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Growling grass frog

The Growling grass frog (Ranoidea raniformis), also known as the southern bell frog, is a large, robust species of in the Pelodryadidae, characterized by its distinctive deep, growling advertisement call and bright to olive-green coloration marked with irregular brown or bronze blotches. Adults typically measure 60–104 mm in snout-to-vent length, with females larger than males, and feature warty skin, prominent tympana, and pale undersides often tinged with yellow or blue on the thighs. Tadpoles are large, up to 110 mm, with a greenish-yellow surface and a prolonged aquatic larval stage lasting 2–15 months. Native to southeastern , including , , , and , the species inhabits a variety of permanent and semi-permanent freshwater bodies such as swamps, marshes, slow-flowing streams, farm dams, and irrigation channels, preferring sites with emergent and submerged vegetation like cumbungi and pondweeds for cover and . It has been introduced to , where small populations persist. occurs primarily in and summer, with males calling diurnally and nocturnally from water's edge or floating vegetation to attract females, who deposit eggs in cohesive masses on the water surface that hatch within 3–4 days. The frog is highly mobile, capable of dispersing up to 1 km in a day, and overwinters in nearby vegetation, logs, or burrows. Once widespread and common, the growling grass frog has undergone severe population declines since the , particularly in upland areas, leading to its classification as Vulnerable on the (as of 2023) and under Australia's Environment Protection and Biodiversity Conservation Act 1999, with populations severely fragmented and continuing to decline. Major threats include and modification from , , and altered hydrology; the amphibian ; predation by such as , foxes, and cats; and on water availability and temperature regimes. efforts focus on , , and threat mitigation through protected areas and plans in regions like greater and southwestern , where some populations remain relatively stable. In 2025, a taxonomic revision reclassified the species from the genus Litoria to Ranoidea within the family Pelodryadidae.

Taxonomy

Classification

The growling grass frog is classified within the domain Eukarya, kingdom Animalia, phylum Chordata, subphylum Vertebrata, class Amphibia, order Anura, family Pelodryadidae, genus Ranoidea, and species R. raniformis. The species was originally described as Chirodryas raniformis by Keferstein in 1867, based on a specimen from Australia with a type locality listed as "Australien" (later specified as possibly New South Wales). Keferstein's description was initially synonymized with Hyla aurea in 1868 but was later recognized as distinct. Following a comprehensive phylogenomic analysis, the species was transferred from the genus Litoria to Ranoidea in a 2025 taxonomic revision of the Pelodryadidae family, reflecting its phylogenetic placement within a clade of predominantly ground-dwelling and semi-aquatic Australian frogs. As of November 2025, this revision is not yet universally adopted, with some sources continuing to use Litoria raniformis. Synonyms include Litoria raniformis (used until 2025), Hyla aurea raniformis, Hyla aurea major, and Ranoidea major. The placement in Pelodryadidae distinguishes it from true hylid frogs of the , as this family represents a distinct Australo-Papuan radiation adapted to diverse and environments.

Etymology

The "growling grass frog" originates from the species' deep, guttural , which resembles a growl, and its frequent occurrence in grassy wetlands. However, the "grass frog" designation is misleading, as the species belongs to the family Pelodryadidae rather than the African grass frog genus Ptychadena. Alternative common names include "southern bell frog," alluding to a resonant, bell-like aspect of its vocalizations in certain contexts; "warty swamp frog," referencing its textured, warty dorsal skin and affinity for swampy environments; and erroneously "," stemming from superficial color similarities and taxonomic confusion with other green-hued species like the (Ranoidea aurea). The binomial scientific name Ranoidea raniformis reflects both morphological and phylogenetic considerations. The Ranoidea, newly erected in 2025 from the former broad Litoria based on genomic , derives from the superfamily Ranoidea and evokes a resemblance to the true frogs of the family Ranidae in habitus and form. The specific epithet raniformis combines the Latin rana (frog) with -formis (in the shape of), denoting "frog-shaped" and underscoring the species' archetypal anuran morphology as perceived in 19th-century . Originally described as Chirodryas raniformis by German zoologist Johann Wilhelm Keferstein in 1867 from specimens collected in , the name has undergone revisions: it was transferred to Litoria in the to align with Australo-Papuan tree frogs, and reclassified to Ranoidea in 2025 amid a major phylogenomic restructuring of the family Pelodryadidae that split Litoria into 35 genera.

