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Description

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Conraua goliath is the largest frog in the world, with some individuals weighing as much as 3.3 kilograms and reaching ~32 cm snout–vent length (Krüger 1912; Sabater-Pi 1985. Channing and Rödel 2019). The largest reported individuals are females, with males ranging from 22 – 32 cm snout–vent length (1.5 – 2.7 kg) and females ranging from 15 – 32 cm snout–vent length (0.6 – 3.3 kg) (Sabater-Pi 1985). 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). Tadpoles have robust and muscular tails and mouths with prominent, heavily keratinized jaw sheaths, 7 rows of keratodonts surrounding the mouth (keratodont formula: 7(5 – 7)/7(1)), and many small papillae, especially around lateral and ventral borders (Lamotte et al. 1959; Channing et al. 2012). The straight, short spiracular tube is located on the left side of the posterior half of the body. The vent tube is nearly one-third the length of the body and opens on the right side of the ventral fin (Channing et al. 2012). 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). 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). 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).The species authority is: 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. The phylogenetic affinities of C. goliath with respect to other species in the genus remain unresolved as there is no comprehensive analysis of Conraua. 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 goliathConraua 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).

References

  • Amiet, J.-L. (1986). ''La batrachofaune sylvicole d’un secteur forestier du Cameroun: la région de Yaoundé.'' Mémoires du Muséum national d’histoire naturelle, Sér. A., Zoologie, 132, 29–42.
  • Amiet, J.-L., Goutte, S. (2017). Chants d'amphibiens du Cameroun. Locus Solus Publishing House, Chateaulin, France.
  • 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,
  • Channing, A., Rödel, M.-O. (2019). . Struik Nature, 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.
  • Gewalt, W (1976). ''Der 1000-Dollar-Frosch aus Kamerun.'' Tier, 16(11), 8-11.
  • Gewalt, W. (1977). ''Einige Bemerkungen über Fang, Transport und Haltung des Goliathfrosches (Conraua goliath Boulenger).'' Zoologische Garten, Neue Folge, Jena, 47(3/4), 161–192.
  • Herrmann, H.-W., Böhme, W., Herrmann, P.A., Plath, M., Schmitz, A., Solbach, M. (2005). ''African biodiversity hotspots: the amphibians of Mt. Nlonako, Cameroon.'' Salamandra, 41(1/2), 61–81.
  • Herrmann, H.-W., Edwards, T. (2006). ''Conraua goliath: skittering locomotion.'' Herpetological Review, 37(2), 202–203.
  • Hutchison, V.H. (1998). The Goliath Frog (Conraua goliath): physiological ecology of the largest anuan. Extended abstracts of the 1998 International Symposium on Animal Adaptation, Institute of Zoology, Academia Sinica, Taipei, Taiwan, Republic of China. 5 pp.
  • Krüger, B. (1912). ''Zur Kenntnis des grössten lebeden Frosches, Rana goliath.'' Blgr. Blätter für Aquarien- und Terrarien-Kunde, 23(24), 383.
  • Lamotte, M. and Perret, J.-L. (1968). ''Revision du genre Conraua Nieden.'' Bulletin de l’Institut fondamental d’Afrique noire, Série A, 30, 1603-1644.
  • Lamotte, M., Perret, J.-L. and Dzieduszycka, S. (1959). ''Contribution à l'étude des batraciens de l'Ouest Africain IX. Les formes larvaires de Petropedetes palmipes, Conraua goliath et Acanthixalus spinosus.'' Bulletin de l'Institut fondamental d'Afrique noire, Série A, 21, 762-776.
  • Notas sobre la ecología de la Rana Gigante de Río Muni (Conraua goliath, Boulenger) (1967). ''.'' Zoo, 1967, 24–25.
  • Pawley, R. (1987). ''The management of the Goliath Frog, Conraua goliath: a goliath problem?'' Bulletin of the Chicago Herpetological Society, 22(5), 96–97.
  • Pawley, R., Hutchison, V.H. (1989). ''Territorial behavior and vocalizing in the Goliath Frog, Conraua goliath, in the Chicago Zoological Park.'' Abstracts of the First World Congress of Herpetology, Canterbury, UK.
  • Perret, J.-L. (1957). ''Observations sur Rana goliath.'' Blgr. Bulletin de la Société Neucthâteloise des Scienes Naturelles, 80, 195–202.
  • Perret, J.-L. (1960). ''Études herpétologiques africaines II.'' Bulletin de la Société Neucthâteloise des Scienes Naturelles, 83, 93–100.
  • Perret, J.-L. and Mertens, R. (1957). ''Étude d’une collection herpétologique faite au Cameroun de 1952 á 1955.'' Bulletin de l’Institut fondamental d’Afrique noire, Série A, 19, 548-601.
  • Ruoso, Cyril. “Taking Down Goliath.” Biographic.com, 8 Mar. 2018, www.biographic.com/posts/sto/taking-down-goliath.
  • Sabater-Pi, J. (1972). ''Der größte Frosch der Erde ist ein fast unbekanntes Tier.'' Das Tier, 1972, 36–37.
  • Sabater-Pi, J. (1985). ''Contribution to the biology of the Giant Frog (Conraua goliath, Boulenger).'' Amphibia-Reptilia, 6(2), 143-153.
  • 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.
  • 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.
  • Zahl, P.A. (1967). ''In quest of the world’s largest frog.'' National Geographic Magazine, 132(1), 146–152.

