The golden frog is active during the day (3) and lives in groups usually consisting of twice as many males as females (4). It is an insectivorous species, feeding on termites, fruit flies, ants and a huge range of other insects (4). Breeding tends to start after the first heavy rains of the year, and when there is plenty of food. Males attract females with their call (4). The male will then rapidly move himself to the female's back, without embracing her (virtual or cephalic amplexus) (6). The females do not lay their eggs in water, but in damp leaf litter, moss or under bark and rocks next to a water source (4). Each clutch contains 20 - 60 white eggs, each one measuring up to 2 mm in diameter (3); they are fertilised by the male immediately after laying (4). Two weeks later the tadpoles hatch out and they either wriggle into water or are are washed into small pools by heavy rain. It takes around 70 days for the tadpoles to metamorphose into froglets which measure 11 mm in length (3). The typical yellow-red colouration is acquired only after some weeks (6). Sexual maturity is reached 12 to 14 months later, and the average life span is eight years (4).

The golden frog is a small poisonous frog which is bright yellow, orange or red with some red colouration on the upper surface of the hind legs (3). It is a member of the Madagascan genus Mantella, the members of which have evolved to be very similar in appearance and behaviour to the very distantly related poison arrow frogs of South America (family Dendrobatidae) (3). In this species, the eyes are typically black, although there may occasionally be golden pigmentation in the upper portion of the iris (3). The legs are short, and the tips of the fingers and toes bear distinct adhesive pads (4). Males are typically smaller than females and have a more angular body shape. The bellies of males are generally lighter in colour than those of females. Two pale-coloured ducts that carry sperm and urine are often visible passing along the belly (4). Males do not call as often as other species of mantella frogs (4); the call is composed of a series of short notes, each of which includes three short clicks (3). The tadpoles of this species do not have external gills, and the eyes are located on the top of the head. Young froglets are olive green in colour with dark marks on the back and the hind limbs feature dark bands (3).

A generally rather small and stout Mantella, 19-24 mm, females rarely can reach up to 31 mm. Dorsally uniformly yellow-orange, in some populations red-orange, often with a translucent shade. Bright red flashmarks present. Iris nearly uniformly black, only a little light pigment in its upper part. Ventrally uniform, similar to dorsal surface but generally somewhat lighter, except red flashmark (extended nearly on the whole tibia). Some inner organs visible through the slightly transparent ventral skin. Similar species: The colouration of M. aurantiaca is unique.

Taken with permission from Glaw and Vences (2007).

Featured in Amazing Amphibians on 29 July 2013

A generally rather small and stout Mantella, 19-24 mm, females rarely can reach up to 31 mm. Dorsally uniformly yellow-orange, in some populations red-orange, often with a translucent shade. Bright red flashmarks present. Iris nearly uniformly black, only a little light pigment in its upper part. Ventrally uniform, similar to dorsal surface but generally somewhat lighter, except red flashmark (extended nearly on the whole tibia). Some inner organs visible through the slightly transparent ventral skin. Similar species: The colouration of M. aurantiaca is unique.

Taken with permission from Glaw and Vences (2007).

Featured in Amazing Amphibians on 29 July 2013

This species has a very restricted distribution in east-central Madagascar, centred on the Torotorofotsy area (c. 7km north-west of Andasibe) and the Andromena Forest at the Samarirana River. Several other small forest fragments north and south of Moramanga are populated by the species as well. Its recorded altitudinal range is 920-960 m asl.

Andranomandry, Andranomena, Torotorofotsy. It occurs between 920-960m aslin primary and secondary rainforest and usually damp, swampy areas, often associated with screw pine (Pandanus) forest (Vences et al. 2008).

Mantella aurantiaca occupy the montane pandanus forests around Andasibe and they are typically found in isolated patches ranging throughout southeastern Madagascar.

Biogeographic Regions: ethiopian (Native )

Other Geographic Terms: island endemic

Andranomandry, Andranomena, Torotorofotsy. It occurs between 920-960m aslin primary and secondary rainforest and usually damp, swampy areas, often associated with screw pine (Pandanus) forest (Vences et al. 2008).

This frog is endemic to central-eastern Madagascar (6), where it occurs in a highly restricted area at elevations over 900m (3).

