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Accurate taxonomy is central to the study of biological diversity, as it provides the needed evolutionary framework for taxon sampling and interpreting results. While the number of recognized species in the class Mammalia has increased through time, tabulation of those increases has relied on the sporadic release of revisionary compendia like the Mammal Species of the World (MSW) series. Here, we present the Mammal Diversity Database (MDD), a digital, publically accessible, and updateable list of all mammalian species, now available online: https://mammaldiversity.org. The MDD will continue to be updated as manuscripts describing new species and higher taxonomic changes are released. Starting from the baseline of the 3rd edition of MSW (MSW3), we performed a review of taxonomic changes published since 2004 and digitally linked species names to their original descriptions and subsequent revisionary articles in an interactive, hierarchical database. We found 6,495 species of currently recognized mammals (96 recently extinct, 6,399 extant), compared to 5,416 in MSW3 (75 extinct, 5,341 extant)—an increase of 1,079 species in about 13 years, including 11 species newly described as having gone extinct in the last 500 years. We tabulate 1,251 new species recognitions, at least 172 unions, and multiple major, higher-level changes, including an additional 88 genera (1,314 now, compared to 1,226 in MSW3) and 14 newly recognized families (167 compared to 153). Analyses of the description of new species through time and across biogeographic regions show a long-term global rate of ~25 species recognized per year, with the Neotropics as the overall most species-dense biogeographic region for mammals, followed closely by the Afrotropics. The MDD provides the mammalogical community with an updateable online database of taxonomic changes, joining digital efforts already established for amphibians (AmphibiaWeb, AMNH’s Amphibian Species of the World), birds (e.g., Avibase, IOC World Bird List, HBW Alive), non-avian reptiles (The Reptile Database), and sh (e.g., FishBase, Catalog of Fishes).
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1
How many species of mammals are there?
Connor J. Burgin,1 JoCelyn P. Colella,1 PhiliP l. Kahn, and nathan S. uPham*
Department of Biological Sciences, Boise State University, 1910 University Drive, Boise, ID 83725, USA (CJB)
Department of Biology and Museum of Southwestern Biology, University of New Mexico, MSC03-2020, Albuquerque, NM 87131,
USA (JPC)
Museum of Vertebrate Zoology, University of California, Berkeley, CA 94720, USA (PLK)
Department of Ecology and Evolutionary Biology, Yale University, New Haven, CT 06511, USA (NSU)
Integrative Research Center, Field Museum of Natural History, Chicago, IL 60605, USA (NSU)
1Co-first authors.
* Correspondent: nathan.upham@yale.edu
Accurate taxonomy is central to the study of biological diversity, as it provides the needed evolutionary framework
for taxon sampling and interpreting results. While the number of recognized species in the class Mammalia has
increased through time, tabulation of those increases has relied on the sporadic release of revisionary compendia
like the Mammal Species of the World (MSW) series. Here, we present the Mammal Diversity Database
(MDD), a digital, publically accessible, and updateable list of all mammalian species, now available online:
https://mammaldiversity.org. The MDD will continue to be updated as manuscripts describing new species and
higher taxonomic changes are released. Starting from the baseline of the 3rd edition of MSW (MSW3), we
performed a review of taxonomic changes published since 2004 and digitally linked species names to their
original descriptions and subsequent revisionary articles in an interactive, hierarchical database. We found 6,495
species of currently recognized mammals (96 recently extinct, 6,399 extant), compared to 5,416 in MSW3 (75
extinct, 5,341 extant)—an increase of 1,079 species in about 13 years, including 11 species newly described
as having gone extinct in the last 500 years. We tabulate 1,251 new species recognitions, at least 172 unions,
and multiple major, higher-level changes, including an additional 88 genera (1,314 now, compared to 1,226 in
MSW3) and 14 newly recognized families (167 compared to 153). Analyses of the description of new species
through time and across biogeographic regions show a long-term global rate of ~25 species recognized per year,
with the Neotropics as the overall most species-dense biogeographic region for mammals, followed closely by the
Afrotropics. The MDD provides the mammalogical community with an updateable online database of taxonomic
changes, joining digital efforts already established for amphibians (AmphibiaWeb, AMNH’s Amphibian Species
of the World), birds (e.g., Avibase, IOC World Bird List, HBW Alive), non-avian reptiles (The Reptile Database),
and fish (e.g., FishBase, Catalog of Fishes).
Una taxonomía que precisamente refleje la realidad biológica es fundamental para el estudio de la diversidad
de la vida, ya que proporciona el armazón evolutivo necesario para el muestreo de taxones e interpretación
de resultados del mismo. Si bien el número de especies reconocidas en la clase Mammalia ha aumentado con
el tiempo, la tabulación de esos aumentos se ha basado en las esporádicas publicaciones de compendios de
revisiones taxonómicas, tales como la serie Especies de mamíferos del mundo (MSW por sus siglas en inglés).
En este trabajo presentamos la Base de Datos de Diversidad de Mamíferos (MDD por sus siglas en inglés):
una lista digital de todas las especies de mamíferos, actualizable y accesible públicamente, disponible en la
dirección URL https://mammaldiversity.org/. El MDD se actualizará con regularidad a medida que se publiquen
artículos que describan nuevas especies o que introduzcan cambios de diferentes categorías taxonómicas. Con la
tercera edición de MSW (MSW3) como punto de partida, realizamos una revisión en profundidad de los cambios
taxonómicos publicados a partir del 2004. Los nombres de las especies nuevamente descriptas (o ascendidas
a partir de subespecies) fueron conectadas digitalmente en una base de datos interactiva y jerárquica con sus
Journal of Mammalogy, 99(1):1–14, 2018
DOI:10.1093/jmammal/gyx147
invited PaPer
© 2018 American Society of Mammalogists, www.mammalogy.org
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descripciones originales y con artículos de revisión posteriores. Los datos indican que existen actualmente 6,495
especies de mamíferos (96 extintas, 6,399 vivientes), en comparación con las 5,416 reconocidas en MSW3 (75
extintas, 5,341 vivientes): un aumento de 1,079 especies en aproximadamente 13 años, incluyendo 11 nuevas
especies consideradas extintas en los últimos 500 años. Señalamos 1,251 nuevos reconocimientos de especies,
al menos 172 uniones y varios cambios a mayor nivel taxonómico, incluyendo 88 géneros adicionales (1,314
reconocidos, comparados con 1,226 en MSW3) y 14 familias recién reconocidas (167 en comparación con 153
en MSW3). Los análisis témporo-geográficos de descripciones de nuevas especies (en las principales regiones
del mundo) sugieren un promedio mundial de descripciones a largo plazo de aproximadamente 25 especies
reconocidas por año, siendo el Neotrópico la región con mayor densidad de especies de mamíferos en el mundo,
seguida de cerca por la region Afrotrópical. El MDD proporciona a la comunidad de mastozoólogos una base de
datos de cambios taxonómicos conectada y actualizable, que se suma a los esfuerzos digitales ya establecidos
para anfibios (AmphibiaWeb, Amphibian Species of the World), aves (p. ej., Avibase, IOC World Bird List, HBW
Alive), reptiles “no voladores” (The Reptile Database), y peces (p. ej., FishBase, Catalog of Fishes).
