Phyllomedusa bicolor is a large, arboreal frog (Boulenger 1882; Lima et al. 2005; Mota et al. 2020). Females range from 111 - 119 mm in snout-vent length, while males range from 91 - 113 mm (Lima et al. 2005; Venâncio and Melo-Sampaio 2010). A truncated snout can be observed in addition to an oblique loreal region. The diameter of the eye is shorter than the snout and bigger than the tympanum. The interorbital space is greater than the upper eyelid (Boulenger 1882). There is a distinct parotoid gland observable from behind the eye and over the tympanum (Boulenger 1882; Lacombe et al. 2000; Lima et al. 2005). Skin covers the tympanic membrane (Wagler 1830).
The fingers are free, and the second and third are larger than the first and fourth respectively; the disks of the fingers are flat and similar in size to the tympanum (Wagler 1830; Boulenger 1882). An inner metatarsal tubercle is faint. When the hind limb is adpressed along the body, the tibio-tarsal articulation reaches the shoulder or tympanum. The toes are also free, with the first and second being equal in length, and with smaller disks than the fingers (Boulenger 1882).
Bony deposits stud the upper surface of P. bicolor and granulations that are visible ventrally and at the lower surface of the thighs (Boulenger 1882). Lipid glands appear in the dorsal and dorsolateral regions, mucous glands appear in ventral surface skin, and serous glands appear all over the body (Lacombe et al. 2000).
Pinto et al. (2013) determined that metamorphosed froglets at Stage 46 have a snout-vent ranging from 21.34 - 22.81 mm.
Phyllomedusa bicolor can be differentiated from other species by coloration. Specifically P. bicolor has dark gray irises and purplish/brown fingers with blue/green discs, while P. tarsius has red-orange irises with black reticulations and white-tipped, brown first and second fingers. Such coloration also contrasts to P. vaillanti, which has silvery-gray irises and orange or dark purple discs on the fingers. Furthermore, P. bicolor has blue-green bellies and purple-edged, white spots on their sides, while P. vaillanti has purple bellies and sides (Lima et al. 2005).
It is unclear whether the following coloration description is in life or preservative. The dorsum of P. bicolor is blue-green, while the ventrum ranges from purplish-white, cream, or yellow-white. Small, purple-edged, white spots are visible laterally along the body and limbs, more dense on the flank and hind legs. Similar spots are also on the chest and lower lips. A narrow line of similar coloration to such spots runs along the outer side of the forearms and tarsus. The fingers themselves are either purplish or brown with blue or green discs. Phyllomedusa bicolor also has dark gray irises (Boulenger 1882; Lima et al. 2005).
At the froglet stage, froglets are green, with sparse, darkly-outlined, and white spots on the chest and hind legs (Pinto et al. 2013).
Regarding size, males are generally smaller than females (Lima et al. 2005; Venâncio and Melo-Sampaio 2010).
Distribution and Habitat
Country distribution from AmphibiaWeb's database: Bolivia, Brazil, Colombia, Ecuador, French Guiana, Guyana, Peru, Suriname, Venezuela
Phyllomedusa bicolor is endemic to the Amazon Basin of Brazil, Peru, Bolivia, Columbia, the Guianan Region of Venezuela and the Guianas, extreme eastern Panama, and the Cerrado of Maranhão, Brazil (Duellman et al. 2016; Venâncio and Melo-Sampaio 2010). Such regions range from 0 - 900 m in elevation, encompassing freshwater wetlands and forests that can receive annual precipitation of 1900 to 2300 mm and warm to hot temperatures. Dry seasons can last from June to October while wet seasons from November to May (Neckel-Oliveira and Wachlevski 2004; Lima et al. 2005; IUCN 2010).
Life History, Abundance, Activity, and Special Behaviors
This species is nocturnal and arboreal (Neckel-Oliveira and Wachlevski 2004).
The frogs can jump, but have been described by Caramaschi and Cruz (2002) to walk slowly on foliage to search for resting places or food, with their bright colors hidden during movement, and aposematically displayed at rest.
Males call from high trees (ranging from 1 - 8 m) and vocalize around 19:00 hours, exhibiting fighting behavior over females (Venâncio and Melo-Sampaio 2010). Phyllomedusa bicolor displays breeding site fidelity, as well as low dispersal and movement patterns (Mota et al. 2020).
Phyllomedusa bicolor possesses three dermal glands: mucous, lipid, and serous glands. The mucous and lipid glands of P. bicolor produce cutaneous secretions for skin homeostasis. Serous glands, meanwhile, produce granules, which start small and fuse into larger granules, meant to store and eventually discharge venom; serous glands are the only integumental glands that contain the peptide I-13, which indicates the presence of biologically active and mature peptides (Barberio et al. 1987; Lacombe et al. 2000). Lacombe et al. (2000) and Venâncio and Melo-Sampaio (2010) both hypothesize that the biologically active peptides produced on the skin of P. bicolor, in addition to a large body size, aid in defense against predators.
