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.     

A survey of shape variation in keratinized labial teeth of anuran larvae as related to phylogeny and ecology

Labial teeth of anuran tadpoles are keratinized structures derived from the activity of a single epidermal cell of the oral labia; they are not homologous with adult anuran teeth, nor with teeth of other vertebrates. The present study comprises a first approach for studying labial tooth shape variation that will be useful for future studies of comparative development and the functional mechanics of feeding structures. We examined interspecific shape variations in the labial teeth of anuran tadpoles and searched for correlations of these variations with ecomorphological guilds and phylogeny. Species ordination shows that important variations at various taxonomic levels are related mainly to the general curvature of the tooth axis, the angle between the labial tooth base and tip, head length and curvature, and sheath width. The teeth of most basal taxa are broad‐based and curved, although some broad‐based teeth also characterize some phthanobatrachian species. Teeth of hyloids and ranoids differ in the oral angle, overall curvature, and sheath width. A phylogenetically independent ecomorphological effect is significant only for lotic suctorial and gastromyzophorous guilds; teeth in these forms have short, thick and curved heads, wide sheaths, and generally acute oral angles. The lack of a significant correlation between labial tooth shape and trophic guilds suggests that labial tooth harvesting ability has a wide latitude that could be particularly functional only under specific circumstances. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 609–625.     

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.     

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.     

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.     

Evolutionary significance of oral morphology in the carnivorous tadpoles of tiger frogs, genus Hoplobatrachus (Ranidae)

We describe the larval stages of three species of the Asian-African tiger frogs Hoplobatrachus chinensis, H. occipitalis and H. tigerinus . The tadpoles of all three species are very similar, with peculiar oral features: (1) double rows of needle-like labial teeth, (2) strong emarginations on the large jaw sheaths and (3) keratinized spurs on the buccal floor. Characters 1 and 2 (and perhaps 3) are probably related to the carnivorous habits of these tadpoles. A molecular phylogeny based on 2430 base pairs of two nuclear and four mitochondrial genes corroborated monophyly of Asian and African Hoplobatrachus, and identified Euphlyctis as their sister group. Tadpoles of the latter genus lack buccal spurs and double labial tooth rows but share large jaw sheaths, the upper with a medial projection. Therefore, the common ancestor of Euphlyctis and Hoplobatrachus probably was also characterized by this state, and may have been facultatively carnivorous. Further carnivorous specializations in Hoplobatrachus could explain why tiger frogs have been so successful in populating arid environments where ponds are at high risk of desiccation. Larval morphology may prove to be the key innovation which enabled them to disperse, in the Late Cenozoic, into their current very wide distribution area in Asia and Africa.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 171–181.     

五种蝌蚪口器及舌鳃骨的结构比较

采用体视显微镜和骨骼双染色法对5种不同栖息环境的无尾两栖动物蝌蚪的口器和舌鳃骨的形态结构特征进行了观察。5种蝌蚪口器由唇齿行、唇乳突和角质颌等组成。舌鳃骨是由关联骨Ⅰ、关联骨Ⅱ、角舌骨、舌鳃骨盘和角鳃骨等骨骼组成。蝌蚪的梅氏软骨若较发达,其摄食方式可能为刮食;蝌蚪的舌鳃骨发达,其摄食方式则可能为滤食。角质颌、唇齿以及角鳃骨上鳃耙的出现显著增强了蝌蚪主动摄食能力和对食物与非食物的主动选择性。     

Variation of apomorphic characters in stream-dwelling tadpoles of the bufonid genus Ansonia (Amphibia: Anura)

Six larval forms of the bufonid genus Ansonia from Borneo share the following synapomorphies: cup-like, ventral oral disc; an expanded post-dental portion of the lower lip, which has a papillate margin; upper keratinized jaw sheath divided; body markedly flattened ventrally; eyes set relatively far back on the body. All of these tadpoles live on the bottom in strong currents, except for larval Ansonia leptopus , which lives in drifts of dead leaves that accumulate in eddies within streams. These larval forms differ among themselves in body shape, development of inframarginal papillae on the lower lip, size of the gap between the keratinized parts of the upper jaw sheath, width of the post-dental portion of the lower lip, relative lengths of upper and lower rows of labial teeth, and arrangement of the gut coils. One form has an abdominal sucker. Changes in these characters are not correlated; the derived condition in one character is not always associated with the derived state in another. Consequently, these tadpoles cannot be arranged in a simple morphocline from least to most derived, again with the exception of A. leptopus , which is the least modified in all respects. Although tadpoles of Ansonia resemble those of the neotropical bufonid genus Atelopus in general specialization for benthic life in flowing water, they differ from that group in body form and details of the oral disc.     

