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''Cryptoprocta spelea'', also known as the giant fossa, is an
extinct species This page features lists of extinct species, organisms that have become extinct, either in the wild or completely disappeared from Earth. In actual theoretical practice, a species not definitely located in the wild in the last fifty years of cur ...
of carnivore from
Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Africa ...
in the family
Eupleridae Eupleridae is a Family (biology), family of carnivorans endemic to Madagascar and comprising 10 known living species in seven genus, genera, commonly known as euplerids, Malagasy mongooses or Malagasy carnivorans. The best known species is t ...
which is most closely related to the
mongoose A mongoose is a small terrestrial carnivorous mammal belonging to the family Herpestidae. This family is currently split into two subfamilies, the Herpestinae and the Mungotinae. The Herpestinae comprises 23 living species that are native to ...
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'' was larger than the fossa, but otherwise similar. The two have not always been accepted as distinct species. When and how ''C. spelea'' became 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 A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
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 in the same places 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.


Taxonomy

In 1902,
Guillaume Grandidier Guillaume Grandidier (1 July 1873 – 13 September 1957) 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 A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
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.Goodman et al., 2004, p. 130
Charles Lamberton Charles Lamberton (23 April 18768 October 1960) was a French paleontologist who lived and studied on the island of Madagascar between 1911 and 1948 and specialized in the recently extinct subfossil lemurs. He made significant contributions towa ...
reviewed subfossil and living ''Cryptoprocta'' in 1939 and agreed with Petit in recognizing 2 species, naming this species from a specimen found at
Ankazoabo Cave Ankazoabo is a town in Atsimo-Andrefana Region, Madagascar and the capital of the district with the same name Ankazoabo (district) Ankazoabo is a district in Atsimo-Andrefana Region, Madagascar. 10700 ha of the district are cultivated. An airpo ...
near the place
Itampolo Itampolo is a town and commune ( mg, kaominina) 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 ...
. The generic name 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.Goodman et al., 2003, p. 1167 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.Goodman et al., 2004, p. 131 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''.Goodman et al., 2004, p. 136 Grandidier had not designated a
type specimen In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to anchor or centralizes the ...
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 In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to anchor or centralizes th ...
). Lamberton recognized a third species, ''Cryptoprocta antamba'', on the basis of a
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bon ...
(lower jaw) with abnormally broad spacing between the condyloid processes at the back. He also referred two
femora The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates with t ...
(upper leg bones) and a
tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...
(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 (1607–1660) was a French governor of Madagascar, born in Orléans in 1607. He was named governor of Madagascar by the French East India Company in 1648. Flacourt restored order among the French soldiers, who had mutinied. ...
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''.Goodman et al., 2004, p. 137 Together, the fossa and ''C. spelea'' form the genus ''Cryptoprocta'' within the family
Eupleridae Eupleridae is a Family (biology), family of carnivorans endemic to Madagascar and comprising 10 known living species in seven genus, genera, commonly known as euplerids, Malagasy mongooses or Malagasy carnivorans. The best known species is t ...
, which also includes the other Malagasy
carnivora Carnivora is a monophyletic order of placental mammals consisting of the most recent common ancestor of all cat-like and dog-like animals, and all descendants of that ancestor. Members of this group are formally referred to as carnivorans, ...
ns—the
falanouc ''Eupleres'' is a genus of two species of mongoose-like euplerid mammal native to Madagascar that are known as falanoucs. They are primarily terrestrial and consume mainly invertebrates. Species *Eastern falanouc, ''Eupleres goudotii'' - mes ...
, the
fanaloka The Malagasy or striped civet (''Fossa fossana''), also known as the fanaloka (Malagasy, ) or jabady, is an Eupleridae, euplerid endemic to Madagascar. It is the monotypic taxon, only species in genus ''Fossa''. The Malagasy civet is a small ma ...
s, and the
Galidiinae Galidiinae is a subfamily of carnivorans that is restricted to Madagascar and includes six species classified into four genera. Together with the three other species of indigenous Malagasy carnivorans, including the fossa, they are currently cl ...
.
DNA sequence DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, guanine, cytosine, and thymine. Th ...
studies suggest that the Eupleridae form a single natural (
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic ...
) group and are most closely related to the
mongoose A mongoose is a small terrestrial carnivorous mammal belonging to the family Herpestidae. This family is currently split into two subfamilies, the Herpestinae and the Mungotinae. The Herpestinae comprises 23 living species that are native to ...
s of Eurasia and mainland Africa.


