Conraua goliath (Boulenger, 1906)
Goliath Frog, Giant Frog
|Species Description: Boulenger, G. A. (1906). "Descriptions of new batrachians discovered by Mr. G.L. Bates in South Cameroon." Annals and Magazine of Natural History, Series 7, 17: 317–323.|
© 2019 Marvin Schäfer and Frogs & Friends e.V. (1 of 51)
The species description by Boulenger (1906 - Annals) was based on one specimen. The depressed head is wider than long with an obtusely pointed snout. The nostril is closer to the snout tip than the eye. The canthus rostralis is obtuse and the loreal region is deeply grooved. The interorbital space is shorter than the length of the upper eyelid. The tympanum is small relative to the large head, and bordered above by a prominent fold or ridge. The short fingers each have one large, flat, oval subarticular tubercle. The first and second fingers are about equal and the third is longer than the fourth (Boulenger 1906 - Annals). The hindlimbs are powerful and long relative to the body length (~150% of snout–vent length; Sabater-Pi 1985). When extended along the body, the tibiotarsal articulation reaches the tip of the snout. The feet are as long as the crus (Boulenger 1906 - Annals) and fully webbed with relatively thick interdigital membranes extending along the full length of the toes and ending at the disc-like toe tips (see photograph in Zahl 1967). The foot has a flat, elliptical inner metatarsal tubercle and no outer tubercle. As with the fingers, the moderate toes have large, flat, oval subarticular tubercles. There is a strong dermal fringe on the outer side of the fifth toe (Boulenger 1906 - Annals). The skin on the dorsum and on top of the limbs is granular with many fine ridges (Pawley 1987). The flanks have small warts. The upper surface of the thighs and tibias have longitudinal granular folds. The belly and throat also have small granules (Boulenger 1906 - Annals).
Conraua goliath can be differentiated from Conraua robusta, the only other large Conraua that co-occurs with C. goliath, via the ventral coloration, which is white or gray in C. robusta (Channing and Rödel 2019). Though some species of Conraua retain lateral line organs as adults, these organs are lost at metamorphosis in C. goliath and C. robusta (as well as the other Central African C. crassipes; Gutsche et al. 2023).
In life, the background dorsal coloration ranges from dark green to orangey green with darker spots on the dorsum. There are irregular cross-bars on the limbs and the posterior surface of the hindlimbs are blackish with white spots. The abdomen and ventral part of the limbs are yellowish orange (Boulenger 1906 - Annals; Channing and Rödel 2019).
Distribution and Habitat
Country distribution from AmphibiaWeb's database: Cameroon, Equatorial Guinea
Adult goliath frogs are found in rapids and cascades of rivers with a sandy bottom and clean, slightly tannic, cool (16 – 22 oC), oxygen-rich waters (Sabater-Pi 1962, 1985; Zahl 1967). Typically, these rivers are large enough to not be enclosed within the forest canopy (Amiet 1986).
Life History, Abundance, Activity, and Special Behaviors
Large adults are often observed on rocks close to or within waterfalls and rapids spaced at a distance of 3 - 5 m, whereas subadults are found more often on mid-stream rocks. Metamorphs and juveniles in pools adjacent to the river that have more slow-moving water (Gonwouo et al. 2022).
In Río Muni of Equatorial Guinea, reproduction occurs during the dry season (July and August) as well as during the short dry season of December and January (Zahl 1967).
Egg masses are deposited underwater and attached to vegetation or stones in pools near rapids (Sabater-Pi 1985; Channing and Rödel 2019). In some cases, frogs (likely males) create nests in existing pools and washouts that may be expanded, whereas in others frogs actively dig depressions into the gravel banks of rivers (Schäfer et al. 2019). In all cases, these pools and depressions are actively cleaned of leaf litter and detritus by the frogs. Most nests contain 150 – 350 eggs, though in some cases might have as many as 2,800 eggs (Schäfer et al. 2019). Multiple nests may be deposited in these pools and depressions, with up to three cohorts of tadpoles being involved in some nests, and tadpoles appear to finish development within these nests (Schäfer et al. 2019). Larval development takes approximately 3 months to go through metamorphosis (Zahl 1967; Sabater-Pi 1985). Female frogs attend these nests for some period of time after oviposition (Schäfer et al. 2019).
