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Batrachia
Batrachia
Batrachia
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Batrachia
Temporal range:
Early Triassic – Present, 250–0 Ma Possible earlier origin based on molecular clock
Australian green tree frog (Litoria caerulea)
Fire salamander (Salamandra salamandra)
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Chordata
Class: Amphibia
Subclass: Lissamphibia
Superorder: Batrachia
Latreille, 1800
Clades

Caudata
Salientia

The Batrachia /bəˈtreɪkiə/ are a clade of amphibians that includes frogs and salamanders, but not caecilians nor the extinct allocaudates.[1] The name Batrachia was first used by French zoologist Pierre André Latreille in 1800 to refer to frogs, but has more recently been defined in a phylogenetic sense as a node-based taxon that includes the last common ancestor of frogs and salamanders and all of its descendants. The idea that frogs and salamanders are more closely related to each other than either is to caecilians is strongly supported by morphological and molecular evidence; they are, for instance, the only vertebrates able to raise and lower their eyes.[2] However, an alternative hypothesis exists in which salamanders and caecilians are each other's closest relatives as part of a clade called the Procera, with frogs positioned as the sister taxon of this group.[3]

Origins

[edit]

The earliest batrachians are the stem-frogs Triadobatrachus and Czatkobatrachus from the Early Triassic, about 250 million years ago. However, several molecular clock estimates place the first appearance of the Batrachia (the time at which frog and salamander lineages diverged from each other) before the Early Triassic. Most estimates place the divergence in the Permian[4] but some put it as far back as 367 million years ago in the Late Devonian (which is when tetrapods are thought to have started to emerge from fishapods).[5] However, there is no evidence of lissamphibians or lissamphibian-like animals in the fossil record at this time. The tetrapod groups that are hypothesized as ancestors of modern amphibians (lepospondyls and amphibamid temnospondyls) appear in the Late Carboniferous, roughly 300 million years ago. Large fossil tetrapod assemblages are known from the Artinskian stage of the Early Permian about 275 million years ago and contain no lissamphibians, suggesting that the Early Permian may be an upper bound for the age of Batrachia.[6]

References

[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Batrachia

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Batrachia is a major within the subclass , consisting of the orders Anura (frogs and toads) and Urodela (salamanders and newts) as sister groups, excluding the worm-like (Gymnophiona). This clade represents the vast majority of extant diversity, with 7,915 species of anurans and 828 species of urodeles, totaling 8,743 species as of November 2025. The evolutionary origins of Batrachia trace back to the Permian period, with stem-group representatives such as Gerobatrachus hottoni appearing in the Early Permian of around 290 million years ago, exhibiting a mosaic of primitive and derived traits shared with both frogs and salamanders. Phylogenetic analyses indicate that the divergence of the anuran and urodelan lineages occurred by the Middle Permian, predating the radiation of crown-group lissamphibians, though the earliest undisputed fossils of frogs date to the and salamanders to the . This timeline challenges some estimates suggesting later origins and supports a temnospondyl ancestry for Batrachia within tetrapods. Batrachia encompasses a remarkable range of adaptations, including the specialized jumping locomotion and vocalization of anurans, contrasted with the often elongated bodies, regeneration abilities, and aquatic or terrestrial lifestyles of urodeles. Ecologically, batrachians are vital components of terrestrial and freshwater ecosystems globally, serving as predators, prey, and indicators of , though many species face threats from habitat loss and . Recent phylogenomic research, incorporating thousands of genes across hundreds of species, has strongly corroborated the of Batrachia and its position as the sister to Gymnophiona, resolving long-standing debates in amphibian systematics.

