Echimyidae Phylogeny
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Echimyidae is the family of neotropical spiny rats and their
fossil 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 ...
relatives. This is the most species-rich family of
hystricognath The Hystricognathi are an infraorder of rodents, distinguished from other rodents by the bone structure of their skulls. The masseter medialis (a jaw muscle) passes partially through a hole below each eye socket (called the infraorbital foramen) ...
rodents. It is probably also the most ecologically diverse, with members ranging from fully
arboreal Arboreal locomotion is the Animal locomotion, locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally, but others are exclusively arboreal. Th ...
to
terrestrial Terrestrial refers to things related to land or the planet Earth. Terrestrial may also refer to: * Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on or near the ground, as opposed to ...
to
fossorial A fossorial () animal is one adapted to digging which lives primarily, but not solely, underground. Some examples are badgers, naked mole-rats, clams, meerkats, and mole salamanders, as well as many beetles, wasps, and bees. Prehistoric eviden ...
to
semiaquatic In biology, semiaquatic can refer to various types of animals that spend part of their time in water, or plants that naturally grow partially submerged in water. Examples are given below. Semiaquatic animals Semiaquatic animals include: * Verte ...
habits. They presently exist mainly in
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
; three members of the family also range into
Central America Central America ( es, América Central or ) is a subregion of the Americas. Its boundaries are defined as bordering the United States to the north, Colombia to the south, the Caribbean Sea to the east, and the Pacific Ocean to the west. ...
, and the
hutias Hutias (known in Spanish as jutía) are moderately large cavy-like rodents of the subfamily Capromyinae that inhabit the Caribbean islands, with most species restricted to Cuba and Hispaniola. Twenty species of hutia have been identified, but at ...
are found in the
Greater Antilles The Greater Antilles ( es, Grandes Antillas or Antillas Mayores; french: Grandes Antilles; ht, Gwo Zantiy; jam, Grieta hAntiliiz) is a grouping of the larger islands in the Caribbean Sea, including Cuba, Hispaniola, Puerto Rico, Jamaica, and ...
in the
Caribbean The Caribbean (, ) ( es, El Caribe; french: la Caraïbe; ht, Karayib; nl, De Caraïben) is a region of the Americas that consists of the Caribbean Sea, its islands (some surrounded by the Caribbean Sea and some bordering both the Caribbean Se ...
. Species of the extinct subfamily Heteropsomyinae formerly lived on
Cuba Cuba ( , ), officially the Republic of Cuba ( es, República de Cuba, links=no ), is an island country comprising the island of Cuba, as well as Isla de la Juventud and several minor archipelagos. Cuba is located where the northern Caribbea ...
,
Hispaniola Hispaniola (, also ; es, La Española; Latin and french: Hispaniola; ht, Ispayola; tnq, Ayiti or Quisqueya) is an island in the Caribbean that is part of the Greater Antilles. Hispaniola is the most populous island in the West Indies, and th ...
, and
Puerto Rico Puerto Rico (; abbreviated PR; tnq, Boriken, ''Borinquen''), officially the Commonwealth of Puerto Rico ( es, link=yes, Estado Libre Asociado de Puerto Rico, lit=Free Associated State of Puerto Rico), is a Caribbean island and Unincorporated ...
in the
Antilles The Antilles (; gcf, label=Antillean Creole, Antiy; es, Antillas; french: Antilles; nl, Antillen; ht, Antiy; pap, Antias; Jamaican Patois: ''Antiliiz'') is an archipelago bordered by the Caribbean Sea to the south and west, the Gulf of Mex ...
, probably until the arrival of Europeans.


