Cryptoprocta spelea
Encyclopedia
Cryptoprocta spelea, also known as the giant fossa, is an extinct
species of carnivore from Madagascar
in the family Eupleridae
, which is most closely related to the mongoose
s and includes all Malagasy carnivorans. It was first described in 1902, and in 1935 was recognized as a separate species from its closest relative, the living fossa
(Cryptoprocta ferox). C. spelea is larger than the fossa, but otherwise similar. The two have not always been accepted as distinct species. When and how the larger form went extinct is unknown; there is some anecdotal evidence, including reports of very large fossas, that there is more than one surviving species.
The species is known from subfossil
bones found in a variety of caves in northern, western, southern, and central Madagascar. In some sites, it occurs with remains of C. ferox, but there is no evidence that the two lived at the same time. Living species of comparably-sized, related carnivores in other regions manage to coexist, suggesting that the same may have happened with both C. spelea and C. ferox. C. spelea would have been able to prey on larger animals than its smaller relative could have, including the recently extinct giant lemurs
.
described subfossil
carnivoran remains from two caves on Madagascar as a larger "variety" of the living fossa
(Cryptoprocta ferox), C. ferox var. spelea. G. Petit, writing in 1935, considered spelea to represent a distinct species. Charles Lamberton reviewed subfossil and living Cryptoprocta in 1939 and agreed with Petit in recognizing two species, naming this species from a specimen found at Ankazoabo Cave near Itampolo
. The specific name spelea means "cave" and was given because of the location of its discovery. However, Lamberton apparently had at most three skeletons of the living fossa, not nearly enough to capture the range of variation in that species, and some later authors did not separate C. spelea and C. ferox as species. Steven Goodman
and colleagues, using larger samples, compiled another set of Cryptoprocta measurements that was published in a 2004 article. They found that some subfossil Cryptoprocta fell outside the range of variation of living C. ferox, and identified those as representing C. spelea. Grandidier had not designated a type specimen for the species, and to maintain C. spelea as the name for the larger form of the fossa, Goodman and colleagues designated a specimen to serve as the type specimen (specifically, a neotype).
Lamberton recognized a third species, Cryptoprocta antamba, on the basis of a mandible
(lower jaw) with abnormally broad spacing between the condyloid process
es at the back. He also referred two femora
(upper leg bones) and a tibia
(lower leg bone) intermediate in size between C. spelea and C. ferox to this species. The specific name refers to the "antamba", an animal allegedly from southern Madagascar described by Étienne de Flacourt
in 1658 as a large, rare, leopard-like carnivore that eats men and calves and lives in remote mountainous areas; it may have been the giant fossa. Goodman and colleagues could not locate Lamberton's material of Cryptoprocta antamba, but suggested that it was based on an abnormal C. spelea. Together, the fossa and C. spelea form the genus Cryptoprocta within the family Eupleridae
, which also includes the other Malagasy carnivora
ns—the falanouc
, the fanaloka
s, and the Galidiinae
. DNA sequence
studies suggest that the Eupleridae form a single natural (monophyletic) group and are most closely related to the mongoose
s of Eurasia and mainland Africa.
account of the fossa, Michael Köhncke and Klaus Leonhardt wrote that the two were morphologically identical. However, remains of C. spelea are larger than any living C. ferox. Goodman and colleagues found that skull measurements in specimens they identified as C. spelea were 1.07 to 1.32 times as large as in adult C. ferox, and postcrania
l measurements were 1.19 to 1.37 times as large. The only specimen of C. spelea in which condylobasal length (a measure of total skull length) could be ascertained measured 153.4 mm (6.04 in), compared to a range of 114.5 to 133.3 mm (4.51 to 5.25 in) in adult C. ferox. Humerus
(upper arm bone) length in twelve C. spelea is 122.7 to 146.8 mm (4.83 to 5.78 in), averaging 137.9 mm (5.43 in), compared to 108.5 to 127.5 mm (4.27 to 5.02 in), averaging 116.1 mm (4.57 in), in the extant fossa. Body mass estimates for C. spelea range from 17 kg (37 lb) to 20 kg (44 lb), and it was among the largest carnivores of the island. By comparison, adult C. ferox range from 5 kg (11 lb) to 10 kg (22 lb).