Etymology and Definition

Etymology

The term Batrachia derives from the Ancient Greek word bátrachos (βάτραχος), meaning "frog," reflecting the group's historical association with frog-like amphibians. The name was first introduced in scientific nomenclature by the French naturalist Alexandre Brongniart in 1800, who used Batraciens to designate a group encompassing frogs (Anura) and salamanders (Urodela), excluding caecilians (Gymnophiona) and treating them as distinct from reptiles. This was promptly latinized to Batrachia (or Batrachii) by Pierre André Latreille in the same year, establishing it as a subclass for tailed and tailless amphibians, deliberately excluding reptiles to clarify their separation within the broader class of Amphibia. Subsequent usage evolved the term's scope; for instance, Blasius Merrem in 1820 expanded Batrachia to include all extant amphibians, incorporating caecilians (Apoda) alongside frogs (Salientia) and salamanders (Gradientia) as a comprehensive class. Over time, the nomenclature shifted in line with advancing classifications, with Batrachia increasingly applied to clades focused on frogs and salamanders in post-19th-century systematics.

Definition

Batrachia is a monophyletic within the subclass , encompassing all extant frogs (order Anura) and salamanders (order Urodela), defined as the of these two orders and all of its descendants. This node-based definition excludes (order Gymnophiona), which form the sister clade to Batrachia within Lissamphibia. Taxonomically, Batrachia holds the rank of superorder under the class Amphibia. The core definition of Batrachia focuses on living taxa and does not incorporate extinct groups such as Allocaudata or , although some phylogenetic hypotheses propose affinities between these fossils and Batrachia. Key synapomorphies uniting Anura and Urodela include the ability to retract the eyeballs into the buccal cavity to assist in prey transport during feeding, a feature absent in , as well as a specific pattern of . These traits underscore the close evolutionary relationship between frogs and salamanders, distinguishing Batrachia from other lineages.

Taxonomy

Historical Classification

The term Batrachia was initially established in the early 19th century as a taxonomic group for amphibians. Pierre André Latreille introduced it in 1800 as a subclass (Batrachii) to encompass tailed and tailless amphibians, distinguishing them within the broader class of reptiles. Contemporaneously, Alexandre Brongniart, in his 1800 essay on reptile classification, employed Batrachia as one of four orders within Reptilia, explicitly separating amphibians from saurians, ophidians, and chelonians based on morphological traits like skin texture and reproduction. By 1820, Blasius Merrem elevated Batrachia to class rank in his systematic attempt, defining it to include all extant amphibians: caecilians (Apoda), salamanders (Gradientia), and frogs (Salientia). Subsequent refinements built upon this foundation. André Marie Constant Duméril, in 1806, organized Batrachia into two orders—Anoures (tailless, now Anura) and Urodèles (tailed, now Urodela)—emphasizing anatomical differences such as tail presence and limb structure while retaining caecilians within the broader amphibian framework. Johann Jakob von Tschudi, in his 1838 classification, further subdivided Batrachia into suborders, including Salientia for frogs and Derotremata for certain salamanders, aiming to reflect graduated morphological series among the groups. In the , Batrachia continued to denote all living amphibians in many systems, with (Gymnophiona) included alongside frogs and salamanders. For instance, G. Kingsley Noble's influential treated the three orders as a monophyletic class Amphibia, synonymous with Batrachia in scope, based on shared larval features and osteological similarities. Post-World War II, debates intensified over amphibian monophyly, with some herpetologists questioning the unity of , frogs, and salamanders due to disparate cranial morphologies and life histories, though traditional inclusive classifications persisted. The advent of cladistic methods in the prompted a shift, with analyses increasingly supporting Batrachia as a excluding , restricted to frogs ( or Anura) and salamanders (Urodela). This usage, common in older texts as a synonym for Salientia + Urodela, reflected emerging evidence of closer affinities between those groups. In modern , Batrachia is similarly limited to frogs and salamanders.

