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 preserve ...
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 foram ...
rodents. It is probably also the most ecologically diverse, with members ranging from fully
arboreal Arboreal locomotion is the 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 (scansorial), but others are exclusively arboreal. The hab ...
to
terrestrial Terrestrial refers to things related to land or the planet Earth, as opposed to extraterrestrial. 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 o ...
to
fossorial A fossorial animal () is one that is adapted to digging and which lives primarily (but not solely) underground. Examples of fossorial vertebrates are Mole (animal), moles, badgers, naked mole-rats, meerkats, armadillos, wombats, and mole salamand ...
to
semiaquatic In biology, being semi-aquatic refers to various macroorganisms that live regularly in both aquatic and terrestrial environments. When referring to animals, the term describes those that actively spend part of their daily time in water (in ...
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 considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
; three members of the family also range into
Central America Central America is a subregion of North America. Its political boundaries are defined as bordering Mexico to the north, Colombia to the southeast, the Caribbean to the east, and the Pacific Ocean to the southwest. Central America is usually ...
, and the
hutia Hutias (known in Spanish as jutía) are moderately large cavy-like rodents of the subfamily Capromyinae that inhabit the Caribbean islands. Most species are restricted to Cuba, but species are known from all of the Greater Antilles, as well as ...
s are found in the
West Indies The West Indies is an island subregion of the Americas, surrounded by the Atlantic Ocean, North Atlantic Ocean and the Caribbean Sea, which comprises 13 independent island country, island countries and 19 dependent territory, dependencies in thr ...
in the
Caribbean The Caribbean ( , ; ; ; ) is a region in the middle of the Americas centered around the Caribbean Sea in the Atlantic Ocean, North Atlantic Ocean, mostly overlapping with the West Indies. Bordered by North America to the north, Central America ...
. Species of the extinct subfamily Heteropsomyinae formerly lived on
Cuba Cuba, officially the Republic of Cuba, is an island country, comprising the island of Cuba (largest island), Isla de la Juventud, and List of islands of Cuba, 4,195 islands, islets and cays surrounding the main island. It is located where the ...
,
Hispaniola Hispaniola (, also ) is an island between Geography of Cuba, Cuba and Geography of Puerto Rico, Puerto Rico in the Greater Antilles of the Caribbean. Hispaniola is the most populous island in the West Indies, and the second-largest by List of C ...
, and
Puerto Rico ; abbreviated PR), officially the Commonwealth of Puerto Rico, is a Government of Puerto Rico, self-governing Caribbean Geography of Puerto Rico, archipelago and island organized as an Territories of the United States, unincorporated territo ...
in the
Antilles The Antilles is an archipelago bordered by the Caribbean Sea to the south and west, the Gulf of Mexico to the northwest, and the Atlantic Ocean to the north and east. The Antillean islands are divided into two smaller groupings: the Greater An ...
.


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'' (bandicoo ...
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'', family Caviidae. Animal fancy, Breeders tend to use the name "cavy" for the ani ...
s and
chinchilla Chinchilla refers to either of two species ('' Chinchilla chinchilla'' and '' Chinchilla lanigera'') of crepuscular rodents of the parvorder Caviomorpha, and are native to the Andes mountains in South America. They live in colonies called "her ...
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 in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
s long enough for the spiny rat to escape. Unlike the tails of some species of
lizard Lizard is the common name used for all Squamata, squamate reptiles other than snakes (and to a lesser extent amphisbaenians), encompassing over 7,000 species, ranging across all continents except Antarctica, as well as most Island#Oceanic isla ...
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, as opposed to extraterrestrial. 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 o ...
,
arboreal Arboreal locomotion is the 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 (scansorial), but others are exclusively arboreal. The hab ...
, or
fossorial A fossorial animal () is one that is adapted to digging and which lives primarily (but not solely) underground. Examples of fossorial vertebrates are Mole (animal), moles, badgers, naked mole-rats, meerkats, armadillos, wombats, and mole salamand ...
. 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 ar ...
-like, with stocky bodies and short tails. Most species do poorly in conditions of high heat and
arid Aridity is the condition of geographical regions which make up approximately 43% of total global available land area, characterized by low annual precipitation, increased temperatures, and limited water availability.Perez-Aguilar, L. Y., Plata ...
ity and are restricted to regions with abundant water. They are almost exclusively
herbivorous A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat n ...
.