Characteristics

In general form, most spiny rats resemble
rat Rats are various medium-sized, long-tailed rodents. Species of rats are found throughout the order Rodentia, but stereotypical rats are found in the genus ''Rattus''. Other rat genera include ''Neotoma'' ( pack rats), ''Bandicota'' (bandicoot ...
s, although they are more closely related to
guinea pig The guinea pig or domestic guinea pig (''Cavia porcellus''), also known as the cavy or domestic cavy (), is a species of rodent belonging to the genus ''Cavia'' in the family Caviidae. Breeders tend to use the word ''cavy'' to describe the ani ...
s and
chinchilla Chinchillas are either of two species (''Chinchilla chinchilla'' and ''Chinchilla lanigera'') of crepuscular rodents of the parvorder Caviomorpha. They are slightly larger and more robust than ground squirrels, and are native to the Andes mount ...
s. Most species have stiff, pointed hairs, or spines, that presumably serve for protection from predators. Many echimyids can break off their tails when attacked. This action may confuse
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
s long enough for the spiny rat to escape. Unlike the tails of some species of
lizard Lizards are a widespread group of squamate reptiles, with over 7,000 species, ranging across all continents except Antarctica, as well as most oceanic island chains. The group is paraphyletic since it excludes the snakes and Amphisbaenia alt ...
s, however, the tails of spiny rats do not regenerate. Therefore, the tactic can only be used once in an individual's lifetime. Most spiny rats are rare and poorly known, but a few are extremely abundant. Various species are respectively
terrestrial Terrestrial refers to things related to land or the planet Earth. Terrestrial may also refer to: * Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on or near the ground, as opposed to ...
,
arboreal Arboreal locomotion is the Animal locomotion, locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally, but others are exclusively arboreal. Th ...
, or
fossorial A fossorial () animal is one adapted to digging which lives primarily, but not solely, underground. Some examples are badgers, naked mole-rats, clams, meerkats, and mole salamanders, as well as many beetles, wasps, and bees. Prehistoric eviden ...
. In general, the arboreal forms are most rat-like in appearance, whilst the burrowing species are more
gopher Pocket gophers, commonly referred to simply as gophers, are burrowing rodents of the family Geomyidae. The roughly 41 speciesSearch results for "Geomyidae" on thASM Mammal Diversity Database are all endemic to North and Central America. They are ...
-like, with stocky bodies and short tails. Most species do poorly in conditions of high heat and
arid A region is arid when it severely lacks available water, to the extent of hindering or preventing the growth and development of plant and animal life. Regions with arid climates tend to lack vegetation and are called xeric or desertic. Most ar ...
ity and are restricted to regions with abundant water. They are almost exclusively
herbivorous A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
.


Systematics

The current taxonomic content of the family Echimyidae has been reshaped over time, and its organization into coherent units stems from two realizations. The first is that
cladistic Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups (" clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived char ...
approaches applied to morphological characters showed that many traits used to define taxa were
plesiomorphic In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral character shared by all members of a clade, which does not distinguish the clade from other clades. Plesiomorphy, symplesiomorphy, apomorphy, and ...
or
homoplastic Homoplasy, in biology and phylogenetics, is the term used to describe a Phenotypic trait, feature that has been gained or lost independently in separate lineages over the course of evolution. This is different from Homology (biology), homology, wh ...
. The second realization came from the advent of
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
analyses of
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 ...
and
protein sequence Protein primary structure is the linear sequence of amino acids in a peptide or protein. By convention, the primary structure of a protein is reported starting from the amino-terminal (N) end to the carboxyl-terminal (C) end. Protein biosynthesi ...
data with probability methods —
maximum likelihood In statistics, maximum likelihood estimation (MLE) is a method of estimation theory, estimating the Statistical parameter, parameters of an assumed probability distribution, given some observed data. This is achieved by Mathematical optimization, ...
and
Bayesian inference Bayesian inference is a method of statistical inference in which Bayes' theorem is used to update the probability for a hypothesis as more evidence or information becomes available. Bayesian inference is an important technique in statistics, a ...
— leading to the identification of robust
clades A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, t ...
and the recognition of higher categorical ranks (see Phylogeny section). The following table recapitulates and compares the taxonomic content of taxa recognized on molecular and traditional basis: the two families Capromyidae and
Myocastoridae Myocastorini is a tribe of echimyid rodents, proposed in 2017, and containing the five extant genera ''Callistomys'', ''Hoplomys'', ''Myocastor'', ''Proechimys'', and ''Thrichomys ''Thrichomys'' is a genus of South American rodents in the fam ...
, the five subfamilies
Echimyinae Echimyinae is a subfamily of rodents belonging to the spiny rats family Echimyidae. It contains 14 arboreal genera—all the members of the tribe Echimyini, plus ''Callistomys''—a few terrestrial genera (''Thrichomys'', ''Hoplomys'', and ''Pro ...
,
Euryzygomatomyinae Euryzygomatinae is a subfamily of rodents, proposed in 2017, and containing three extant genera of spiny Echimyidae: '' Clyomys'', '' Euryzygomatomys'', and ''Trinomys''. Members_of_this_echimyid_subfamily_all_share_an_origin_in_the_eastern_part ...
,
Capromyinae Hutias (known in Spanish as jutía) are moderately large cavy-like rodents of the subfamily Capromyinae that inhabit the Caribbean islands, with most species restricted to Cuba and Hispaniola. Twenty species of hutia have been identified, but a ...
, Dactylomyinae,
Eumysopinae Echimyidae is the family of neotropical spiny rats and their fossil relatives. This is the most species-rich family of Hystricognathi, hystricognath rodents. It is probably also the most ecologically diverse, with members ranging from fully ar ...
, and the four tribes
Echimyini Echimyini is a tribe of echimyid rodents, proposed in 2016, and containing 13 extant genera: all of the tree rats ''Echimys'', ''Phyllomys'', '' Makalata'', ''Pattonomys'', '' Toromys'', ''Diplomys'', ''Santamartamys'', and ''Isothrix'', the long ...
,
Myocastorini Myocastorini is a tribe of echimyid rodents, proposed in 2017, and containing the five extant genera ''Callistomys'', ''Hoplomys'', ''Myocastor'', ''Proechimys'', and ''Thrichomys ''Thrichomys'' is a genus of South American rodents in the fam ...
, Capromyini, and Plagiodontini.