Description

Adult morphology

The growling grass frog (Ranoidea raniformis), also known as the southern bell frog, is a large with adults reaching a snout-to-vent length (SVL) of up to 100 mm, though females are typically larger (average 70–100 mm) than males (average 50–70 mm). The body is robust, with long hind legs adapted for jumping and swimming, fully on the hind feet, and expanded adhesive discs on the toe tips that facilitate climbing on vegetation and surfaces. The eyes are prominent with horizontal pupils, and a distinct (eardrum) is visible behind each eye. Dorsally, the skin is warty and granular, featuring numerous small tubercles and shallow bumps that give a rough , often accented by a pale longitudinal stripe along the midline and dorso-lateral folds running from behind the eyes to the . Coloration varies but is typically mottled bright to or on the back, with irregular dark blotches; the venter is smooth and cream to white. Hidden surfaces, such as the thighs and , display striking blue- flash colors, sometimes flecked with yellow, which are revealed during movement. Sexual dimorphism is evident in size, with females larger than males, and in breeding males, which develop black nuptial pads on the inner thumbs for amplexus and a mottled dark throat. Compared to the similar green and golden bell frog (Ranoidea aurea), the growling grass frog has a rougher, more warty dorsal skin with fewer prominent ridges and less extensive yellow markings along the sides.

Larval stage

The tadpoles of the growling grass frog (Ranoidea raniformis) reach a total length of 80–110 mm, with occasional individuals up to 130–150 mm, featuring an elongated body and a deep, powerful tail fin that facilitates agile swimming near the water surface. Newly hatched tadpoles are greyish-black or dark brown, while older larvae exhibit variable coloration ranging from dark velvety black-brown to green, light yellowish-grey, or pinkish-grey with yellowish tail fins; individuals approaching often develop a yellowish-pink sheen or golden markings along the sides. The belly is typically translucent, allowing visibility of internal structures. Key morphological features include a large oral disc adapted for filter-feeding and , equipped with two upper and three lower rows of labial teeth—the inner rows divided—and bordered by a narrow band of papillae; are present only briefly after hatching and are soon resorbed. Eggs hatch 2–4 days after deposition, with tadpoles completing in approximately 3 months under warm conditions (around 23°C), though the larval period can extend to 2–15 months in cooler regions where development is delayed. These tadpoles demonstrate notable adaptations for in temperate environments, including the to overwinter in a dormant state during cold periods, tolerating low temperatures that prolong the larval phase without mortality. They are also physiologically equipped for aquatic life, excreting as (ammonotelic) to maintain balance in , and their active helps them access oxygenated surface layers in potentially stagnant habitats.

Distribution and habitat

Geographic range

The growling grass frog (Litoria raniformis) is native to south-eastern , with its distribution spanning from the south-eastern corner of —particularly along the —throughout most of and , and extending to the eastern half of . The species is absent from arid interior regions of the continent and occurs at elevations ranging from to approximately 1,300 m. Historically, the frog was once widespread in wetlands across a broad area from in to in , representing one of the most common species in these habitats prior to the late 1970s. Since the 1980s, its range has contracted markedly, with notable disappearances from higher elevations, the Australian Capital Territory, and parts of the Mt Lofty Ranges and Plains in . Current native populations are fragmented, particularly in urbanizing areas around where habitat isolation has led to isolated remnants, while more stable groups persist in rural wetlands such as those in north-western and south-western , the Lower Murrumbidgee and systems in , and eastern . Outside its native range, L. raniformis was introduced to New Zealand in 1867 on the South Island from Melbourne stock and has since become established across wetlands in both the North and South Islands. Populations in New Zealand remain widespread and generally stable, though some localized declines have occurred, particularly in North Island wetlands influenced by chytrid fungus.

