Cryptoprocta spelea

In 1902, Guillaume Grandidier listed subfossil carnivoran remains from two caves on Madagascar as a larger "variety" of the living fossa (Cryptoprocta ferox), C. ferox var. spelea, without a detailed description or holotype designation. In 1935, Gabriel Petit considered spelea to represent a distinct species, but did not assign any specimen as the holotype. Charles Lamberton reviewed the subfossil and living Cryptoprocta remains in 1939, and assigned other specimens found in different localities to C. spelea, agreeing with Petit in the recognition of two species. The generic name Cryptoprocta translates to "hidden anus" referring to the fact that the anus is hidden by anal sacs in C. ferox. 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 two distinct 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 no material described by Grandidier that is associated with the name spelea can be located in the collections of the National Museum of Natural History, France (MNHN). To maintain C. spelea as the name for the larger form of the fossa, Goodman and colleagues proposed a well-preserved specimen MNHN CG 1977.755 to be designated as the neotype. Lamberton recognized a third species, Cryptoprocta antamba, on the basis of a mandible (lower jaw) with abnormally broad spacing between the condyloid processes 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 carnivorans—the falanouc, the fanalokas, and the Galidiinae. DNA sequence studies suggest that the Eupleridae form a single natural (monophyletic) group and are most closely related to the mongooses 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 Species 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 spelea were 1.07 to 1.32 times as large as in adult C. ferox, and postcranial 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 , compared to a range of in adult C. ferox. Humerus (upper arm bone) length in twelve C. spelea is , averaging , compared to , averaging , in the extant fossa. Body mass estimates for C. spelea range from to , and it was among the largest carnivores of the island. By comparison, adult C. ferox range from to . ==Distribution and ecology==

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 prey it may have eaten some of the big, now extinct subfossil lemurs 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 hippopotamuses. ==Extinction==

(Cryptoprocta ferox) is a smaller relative of C. spelea that still survives. The specific reason and time for the extinction of C. spelea is not known. Goodman et al. (2004) suggested that C. spelea went extinct before 1400, while the IUCN Red List listed its time of extinction as before 1658. 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 the alleged record of a , fossa captured at Morondava in 1954. Goodman and colleagues suggested that further research may demonstrate that there is more than one species of fossa yet alive. C. spelea is the only extinct carnivoran mammal known from Madagascar; recently extinct (non-carnivoran) Madagascan animals also include large lemurs, elephant birds, and Malagasy hippopotamuses. The extinction of C. spelea may have changed predation dynamics on Madagascar due to Madagascar's larger terrestrial vertebrates experiencing the lack of predator pressure. ==References==