Phyllomedusa bicolor follow Haddad and Prado’s reproductive Mode 24 (2005), where eggs hatch as exotrophic tadpoles that fall into still freshwater. A jelly-like mass of eggs with a mean of 859.5 eggs per clutch, is laid by females in leaf nests hanging over water. These nests are created by males roughly folding/joining 2 or 3 leaves and range from 1.4 - 2.8 m above the water (Neckel-Oliveira and Wachlevski 2004; Lima et al. 2005; Mota et al. 2020). Phyllomedusa bicolor has the largest clutch size of the Amazonian arboreal anuran nesters. As anuran eggs are a high-protein energy source, P. bicolor clutches have predators of staphylinid beetles, phorid flies, and mammals; due to the large size of the eggs, P. bicolor egg predation by mammals is unique when compared to P. tarsius and P. tomopterna (Neckel-Oliveira and Wachlevski 2004).
Phyllomedusa bicolor has exotrophic larvae that inhabit non-flowing systems of water. As suspension-raspers, they partially feed through rasping submerged surfaces or filtering suspended particles from water columns (McDiarmid and Altig 1999).
Described at Stage 36 by Pinto et al. (2013), but with information on Stages 24 - 40 available in the text, the total length of the larvae is between 41.32 - 46.17 mm, with the body roughly 40% of that length and the tail roughly 60%. From a lateral view, the body is triangular, and from a dorsal view, the body is elongated. The snout is truncated both laterally and dorsally. The nostrils are small and ovoid, positioned anterolaterally near the snout, with an internarial distance ~30% greater than eye diameter. The larvae have large eyes (a diameter of 2.28 - 2.46 mm), which are laterally positioned and directed. The interorbital distance is ~70% greater than eye diameter. The oral disc, with a width ranging from 4.00 - 4.50 mm, is anteriorly directed and ventrally emarginated. The marginal papillae is uniseriate and elongate, leaving a wide gap on the anterior labium, which is roughly two-thirds of the first anterior labial tooth row. Smaller, submarginal papillae are laterally present on the posterior and anterior labium. The labial tooth row formula is 2(2)/3(1). Anterior rows 1 and 2 are of similar length, while the second posterior row is shorter than the first posterior row, but double the length of the third posterior row. The second anterior row has a wide gap, while first posterior row has a small gap. Additionally, on the first posterior row, there are around 45 labial teeth per millimeter. The upper and lower jaw sheaths are wide with small, triangular-shaped, fine serrations. However, the upper jaw sheath is M-shaped, while lower is V-shaped. On the lower jaw sheath, there are around 27 serrations per millimeter (Pinto et al. 2013).
A posteriorly directed, singular spiracle is nearly ventral and has an absent inner wall. The spiracular opening is large, not visible dorsally, and located below the middle of the body. Attached to the right side of the ventral fin is the larvae’s short vent tube. The ventral fin itself is slightly arched, coming from the body terminus and consistent in height for two-thirds of its anterior, until it gradually tapers to the tip. The dorsal fin originates from the tail-body junction, featuring a unique cord (thickening on the edge of the fin) that is present throughout all stages. The upper fin is generally shallow, but is at its highest point toward the posterior, and extends nearly to the end of the tail. The caudal musculature is broader than the dorsal fin and gradually tapers to a pointed tip. Maximum tail height ranges from 8.00 - 9.44 mm, which is roughly double the maximum height of the tail musculature (Pinto et al. 2013).
In life, the tadpoles have orange dorsums and anterior parts of their body, in addition to a silver belly. The tail musculature and the dorsal fin’s unique cord is a paler orange, and the fins are a translucent orange. Their irises are also silver. In preservation, the light yellow body has a brownish gray dorsum, gray lateral side, and gray belly. The tail musculature and the dorsal fin’s unique cord is pale brown, while the casual musculature’s dorsal surface is brown. Their fins are translucent as well. Through the skin, intestines cannot be observed (Pinto et al. 2013).
From Stages 24 to 40, the larvae undergo changes in labial tooth row formula, vent tube distinction, and skin texture. The labial tooth row formula at Stage 24 is 1(1)/2(1) and 2(2)/3(1) or 2(2)/3(1-2) at Stages 26, 27, 29, and 34. After Stage 24, two anterior tooth rows and three posterior tooth rows stabilize, with posterior gaps only varying at advanced developmental stages. From early developmental stages (Stages 24 - 25), the vent tube is indistinct and the larvae are more slender, less pigmented, and lighter in tail musculature. From Stages 36, 37, 38, and 40, the lateral surfaces develop a grainy texture from the postorbital region to body terminus, with such a texture scarce ventrally (Pinto et al. 2013).