Preferred habitats

The growling grass frog (Litoria raniformis) primarily inhabits permanent or semi-permanent freshwater bodies, including swamps, ponds, lakes, farm dams, irrigation channels, quarries, and slow-flowing rivers or streams. These environments provide essential conditions for breeding and survival, often featuring large, continuous areas that undergo regular flooding cycles. Vegetation plays a critical role in habitat selection, with the frog associating closely with emergent reeds such as common reed (), cumbungi (Typha orientalis), and tall spike-rush (Eleocharis sphacelata), as well as submerged plants like curly pondweed () and hornwort (). Floating vegetation, including duckweed (Lemna spp.) and ferny azolla (), offers additional cover for shelter and breeding sites, while grassy margins around water edges support basking and foraging activities. High aquatic vegetation cover—typically 20-80% depending on wetland permanence—is preferred to enhance occupancy probabilities. Microhabitat preferences emphasize sunny, shallow edges of water bodies for , where adults can bask during the day and engage in nocturnal calling and foraging among emergent or floating vegetation, bank-side rocks, or open pasture. The shows tolerance for brackish conditions, with electrical conductivity up to approximately 10,000 μS/cm, allowing occupation of slightly saline wetlands in certain regions. Habitat use varies by life stage: adults occupy vegetated wetlands for shelter under logs, rocks, or dense , particularly during non-breeding periods, while dispersing to flooded areas in and summer for . Tadpoles develop in open still or slow-moving water, requiring permanent or seasonally flooded sites free of for successful , which can take 2–15 months, depending on temperature and location. Juveniles seek moist refugia in wetlands post-, rapidly growing to maturity within about four months and utilizing similar vegetated margins as adults. Key habitat requirements include water depths of 0.5-2 meters to support growth and development, with deeper areas exceeding 1.5 meters ideal for sustained quality. Connectivity to floodplains is essential, facilitating breeding cues and dispersal corridors between populations, often within a 1 km radius for optimal occupancy.

Ecology and behavior

Diet and foraging

The growling grass frog (Litoria raniformis) is an opportunistic with a varied diet consisting primarily of such as (including crickets, beetles, moths, and cicadas), spiders, and crustaceans, supplemented by small vertebrates like tadpoles, , and other frogs, including conspecifics. Less commonly, it preys on and . Tadpoles are also herbivorous-detritivorous, feeding on and before incorporating small or engaging in as they grow. As a sit-and-wait , the growling grass frog forages mainly at night, using visual cues to detect prey from perches on emergent , rocks, or bare near edges. It employs lunge strikes to capture prey within reach, often swallowing items whole. activity peaks in late summer and autumn, aligning with higher prey availability in wetlands. In wetland ecosystems, the growling grass frog serves as a mid-level predator, helping regulate invertebrate populations and serving as prey for larger animals like birds and mammals.

Reproduction

The growling grass frog (Litoria raniformis) breeds during spring and early summer in its native range, typically from August or September to January or February, with occasional extension into March or April depending on environmental conditions. Breeding activity is triggered by rainfall events that fill or replenish wetlands, combined with water temperatures between 11°C and 16°C, which stimulate chorusing behavior among males. In southern Australia, peak breeding coincides with warmer months when daytime air temperatures often exceed 15°C, promoting female responsiveness to male calls. Mating is initiated by males calling from positions floating in open water or at the edges of wetlands, producing a characteristic growl or to attract females, often in large choruses that occur day or night. Once a female approaches, the male clasps her in axillary (pectoral) , a firm embrace around the upper body that stimulates egg release and deposition. The exhibits a polygamous , with males capable of with multiple females during a single breeding season, and females potentially laying more than one if conditions allow repeated matings. Females deposit eggs in gelatinous, floating rafts or mats attached to submerged , with each containing 1,900 to 4,000 pigmented eggs measuring 1.3–1.5 mm in diameter within 2–2.5 mm capsules. These rafts typically sink within 12 hours after deposition, allowing eggs to develop in shallow water. There is no , with eggs and resulting tadpoles left to develop independently. Sexual maturity is reached after 2 years of age, corresponding to a snout-vent length of approximately 55 mm. In the wild, individuals have a lifespan of up to 4–5 years, though most populations show a majority of adults aged 2–3 years due to environmental pressures.