Adult snout vent lengths (SVLs) are typically 1.25 inches, although 1.5 inch SVLs have been reported. These frogs exhibit brilliant golden-orange coloration that is impossible to miss. They occasionally have red flash marks on the inner portion of the hind legs. The eyes of this species are jet black. The legs are very short with distinct adhesive disks found on the fingers and toes (Badger, 1995). There is sexual dimorphism. Males are generally smaller, slimmer and more angular in build than females, and tend not to call as much as other species of male mantellas. The male's ventral surface is lighter in color and therefore causes the seminiferous ducts (narrow pair of pale lines) to be visible. These ducts hava a dual purpose in males, carrying both sperm and urine. Females also have these ducts but they are, for the most part, concealed by the uterus and oviducts. The ducts in females do not carry sperm but they still function in urine transportation (Staniszewski, 1997).

Average length: 3.175 cm.

Other Physical Features: ectothermic ; heterothermic ; bilateral symmetry ; poisonous

Sexual Dimorphism: female larger; sexes shaped differently

Habitat and Ecology

Systems

• Terrestrial
• Freshwater

Golden mantellas are considered an "upland" species, due to the fact that they are found at an altitude of about 900 meters. The climate is typically moist, humid and temperate. They usually inhabit mossy or grassy mounds of forest debris that border shallow swampy waters.

Range elevation: 900 (high) m.

Habitat Regions: tropical ; terrestrial

Terrestrial Biomes: forest ; rainforest

Wetlands: swamp

Inhabits Pandanus forests where it is found in sunny areas. They tend to occur amongst vegetation in swampy sites (3).

Mantella aurantiaca are entirely insectivorous. A diet commonly consists of termites (Isoptera), ants (Formicidae), fruitflies (Drosphila), and just about any other arthropod that can be fit into the mouth. Golden mantellas are known for attempting to eat anything, even if the taste is repulsive (Bartlett, 1996).

Animal Foods: insects; terrestrial non-insect arthropods

Primary Diet: carnivore (Insectivore )

The brilliant colors exhibited by M. aurantiaca can be attributed to a phenomenon called aposematic coloration, where toxic or dangerous animals use bright colors or marking to advertise their toxicity to potential predators. Golden mantellas have toxic skin secretions, protecting them from most predators.

Anti-predator Adaptations: aposematic

Golden mantellas use auditory cues, and may also use visual or chemical cues to communicate. They use their vision to locate prey.

Communication Channels: tactile ; acoustic

Perception Channels: visual ; acoustic ; vibrations

Eggs undergo rapid terrestrial development (2-6 day hatch period) and the newly hatched tadpoles either wriggle to the nearby water source or are washed into the water by storms (Bartlett, 1996). Tadpoles typically metamorphose into froglets 6 to 8 weeks after hatching. The tadpoles are primarily herbivores, feeding on algae and detritus, although some meat matter may be incorporated into the diet. Once the tadpoles become froglets, they are usually 10 to 14 mm in SVL and begin feeding on the more typical adult insect (springtails and small aphids) (Staniszewski. 1997). Sexual maturity is reached in 12 to 14 months.

Development - Life Cycle: metamorphosis

Mantella aurantiaca typically has a life span of 8 years.

Average lifespan

Status: wild: 8 years.

Average lifespan

Sex: female

Status: captivity: 3.7 years.

Maximum longevity: 10.9 years (captivity)

Males use a call consisting of a series of short notes, with three clicks per note, to attract female mates. Territorial aggression does occur in both sexes during this time, but especially in males. Intruders are sometimes grabbed around the upper body or head and are typically pushed away. The actual courtship process of this species is rather secretive and usually takes place under bark, logs, or rocks. If a non-gravid femaile is amplexed, she will flick her legs and back flip until the male releases her.

Mating System: polygynandrous (promiscuous)

Mating usually occurs when there has been an abundant amount of food available and when the first substantial rain comes. Once amplexus is successfully underway, a suitable egg-laying site will be searched for. These sites usually include damp moss, crevices in logs, underneath damp bark or rocks and are always adjacent to a water source (Staniszewski, 1997). Clutches consist of 12 to 30 eggs, 2 to 3 mm in diameter and are immediately fertilized by a male, although fertilization can occur up to 2 days later and by multiple males.

