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Tendencias en la investigacion sobre especies terrestres del orden Carnivora.



The members of the order Carnivora display great diversity in their demography, body size, feeding habits, behavior, home range, and habitats (Gittleman et al., 2001). Frequently, members of this order are essential in determining the structure of the terrestrial communities, primarily as a result of their ecological role in trophic webs and their effects on the different groups present in the environments they inhabit (Terborgh et al., 1999; Gittleman and Gompper, 2005).

Many conservation strategies may be based on carnivorous mammals (Loyola et al., 2008). Nevertheless, for these strategies to be effective, they require reliable information on the population dynamics, distribution, and ecological requirements of the species of interest, as well as the impact they have on their ecosystems (Ray et al., 2005).

However, the degree of information for each species is often not homogeneous (Ginsber, 2001), and the compilation and analysis of the issues addressed in the scientific literature are available only for some groups. In previous works, some studies have analyzed the information about particular groups in specific regions, such as those conducted for the family Canidae and Felidae in South America (Medel and Jaksic, 1988; Lucherini et al., 2004; Clavijo and Ramirez, 2009). For some species, such as puma Puma concolor (Lopez-Gonzalez and Gonzalez, 1998), polecat Mustela putorius (Blandford, 2008), genet Genetta genetta (Virgos et al., 1999), and European badger Meles meles (Rope and Mickevicius, 1995), reviews of particular issues, such as diet, are available.

These studies show the advance of knowledge in one or more disciplines, as well as areas where more research is needed for each group. But there is no previous work to provide a perspective on this situation for the order Carnivora as a whole. This work aims to provide information on trends in research on the order Carnivora in terms of topics and species that have been addressed in the scientific literature in the last three decades, and to make recommendations to guide future research.


We reviewed the contents of fifteen internationally circulating scientific journals and recorded the number of articles published between 1980 and 2010 that made reference to one or more species of the order Carnivora. This was considered as the adequate period of time to provide an overview on recent research with this group.

The journals consulted were Studies on Neotropical Fauna and Environmental, The Southwestern Naturalist, Journal of Animal Ecology, Journal of Zoology, Journal of Mammalogy, Mammalia, Mammal Review, Mastozoologia Neotropical, Small Carnivore Conservation, Canid News, Cat News, African Journal of Ecology, Journal of East African Natural History, South African Journal of Wildlife Research, and Raffles Bulletin of Zoology. These journals were selected because of their relevance, availability, and because they published a wide range of topics, including diversity in general (two journals), zoology (six), mammals (four), and carnivores specifically (three). To avoid bias in the percentage of subjects covered, we did not include journals that address a specific topic such as conservation, distribution or diseases, to name a few. Although these journals are edited mainly in Europe (9), the Americas (3), and Africa (3), they include studies addressing species from around the world.

For each article, we recorded the species, general topic, and country in which the study was performed. The articles were classified into 16 topics (Table 1), according to the content expressed in the title and abstract. In the classification of the topics we followed Guevara Chumacero et al. (2005), with some modifications.

The studies were analyzed on both a global level and by continent, as well as by the most studied topics, species, and families, with respect to the total number of published articles. As species richness varies across continents, the number of articles published on each continent was expressed as a proportion (PAC = the number of articles published in each continent/the number of species present in that continent). Likewise, when comparing the number of studies for each family, the number of contributions referring to a given family was expressed as a proportion of the number of species each family contains (PAF = the number of articles published per family/the number of species per family). In addition, the publication Mammalian Species, which includes monographs of species from around the world, was reviewed to get an idea of rogress in knowledge of species of this order. We considered that the species included in this publication are relatively well understood in terms of natural history, distribution and ecology. In this study we followed the taxonomy proposed by Wilson and Mittermeier (2009).


We recorded a total of 2117 studies and of these, the 14.8% were published in the 1980s, the 29.9% in the 1990s, and the 55.3% in the 2000s. Of the 2117 studies found, 758 (35.8%) were performed in the Americas, 476 (22.48%) in Europe, 390 (18.42%) in Africa, and 417 (19.7%) in Asia. The remaining (3.6%) were works spanning more than one continent (72 studies), or were conducted in Oceania (five studies). The most studied topics were distribution, diet, ecology, and natural history, both globally and in each continent (Table 1). In contrast, the topics less addressed were taxonomy, evolution, genetics and zoosanitary issues. Considering the number of contributions as a proportion with respect to the number of carnivore species present on each continent, Europe displayed the highest value (PAC = 20.7), followed by the Americas (PAC = 9.5). Asia and Africa displayed low and similar values (PAC = 4.0 and PAC = 4.6, respectively).

