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Maor et al. Evolution of activity patterns in mammals 1 Temporal niche expansion in mammals from a nocturnal ancestor after 1 dinosaur extinction 2 3 Roi Maor 1,2, *, Tamar Dayan 1,3 , Henry Ferguson-Gow 2 and Kate E. Jones 2,4, * 4 5 1 School of Zoology, George S. Wise Faculty of Life Science, Tel Aviv University, Tel Aviv, 6 6997801, Israel. 7 2 Centre for Biodiversity and Environment Research, Department of Genetics, Evolution and 8 Environment, University College London, Gower Street, London, WC1E 6BT, United 9 Kingdom. 10 3 The Steinhardt Museum of Natural History, Tel Aviv University, Tel Aviv 6997801, Israel. 11 4 Institute of Zoology, Zoological Society of London, Regent’s Park, London, NW1 4RY, 12 United Kingdom. 13 * Correspondence and requests for materials should be addressed to RM 14 ([email protected]) or KEJ ([email protected]). 15

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Page 1: 1 Temporal niche expansion in mammals from a nocturnal ...€¦ · Maor et al. Evolution of activity patterns in mammals 5 91 before the extinction of dinosaurs, calling for a wider

Maor et al. Evolution of activity patterns in mammals

1

Temporal niche expansion in mammals from a nocturnal ancestor after 1

dinosaur extinction 2

3

Roi Maor1,2,*, Tamar Dayan1,3, Henry Ferguson-Gow2 and Kate E. Jones2,4,* 4

5

1 School of Zoology, George S. Wise Faculty of Life Science, Tel Aviv University, Tel Aviv, 6

6997801, Israel. 7

2 Centre for Biodiversity and Environment Research, Department of Genetics, Evolution and 8

Environment, University College London, Gower Street, London, WC1E 6BT, United 9

Kingdom. 10

3 The Steinhardt Museum of Natural History, Tel Aviv University, Tel Aviv 6997801, Israel. 11

4 Institute of Zoology, Zoological Society of London, Regent’s Park, London, NW1 4RY, 12

United Kingdom. 13

* Correspondence and requests for materials should be addressed to RM 14

([email protected]) or KEJ ([email protected]). 15

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Most modern mammals, including strictly diurnal species, exhibit sensory adaptations 16

to nocturnal activity, thought to be the result of a prolonged nocturnal phase or 17

'bottleneck' during early mammalian evolution. Nocturnality may have allowed 18

mammals to avoid antagonistic interactions with diurnal dinosaurs during the Mesozoic. 19

However, understanding the evolution of mammalian activity patterns is hindered by 20

scant and ambiguous fossil evidence. While ancestral reconstructions of behavioural 21

traits from extant species have the potential to elucidate these patterns, existing studies 22

have been limited in taxonomic scope. Here, we use an extensive behavioural dataset for 23

2415 species from all extant orders to reconstruct ancestral activity patterns across 24

Mammalia. We find strong support for the nocturnal origin of mammals and the 25

Cenozoic appearance of diurnality, although cathemerality (mixed diel periodicity) may 26

have appeared in the late Cretaceous. Simian primates are among the earliest mammals 27

to exhibit strict diurnal activity, some 52-33 Million years ago (Mya). Our study is 28

consistent with the hypothesis that temporal partitioning between early mammals and 29

dinosaurs during the Mesozoic led to a mammalian nocturnal bottleneck, but also 30

demonstrates the need for improved phylogenetic estimates for Mammalia. 31

Species exhibit characteristic patterns of activity distribution over the 24-hour (diel) 32

cycle, and as environmental conditions may change radically, yet predictably between day 33

and night, activity patterns allow individuals to anticipate fluctuations, and time activity 34

optimally1,2. Physiological and behavioural adaptations to different activity patterns are 35

important contributors to individual fitness3, and therefore to species evolutionary success4,5. 36

Moreover, long-term shifts in activity patterns may reveal shifts in selective regimes, caused 37

by changes in biotic and abiotic conditions5-7. Although mammals exhibit striking 38

morphological, behavioural and ecological niche diversity8, the distribution of mammalian 39

activity patterns is strongly biased towards nocturnality9. Additionally, most mammalian 40

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species, including strictly diurnal ones, exhibit visual adaptations to nocturnal activity that are 41

similar to those of nocturnal birds and reptiles10. For example, mammals (except Haplorrhine 42

primates) lack a fovea – an area in the retina that enables very high visual acuity found in fish, 43

reptiles, and birds that are diurnal visual predators11. Most mammalian eyes have high ratios 44

of corneal diameter to axial ocular length which favour sensitivity to low-light over visual 45

acuity, and are comparable to those found in nocturnal reptiles and birds10. Compared to all 46

other vertebrates, mammals also exhibit reduced diversity of active photoreceptors which 47

allow colour perception in bright environments12,13. Many day-active mammals (e.g. 48

ungulates, carnivores) have rod-dominated retinae, i.e. have eyes better suited for low-light 49

conditions (night vision), although ratios of retinal rod and cone ratios show high 50

interspecific variability14. There is also evidence that enhanced olfactory sensitivity15, broader 51

frequency range hearing16, and sophisticated whisker-mediated tactile perception17 may have 52

evolved in mammals to compensate for insufficient visual information10,13. 53

In his seminal work, Walls11 noted the differences between mammals and other 54

(mostly diurnal) amniotes in eye shape, retinal composition and visual pathways. He 55

proposed that the predominance of nocturnal adaptations in mammals may be the result of a 56

prolonged nocturnal phase in the early stages of mammalian evolution, after which emerged 57

the more diverse patterns observed today11,13. This ‘nocturnal bottleneck’ hypothesis suggests 58

that mammals were restricted to nocturnal activity by antagonistic interactions with the 59

ecologically dominant diurnal dinosaurs during the Mesozoic11,13,18. The Cretaceous-60

Paleogene (K-Pg) mass extinction event circa 66Mya, led to the extinction of all non-avian 61

dinosaurs along with the marine- and flying reptiles, and the majority of other vertebrates, 62

and invertebrate and plant taxa19,20. This event marks the end of the Mesozoic ‘reign of 63

dinosaurs’ and the transition to the mammal-dominated Cenozoic fauna. If an antagonistic 64

interaction with dinosaurs was an important factor in restricting early mammals to nocturnal 65

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activity, then the vast majority of, if not all Mesozoic mammals are expected to have been 66

nocturnal, and diurnal mammals would have only appeared after the K-Pg mass extinction 67

event. 68

Support for the nocturnal bottleneck hypothesis is drawn from anatomical and 69

morphological studies10,11, and increasingly from molecular studies12,13, but remains indirect. 70

For example, some Synapsids, the non-mammalian lineage ancestral to mammals, were 71

adapted to nocturnal activity >300Mya, suggesting that nocturnality, a relatively rare state in 72

amniotes, may have already characterised the Palaeozoic precursors of mammals21. However, 73

inferring activity patterns from fossil morphology may be unreliable22,23, particularly as all 74

modern mammals (except Haplorrhine primates) have nocturnal-type ocular and cranial 75

morphologies (e.g. high corneal diameter to axial length ratios, a large binocular visual field 76

overlap) regardless of their activity pattern10,23. Evidence from histological and molecular 77

studies of the evolutionary development of mammalian eyes indicate that nocturnal 78

adaptations preceded diurnal ones12,24, but this does not help elucidate questions around the 79

timing of these adaptations. 80

Ancestral reconstructions of behavioural traits using a phylogenetic comparative 81

approach may help to understand both the pattern and timing of the evolution of activity 82

patterns in mammals since activity patterns have been shown to be genetically determined25 83

yet responsive to selective pressures2. However, phylogenetic studies of mammalian activity 84

patterns so far have mostly focused on two mammalian orders – primates26-28 and rodents29. 85

Primate activity patterns have been studied extensively, and some evidence suggests that 86

primate diurnality originated in the most recent common ancestor (MRCA) of suborder 87

Haplorrhini (all monkeys, apes and tarsiers)5 in the Mesozoic30,31. It is conceivable, although 88

thus far not tested, that diurnal diversifications in other orders of Mesozoic origins, e.g. 89

Scandentia (treeshrews), Macroscelidea (elephant shrews) and Rodentia, could have occurred 90

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before the extinction of dinosaurs, calling for a wider examination of how activity patterns 91

evolved across mammals. 92

Here, we use an extensive dataset of activity patterns for 2415 mammal species, 93

representing 135 of the 148 extant families and all extant orders (Supplementary Table 1) to 94

investigate ancestral activity patterns in mammals, and to understand the timings of the 95

appearance of mammal diurnality. We assign species to one of five activity patterns: (i) 96

nocturnal – active only or mostly in the dark; (ii) diurnal – active only or mostly during 97

daylight hours; (iii) cathemeral – active both during the day and during the night; (iv) 98

crepuscular – active only at twilight, around sunrise and/or sunset; and (v) ultradian – active 99

in cycles of a few hours (see Methods). We map the three main activity patterns (nocturnal, 100

cathemeral, and diurnal) onto two phylogenetic frameworks representing two of the main 101

hypotheses of mammalian evolutionary history for our analyses, termed here short-fuse (SF) 102

following31 updated by32, and long-fuse (LF) phylogenies (adapted from30) (Fig. 1). We then 103

use reversible jump Markov Chain Monte Carlo (rjMCMC) methods33 to estimate transition 104

rates between different activity states, and to infer the posterior probability (PP) of character 105

states at each node in the phylogenies. This allows us to examine the evolution of activity 106

patterns of mammals, and to test the main predictions of the nocturnal bottleneck hypothesis; 107