Extant genera

* Family Echimyidae - spiny rats


Extinct genera

* Family Echimyidae - spiny rats ** †'' Cercomys'' ** †'' Maruchito'' ** †'' Paulacoutomys'' ** †'' Proclinodontomys'' ** †'' Willidewu'' ** Subfamily † Adelphomyinae *** †'' Adelphomys'' *** †'' Deseadomys'' *** †'' Paradelphomys'' *** †'' Stichomys'' *** †'' Xylechimys'' ** Subfamily † Heteropsomyinae - extinct West Indian echimyids *** †''
Boromys ''Boromys'' is an extinction, extinct genus of Cuban rodents in the family Echimyidae. It contains the following species: * Oriente cave rat (''Boromys offella'') * Torre's cave rat (''Boromys torrei'') References

Boromys, Prehistoric ...
'' *** †''
Brotomys The Hispaniolan edible rat (''Brotomys voratus'') is a recently extinct species of rodent in the family Echimyidae. It is the only species in the genus ''Brotomys''. It was endemic to the island of Hispaniola in the Caribbean, in what is today ...
'' *** †''
Heteropsomys ''Heteropsomys'' is an extinct genus of rodent in the family Echimyidae Echimyidae is the family of neotropical spiny rats and their fossil relatives. This is the most species-rich family of hystricognath rodents. It is probably also the mo ...
'' *** †'' Puertoricomys'' ** Subfamily
Eumysopinae Echimyidae is the family of neotropical spiny rats and their fossil relatives. This is the most species-rich family of Hystricognathi, hystricognath rodents. It is probably also the most ecologically diverse, with members ranging from fully ar ...
*** †'' Acarechimys'' *** †'' Chasichimys'' *** †'' Eumysops'' *** †'' Palaeoechimys'' *** †'' Pampamys'' *** †'' Pattersomys'' *** †'' Protacaremys'' *** †'' Protadelphomys'' *** †'' Sallamys'' **Subfamily
Capromyinae Hutias (known in Spanish as jutía) are moderately large cavy-like rodents of the subfamily Capromyinae that inhabit the Caribbean islands, with most species restricted to Cuba and Hispaniola. Twenty species of hutia have been identified, but a ...
- hutias ***†''
Hexolobodon The imposter hutia (''Hexolobodon phenax'') is an extinct species of rodent in the hutia subfamily (Capromyinae). It is the only species in the genus ''Hexolobodon'' and tribe Hexolobodontini. It was found only on the Caribbean island of Hispanio ...
'' ***†''
Hyperplagiodontia ''Hyperplagiodontia'', rarely called the wide-toothed hutia, is an extinct genus of hutia which contains a single species, ''Hyperplagiodontia araeum''. The species was originally described as a member of the genus ''Plagiodontia'' along with the ...
'' ***†''
Isolobodon ''Isolobodon'' is an extinct genus of rodent in the subfamily Capromyinae Hutias (known in Spanish as jutía) are moderately large cavy-like rodents of the subfamily Capromyinae that inhabit the Caribbean islands, with most species restricted ...
'' ***†''
Rhizoplagiodontia Lemke's hutia (''Rhizoplagiodontia lemkei'') is an extinct species of rodent in the subfamily Capromyinae. It is monotypic within the genus ''Rhizoplagiodontia''. It was endemic to Hispaniola (the Dominican Republic and Haiti). Its natural habita ...
''


About ''Chaetomys''