The families with fewer studies were Eupleridae, Mephitidae and Herpestidae, and the families most studied were Mustelidae, Canidae and Felidae, both globally and by continents (Table 2). The articles published on the global level made reference to 196 of the 245 known terrestrial species of Carnivora in the world (Wilson and Mittermeier, 2009). Of these 196 species, the most studied were the European badger M. meles and the coyote Canis latrans, followed by the red fox Vulpes vulpes, and the river otter Lutra lutra, the lion Panthera leo, the wolf Canis lupus, the leopard Panthera pardus, and the puma (Table 3).

From a total of 73 species registered in this review for the Americas, the most studied species were the coyote (84 articles), the puma (55), the black bear Ursus americanus (51), the jaguar Panthera onca (41) and the wolf (38). In Europe, we recorded 35 species, included 11 non-native species, as for example the giant panda Ailuropoda melanoleuca native of Asia, and the fosa Cryptoprocta ferox native of Africa. The most studied species were the European badger (118 articles), the river otter (67), and the red fox (48). In Asia, the leopard and the tiger Panthera tigris (27 articles, each), the giant panda (13 articles), the European mink Mustela lutreola (11), and the Owston's palm civet Chrotogale owstoni (10) were the most studied out of a total of 83 registered species. In Africa, the most studied species were the lion (61 articles), the spotted hyena Crocuta crocuta (42), the cheetah Acinonyx jubatus (35), the African wild dog Lycaon pictus (34), and the leopard (28) out of a total of 60 registered species.

Of the 848 monographs published in Mammalian Species, 105 belong to the order Carnivora. Thus, 42.9% of the carnivores have a monograph: 28% of the monographs corresponds to mustelids (29 species), 26% to canids (28 species), 12% to felines (13 species), and 10% to herpestids (11 species). The remaining 24% is distributed among the other families. For the families Canidae and Hyaenidae, between 80 y 75% of their species already have a monograph in this journal; for the families Ursidae, Mustelidae, and Mephitidae, between 50 and 60%; for the families Procyonidae, Felidae and Herpestidae, between 40 and 30%, and for the families Eupleridae and Viverridae, about 10%. No species of the families Nandiniidae and Prionodontidae (one and two species, respectively), have monographs to the date of this review.


Although the thematic analysis presented here does not represent the entirety of the studies performed on species of the order Carnivora on the global level, and although we did not take into account information published in regional journals, we think it does give a realistic and updated view of the research trends for this order.

Field work for the articles included in this study was concentrated in Europe and the Americas, leaving the remaining continents with a low proportion of studies. The greatest proportion of studies was carried out in Europe (PAC = 20.7), despite the fact that this continent has fewer terrestrial species (24 species). In contrast, the proportion of studies carried out in Asia was low (PAC = 4.0) in spite of the fact that the species richness of this continent is quite high (104 species).

The production of published articles on a global scale has principally focused on few families. Of the 13 terrestrial families that make up the order Carnivora, just three summed 70.3% of the total number of publications: Felidae, Mustelidae and Canidae. Of the remaining ten families, six (Eupleridae, Herpestidae, Mephitidae, Nandiniidae, Viverridae, and Prionodontidae) were scarcely studied. This low number of studies may be due to the species richness of each family. However, the ursids, despite having the same species richness as euplerids (eight species), have a considerably larger proportion of studies. In contrast, although the viverrids constitute almost the same number of species as canids (34 vs. 35 species, respectively), they have a considerably smaller proportion of studies (3.1 vs. 13.1).

In addition, we found a wide variety of species groups that have not been researched within the areas in which they are distributed. For example, the proportions of studies generated on the families Viverridae, Herpestidae, and Eupleridae were low in Africa or in Asia, although these two continents include most of the range of these three families. The families Procyonidae and Mephitidae faced the same situation in the Americas. Heterogeneous representation of families was also seen in the percentage of appearance in Mammalian Species. More than half of the 105 species with a monograph in the journal belong to the families Canidae and Mustelidae, whereas the families Herpestidae, Procyonidae, Viverridae, and Eupleridae are little represented. That is, close to 70% of the species that makes up these families lack of the general descriptions provided by this journal.