Modern Placement

In contemporary , Batrachia is classified as a superorder within the subclass of the class Amphibia, comprising the orders Anura (frogs) and (salamanders), to the exclusion of Gymnophiona (). The complete hierarchical lineage is cellular organisms > Eukaryota > Opisthokonta > Metazoa > > > Deuterostomia > Chordata > Craniata > Vertebrata > > > Euteleostomi > > Dipnotetrapodomorpha > Tetrapoda > Amphibia > > Batrachia. The name Batrachia was originally proposed by Latreille in 1800 as a group for frogs and salamanders, but it has been redefined in modern to denote the monophyletic uniting Anura and based on shared morphological and molecular synapomorphies, such as pedicellate teeth and a specific vertebral structure. This placement aligns with the Batrachia hypothesis, strongly supported by multilocus molecular analyses that recover frogs and salamanders as sister groups with high . Batrachia is recognized in major databases, including the NCBI Taxonomy (ID 41666) as a superorder, and the Amphibian Species of the World database maintained by the American Museum of Natural History, which acknowledges its usage within Lissamphibia despite some historical synonymy with the broader subclass. In rank-free phylogenetic systems under the PhyloCode, Batrachia is often treated as an unranked clade to emphasize evolutionary relationships over Linnaean ranks.

Phylogeny

Position in Lissamphibia

Batrachia, comprising the orders Anura (frogs) and (salamanders), is positioned as the to Gymnophiona () within the crown-group , the clade encompassing all living amphibians. This arrangement reflects the of , which includes Batrachia and Gymnophiona as its two primary subclades, a consensus supported by both molecular and evidence. The of is further reinforced by shared morphological synapomorphies, such as pedicellate, bicuspid teeth, and a bifid in larval stages, which distinguish the group from stem tetrapods. Key phylogenetic evidence for this placement includes molecular analyses that consistently recover Batrachia as a clade diverging from in the Late to Early Permian. estimates indicate the split between Batrachia and Gymnophiona occurred approximately 300–350 million years ago (Ma), aligning with the earliest divergences within crown during the late . support comes from stem batrachians like Gerobatrachus hottoni from the Early Permian, which exhibits traits bridging anuran and caudate morphologies and positions Batrachia as a derived within . Alternative hypotheses, such as the "Procera" clade uniting and Gymnophiona to the exclusion of Anura, have been proposed based on limited early molecular datasets but are rejected by the majority of subsequent studies favoring Batrachia due to stronger congruence across genomic and morphological data. For instance, comprehensive phylogenomic analyses and large-scale molecular phylogenies have upheld Batrachia as the dominant topology. Recent studies, including a 2023 analysis of stem caecilians and 2025 re-examinations of early frog fossils, continue to support this placement within a dissorophoid temnospondyl ancestry. Regarding stem lissamphibians, temnospondyls are excluded from the crown group, with only certain dissorophoid temnospondyls considered as stem taxa giving rise to ; Batrachia emerges as a derived within this crown radiation, distinct from broader temnospondyl diversity.

Internal Relationships

The internal phylogeny of Batrachia is defined by the basal divergence between its two extant orders, Anura (frogs and toads) and (salamanders and newts), which represent the primary lineages within this . This split is estimated to have occurred in the Middle Permian (~265 million years ago), based on phylogenetic analyses of fossil evidence such as Gerobatrachus hottoni. Phylogenetic reconstructions consistently depict Anura and as sister taxa, forming a monophyletic group exclusive of (Gymnophiona), with shared derived traits such as pedicellate teeth and a specific vertebral structure supporting this topology. Consensus on Batrachia's internal relationships has been reached through integrated morphological and molecular evidence. Morphological studies, including analyses of the salientian stem lineage—encompassing early frog-like forms—highlight skeletal adaptations like elongated ilia and reduced tail vertebrae that unite Anura and as closer relatives than either is to gymnophionans. Seminal work by Milner (1983) on the origins of salientians provided early morphological support for this batrachian framework by tracing anuran evolution from stem forms. Complementing this, molecular phylogenies utilizing (mtDNA) and nuclear genes have robustly confirmed Batrachia and the Anura-Caudata linkage, with high bootstrap support in maximum likelihood trees derived from multi-gene datasets. San Mauro's (2010) multilocus analysis, incorporating 23 genes totaling over 9,000 base pairs, reinforced these findings through , estimating the divergence while underscoring the clade's ancient origins. Extinct relatives further illuminate Batrachia's internal structure, with albanerpetontids often positioned as a potential outgroup or stem batrachian lineage in phylogenetic analyses. These small, lizard-like amphibians, known from to fossils, share features like fused frontals and angular jaws with batrachians but retain plesiomorphic traits such as epipterygoids, suggesting they branched off near the base of Batrachia or as its immediate sister . Cladistic studies place albanerpetontids outside crown Batrachia but within , providing critical context for the clade's early diversification without altering the core Anura-Caudata topology.