Key words: biodiversity, conservation, extinction, taxonomy
Species are a fundamental unit of study in mammalogy. Yet spe-
cies limits are subject to change with improved understanding of
geographic distributions, field behaviors, and genetic relation-
ships, among other advances. These changes are recorded in a
vast taxonomic literature of monographs, books, and periodi-
cals, many of which are difficult to access. As a consequence,
a unified tabulation of changes to species and higher taxa has
become essential to mammalogical research and conservation
efforts in mammalogy. Wilson and Reeder’s 3rd edition of
Mammal Species of the World (MSW3), published in November
2005, represents the most comprehensive and up-to-date list of
mammalian species, with 5,416 species (75 recently extinct,
5,341 extant), 1,229 genera, 153 families, and 29 orders. That
edition relied on expertise solicited from 21 authors to deliver
the most comprehensive list of extant mammals then availa-
ble. However, the episodic release of these massive anthologies
(MSW1—Honacki et al. 1982; MSW2—Wilson and Reeder
1993; MSW3—Wilson and Reeder 2005) means that taxo-
nomic changes occurring during or soon after the release of a
new edition may not be easily accessible for over a decade. For
example, MSW3, compared to MSW2, resulted in the addition
of 787 species, 94 genera, and 17 families compared to MSW2
(Solari and Baker 2007). Since the publication of MSW3, there
has been a steady flow of taxonomic changes proposed in peer-
reviewed journals and books; however, changes proposed more
than a decade ago (e.g., Carleton et al. 2006; Woodman et al.
2006) have yet to be incorporated into a Mammalia-wide refer-
ence taxonomy. This lag between the publication of taxonomic
changes and their integration into the larger field of mammal-
ogy inhibits taxonomic consistency and accuracy in mam-
malogical research, and—at worst—it can impede the effective
conservation of mammals in instances where management deci-
sions depend upon the species-level designation of distinctive
evolutionary units.
The genetic era has catalyzed the discovery of morphologi-
cally cryptic species and led to myriad intra- and interspecific
revisions, either dividing species (splits) or uniting them (lumps).
Many groups of mammals are taxonomically complex and in
need of further revision, especially those that have received
relatively little systematic attention or are morphologically or
behaviorally cryptic (e.g., shrews, burrowing mammals). For
example, the phylogenetic placement of tenrecs and golden
moles (families: Tenrecidae and Chrysochloridae) has long been
a point of taxonomic contention, having variously been included
within Insectivora, Eulipotyphla, and Lipotyphla. Taxonomic
assignment of this group was only conclusively resolved when
genetic data (Madsen et al. 2001; Murphy et al. 2001), as corrob-
orated by morphology (Asher et al. 2003), aligned Tenrecidae
and Chrysochloridae in the order Afrosoricida and found it
allied to other African radiations in the superorder Afrotheria
(Macroscelidea, Tubulidentata, Hyracoidea, Proboscidea,
Sirenia). As analytical methods evolve and techniques become
more refined, mammalian taxonomy will continue to change,
making it desirable to create an adjustable list of accepted spe-
cies-level designations and their hierarchical placement that can
be updated on a regular basis. Such a list is needed to promote
consistency and accuracy of communication among mammalo-
gists and other researchers.
Here, using MSW3 as a foundation, we provide an up-to-
date list of mammal species and introduce access to this spe-
cies list as an amendable digital archive: the Mammal Diversity
Database (MDD), available online at http://mammaldiversity.
org. We compare our list to that of MSW3 to quantify changes
in mammalian taxonomy that have occurred over the last
13 years and evaluate the distribution of species diversity and
new species descriptions across both geography and time. We
intend the MDD as a community resource for compiling and
disseminating published changes to mammalian taxonomy in
real time, rather than as a subjective arbiter for the relative
strength of revisionary evidence, and hence defer to the peer-
reviewed literature for such debates.
Materials and Methods
Starting from those species recognized in MSW3, we reviewed
> 1,200 additional taxonomic publications appearing after
MSW3’s end-2003 cutoff date in order to compile a list of
every recognized mammal species. In addition to evaluating
peer-reviewed manuscripts, other major references included the
Handbook of the Mammals of the World volumes 1–6 (Wilson
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and Mittermeier 2009, 2011, 2014, 2015; Mittermeier et al.
2013; Wilson et al. 2016), Mammals of South America volumes
1 and 2 (Gardner 2007; Patton et al. 2015), Mammals of Africa
volumes 1–6 (Kingdon et al. 2013), Rodents of Sub-Saharan
Africa (Monadjem et al. 2015), Taxonomy of Australian
Mammals (Jackson and Groves 2015), and Ungulate Taxonomy
(Groves and Grubb 2011). We linked each species to its pri-
mary, descriptive publication and if a species was taxonomi-
cally revised since 2004, the associated revisionary publications
also were linked. The list was curated for spelling errors and
compared to the species recognized in MSW3 to determine the
total change in the number of recognized species over the inter-
val 1 January 2004 to 15 August 2017; the latter date was our
cutoff for reviewing literature. As with MSW3 and the IUCN
(2017) RedList, species totals for the MDD include mamma-
lian species that have gone extinct during the last 500 years,
an arbitrary period of time used to delimit species “recently
extinct”. The IUCN taxonomy was downloaded on 28 June
2017.