Cryptoprocta spelea is the only extinct member of the order Carnivora
known from Madagascar; recently extinct Madagascan animals also include at least 17 species of lemur
s, most of which are larger than the living forms, as well as elephant bird
s and Malagasy hippopotamus
es, among others. Subfossil remains of the giant fossa have been found in Holocene
cave sites from the northern end of Madagascar along the west coast to the far south, and in the central highlands. Some sites have yielded both C. spelea and smaller remains referable to the living species, C. ferox; however, lack of robust stratigraphic
knowledge and no available radiocarbon dating
on subfossil Cryptoprocta bones makes it uncertain whether the two species lived in the same region at the same time. The size ratio between the two species is within the range of ratios seen between similar-sized living cats
and mongooses found in the same areas, suggesting that the two species may have been able to occur together.
With its large size and massive jaws and teeth, C. spelea was a formidable, "puma-like" predator, and in addition to smaller lemurids, it may have eaten some of the big, now extinct subfossil lemur
s that would have been too large for C. ferox. No subfossil evidence has been found to definitively show that lemurs were its prey; this assumption is based on the diet of the smaller, extant species of fossa. Other possible prey include tenrecs, smaller euplerids, and even young Malagasy hippopotamus
es. Its extinction may have changed predation dynamics on Madagascar.
The IUCN Red List
currently lists C. spelea as an extinct species; why and when it went extinct remains unknown. However, local people on Madagascar often recognize two forms of fossa, a larger fosa mainty (or "black Cryptoprocta") and a smaller fosa mena (or "reddish Cryptoprocta"). There are also some anecdotal records of very large living fossas, such as a 2-m (7 ft), 30-kg (70 lb) fossa at Morondava
. Goodman and colleagues suggested that further research may demonstrate that there is more than one species of fossa yet alive.
Extinction
In biology and ecology, extinction is the end of an organism or of a group of organisms , normally a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and recover may have been lost before this point...
species of carnivore from Madagascar
Madagascar
The Republic of Madagascar is an island country located in the Indian Ocean off the southeastern coast of Africa...
in the family Eupleridae
Eupleridae
The family Eupleridae is a group of carnivorans endemic to Madagascar and comprising 10 known species in seven genera. Probably the best known species is the Fossa , in the subfamily Euplerinae...
, which is most closely related to the mongoose
Mongoose
Mongoose are a family of 33 living species of small carnivorans from southern Eurasia and mainland Africa. Four additional species from Madagascar in the subfamily Galidiinae, which were previously classified in this family, are also referred to as "mongooses" or "mongoose-like"...
s and includes all Malagasy carnivorans. It was first described in 1902, and in 1935 was recognized as a separate species from its closest relative, the living fossa
Fossa (animal)
The fossa is a cat-like, carnivorous mammal that is endemic to Madagascar. It is a member of the Eupleridae, a family of carnivorans closely related to the mongoose family . Its classification has been controversial because its physical traits resemble those of cats, yet other traits suggest a...
(Cryptoprocta ferox). C. spelea is larger than the fossa, but otherwise similar. The two have not always been accepted as distinct species. When and how the larger form went extinct is unknown; there is some anecdotal evidence, including reports of very large fossas, that there is more than one surviving species.
The species is known from subfossil
Subfossil
Subfossil refers to remains whose fossilization process is not complete, either for lack of time or because the conditions in which they were buried were not optimal for fossilization....
bones found in a variety of caves in northern, western, southern, and central Madagascar. In some sites, it occurs with remains of C. ferox, but there is no evidence that the two lived at the same time. Living species of comparably-sized, related carnivores in other regions manage to coexist, suggesting that the same may have happened with both C. spelea and C. ferox. C. spelea would have been able to prey on larger animals than its smaller relative could have, including the recently extinct giant lemurs
Subfossil lemur
Subfossil lemurs are lemurs from Madagascar that are represented by recent remains dating from nearly 26,000 years ago to approximately 560 years ago. They include both living and extinct species, although the term more frequently refers to the extinct giant lemurs...