Phyllomedusa bicolor larvae, when compared to other Phyllomedusa larvae, similarly feature triangular bodies with anteroventral mouths and small nostrils near their snout. The larvae also have heavy caudal musculature, ventral fins deeper than the shallow dorsal fin, and a sinistral, nearly ventral spiracle. But, the tadpoles of P. bicolor are distinguishable from all closely related species, except P. vaillanti through their dorsal fin’s unique cord, which is visible at all stages. Phyllomedusa bicolor larvae can also be distinguished from P. bahiana larvae by their smaller size, papillae disposition, and coloration in life (where P. bahiana has marginal and sub-marginal papillae regularly and laterally positioned in double rows, and dorsolaterally on portions of the oral disc, a pale beige body and tail musculature, and very dark brown punctuations, giving a dark appearance, on the dorsal and lateral surfaces). A smaller size and coloration distinguish P. bicolor from Phyllomedusa boliviana, P. tomopterna, P. trinitatis, and P. tetraploidea (where, in life, P. boliviana has gold and black pigments with a transparent dorsum, a transparent belly, and gold lateral surfaces, P. tomopterna has a transparent whitish olive body, silver/yellow/yellowish tan abdomen, whitish/transparent tail with a diffuse orange patch in the middle of the lower fin, P. trinitatis has a distinct cluster of black spots on the ventral fin, which expand to form a black bar across the tail, and P. tetraploidea (unclear if in life) has a rectangular dark spot between its eyes). Phyllomedusa bicolor is differentiated from P. camba by their smaller size, depth of fins, labial tooth row formula, and coloration in life (where P. camba has ventral and dorsal fins of similar depth, a labial tooth row formula of 2(2)/3, dorsally black bodies that become dark bluish gray ventrally). Furthermore, P. bicolor is differentiated from P. vaillanti by their smaller size, labial tooth row formula, size of labial tooth row, and coloration in life (where P. vaillanti has a labial tooth row formula of 2(2)/3 or 2(2)/3(1) with posterior 2 slightly greater than posterior 3, a black spot between the eyes, a gray-greenish body and tail and whitish belly in eastern Brazilian Amazon, and a dull green dorsum, pale green musculature, and white belly in Peru) (Pinto et al. 2013). Larger size, vent tube size and position, and coloration in life separates P. bicolor from P. atelopoides (where P. atelopoides has a longer vent tube that is sinistral to the ventral fin, a black brownish body, silvery blue belly, and pale lime green snout). Phyllomedusa bicolor larvae have a larger size, distribution of submarginal papillae, and coloration when compared to P. iheringi (where P. iheringi does not have submarginal papillae, but does feature four dark spots and a trapezoidal blotch on its head). The size of their labial tooth row distinguishes P. bicolor from P. burmeisteri and P. distincta (where in both species the second posterior tooth row is more than double the length of third posterior). Labial tooth row formula and distribution of marginal papillae separates P. bicolor from P. hypochondrialis (where P. hypochondrialis, in Venezuela, has a labial tooth row formula of 2(2)/3 and a gap in the marginal papillae of its posterior labium). In addition, P. bicolor is separated from P. palliata by their labial tooth row formula and coloration in life (where P. palliata has a labial tooth row formula of 2(2)/3 and bluish gray body and tail with a iridescent bluish green sheen on its belly). Only coloration in life distinguishes P. bicolor from P. tarsius (where P. tarsius has a transparent olive/whitish body and a silver/pale metallic green belly). Lastly, P. bicolor is differentiated from P. venusta by the size of papillae gap (where P. venusta has a gap between papillae that is roughly one-third of the first anterior row) (Pinto et al. 2013).
Regarding development, Neckel-Oliveira and Wachlevski (2004) observed that embryonic development was quick, lasting 10.9 - 12.1 days, before the larvae hatch with external gills, falling into the water below, and developing into aquatic tadpoles.
The development of the mucous, serous, and lipid glands were also observed by Lacombe et al. (2000) on the P. bicolor tadpole skin. The three glands appear as small granular structures in pigmented layers that are distinct on the dorsal skin. The structures, particularly the serous glands, develop into larger acini later. The lipid glands develop later during ontogenesis (Lacombe et al. 2000).
Trends and Threats
Phyllomedusa bicolor is threatened by logging and wood harvesting, but conservation sites have been established in the species’ distribution (IUCN 2010).
