Aldabrachampsus
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''Aldabrachampsus'' is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of small horned crocodile known from fragmentary remains. It lived during the
Pleistocene The Pleistocene ( ; referred to colloquially as the ''ice age, Ice Age'') is the geological epoch (geology), epoch that lasted from to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fin ...
on
Aldabra Aldabra, the world's second-largest coral atoll (the largest is Kiritimati), is located east of the continent of Africa. It is part of the Aldabra Group of islands in the Indian Ocean that are part of the Outer Islands (Seychelles), Outer Islands ...
Atoll,
Seychelles Seychelles (, ; ), officially the Republic of Seychelles (; Seychellois Creole: ), is an island country and archipelagic state consisting of 155 islands (as per the Constitution) in the Indian Ocean. Its capital and largest city, Victoria, ...
in the western
Indian Ocean The Indian Ocean is the third-largest of the world's five oceanic divisions, covering or approximately 20% of the water area of Earth#Surface, Earth's surface. It is bounded by Asia to the north, Africa to the west and Australia (continent), ...
. The name ''Aldabrachampsus dilophus'' means "Two-crested crocodile from Aldabra". It was a small animal, reaching a length of , comparable in size to the smallest extant crocodilians.


Description

''Aldabrachampsus'' is only known from fragmentary cranial and mandibular remains including the
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammals h ...
,
frontal bone In the human skull, the frontal bone or sincipital bone is an unpaired bone which consists of two portions.'' Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bo ...
, parietal, dentaries and squamosals among others. Some postcranial bones are also known including several
vertebrae Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spinal ...
and a
femur The femur (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg. The Femo ...
. Like most other crocodilians ''Aldabrachampsus'' possessed five teeth situated in the premaxilla, however the second to fifth are all aligned in a straight line rather than following a convex path. Following the premaxillary teeth the rostrum of the crocodile constricted, creating a notch for one of the dentary teeth. There is a toothless region (diastema) present between the second and third alveoli of the dentary. The orbital margins of both the frontal and prefrontal bone are raised. The main body of the frontal is shaped like a
pentagon In geometry, a pentagon () is any five-sided polygon or 5-gon. The sum of the internal angles in a simple polygon, simple pentagon is 540°. A pentagon may be simple or list of self-intersecting polygons, self-intersecting. A self-intersecting ...
with a distinctly V-shaped suture with the parietals, excluding the frontal from contributing to the
supratemporal fenestrae Temporal fenestrae are openings in the Temple (anatomy), temporal region of the skull of some Amniote, amniotes, behind the Orbit (anatomy), orbit (eye socket). These openings have historically been used to track the evolution and affinities of re ...
. The anterior process of the frontal bone is slender and notably protrudes from the body following a sudden sharp constriction. Of the parietal only the posterior section is known, which shows a flat surface and a deep notch in its rear margin exposing the occipital region. The squamosal bone widens towards its suture with the
postorbital The ''postorbital'' is one of the bones in vertebrate skulls which forms a portion of the dermal skull roof and, sometimes, a ring about the orbit. Generally, it is located behind the postfrontal and posteriorly to the orbital fenestra. In some ve ...
and the bone bears a horn-like projection that faces outwards. Similar structures are known in the
Madagascar Madagascar, officially the Republic of Madagascar, is an island country that includes the island of Madagascar and numerous smaller peripheral islands. Lying off the southeastern coast of Africa, it is the world's List of islands by area, f ...
crocodile ''
Voay ''Voay'' is an extinct genus of crocodile from Madagascar that lived during the Late Pleistocene to Holocene, containing only one species, ''V. robustus''. Numerous subfossils have been found, including complete skulls, noted for their distinctiv ...
'', '' Crocodylus anthropophagus'' from
Tanzania Tanzania, officially the United Republic of Tanzania, is a country in East Africa within the African Great Lakes region. It is bordered by Uganda to the northwest; Kenya to the northeast; the Indian Ocean to the east; Mozambique and Malawi to t ...
and even alligatoroids like '' Acresuchus'' and ''
Ceratosuchus ''Ceratosuchus'' ("horned crocodile") is an extinct genus of alligatorine crocodylian from latest Paleocene rocks of Colorado's Piceance Basin and earliest Eocene rocks of Wyoming's Bighorn Basin in North America, a slice of time known as the ...
''. In modern species both
Siamese crocodile The Siamese crocodile (''Crocodylus siamensis'') is a medium-sized freshwater crocodile native to Indonesia (Borneo and possibly Java), Brunei, East Malaysia, Laos, Cambodia, Myanmar, Thailand and Vietnam. The species is critically endangered and ...
s and Cuban crocodiles share similar squamosal horns. However ''Aldabrachampsus'' differs from these taxa in two ways. The horns are notably less pointed, showing a more rounded morphology, and their apex is located towards the halfway point of the bone, rather than the squamosal horns of other crocodilians which reach their apex towards the posterior. The vertebrae show that the neural centra and neural arches were at least partly fused with the sutures beginning to close. This indicates that the animal's these bones belonged to had almost reached their adult size. With a length of the vertebrae suggest that the animal, assuming similar proportions to modern crocodiles, was relatively small in size, approximately long. Extrapolating the size based on the skull remains gives similar results ranging from . This size would be similar to the smallest extant crocodilians, the South American dwarf caimans and African dwarf crocodiles, alongside extinct dwarf mekosuchines like '' Volia'' and '' Mekosuchus''. Some crocodilian remains found later might indicate the presence of a larger bodied and more robust animal on the island, it is however not clear if these fossils belong to a large specimen or species of ''Aldabrachampsus'' or a taxon previously not known from the island like the
Nile crocodile The Nile crocodile (''Crocodylus niloticus'') is a large crocodilian native to freshwater habitats in Africa, where it is present in 26 countries. It is widely distributed in sub-Saharan Africa, occurring mostly in the eastern, southern, and ce ...
or an osteolaemine related to ''Voay''. These remains indicate a crocodile with a body length of .