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Distribution and Habitat

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The distribution of Conraua goliath is limited to a small region of Central Africa in the countries of Cameroon and Equatorial Guinea. In this region, C. goliath occurs in the tropical lowland forests of the Lower Guinean Forest Zone, extending 150 – 200 km inland. Its distribution extends from the Sanaga river basin in Cameroon in the north to the Benito river basin in the Republic of Equatorial Guinea in the south, and is found in rivers including the Benito, Kienké, Laña, Nkebe, Ntem, Nyong, Sanaga, and Uoro (Boulenger 1906 - Annals; Perret 1957; Sabater-Pi 1985; Lasso et al. 2002; Herrmann et al. 2005). Adult goliath frogs are found in rapids and cascades of rivers with a sandy bottom and clean, slightly tannic, cool (16 – 22 C), 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).
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Life History, Abundance, Activity, and Special Behaviors

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This species has a limited geographic distribution, narrow ecological requirements, is actively hunted by peoples in both Cameroon and Equatorial Guinea, and is threatened by both the conversion of forests into agricultural lands and potentially hydroelectric dams. There are currently ex situ conservation efforts beginning in Cameroon in an attempt to rear this species in captivity (Ruoso 2018).
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Life History, Abundance, Activity, and Special Behaviors

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While adults of C. goliath are generally nocturnal, during the day adults may bask on rocks emerging from streams and rivers for up to a half hour at a time before returning to the water (Sabater-Pi 1985; Channing and Rödel 2019). The escape behavior of individuals in or near rivers is to jump into the water, typically with a single leap. In some cases, however, individuals have been observed to skitter across the surface of the water with as many as seven consecutive short leaps over as much as 3.5 m (Herrmann and Edwards 2006). On land, however, individuals typically appear exhausted after five or six leaps (Sabater-Pi 1967) making them easy to capture by humans. 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). 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, a high elevation Angolan endemic. Thus, the identify of this aquatic plant remains unknown, though is plausibly a member of the Podostemaceae (M. Cheek, pers. comm. to D.C. Blackburn). 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.
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Relation to Humans

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The Goliath frog is hunted frequently for food, including using poisons, nets, traps, and snares, including above oviposition sites (Perret 1957; Sabater-Pi 1985; Schäfer et al. 2019). As of 2019, it remains the only amphibian species afforded special protection status in Cameroon, which is why it subject to the US Lacey Act even though it is not protected by CITES.
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Lifespan, longevity, and ageing

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Maximum longevity: 5.9 years (captivity) Observations: Weighting over 3Kgs, this is considered the largest frog in the world (http://amphibiaweb.org/).
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Habitat

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Goliath frogs are found in or near fast-flowing rivers and waterfalls in dense, humid and relatively high-temperature rain forests. The water is usually clean, oxygen rich, slightly acidic and usually around 67°F. The presence of larval food also determines where goliath frogs occur.

Habitat Regions: tropical ; freshwater

Terrestrial Biomes: rainforest

Aquatic Biomes: rivers and streams

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Khandelwal, S. 2008. "Conraua goliath" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Conraua_goliath.html
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Sunaina Khandelwal, University of Maryland, Baltimore County
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Kevin Omland, University of Maryland, Baltimore County
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Associations

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The greenish brown dorsal skin of goliath frogs camouflages them well with the wet-moss covered rocks that adult frogs bask on. Goliath frogs are only known to be preyed on by humans, although other predators are possible.

Known Predators:

  • humans (Homo sapiens)

Anti-predator Adaptations: cryptic

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Khandelwal, S. 2008. "Conraua goliath" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Conraua_goliath.html
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Sunaina Khandelwal, University of Maryland, Baltimore County
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Morphology

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Goliath frogs are the largest frogs on Earth. They are from 17 to 32 cm long and from 600 to 3250 grams in weight. They have a flattened, wide body with a triangular head. Their dorsal skin is granular and greenish brown and the ventral skin is a lighter yellowish green. The hind legs are long, front legs are shorter and stout, and all of the feet are extensively webbed. Their eyes can be nearly 2.5 cm in diameter.