We considered “de novo” species descriptions to be those
species recognized since MSW3 and named with novel spe-
cies epithets (post-MSW3 proposal date), whereas “splits” are
species established by resurrecting an existing name (i.e., ele-
vated subspecies or synonym, and pre-MSW3 proposal). We
based these 2 bins of new species on the epithet authority year
to enable downstream analyses of species discovery trends.
However, we acknowledge that this categorization is not precise
regarding the more complex (and biologically interesting) issue
of how many species were derived from new field discover-
ies of distinctive populations versus the recognition of multiple
species within named forms (Patterson 1996). Nevertheless, we
expected the de novo category to encompass those field dis-
coveries along with other types of species descriptions, and the
splits category to encompass instances where existing names
are elevated or validated, both of which are categories warrant-
ing future investigation.
In addition to taxonomic ranks (order, family, genus, species)
and primary data links, MDD species information includes
the year of description, scientific authority, and geographic
occurrence by biogeographic region. Here, we approximate
the biogeographic realms defined by the World Wildlife Fund
(Olson and Dinerstein 1998; Olson et al. 2001), with the excep-
tion that we classified countries split across multiple biogeo-
graphic realms as belonging exclusively to the realm covering
the majority of that country. We defined the Nearctic realm as
all of North America, including Florida, Bermuda, and all of
Mexico. The Neotropical realm included all of South America,
Central America, and the insular Caribbean. The Palearctic
realm included all of Europe, northern Asia (including all of
China), Japan, and northern Africa (Egypt, Algeria, Tunisia,
Morocco, Western Sahara, Canary Islands, and the Azores). The
Indomalayan realm included southern and southeastern Asia
(Pakistan, India, Nepal, Bhutan, Vietnam, Laos, Myanmar)
and all islands west of Sulawesi including the Greater Sundas
and Philippines. The Afrotropical realm included all of sub-
Saharan Africa and the Arabian Peninsula, plus Madagascar
and the nearby Indian Ocean islands (e.g., Comoros, Mauritius,
Seychelles). We grouped the Australasian and Oceanian
realms to include a single category for Australia, New
Zealand, Sulawesi, and the islands east of Sulawesi, including
Melanesia, Polynesia, Micronesia, Hawaii, and Easter Island,
but excluding the Palearctic Japanese Bonin Islands. There are
no terrestrial mammal species native to Antarctica. Open-water
and coastal marine species, including the few Antarctic breed-
ing species (e.g., leopard seals, Hydrurga), were grouped sep-
arately. Freshwater species (e.g., river dolphins, river otters)
were sorted by their resident landmass.
Based on our newly curated list, we calculated the number
of new species described each decade since the origin of bi-
nomial nomenclature (Linnaeus 1758) to determine the major
eras of species discovery and taxonomic description. The year
1758 includes all the species described by Linnaeus that are
still currently recognized. For each biogeographic realm, we
calculated the total number of mammalian species recognized
and the number of new species recognized since 2004. Note
that the recognition of new species in a particular region can re-
flect greater research efforts per region or taxon and thus cannot
be extrapolated to the expected number of undiscovered species
in that region. We scaled the number of species by regional land
area (km2World Atlas 2017) to determine the most species-
dense region.
results
The MDD currently lists 6,495 valid species of mammals
(6,399 extant, 96 recently extinct), which is 1,079 more spe-
cies than were recognized in MSW3 (1,058 extant and 21
extinct) and a 19.9% increase in species during about 13 years
(Table 1). The MDD recognizes 1,251 new species described
since MSW3 in categories of splits (720 species; 58%) and de
novo species descriptions (531 species; 42%), indicating that
at least 172 species were lumped together since the release of
MSW3. The MDD documents a total of 1,314 genera (increas-
ing by 88 from MSW3), 167 families (increasing by 14), and
27 orders (decreasing by 2). The MDD also includes 17 domes-
ticated species in the listing to facilitate the association of
Table 1.—Comparison of Mammal Diversity Database (MDD)
taxonomic totals and those of Mammal Species of the World (MSW)
editions 1–3 and the International Union of Conservation of Nature
(IUCN) RedList, version 2017-1.
Taxa MSW1 MSW2 MSW3 IUCN MDD
1982 1993 2005 2017 This study
Species
Total 4,170 4,631a5,416 5,560 6,495
Extinct NA NA 75 85b96
Living NA NA 5,341 5,475 6,399
Living wild NA NA 5,338 5,475 6,382
Genera 1,033 1,135 1,230 1,267 1,314
Families 135 132 153 159 167
Orders 20 26 29 27 27
aCorrected total per Solari and Baker (2007).
bExtinct IUCN mammals include both “EX” (extinct) and “EW” (extinct in
the wild).
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4 JOURNAL OF MAMMALOGY
these derivatives of wild populations with their often abundant
trait data (e.g., DNA sequences, reproductive data). Details
of the full MDD version 1 taxonomy, including associated
citations and geographic region assignments, are provided in
Supplementary Data S1.
The largest mammalian families are in the order Rodentia—
Muridae (834 species versus 730 in MSW3) and Cricetidae
(792 species versus 681 in MSW3)—followed by the chi-
ropteran family Vespertilionidae (493 species versus 407 in
MSW3) and the eulipotyphlan family Soricidae (440 species
versus 376 in MSW3). Unsurprisingly, the 2 most speciose
orders (Rodentia and Chiroptera) witnessed the most species
additions: 371 and 304 species, respectively. The most speciose
rodent family besides Muridae and Cricetidae is Sciuridae (298
species) and 6 rodent families are monotypic: Aplodontiidae,
Diatomyidae, Dinomyidae, Heterocephalidae, Petromuridae,
and Zenkerellidae. The most speciose chiropteran families
along with Vespertilionidae are Phyllostomidae (214 species)
and Pteropodidae (197 species), whereas there is only 1 mono-
typic bat family: Craseonycteridae.
The increased number of recognized genera to 1,314 (from
1,230 in MSW3) results from the demonstrated paraphyly
of several speciose and widely distributed former genera.