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 ( ; from Ancient Greek 'branch') 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 ...
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, an ...
or homoplastic. The second realization came from the advent of
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 ...
analyses of
DNA sequence A nucleic acid sequence is a succession of bases within the nucleotides forming alleles within a DNA (using GACT) or RNA (GACU) molecule. This succession is denoted by a series of a set of five different letters that indicate the order of the nu ...
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 biosynthe ...
data with probability methods —
maximum likelihood In statistics, maximum likelihood estimation (MLE) is a method of estimating the parameters of an assumed probability distribution, given some observed data. This is achieved by maximizing a likelihood function so that, under the assumed stati ...
and
Bayesian inference Bayesian inference ( or ) is a method of statistical inference in which Bayes' theorem is used to calculate a probability of a hypothesis, given prior evidence, and update it as more information becomes available. Fundamentally, Bayesian infer ...
— leading to the identification of robust
clades In biology, a clade (), also known as a monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach to taxonomy ...
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''. Definition Myocastorini members share long upper incisor ro ...
, 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 ''Pr ...
,
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 par ...
,
Capromyinae Hutias (known in Spanish as jutía) are moderately large cavy-like rodents of the subfamily Capromyinae that inhabit the Caribbean islands. Most species are restricted to Cuba, but species are known from all of the Greater Antilles, as well as Th ...
, Dactylomyinae,
Eumysopinae 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 most ecologically diverse, with members ranging from fully arboreal to terre ...
, 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 ...
,
Myocastorini Myocastorini is a tribe of echimyid rodents, proposed in 2017, and containing the five extant genera '' Callistomys'', '' Hoplomys'', '' Myocastor'', ''Proechimys ''Proechimys'' is a genus of South American South America is a contine ...
, 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 extinct genus of Cuban rodents 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 ...
'' *** †''
Brotomys The Hispaniolan edible rat (''Brotomys voratus'') is a Holocene extinction, recently extinct species of rodent in the family Echimyidae. It is the only species in the genus ''Brotomys''. It was Endemism, endemic to the island of Hispaniola in the ...
'' *** †''
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 m ...
'' *** †'' Puertoricomys'' ** Subfamily
Eumysopinae 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 most ecologically diverse, with members ranging from fully arboreal to terre ...
*** †'' 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. Most species are restricted to Cuba, but species are known from all of the Greater Antilles, as well as Th ...
- 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. Most species are restricted t ...
'' ***†''
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'', 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 postcanines 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 or phylogeny is a graphical representation which shows the evolutionary history between a set of species or Taxon, taxa during a specific time.Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, M ...
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 used ...
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 ''Pr ...
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 par ...
, ''
Carterodon Owl's spiny rat (''Carterodon sulcidens'') is a rodent species in the family Echimyidae found in Brazil. It is the only species in the genus ''Carterodon''. Owl's spiny rat has evolved characteristics such as a heightened ability to dig in open ...
'', and the Capromyidae. The first major clade contains a majority of arboreal genera (e.g., ''
Phyllomys ''Phyllomys'' is a genus of arboreal Echimyidae, 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, ...
'', ''
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 these ...
''), a few terrestrial taxa (e.g., ''
Proechimys ''Proechimys'' is a genus of South American South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described ...
''), 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. It is the only species in the genus ''Carterodon''. Owl's spiny rat has evolved characteristics such as a heightened ability to dig in open ...
''), 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 Most of the Caribbean countries are islands in the Caribbean Sea, with only a few in inland lakes. The largest islands include Cuba, Hispaniola, Jamaica and Puerto Rico. Some of the smaller islands are referred to as a ''rock'' or ''reef.'' ''I ...
(Capromyidae).


References and notes

{{Authority control Rodent families Extant Chattian first appearances Taxa named by John Edward Gray