(i) the most recent common ancestor to all extant mammals was nocturnal, and (ii) mammal 108

diurnality first emerged in the Cenozoic. 109

Results 110

We find that the modal values of PPNoct (posterior probability of nocturnality) at the ancestral 111

node of extant mammals were 0.74 (Credible Interval, CrI 0.71-0.76) and 0.59 (CrI 0.54-112

0.64) for SF and LF phylogenies, respectively, offering support for a noctural ancestor (Fig. 113

2). In contrast, a cathemeral or a diurnal ancestral state is much less well supported: modal 114

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value of PPCath (posterior probability of cathemerality) = 0.24 (CrI 0.23-0.26) and 0.31 (CrI 115

0.29-0.33) for SF and LF, respectively, or PPDiur (posterior probability of diurnality) = 0.02 116

(CrI 0.01-0.03) SF and 0.1 (CrI 0.07-0.14) LF (Fig. 2). The narrow and non-overlapping 117

distributions of PP values across the activity pattern reconstructions indicate that our results 118

are consistent and robust across samples of the rjMCMC chains, although the distributions 119

are wider using the LF phylogeny (Fig. 2). 120

The first strong evidence (where the reconstructed activity pattern was supported by 121

modal PP values >0.67) in mammals of an expansion of temporal niche into cathemerality, is 122

in the early Paleogene (Cenozoic) for the SF phylogeny (no later than 65.8Mya), or in the late 123

Cretaceous (Mesozoic) for the LF phylogeny (no later than 74.7Mya) (Figs. 3 and 4). 124

Although the LF phylogeny supports a Mesozoic shift to cathemerality, the modal PP values 125

of the remaining 41 Mesozoic nodes were either nocturnal (23 nodes), or unclear – where all 126

three activity patterns were supported by modal PP values <0.67 (18 nodes). Using the SF 127

phylogeny, we reconstruct the first transition to cathemerality in the MRCA of order 128

Cetartiodactyla (cetaceans and even-toed ungulates). This taxa was likely to be cathemeral 129

(PPCath = 0.79 CrI 0.72-0.87), and almost certainly exhibited considerable daytime activity 130

(PPNoct = 0.02 CrI 0.01-0.04) (Fig. 3). Using the LF phylogeny, the first cathemeral transition 131

was in the MRCA of families Soricidae (shrews) and Talpidae (moles) (PPCath = 0.81 CrI 132

0.61-0.91; PPDiur = 0.07 CrI 0.03-0.15) (Fig. 4). 133

Evidence of the evolution of diurnality (modal PP values >0.67) first appears in the 134

early Paleogene (no later than 52.4Mya or 63.8Mya for SF and LF phylogeny, respectively) 135

(Figs. 3 and 4). Using the SF phylogeny, we reconstruct transition to diurnality in the MRCA 136

of the Simiiformes (all monkeys and apes) (PPDiur = 0.76, CrI 0.75-0.78; PPCath = 0.23, CrI 137

0.22-0.25) (Fig.3). Using the LF phylogeny, the first taxon to exhibit diurnal activity was the 138

MRCA of the family Macroscelididae (elephant shrews) (PPDiur = 0.77, CrI 0.76-0.80; PPCath 139

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= 0.22, CrI 0.19-0.23; 63.8Mya), followed by the MRCA of families Ctenodactylidae (comb 140

rats, Rodentia) (PPDiur = 0.76; CrI 0.73-0.78; 61.6Mya), Camelidae (Cetartiodactyla) (PPDiur = 141

0.74, CrI 0.72-0.77; 59.6Mya), and Tupaiidae (treeshrews, Scandentia) (PPDiur = 0.99, CrI 142

0.99-0.99; 51.1Mya) in rapid succession (Fig. 4). 143

For both SF and LF phylogenies, we find that transition rates from a cathemeral pattern to 144

either noctural or diurnal are about three times higher than the transition rates from either 145

nocturnal or diurnal to cathemeral (Table 1). Furthermore, the transition rates in the SF 146

reconstruction are three orders of magnitude lower than the respective rates in the LF 147

reconstruction. 148

Discussion 149

We have shown that extant mammals likely originated from a nocturnal ancestor, and that 150

these ancestors remained nocturnal throughout the Mesozoic until either 9 Million years 151

(Myr) before the K-Pg event (LF reconstruction), or just after it (SF reconstruction). On 152

balance, our evidence suggests that mammals likely remained nocturnal throughout the 153

Mesozoic as nocturnal activity is strongly supported at most Mesozoic nodes in both SF and 154

LF reconstructions. We find strong evidence that the shift to strict diurnality occurred after 155

the K-Pg event (both SF and LF reconstructions), although cathemerality may have appeared 156

in the late Cretaceous (74.7Mya LF reconstruction). Combined with other sources of 157

evidence, such as the morphology of mammalian eyes10,23, composition and reduced diversity 158

of retinal photoreceptors12,13,24,34, and the emphasis on alernative sensory systems11,15-17, our 159

analysis helps to further establish the nocturnal ancestry of mammals and that diurnality only 160

orignated in mammals after the dissapearance of the dinosaurs, as predicted by the nocturnal 161

bottleneck hypothesis. 162

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Even if we accept the appearance of cathemeral mammals as an expansion of the 163

temporal niche before the K-Pg event, it does not necessarily provide strong evidence against 164

the nocturnal bottleneck hypothesis. Declines in dinosaur diversity long before the K-Pg 165

event have been suggested, either globally, starting at least 40Myr before the K-Pg event35, or 166

locally – herbivorous dinosaurs in present-day North America were declining for up to 167

15Myr prior to the event20. In contrast, fossils show that mammals had evolved considerable 168

eco-morphological diversity as early as the mid-Jurassic period (174-164 Mya), and 169

diversified along all axes of the ecological niche36,37, except the temporal axis. Moreover, 170

extensive mammal radiations occurred following the Cretaceous Terrestrial Revolution (KTR, 171

120-80Mya), whereby angiosperms rose to dominate the global flora, and revolutionised eco-172

space30,38,39. Under such conditions, a partial invasion of mammals into the temporal niche of 173

declining dinosaurs does not violate the assumption of temporal partitioning. Indeed, 174

evidence of a shift in retinal opsin sensitivity (linked to more diurnal activity patterns) in 175

some mammalian clades (Cetartiodactyls, primates, carnivores, and some Afrotheria orders) 176

more than 70Mya24,34, offers further support for a transition occurring during this period. 177

The MRCA of infraorder Simiiformes (monkeys and apes) was among the first taxa to 178

have evolved diurnality (52.4Mya, SF reconstruction), and this is consistent with their 179

evolution of diurnally-adapted vision, specifically trichromacy and a low ratio of corneal 180

diameter to axial length10,12,23 – unique in mammals. Other diurnal clades such as squirrels 181

(Sciuridae) and elephant-shrews (Macroscelididae) evolved at about the same time as the 182

Simiiformes30,31 and presumably had similar opportunity to evolve comparable visual 183

adaptations to diurnality. However, these groups rely on high ratios of retinal cones to rods 184

for daylight vision14, suggesting that diurnality in Simiiformes may have evolved 185

considerably earlier than the minimum date of 52.4Mya. Simiiformes lie on an evolutionary 186

branch that originates 83.2Mya (SF), when they diverged from tarsiers – their closest living 187

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relatives in the suborder Haplorrhini. Tarsiers are strictly nocturnal, but share with the 188

Simiiformes several adaptations for high visual acuity, typical to diurnal vision28,40. The 189

morphological and physiological adaptations to nocturnality in tarsiers are unlike those of any 190

other nocturnal primate, suggesting that tarsiers originated from a diurnal ancestor, the 191

MRCA of Haplorrhini, and secondarily adapted to nocturnal life5,6. The Haplorrhine MRCA 192

was a Mesozoic species that lived until 83.2Mya (SF) or 78.1Mya (LF). This would imply 193

that Mesozoic mammals were able to break out of the nocturnal bottleneck and endure direct 194

interaction with dinosaurs following the KTR. Nevertheless, both reconstructions here, as 195

well as other reconstructions of primate activity patterns based on different sets of data, 196

including data on visual physiology, find weak or no evidence to the diurnality of the 197

Haplorrhine MRCA26-28. 198

There are other uncertainties around the dates for three of the four taxa identified as 199

shifting to diurnality within 7Myr after the K-Pg in the LF reconstruction (Macroscelididae, 200

Ctenodactylidae, Camelidae). This is due to how we re-scaled the terminal-branches in30 to 201

produce the species-level LF phylogeny. However, according to the dates given in30 and 202

additional studies supporting the LF hypothesis41-44, these families originated in the Cenozoic, 203

so our prediction of Cenozoic origins to mammal diurnality remains intact. The MRCA of 204

Tupaiidae (Scandentia) and their closest living relative – the nocturnal Ptilocercidae (Pen-205

tailed tree shrews, a monotypic family) – has been placed in the Cenozoic, 60.1 Mya30 The 206

LF reconstruction shows that this species was probably diurnal or cathemeral, but neither 207

pattern was supported by PP values >0.67. 208

On both SF and LF reconstructions, the rates of transition from cathemeral activity to 209

either nocturnal or diurnal imply that the diurnal and nocturnal niches may be more 210

favourable for mammals. However, our results unequivocally support the persistence of 211

cathemerality in mammals since the K-Pg. In primates, cathemerality has been argued 212