Apoptotic cell death in the central nervous system of Bufo arenarum tadpoles induced by cypermethrin

Tadpoles of the toad Bufo arenarum treated with cypermethrin (CY) at concentrations above 39 μg CY/L showed dose-dependent apoptotic cell death in immature cells of the central nervous system as demonstrated by morphometric analysis, the TUNEL method, and DNA fragmentation assay. Light-and electron-microscopic studies showed structural alterations in the intermediate and marginal layers of the brain. Immature cerebral tissue showed cellular shrinkage, nuclear fragmentation and increase of intercellular spaces. In this study we demonstrated high toxicity of CY to larval stages of Bufo arenarum. Our results show that doses lower than those used in routine insecticide applications can cause massive apoptosis in the immature cells of the central nervous system. These results coincide with our previous studies in Physalaemus biligonigerus, confirming the severe toxic effects of CY to the central nervous system of anuran species from Argentina. This may increase the mortality index in wild animals and contribute to the loss of biodiversity in our agroecosystems. We postulate that CY induces apoptosis in central nervous system cells of Bufo arenarum tadpoles by specific neurotoxic mechanisms.     

Cranial anatomy of Shunosaurus, a basal sauropod dinosaur from the Middle Jurassic of China

Shunosaurus, from the Middle Jurassic of China, is probably the best‐known basal sauropod and is represented by several complete skeletons. It is unique among sauropods in having a small, bony club at the end of its tail. New skull material provides critical information about its anatomy, brain morphology, tooth replacement pattern, feeding habits and phylogenetic relationships. The skull is akinetic and monimostylic. The brain is relatively small, narrow and primitively designed. The tooth replacement pattern exhibits back to front replacement waves in alternating tooth position. The teeth are spatulate, stout and show well‐developed wear facets indicative of coarser plant food. Upper and lower tooth rows interdigitate and shear past each other. Tooth morphology, skull architecture, and neck posture indicate that Shunosaurus was adapted to ground feeding or low browsing. Shunosaurus exhibits the following cranial autapomorphies: emargination of the ventral margin of the jugal/quadratojugal bar behind the tooth row; postorbital contains a lateral pit; vomers do not participate in the formation of the choanae; pterygoid is extremely slender and small with a dorsal fossa; quadrate ramus of the pterygoid is forked; quadratojugal participates in the jaw articulation; tooth morphology is a combination of cylindrical and spatulate form; basipterygoid process is not wrapped by the caudal process of the pterygoid; trochlear nerve has two exits; occlusal level of the maxillary tooth row is convex downward, whereas that of the dentary is concave upward, acting like a pair of garden shears; dentary tooth count is 25 or more; and the replacing teeth invade the labial side of the functional teeth. Cranial characters among the basal sauropods are reviewed. As Shunosaurus is the earliest sauropod for which cranial remains are known, it occupies an important position phylogenetically, showing the modification of skull morphology from the prosauropod condition. Although the skull synapomorphies of Sauropoda are unknown at present, 27 cranial synapomorphies are known for the clade Eusauropoda. © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society, 2002, 136 , 145?169.     

Addendum to the Morphology of Steinernema scapterisci

This paper provides additional information on the morphology of Steinernema scapterisci. For females, the amphids are shown for the first time, and clearer scanning electron microscope (SEM) photographs of the six labial and four cephalic papillae are presented. For males, six labial and four cephalic papillae, one or two pairs of additional genital papillae, and the death shape are shown. For infective juveniles, six labial and four cephalic papillae and an elevated oral disc are shown.     

Plasticity and constraints on feeding kinematics in anuran larvae.