Vocalization

The primary vocalization of the growling grass frog (Litoria raniformis) is the advertisement call produced by males, consisting of a deep growl typically lasting about 1 second and described onomatopoeically as "crawark-crawark-crok-crok." This call features a long introductory note followed by two or three short grunts, with dominant frequencies around 1200 Hz and a frequency range of 300–3000 Hz, including secondary peaks at approximately 500 Hz and 1300 Hz. The call's low-frequency components facilitate long-distance transmission in wetland environments, allowing choruses to be audible over 1 km. Call variations include the growl, which may extend to about 3 seconds in some instances, and a faster variant comprising a train of rapid grunts lasting 0.26–1.07 seconds (mean 0.58 seconds), both serving as advertisement signals in choruses. Males also produce encounter calls, such as short grunts (duration 0.19 seconds, pulse rate 25.7 Hz, dominant 600–800 Hz), during interactions with other males, potentially for territorial defense. Release calls, manifested as high-pitched squeals (~1 second, emphasized at 3.5–6 kHz), are emitted by both sexes in distress, such as when grasped by predators or conspecifics. These variations enable species-specific communication, distinguishing L. raniformis calls from those of the closely related green and golden bell frog (Litoria aurea), which features a distinct drone absent in L. raniformis. Males typically produce these calls while floating in open water or on emergent vegetation, primarily at or after dark during the breeding season from to , when choruses form in response to suitable conditions. Calling activity peaks in warmer months and increases with rising air temperatures, with choruses exhibiting both discrete bursts and prolonged sessions lasting up to several hours. These vocalizations play a key role in during . Recordings of L. raniformis calls have been available since the , with early analyses contributing to taxonomic and bioacoustic studies, and they remain essential for contemporary population surveys through call playback and monitoring techniques.

Conservation

The growling grass frog (Litoria raniformis) is classified as Vulnerable on the IUCN Red List, with the assessment conducted in 2023 under criteria A2ace, indicating an observed, estimated, projected, or suspected population reduction of at least 30% over three generations due to habitat loss, disease, and other factors. In Australia, the species holds national Vulnerable status under the Environment Protection and Biodiversity Conservation Act 1999 (EPBC Act). Regionally, it is listed as Vulnerable under Victoria's Flora and Fauna Guarantee Act 1988, as per the February 2024 threatened list. In Tasmania, it is also Vulnerable under the Threatened Species Protection Act 1995, though populations there have shown relative stability compared to mainland declines, with an estimated 5,000–10,000 adults persisting. Population trends indicate a marked decline across the species' range since the 1980s and 1990s, with reductions exceeding 50% in some areas, including local extirpations from urban and parts of the Australian Capital Territory (). Remaining populations are highly fragmented, with global mature individual estimates fewer than 10,000, concentrated in isolated sites. Some stabilization has been observed in managed wetlands through targeted . Monitoring efforts, including annual call playback surveys and auditory detection, have tracked these trends since the 1990s, particularly in and , providing data on occupancy and abundance changes.