Freshwater shrimp have been observed feeding on eggs of C. goliath within the nests constructed by adults (Schäfer et al. 2019).
The diets of adults include insects (beetles, ants, dragonflies, caddisflies), millipedes, freshwater shrimp, spiders, scorpions, freshwater snails, as well as larval and adult anurans (Perret 1957, 1960; Sabater-Pi 1962, 1967, 1985; Gewalt 1977). In captivity, when fed young mice, frogs have been observed to feed out of the water, though will also feed while their bodies are submerged in the water, capturing prey on the water’s surface (Pawley 1987). Suction-feeding, however, has not been reported in these aquatic frogs.
Despite previous assertions that adults do not call (Perret 1957; Sabater-Pi, 1985), C. goliath has a repertoire of call types, including brief whistles, chirping, and even a “sighing deep roar” (Channing and Rödel 2019). The advertisement call of Conraua goliath is a high frequency sound of long duration (4.1 kHz, and up to 1.5 minutes; Pawley and Hutchison 1989; Hutchison 1998; Amiet and Goutte 2017). Strangely, claims were made in jest during the early 20th-century that this very large frog might have a high frequency call (Claybird and Wanawiwi 1939), and this appears to be the case. Similar to C. crassipes, the advertisement call of C. goliath is produced with the mouth open (Hutchison 1998).
In captivity, males have been observed to exhibit territorial behaviors including lunges at one another, wrestling venter to venter, and biting (Pawley and Hutchison, 1989). These may be sometimes accompanied by vocalizations varying between guttural calls (1.6 – 2.3 kHz) and high-frequency squeaks (4.5 – 5.8 kHz) (Pawley and Hutchison 1989; Hutchison, 1998). These interactions were only observed during the day and ranged from occurring once per week to as rarely as every three or four months.
In life, the coloration of the body of tadpoles is a somber grey with darker round spots on the dorsal surface, with equally large but more irregular spots on the tail. The spiracular tube is unpigmented (Lamotte et al. 1959; Channing et al. 2012).
Sabater-Pi (1985) reported that tadpoles of C. goliath feed on the leaves of the aquatic plant "Dicraea warmingii"; the genus Dicraea is currently considered a junior synonym of the genus Podostemum (Podostemaceae), which does not occur in Africa, and the taxon to which Sabater-Pi (1985) is likely referring, Inversodicraea warmingiana, is a high elevation Angolan endemic. Thus, the identity of this aquatic plant remains unknown, though is plausibly a member of the Podostemaceae (M. Cheek, pers. comm. to D.C. Blackburn).
Trends and Threats
Relation to Humans
Possible reasons for amphibian decline
General habitat alteration and loss
A recent phylogenetic study of relationships among species of Conraua (Blackburn et al. 2020) found a strongly supported sister relationship between C. goliath and C. robusta; both are large-bodied species that can occur in syntopy in Cameroon. Recent phylogenomic analysis have recovered Conrauidae, which contains only the genus Conraua, as the sister lineage of the Petropedetidae (Feng et al. 2017; Yuan et al. 2019). Conrauidae and Petropedetidae are, in turn, sister to the Pyxicephalidae.
Since their discovery, goliath frogs have proved fascinating to scientists and the public alike (Boulenger 1906 - Proceedings; Krüger, 1912), inspiring work both in the field and at zoos and aquaria (Gewalt, 1976, 1977; Pawley, 1987) and many popular articles (Sabater-Pi, 1962, 1967, 1972; Zahl, 1967; Ruoso, 2018). Some of what we know of the basic biology of these large, charismatic frogs is due to the study of captive animals (e.g., Pawley and Hutchison, 1989; Hutchison, 1998).
Hutchison (1998) found C. goliath to have a somewhat lower metabolic rate than other ranoid frogs. The volume of air brought in during a breath is typically small, but the frogs take frequent breaths and oxygen uptake is higher than other frogs of similar mass.