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adaptive under fluctuating environmental conditions26,45 and cathemeral species show higher 213

speciation rates (although lower overall diversification rates) compared to nocturnal and 214

diurnal species27. If these patterns are also true for the rest of Mammalia, they could explain 215

the persistence of mammal cathemerality against the net outflow of species and slow 216

diversification rates. In Lepidoptera (moths and butterflies), the persistence of a mixed 217

(cathemeral) diel activity pattern has been argued to be the result of conflicting predation 218

pressures, from bats during the night and birds during the day46. Hence, cathemeral activity 219

may be preferred when strong selective forces are acting in opposite directions. The 220

appearance of mammal cathemerality may have been due to high nocturnal predation risk on 221

one side (perhaps from other mammals making the nocturnal niche less advantageous), and 222

the difficulties of adapting to a diurnal niche on the other. 223

The higher transition rates for the LF tree are likely a result of the method we used to 224

construct the species-level LF phylogeny, i.e. re-scaling the branch lengths of species-level 225

clades from the SF phylogeny31 to maintain the length of the corresponding terminal branch 226

provided by30. SF branch lengths were usually scaled down in this process, because the SF 227

generally estimates older divergence dates than the LF, reflecting the difference between the 228

two phylogenetic models. A consequence of our grafting procedure is that a band of 229

artificially short branches is formed near these graft points, which implies rapid change. 230

Higher rates allow for more change along tree branches, and reduce the precision of the 231

results, which probably contributed to our LF reconstruction yielding fewer decisive 232

predictions and lower statistical support compared with the SF reconstruction (Figs. 2, 3 and 233

4). Whilst a direct comparison of transition rates between the two phylogenetic hypotheses is 234

therefore precluded, the broad pattern of transitions (i.e. low transition rates into 235

cathemerality and high transition rates out of it in either direction) is supported in both 236

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analyses, as is the general pattern of temporal niche evolution that emerges from the node 237

reconstructions. 238

Although we have demonstrated the importance of the phylogenetic comparative 239

approach to the investigation of the evolution of behavioural traits in mammals, ancestral 240

reconstruction methods rely heavily on the accuracy of phylogenetic estimates. The LF 241

hypothesis of mammalian evolutionary history is well supported30,41,44, but phylogenetic 242

estimates are only available at family-level, and further modification was required to add the 243

species-level information for our analysis. Despite the attention attracted recently by studies 244

of mammalian phylogenies30,41,44,47, only the SF hypothesis is represented by a species-level 245

phylogeny, making the incorporation of the LF hypothesis and the explosive model 246

problematic for phylogenetic comparative analyses that are based on detailed species-level 247

data. 248

In conclusion, we argue that the activity patterns of Mesozoic mammals are consistent 249

with the prediction of temporal partitioning, and that the gradual acquisition of daytime 250

activity in mammals, first cathemerality then diurnality, coincided with the decrease in 251

pressure from dinosaurs, whether due to their decline or extinction. Given the current 252

evidence, temporal partitioning within Mesozoic amniotes mostly followed the phylogenetic 253

(mammal-archosaur) division, but while some dinosaurs invaded the nocturnal niche22, we 254

find little support for Mesozoic mammals invading the diurnal niche. The constraints on 255

mammals becoming diurnal during the Mesozoic would have been strong enough to 256

counteract the ecological pressure to diversify, following at least 100Myr of mammalian 257

sensory and eco-morphological radiations that sub-divided their nocturnal niches. Mammals 258

diversified rapidly once they expanded outside the nocturnal niche, but whether invading the 259

diurnal niche facilitated mammals’ Cenozoic success remains to be answered. 260

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Methods 261

Data. We collated activity records for 2415 mammal species, representing all 29 262

extant orders and 135 of 148 extant families from the PanTHERIA database8, and from 263

published sources such as research articles, field guides, and encyclopaedias (Supplementary 264

Table 1). To achieve maximal representation of taxonomic diversity, we specifically targeted 265

under-represented orders, and repeated the process for under-represented families. 266

Nonetheless, any records we found in this process were incorporated into our data set, 267

whether of a target taxon or not, unless a similar record (same species and activity pattern) 268

was previously obtained. Although activity pattern data was only available for just under half 269

of all known species48 (44.6%), 91.2% of families were represented in the database. The most 270

under-represented taxa were the largest orders (Rodentia 59% missing species, Chiroptera 271

74% and Soricomorpha 82%). Bats are almost entirely nocturnal, and Soricomorpha is 272

predominantly cathemeral (except the nocturnal Erinaceomorpha). In rodents too, activity 273

patterns closely follow phylogeny29. Therefore, the inclusion of the missing species would 274

likely have only a minor effect, if any, on the character transition rate matrix and the overall 275

reconstruction results. 276

We assigned each species into one of five activity patterns: (i) nocturnal – active only 277

or mostly in the dark; (ii) diurnal – active only or mostly during daylight hours; (iii) 278

cathemeral – active both during the day and during the night; (iv) crepuscular – active only at 279

twilight, around sunrise and/or sunset; and (v) ultradian – active in cycles of a few hours. We 280

considered species nocturnal or diurnal based on qualitative descriptions in sources, as 281

precise quantitative measurements are rare, where species described as ‘nocturnal’ or ‘active 282

at night’ were assigned to nocturnal and species described as ‘diurnal’ or ‘active during 283

daylight’ were assigned to diurnal. We also categorised species to these two categories if 284

those descriptions were preceded by ‘only’, ‘exclusively’, ‘strictly’, ‘mostly’, 285

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‘predominantly’, almost exclusively’, or ‘mainly. Species which were described as ‘nocturnal 286

and diurnal’, ‘active day and night’, ‘active at all hours’, ‘arrhythmic’, ‘nocturnal in summer 287

and diurnal in winter’ were assigned as having a cathemeral activity pattern. Crepuscular 288

activity was assigned to species described as ‘mostly or mainly or predominantly crepuscular’, 289

‘active at dusk’, ‘active at dusk and dawn’, ‘around sunrise and sunset’, ‘activity peaks in late 290

afternoon or early evening’. Ultradian patterns were assigned when species were described 291

as ‘ultradian’ or the source described several rhythmic cycles of activity and rest over a 24-292

hour period. We follow the taxonomy and species binomials in Mammal Species of the 293

World, 3rd Edition48, with one exception: we use Cetartiodactyla, instead of separate orders 294

Artiodactyla and Cetacea, following49,50. We resolved conflicts where sources disagreed on 295

species activity pattern as follows: (i) records of crepuscular activity (dusk or dawn), when in 296

conjunction with nocturnal or diurnal activity, were changed to nocturnal or diurnal, 297

respectively; (ii) records from complied sources were preferred over localised studies (which 298

are prone to idiosyncrasies); and (iii) records from more recent sources were preferred. This 299

left 29 species unresolved and these species were excluded from subsequent analyses, giving 300

a total number of species = 2386 (1426 nocturnal, 615 diurnal, 322 cathemeral, 22 301

crepuscular, and one ultradian species). 302

Phylogenetic framework. We used two phylogenetic frameworks representing two 303

of the main hypotheses of mammalian evolutionary history for our analyses: the short-fuse 304

(SF) hypothesis is represented by the species-level "best dates" supertree31 updated from32, 305

and the long-fuse (LF) hypothesis is represented by the amino-acid supermatrix phylogeny30 306

(Fig.1). The SF hypothesis asserts that the most recent common ancestor (MRCA) of all 307

extant mammals diverged into its daughter lineages (Prototheria and Theria) in the mid-308

Jurassic, 166.2Mya, whereas according to the LF hypothesis this divergence took place in the 309

late-Triassic, 217.8Mya. Both hypotheses agree that multiple extant lineages diverged in the 310

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Cretaceous and survived the K-Pg event (Fig. 1), but the SF hypothesis posits that intra-311

ordinal divergence of placental mammals had already begun prior to the K-Pg event, while 312

the LF hypothesis places intra-ordinal divergence in the Cenozoic. A third evolutionary 313

hypothesis, the explosive model, is supported by fossil evidence and morphological data47, 314

but has been criticised for implying impossibly-high rates of evolution in the early-Cenozoic 315

radiation of placental mammals, and for other problems41,51, so we do not consider it here. 316

Here, we represent the LF hypothesis using the family-level supermatrix phylogeny30 317

(downloaded from TreeBASE: http://purl.org/phylo/treebase/phylows/study/TB2:S11872 on 318

01MAR2015). For our analyses we rendered it ultrametric, i.e. all the tips (species) of the tree 319

are equidistant from the root, so that branch lengths are proportional to time. The LF 320

hypothesis has recently gained support from several studies41-44, but it lacks species-level 321

resolution, which is essential for our analysis. We therefore used each terminal branch of the 322

supermatrix phylogeny (representing a taxonomic family) as a root branch onto which we 323

appended the internal branching pattern of the family, as given in31 updated from32. In order 324

to retain the original LF timeline, we scaled the appended branching pattern to 85% of its 325

original supermatrix phylogeny branch length, and the root branch completed the remaining 326

15%. Other proportions, for example 70:30 or 50:50 branch scaling would have compressed 327

intra-family branching patterns, resulting in branch lengths that were very different from their 328

original values. For this process we used functions from packages ape52 and phangorn53 in R 329

version 3.2.354. Species that we had data for but that were absent from the phylogenetic 330

frameworks were omitted from the analyses: 33 species from the SF phylogeny, and an 331

additional 38 species and 3 families missing from the LF phylogeny. as families Aotidae, 332

Pitheciidae and Lepilemuridae (Primates) were not originally included in the supermatrix 333

phylogeny30. It is unlikely that the omission of these three families would have had an impact 334

on our analysis, as of these three families, Pitheciidae and Lepilrmuridae are entirely diurnal 335