.
Taxonomy
In 1902, Guillaume GrandidierGuillaume Grandidier
Guillaume Grandidier was a French geographer , ethnologist , zoologist who studied the island of Madagascar.He was the son of the wealthy industrialist Alfred Grandidier also a zoologist and expert on Madagascar...
described subfossil
Subfossil
Subfossil refers to remains whose fossilization process is not complete, either for lack of time or because the conditions in which they were buried were not optimal for fossilization....
carnivoran remains from two caves on Madagascar as a larger "variety" of the living fossa
Fossa (animal)
The fossa is a cat-like, carnivorous mammal that is endemic to Madagascar. It is a member of the Eupleridae, a family of carnivorans closely related to the mongoose family . Its classification has been controversial because its physical traits resemble those of cats, yet other traits suggest a...
(Cryptoprocta ferox), C. ferox var. spelea. G. Petit, writing in 1935, considered spelea to represent a distinct species. Charles Lamberton reviewed subfossil and living Cryptoprocta in 1939 and agreed with Petit in recognizing two species, naming this species from a specimen found at Ankazoabo Cave near Itampolo
Itampolo
Itampolo is a town and commune in southwestern Madagascar. It belongs to the district of Ampanihy, which is a part of Atsimo-Andrefana Region. The population of the commune was estimated to be approximately 32,000 in 2001 commune census....
. The specific name spelea means "cave" and was given because of the location of its discovery. However, Lamberton apparently had at most three skeletons of the living fossa, not nearly enough to capture the range of variation in that species, and some later authors did not separate C. spelea and C. ferox as species. Steven Goodman
Steven Goodman
Steven Goodman is an American Conservation Biologist, and field biologist on staff in the Department of Zoology at the Field Museum of Natural History....
and colleagues, using larger samples, compiled another set of Cryptoprocta measurements that was published in a 2004 article. They found that some subfossil Cryptoprocta fell outside the range of variation of living C. ferox, and identified those as representing C. spelea. Grandidier had not designated a type specimen for the species, and to maintain C. spelea as the name for the larger form of the fossa, Goodman and colleagues designated a specimen to serve as the type specimen (specifically, a neotype).
Lamberton recognized a third species, Cryptoprocta antamba, on the basis of a mandible
Mandible
The mandible pronunciation or inferior maxillary bone forms the lower jaw and holds the lower teeth in place...
(lower jaw) with abnormally broad spacing between the condyloid process
Condyloid process
The condyloid process is part of the mandible and is thicker than the coronoid, and consists of two portions: the condyle, and the constricted portion which supports it, the neck.-Condyle :...
es at the back. He also referred two femora
Femur
The femur , or thigh bone, is the most proximal bone of the leg in tetrapod vertebrates capable of walking or jumping, such as most land mammals, birds, many reptiles such as lizards, and amphibians such as frogs. In vertebrates with four legs such as dogs and horses, the femur is found only in...
(upper leg bones) and a tibia
Tibia
The tibia , shinbone, or shankbone is the larger and stronger of the two bones in the leg below the knee in vertebrates , and connects the knee with the ankle bones....
(lower leg bone) intermediate in size between C. spelea and C. ferox to this species. The specific name refers to the "antamba", an animal allegedly from southern Madagascar described by Étienne de Flacourt
Étienne de Flacourt
Étienne de Flacourt was a French governor of Madagascar, born at Orléans in 1607. He was named governor of Madagascar by the French East India Company in 1648....
in 1658 as a large, rare, leopard-like carnivore that eats men and calves and lives in remote mountainous areas; it may have been the giant fossa. Goodman and colleagues could not locate Lamberton's material of Cryptoprocta antamba, but suggested that it was based on an abnormal C. spelea. Together, the fossa and C. spelea form the genus Cryptoprocta within the family Eupleridae
Eupleridae
The family Eupleridae is a group of carnivorans endemic to Madagascar and comprising 10 known species in seven genera. Probably the best known species is the Fossa , in the subfamily Euplerinae...