Several recent videos highlighting behaviors in Conraua goliath
Conraua goliath in the wild [in Spanish]:
Conraua goliath guarding nest in the wild:
This species was featured as News of the Week on October 15, 2019:
Despite having been described in 1906, little is known about the natural history of the largest frog in the world, Conraua goliath. In their recent paper, Schäfer et al. (2019) shed light on the species' reproductive behavior by documenting their construction of nest sites. The authors described three types of nesting sites they found in West Cameroon that protect developing offspring from river torrents and predators: rock pools, existing washouts, and dug-out depressions in gravel riverbanks. The different types of nest sites have differing levels of construction effort and risk of flooding or drying. However, in each of these sites, the breeding adults cleared the area of detritus and leaf-litter and deposited eggs on multiple occasions. Camera traps showed that adults guarded nests at night, which is consistent with local knowledge. The authors speculate that because large, heavy objects must be moved for nest construction, this type of nest construction may have favored large size in this species (Written by Ann T. Chang).
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Amiet, J.-L., Goutte, S. (2017). Chants d'amphibiens du Cameroun. Locus Solus Publishing House, Chateaulin, France. [link]
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Boulenger, G. A. (1906). ''Descriptions of new batrachians discovered by Mr. G.L. Bates in South Cameroon.'' Annals and Magazine of Natural History, Series 7, 17, 317-323.
Boulenger, G.A. (1906). ''Exhibition of a specimen of, and remarks upon, a giant frog from Cameroon.'' Proceedings of the Zoological Society of London, [link]
Channing, A., Rödel, M.-O. (2019). Field Guide to the Frogs and Other Amphibians of Africa. Penguin Random House South Africa, Cape Town, South Africa.
Feng, Y.-J., Blackburn, D.C., Liang, D., Hillis, D.M., Wake, D.B., Cannatella, D.C., Zhang, P. (2017). ''Phylogenomics reveals rapid, simultaneous diversification of three major clades of Gondwanan frogs at the Cretaceous–Paleogene boundary.'' Proceedings of the National Academy of Sciences, USA, 114(29), E5864-E5870. [link]
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Gonwouo, N.L., Schäfer, M., Tsekané, S.J., Hirschfeld, M., Tchassem, F.A.M., Rödel, M.-O. (2022). “Goliath Frog (Conraua goliath) abundance in relation to frog age, habitat, and human activity.” Amphibian & Reptile Conservation, 16(2), e319. [link]
Gutsche, A., Lingelbach, K., Neira-Salamea, K.D., Rödel, M.-O. (2023). “Postmetamorphic retention of a lateral line system in African slippery frogs (Amphibia: Anura: Conraua).” Zoologischer Anzeiger, 304, 21–31. [link]
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Schäfer, M., Tsekané, S.J., Tchassem, F.A.M., Drakulić, S., Kameni, M., Gonwouo, N.L., Rödel, M.-O. (2019). ''Goliath frogs build nests for spawning – the reason for their gigantism?'' Journal of Natural History, 53(21–22), 1263–1276. [link]
Yuan, Z.-Y., Zhang, B.-L., Raxworthy, C.J., Weisrock, D.W., Hime, P.M., Jin, J.-Q., Lemmon, E.M., Lemmon, A.R., Holland, S.D., Kortyna, M.L., Zhou, W.-W., Peng, M.-S., Che, J., Prendini, E. (2019). ''Natatanuran frogs used the Indian Plate to step-stone disperse and radiate across the Indian Ocean.'' National Science Review, 6, 10–14. [link]
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Originally submitted by: David C. Blackburn (first posted 2001-02-26)
Edited by: Ann T. Chang, Michelle S. Koo (2023-05-31)
Species Account Citation: AmphibiaWeb 2023 Conraua goliath: Goliath Frog <https://amphibiaweb.org/species/4691> University of California, Berkeley, CA, USA. Accessed Sep 29, 2023.
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Citation: AmphibiaWeb. 2023. <https://amphibiaweb.org> University of California, Berkeley, CA, USA. Accessed 29 Sep 2023.
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