This includes Spermophilus, which was split into 8 dis-
tinct genera (Spermophilus, Urocitellus, Callospermophilus,
Otospermophilus, Xerospermophilus, Ictidomys, Poliocitellus,
and NotocitellusHelgen et al. 2009) and Oryzomys, which
was split into 11 genera (Oryzomys, Aegialomys, Cerradomys,
Eremoryzomys, Euryoryzomys, Hylaeamys, Mindomys,
Nephelomys, Oreoryzomys, Sooretamys, and Transandinomys
Weksler et al. 2006). Many smaller generic splits broke 1 genus
into 2 or more genera and often involved the naming of a new
genus, such as with Castoria (formerly AkodonPardiñas
et al. 2016), Paynomys (formerly ChelemysTeta et al. 2016),
and Petrosaltator (formerly ElephantulusDumbacher 2016).
Other genera were described on the basis of newly discovered
taxa, such as Laonastes (Jenkins et al. 2005), Xeronycteris
(Gregorin and Ditchfield 2005), Rungwecebus (Davenport
et al. 2006), Drymoreomys (Percequillo et al. 2011), and
Paucidentomys (Esselstyn et al. 2012). The most speciose cur-
rently recognized genera are Crocidura (197 species), Myotis
(126 species), and Rhinolophus (102 species). These also are
the only genera of mammals that currently exceed 100 recog-
nized and living species, with Rhinolophus reaching this level
only recently.
Higher-level taxonomy also was significantly altered since
2004, with the recognition of 14 additional families and 2
fewer orders than MSW3. In the MDD, we included 3 families
(†Megaladapidae, †Palaeopropithecidae, †Archaeolemuridae)
that were not in MSW3 but that may have gone extinct in the
last 500 years (McKenna and Bell 1997; Montagnon et al. 2001;
Gaudin 2004; Muldoon 2010). The net addition of 11 other
families in the MDD are the result of taxonomic splits and new
taxon discoveries, as well as families lumped since MSW3.
For example, Dipodidae was split into 3 families (Dipodidae,
Zapodidae, Sminthidae—Lebedev et al. 2013), Hipposideridae
into 2 (Hipposideridae, Rhinonycteridae—Foley et al. 2015), and
Bathyergidae into 2 (Bathyergidae, Heterocephalidae—Patterson
and Upham 2014). One family, Diatomyidae, was added based
on a species discovery (Laonastes aenigmamusJenkins et al.
2005), although it was already known as a prehistorically extinct
family (Dawson et al. 2006). Additional newly recognized
families are Chlamyphoridae, Cistugidae, Kogiidae, Lipotidae,
Miniopteridae, Pontoporiidae, Potamogalidae, Prionodontidae,
and Zenkerellidae. Three families recognized in MSW3 have
since been subsumed: Myocastoridae and Heptaxodontidae inside
Echimyidae (Emmons et al. 2015), and Aotidae inside Cebidae
(Schneider and Sampaio 2015; Dumas and Mazzoleni 2017).
Note that Capromyidae is still recognized at the family level
(Fabre et al. 2017). The order Cetacea also experienced major revi-
sions, and is now included within the order Artiodactyla based on
genetic and morphological data (Gatesy et al. 1999; Adams 2001;
Asher and Helgen 2010). Soricomorpha and Erinaceomorpha
also are grouped together in the order Eulipotyphla, given their
shared evolutionary history demonstrated by genetic analyses
(Douady et al. 2002; Meredith et al. 2011).
On average, since 1758, 24.95 species have been described
per decade, including 3 major spikes in species recognition in the
1820–1840s, 1890–1920s, and 2000–2010s (Fig. 1). These bursts
of systematic and taxonomic development were followed by 2
major troughs from about 1850–1880 and 1930–1990 (Fig. 1).
Currently, we detect an accelerating rate of species description
per decade, increasing from the 1990s (207 species), 2000s (341
species), and 2010s so far (298 species). A linear regression on
these data suggests that if trends in mammalian species discov-
ery continue, 120.46 species are yet to be discovered this decade,
potentially resulting in a total of 418 new species to be recog-
nized between 2010 and 2020 (R2 = 0.97, P < 0.000; Fig. 1).
Across biogeographic regions, the Neotropics harbors the
greatest number of currently recognized mammalian species
(1,617 species), followed by the Afrotropics (1,572 species),
and the Palearctic (1,162 species), whereas Australasia-Oceania
has the least (527 species) (Fig. 2). The Neotropics also has
the most newly recognized species (362 species—169 de novo
and 193 split), again followed by the Afrotropics (357 spe-
cies—158 de novo and 199 split), and with the fewest new spe-
cies described from Australasia-Oceania (48 species—18 de
novo and 30 split). Other categories included the marine (124
total species—4 de novo and 5 split), domesticated (17 total spe-
cies—0 de novo and 2 split), and extinct (96 total species—7 de
novo and 4 split; Fig. 2; Table 2) categories. When weighting
the biogeographic realms by land area, we find the Neotropics
and Afrotropics are also the most species-dense biogeographic
regions (85.1 and 71.1 species per km2, respectively), followed
closely by Australasia-Oceania (61.4 species per km2; Table 2).
In all realms except the Indomalayan, more species were recog-
nized via taxonomic splits than by de novo descriptions.
discussion
Mammalogists have a collective responsibility to serve the most
current taxonomic information about mammalian biodiversity
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INVITED PAPER—MAMMALIAN SPECIES DIVERSITY 5
to the general public. The need for mammalian taxonomy to
reflect our current understanding of species boundaries and
evolutionary relationships is only expected to grow as efforts to
synthesize “big data” increase in frequency, scope, and sophis-
tication. Studies at this macroscale address major questions
in evolution, ecology, and biodiversity conservation across
the tree of life (e.g., Rabosky et al. 2012; Hedges et al. 2015;
Hinchliff et al. 2015), yielding results relevant to global issues
of sustainability that require our best data on biodiversity
(Pascual et al. 2017). Mammalogists, in turn, benefit from easy
access to this biodiversity data for purposes of study design,
classroom teaching, analyses, and writing. The release of the
MDD therefore addresses a key need in the mammalogical and
global biodiversity communities alike. Whether we study the
behavioral ecology of desert rodents or the macroevolution of
tetrapods, biologists collectively need accurate measurements
of species diversity—the most commonly assessed (but not the
only) dimension of biodiversity (Jarzyna and Jetz 2016).
The MDD represents the most comprehensive taxonomic
compendium of currently recognized mammals, documenting
Fig. 1.—Cumulative and decadal descriptions of taxonomically valid extant mammal species from 1758 to 15 August 2017.