The bristle-spined rat, ''
Chaetomys subspinosus The bristle-spined rat (''Chaetomys subspinosus'') is an arboreal rodent from the Atlantic forest in eastern Brazil. Also known as the bristle-spined porcupine or thin-spined porcupine, it is the only member of the genus ''Chaetomys'' and the s ...
'', has sometimes been classified in Echimyidae, although traditionally considered a member of the New World porcupine family Erethizontidae. The classification with Echimyidae is supported by similarities in the
cheek teeth Cheek teeth or post-canines comprise the molar and premolar teeth in mammals. Cheek teeth are multicuspidate (having many folds or tubercles). Mammals have multicuspidate molars (three in placentals, four in marsupials, in each jaw quadrant) and ...
structure. Like all living caviomorphs except erethizontids, ''Chaetomys'' seems to lack posterior carotid foramina, and together with all echimyids and in contrast to all other caviomorphs, ''Chaetomys'' seems to retain the otherwise deciduous premolars (dP4). Some of these characters have been, however, reinterpreted as evidence for affinities between ''Chaetomys'' and the Erethizontidae. A molecular
phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...
based on the
mitochondrial A mitochondrion (; ) is an organelle found in the cells of most Eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is use ...
gene coding for cytochrome ''b'' combined to karyological evidence actually suggests ''Chaetomys'' is more closely related to the Erethizontidae than to the Echimyidae, although it branches as the sister group to the rest of the Erethizontidae.


Phylogeny

The phylogenetic tree of the Echimyidae shows a major split between the subfamily
Echimyinae Echimyinae is a subfamily of rodents belonging to the spiny rats family Echimyidae. It contains 14 arboreal genera—all the members of the tribe Echimyini, plus ''Callistomys''—a few terrestrial genera (''Thrichomys'', ''Hoplomys'', and ''Pro ...
and an assemblage containing the
Euryzygomatomyinae Euryzygomatinae is a subfamily of rodents, proposed in 2017, and containing three extant genera of spiny Echimyidae: '' Clyomys'', '' Euryzygomatomys'', and ''Trinomys''. Members_of_this_echimyid_subfamily_all_share_an_origin_in_the_eastern_part ...
, ''
Carterodon Owl's spiny rat (''Carterodon sulcidens'') is a rodent species in the family Echimyidae found in Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and ...
'', and the Capromyidae. The first major clade contains a majority of arboreal genera (e.g., ''
Phyllomys ''Phyllomys'' is a genus of arboreal spiny rat, geographically restricted to the forests of eastern Brazil. The etymology of the genus name derives from the two ancient greek words (), meaning "plant leaf", and (), meaning "mouse, rat". Phy ...
'', ''
Dactylomys ''Dactylomys'' is the genus of South American bamboo rats They are arboreal members of the family Echimyidae. Systematics The genus name ''Dactylomys'' derives from the two Ancient Greek words (), meaning "finger", and (), meaning "mouse, rat" ...
'', and ''
Mesomys ''Mesomys'' is a genus of South American spiny rats in the family Echimyidae. The genus name ''Mesomys'' derives from the two ancient greek words (), meaning "middle of, between", and (), meaning "mouse, rat". It refers to the fact that this ...
''), a few terrestrial taxa (e.g., ''
Proechimys ''Proechimys'' is a genus of South American spiny rats of the family Echimyidae. All species of the genus are terrestrial. In the lowland Neotropical forests, ''Proechimys'' rodents are often the most abundant non-volant mammals. They are recogni ...
''), and a subaquatic one (''
Myocastor ''Myocastor'' is a genus of rodent that contains the living nutria (or coypu), as well as several fossil species. Taxonomy Due to similar cranial morphology, the nutria was once considered a close relative of the Caribbean hutias and placed toge ...
''). The second major clade includes fossorial genera (e.g., ''
Euryzygomatomys ''Euryzygomatomys'' is a genus of South American rodents, commonly called guiaras, in the family Echimyidae. It contains two extant and one fossil species, found in Argentina, Brazil and Paraguay. They are as follows: * Brandt's guiara (''Euryzy ...
'' or ''
Carterodon Owl's spiny rat (''Carterodon sulcidens'') is a rodent species in the family Echimyidae found in Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and ...
''), a terrestrial one (''
Trinomys The Atlantic spiny rats are all found in the genus ''Trinomys''. They are a group of South American spiny-rats in the family Echimyidae. Extant species of ''Trinomys'' Based on Natureserve.InfoNatura: Animals and Ecosystems of Latin America e ...
''), and members inhabiting the Caribbean islands (Capromyidae).


References and notes

{{Taxonbar">from=Q852726 Rodent families Hystricognath rodents Extant Chattian first appearances Taxa named by John Edward Gray