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and nocturnal, respectively, and conform to the activity pattern of the respectively clades 336

within which they are nested. Aotidae, on the other hand, is nocturnal. While this can 337

potentially alter ancestral reconstruction results, Aotidae is nested within the otherwise 338

exclusively diurnal Platyrrhini (new world monkeys)27, so its effect on the LF reconstruction 339

is would be minimal beyond the node immediately ancestral to Aotidae. 340

Analyses. We used BayesTraits v333 to reconstruct the evolution of mammalian 341

activity patterns. BayesTraits implements Markov Chain Monte Carlo (MCMC) methods to 342

sample from the posterior distributions of transition rates for a transition matrix describing 343

the evolution of a discrete character. The obtained posterior distribution allows the user to 344

infer the posterior probability of each character state at the root and at each internal node of 345

the phylogeny. By employing reversible jump MCMC (rjMCMC), BayesTraits is also able to 346

sample from the posterior distribution of model configurations and optimise the number of 347

parameters in the model. This removes the need for comparing models with different number 348

of parameters by sampling from model space and parameter space concurrently55. 349

We only consider the three main activity patterns across mammals in our analysis 350

(nocturnal diurnal and cathemeral) in order to reduce the complexity of the model and 351

increase its biological interpretability (four transition rates instead of 16). Additionally, we 352

remove ultradian activity patterns as these are mostly found with polar and subterranean 353

species, where the 24-hour cycle is of reduced importance. This made the total number 354

species used as 2330 species, 135 families (nocturnal species = 1399, diurnal = 610, and 355

cathemeral = 321), and 2292 species, 132 families (nocturnal species = 1384, diurnal = 588, 356

and cathemeral = 320) for the SF and LF analysis, respectively. We use an ordered model of 357

trait evolution: Nocturnal↔Cathemeral↔Diurnal, whereby direct Nocturnal↔Diurnal 358

transitions are not allowed (set to zero). A transition from diurnal to nocturnal (or vice versa) 359

would therefore involve at least two ‘steps’, passing through cathemeral, although both steps 360

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may occur along the same branch. This ordered model reflects the continuous and mutually-361

exclusive nature of morphological and histological adaptations to diurnality and nocturnality 362

(e.g. retinal rod to cone ratio, corneal diameter to axial length ratio, front-facing versus 363

lateral-facing eye sockets) , while cathemerality involves an intermediate state of the relevant 364

phenotypes23,56. Our underlying hypothesis is that during shifts from diurnality to nocturnality 365

(and vice versa) species go through a phase of cathemeral capability, where they are equally 366

well adapted to both. All other transition rates were free to take any value. We used rjMCMC 367

to estimate the optimal model configuration55. As activity pattern in our analyses was not a 368

binary trait, we used the ‘multistate’ mode of BayesTraits to sample from the posterior 369

distribution of transition rates between activity pattern categories. For each phylogeny, we 370

opted for the reversible-jump MCMC procedure, and set a wide uniform prior, bounded 371

between 0 and 100 for all transition rates, to ensure that our prior did not have a strong effect 372

on the nature of the posterior. Each rjMCMC chain was run until convergence was reached 373

(at least one million iterations), after which point the chains were sampled every 4000 374

iterations until a posterior of 1000 samples was obtained. We chose this wide sampling 375

interval in order to minimise autocorrelation in our posterior samples. We ran twelve 376

replicates of each chain (corresponding a phylogeny) in order to ensure consistency, and that 377

each independent run converged on the same posterior distribution. The marginal likelihoods 378

of each chain were calculated using the stepping stone sampler57 as implemented in 379

BayesTraits (500 stones, 1000 iterations per stone) and compared between independent 380

replicates to ensure consistency. 381

In order to estimate the character state at each internal node, we used the modal value 382

of the PP of each character state, calculated as the peak value of the kernel density of each 383

posterior distribution. For each posterior probability distribution, we report the 95% Credible 384

Interval (CrI), the highest density interval covering 95% of the posterior distribution. We 385

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used the R package phytools 58 to plot the PP values of each node on the mammal 386

phylogenies (Figs. 3 and 4). To measure the accumulation of mammalian temporal niches 387

over time, we calculated the running total of nodes (lineages) where an activity pattern was 388

supported with PP >0.67, and plotted this along the mammal evolution timeline (Figs. 3 and 389

4). A confidence threshold of 0.67 means that the PP values of the best-supported state is at 390

least 0.34 higher (or twice as likely) than the second most probable state. The PP distributions 391

of either state would have to be extremely flat to make the difference between two peak 392

values smaller than two standard deviations. The threshold of 0.67 thus ensures small to no 393

overlap between two distributions. 394

Estimates of character transition rates and reconstructions of ancestral states can be 395

inaccurate if certain character states lead to very different diversification rates59, and methods 396

such as BayesTraits (BT) do not account for the effects of character states on diversification 397

rates. We reanalysed our data to investigate the robustness of our analysis with an additional 398

method, Multistate Speciation and Extinction (MuSSE60), to control for differences in 399

diversification rates. However, this method requires fully bifurcating phylogenetic trees, or, if 400

polytomies are present, that all branches in the phylogenies descending from them are 401

collapsed60. To enable a MuSSE reconstruction, we used Maximum Clade Credibility (MCC) 402

implemented in the R package phangorn53 to summarise a single, fully-bifurcating tree from 403

a distribution of 100 fully bifurcating trees61 randomly derived from the SF phylogeny used 404

in the BT analysis. We could only perform this analysis on the SF phylogeny as the mosaic 405

nature of the LF phylogeny meant that the resulting tree from random resolution was very 406

similar to the SF tree. We acknowledge that random resolution of polytomies may result in 407

unlikely topologies and incorrect branch lengths, but is a pragmatic solution to the 408

incompleteness of mammalian phylogenetic information available. As the results of the 409

MuSSE reconstruction are very similar to those obtained by the BT analysis, and do not 410

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change our overall conclusions (Supplementary Figure 1, Supplementary Table 2), our results 411

are likely robust to the differential diversification rates in activity patterns. 412

Code Availability. Computer code essential for replicating the results in this study is 413

available on Figshare (doi: 10.6084/m9.figshare.4797367). 414

Data Availability. The authors declare that all data supporting the findings of this study are 415

available within the paper and its supplementary information files. All data is available on 416

Figshare a (doi: 10.6084/m9.figshare.4775416; doi:10.6084/m9.figshare.4774648). Reprints 417

and permissions information are available at www.nature.com/reprints. 418

419

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584

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Supplementary Information is available in the online version of the paper. 585

586

Acknowledgements We thank TCD Lucas, S Meiri, EE Dyer, O Comay and I Pizer-Mason 587

for technical assistance and providing data, and N Kronfeld-Schor for discussion. This work 588

was funded with support from ISF grant No. 785/09 (TD), Tel Aviv University GRTF fund 589

and Naomi Kadar Foundation (RM), and NERC Open CASE PhD studentship 590

(NE/H018565/1) (HFG). 591

592

Author Contributions RM, TD and KEJ developed the overall study design. RM collected 593

and processed the data, and carried out the analyses with assistance from HFG. RM and KEJ 594

led on the writing of the manuscript with significant contributions from all authors. 595

596

Competing Financial Interests The authors declare no competing financial interests. 597

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Figure 1. Activity patterns distribution across (a) the short-fuse (SF), and (b) the long-598

fuse (LF) estimates of mammalian evolution. Species activity patterns are denoted by 599

different colours in the perimeter circle, where nocturnal is denoted as blue; diurnal yellow; 600

cathemeral green; and ambiguous magenta. Branch colours represent taxonomy, where 601

Marsupials are pink; Afrotheria brown; Soricomorpha+Erinaceomorpha green; Chiroptera 602

blue; Cetartiodactyla yellow; Carnivora grey; Primates purple; Rodentia orange; and all other 603

orders are black. Mesozoic and Cenozoic eras are denoted by blue and white backgrounds, 604

respectively. SF phylogeny follows31 updated by32, and LF phylogeny is adapted from30 (see 605

Methods). Branch lengths are proportional to time (Myr). 606

Figure 2. Posterior probability (PP) density of ancestral activity patterns 607

reconstructions of the most recent common ancestor of crown-group Mammalia from 608

(a) SF and (b) LF phylogenies. Distribution curves are calculated from 1000 post-burnin 609

rjMCMC samples, and modal PP values for each distribution are shown in bold. Colours 610

correspond to activity patterns. 611

Figure 3. Reconstruction of ancestral activity patterns and character accumulation, 612

across the ‘short fuse’ (SF) hypothesis of mammalian evolution. (a) Ancestral activity 613

pattern reconstruction across the SF phylogeny31 updated by32. Pie charts correspond to 614

ancestral reconstructions at each node, and colours denote the proportional value of the 615

posterior probability (PP) of each activity pattern, where nocturnal is blue; cathemeral green; 616

and diurnal yellow. Shading denotes geological era. Branch lengths are proportional to time, 617

with branches younger than 45Mya replaced with wedges for visualisation purposes. The red 618

dashed line represents the K-Pg boundary. (b) Lineages through time plot for activity patterns. 619

The predominant activity pattern was assigned to each node based on PP values, with a 620