Description

Although some morphological differences between the two fossa species have been described,Lamberton, 1939, p. 182 these may be
allometric Allometry is the study of the relationship of body size to shape, anatomy, physiology and finally behaviour, first outlined by Otto Snell in 1892, by D'Arcy Thompson in 1917 in ''On Growth and Form'' and by Julian Huxley in 1932. Overview Allome ...
(growth-related), and in their 1986 ''
Mammalian Species ''Mammalian Species'' is a peer-reviewed scientific journal published by Oxford University Press on behalf of the American Society of Mammalogists. The journal publishes accounts of 12–35 mammal species yearly. The articles summarize the current ...
'' account of the fossa, Michael Köhncke and Klaus Leonhardt wrote that the two were morphologically identical.Köhncke and Leonhardt, 1986, p. 2 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
postcrania Postcrania (postcranium, adjective: postcranial) in zoology and vertebrate paleontology is all or part of the skeleton apart from the skull. Frequently, fossil remains, e.g. of dinosaurs or other extinct tetrapods, consist of partial or isolated s ...
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 The humerus (; ) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of a roun ...
(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.Goodman et al., 2004, table 1 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 and ecology

Subfossil remains of the giant fossa have been found in
Holocene The Holocene ( ) is the current geological epoch. It began approximately 11,650 cal years Before Present (), after the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene togeth ...
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 is a branch of geology concerned with the study of rock layers ( strata) and layering (stratification). It is primarily used in the study of sedimentary and layered volcanic rocks. Stratigraphy has three related subfields: lithostr ...
knowledge and no available
radiocarbon dating Radiocarbon dating (also referred to as carbon dating or carbon-14 dating) is a method for determining the age of an object containing organic material by using the properties of radiocarbon, a radioactive isotope of carbon. The method was de ...
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 The cat (''Felis catus'') is a domestic species of small carnivorous mammal. It is the only domesticated species in the family Felidae and is commonly referred to as the domestic cat or house cat to distinguish it from the wild members of ...
and
mongooses A mongoose is a small terrestrial carnivorous mammal belonging to the family Herpestidae. This family is currently split into two subfamilies, the Herpestinae and the Mungotinae. The Herpestinae comprises 23 living species that are native to s ...
found in the same areas, suggesting that the two species may have been able to occur together.Goodman et al., 2004, p. 141 With its large size and massive jaws and teeth,Goodman et al., 2004, p. 138 ''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 lemur Subfossil lemurs are lemurs from Madagascar that are represented by recent (subfossil) remains dating from nearly 26,000 years ago to approximately 560 years ago (from the late Pleistocene until the Holocene). They include both extant ...
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
tenrec A tenrec is any species of mammal within the afrotherian family Tenrecidae endemic to Madagascar. Tenrecs are wildly diverse; as a result of convergent evolution some resemble hedgehogs, shrews, opossums, rats, and mice. They occupy aquatic, a ...
s, smaller euplerids, and even young Malagasy hippopotamuses.


Extinction

Why and when ''C. spelea'' went extinct is not known; it is possible that ''C. spelea'' went extinct before 1400. ''C. spelea'' is on the
IUCN Red List The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biol ...
. 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 (, from mg, morona lava "long coast") 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 . Its population as of the 2018 census, was 53,510. Population T ...
. 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
carnivora Carnivora is a monophyletic order of placental mammals consisting of the most recent common ancestor of all cat-like and dog-like animals, and all descendants of that ancestor. Members of this group are formally referred to as carnivorans, ...
n mammal known from Madagascar; recently extinct (non-carnivoran) Madagascan animals also include large
lemur Lemurs ( ) (from Latin ''lemures'' – ghosts or spirits) are wet-nosed primates of the superfamily Lemuroidea (), divided into 8 families and consisting of 15 genera and around 100 existing species. They are endemic to the island of Madaga ...
s,
elephant bird Elephant birds are members of the extinct ratite family Aepyornithidae, made up of flightless birds that once lived on the island of Madagascar. They are thought to have become extinct around 1000-1200 CE, probably as a result of human activit ...
s, and Malagasy hippopotamuses.Burney et al., 2004, p. 25 The extinction of ''C. spelea'' may have changed predation dynamics on Madagascar.


References


Literature cited

* * * * * * * * * * * * * {{Taxonbar, from=Q1716674 Euplerids Extinct carnivorans Holocene extinctions Prehistoric animals of Madagascar