Evolutionary History

Origins

The origins of the Batrachia clade, encompassing frogs (Anura) and salamanders (Caudata), are inferred primarily through molecular clock analyses and phylogenetic reconstructions, as direct fossil evidence remains sparse for the critical early phases. Molecular dating estimates place the divergence of crown-group Batrachia—the split between the lineages leading to modern Anura and Caudata—between approximately 250 and 300 million years ago (Ma) during the Late Carboniferous to Permian. Most estimates place the appearance of stem-Batrachia in the Permian around 290 Ma, though some molecular clock analyses suggest earlier origins in the Late Devonian (~370 Ma). These timelines are derived from multilocus datasets and relaxed clock models calibrated against key fossil nodes, highlighting a rapid early diversification within Lissamphibia that predates the breakup of the supercontinent Pangaea. The environmental context for Batrachia's emergence points to freshwater habitats prevalent in the Paleozoic Era, where early ancestors transitioned from aquatic to semi-aquatic lifestyles amid fluctuating continental conditions. These origins likely coincided with periods of environmental instability, including mass extinctions such as the Late Devonian and the broader Carboniferous-Permian transitions, which reshaped aquatic ecosystems and favored adaptations like permeable skin and larval aquatic stages in precursor lineages. Batrachians are thought to have adapted from more generalized forebears in shallow lakes, streams, and swamps, where features such as systems and supported survival in oxygen-variable waters before evolving enhanced terrestrial capabilities. Precursor groups for Batrachia are debated but commonly traced to dissorophoid temnospondyls within the Temnospondyli subclass, with alternative hypotheses linking them to lepospondyl amniote relatives; no unambiguous crown-Batrachia fossils are known from the Permian, though inferences draw from broader lissamphibian phylogenetic patterns. A key debate centers on the exact ancestral morphology, with taxa like the Early Permian Gerobatrachus hottoni—an amphibamid temnospondyl exhibiting a mosaic of salientian and caudatan traits, such as short trunk vertebrae and robust otic notches—proposed as a potential stem batrachian that bridges Paleozoic amphibians to Mesozoic crown forms. This temnospondyl origin supports a monophyletic Lissamphibia emerging from aquatic dissorophoids, contrasting polyphyletic models that split Batrachia and caecilians across temnospondyl and lepospondyl lines.