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minimum value of 0.67. Nodes with reconstructed activity pattern PP values of <0.67 were 621

excluded from the lineages through time plot. 622

Figure 4. Reconstruction of ancestral activity patterns and character accumulation, 623

across the ‘long fuse’ (LF) hypothesis of mammalian evolution. (a) Ancestral activity 624

pattern reconstruction across the LF phylogeny adapted from30. Pie charts correspond to 625

ancestral reconstructions at each node, and colours denote the proportional value of the 626

posterior probability (PP) of each activity pattern, where nocturnal is blue; cathemeral green; 627

and diurnal yellow. Shading denotes geological era. Branch lengths are proportional to time, 628

with branches younger than 45Mya replaced with wedges for visualisation purposes. The red 629

dashed line represents the K-Pg boundary. (b) Lineages through time plot for activity patterns. 630

The predominant activity pattern was assigned to each node based on PP values, with a 631

minimum value of 0.67. Nodes with reconstructed activity pattern PP values of <0.67 were 632

excluded from the lineages through time plot. 633

634

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Maor et al. Evolution of activity patterns in mammals

29

Table 1. Character transition rate matrix for SF and LF ancestral activity pattern 635

reconstructions. Transition rates are from the state in the column to the state in the row and 636

represent model posterior values. Direct transitions between nocturnal and diurnal were not 637

allowed (0) under our character state transition model. 638

639

Phylogeny Transition rates

Short fuse

Nocturnal Cathemeral Diurnal Nocturnal - 0.01 0 Cathemeral 0.03 - 0.03 Diurnal 0 0.01 -

Long fuse

Nocturnal Cathemeral Diurnal Nocturnal - 1.97 0 Cathemeral 7.46 - 7.41 Diurnal 0 1.96 -

640

Page 30: 1 Temporal niche expansion in mammals from a nocturnal ...€¦ · Maor et al. Evolution of activity patterns in mammals 5 91 before the extinction of dinosaurs, calling for a wider

Tachyglossus aculeatus

Zaglossus bruijni

Ornithorhynchus anatinus

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mal

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pum

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pla

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spinosis

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subsp

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melanony

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ruddi

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mys

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ohyd

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Arvi

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us

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xulans

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Akodon affinisAkodon albiventerAkodon azaraeAkodon cursorAkodon iniscatusAbrothrix olivaceusNecromys urichiAkodon boliviensis

Akodon dayiAkodon molinaeAkodon varius

Thaptomys nigritaAbrothrix andinusAbrothrix jelskii

Auliscomys boliviensisLoxodontomys micropusAuliscomys pictusAuliscomys sublimis

Phyllotis caprinusPhyllotis darwiniPhyllotis magisterPhyllotis osgoodiPhyllotis osilaePhyllotis xanthopygus

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Andinomys edax

Kunsia frontoKunsia tomentosusScapteromys tumidus

Necromys lasiurusNecromys obscurus

Calomys hummelinckiChelemys macronyxChinchillula sahamae

Delomys dorsalisDelomys sublineatus

Eligmodontia typusGeoxus valdivianus

Holochilus brasiliensisHolochilus chacariusLundomys molitorHolochilus sciureus

Irenomys tarsalisMelanomys caliginosus

Neacomys guianaeNeacomys pictusNeacomys spinosusNeacomys tenuipes

Nectomys palmipesNectomys rattusNectomys squamipes

Neotomys ebriosusOecomys bicolor

Oligoryzomys chacoensisOligoryzomys delticolaOligoryzomys destructorOligoryzomys flavescensOligoryzomys fulvescensOligoryzomys microtisOligoryzomys nigripes

Oryzomys alfaroiOryzomys bolivarisOryzomys angouyaOryzomys megacephalusOryzomys chapmaniOryzomys couesiOryzomys russatusOryzomys macconnelliOryzomys melanotisOryzomys nitidusOryzomys palustrisOryzomys rostratusOryzomys subflavusOryzomys talamancae

Oxymycterus nasutusOxymycterus rufus

Punomys lemminusReithrodon auritus

Rhipidomys austrinusRhipidomys caucensisRhipidomys couesiRhipidomys fulviventerRhipidomys latimanusRhipidomys leucodactylusRhipidomys macconnelliRhipidomys mastacalisRhipidomys nitelaRhipidomys ochrogasterRhipidomys venezuelaeRhipidomys venustusRhipidomys wetzeli

