚪.由此证明凹耳蛙不具有湍蛙型蝌蚪,应不属于湍蛙属蛙类.对凹耳蛙成体和蝌蚪特征的分析表明其与蛙亚科其他属及所辖种不同,建议另立新属.     

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     

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.     

Karyotyp und DNS-Gehalt von Bufo bufo,B. viridis,B. bufo × B. viridis und B. calamita (Amphibia,Anura)

The karyotypes in spermatogonial and leukocyte metaphases of the toads Bufo bufo, B. viridis and B. calamita (all 2n=22) were analysed and the DNA content of colchicine treated and Feulgen stained spermatogonial metaphase chromosomes measured microspectrophotometrically. The toad species possess similar karyotypes, but the chromosomes of B. bufo are somewhat longer than the chromosomes of B. viridis and B. calamita. All chromosomes of B. bufo contain significantly more than, but in no case twice as much DNA as their homologues in the other two species. Eight chromosomes of B. bufo contain 30–40%, three about 50% more DNA than their homologues in B. viridis. Exactly the same DNA-differences between both sets of chromosomes were found in B. bufo × B. viridis hybrids. Significant differences in the DNA amount of B. viridis and B. calamita exist only between the large chromosomes of these species. The ratio of the total DNA amount of the genomes in the three species is 1.49∶1.07∶1. These DNA-differences between the three toad species are confirmed by microspectrophotometric DNA measurements of their erythrocyte nuclei. It is supposed that these interspecific differences in DNA content of the toads are not a consequence of differential polyteny but are caused during the evolution process by local increase in DNA in all chromosomes of B. bufo and in the large chromosomes of B. viridis.     

Morphology of the oral disc of Bufo bufo (Salientia: Bufonidae) tadpoles

The fully developed oral disc of the tadpole of Bufo bufo consists of dorsal and ventral labia bearing, respectively, two and three ridges bearing numerous horny denticles, a horny beak provided with jaw sheath serrations, and large lateral papillae that are borne by two cutaneous plicae. As development progresses toward metamorphosis, these structures gradually regress until they disappear. Each cusped clavate labial denticle adheres, by means of a thin peduncle, to a similar labial denticle fixed in the lip and formed by a group of three or four cells that keratinize gradually and thus present remarkable differences in their morphology. Once all the cells of a group have been converted into horny tissue, the denticle sheds and is replaced by the underlying one. The beak serrations also are horny structures; each consists of a columnar band of cells which undergoes a gradual keratinization. The horny cells that detach themselves at intervals, being replaced by those of the underlying anlagen. The labial denticles and the beak serrations keratinize in two distinct ways. In the former, the desmosomal filaments appear to play an important role whereas, in the latter, the keratin seems to be synthesized “ex novo” by the ribosomes.