Range mass: 600 to 3250 g.

Range length: 17 to 32 cm.

Other Physical Features: ectothermic ; heterothermic ; bilateral symmetry

Sexual Dimorphism: sexes alike

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Khandelwal, S. 2008. "Conraua goliath" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Conraua_goliath.html
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Sunaina Khandelwal, University of Maryland, Baltimore County
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Life Expectancy

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Lifespan of goliath frogs is not well documented, although individuals in the wild seem to live longer than those in captivity.

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Khandelwal, S. 2008. "Conraua goliath" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Conraua_goliath.html
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Sunaina Khandelwal, University of Maryland, Baltimore County
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Distribution

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The geographic range of goliath frogs is restricted to coastal, equatorial Guinea and Cameroon in West Africa.

Biogeographic Regions: ethiopian (Native )

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Khandelwal, S. 2008. "Conraua goliath" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Conraua_goliath.html
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Sunaina Khandelwal, University of Maryland, Baltimore County
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Kevin Omland, University of Maryland, Baltimore County
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Trophic Strategy

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Adult goliath frogs feed on insects, crustaceans, fish, mollusks, small mammals and amphibians, including newts, salamanders, and smaller frogs. Goliath tadpoles are vegetarians and feed only on one aquatic plant, Dicraea warmingii, found only near waterfalls and streams.

Animal Foods: mammals; amphibians; insects; terrestrial non-insect arthropods; mollusks

Plant Foods: algae

Primary Diet: carnivore (Insectivore , Eats non-insect arthropods, Molluscivore )

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Khandelwal, S. 2008. "Conraua goliath" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Conraua_goliath.html
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Sunaina Khandelwal, University of Maryland, Baltimore County
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Kevin Omland, University of Maryland, Baltimore County
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Associations

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Being carnivores, frogs occupy an intermediate position in food webs and their numbers can be indicators of the health of a broader ecosystem. Generally frogs regulate insect populations and also feed on other frogs, fish, small mammals and other small amphibians. Frogs in turn serve as food for birds, reptiles and mammals.

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Khandelwal, S. 2008. "Conraua goliath" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Conraua_goliath.html
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Sunaina Khandelwal, University of Maryland, Baltimore County
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Kevin Omland, University of Maryland, Baltimore County
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Benefits

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Because of their size, goliath frogs are valued for the international pet and zoo trade. They are also a valuable food source for local communities.

Positive Impacts: pet trade ; food

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Khandelwal, S. 2008. "Conraua goliath" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Conraua_goliath.html
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Sunaina Khandelwal, University of Maryland, Baltimore County
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Kevin Omland, University of Maryland, Baltimore County
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Benefits

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There are no known adverse effects of goliath frogs on humans

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Khandelwal, S. 2008. "Conraua goliath" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Conraua_goliath.html
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Sunaina Khandelwal, University of Maryland, Baltimore County
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Life Cycle

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Goliath frog tadpoles are not much larger than those of other frogs, but the majority of growth occurs in the first 75 to 90 days of development. The goliath frog's young tadpoles are not much larger than other frogs'. So the majority of the growth process occurs during the 75-90 days. As in other frogs, goliath frog larvae undergo metamorphosis to become adult frogs.

Development - Life Cycle: metamorphosis

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Khandelwal, S. 2008. "Conraua goliath" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Conraua_goliath.html
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Sunaina Khandelwal, University of Maryland, Baltimore County
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Conservation Status

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The IUCN sites goliath frogs as an endangered species because of a 50% decline in population size in the last three generations. The most significant threats to the species include hunting for food, destruction of forest habitat because of human settling, sedimentation, commercial logging and agriculture, and exportation for zoos, the pet trade, and competitive frog races. Some of the goliath frogs natural habitats have been declared protected areas including the Monte Alen National Park in Equatorial Guinea. Three wildlife sanctuaries in Littoral Province have also been approved as protected areas in Cameroon. The Equatorial Guinean government has set a limit of 300 on the frogs that may be exported per year. However, local conservation efforts and captive-breeding programs also need to be considered in order to maintain population sizes.

US Federal List: threatened

CITES: no special status

IUCN Red List of Threatened Species: endangered

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Khandelwal, S. 2008. "Conraua goliath" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Conraua_goliath.html
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Sunaina Khandelwal, University of Maryland, Baltimore County
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Kevin Omland, University of Maryland, Baltimore County
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Behavior

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Goliath frogs do not have vocal sacs and are often considered mute. As a mating call, goliath frogs hold their mouths open and make a whistling noise.