Tadpoles of the majority of anuran species have tiny, anatomically complex mouths. In most species the larval jaws are keratinized sheaths (beaks) overlying infrarostral cartilages. Surrounding the beak is a flexible oral disc and transverse rows of small, keratinized denticles. We used high-speed videography (250, 500 and 1000 frames per second) of Rana catesbeiana tadpoles to observe the kinematics of these mouthparts in feeding and breathing. Tadpoles can protract and retract their jaws as well as make them wider and narrower with each gape cycle. We demonstrate that during air-breathing, movement of the oral disc helps surfacing tadpoles to capture air quickly by preventing water from coming into the mouth. For our feeding study, we observed tadpoles as they grazed on both clean and algal covered glass surfaces. As the jaws close, the lower beak narrows to a greater degree when it encounters resistance. The denticle rows are used to both anchor the mouth and rasp surfaces during feeding. The hyperkinetic mouth parts of tadpoles permit grazing on non-planar surfaces of variable resistance. A trade-off in having such mobile jaws is loss of stability; no generalized tadpoles can generate great forces with their jaws, which would be necessary to subdue and dismember large tough prey. The feeding system of tadpoles is built out of soft tissues (such as cartilage and keratin) that can be shed (the keratinized sheaths) or remodeled (the underlying infrarostral cartilage) quickly, thus facilitating metamorphosis.     

Agropastoral activities increase fluctuating asymmetry in tadpoles of two neotropical anuran species

Agriculture and pasture activities are the main drivers for habitat reduction, directly affecting amphibian assemblages. In the Cerrado, the progress of agricultural and pasture areas negatively affects aquatic environments and their organisms, once the suppression of marginal vegetation reduces the natural protection against allochtone stressors. One possible way to measure the stress level is the Fluctuating Asymmetry (FA), which is calculated based on the deviations in the development of bilateral morphological traits of the organism. Herein, we evaluated whether environments with a higher degree of agropastoral influence and reduced marginal vegetation can increase FA levels in tadpoles of two common and widely distributed anuran species in the Cerrado (Physalaemus cuvieri and Scinax fuscomarginatus). We sampled and classified water bodies according to the percentage of agropastoral land use and marginal vegetation, and measured four morphological traits of tadpoles to evaluate the degree of FA. We found that in environments with intensive agropastoral land use, tadpoles of P. cuvieri and S. fuscomarginatus had higher FA in nostril‐snout distance (NSD). In environments with reduced marginal vegetation, tadpoles of S. fuscomarginatus had higher FA in eye width (EW), but no effect was detected for tadpoles of P. cuvieri. These morphological traits (i.e. nostrils and eyes) are associated to individual fitness of tadpoles. Thus, these developmental deviations can affect species fitness and population homeostasis over time, contributing to generate stochastic population dynamics and increasing the vulnerability of native species to local extinctions. The use of FA as a tool to measure environmental impact on species with potential to be used as bioindicators can contribute to generate and test hypothesis when integrated with long‐term studies.     

Stephanofilaria thelazioides n. sp. (Nematoda: Filariidae) from a hippopotamus and its affinities with the species parasitic in the African black rhinoceros

Stephanofilaria thelazioides n. sp. (Filarioidea: Filariidae: Stephanofilariinae) is described from a hippopotamus Hippopotamus amphibius. This nematode is close to S. dinniki Round, 1964, a parasite of the black rhinoceros Diceros bicornis in Africa, but differs from it in the number of cuticular spines surrounding the mouth, the arrangement of the cloacal papillae and the measurements of the spicules, gubernaculum and microfilariae. Species of the genus Stephanofilaria possess spines on the head which have been derived by modification of the sensory papillae. S. thelazioides is the most primitive species of the genus and has the least modified arrangement of these papillae, with six bifid internal labial spines, four bifid external labial spines and four cephalic papillae. The genus appears to have diversified in various mammals which have in common a thick skin, such as rhinoceroses, elephants, buffaloes and now the hippopotamus. It appears to have become adapted secondarily to domestic bovines, initially in Asia and subsequently in North America.     