Fig. 2.—The number of mammalian species distributed in each biogeographical region: Palearctic, Afrotropic, Indomalayan, Nearctic, Neotropic,
and Australasia-Oceania (i.e., Aust-Oceania), with marine, extinct, and domestic species in separate categories. Each group is divided into species
recognized in both MSW3 and MDD, and new species in the MDD in categories of newly coined species epithet (de novo) versus existing species
epithet (splits). The dot within each bar indicates the relative species density per km2 land area, values are available in Table 2. MDD = Mammal
Diversity Database; MSW3 = 3rd edition of Mammal Species of the World.
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6 JOURNAL OF MAMMALOGY
6,399 extant species (Tables 1 and 3) as well as 96 recently
extinct species for a total of 6,495 species. This database is
updateable and digitally searchable, tracking primary sources
of species descriptions and phylogenetic studies of higher-level
(genus or family) taxonomic changes and compiling them into
a single listing. The MDD thus closes the gap between pro-
posed taxonomic changes and integration into a broader under-
standing of mammalian diversity, and it then distributes this
information to the scientific community and lay public as it is
published in scientific literature. We aim for the MDD to build
on this capacity as a record keeper to be a resource for hosting
histories of taxonomic change. For example, the MDD records
both the description of Tapirus kabomani (Cozzuol et al. 2013)
and the later synonymy of this taxon under T. terrestris (Voss
et al. 2014). Likewise, the revision of Spermophilus ground
squirrels into 8 genera (Helgen et al. 2009) altered the binomial
names of 28 species, a rearrangement that usefully established
generic monophyly, but one that has not been readily summa-
rized for workers without easy access to libraries. The MDD
compiles data on genus transfers published since 2004 across
all of Mammalia, helping to release researchers from undertak-
ing piecemeal taxonomic updates for their projects.
Preliminary findings from the MDD compilation indicate
that Primates has been a nexus of new species discovery, which
is unexpected given their large body sizes. An incredible 148
primate species have been recognized since the publication of
MSW3, including 67 de novo and 81 splits (Tables 1 and 3), a
taxonomic outcome that is striking for our closest human rela-
tives. Taxonomic revisions have centered around New World
monkey families (Cebidae—Boubli et al. 2012; Pitheciidae—
Marsh 2014) and many de novo species descriptions also
occurred among Malagasy lemurs (Cheirogaleidae—Lei et al.
2014; Lepilemuridae—Louis et al. 2006). However, persis-
tent taxonomic uncertainty within the family Cercopithecidae
(Groves 2007a, 2007b; Mittermeier et al. 2013) suggests that
the species-level diversity of Primates is not yet stable and will
continue to fluctuate.
Among other taxonomic changes, the MDD documents the
addition of 371 species of Rodentia, 304 species of Chiroptera,
86 species of Eulipotyphla, and 227 species of Artiodactyla,
including many species from historically well-studied geo-
graphic regions (Table 2; Rausch et al. 2007; Castiglia et al.
2017). While the addition of > 300 species each of rodents and
bats is unsurprising given their existing diversity, these clades
may reasonably contain disproportionally high levels of cryptic
diversity (e.g., Ruedi and Mayer 2001; Belfiore et al. 2008),
and thus the application of genetic sequence data may continue
to yield greater insights. Within Eulipotyphla (most particularly
in shrews), we expect that the discovery of new species will
continue given their rate of recent discoveries and frequency of
morphological crypsis (Esselstyn et al. 2013). The species rich-
ness in Sorex (86 species) and Crocidura (197 species) suggests
that genus-level revisions are needed and, when conducted, are
likely to yield further taxonomic rearrangements (Castiglia
et al. 2017; Matson and Ordóñez-Garza 2017).
The MDD includes a total of 465 species of non-cetacean
Artiodactyla and Perissodactyla recognized by Groves and
Grubb (2011) with select modifications based on taxonomic
refinements published after the release of the latter (e.g., 4 spe-
cies of Giraffa [Bercovitch et al. 2017] versus 8 [Groves and
Grubb 2011]). This total compares to 240 species in these or-
ders recognized in MSW3 (> 93% increase). Although some
researchers have argued that the changes proposed by Groves
and Grubb (2011) exemplify an extreme form of taxonomic in-
flation (Lorenzen et al. 2012; Zachos et al. 2013; Harley et al.
2016), the increase in species richness is comparable to concur-
rent rates of increase in the richness of Rodentia, Chiroptera,
Eulipotyphla, and Primates. For now, inclusion of the tax-
onomy of Groves and Grubb (2011) in the MDD ensures that
these taxa are vetted by the greater mammalogical community
using multiple tiers of evidence (de Queiroz et al. 2007; Voss
et al. 2014).
Following the publication of Linnaeus’s 10th edition of
Systema Naturae in 1758, the number of described species of
mammals has increased at various rates, punctuated by factors
including the efforts of prolific systematists and world events
(Fig. 1). For example, Oldfield Thomas (1858–1929) of the
British Museum (now the Natural History Museum, London),
considered one of the “greatest taxonomists […] who ever
lived” (Flannery 2012), was responsible for nearly 3,000 new
names for genera, species, and subspecies (Hill 1990). In turn,
reduced rates of species descriptions in the mid-20th cen-
tury may be linked to periods of political instability and lim-
ited scientific activity during World War I (1914–1918) and II
(1939–1945). Methodological innovations such as polymer-
ase chain reaction (PCR—Mullis et al. 1989) may have driven
Table 2.—The total number of mammal species in the Mammal Diversity Database (MDD) as compared to Mammal Species of the World, vol-
ume 3 (MSW3) that live within each biogeographic realm and those belonging to domestic and extinct categories. Numbers correspond to Fig. 2.
Note that some species are found within multiple regions, so column totals do not correspond to taxonomic totals.
Category Total species Shared with MSW3 De novo Split Area (million km2) Density (species/km2)
Neotropic 1,617 1,255 169 193 19.0 85.1
Afrotropic 1,572 1,215 158 199 22.1 71.1
Palearctic 1,162 938 48 176 54.1 21.5
Indomalaya 954 774 97 83 7.5 12.7
Nearctic 697 628 15 54 22.9 30.4
Aust-Oceania 527 479 18 30 8.6 61.4
Marine 124 115 4 5
Domestic 17 15 2
Extinct 96 85 7 4
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INVITED PAPER—MAMMALIAN SPECIES DIVERSITY 7
Table 3.—Totals of the genera and species per families and orders currently listed in the Mammal Diversity Database (MDD) online compila-
tion, along with new species described since Mammal Species of the World volume 3 (MSW3) in categories of split or de novo, based on whether
the specific epithet already existed or was newly coined, respectively.