Breeding interval: Breeding occurs when food is abundant, depending on rainfall.

Breeding season: Breeding is opportunistic, occurring when conditions become favorable.

Range number of offspring: 12 to 30.

Range age at sexual or reproductive maturity (female): 12 to 14 months.

Range age at sexual or reproductive maturity (male): 12 to 14 months.

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

Once the eggs are laid, parent golden mantellas have no further involvement in the development of their young.

Parental Investment: pre-fertilization (Provisioning, Protecting: Female)

Year Assessed

Assessor/s

Reviewer/s

Contributor/s

Several human-caused factors are causing a decline in native populations of the golden mantella. Overcollection for the pet industry, introduced predatory species, major deforestation in Madagascar and human encroachment are all among the leading causes fueling this raging decline.

CITES: appendix ii

IUCN Red List of Threatened Species: critically endangered

Classified as Vulnerable (VU A1cd) by the IUCN Red List 2003 (1) and listed on Appendix II of CITES (3). A revision of IUCN categorisation for the Malagasy amphibians led to inclusion of Mantella aurantiaca in the “critically endangered” category (Global Amphibian Assessment, Madagascar Working Group, unpublished) (6).

Population

Population Trend

The occurrence of this species is correlated with humid Pandanus forest, where it can be found in sun-exposed sites (Glaw and Vences 2006). Here Mantella aurantiaca can be heard calling from the ground, among vegetation in swampy areas (Glaw and Vences 2006). It is active during the day (Blommers-Schlösser and Blanc 1991; Glaw and Vences, pers. obs.).

Calls: Irregular series of short chirping notes. The call (from Andasibe) consists of a series of short notes (duration 50-60 ms). An oscillogram of one note shows that it consists of 3 very short clicks. Frequency ranges from about 4.5-6.5 kHz, dominant frequency is about 5.3kHz. Another analysis confirms that notes consist of 3 clicks, but shows a frequency band between 3 and 7 kHz (Glaw and Vences 1994; Ahl 1929). The call has also been described as an irregular succession of short chirps (Glaw and Vences 2006).

Eggs and tadpoles (from near Andasibe): Eggs are deposited in moist leaf litter outside of water. One clutch consists of 20-60 whitish eggs (diameter 1.5-2mm). Embryogenesis lasts 14 days and the tadpoles are flooded into small pools by heavy rain. The tadpoles develop within about 70 days into froglets measuring 11 mm (Glaw and Vences 1994). Frogs of the genus Mantella are sexually mature within a year after metamorphosis and thus have short generation times (Glaw et al. 2000).

The adult colouration is probably aposematic, as M. aurantiaca contains a variety of toxins in the skin, including pumiliotoxin, allopumiliotoxin and homopumiliotoxin alkaloids, as well as pyrrolizidines, indolizidines and quinolizidines (Daly et al. 1984, 1996; Garraffo et al. 1993). This toxicity is derived from dietary sources, as is the case with dendrobatid frogs (Daly et al. 1997). Captive-bred M. aurantiaca lack toxicity when fed non-toxic arthropods, but readily accumulate alkaloids when fed alkaloid-dusted fruit flies (Daly et al. 1997).

The occurrence of this species is correlated with humid Pandanus forest, where it can be found in sun-exposed sites (Glaw and Vences 2006). Here Mantella aurantiaca can be heard calling from the ground, among vegetation in swampy areas (Glaw and Vences 2006). It is active during the day (Blommers-Schlösser and Blanc 1991; Glaw and Vences, pers. obs.).

Calls: Irregular series of short chirping notes. The call (from Andasibe) consists of a series of short notes (duration 50-60 ms). An oscillogram of one note shows that it consists of 3 very short clicks. Frequency ranges from about 4.5-6.5 kHz, dominant frequency is about 5.3kHz. Another analysis confirms that notes consist of 3 clicks, but shows a frequency band between 3 and 7 kHz (Glaw and Vences 1994; Ahl 1929). The call has also been described as an irregular succession of short chirps (Glaw and Vences 2006).