Fossil Record

The fossil record of Batrachia, encompassing frogs () and salamanders (), is notably sparse, reflecting the challenges of preservation in the geological record due to their small size, delicate skeletons, and predominantly aquatic or semi-aquatic lifestyles. Stem-batrachians are known from the Early Permian, such as Gerobatrachus hottoni (~290 Ma, ), exhibiting mixed anuran-urodelan traits. The earliest known fossils attributable to crown Batrachia appear in the , marking the initial diversification of stem-group members shortly after the Permian-Triassic mass extinction. massinoti, discovered in and dated to approximately 250 million years ago (Ma), represents a stem salientian with primitive features such as a elongated trunk with 14 presacral vertebrae (compared to the nine typical of modern frogs) and reduced tail, providing key evidence for the early evolution of frog-like body plans. Similarly, Czatkobatrachus polonicus from Early Triassic deposits in exhibits salientian affinities, including a shortened trunk and ilium morphology suggestive of jumping adaptations, further supporting a rapid post-extinction radiation of batrachians in high-latitude environments. The record remains limited, with fewer than a dozen well-documented taxa for crown groups, underscoring significant gaps particularly for early s. The record begins in the with stem forms like Triassurus sixtelae (~230 Ma, ), and crown-group caudates appear by the , such as Marmorerpeton wakei and Kokartus honorarius (~166 Ma). Karaurus sharovi from () sediments in (about 155 Ma) represents an early crown caudate, characterized by a -like body with well-developed limbs and a tail fin, indicative of an aquatic lifestyle. For frogs, Prosalirus bitis from the Kayenta Formation in (approximately 190 Ma) is the oldest known crown salientian, featuring advanced traits like a bicondylar sacro-urostylic articulation that enabled powerful leaps, bridging stem and modern anuran morphology. Overall, batrachian fossils are rare outside of Laurasian localities, likely biased by depositional environments favoring exceptional preservation in lagoonal or karstic settings. The era witnesses a marked expansion in batrachian diversity following the Cretaceous-Paleogene (K-Pg) extinction, with fossils becoming more abundant and geographically widespread, reflecting recovery and into varied niches. Palaeobatrachids, an extinct family of frogs, are prominent in Eocene deposits across , such as those from the Geiseltal and Messel sites in (around 45-40 Ma), where well-preserved skeletons reveal specialized aquatic adaptations like elongated bodies and reduced limbs, contrasting with the more terrestrial modern anurans. This post-K-Pg proliferation includes diverse assemblages in North American and Eurasian lagerstätten, though the total described fossil taxa for Batrachia numbers in the hundreds, highlighting ongoing challenges in sampling and identification. These fossils collectively imply that batrachians achieved much of their modern morphological disparity by the , with implications for understanding resilience to mass extinctions.

Characteristics

Shared Traits

Batrachia, the clade comprising frogs (Anura) and salamanders (), exhibits several morphological and physiological features characteristic of the group, though its is primarily supported by phylogenetic analyses rather than unique morphological synapomorphies. One prominent feature is pedicellate dentition, where teeth consist of two distinct parts—a basal pedicel and a —separated by an unmineralized zone, allowing flexibility and replacement. This structure is evident in marginal teeth along the jaws and supports feeding adaptations in both anurans and urodeles. Members of Batrachia possess the levator bulbi muscle, a specialized ocular muscle that enables protrusion of the eyes in anurans and urodeles, aiding in swallowing by compressing the buccal cavity and facilitating prey manipulation. Larval stages in Batrachia exhibit characteristic external, bushy gills that facilitate aquatic respiration. These gills are filiform and highly branched, maximizing surface area for oxygen exchange during the vulnerable early . Compared to , which retain embedded dermal scales, Batrachia show a complete loss of dermal scales, resulting in a smooth, glandular skin that supports and moisture retention but increases vulnerability to . Sensory adaptations in Batrachia include the lateral line system, retained in aquatic larvae and some adults, which consists of neuromasts detecting water movements and vibrations for navigation and prey detection. Electroreception is absent in Batrachia, with reliance on mechanosensory cues. The clade demonstrates remarkable size variation, from the diminutive frog Paedophryne amauensis at approximately 8 mm snout-vent length to the giant salamander Andrias davidianus reaching up to 1.8 m in total length, reflecting a shared potential for miniaturization and gigantism influenced by habitat and life history.