Communication Channels: visual ; acoustic

Perception Channels: visual ; tactile ; acoustic

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Khandelwal, S. 2008. "Conraua goliath" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Conraua_goliath.html
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Sunaina Khandelwal, University of Maryland, Baltimore County
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Kevin Omland, University of Maryland, Baltimore County
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Reproduction

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Little is known about goliath frog mating systems. The little research that has been done suggests that male goliath frogs perform the calling ritual differently from most other frogs. Instead of sucking in air into vocal sacs and blowing it out to make calls, goliath frogs hold the mouth open to make a long whistling noise. Goliath frogs do not have vocal sacs. Females then follow this noise to the male.

Mating System: polygynous

Female goliath frogs lay several hundred eggs at a time that are about 3.5 mm in diameter. They stick to vegetation at the bottom of the rivers and streams in which they reproduce. The eggs then hatch into tadpoles that grow to about 5 cm long over the next 75 to 90 days. There is no information about the seasonality or frequency of breeding or time to sexual maturity.

Key Reproductive Features: iteroparous ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; fertilization (External ); oviparous

There is no parental investment in goliath frog young after the female deposits the eggs after fertilization.

Parental Investment: no parental involvement; pre-fertilization (Provisioning, Protecting: Female)

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Khandelwal, S. 2008. "Conraua goliath" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Conraua_goliath.html
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Sunaina Khandelwal, University of Maryland, Baltimore County
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Biology

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Unlike most other frogs and toads, goliath frogs have no vocal sac and therefore courtship does not involve displaying any calls (2). Females lay large clutches of several hundred eggs onto vegetation on the river bottom (3). Once the tadpoles hatch they feed on the food plant, which is only located near to waterfalls and rapids (2). It takes around 85 to 95 days for complete metamorphosis into the adult form (3). Adults feed on insects, crustaceans and fish (2).
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Conservation

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The goliath frog is not currently protected under any trade restrictions and collection continues to threaten remaining populations. Captive breeding programmes have not proven successful and the only method of effectively preserving this amphibious giant is to safeguard areas of remaining habitat (2).
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Description

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The massive goliath frog, as its name alludes, is the largest frog in the world with individuals weighing over three kilograms (2). The granular skin is greenish in colour with a yellowish-orange underside, and the feet and hands are webbed (3).
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Habitat

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Inhabits fast-flowing rainforest rivers and cascades, which have sandy bottoms and rocks covered with Dicraea warmingii warmingii, the primary food source for the goliath frog tadpoles (3).
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Range

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Found in a narrow range in Equatorial Guinea and Cameroon, south from the basin of the Sanaga River to the Benito, Wolo and Mbini Rivers (3).
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Status

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Classified as Endangered (EN) on the IUCN Red List 2007 (1).
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Threats

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Much of the dense rainforest of the goliath frog's habitat has been deforested for timber and to make way for agriculture. The construction of dams also threatens the breeding habitat of these frogs and this species is particularly vulnerable to habitat alteration due to its highly restricted range (2). Additionally, goliath frogs are considered a delicacy and are collected by local people for food (3); their large size has also encouraged collection for the pet trade in the past (2).
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Goliath frog

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The goliath frog (Conraua goliath), otherwise known commonly as the giant slippery frog and the goliath bullfrog, is a species of frog in the family Conrauidae. The goliath frog is the largest living frog.[3][4] Specimens can grow up to 32 centimetres (12.6 in) in length from snout to vent, and weigh up to 3.25 kilograms (7.2 lb). This species has a relatively small habitat range in Cameroon and Equatorial Guinea.[5] Its numbers are dwindling due to habitat destruction and its collection for food and the pet trade.[1]

Description

Sexual dimorphism between the sexes is minimal.[6] In a sample of 15 individuals, weights ranged between 600 and 3,250 g (1.3 and 7.2 lb), and snout-vent lengths were between 17 and 32 cm (6.7 and 12.6 in).[5] Their eyes can be nearly 2.5 cm (1.0 in) in diameter. The conspicuous tympanum has a diameter around 0.5 cm (0.20 in) and is separated from the eye by about 5 cm (2.0 in) in adults. Goliath frog eggs and tadpoles are about the same size as other frogs despite their very large adult form.

A lateral fold extends from the eye to the posterior portion of the tympanum. Toes are fully webbed, with large interdigital membranes extending down to the toe tips. The second toe is the longest. The skin on the dorsum and on top of the limbs is granular. Dorsal coloration is green sienna, while the abdomen and ventral part of the limbs are yellow/orange. They have acute hearing, but no vocal sac, and also lack nuptial pads.