Scolomys melanopsScolomys ucayalensis

Sigmodontomys alfari

Wiedomys pyrrhorhinos

Wilfredomys oenaxZygodontomys brevicauda

Ichthyomys pittieriIchthyomys stolzmanni

Ichthyomys tweediiNeusticomys monticolus

Rheomys raptorSigmodon alstoniSigmodon arizonae

Sigmodon hispidusSigmodon toltecusSigmodon ochrognathus

Nyctomys sumichrasti

Otonyctomys hatti

Ototylomys phyllotis

Tylomys miraeTylomys nudicaudus

Tylomys tumbalensis

Tylomys watsoni

Baiomys musculus

Baiomys taylori

Scotinomys teguina

Ochrotomys nuttalli

Habromys lophurus

Neotomodon alstoni

Podomys floridanus

Osgoodomys banderanus

Peromyscus attwateri

Peromyscus aztecus

Peromyscus boylii

Peromyscus bullatus

Peromyscus californicus

Peromyscus caniceps

Peromyscus crinitus

Peromyscus dickeyi

Peromyscus difficilis

Peromyscus eremicus

Peromyscus eva

Peromyscus furvus

Peromyscus gossypinus

Peromyscus grandis

Peromyscus gratus

Peromyscus guardia

Peromyscus guatemalensis

Peromyscus gymnotis

Peromyscus hooperi

Peromyscus interparietalis

Peromyscus leucopus

Peromyscus levipes

Peromyscus madrensis

Peromyscus maniculatus

Peromyscus mayensis

Peromyscus megalops

Peromyscus mekisturus

Peromyscus melanocarpus

Peromyscus melanophrys

Peromyscus melanotis

Peromyscus melanurus

Peromyscus merriami

Peromyscus mexicanus

Peromyscus nasutus

Peromyscus ochraventer

Peromyscus keeni

Peromyscus pectoralis

Peromyscus perfulvus

Peromyscus polionotus

Peromyscus polius

Peromyscus pseudocrinitus

Peromyscus sejugis

Peromyscus simulus

Peromyscus slevini

Peromyscus spicilegus

Peromyscus stephani

Peromyscus stirtoni

Peromyscus truei

Peromyscus winkelmanni

Peromyscus yucatanicus

Peromyscus zarhynchusOnychomys leucogaster

Onychomys torridus

Isthmomys flavidus

Isthmomys pirrensis

Reithrodontomys brevirostris

Reithrodontomys darienensis

Reithrodontomys gracilis

Reithrodontomys mexicanus

Reithrodontomys paradoxus

Reithrodontomys spectabilis

Reithrodontomys creper

Reithrodontomys microdon

Reithrodontomys rodriguezi

Reithrodontomys tenuirostrisReithrodontomys fulve

scens

Reithrodontomys hirsutus

Reithrodontomys burti

Reithrodontomys chrysopsis

Reithrodontomys humulis

Reithrodontomys megalotis

Reithrodontomys montanus

Reithrodontomys raviventris

Reithrodontomys sumichrasti

Reithrodontomys zacatecae

Xenomys nelsoni

Neotoma albigula

Neotoma bryanti

Neotoma devia

Neotoma floridana

Neotoma magisterNeotoma lepida

Neotoma martinensis

Neotoma mexicana

Neotoma micropus

Neotoma phenax

Neotoma cinerea

Neotoma fuscipes

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Mer

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Mer

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Mer

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Mer

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Psa

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Psa

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Des

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Des

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Den

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Ste

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Cha

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Dip

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Dip

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Mic

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Tho

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Thom

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Geo

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Geo

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Geo

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Het

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Het

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Het

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Cra

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Octodon bridgesi

Octodon degus

Massoutiera m

zabi

Pectinator spekei

Ctenodactylus gundi

Ctenodactylus vali

Felovia vae

Tympanoctom

ys barrerae

Octom

ys mim

ax

Ctenom

ys haigi

Ctenom

ys opimus

Ctenom

ys torquatus

Ctenom

ys talarum

Abrocom

a bennettii

Hydrochoerus hydrochaeris

Dinom

ys branickii

Petrom

us typicus

Kerodon rupestrisC

avia porcellus

Cavia aperea

Microcavia australis

Galea m

usteloides

Galea spixii

Galea flavidens

Myoprocta acouchy

Dasyprocta punctata

Dasyprocta leporina

Dasyprocta azarae

Dasyprocta coibae

Dasyprocta cristata

Dasyprocta fuliginosa

Dasyprocta guam

ara

Dasyprocta kalinow

skii

Dasyprocta m

exicana

Dasyprocta prym

nolopha

Dasyprocta ruatanica

Myocastor coypus

Trinomys dim

idiatusH

oplomys gym

nurus

Proechim

ys quadruplicatusP

roechimys sim

onsi

Euryzygom

atomys spinosus

Lonchothrix emiliae

Proechim

ys longicaudatus

Proechim

ys trinitatus

Carterodon sulcidens

Chaetom

ys subspinosus

Proechim

ys brevicauda

Proechim

ys canicollis

Proechim

ys guyannensis

Proechim

ys cuvieri

Proechim

ys semispinosus

Proechim

ys guairae

Proechim

ys hoplomyoides

Proechim

ys magdalenae

Proechim

ys oconnelli

Proechim

ys poliopus

Proechim

ys steerei

Makalata didelphoides

Echim

ys chrysurusP

hyllomys blainvillii

Phyllom

ys brasiliensisP

hyllomys dasythrix

Makalata grandis

Phyllom

ys lamarum

Phyllom

ys nigrispinusE

chimys sem

ivillosusP

hyllomys thom

asi

Isothrix bistriataIsothrix pagurus

Diplom

ys canicepsD

iplomys labilis

Diplom

ys rufodorsalis

Kannabateom

ys amblyonyx

Dactylom

ys dactylinusD

actylomys boliviensis

Olallam

ys edaxM

esomys hispidus

Mesom

ys lenicepsM

akalata obscuraM

esomys stim

ulax

Caprom

ys piloridesG

eocapromys brow

nii

Geocaprom

ys ingrahami

Mesocaprom

ys melanurus

Plagiodontia aedium

Dolichotis patagonum

Dolichotis salinicola

Cuniculus paca

Cuniculus taczanow

skii

Trichys fasciculata

Hystrix africaeaustralis

Hystrix cristata

Hystrix brachyura

Hystrix indica

Atherurus m

acrourus

Atherurus africanus

Thryonom

ys swinderianus

Thryonom

ys gregorianus

Heterocephalus glaber

Heliophobius argenteocinereus

Cryptom

ys damarensis

Cryptom

ys hottentotus

Cryptom

ys mechow

i

Georychus capensis

Bathyergus suillus

Erethizon dorsatum

Sphiggurus insidiosus

Sphiggurus villosus

Sphiggurus m

exicanus

Sphiggurus spinosus

Sphiggurus vestitus

Coendou bicolor

Coendou prehensilis

Coendou rothschildi

Lagostomus m

aximus

Chinchilla lanigera

Chinchilla chinchilla

Lagidium peruanum

Lagidium viscacia

Lagidium w

olffsohni

Spalacopus cyanus

Octodontom

ys gliroides

Aconaem

ys sagei

Aconaem

ys fuscus

Aplodontia rufa

Marm

ota menzbieri

Marm

ota caudata

Marm

ota himalayana

Marm

ota marm

ota

Marm

ota monax

Marm

ota flaviventris

Marm

ota olympus

Marm

ota caligata

Marm

ota vancouverensis

Spermophilus colum

bianus

Spermophilus parryii

Spermophilus richardsonii

Spermophilus arm

atus

Spermophilus tridecem

lineatus

Spermophilus franklinii

Spermophilus washingtoni

Spermophilus spilosom

a

Spermophilus beecheyi

Spermophilus citellus

Spermophilus suslicus

Spermophilus beldingi

Spermophilus adocetus

Spermophilus m

ollis

Spermophilus brunneus

Spermophilus tow

nsendii

Spermophilus tereticaudus

Spermophilus m

ohavensis

Spermophilus saturatus

Spermophilus m

exicanus

Spermophilus elegans

Spermophilus lateralis

Spermophilus variegatus

Cynomys gunnisoni

Cynomys leucurus

Cynomys parvidens

Cynomys ludovicianus

Cynomys m

exicanus

Amm

ospermophilus leucurus

Amm

ospermophilus nelsoni

Amm

ospermophilus interpres

Amm

ospermophilus harrisii

Tamias striatus

Tamias sibiricus

Tamias am

oenus

Tamias quadrim

aculatus

Tamias canipes

Tamias rufus

Tamias quadrivittatus

Tamias dorsalis

Tamias palm

eri

Tamias um

brinus

Tamias townsendii

Tamias senex

Tamias m

erriami

Tamias obscurus

Tamias ruficaudus

Tamias ochrogenys

Tamias speciosus

Tamias m

inimus

Tamias alpinus

Tamias panam

intinus

Myosciurus pum

ilio

Heliosciurus ruwenzorii

Heliosciurus gambianus

Heliosciurus mutabilis

Heliosciurus punctatus

Heliosciurus rufobrachium

Epixerus ebii

Protoxerus stangeriFunisciurus pyrropus