Threats

The growling grass frog (Litoria raniformis) faces significant habitat loss primarily due to , , and wetland drainage, which have drastically reduced available breeding sites across its range. In the southern Murray-Darling Basin, more than 80% of native vegetation has been cleared, severely impacting wetland habitats essential for the species. Urban development around has further exacerbated habitat degradation and fragmentation, isolating populations and limiting dispersal. River regulation and altered water management practices, including dam construction and diversions, have disrupted natural flooding cycles critical for , as the species relies on seasonal inundation of wetlands. Prolonged droughts, intensified by these modifications, further dry out temporary water bodies, preventing and leading to population crashes. Chytridiomycosis, caused by the chytrid fungus , has been a major driver of die-offs since the 1990s, with the growling grass frog showing high susceptibility to this skin infection that disrupts electrolyte balance and leads to . The disease emerged widely in during this period, contributing to rapid declines in the species' abundance. Introduced predatory fish, particularly the eastern gambusia (Gambusia holbrooki), pose a direct threat by consuming eggs and tadpoles in breeding wetlands, with this proliferating in modified waterways. from agricultural pesticides, herbicides, and fertilizers contaminates water bodies, impairing frog development, reducing breeding success, and increasing vulnerability to diseases. compounds these issues by exacerbating wetland drying through reduced rainfall and higher evaporation rates, while elevated ultraviolet-B radiation from may harm basking individuals. In urban fringes such as those around , these threats interact synergistically: increases exposure to predators and pollutants, while altered and disease amplify mortality rates, creating compounded risks for remnant populations.

Conservation efforts

restoration efforts for the growling grass frog (Litoria raniformis) focus on rehabilitating wetlands through revegetation and hydrological management to support breeding and dispersal. In , projects emphasize re-vegetating emergent reeds and aquatic in degraded wetlands to provide and foraging sites, with positive responses observed in restored sites featuring floating or emergent vegetation. The Melbourne Strategic Assessment program collaborates with Melbourne Water to create new breeding wetlands and secure areas, enhancing between fragmented populations in urbanizing landscapes. In the Murray-Darling Basin, environmental watering regimes simulate natural flooding to inundate floodplains, triggering breeding events and maintaining habitats critical for the species. Policy protections underpin these initiatives, with the species listed as vulnerable under the federal Environment Protection and Biodiversity Conservation (EPBC) Act 1999, mandating environmental impact assessments for actions that may significantly affect its habitat. In , the Flora and Fauna Guarantee (FFG) Act 1988 supports action statements that designate reserves and guide land management to protect key sites, including farm dams and swamps in southwestern regions. Disease management targets , a major threat, through protocols in translocation programs that include , antifungal treatments like baths, and site to prevent spread. Ongoing research explores chytrid resistance in populations, with studies assessing infection dynamics in cooler climates to inform refugia creation. Monitoring and research efforts involve community-based surveys using the FrogID app to track distribution and abundance, building public awareness and data for . Genetic studies, such as a 2015 analysis of Melbourne's urban-fringe populations, reveal low connectivity and diversity, guiding restoration to enhance via habitat corridors. Success stories include population stabilization in managed farm dams modified with aquatic vegetation and refuge zones, where enhanced hydroperiods have supported persistent breeding. Reintroduction trials, such as those at Winton Wetlands in Victoria, have established new populations in restored habitats absent since 1970, demonstrating viability through captive-bred releases and monitoring.