Genera Species New species since MSW3
Splits De novo
Class Mammalia 1,314 6,495 720 531
Subclass Prototheria 3 5
Order Monotremata 3 5
Family Ornithorhynchidae 1 1
Family Tachyglossidae 2 4
Subclass Theria 1,311 6,490 720 531
Infraclass Marsupialia 91 379 32 29
Order Didelphimorphia 18 111 15 18
Family Didelphidae 18 111 15 18
Order Paucituberculata 3 7 1
Family Caenolestidae 3 7 1
Order Microbiotheria 1 3 2
Family Microbiotheriidae 1 3 2
Order Notoryctemorphia 1 2
Family Notoryctidae 1 2
Order Dasyuromorpha 19 78 5 5
Family Dasyuridae 17 76 5 5
Family Myrmecobiidae 1 1
Family †Thylacinidae 1 1
Order Peramelemorphia 8 23 1 1
Family †Chaeropodidae 1 1
Family Peramelidae 6 20 1 1
Family Thylacomyidae 1 2
Order Diprotodontia 41 155 11 2
Family Acrobatidae 2 3 1
Family Burramyidae 2 5
Family Hypsiprymnodontidae 1 1
Family Macropodidae 13 67 3
Family Petauridae 3 12 1
Family Phalangeridae 6 30 3 1
Family Phascolarctidae 1 1
Family Potoroidae 4 12 1
Family Pseudocheiridae 6 20 3
Family Tarsipedidae 1 1
Family Vombatidae 2 3
Infraclass Placentalia 1,220 6,111 684 502
Superorder Afrotheria 34 89 8 6
Order Tubulidentata 1 1
Family Orycteropodidae 1 1
Order Afrosoricida 20 55 1 3
Family Chrysochloridae 10 21
Family Potamogalidaea2 3
Family Tenrecidae 8 31 1 3
Order Macroscelidea 5 20 2 3
Family Macroscelididae 5 20 2 3
Order Hyracoidea 3 5 1
Family Procaviidae 3 5 1
Order Proboscidea 2 3
Family Elephantidae 2 3
Order Sirenia 3 5
Family Dugongidae 2 2
Family Trichechidae 1 3
Superorder Xenarthra 14 30
Order Cingulata 9 20
Family Chlamyphoridaeb8 13
Family Dasypodidae 1 7
Order Pilosa 5 10
Family Bradypodidae 1 4
Family Cyclopedidae 1 1
Family Megalonychidae 1 2
Family Myrmecophagidae 2 3
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8 JOURNAL OF MAMMALOGY
Genera Species New species since MSW3
Splits De novo
Superorder Euarchontoglires 616 3,194 285 249
Order Scandentia 4 24 4
Family Ptilocercidae 1 1
Family Tupaiidae 3 23 4
Order Dermoptera 2 2
Family Cynocephalidae 2 2
Order Primates 84 518 81 67
Family †Archaeolemuridaec1 2
Family Atelidae 4 25 3
Family Cebidaed11 89 27 2
Family Cercopithecidae 23 160 24 5
Family Cheirogaleidae 5 40 1 20
Family Daubentoniidae 1 1
Family Galagidae 6 20 2 2
Family Hominidae 4 7
Family Hylobatidae 4 20 3 2
Family Indriidaee3 19 2 6
Family Lemuridae 5 21 2
Family Lepilemuridae 1 26 16
Family Lorisidae 4 15 6 1
Family †Megaladapidaec1 1
Family †Palaeopropithecidaec1 1
Family Pitheciidae 7 58 9 9
Family Tarsiidae 3 13 2 4
Order Lagomorpha 13 98 10 1
Family Leporidae 11 67 5 1
Family Ochotonidae 1 30 5
Family †Prolagidae 1 1
Order Rodentia 513 2,552 190 181
Family Abrocomidae 2 10
Family Anomaluridae 2 6
Family Aplodontiidae 1 1
Family Bathyergidae 5 21 3 4
Family Calomyscidae 1 8
Family Capromyidae 7 17
Family Castoridae 1 2
Family Caviidae 6 21 3
Family Chinchillidae 3 7 1
Family Cricetidae 145 792 75 61
Family Ctenodactylidae 4 5
Family Ctenomyidae 1 69 5 6
Family Cuniculidae 1 2
Family Dasyproctidae 2 15 2 1
Family Diatomyidaef1 1 1
Family Dinomyidae 1 1
Family Dipodidae 13 37 3
Family Echimyidaeg25 93 6 3
Family Erethizontidae 3 17 1 2
Family Geomyidae 7 41 8 1
Family Gliridae 9 29 1
Family Heterocephalidaeh1 1
Family Heteromyidae 5 66 6 2
Family Hystricidae 3 11
Family Muridae 157 834 41 84
Family Nesomyidae 21 68 1 6
Family Octodontidae 7 14 1
Family Pedetidae 1 2
Family Petromuridae 1 1
Family Platacanthomyidae 2 5 2 1
Family Sciuridae 62 298 18 5
Family Sminthidaei1 14 2
Table 3.Continued
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INVITED PAPER—MAMMALIAN SPECIES DIVERSITY 9
Genera Species New species since MSW3
Splits De novo
Family Spalacidae 7 28 8
Family Thryonomyidae 1 2
Family Zapodidaei3 12 6 1
Family Zenkerellidaej1 1
Superorder Laurasiatheria 556 2,798 399 247
Order Eulipotyphlak56 527 23 63
Family Erinaceidae 10 24
Family †Nesophontidae 1 6
Family Solenodontidae 1 3
Family Soricidae 26 440 16 55
Family Talpidae 18 54 7 8
Order Chiroptera 227 1,386 130 174
Family Cistugidael1 2
Family Craseonycteridae 1 1
Family Emballonuridae 14 54 3
Family Furipteridae 2 2
Family Hipposideridae 7 88 6 8
Family Megadermatidae 5 6 1
Family Miniopteridael1 35 7 9
Family Molossidae 19 122 12 13
Family Mormoopidae 2 17 8
Family Mystacinidae 1 2
Family Myzopodidae 1 2 1
Family Natalidae 3 11 3
Family Noctilionidae 1 2
Family Nycteridae 1 16
Family Phyllostomidae 62 214 22 37
Family Pteropodidae 45 197 5 12
Family Rhinolophidae 1 102 10 14
Family Rhinonycteridaem4 9 1 3
Family Rhinopomatidae 1 6 1 1
Family Thyropteridae 1 5 2
Family Vespertilionidae 54 493 55 70
Order Carnivora 130 305 23 2
Family Ailuridae 1 2 1
Family Canidae 13 39 3
Family Eupleridae 7 8
Family Felidae 14 42 5
Family Herpestidae 16 36 2
Family Hyaenidae 3 4
Family Mephitidae 4 12 1
Family Mustelidae 23 64 5 1
Family Nandiniidae 1 1
Family Odobenidae 1 1
Family Otariidae 7 16
Family Phocidae 14 19
Family Prionodontidaen1 2
Family Procyonidae 6 14 2 1
Family Ursidae 5 8
Family Viverridae 14 37 4
Order Pholidota 3 8
Family Manidae 3 8
Order Perissodactyla 8 21 4
Family Equidae 1 12 4
Family Rhinocerotidae 4 5
Family Tapiridae 3 4
Order Artiodactylao132 551 219 8
Family Antilocapridae 1 1