Classification

The fragmentary nature of ''Aldabrachampsus'' has left its exact placement within
Crocodilia Crocodilia () is an order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorph pseudosuchia ...
somewhat ambiguous.
Phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
analysis suggests it is part of the
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Crocodylidae Crocodiles (family Crocodylidae) or true crocodiles are large, semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term "crocodile" is sometimes used more loosely to include all extant membe ...
, composed of the
subfamilies In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily names with "-oideae", and zool ...
Crocodylinae and Osteolaeminae. Although this relationship is recovered by the strict consensus tree, which results in a large
polytomy An internal node of a phylogenetic tree is described as a polytomy or multifurcation if (i) it is in a rooted tree and is linked to three or more child subtrees or (ii) it is in an unrooted tree and is attached to four or more branches. A tree ...
rendering ''Aldabrachampsuss exact position unknown, other trees suggest potentially different relationships. In some trees, ''Aldabrachampsus'' is recovered as a
sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
to ''
Voay ''Voay'' is an extinct genus of crocodile from Madagascar that lived during the Late Pleistocene to Holocene, containing only one species, ''V. robustus''. Numerous subfossils have been found, including complete skulls, noted for their distinctiv ...
'', while others find it to be a sister to '' Crocodylus palaeindicus''. However, both these positions are weakly supported, based on a single morphological character (the exposed supraoccipital), which is expressed differently between the different crocodiles in question. Brochu, however, argues that despite being based on the same character, a relationship to ''Voay'' (then ''Crocodylus robustus'') within Osteolaeminae was more likely due to the anatomy of the
nare Nare may refer to: People * Daniel Nare, Botswanan football manager * Tadu Teshome Nare (born 2001), Ethiopian runner * Théophile Naré (born 1966), Catholic bishop of Kaya, Burkina Faso Other meanings * Nareh (given name) (or Nare, Nara), Armeni ...
s,
palate The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly sep ...
and the
squamosal The squamosal is a skull bone found in most reptiles, amphibians, and birds. In fishes, it is also called the pterotic bone. In most tetrapods, the squamosal and quadratojugal bones form the cheek series of the skull. The bone forms an ancestra ...
horns, although they differ in shape.


Evolution

The first remains of ''Aldabrachampsus'' were found in Quaternary deposits of Point Hodoul at the eastern end of the Aldabra Atoll. The limestone cavities that yielded these remains also preserved the fossils of lizards, tortoises, birds and land snails and date to approximately 118,000 years ago. Just before the appearance of ''Aldabrachampsus'' the atoll was entirely submerged by water, which suggests that rather than being an endemic species affected by
island dwarfism Insular dwarfism, a form of phyletic dwarfism, is the process and condition of large animals evolving or having a reduced body size when their population's range is limited to a small environment, primarily islands. This natural process is disti ...
, this crocodile was already of small stature by the time it arrived on the island. The fragmentary nature and poorly resolved phylogenetic relationships however make it difficult to determine where ''Aldabrachampsus'' originated. It is possible that the genus arrived in the Aldabra Atoll from the east, assuming close relationship with ''Crocodylus palaeindicus'', however Brochu again suggests that a relationship to ''Voay'' and thus an African or Malagasy origin is more likely. A Malagasy origin would be consistent with the proximity between the atoll and Madagascar, which has led to faunal exchanges between the islands, especially prominent in regards to tortoises. Given the small stature of ''Osteolaemus'', it is possible that minute bodysize is ancestral to this grouping and was simply reversed by ''Voay'' on Madagascar. Another hypothesis meanwhile suggests that smaller bodysize is a trait evolved by ''Osteolaemus'' and ''Aldabrachampsus'' independently of one another.


Paleoecology

Fossil discoveries made at a dried up pond on Grande Terre Island corresponding to the Late Pleistocene Aldabra Limestone show evidence of a predator-prey relationship between crocodilians and the native giant tortoise '' Aldabrachelys gigantea''. Several tortoise shell fragments from the site are known to bear distinct feeding marks in the form of unhealed puncture wounds inflicted to the nuchal armor, plastron and the pubis of the tortoise. It is possible that a crocodile attacked and killed a tortoise head on as it was drinking at a watering hole, puncturing the nuchal plate at the front of the shell rather than attacking the high, better defended sides of the turtle. Another possibility is that the tortoise was not killed by crocodiles and instead only scavenged on. Again the punctures on the nuchal shield are caused by the fact that this particular area allows for better access at the meat beneath the shell. However, as the crocodile remains found alongside the turtle bones are notably larger than any previously recorded ''Aldabrachampsus'' specimens and very fragmentary, it is not certain if they belong to the same genus or instead represent a second species of crocodile native to the island.


References

{{Taxonbar, from=Q4713415, from2=Q15041820 Crocodylidae Pleistocene crocodylomorphs Monotypic prehistoric reptile genera Prehistoric pseudosuchian genera Fossil taxa described in 2006