, which also includes the other Malagasy carnivora
Carnivora
The diverse order Carnivora |Latin]] carō "flesh", + vorāre "to devour") includes over 260 species of placental mammals. Its members are formally referred to as carnivorans, while the word "carnivore" can refer to any meat-eating animal...
ns—the falanouc
Falanouc
The Falanouc is a rare mongoose-like mammal endemic Malagasy euplerid .It is classified alongside its closest living relative, the Fanaloka, in the subfamily Euplerinae. The Falanouc has several peculiarities which merit its independent classification...
, the fanaloka
Fanaloka
The Malagasy or striped civet , also known as the fanaloka , is a rare euplerid endemic to Madagascar....
s, and the Galidiinae
Galidiinae
Galidiinae is a subfamily of carnivorans restricted to Madagascar which includes six species classified into four genera. Together with the three other species of indigenous Malagasy carnivorans, including the fossa, they are currently classified in the family Eupleridae within the suborder...
. DNA sequence
DNA sequence
The sequence or primary structure of a nucleic acid is the composition of atoms that make up the nucleic acid and the chemical bonds that bond those atoms. Because nucleic acids, such as DNA and RNA, are unbranched polymers, this specification is equivalent to specifying the sequence of...
studies suggest that the Eupleridae form a single natural (monophyletic) group and are most closely related to the mongoose
Mongoose
Mongoose are a family of 33 living species of small carnivorans from southern Eurasia and mainland Africa. Four additional species from Madagascar in the subfamily Galidiinae, which were previously classified in this family, are also referred to as "mongooses" or "mongoose-like"...
s of Eurasia and mainland Africa.
Description
Although some morphological differences between the two fossa species have been described, these may be allometric (growth-related), and in their 1986 Mammalian SpeciesMammalian Species
Mammalian Species is a peer-reviewed scientific journal published by the American Society of Mammalogists. The journal publishes accounts of 25–35 mammal species yearly. The articles summarize the current literature about each mammal and its systematics, genetics, fossil history, distribution,...
account of the fossa, Michael Köhncke and Klaus Leonhardt wrote that the two were morphologically identical. However, remains of C. spelea are larger than any living C. ferox. Goodman and colleagues found that skull measurements in specimens they identified as C. spelea were 1.07 to 1.32 times as large as in adult C. ferox, and postcrania
Postcrania
Postcrania[p] in zoology and vertebrate paleontology refers to all or part of the skeleton apart from the skull. Frequently, fossil remains, e.g...
l measurements were 1.19 to 1.37 times as large. The only specimen of C. spelea in which condylobasal length (a measure of total skull length) could be ascertained measured 153.4 mm (6.04 in), compared to a range of 114.5 to 133.3 mm (4.51 to 5.25 in) in adult C. ferox. Humerus
Humerus
The humerus is a long bone in the arm or forelimb that runs from the shoulder to the elbow....
(upper arm bone) length in twelve C. spelea is 122.7 to 146.8 mm (4.83 to 5.78 in), averaging 137.9 mm (5.43 in), compared to 108.5 to 127.5 mm (4.27 to 5.02 in), averaging 116.1 mm (4.57 in), in the extant fossa. Body mass estimates for C. spelea range from 17 kg (37 lb) to 20 kg (44 lb), and it was among the largest carnivores of the island. By comparison, adult C. ferox range from 5 kg (11 lb) to 10 kg (22 lb).