The studies published over the last three decades are concentrated on few species, generally large in size, habitat generalists, or charismatic, such as M. meles, L. lutra, C. latrans, P leo, and A. melanoleuca. In contrast, other species groups, generally little charismatic, small sized, or those that may be difficult to observe and study due to their habitats or behavior, have received less attention. This is the case of many small cats, the procyonids Nasuella olivacea, Bassaricyon alleni, and B. gabbii, the canid Nyctereutes procyonoides, the mustelids M. felipei, and M. lutreolina, and the viverrid G. thierryi, to mention a few.

In terms of research topics, this review revealed that the distribution and diet were the most recurrent aspect in studies within the order Carnivora, especially for the families Felidae, Mustelidae and Canidae. However, for some species even basic aspects such as natural history, breeding behavior and habitat association remain unknown (Wilson and Mittermeier, 2009), among which are the herpestids Helogale hirtula and Herpestes semitorquatus, the mustelids M. felipei and M. lutreolina, or the viverrids G. bourloni , G. poensis, and G. thierryi.

Added to this, the lack of knowledge of some aspects has contributed to discrepancies in the systematics of some species of the order Carnivora. In such circumstances are South American cats, which have a long history of changes in their taxonomy, even in recent years (Clavijo and Ramirez, 2009; Wilson and Mitttermeier, 2009). A similar problem also face the procyonids Procyon insularis, Nasua nelsoni (Cuaron et al., 2004), B. beddardi, B. lasius, and B. pauli, or the mustelid Lutra nippon (Wozencraft, 2005; Wilson and Mittermeier, 2009). These differences are also present at the family level in various classifications, such as in the case of Prionodontidae (Wozencraft, 2005; Schipper et al., 2008; Wilson and Mittermeier, 2009). This suggests that studies on genetics, evolution, taxonomy, and morphology are required to help the clarification of the systematics of the order Carnivora (Cuaron et al., 2004; Clavijo and Ramirez, 2009).

Although some authors consider that conservation efforts do not require an understanding of all aspects for the species of interest (Ginsber, 2001), knowledge of ecological requirements is necessary, especially in cases where the risk of extinction is found to be significant, to appropriately channel conservation actions and resources (Ginsberg, 2001; Meiri, 2005; Jenning and Veron, 2009). However, the lack of studies on groups of species may hamper the accurate identification of their conservation status within each region, there may be discrepancies in terms of risk category assessment between the local conservation perspective and the global conservation perspective (Dalerum et al., 2009). As a result of this situation, only a few of the species in this order have been classified in a higher risk category in the international context (Schipper et al., 2008; IUCN, 2010). Therefore it is advisable to collect information on various aspects of threatened species in order to make it possible to reevaluate the conservation status (Schipper et al., 2008).


Some research indicates that the regions of Asia, Africa and South America have high species richness (Mills et al., 2001; Dalerum et al., 2009). However, is in these places where a large number of species are at risk (Servheen et al., 1999; Cardillo et al., 2004; Mills et al., 2005). Despite this diversity and degree of threat, these regions do not excel in the number of published studies. It would be important to increase the number of studies on native species of these regions before the environmental changes drastically affect these populations.

Future research should be directed towards addressing topics other than natural history, such as genetics, habitat selection, activity patterns, or zoosanitary issues. Similarly, it is advisable to expand the scope of future studies to include little studied groups, especially threatened species, small-sized species, and species which are not very abundant or charismatic. Given this situation, the study of aspects such as population trends or demography, as well as community level ecology, is needed in order to understand under what circumstances or through what ecological patterns, communities are developed and maintained, with the aim of maximizing the effectiveness of conservation actions and resources.

Recibido 10 octubre 2012. Aceptado 5 abril 2013. Editor asociado: F Prevosti

Gabriela Perez-Irineo and Antonio Santos-Moreno

Laboratorio de Ecologia Animal, Centro Interdisciplinario de Investigacion para el Desarrollo Integral Regional, Unidad Oaxaca, Instituto Politecnico Nacional, Hornos 1003, 71230 Santa Cruz Xoxocotlan, Oaxaca, Mexico [Correspondence: Gabriela Perez Irineo <>].