Habitat and distribution

The Goliath frog is mainly found near waterfalls in Equatorial Guinea and Cameroon. Their habitat is divided into two main seasons: the dry season which occurs from November to April and the rainy season which occurs from May to October.[7]

The Goliath frog is one of the largest frogs and since it has such a large size, it has an extremely selective distribution. This species is primarily located in a dense equatorial forest fringe which is somewhat parallel to the coast and surrounded by rivers. The Goliath frog was located in Sanaga Basin mainly appearing near the Nachtigal cascades and in the Sakbayeme rapids, in Kienke Basin, Ntem Basin mainly being located near the rapids of the Mensolo and Nsana, in Mbía Basin where it was found to be very abundant in the rapids and cascades. These distribution patterns emphasize its limited environment which tends to have a clear preference for water territories.[8]

Conservation

Preserved specimen at the Harvard Museum of Natural History

The primary threat to the goliath frog is hunting, as it is considered a food source in its native range.[1] The IUCN has highlighted the need for conservation measures, in cooperation with local communities, to make sure the hunting is at sustainable levels.[1] To a lesser extent they are also threatened by habitat loss and degradation.[1] They were extensively exported to zoos and the pet trade, but have proven shy and nervous in captivity.[9] Although captives may live longer than their wild counterparts, the species has not been bred in captivity.[9][10] Due to their classification as an endangered species, the Equatorial Guinean government has declared that no more than 300 goliaths may be exported per year for the pet trade, but few now seem to be exported from this country.[3]

Diet

It was determined in a study that the goliath frog consumes a wide variety of food which suggests that the frog is omnivorous with a carnivorous preference. Their prey are terrestrial, aquatic, and semi-aquatic, indicating that they hunt both on land and in water. Food preferences were different among the different weight groups of each frog which could be correlated to different stages of development. For frogs who weigh less than 400 g (14 oz) (could be equivalent to the young), they consumed annelids, arachnids, myriapods, insects, crustaceans, gastropods, and reptiles. For frogs who weigh more than 1,000 g (35 oz) (could be equivalent to a fully developed frog), they consumed arachnids, myriapods, insects, crustaceans, and gastropods with a significantly higher occurrence of myriapods. This shows that annelids and reptiles are present only in the diet of low weight frogs which emphasizes the more diversified diet for younger goliath frogs.[11] Fully developed frogs are also believed to prey on fish, small mammals, and amphibians, including newts, salamanders, and smaller frogs.[12]

Reproduction

Like most amphibians, water is vital for their reproduction. Because the goliath frog lacks a vocal sac, it does not produce mating calls, a behavior generally present in frogs and toads.[5] The egg masses consist of several hundred to a few thousand eggs, approximately 3.5 mm (0.14 in) each, and often attached to aquatic vegetation.[13] There are three main nest types, all semi-circular in shape and located in or near a river: In the first they clear a section in a river pool. In the second they expand an already existing pool, daming it off from the river. In the final they dig a pool, roughly 1 m (3 ft) wide and 10 cm (4 in) deep, sometimes moving quite large stones in the process. This may partially explain the goliath frog's large size, as larger frogs may be more successful at moving heavy objects when constructing their nests. Adults have also been shown to guard the nests at night. Although not confirmed, there are indications that the nest is constructed by the male, while the female guards the nest with the eggs.[13] Larval development takes between 85 and 95 days.

Life cycle

While the reproductive behavior for this species is mostly not well-known, studies by Lamotte, Perret, and Zahl have allowed an overall chronological table of larval development to be created.[5]

Typically after 24 hours, the cover of the egg mass becomes yellow and the eggs become dark gray-brown. These egg masses were found attached to the bottom of plants. [5]

First Month: no organs were differentiated and only the ocular region showed significant pigmentation and transparent external gills. The lengths were 8–10 mm (0.31–0.39 in) and while the body/tail appeared to be slightly pigmented, the abdomen was always nearly white.[5]

Second Month: The activity of the tadpoles increased greatly for they are now beginning to feed on leaves. Furthermore, they have developed a denser pigmentation, and the spiraculum and anal tube are now beginning to become visible. Additionally, the mouth and the eyes are beginning to function. Sizes range 19–21 mm (0.75–0.83 in). As the month continues, pigmentation is becoming more intense via the presence of nearly black spots, two rows of teeth are developing on both the upper and lower lip, feeding increases greatly, and their size becomes 24 mm (0.94 in).[5]

