This paper presents larval evidence and evaluates its contribution to the discussion of frog phylogeny; 136 larval characters, 6 reproductive biology characters, and 14 adult morphology characters were scored for 81 frog and 4 caudate species. More than 90% of the data matrix entries represent original data derived from personal direct examination of specimens. Some larval characters are described for the first time and many others have not been assessed for specific taxa or in a broad phylogenetic context before. Homoplasy appears common in this and other amphibian morphological data sets. The data supported and confirmed various wellknown clades, among others the Anura, Bufonidae, Ceratophryinae, Discoglossidae, Dendrobatidae, Hyperoliidae, Microhylidae, South American microhylids, Phyllomedusinae, Pseudinae, Pipoidea, Pipidae, and Scoptanura. The Ascaphidae was sister group to all other anurans and the Pipoidea was placed more basally than in some previous analyses. The Eurasian pelobatids formed a clade, whereas Spea and Pelodytes did not group robustly with them. Pelobatoid frogs emerged as a paraphyletic ''transitional'' assemblage including Heleophryne. The resolution of basal neobatrachian splits remained labile, although some subclades within the Neobatrachia were robustly supported. The ''Hylidae'' was paraphyletic, and hyline species were paraphyletic with respect to the Pseudinae. Hemisus clearly was in a clade with the Hyperoliidae and is proposed to be included in that family. Scaphiophryne was confirmed as basal taxon within the Microhylidae. Compared to the larval stages of the most recent common ancestor of anurans, members of the Scoptanura (microhylids except scaphiophrynines) have accumulated the highest number of apomorphic character states in anuran evolution.
The larval neurocranium and visceral arches of seven dendrobatid species representing four genera are described, based on cleared-and-stained and serially sectioned specimens. A variety of characters is shared by all seven species. Larval features do not substantiate the assumption of close ranoid affinities of the Dendrobatidae. Instead dendrobatid larvae share features such as the special quadripartite cartilago suprarostralis, the lack of the larval processus oticus, the presence of three foramina acustica, and the lack of a foramen perilymphaticum accessorius with many bufonoid larvae. The first of these characters is unique to bufonids, hylids, dendrobatids, and some New World leptodactylids; the other characters also occur in pelobatids and are presumably plesiomorphic for the Neobatrachia. The free proximal ends of Ceratobranchialia II and III are an autapomorphy of the Dendrobatidae supporting the monophyly of the family. Some features of the cranium are paedomorphic: low cartilago orbitalis, lack of connection between cartilage orbitalis and otic capsule (most species), and vestigal taeniae tecti. New anatomical terms are introduced.
Summary A phylogenetic parsimony analysis of fifty‐four larval characters of Myxophaga (excluding Lepiceridae) resulted in two minimal length cladograms. The monophyly of Torridincolidae, Hydroscaphidae and Microsporidae is supported by several autapomorphies: miniaturization, flattened body with laterally extended tergites, broadened head, scale‐like surface structures, broad tentorial bridge, disc‐shaped labral sensilla, spiracular gills and pupation in the last larval exuviae. Hydroscaphidae are the sister group of Microsporidae. Larvae of both families are characterized by semi‐entognathous mouthparts, tergites with posterior rows of lancet‐shaped setae, claws with flattened basal spines and balloon‐shaped spiracular gills. The monophyly of all families is supported by autapomorphies. Torridincolidae excluding Delevea is defined as a monophylum by four derived character states: body ovoid, thorax semicircular and as long as abdomen, labral sensilla fused and abdominal sternite IX distinctly reduced and triangular. The monophyly of Torridincolinae (sensu Endrödy‐Younga 1997b) is supported by two autapomorphies. The proposed branching pattern suggests that the early representatives of Myxophaga (excluding Lepiceridae) were living in aquatic conditions with a preference for hygropetric habitats. The tendency to live on rocks in running water and miniaturization have played an important role in myxophagan evolution.