The authors would like to thank the Mexican Consejo Nacional de Ciencia y Tecnologia and Instituto Politecnico Nacional for its generous funding of the graduate studies of the first author, and G. Ramos-Fernandez for his valuable comments on this work.


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Table 1
Percentage of articles on different research topics for
the order Carnivora, from 1980 to 2010.

Topic                Subtopic                             The

Abundance            Density, relative abundance          3.3
Home Range           Territory                            4.0
Diet                 Preference, habits                   14.7
Natural History      Metabolism, reproduction,            12.8
Conduct              Activity patterns, behavior          8.5
Conservation         Species at risk, programs, risk      6.6
Distribution         New records, zoogeography,           13.7
Ecology              Diversity, predation,                14.7
Evolution            Phylogeny, paleontology              1.8
Genetics             Alleles, genetic variation           4.9
Zoosanitary Issues   Diseases, zoonosis, parasites        1.5
Methods              Efficiency, comparisons, new         2.1
Morphology           Morphometry, size, weight            5.6
Taxonomy             Classification, systematic           1.3
Habitat Use          Selection, preference                2.9
Other                Ethnozoology, hunting/poaching,      1.6
                       literature reviews

Topic                Africa   Asia   Europe   Total

Abundance             2.1     2.8     5.3      3.5
Home Range            2.3     0.7     2.9      2.6
Diet                  17.4    7.9     16.0    13.7
Natural History       9.0     5.7     13.0    10.7
Conduct               12.3    4.8     7.1      7.9
Conservation          12.1    18.2    8.4     10.6
Distribution          14.4    32.6    14.9    17.4
Ecology               9.5     11.2    8.0     11.1
Evolution             0.8     0.4     0.6      1.5
Genetics              1.8     0.9     3.2      3.1
Zoosanitary Issues    2.8     1.6     5.0      2.6
Methods               3.8     2.9     5.3      3.4
Morphology            3.4     2.9     6.7      5.7
Taxonomy              0.3     2.4     0.0      1.1
Habitat Use           5.9     1.6     1.9      2.9
Other                 2.1     3.2     1.7      2.2

Table 2
Species richness and the number of published works per family
of mammalian carnivores between 1980 and 2010 on the global
level and by continent. References: a = total species richness,
b = number of species with at least one work, c = number of
published articles, d = proportion of studies (PAF = the number
of articles published per family/number of species of the

                       Globally              The Americas

                  a     b     c      d     a    b     c     d

Ailuridae         1     i     10     10
Canidae          35    30    457    13.1   18   17   267   14.8
Euplerida         8     6     13    1.6
Felidae          37    36    530    14.3   15   11   152   10.1
Herpestidae      34    20     58    1.7         1     3
Hyaenida          4     4     52     13
Mephitidae       12     9     27    2.2    10   8    26    2.6
Mustelidae       57    46    501    8.8    21   21   111   5.3
Nandiniidae       1     1     3      3
Prionodontidae    2     2     7     3.5
Procyonidae      12    11     82    6.8    12   11   74    6.2
Ursidae           8     8    134    16.7   4    4    86    21.5
Viverridae       34    22    107    3.1
Order                        136                     39
TOTAL            245   196   2117          80   73   758

                        Europe               Africa

                 a    b     c     d     a    b     c     d

Ailuridae                                    1     1
Canidae          4    7    76     19    14   9    82    5.8
Euplerida             2     3           8    6    10    1.2
Felidae          4    5    49    16.3   10   10   139   13.9
Herpestidae      2    3     6     6     25   13   29    1.7
Hyaenida                                4    4    49    12.2
Mephitidae            1     2
Mustelidae       11   11   292   24.3   7    5    25    3.6
Nandiniidae                             1    1     3     3
Procyonidae                 1
Ursidae          2    3    21    10.5
Viverridae       1    3    11     11    16   11   22    1.4
Order                      15                     30
TOTAL            24   35   476          85   60   390


                  a    b     c     d

Ailuridae         1    1     9     9
Canidae          11    11   37    3.3
Felidae          21    20   155   7.4
Herpestidae       9    7    20    2.2
Hyaenida          1    1     2     2
Mephitidae        2
Mustelidae       33    21   60    1.8
Prionodontidae    2    2     7    3.5
Ursidae           6    5    23    3.8
Viverridae       18    15   70    3.9
Order                       34
TOTAL            104   83   417