Third Month: The posterior legs are now beginning to form and the length of the tadpoles are around 40mm. As the month progresses, the posterior legs become larger and joints/fingers are becoming more distinguishable. The total length of the tadpoles is now 45 mm. Finally, at the end of the month, anterior legs have fully appeared, posterior legs are grown with long and powerful fingers accompanying this growth, the mouth has become an arched slot line, tail regression has begun, and these tadpoles begin to put their head out of the water in order to breathe.[5]

Fourth Month: All of the specimens in the experiment reached the final stage of metamorphosis. The tail has been either completely or nearly reabsorbed, the shape and color of adults is obtained with slightly lighter and greener pigmentation, the total length is 35 mm (1.4 in). The large size characteristic of these frogs is not obtained yet. The entire process of larval development takes approximately 85–95 days to complete. [5]

Longevity

The goliath frog can live up to 15 years in the wild. In captivity, they can live up to 21 years. It is preyed upon by snakes, Nile crocodiles, Nile monitors, and humans, among other predators.

Parental care

Nesting patterns

The goliath frog creates nests as sites for their offspring as a form of parental care. There are three main types of nests: type 1 mainly contains rock pools that were cleared from detritus and leaf litter, type 2 contains existing washouts at riverbanks, and type 3 were depressions dug by the frogs into the gravel riverbanks. Each nest type contains advantages and disadvantages. Nest type 1 was the easiest to create since only cleaning of the substrate was required to create the nest. Consequently, these types of nests were the least reliable since they were usually positioned in the river bed which makes them extremely vulnerable to being washed out by the rising water levels and to having predators enter the nest. Both nest types 2 and 3 were less likely to be washed out, however, they have an increased risk for being dried-up during the dry season. Consequently, while each nest promotes clear advantages, nests are typically constructed depending on the environment cues (whether it is the dry season or the rainy season). All nest types can be used several times, and can consist of three distinct cohorts of tadpoles.[14] The construction of these nests can also explain how the goliath frog became the largest frog. Digging out these nests which exceed 1 m in diameter is an extremely arduous task. Other species which perform this task are also quite large in size. This includes: male African Bullfrogs, Gladiator Frogs, and the Bornean Giant River Frog.[13]

Typically, the goliath frog attaches its eggs either underwater, in small groups to rocks, or in gravel or larger pieces of wood. The construction of the nests may help reduce predation for it would be more difficult for the fish and shrimp (species which typically eat the eggs) to find the eggs and it may prevent the eggs from being washed away by the rapid current. However, in contrast, the changing water levels may also cause an increase in predation, cause more of the eggs to spill out, and increase tadpole morality as well since the tadpoles and eggs may remain trapped within the nests. Additionally, these nests allow the Goliath frogs to become less dependent on existing structures for egg deposition which can allow these frogs to prolong their breeding season and also increase the amount of suitable breeding sites (they are determined to be suitable by the absence of predators or water presence since water is required for the offspring to survive).[13]

The process of constructing a nest for the offspring is used as a method of promoting a male’s reproductive benefits to the females. It also serves as the main parental investment since once the female deposits the eggs after fertilization, there is no more parental investment. [13]

Enemies

Parasites

The goliath frog is endangered due to deforestation, overhunting, and parasites. One particular parasite is one species of microfilaria which belongs to the Icoseilla genus. This parasite is often found within the blood and lymphatic system and its spread throughout the lymphatic system can cause lethargy and mortality when the infection is serious. This parasite was more prevalent during the dry season which is primarily due to the water speed decreasing thereby allowing more potential breeding sites created. Since there are more breeding sites (it is important to note that their primary habitat is near rivers and waterfalls, but in the dry season, they tend to create breeding sites in areas that contain less water), there would be more opportunity for the mosquitos to infect the frogs thereby increasing the contagion rate. Furthermore, there was a positive relationship detected between host size and parasite abundance: the greater the size of the host, the more intense the infection would be. Within this species, male goliath frogs were found to be more significantly infected when compared to the females which can be due to the weight difference between the two. Additionally, as within most species, as age increases, the severity of the infection will also increase.[15]