Suctorial anuran larvae are highly specialized for living in fast‐flowing waters, using their oral disks as adhesive organs to attach to the substrate. The cranial musculoskeletal structure of suctorial larvae of Litoria nannotis, L. rheocola, and Nyctimystes dayi (Hylidae: Pelodryadinae) were compared with congenerics with pond‐type larvae (L. caerulea, L. genimaculata, L. xanthomera). Data from two other neobatrachian species with suctorial larvae (Boophis sp., Hyla armata) as well as published descriptions were taken into account. Suctorial tadpoles evolved several times independently in the Neobatrachia and share various features, irrespective of their phylogenetic position. These include the following. Cornua trabeculae are expanded anteriorly and sometimes fused. The lower jaws are robust. The greatest width of the skull is at the level of the jaw articulation. The upper jaw cartilages are partially or fully fused. The palatoquadrate is robust and connected to the skull by a wide commissura quadratocranialis anterior, processus oticus, processus basalis (in some species), and processus ascendens (vestigial or absent in some species). A processus ventralis quadrati provides an extended area of origin for the m. orbitohyoideus. The m. rectus abdominis inserts far anterior and acts on the cranium. The insertion of the epaxial musculature is shifted anteriorly to the anterior parts of the otic capsule. The mm. diaphragmatobranchialis and rectus cervicis cross at their origins. The origin of the m. levator mandibulae anterior has shifted posteriorly. The branchial basket is relatively small and the ceratohyal area is large. Multiple convergent evolution of these features suggests that they may be causally associated with the suctorial mode of larval life. Aside from these characters, however, the suctorial and pond‐type neobatrachian species are remarkably similar in their jaw musculature and hyobranchial musculoskeletal composition. In some features, Ascaphus truei differs significantly from the neobatrachian suctorial species, indicating the influence of the historically distant separation of the two taxa. A novel modification of the upper jaw abduction mechanism has evolved in L. nannotis, L. rheocola, and N. dayi. It involves an adrostral cartilage as a pushing‐rod element. This mechanism and unique structural similarities of the cartilago labialis superior gives support to the preliminary assumption that the nannotis species group is more closely related to the suctorial Nyctimystes dayi than it is to other Litoria species with pond‐type larvae. Suctorial larvae presumably were present in the most recent common ancestor of the Litoria nannotis group and Nyctimystes dayi. J. Morphol. 238:109–141, 1998. © 1998 Wiley‐Liss, Inc.
The speed of larval development of the South African bullfrog, Pyxicephalus adspersus, is exceptional among living frogs. Embryonic development, larval phase, and metamorphosis can be completed in 17 days at a temperature of 29°C. The metamorphosis only takes 5 days. The present study shows that, despite the unusually short larval phase in P. adspersus, the state of skeletal differentiation reached at the end of metamorphosis is similar to that of other frog species. There is no shift of cranial bone formation postmetamorphosis as could have been expected and is known from other species. The majority of compared species are particularly similar in the sequence of bone formation in the postcranial skeleton. However, there are clear differences among species in the timing of these events relative to the larval growth trajectory, absolute time, and certain developmental markers, such as external limb differentiation. For example, skeletogenesis and externally visible limb differentiation are only loosely integrated. Interspecific comparisons show that, in P. adspersus, the early onset of skeletal ossification is an unusual feature among frogs. Freshly metamorphosed froglets of P. adspersus are already distinct from comparable stages of other species in having strong jaws, fang-like teeth, and a squamosal-maxilla contact. The latter stabilizes the maxillary arcade and the suspensorium and might relate to the ability to catch and swallow very large vigorous prey, such as siblings, shortly after metamorphosis. The presence of a complete set of dermatocranial elements and postmetamorphic ossification of only the sphenethmoid and operculum are considered plesiomorphic features, whereas the much less completely ossified skulls of metamorphosed froglets, particularly in Bufo and Hamptophryne, are likely apomorphic developmental traits within the Anura.& b d y :Larval and metamorphic skeletal development in the fast-developing frog Pyxicephalus adspersus (Anura, Ranidae)
Anurans (frogs and toads) are unique among land vertebrates in possessing a free-living larval stage that, parallel to adult frogs, diversified into an impressive range of ecomorphs. The tempo and mode at which tadpole morphology evolved through anuran history as well as its relationship to lineage diversification remain elusive. We used a molecular phylogenetic framework to examine patterns of morphological evolution in tadpoles in light of observed episodes of accelerated lineage diversification. Our reconstructions show that the expansion of tadpole morphospace during the basal anuran radiation in the Triassic/Early Jurassic was unparalleled by the basal neobatrachian radiation in the Late Jurassic/Early Cretaceous or any subsequent radiation in the Late Cretaceous/Early Tertiary. Comparative analyses of radiation episodes indicate that the slowdown of morphospace expansion was caused not only by a drop in evolutionary rate after the basal anuran radiation but also by an overall increase in homoplasy in the characters that did evolve during later radiations. The overlapping sets of evolving characters among more recent radiations may have enhanced tadpole diversity by creating unique combinations of homoplastic traits, but the lack of innovative character changes prevented the exploration of fundamental regions in morphospace. These complex patterns transcend the four traditionally recognized tadpole morphotypes and apply to most tissue types and body parts.
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