Table 3
Species of the order Carnivora with the greatest quantity of
published works from 1980 to 2010. References: A = refers to
whether the species has a review in Mammalian Species,
B = number of species by family to have a review within
Mammalian Species, C = the value of B expresses as a

Family           Species                    Number of   Mammalian
                                            Articles     Species


Ailuridae        Ailurus fulgens               10           X

Canidae          Canis latrans                 84           X
                 Vulpes vulpes                 72           X
                 Canis lupus                   60           X
                 Lycaon pictus                 35
                 Vulpes macrotis               23           X
                 Lycalopex culpaeus            19           X
                 Vulpes lagopus                17           X
                 Canis mesomelas               14           X
                 Chrysocyon brachiurus         14

Eupleridae       Crytoprocta ferox              5
                 Mungotictis decemlineata       3

Felidae          Panthera leo                  62           X
                 Panthera pardus               57
                 Puma concolor                 55           X
                 Acinonyx jubatus              45           X
                 Panthera onca                 41           X
                 Lynx rufus                    33           X
                 Panthera tigris               31           X
                 Lynx lynx                     25           X
                 Leopardus pardalis            16           X

Herpestidae      Ichneumia albicaudata          4

Hyaenidae        Crocuta crocuta               17
                 Hyaena brunnea                 3           X

Mephitidae       Mephitis mephitis             11           X
                 Spilogale putorius             7           X

Mustelidae       Meles meles                   123
                 Lutra lutra                   70
                 Neovison vison                38           X
                 Mustela lutreola              32           X
                 Martes pennanti               13           X
                 Martes americana              12           X
                 Mustela putorius              12
                 Martes martes                 11
                 Gulo gulo                     10           X

Nandiniidae      Nandinia binotata              3

Prionodontidae   Prionodon pardicolor           6
                 Prionodon linsang              1

Procyonidae      Procyon lotor                 32           X
                 Nasua narica                   9           X

Ursidae          Ursus americanus              51           X
                 Ursus arctos                  24           X
                 Ursus maritimus               22           X
                 Ailuropoda melanoleuca        15           X

Viverridae       Genetta genetta               14           X
                 Chrotogale owstoni            13
                 Viverricula indica             6

Family           Species                        Mammalian Species

                                             Number of     Presence
                                              Species     Percentage
                                            Present (B)      (C)

Ailuridae        Ailurus fulgens                 1           100

Canidae          Canis latrans                  28            80
                 Vulpes vulpes
                 Canis lupus
                 Lycaon pictus
                 Vulpes macrotis
                 Lycalopex culpaeus
                 Vulpes lagopus
                 Canis mesomelas
                 Chrysocyon brachiurus

Eupleridae       Crytoprocta ferox               1           12.5
                 Mungotictis decemlineata

Felidae          Panthera leo                   13           35.1
                 Panthera pardus
                 Puma concolor
                 Acinonyx jubatus
                 Panthera onca
                 Lynx rufus
                 Panthera tigris
                 Lynx lynx
                 Leopardus pardalis

Herpestidae      Ichneumia albicaudata          11           32.3

Hyaenidae        Crocuta crocuta                 3            75
                 Hyaena brunnea

Mephitidae       Mephitis mephitis               6            50
                 Spilogale putorius

Mustelidae       Meles meles                    29           50.8
                 Lutra lutra
                 Neovison vison
                 Mustela lutreola
                 Martes pennanti
                 Martes americana
                 Mustela putorius
                 Martes martes
                 Gulo gulo

Nandiniidae      Nandinia binotata               0            0

Prionodontidae   Prionodon pardicolor            0            0
                 Prionodon linsang

Procyonidae      Procyon lotor                   5           41.7
                 Nasua narica

Ursidae          Ursus americanus                5           62.5
                 Ursus arctos
                 Ursus maritimus
                 Ailuropoda melanoleuca

Viverridae       Genetta genetta                 3           8.8
                 Chrotogale owstoni
                 Viverricula indica
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Title Annotation:texto en ingles
Author:Perez-Irineo, Gabriela; Santos-Moreno, Antonio
Publication:Mastozoologia Neotropical
Date:Jun 1, 2013
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