There are also parasites that mainly target the gastro-intestinal tract of this frog. These parasites are called helminth parasites which are worm-like parasites divided into three main groups: flukes which are leaf-shaped flatworms, tapeworms which are elongated flatworms that inhabit extraintestinal tissues, and roundworms which inhabit intestinal and extraintestinal sites. The Goliath frog, however, was mainly infected by Nematodes (90.5%) which is a specific type of roundworm. The helminth species discovered within the Goliath frog was extremely similar to those discovered in amphibian hosts in other African countries emphasizing that its location/habitat is the main cause of the prevalence of this predator. However, the Goliath frog was also infected by the nymph of Sebekia sp which could be primarily due to these frogs sharing the same habitat of crocodiles (definitive host) and fishes (intermediate host). Frogs that originate in Loum and Yabassi which are places within Cameroon had the largest variety of helminth species whereas frogs from Nkondjock had the smallest variety. This could be explained by the difference in agricultural activities, deforestation, and poaching. Liver weight was examined to reveal that there is a higher accumulation of toxic products at Loum due to the significant increase in agriculture practice. Consequently, land use effects and their impact on water habitats plays a significant role in the pattern of parasitism and land use by goliath frogs. Additionally, the direct life cycles of the helminths may play an important role in species diversity Parasites which have direct life cycles spend a majority of their adult life within one host which allows their offspring to be spread from one host to another. These types of parasites also often lack an intermediate stage which means that they must be able to survive in the outside environment as well and be able to establish themselves within a new host. This ability to adapt to new environments contributes greatly to the complexity of helminth communities of goliath frogs.[16][17][18]

Diseases: Chytridiomycosis

Chytridiomycosis is an infectious disease which affects many different species of amphibians worldwide. It is caused by the chytrid fungus which has the capability of causing random deaths due to its high mortality rate. This fungus tends to invade the surface layers of the skin thereby causing damage to the outer keratin layer (this fungus only attacks the parts of the frog’s skin that contains keratin). As the tadpole continues to grow, more keratin becomes present on the skin thereby allowing the fungus to spread to many parts of the body resulting in the death of the tadpole. Amphibian skin is vital because it is physiologically active meaning it plays important roles in regulating respiration, water, and electrolytes. While the method of how this fungus kills frogs is not known, it can be hypothesized that its invasion through the skin is related since it can cause an electrolyte depletion, osmotic imbalance, and make it more difficult for the frog to breathe. When infected with this fungus, a frog may have discolored skin, peeling on the outside layers of its skin, be sluggish, and have its legs spread slightly away from itself. This fungus is also transmittable since it can be directly transferred through contact between frogs and tadpoles or through exposure to infected water. Thus, it is highly spreadable and with its high mortality rate among frogs, it is extremely deadly.[19][20]

Physiology

The goliath frog contains extensive skin folds to promote respiratory gas exchange at high altitudes. Additionally, the lungs within these frogs are reduced to about one-third of the volume of other frogs and they also contain a smaller heart. This is primarily due to their difference in predation methods. Goliath frogs are typically sit-and-wait predators. This means that they tend to capture/trap their prey either by luring them or using elements of surprise by acting extremely stealthy. As a result, they have a reduced metabolic rate and a unique method of breathing. When attempting to breathe, each buccal movement (a method in which the mouth expands and contracts in order to promote the movement of air into the lungs) pumps air at a rapid rate and the process of getting the oxygen removed from the air is slightly more efficient in this species of frog. These adaptations are very useful for the production of territorial and reproductive calls created by these frogs. Typically, the call of the goliath frog which is produced by the frog opening its mouth is admitted at a high frequency of 4.4 kHz. Furthermore, the goliath frog does not contain a vocal sac which causes the process for making noises/calls for reproduction to be different than most frogs.[21]

Interactions with humans

While effects of climate change, agriculture, and deforestation greatly affect this species to the extent of making them endangered, they are also mainly captured by the locals who use lanterns to get their attention and then shortly after, they immobilize them using meshed nets. In Nkombia, they are also captured with nets during the day while they are resting on the rocks. However, this method of capture is not very effective for they are able to escape quite well due to their ability to jump 1.2 to 3 m (3.9 to 9.8 ft) high. Humans are the main predators for this species and are the main cause of their endangerment. In order to save the species, hunting and evironmental destruction should be limited.

See also

  • Helmeted water toad (Calyptocephalella gayi) – A South American species that can reach a similar length, but with a lower maximum weight
  • Lake Junin giant frog (Telmatobius macrostomus) – A South American species that is the world's largest exclusively aquatic frog