Funisciurus anerythrus

Funisciurus congicus

Funisciurus isabella

Funisciurus lemniscatus

Paraxerus cepapi

Paraxerus flavovittis

Paraxerus lucifer

Paraxerus palliatus

Paraxerus poensis

Spermophilopsis leptodactylus

Atlantoxerus getulus

Xerus rutilus

Xerus erythropus

Xerus inauris

Xerus princeps

Tamiasciurus hudsonicus

Tamiasciurus douglasii

Sciurus carolinensis

Sciurus niger

Sciurus granatensis

Sciurus flam

mifer

Sciurus variegatoides

Sciurus anom

alus

Sciurus stram

ineus

Sciurus lis

Sciurus igniventris

Sciurus spadiceus

Sciurus arizonensis

Sciurus oculatus

Sciurus griseus

Sciurus alleni

Sciurus richm

ondi

Sciurus colliaei

Sciurus sanborni

Sciurus aureogaster

Sciurus pyrrhinus

Sciurus aestuans

Sciurus vulgaris

Sciurus ignitus

Sciurus yucatanensis

Sciurus gilvigularis

Sciurus pucheranii

Sciurus nayaritensis

Sciurus deppei

Microsciurus flaviventer

Microsciurus m

imulus

Microsciurus santanderensis

Microsciurus alfari

Syntheosciurus brochus

Pterom

ys volans

Pterom

ys mom

onga

Petaurista petaurista

Petaurista philippensisPetaurista alborufus

Petaurista nobilis

Petaurista magnificus

Petaurista elegans

Petaurista xanthotis

Trogopterus xanthipes

Pterom

yscus pulverulentus

Aerom

ys tephromelas

Aerom

ys thomasi

Eupetaurus cinereus

Aeretes m

elanopterus

Glaucom

ys volans

Glaucom

ys sabrinus

Eoglaucom

ys fimbriatus

Hylopetes lepidus

Petinomys setosus

Petinomys crinitus

Petinomys genibarbis

Petinomys hageni

Petinomys sagitta

Petinomys vorderm

anni

Hylopetes alboniger

Hylopetes nigripes

Hylopetes spadiceus

Petaurillus kinlochii

Petaurillus hosei

Iomys horsfieldiiR

atufa bicolor

Ratufa affinis

Ratufa indica

Ratufa m

acroura

Callosciurus prevostii

Callosciurus caniceps

Callosciurus finlaysonii

Callosciurus orestes

Callosciurus adam

si

Callosciurus notatus

Callosciurus inornatus

Callosciurus baluensis

Callosciurus phayrei

Callosciurus m

elanogaster

Callosciurus quinquestriatus

Lariscus insignis

Lariscus obscurus

Rhinosciurus laticaudatus

Funambulus palm

arum

Funambulus pennantii

Tamiops m

cclellandii

Tamiops m

aritimus

Tamiops rodolphii

Tamiops sw

inhoei

Drem

omys rufigenis

Drem

omys lokriah

Nannosciurus m

elanotisSundasciurus hippurus

Sundasciurus lowii

Sundasciurus tenuis

Callosciurus nigrovittatus

Callosciurus pygerythrus

Sciurillus pusillus

Dryomys laniger

Dryomys nitedula

Chaetocauda sichuanensis

Eliomys melanurus

Eliomys quercinus

Selevinia betpakdalaensis

Muscardinus avellanarius

Glis glis

Glirulus japonicus

Graphiurus christyi

Graphiurus kelleni

Graphiurus lorraineus

Graphiurus murinus

Graphiurus ocularis

Graphiurus platyops

Graphiurus surdus

Graphiurus crassicaudatus

Graphiurus nagtglasii

Nesolagus netscheri

Sylvilagus aquaticus

Sylvilagus brasiliensis

Sylvilagus palustris

Sylvilagus audubonii

Sylvilagus bachmani

Caprolagus hispidus

Poelagus marjorita

Lepus alleni

Lepus callotis

Lepus flavigularis

Lepus californicus

Lepus americanus

Lepus arcticus

Lepus timidus

Lepus townsendii

Lepus brachyurus

Lepus europaeus

Lepus comus

Lepus capensis

Lepus saxatilis

Lepus microtis

Lepus hainanus

Lepus nigricollis

Oryctolagus cuniculusBunolagus monticularis

Romerolagus diazi

Pentalagus furnessi

Pronolagus crassicaudatus

Pronolagus rupestris

Pronolagus randensis

Ochotona collaris

Ochotona princeps

Ochotona koslowi

Ochotona rufescens

Ochotona macrotis

Ochotona roylei

Ochotona curzoniae

Allenopithecus nigroviridis

Cercopithecus ascanius

Cercopithecus cephus

Cercopithecus erythrogaster

Cercopithecus erythrotis

Cercopithecus petaurista

Cercopithecus sclateri

Cercopithecus mitis

Cercopithecus albogularis

Cercopithecus nictitans

Cercopithecus campbelli

Cercopithecus mona

Cercopithecus pogonias

Cercopithecus wolfi

Cercopithecus denti

Cercopithecus diana

Cercopithecus dryas

Cercopithecus neglectus

Cercopithecus hamlyniCercopithecus lhoesti

Cercopithecus preussi

Cercopithecus solatus

Chlorocebus aethiops

Chlorocebus pygerythrus

Chlorocebus sabaeus

Chlorocebus tantalusErythrocebus patas

Miopithecus talapoin

Cercocebus agilis

Cercocebus galeritus

Cercocebus torquatus

Cercocebus atys

Mandrillus leucophaeus

Mandrillus sphinx

Lophocebus albigena

Lophocebus aterrimus

Papio hamadryas

Papio anubis

Papio cynocephalus

Papio papio

Papio ursinus

Theropithecus gelada

Macaca arctoides

Macaca assamensis

Macaca thibetana

Macaca radiata

Macaca sinica

Macaca cyclopis

Macaca mulatta

Macaca fuscata

Macaca fascicularis

Macaca maura

Macaca nigra

Macaca nigrescens

Macaca ochreata

Macaca tonkeana

Macaca hecki

Macaca nemestrina

Macaca leonina

Macaca pagensis

Macaca silenus

Macaca sylvanus

Colobus angolensis

Colobus guereza

Colobus polykomos

Colobus vellerosusColobus satanas

Piliocolobus badius

Piliocolobus preussi

Piliocolobus kirkii

Piliocolobus pennantii

Piliocolobus rufomitratus

Procolobus verus

Simias concolor

Nasalis larvatus

Presbytis comata

Presbytis hosei

Presbytis thomasi

Presbytis femoralis

Presbytis siamensis

Presbytis melalophos

Presbytis frontata

Presbytis potenziani

Presbytis rubicunda

Semnopithecus entellus

Trachypithecus auratus

Trachypithecus cristatus

Trachypithecus francoisi

Trachypithecus delacouri

Trachypithecus laotum

Trachypithecus poliocephalus

Trachypithecus geei

Trachypithecus pileatus

Trachypithecus obscurus

Trachypithecus phayrei

Trachypithecus johnii

Trachypithecus vetulusRhinopithecus avunculus

Rhinopithecus bieti

Rhinopithecus brelichi

Rhinopithecus roxellana

Pygathrix nemaeus

Gorilla gorilla

Homo sapiens

Pan paniscusPan troglodytesPongo pygmaeusPongo abelii

Hylobates agilisHylobates lar

Hylobates moloch

Hylobates muelleri

Hylobates pileatus

Hylobates klossii

Bunopithecus hoolock

Symphalangus syndactylus

Nomascus concolor

Nomascus leucogenys

Nomascus gabriellae

Alouatta belzebulAlouatta caraya

Alouatta coibensisAlouatta palliata

Alouatta guaribaAlouatta pigra

Alouatta saraAlouatta seniculus

Ateles belzebuth

Ateles chamekAteles marginatusAteles fuscicepsAteles geoffroyiAteles paniscusBrachyteles arachnoidesBrachyteles hypoxanthusOreonax flavicaudaLagothrix lagotricha

Cacajao calvusCacajao melanocephalusChiropotes albinasusChiropotes satanas

Pithecia aequatorialis

Pithecia albicansPithecia irrorataPithecia monachusPithecia pithecia

Callicebus brunneusCallicebus hoffmannsi

Callicebus cinerascensCallicebus moloch

Callicebus caligatusCallicebus cupreusCallicebus dubius

Callicebus donacophilusCallicebus modestusCallicebus oenantheCallicebus olallaeCallicebus torquatus

Aotus azaraeAotus lemurinusAotus hershkovitziAotus miconaxAotus nancymaaeAotus nigricepsAotus trivirgatusAotus vociferans

Callicebus personatus

Callimico goeldii

Callithrix argentataCallithrix emiliae

Callithrix intermediaCallithrix leucippe

Callithrix humeralifera

Callithrix chrysoleucaCallithrix pygmaea

Callithrix auritaCallithrix flaviceps

Callithrix geoffroyiCallithrix kuhlii

Callithrix jacchusCallithrix penicillataLeontopithecus caissara

Leontopithecus chrysopygus

Leontopithecus rosalia

Leontopithecus chrysomelas

Saguinus bicolorSaguinus midas

Saguinus geoffroyiSaguinus oedipus

Saguinus leucopusSaguinus imperator

Saguinus labiatusSaguinus mystax

Saguinus fuscicollisSaguinus tripartitusSaguinus nigricollis

Saguinus inustusCebus albifronsCebus capucinusCebus olivaceusCebus apella

Cebus xanthosternos

Saimiri boliviensisSaimiri vanzolinii

Saimiri oerstediiSaimiri sciureusSaimiri ustus

Tarsius bancanusTarsius dentatusTarsius pumilusTarsius tarsierTarsius syrichta

Allocebus trichotisMirza coquereli

Microcebus murinusMicrocebus griseorufus

Microcebus myoxinusMicrocebus rufus

Cheirogaleus majorCheirogaleus crossleyi

Cheirogaleus mediusPhaner furcifer

Avahi lanigerAvahi occidentalis

Propithecus diademaPropithecus edwardsi

Propithecus perrieri

Propithecus tattersalli

Propithecus verreauxiPropithecus coquereli

Indri indriDaubentonia madagascariensis

Eulemur coronatus

Eulemur fulvusEulemur albocollarisEulemur collarisEulemur rufusEulemur sanfordiEulemur macacoEulemur mongozEulemur rubriventer