Eggs and tadpoles (from near Andasibe): Eggs are deposited in moist leaf litter outside of water. One clutch consists of 20-60 whitish eggs (diameter 1.5-2mm). Embryogenesis lasts 14 days and the tadpoles are flooded into small pools by heavy rain. The tadpoles develop within about 70 days into froglets measuring 11 mm (Glaw and Vences 1994). Frogs of the genus Mantella are sexually mature within a year after metamorphosis and thus have short generation times (Glaw et al. 2000).

The adult colouration is probably aposematic, as M. aurantiaca contains a variety of toxins in the skin, including pumiliotoxin, allopumiliotoxin and homopumiliotoxin alkaloids, as well as pyrrolizidines, indolizidines and quinolizidines (Daly et al. 1984, 1996; Garraffo et al. 1993). This toxicity is derived from dietary sources, as is the case with dendrobatid frogs (Daly et al. 1997). Captive-bred M. aurantiaca lack toxicity when fed non-toxic arthropods, but readily accumulate alkaloids when fed alkaloid-dusted fruit flies (Daly et al. 1997).

Major Threats

Critically Endangered: area of occupancy is probably less than 10km². Its distribution is severely fragmented, and the extent of its forest habitat in east-central Madagascar is declining, and the number of mature individuals might also be declining through over-exploitation.It does not occur in protected areas, but it is found near to the Réserve Spéciale d’Analamazaotra. This species is being maintained in captivity by about 35 zoos and other institutions and is being bred in captivity by public institutions and many private individuals (Vences et al. 2008).

Critically Endangered: area of occupancy is probably less than 10km². Its distribution is severely fragmented, and the extent of its forest habitat in east-central Madagascar is declining, and the number of mature individuals might also be declining through over-exploitation.It does not occur in protected areas, but it is found near to the Réserve Spéciale d’Analamazaotra. This species is being maintained in captivity by about 35 zoos and other institutions and is being bred in captivity by public institutions and many private individuals (Vences et al. 2008).

This species is very popular in the pet-trade and over-collection of individuals is still carried out at some areas. Although at present there are no signs of reduction of population size due to this take off, it must be carefully monitored in the future to assure the survivorship of the species (6). Furthermore, large-scale deforestation, predation by introduced species, and encroachment by humans are all posing threats to this frog (4).

Conservation Actions

All mantella frogs are listed on Appendix II of the Convention on International Trade in Endangered Species (CITES), which controls international trade in these species (2). The golden frog is bred in captivity in a number of zoos and other breeding facilities, including the Detroit Zoological Institute's National Amphibian Conservation Centre, which breeds hundreds each year for use in breeding programmes in other institutions (5). The provision of captive-bred individuals to zoos reduces the stress on wild populations caused by collection. Captive breeding programmes may also provide a genetic reservoir of a species to safeguard it should the population undergo a drastic decline or even become extinct; they also provide sources of animals for reintroduction to the wild and allow research to be carried out (5). Although captive breeding is an important facet of any conservation programme, protection of the remaining wild populations and the habitat on which they depend is of great importance (7).

Golden mantellas are toxic, although humans would not typically be exposed to this toxin if they are not harassing these frogs.

Negative Impacts: injures humans (poisonous )

Many individuals are captured from the wild for the pet trade while some are now captive bred.

This species is collected for the pet trade and is commonly bred in captivity (Glaw et al. 2000). Commercial export of M. aurantiaca began in the late 1980s or early 1990s, but was halted in 2002 (CITES Secretariat 2008). Import of wild-caught specimens to the European Union has been banned since 2006 (CITES Secretariat 2008). There is thought to be little illegal trade of wild-caught specimens owing to the low price paid to Malagasy collectors plus the high sensitivity of mantellas to transport conditions (CITES Secretariat 2008).

This species is collected for the pet trade and is commonly bred in captivity (Glaw et al. 2000). Commercial export of M. aurantiaca began in the late 1980s or early 1990s, but was halted in 2002 (CITES Secretariat 2008). Import of wild-caught specimens to the European Union has been banned since 2006 (CITES Secretariat 2008). There is thought to be little illegal trade of wild-caught specimens owing to the low price paid to Malagasy collectors plus the high sensitivity of mantellas to transport conditions (CITES Secretariat 2008).