References

  1. ^ a b c d e IUCN SSC Amphibian Specialist Group (2019). "Conraua goliath". IUCN Red List of Threatened Species. 2019: e.T5263A96062132. doi:10.2305/IUCN.UK.2019-1.RLTS.T5263A96062132.en. Retrieved 19 November 2021.
  2. ^ "Conraua goliath". Wikispecies.
  3. ^ a b "Goliath Frog". The American Museum of Natural History. 2007. Retrieved 9 February 2012.
  4. ^ Macle, Adam (22 January 2017). "10 Largest Frogs and Toads in the World". Gone Froggin. Retrieved 29 May 2021.
  5. ^ a b c d e f g h i Sabater-Pi, J. (1985). "Contribution to the biology of the Giant Frog (Conraua goliath, Boulenger)". Amphibia-Reptilia. 6 (1): 143–153. doi:10.1163/156853885x00047.
  6. ^ Halliday, T. (2016). The Book of Frogs: A Life-Size Guide to Six Hundred Species from around the World. University of Chicago Press. p. 527. ISBN 978-0226184654.
  7. ^ Nguete Nguiffo, Daniel; Wondji, Charles S.; Pone Wabo, Josué; Mpoame, Mbida (2019). "Microfilariae infestation of goliath frogs (Conraua goliath) from Cameroon". PLOS ONE. 14 (5): e0217539. Bibcode:2019PLoSO..1417539N. doi:10.1371/journal.pone.0217539. PMC 6541376. PMID 31141563.
  8. ^ Sabater-Pi, Jordi (1985). "Contribution to the Biology of the Giant Frog (Conraua goliath, Boulenger)". Amphibia-Reptilia. 6 (2): 143–153. doi:10.1163/156853885X00047.
  9. ^ a b Indiviglio, Frank (31 January 2013). "The World's Largest Frog – Working with the Massive Goliath Frog". thatpetplace. Retrieved 4 November 2017.
  10. ^ "Goliath Frog is a Disappearing Giant". Frogsource. Retrieved 4 November 2017.
  11. ^ https://journals.univ-tlemcen.dz/GABJ/index.php/GABJ/article/view/177/131
  12. ^ "Conraua goliath". Animal Diversity Web.
  13. ^ a b c d e Schäfer, Marvin; Tsekané, Sedrick Junior; Tchassem, F. Arnaud M.; Drakulić, Sanja; Kameni, Marina; Gonwouo, Nono L.; Rödel, Mark-Oliver (2019). "Goliath frogs build nests for spawning – the reason for their gigantism?". Journal of Natural History. 53 (21–22): 1263–1276. doi:10.1080/00222933.2019.1642528. S2CID 202012353.
  14. ^ Schäfer, Marvin; Tsekané, Sedrick Junior; Tchassem, F. Arnaud M.; Drakulić, Sanja; Kameni, Marina; Gonwouo, Nono L.; Rödel, Mark-Oliver (11 June 2019). "Goliath frogs build nests for spawning – the reason for their gigantism?". Journal of Natural History. 53 (21–22): 1263–1276. doi:10.1080/00222933.2019.1642528. ISSN 0022-2933. S2CID 202012353.
  15. ^ Nguete Nguiffo, Daniel; Wondji, Charles S.; Pone Wabo, Josué; Mpoame, Mbida (29 May 2019). "Microfilariae infestation of goliath frogs (Conraua goliath) from Cameroon". PLOS ONE. 14 (5): e0217539. Bibcode:2019PLoSO..1417539N. doi:10.1371/journal.pone.0217539. ISSN 1932-6203. PMC 6541376. PMID 31141563.
  16. ^ Castro, Gilbert A. (1996), Baron, Samuel (ed.), "Helminths: Structure, Classification, Growth, and Development", Medical Microbiology (4th ed.), Galveston (TX): University of Texas Medical Branch at Galveston, ISBN 978-0-9631172-1-2, PMID 21413320, retrieved 29 November 2022
  17. ^ Daniel, Nguiffo Nguete; Josue, Wabo Pone; Mpoame, Mbida (1 July 2015). "Gastro-intestinal helminths of goliath frogs (Conraua goliath) from the localities of Loum, Yabassi and Nkondjock in the Littoral Region of Cameroon". Global Ecology and Conservation. 4: 146–149. doi:10.1016/j.gecco.2015.06.009. ISSN 2351-9894.
  18. ^ "parasitic disease - Parasite life cycles | Britannica". www.britannica.com. Retrieved 29 November 2022.
  19. ^ Heritage., Natural Heritage Trust (Australia) Australia. Department of the Environment and (2004). Chytridiomycosis (amphibian chytrid fungus disease). Dept. of the Environment and Heritage, Natural Heritage Trust. OCLC 62538132.
  20. ^ "Frog chytrid fungus". NSW Environment and Heritage. Retrieved 29 November 2022.
  21. ^ "Buccal Pumping - an overview | ScienceDirect Topics".
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Goliath frog: Brief Summary

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The goliath frog (Conraua goliath), otherwise known commonly as the giant slippery frog and the goliath bullfrog, is a species of frog in the family Conrauidae. The goliath frog is the largest living frog. Specimens can grow up to 32 centimetres (12.6 in) in length from snout to vent, and weigh up to 3.25 kilograms (7.2 lb). This species has a relatively small habitat range in Cameroon and Equatorial Guinea. Its numbers are dwindling due to habitat destruction and its collection for food and the pet trade.

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