Family Balaenidae 2 4
Family Balaenopteridae 2 8 1
Family Bovidae 54 297 152 2
Family Camelidae 2 7 1
Table 3.Continued
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10 JOURNAL OF MAMMALOGY
later bursts of species descriptions by allowing morphologically
cryptic but genetically divergent evolutionary lineages to be
recognized as species. For example, over one-half of the spe-
cies described since 2004 appear to have stemmed from taxo-
nomic splits (~58%), many based in part or whole on genetic
data, to go with at least 172 species unions (lumps) during the
same period. As we continue to progress within the genomic era,
where data on millions of independent genetic loci can be read-
ily generated for taxonomic studies, there is a growing under-
standing that hybridization and introgression commonly occur
among mammalian species that may otherwise maintain genetic
integrity (e.g., Larsen et al. 2010; Miller et al. 2012; vonHoldt
et al. 2016). Characterizing species and their boundaries using
multiple tiers of evidence will continue to be essential given the
profound impact of species delimitation on legislative decisions
(e.g., U.S. Endangered Species Act of 1973—Department of the
Interior, U.S. Fish and Wildlife Service 1973).
At the current rate of taxonomic description of mammals
(~25 species/year from 1750 to 2017), we predict that 7,342
mammalian species will be recognized by 2050 and 8,590 by
2100. Alternatively, if we consider the increased rate of taxo-
nomic descriptions since the advent of PCR (~30 species/year
from 1990 to 2017), our estimates increase to 7,509 species
recognized by 2050 and 9,009 by 2100. These estimates sur-
pass Reeder and Helgen’s (2007) prediction of > 7,000 total
mammalian species, but echo their observation that mammals
contain considerably greater species diversity than is com-
monly recognized. Remarkably, the same estimate of ~25 spe-
cies/year was derived somewhat independently from tracking
14 estimates of global diversity (1961–1999—Patterson 2001)
and from species-level changes between MSW2 and MSW3
(Reeder and Helgen 2007), thereby affirming the robustness of
that estimate across both data sources and eras.
Assumed in all taxonomic forecasts is the stability of global
ecosystems, scientific institutions, and natural history collections.
With mammals being disproportionately impacted by human-
induced extinctions (Ceballos et al. 2017), especially in insular
regions like the Caribbean (Cooke et al. in press), efforts to protect
threatened habitats and their resident mammalian species are key
to the continued persistence, existence, and discovery of mammals.
The Neotropics is the most species-dense biogeographic region in
the world, followed closely by the Afrotropics and Australasia-
Oceania, the latter of which is one of the least explored terrestrial
regions on Earth, with the second fewest de novo species descrip-
tions (18 species; Table 2). Inventory efforts may thus be fruitfully
prioritized in northern Australia, Melanesia, Sulawesi, and other
oceanic islands east of Wallace’s Line. However, we note that
obtaining collecting permissions is a barrier to species description
in any region. The continued description and discovery of mamma-
lian species diversity hinges on investment in both natural history
collecting and in the physical collections that house the specimens
essential for taxonomic research. Natural history collections are
Table 3.Continued
Genera Species New species since MSW3
Splits De novo
Family Cervidae 18 93 43
Family Delphinidae 17 40 3 3
Family Eschrichtiidae 1 1
Family Giraffidae 2 5 3
Family Hippopotamidae 2 4
Family Iniidae 1 3 1 1
Family Kogiidaep1 2
Family Lipotidaeq1 1
Family Monodontidae 2 2
Family Moschidae 1 7
Family Neobalaenidae 1 1
Family Phocoenidae 3 7 1
Family Physeteridae 1 1
Family Platanistidae 1 1
Family Pontoporiidaeq1 1
Family Suidae 6 28 11
Family Tayassuidae 3 5 2
Family Tragulidae 3 10 1 1
Family Ziphiidae 6 22 1
aSplit from Tenrecidae.
bSplit from Dasypodidae.
cRecently extinct families not included in MSW3.
dIncludes Aotidae and Callitrichidae.
eWas spelled as “Indridae” in MSW3.
fRecognized as extant based on Laonastes aenigmamus.
gIncludes Heptaxodontidae and Myocastoridae.
hSplit from Bathyergidae.
iSplit from Dipodidae.
jSplit from Anomaluridae.
kIncludes Soricomorpha and Erinaceomorpha.
lSplit from Vespertilionidae.
mSplit from Hipposideridae.
nSplit from Felidae.
oIncludes Cetacea.
pSplit from Physeteridae.
qSplit from Iniidae.
†Extinct.
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INVITED PAPER—MAMMALIAN SPECIES DIVERSITY 11
repositories for the genetic and morphological vouchers used to
describe every new species listed in the MDD, a fact that high-
lights the indispensable role of museums and universities in under-
standing species and the ecosystems in which they live (McLean
et al. 2015). As our planet changes, the need to support geographi-
cally broad and site-intensive biological archives only grows in rel-
evance. Collections represent time series of change in biodiversity
and often harbor undiscovered species (e.g., Helgen et al. 2013),
including those vulnerable or already extinct.