Relation to Humans
The secretions of P. bicolor are commonly referred to as Sapo or Kambô and utilized in hunting rituals by indigenous people from the Southwestern Amazon basin, in particular Brazilians and the Matses Indians of Peru, (Erspamer et al. 1993; Aquila et al. 2017; Vega et al. 2020; Majić et al. 2021). The secretion is meant as a remedy to panema (bad luck in hunting) by improving luck and strength, in addition to simply being used for salutogenic or detoxification purposes (Aquila et al. 2017; Majić et al. 2021). To collect such secretions, P. bicolor is usually caught and kept for three days; its back and limbs are gently and periodically scraped with a stick, which is placed in a leaf bag over a fire, then used again to keep scrapping. After three days, the frog is respectfully released and the stick is covered in a yellow substance, which is mixed with saliva to be used and applied to intentionally burned wounds (Erspamer et al. 1993; Aquila et al. 2017). Kambô has gained popularity in alternative therapy clinics by Western users (Mota et al. 2020; Majić et al. 2021). Phyllomedusa bicolor’s secretion itself contains up to 16 bioactive peptides which can function as opioids, vasoactive molecules, antimicrobials, antiprotozoals, or μ-opioid receptors with analgesic properties (Aquila et al. 2017; Vega et al. 2020; Majić et al. 2021). These biopeptides potentially affect the cardiovascular system of users, causing intense cardiovascular and gastrointestinal symptoms (ranging from vomiting, hot flashes, diarrhea, racing heart, nausea, sweating, etc.) and mild psychoactive effects, as provided by anecdotal observations (notably a feeling that the spirit of the frog is with them) (Erspamer et al. 1993; Majić et al. 2021). Hypotheses suggest that ceruletide and sauvagine may cause central effects, while ceruletide, phyllokinin, phyllomedusin, sauvagine and opioid peptides cause peripheral effects (Majić et al. 2021). Majić et al. (2021) believe that more research is indeed to investigate the effects of the secretion and determine any possible health concerns.
Two studies, one utilizing four mitochondrial genes and five nuclear genes (Faivovich et al. 2005) and another utilizing four mitochondrial genes and three nuclear genes (Wiens et al. 2006) determined that the Phyllomedusa genus is monophyletic. This was later supported by Duellman et al. (2016) through analysis of two mitochondrial genes and ten addition nuclear loci. Faivovich et al. (2005), further found that P. bicolor and P. vallanti are sister taxa, which is supported by the fact that both feature osteoderms (dermal ossifications) that protrude beyond the skin (Ruibal and Shoemaker 1984; Faivovich et al. 2010). However, as Faivovich et al. (2005), Wiens et al. (2006), and Duellman et al. (2016) were more broad scale analyses, it is unclear which species is unequivocally the most closely related to P. bicolor.
Mota et al. (2020), on the other hand, used nucleotide sequences for the genes 16S, Cytb, and RAG-1 to determine that P. bicolor likely has two cryptically divergent lineages corresponding to a geographic division in the eastern and western regions of Amazonia, in addition to the Amazon’s well-defined climate gradient. The eastern Amazonia likely is compromised of populations in the Guiana Shield and the Brazilian Shield.
The name of the genus Phyllomedusa is derived from the Greek, “phyllo”, which means “leaf/foliage,” and “medousa,” which means “queen/protector.” Therefore, Phyllomedusa means “queen/protector of the foliage” and alludes to the green color and large size of members of the genus (Caramaschi and Cruz 2002).
Boddaert (1772) originally named Phyllomedusa bicolor as Rana bicolor until Wagler (1830) proposed the species be placed in Phyllomedusa (Caramaschi and Cruz 2002).
In analysis of the ultrastructure sperm of P. bicolor, as compared to P. hypochondrialis and P. tarsius, P. bicolor was determined to have similar structures to P. tarsius. Relative to P. hypochondrialis, both species had a large spermatozoon with a reduced nuclear space, an abruptly ending anterior region in the nucleus, and a larger axial fibre; the subacrosomal cones, which fill the space between the acrosome vesicle and nucleus, are also transversely less electron-dense and more granular (Costa et al. 2005).
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Originally submitted by: Hong Nguyen (2023-03-01)
Trends and threats by: Hong Nguyen
Relation to humans by: Hong Nguyen
Edited by: Kellie Whittaker, Ann T. Chang (2023-03-01)
Species Account Citation: AmphibiaWeb 2023 Phyllomedusa bicolor: Waxy-Monkey Treefrog; Giant Monkey Frog; Dow Kiet; Kambô Frog; Sapo; Amazonian Giant Maki Frog <https://amphibiaweb.org/species/642> University of California, Berkeley, CA, USA. Accessed Jun 10, 2023.
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Citation: AmphibiaWeb. 2023. <https://amphibiaweb.org> University of California, Berkeley, CA, USA. Accessed 10 Jun 2023.
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