Distribution, ecology, and extinction
| Site | spe. | fer. |
|---|---|---|
| Ampasambazimba | + | |
| Ankarana | + | + |
| Ankazoabo Ankazoabo Ankazoabo is a town in Atsimo-Andrefana Region, Madagascar.... |
+ | |
| Antsirabe Antsirabe Antsirabe is the third largest city in Madagascar and has a population of approximately 183,000. It's the capital of the Vakinankaratra region... |
+ | + |
| Behova | + | + |
| Beloha Beloha Beloha is a town in Androy Region, Madagascar.Beloha is a town in Androy Region, Madagascar.Beloha is a town in Androy Region, Madagascar.... |
+ | + |
| Belo sur Mer Belo sur Mer Belo sur Mer is a town and commune in Madagascar. It belongs to the district of Morondava, which is a part of Menabe Region. The population of the commune was estimated to be approximately 8,000 in 2001 commune census.... |
+ | + |
| Bemafandry | + | |
| Betioky Betioky Betioky is a town in Atsimo-Andrefana Region, Madagascar.... |
+ | |
| Lakaton'ny akanga | + | |
| Lelia | + | |
| Manombo | + | + |
| Tsiandroina | + | |
| Tsiravé | + | |
Abbreviations:
|
||
Cryptoprocta spelea is the only extinct member of the order Carnivora
Carnivora
The diverse order Carnivora |Latin]] carō "flesh", + vorāre "to devour") includes over 260 species of placental mammals. Its members are formally referred to as carnivorans, while the word "carnivore" can refer to any meat-eating animal...
known from Madagascar; recently extinct Madagascan animals also include at least 17 species of lemur
Lemur
Lemurs are a clade of strepsirrhine primates endemic to the island of Madagascar. They are named after the lemures of Roman mythology due to the ghostly vocalizations, reflective eyes, and the nocturnal habits of some species...
s, most of which are larger than the living forms, as well as elephant bird
Elephant bird
Elephant birds are an extinct family of flightless birds found only on the island of Madagascar and comprising the genera Aepyornis and Mullerornis.-Description:...
s and Malagasy hippopotamus
Malagasy Hippopotamus
Several species of Malagasy Hippopotamus lived on the island of Madagascar but are now believed to be extinct. The animals were very similar to the extant Hippopotamus and Pygmy Hippopotamus...
es, among others. Subfossil remains of the giant fossa have been found in Holocene
Holocene
The Holocene is a geological epoch which began at the end of the Pleistocene and continues to the present. The Holocene is part of the Quaternary period. Its name comes from the Greek words and , meaning "entirely recent"...
cave sites from the northern end of Madagascar along the west coast to the far south, and in the central highlands. Some sites have yielded both C. spelea and smaller remains referable to the living species, C. ferox; however, lack of robust stratigraphic
Stratigraphy
Stratigraphy, a branch of geology, studies rock layers and layering . It is primarily used in the study of sedimentary and layered volcanic rocks....
knowledge and no available radiocarbon dating
Radiocarbon dating
Radiocarbon dating is a radiometric dating method that uses the naturally occurring radioisotope carbon-14 to estimate the age of carbon-bearing materials up to about 58,000 to 62,000 years. Raw, i.e. uncalibrated, radiocarbon ages are usually reported in radiocarbon years "Before Present" ,...
on subfossil Cryptoprocta bones makes it uncertain whether the two species lived in the same region at the same time. The size ratio between the two species is within the range of ratios seen between similar-sized living cats
Felidae
Felidae is the biological family of the cats; a member of this family is called a felid. Felids are the strictest carnivores of the thirteen terrestrial families in the order Carnivora, although the three families of marine mammals comprising the superfamily pinnipedia are as carnivorous as the...
and mongooses found in the same areas, suggesting that the two species may have been able to occur together.