Captivity

As a pet

Keeping grass frogs (Litoria raniformis, also known as southern bell frogs) as s is subject to strict legal regulations in due to their protected status under state acts. In most Australian states and territories, such as and , native frogs are protected , requiring a specific license or permit—such as a biodiversity conservation license or frog keeper license—to possess them legally. Wild capture is illegal across all states, as frogs are considered protected species under legislation like the National Parks and Wildlife Act in . In , where the species is an introduced and naturalised species considered a potential vector for diseases like to native frogs, it is listed in Schedule 5 of the Act 1953, meaning it is not protected. Keeping as s is permitted, but release into the wild is illegal to prevent ecological risks. Suitable housing for pet growling grass frogs must mimic their semi-aquatic , with a large providing both land and areas to accommodate their active, ground-dwelling nature. A minimum size of 60 cm length by 45 cm width by 45 cm height is recommended for one to two adults, allowing space for a shallow (at least one-third of the floor area, 10-15 cm deep) for and soaking, alongside a dry land section with like sphagnum moss or leaf litter for burrowing. The setup should include UVB lighting (5.0 spectrum, 10-12 hours daily) to support synthesis, as well as multiple hiding spots such as cork bark, PVC pipes, or live plants like for security and climbing. is critical, requiring dechlorinated, filtered changed weekly to prevent bacterial growth, with a gentle system to maintain slow-moving conditions. Basic care involves a diet of live insects to meet their carnivorous needs, fed every 2-3 days to avoid obesity. Appropriate prey includes gut-loaded crickets, dubia roaches, earthworms, and locusts dusted with calcium and multivitamin supplements, comprising 10-20% of the frog's body weight per feeding session. Environmental parameters must be maintained at 20-30°C (with a warm basking spot up to 27°C to reduce chytrid susceptibility) using under-tank heaters or ceramic heat emitters, and humidity levels of 70-90% achieved through daily misting and the water feature. Annual veterinary health checks are essential, focusing on screening for chytridiomycosis via skin swabs, as captive amphibians are prone to this fungal disease; early detection allows antifungal treatments like itraconazole baths. Specimens must be sourced exclusively from captive-bred stock obtained through licensed breeders or registered suppliers, as wild collection is prohibited in and contributes to population declines. In , sourcing is similarly restricted to prevent further spread of the invasive population. Challenges in captive care include the species' high susceptibility to from improper handling or enclosure disturbances, which can suppress immunity and lead to or refusal to eat. Additionally, they are vulnerable to infectious diseases like and bacterial dermatitis (red leg syndrome) if temperatures drop below 23°C or humidity fluctuates, necessitating vigilant monitoring and for new additions. Poor or inadequate UVB exposure can also cause , underscoring the need for experienced keepers.

Breeding programs

Captive breeding programs for the growling grass frog (Litoria raniformis) are primarily managed by zoos affiliated with the Zoo and Aquarium Association (ZAA), the Research Centre, and conservation organizations such as Winton Wetlands and Nature Glenelg Trust, in collaboration with government agencies like the Victorian Department of Energy, Environment and Climate Action. These initiatives aim to produce disease-free individuals for reintroduction to bolster declining wild populations, with a focus on maintaining to support long-term viability. Breeding techniques include the use of simulated enclosures to mimic natural habitats, such as outdoor with aquatic and controlled water levels at facilities like Winton Wetlands' quarantine and laboratory setup. Hormone induction, involving agonists and antagonists, has been successfully applied to stimulate spermiation, courting , and spawning in captive adults. Genetic strategies, including individuals from diverse wild sources, help prevent , as outlined in ZAA protocols that have achieved breeding up to the F4 generation. Head-starting programs raise tadpoles to juvenile stages in controlled environments to improve overwintering survival before release. Success has been demonstrated through consistent breeding in captivity over the past 15 years, with programs like 's Clayton Bay facility raising initial batches of 35 tadpoles to maturity and preparing for production of hundreds to thousands more for release. At Winton Wetlands, 30 wild-sourced adults were translocated in 2023, quarantined, and moved to breeding enclosures, where calling and mating behaviors were observed in subsequent seasons, establishing a foundational captive population. As of the 2024/2025 breeding season at Winton Wetlands, increased breeding behavior was observed among the captive population, though no egg laying occurred; the team remains optimistic for future spawning. These efforts have contributed to reintroductions, such as the planned release of captive-bred juveniles into Victorian wetlands to restore local populations. Challenges include the high risk of chytrid fungus () infection, necessitating strict and measures during translocation and rearing. Post-release survival rates remain low, estimated at 4-14% during active seasons based on mark-recapture studies, due to predation, quality, and exposure in the wild. Limited funding and expertise for long-term husbandry further constrain program scale. These breeding programs integrate with broader wild by releasing head-started froglets into restored wetlands, such as those enhanced under the Melbourne Strategic Assessment, to support connectivity and research on threat mitigation. Monitoring via audio detection and genetic sampling informs , linking captive outputs to habitat protection efforts.

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