Acting under the supervision of the American Society
of Mammalogists’ Biodiversity Committee, the MDD has
a 2018–2020 plan to further integrate synonym data, track
Holocene-extinct taxa, and add links to outside data sources.
While full synonymies are not feasible, inclusion of common
synonyms will facilitate tracking taxonomic changes through
time, especially within controversial groups (e.g., Artiodactyla
and Perissodactyla—Groves and Grubb 2011). Controversial
taxonomic assignments also will be “flagged” as tentative
or pending further scientific investigation. The MDD aims
to link taxon entries to a variety of relevant per-species and
per-higher taxon data pages on other web platforms, includ-
ing geographic range maps, trait database entries, museum
records, genetic resources, and other ecological information.
Mammalian Species accounts, published by the American
Society of Mammalogists since 1969 and consisting of over
950 species-level treatments, will be linked to relevant MDD
species pages, including synonym-based links. In this manner,
the MDD’s efforts parallel initiatives in other vertebrate taxa
to digitize taxonomic resources (amphibians—AmphibiaWeb
2017; Amphibian Species of the World—Frost 2017; birds:
Avibase—LePage et al. 2014; IOC World Bird List—Gill
and Donsker 2017; the Handbook of the Birds of the World
Alive—del Hoyo et al. 2017; non-avian reptiles, turtles, croco-
diles, and tuatara—Uetz et al. 2016; and bony fish: FishBase—
Froese and Pauly 2017; Catalog of Fishes—Eschmeyer et al.
2017). The new mammalian taxonomic database summarized
herein aims to advance the study of mammals while bringing
it to par with the digital resources available in other tetrapod
clades, to the benefit of future mammalogists and non-mam-
malogists alike.
acknowledgMents
We are grateful to the American Society of Mammalogists for
funding this project, and as well as for logistical support from the
NSF VertLife Terrestrial grant (#1441737). We thank J. Cook,
D. Wilson, B. Patterson, W. Jetz, M. Koo, J. Esselstyn, E. Lacey,
D. Huckaby, L. Ruedas, R. Norris, D. Reeder, R. Guralnick, J.
Patton, E. Heske, and other members of the ASM Biodiversity
Committee for advice, support, and input about this initiative.
suppleMentary data
Supplementary data are available at Journal of Mammalogy online.
Supplementary Data SD1.— Details of the full Mammal
Diversity Database (MDD) version 1 taxonomy, including
associated citations and geographic regions.
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The Council of Higher Education’s (CoHE) National Thesis Center database was searched for postgraduate theses on mammalian animals using a variety of keywords. The theses were classified according to their year of completion, university name, institution, department, level (master’s/PhD), geographical region, province, gender, open access status, subject, animal group, keyword, advisor title, and page range. Between 1976 and 2020, 593 theses on mammalian animals in the domains of science, health, and social sciences were completed in Turkey’s 52 provinces, 88 universities, 12 institutions, and 92 departments. 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Mammal, Mus, wild, stress, Miocene, water, polymerase chain reaction, rat, prey, predator, evolution, Microtus, morphometry, Myotis, oxidative, antioxidant, rattus genetic, banding, mammals, diversity, camera-trap, chromosome, and fauna’ are some of the keywords that appear in thesis abstracts. Rodentia, Carnivora, Artiodactyla, and Chiroptera are the most studied orders. 53.8 percent of academics with the title of professor, 28.2 percent with the title of associate professor, and 17% with the title of doctor lecturer supervised postgraduate theses. This study aimed to examine the history and current state of postgraduate theses on mammalian species in Turkey. The findings are intended to be utilized as a reference for a variety of bibliometric evaluations and research on mammalian animals in the coming years. Keywords: Biology, Database, Gender, Open access
... Unsurprisingly, Brazil leads this region in terms of mammals total biodiversity (Quintela et al., 2020). This rich wild fauna plays an important role in maintaining ecosystems and in the food security and income for thousands of people (Burgin et al., 2018;Ripple et al., 2016;Souto et al., 2019;van Vliet et al., 2015). However, there is an ongoing process of biodiversity loss (Butchart et al., 2010) and, consequently, of the increasing difficulty of human contact with nature, especially in urban centers (Tavares et al., 2012). ...
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... The geographical context of NW Argentina, in the south of the Central Andes, places this region in an area with the highest richness of rodents of South America (Maestri and Patterson, 2016). Additionally, the Order Rodentia is the most diverse group of mammals (with 40% of all known species) and constitutes more than half of all Neotropical mammals (Burgin et al., 2018). These traits make of rodents an excellent biogeographical indicator (Ferro andBarquez, 2009, 2014;Sandoval and Ferro, 2014). ...
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In a recent paper in Current Biology, Fennessy and colleagues [1] conclude that there are four species of giraffe and that their numbers are declining in Africa. Giraffes (Giraffa camelopardalis) are presently classified as one species, with nine subspecies, which are considered ‘Vulnerable’ on the IUCN Red List [2]. The present consensus of one species divided into nine subspecies has previously been questioned (Supplemental information), and Fennessy and colleagues [1] provide another viewpoint on giraffe taxonomy. The fundamental reason for different taxonomic interpretations is that they are based upon different datasets that adopt different statistical techniques and follow different criteria for nomenclature.
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Platyrrhini are a group of Neotropical primates living in central and south America, and have been extensively studied through morphological and molecular data in order to shed light on their phylogeny and evolution. Agreement on the main clades of Neotropical primates has been reached using different approaches, but many phylogenetic nodes remain under discussion. Contrasting hypotheses have been proposed, presumably due to different markers and the presence of polymorphisms in the features considered; furthermore, neither Neotropical primate biodiversity nor their taxonomy are entirely known. In our perspective, a cytogenetic approach can help by making an important contribution to the evaluation of the phylogenetic relationships among Platyrrhini. In this work, molecular cytogenetic data regarding the principal nodes of the Neotropical monkey tree have been reviewed; classical cytogenetic data have also been considered, especially when other data have proven elusive, permitting us to discuss highly derived karyotypes characterized by a wide range of diploid numbers of chromosomes and variable chromosomal evolution with different rearrangement and polymorphism rates.
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