New data on morphology of the African scraping feeder Varicorhinus beso (Osteichthyes: Cyprinidae) with the special reference to specialized traits

Ten samples of Varicorhinus beso, the type species of the genus from East Africa were studied using a large set of morphological characters to evaluate the degree of specialization. It is found that V. beso is highly specialized scraping feeder that possesses many specialized trails: short and wide head, wide inferior mouth with keratinized scraping edge on the lower jaw, modified bones of mouth apparatus, high number of the gill rakers with papillae between rows of the rakers, strongly elongated intestine, and one pair of minute barbels (contrary to scraping trophic forms of barbs possessing two pairs). Despite specialization, V. beso is morphologically rather variable, and some divergence was detected between populations studied.     

Plate-like permanent dental laminae of upper jaw dentition in adult gobiid fish, Sicyopterus japonicus

Sicyopterus japonicus (Teleostei, Gobiidae) possesses a unique upper jaw dentition different from that known for any other teleosts. In the adults, many (up to 30) replacement teeth, from initiation to attachment, are arranged orderly in a semicircular-like strand within a capsule of connective tissue on the labial side of each premaxillary bone. We have applied histological, ultrastructural, and three-dimensional imaging from serial sections to obtain insights into the distribution and morphological features of the dental lamina in the upper jaw dentition of adult S. japonicus. The adult fish has numerous permanent dental laminae, each of which is an infolding of the oral epithelium at the labial side of the functional tooth and forms a thin plate-like structure with a wavy contour. All replacement teeth of a semicircular-like strand are connected to the plate-like dental lamina by the outer dental epithelium and form a tooth family; neighboring tooth families are completely separated from each other. The new tooth germ directly buds off from the ventro-labial margin of the dental lamina, whereas no distinct free end of the dental lamina is present, even adjacent to this region. Cell proliferation concentrated at the ventro-labial margin of the dental lamina suggests that this region is the site for repeated tooth initiation. During tooth development, the replacement tooth migrates along a semicircular-like strand and eventually erupts through the dental lamina into the oral epithelium at the labial side of the functional tooth. This unique thin plate-like permanent dental lamina and the semicircular-like strand of replacement teeth in the upper jaw dentition of adult S. japonicus probably evolved as a dental adaptation related to the rapid replacement of teeth dictated by the specialized feeding habit of this algae-scraping fish.     

Development of the nidicolous tadpoles of Eupsophus emiliopugini (Anura: Cycloramphidae) until metamorphosis,with comments on systematic relationships of the species and its endotrophic developmental mode

Vera Candioti, M.F., Nuñez, J.J. and Úbeda, C. 2011. Development of the nidicolous tadpoles of Eupsophus emiliopugini (Anura: Cycloramphidae) until metamorphosis, with comments on systematic relationships of the species and its endotrophic developmental mode. —Acta Zoologica (Stockholm) 92 : 27–45. Species of Eupsophus are unique within Alsodinae in having nidicolous tadpoles. They are characterized by traits typical of generalized exotrophic (e.g., oral disc and spiracular tube) and endotrophic larvae (e.g., scant pigmentation and large hind limbs). The larval morphology and development of E. emiliopugini, including external, buccal, and musculoskeletal features, is described herein. Like the larvae of other alsodines, these larvae have four lingual and four infralabial papillae, quadratoethmoid process, and an m. rectus cervicis with a double insertion. Among the traits exclusive to the genus are: the absence of the pseudopterygoid process and quadrato‐orbital commissure; presence of the m. subarcualis rectus I with two slips; and presence of the m. subarcualis rectus II–IV inserting on Ceratobranchial II. The development and metamorphosis of Eupsophus include some characters that develop later (e.g., degeneration of mouthparts and chondrocranium with minimum calcification), characters that develop earlier (e.g., hind‐limb appearance and jaw and suspensorium ossification), and characters that develop at the same time (e.g., most external features and cranial muscles) than in most exotrophic species. Some distinctive characters (third lower labial ridge absent, general configuration of the hyobranchial skeleton, skeletal development with retention of larval traits) resemble those of other endotrophic species, and the precocious ossification of jaws and suspensorium is shared with several direct‐developing species among recent amphibians.