Group-Specific Adaptations

Members of the order Anura exhibit specialized adaptations for saltatory locomotion, characterized by elongated hindlimbs that enable powerful jumps, often several times their body length, while forelimbs are reduced in size for landing support. These limb modifications, combined with a shortened body and fused sacral vertebrae forming a urostyle, facilitate efficient terrestrial and semi-aquatic movement, distinguishing anurans from other lissamphibians. Additionally, many male anurans possess vocal sacs—expandable throat pouches that amplify mating calls to attract females and establish territory, a feature absent in caudates. Some anuran have evolved direct development, bypassing a free-living larval stage to produce miniature adults, which enhances survival in arid or predator-rich environments. In contrast, members of the order retain a throughout adulthood, which aids in during and balance on land, reflecting their predominantly aquatic or semi-aquatic lifestyles. are renowned for their regenerative capabilities, including the ability to regrow entire limbs, , and portions of the , a process mediated by of cells at the injury site to form a . This regeneration is exemplified in species like the (Ambystoma mexicanum), which also demonstrates paedomorphosis through , retaining larval features such as and aquatic habits into sexual maturity. Reproductive strategies further highlight these divergences: anurans typically employ during , with eggs often deposited in foam nests or gelatinous masses to protect them from and predators. Caudates, however, utilize via spermatophores—gelatinous packets of sperm deposited by males and retrieved by females—leading to diverse modes including and, in some species like certain , where embryos develop within the female. Ecologically, anurans occupy a broader range of niches, with many species adapted to terrestrial or arboreal habitats through adhesive toe pads and enhanced , contributing to their global diversity in forests and grasslands. Caudates, by comparison, are more frequently confined to moist, aquatic or semi-aquatic environments, leveraging their permeable and for underwater respiration and locomotion, though some terrestrial forms exist in humid microhabitats.

Diversity

Composition

Batrachia, the clade encompassing frogs and salamanders, is composed of two extant orders: Anura (frogs and toads) and (salamanders and newts). Anura is the dominant group, containing 7,915 species as of November 2025 distributed across 57 families, including prominent examples such as Ranidae (true frogs) and Bufonidae (toads). In contrast, includes about 828 species organized into 10 families, with notable representatives like (newts and salamanders) and Cryptobranchidae (). The total living diversity of Batrachia exceeds 8,700 species, with Anura accounting for roughly 91% and the remaining 9%. This numerical imbalance highlights Anura's extensive radiation, particularly within the subclade , which encompasses the majority of anuran species. Crown-group Batrachia lacks major extinct orders, though stem taxa such as the Triadobatrachus massinoti represent primitive batrachians that bridge amphibians and modern forms. In terms of family distribution, Anura achieves a near-global presence, occurring on every continent except , while Caudata species are predominantly found in the , with limited representation in southern regions. Batrachia as a whole forms a monophyletic group within , diverging from Gymnophiona early in amphibian evolution.

Distribution

Batrachia exhibit a across all continents except and most oceanic islands, reflecting their adaptability to a wide array of environments worldwide. The group's overall range is marked by highest in tropical regions, particularly the in and , where environmental conditions support prolific and coexistence. This tropical concentration underscores the clade's evolutionary success in humid, resource-rich ecosystems, with over 7,000 species of Anura and approximately 800 of contributing to this global pattern. Anura, comprising frogs and toads, are ubiquitous and occupy an exceptionally broad spectrum of habitats, from arid deserts exemplified by Breviceps species in southern Africa's coastal dunes and sandy plains, to high-altitude montane zones above 4,000 meters. Certain anurans, such as the cane toad (Rhinella marina), have been widely introduced beyond their native Neotropical range and established invasive populations in Australia, parts of North and South America, and oceanic islands, demonstrating their ecological versatility. In contrast, Caudata, or salamanders, display a more temperate bias within the Holarctic realm, with peak diversity in eastern North America and the mountainous regions of Eurasia, and complete absences from Australia, sub-Saharan Africa, and the Afrotropics more broadly. Habitat preferences among Batrachia emphasize freshwater systems for reproduction, including , , and wetlands, alongside terrestrial and arboreal niches for foraging and shelter. Their characteristic biphasic life cycle—featuring aquatic larval stages and primarily terrestrial adult phases—facilitates occupation of diverse ecological roles but renders them vulnerable to disruptions in interconnected aquatic-terrestrial habitats.

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