With its large size and massive jaws and teeth, C. spelea was a formidable, "puma-like" predator, and in addition to smaller lemurids, it may have eaten some of the big, now extinct subfossil lemur
Subfossil lemur
Subfossil lemurs are lemurs from Madagascar that are represented by recent remains dating from nearly 26,000 years ago to approximately 560 years ago. They include both living and extinct species, although the term more frequently refers to the extinct giant lemurs...
s that would have been too large for C. ferox. No subfossil evidence has been found to definitively show that lemurs were its prey; this assumption is based on the diet of the smaller, extant species of fossa. Other possible prey include tenrecs, smaller euplerids, and even young Malagasy hippopotamus
Malagasy Hippopotamus
Several species of Malagasy Hippopotamus lived on the island of Madagascar but are now believed to be extinct. The animals were very similar to the extant Hippopotamus and Pygmy Hippopotamus...
es. Its extinction may have changed predation dynamics on Madagascar.
The IUCN Red List
IUCN Red List
The IUCN Red List of Threatened Species , founded in 1963, is the world's most comprehensive inventory of the global conservation status of biological species. The International Union for Conservation of Nature is the world's main authority on the conservation status of species...
currently lists C. spelea as an extinct species; why and when it went extinct remains unknown. However, local people on Madagascar often recognize two forms of fossa, a larger fosa mainty (or "black Cryptoprocta") and a smaller fosa mena (or "reddish Cryptoprocta"). There are also some anecdotal records of very large living fossas, such as a 2-m (7 ft), 30-kg (70 lb) fossa at Morondava
Morondava
Morondava is a city located in Menabe Region, of which it is the capital, in Madagascar. It is located in the delta of the Morandava River at .-Communication:...
. Goodman and colleagues suggested that further research may demonstrate that there is more than one species of fossa yet alive.
Literature cited
- Alcover, J.A. and McMinn, M. 1994. Predators of vertebrates on islands (subscription required). BioScience 44(1):12–18.
- Burness, G.P., Diamond J. and Flannery, T. 2001. Dinosaurs, dragons, and dwarfs: The evolution of maximal body size (subscription required). Proceedings of the National Academy of Sciences 98(25):14518–14523.
- Burney, D.A., Burney, L.P., Godfrey, L.R., Jungers, W.L., Goodman, S.M., Wright, H.T. and Jull, A.J.T. 2004. A chronology for late prehistoric Madagascar (subscription required). Journal of Human Evolution 47(1–2):25–63.
- Colquhoun, I.C. 2006. Predation and cathemerality: Comparing the impact of predators on the activity patterns of lemurids and ceboids (subscription required). Folia Primatologica 77(1–2):143–165.
- Garbutt, N. 2007. Mammals of Madagascar: A Complete Guide. A & C Black, 304 pp. ISBN 978-0-7136-7043-1
- Goodman, S.M. 2003. Predation on lemurs. Pp. 1221–1228 in Goodman, S.M. and Benstead, J.P. (eds.). The Natural History of Madagascar. University of Chicago Press, 1728 pp. ISBN 0-226-30306-3
- Goodman, S.M., Ganzhorn, J.U. and Rakotondravony, D. 2003. Introduction to the mammals. Pp. 1159–1186 in Goodman, S.M. and Benstead, J.P. (ed.). The Natural History of Madagascar. University of Chicago Press, 1728 pp. ISBN 0-226-30306-3
- Goodman, S.M., Rasoloarison, R.M. and Ganzhorn, J.U. 2004. On the specific identification of subfossil Cryptoprocta (Mammalia, Carnivora) from Madagascar. Zoosystema 26(1):129–143.
- Hoffman, M. 2008. . In IUCN. IUCN Red List of Threatened Species. Version 2009.2. <www.iucnredlist.org>. Downloaded on May 20, 2010.
- Köhncke, M. and Leonhardt, K. 1986. Cryptoprocta ferox. Mammalian Species 254:1–5.
- Lamberton, C. 1939. Les Cryptoprocta fossiles. Mémoires de l'Académie malgache 27:155–193.
- Turvey, S.T. 2009. Holocene Extinctions. Oxford University Press US, 359 pp. ISBN 978-0-19-953509-5
- Wroe, S., Field, J., Fullagar, R. and Jermiin, L.S. 2004. Megafaunal extinction in the late Quaternary and the global overkill hypothesis (subscription required). Alcheringa 28(1):291–331.