Hapalemur aureus

Hapalemur griseusHapalemur alaotrensisHapalemur occidentalis

Prolemur simus

Lemur catta

Varecia variegataVarecia rubra

Lepilemur dorsalisLepilemur edwardsiLepilemur leucopusLepilemur microdon

Lepilemur mustelinusLepilemur ruficaudatus

Lepilemur septentrionalisLepilemur ankaranensis

Arctocebus aureus

Arctocebus calabarensis

Perodicticus potto

Loris tardigradus

Loris lydekkerianus

Nycticebus coucang

Nycticebus pygmaeus

Euoticus elegantulusEuoticus pallidus

Galago gallarumGalago matschiei

Galago moholiGalago senegalensis

Otolemur crassicaudatusOtolemur garnettii

Galago alleniGalago demidoff

Galago zanzibaricus

Galeopterus variegates

Cynocephalus volans

Ptilocercus lowiiAnathana elliotiDendrogale melanura

Dendrogale murina

Tupaia belangeri

Tupaia dorsalisTupaia glis

Tupaia gracilis

Tupaia javanica

Tupaia longipesTupaia minor

Tupaia montana

Tupaia nicobarica

Tupaia palawanensisTupaia picta

Tupaia splendidulaTupaia tana

Addax

nas

omac

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Ory

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Ory

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Ory

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Hippo

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Hippo

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Alcela

phus

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Alcela

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Conno

chae

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Conno

chae

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taur

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Damal

iscus

luna

tus

Damal

iscus

pyg

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Amm

otra

gus

lerv

ia

Budor

cas t

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Ovis a

mm

on

Ovis a

ries

Ovis d

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Capra

cauc

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Capra

falco

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Capra

hirc

us

Capra

pyr

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Capra

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Hemitr

agus

hylo

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Hemitr

agus

jaya

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Hemitr

agus

jem

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Pseud

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Pseud

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Rupica

pra

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Rupica

pra

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Capric

orni

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Capric

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s m

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dwar

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Ovib

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Ore

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Aepy

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Antid

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s m

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Antil

ope

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Gaz

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ben

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Gaz

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sub

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Gaz

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dor

cas

Gaz

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gaz

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Nan

ger d

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Nan

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Nan

ger g

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Eudo

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thom

soni

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Lito

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walle

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Saig

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loph

us c

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Cepha

loph

us o

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Cepha

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us d

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Cepha

loph

us s

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Cepha

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us s

padi

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Cepha

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us je

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Cepha

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us n

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Cepha

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us n

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Cepha

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us ru

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Cepha

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us le

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Cepha

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Sylvi

capr

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Phila

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Phila

ntom

ba m

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Dor

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gus

meg

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Rap

hice

rus

cam

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Rap

hice

rus

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Rap

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rus

mel

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Kobu

s el

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prym

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Kobu

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Red

unca

aru

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Red

unca

redu

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Red

unca

fulvo

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Pele

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Mad

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Mad

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Mad

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Mad

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Neo

tragu

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Neo

tragu

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Neo

tragu

s py

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Ore

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Our

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our

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Proc

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Biso

n bi

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Biso

n bo

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Bos

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Bos

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Bos

front

alis

Bos

java

nicu

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Buba

lus

buba

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Buba

lus

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corn

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Buba

lus

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Buba

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Bos

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Tetra

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Taur

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Trag

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Trag

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anga

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Trag

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eury

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Trag

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spek

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Trag

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Trag

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Alc

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Bla

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Hip

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Hip

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Ozo

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Maz

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Maz

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Odo

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us v

irgin

ianu

sP

udu

puda

Ran

gife

r tar

andu

s

Axi

s ax

is

Axi

s ca

lam

iane

nsis

Axi

s ku

hlii

Axi

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rcin

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Ruc

ervu

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vauc

elii

Cer

vus

elap

hus

Ruc

ervu

s el

dii

Cer

vus

nipp

on

Rus

a tim

oren

sis

Rus

a un

icol

orM

untia

cus

athe

rode

s

Mun

tiacu

s m

untja

k

Mos

chus

ber

ezov

skii

Mos

chus

chr

ysog

aste

r

Mos

chus

fusc

us

Mos

chus

mos

chife

rus

Ant

iloca

pra

amer

ican

a

Gira

ffa c

amel

opar

dalis

Oka

pia

john

ston

i

Hye

mos

chus

aqu

atic

us

Mos

chio

la m

emin

na

Trag

ulus

java

nicu

s

Trag

ulus

nap

u

Cep

halo

rhyn

chus

hec

tori

Ste

nella

coe

rule

oalb

a

Sot

alia

fluv

iatil

is

Ste

nella

long

irost

ris

Inia

geo

ffren

sis

Esc

hric

htiu

s ro

bust

us

Hex

apro

todo

n lib

erie

nsis

Hip

popo

tam

us a

mph

ibiu

s

Bab

yrou

sa b

abyr

ussa

Hyl

ocho

erus

mei

nert

zhag

eni

Pota

moc

hoer

us la

rvat

us

Pota

moc

hoer

us p

orcu

s

Sus

bar

batu

s

Sus

ceb

ifron

s

Sus

cel

eben

sis

Sus

phi

lippe

nsis

Sus

sal

vani

usS

us s

crof

aP

haco

choe

rus

aeth

iopi

cus

Pha

coch

oeru

s af

rican

us

Cat

agon

us w

agne

ri

Taya

ssu

peca

riPe

cari

taja

cuC

amel

us d

rom

edar

ius

Lam

a gl

ama

Vic

ugna

vic

ugna

Cer

atot

heriu

m s

imum

Dic

eros

bic

orni

s

Dic

eror

hinu

s su

mat

rens

is

Rhi

noce

ros

unic

orni

sTa

piru

s ba

irdii

Tapi

rus

pinc

haqu

e

Tapi

rus

terr

estr

isTa

piru

s in

dicu

sE

quus

asi

nus

Equ

us z

ebra

Equ

us h

emio

nus

Equ

us c

abal

lus

Mustela lutreola

Mustela lutreolina

Mustela nudipes

Mustela sibirica

Mustela eversmanii

Mustela nigripes

Mustela putorius

Mustela altaica

Mustela erminea

Mustela frenata

Mustela nivalis

Mustela africana

Neovison macrodon

Neovison vison

Martes americana

Martes melampus

Martes zibellina

Martes martes

Martes foina

Martes flavigula

Martes gwatkinsii

Martes pennanti

Gulo gulo

Eira barbara

Galictis cuja

Galictis vittata

Lyncodon patagonicus

Ictonyx libyca

Ictonyx striatus

Vormela peregusna

Poecilogale albinucha

Mellivora capensis

Arctonyx collarisMeles meles

Meles anakuma

Mydaus javanensis

Mydaus marchei

Melogale everetti

Melogale moschata

Melogale orientalis

Melogale personata

Lontra felina

Lontra provocax

Lontra longicaudis

Lontra canadensis

Lutra lutra

Aonyx capensis

Aonyx cinereaHydrictis maculicollis

Lutrogale perspicillata

Pteronura brasiliensis

Enhydra lutris

Conepatus chinga

Conepatus leuconotus

Conepatus humboldtii

Conepatus semistriatusMephitis macroura

Mephitis mephitisSpilogale putorius

Spilogale gracilis

Spilogale pygmaea

Taxidea taxus

Procyon cancrivorus

Procyon lotor

Procyon pygmaeus

Nasua narica

Nasua nasua

Nasuella olivacea

Bassariscus astutus

Bassariscus sumichrasti

Bassaricyon beddardi

Bassaricyon gabbii

Bassaricyon lasius

Bassaricyon pauli

Potos flavus

Ailurus fulgens

Odobenus rosmarus

Arctocephalus australis

Arctocephalus forsteri

Arctocephalus galapagoensis

Arctocephalus gazella

Arctocephalus pusillus

Neophoca cinerea

Eumetopias jubatus

Zalophus californianus

Callorhinus ursinus

Pusa sibiricaPhoca largha

Phoca vitulina

Hydrurga leptonyx

Lobodon carcinophaga

Leptonychotes weddellii

Mirounga angustirostris

Mirounga leonina

Monachus schauinslandi

Monachus monachus

Ursus arctos

Ursus maritimus

Ursus thibetanus

Helarctos malayanus

Melursus ursinus

Ursus americanus

Tremarctos ornatus

Ailuropoda melanoleuca

Canis lupus

Canis latrans

Canis simensis

Canis adustus

Canis aureus

Lycalopex culpaeus

Lycalopex griseus

Lycalopex gymnocercus

Lycalopex sechurae

Lycalopex vetulus

Atelocynus microtis

Cerdocyon thous

Chrysocyon brachyurus

Cuon alpinus

Lycaon pictus

Speothos venaticus

Nyctereutes procyonoides

Vulpes corsac

Vulpes rueppellii

Vulpes vulpes

Vulpes lagopusVulpes velox

Vulpes macrotis

Vulpes cana

Vulpes zerda

Vulpes bengalensis

Vulpes chama

Vulpes pallida

Urocyon cinereoargenteus

Urocyon littoralis

Otocyon megalotis

Panthera leo

Panthera pardus

Panthera onca

Panthera tigris

Uncia uncia

Neofelis nebu

losa

Pardofelis m

armorata

Lynx c

anadensis

Lynx l

ynx

Lynx p

ardinu

s

Lynx r

ufus

Catopuma temminck

ii

Profelis

aurata

Leopardus t

igrinus

Leop

ardu

s geo

ffroy

i

Leop

ardu

s guig

na

Leop

ardu

s colo

colo

Leop

ardu

s jac

obitu

s

Leop

ardu

s par

dalis

Leop

ardu

s wied

ii

Felis m

arga

rita

Felis n

igripe

s

Felis s

ilves

tris

Felis b

ieti

Felis c

haus

Felis m

anul

Carac

al ca

raca

l

Lepta

iluru

s ser

val

Priona

iluru

s ben

galen

sis

Priona

iluru

s irio

moten

sis

Priona

iluru

s vive

rrinu

s

Priona

iluru

s rub

igino

sus

Priona

iluru

s plan

iceps

Puma

yago

uaro

undi

Puma

conc

olor

Acinon

yx ju

batu

s

Crocu

ta cr

ocut

a

Hyaen

a hy

aena

Hyaen

a br

unne

a

Prote

les cr

istat

a

Helogale hirtula

Helogale parvula

Mungos gambianus

Mungos mungo

Crossarchus alexandri

Crossarchus ansorgei

Crossarchus obscurus

Ichneumia albicauda

Suricata suricatta

Bdeogale crassicauda

Bdeogale jacksoni

Bdeogale nigripes

Liberiictis kuhni

Cynictis penicillata

Paracynictis selousi

Herpestes edwardsi

Herpestes javanicus

Herpestes brachyurus

Herpestes fuscus

Herpestes urva

Herpestes ichneumon

Herpestes naso

Herpestes semitorquatus

Herpestes smithii

Herpestes vitticollis

Galerella pulverulenta

Galerella sanguinea

Atilax p

aludinosus

Dologale dybowski

i

Rhynch

ogale melleri

Galidictis fa

sciata

Galidictis g

randidieri

Galidia elegans

Mungotictis d

ecemlineata

Salanoia conco

lor

Genetta abyss

inica

Genetta angolensis

Genetta m

aculata

Genetta genetta

Genetta tig

rina

Genetta se

rvalin

a

Genetta cr

istata

Prionodon lin

sang

Prionodon pardico

lor

Poiana richardso

nii

Genetta pisc

ivora

Viverra

cive

ttina

Viverra

megasp

ila

Viverra

tangalunga

Viverra

zibetha

Viverric

ula indica

Civettic

tis ci

vetta

Paradoxurus j

erdoni

Paradoxurus z

eylonensis

Paradoxurus h

ermaphroditu

s

Arctict

is binturong

Paguma larva

ta

Macrogalid

ia muss

chenbroekii

Arctogalid

ia trivir

gata

Nandinia binotata

Chrotogale owstoni

Diplogale hosei

Hemigalus derbya

nus

Cynogale bennettii

Eupleres goudotii

Fossa fo

ssana

Cryptoprocta

ferox

Man

is cr

assic

auda

ta

Man

is pe

ntad

actyl

a

Man

is te

mm

inckii

Nyctim

ene robinsoni

Cynopterus sphinx

Megaloglossus w

oermanni

Eidolon helvum

Eidolon dupreanum

Myonycteris torquata

Rousettus aegyptiacus

Rousettus leschenaultii

Rousettus m

adagascariensis

Scotonycteris zenkeri

Nanonycteris veldkam

pii

Hypsignathus m

onstrosus

Epom

ops franqueti

Micropteropus pusillus

Epom

ophorus wahlbergi

Epom

ophorus angolensis

Epom

ophorus gambianus

Macroglossus m

inimus

Syconycteris australis

Dobsonia m

oluccensis

Pteropus m

elanotusP

teropus hypomelanus

Pteropus alecto

Pteropus dasym

allus

Pteropus livingstonii

Pteropus conspicillatus

Pteropus sam

oensis

Pteropus tonganus

Pteropus poliocephalus

Pteropus m

olossinus

Pteropus giganteus

Pteropus vam

pyrus

Pteropus scapulatus

Pteropus insularisS

accolaimus peli

Saccolaim

us flaviventrisS

accolaimus m

ixtusS

accolaimus saccolaim

us

Taphozous hamiltoni

Taphozous nudiventris

Taphozous theobaldi

Taphozous hilli

Taphozous australisTaphozous georgianusTaphozous kapalgensis

Taphozous hildegardeaeTaphozous longim

anusTaphozous m

auritianusTaphozous perforatusTaphozous m

elanopogon

Mosia nigrescens

Em

ballonura atrataE

mballonura beccarii

Em

ballonura dianaeE

mballonura furax

Em

ballonura raffrayanaE

mballonura sem

icaudata

Coleura afra

Coleura seychellensis

Em

ballonura alectoE

mballonura m

onticola

Rhynchonycteris naso

Cyttarops alecto

Diclidurus isabellus

Diclidurus scutatus

Diclidurus albus

Diclidurus ingens

Centronycteris m

aximiliani

Corm

ura brevirostris

Saccopteryx bilineata

Saccopteryx leptura

Saccopteryx gym

nuraS

accopteryx canescens

Balantiopteryx io

Balantiopteryx infusca

Balantiopteryx plicata

Peropteryx leucoptera

Peropteryx kappleri

Peropteryx m

acrotis

Craseonycteris thonglongyai

Cardioderm

a corLavia fronsM

acroderma gigas

Nycteris thebaica

Nycteris grandis

Nycteris hispida

Rhinolophus blasii

Rhinolophus capensis

Rhinolophus clivosus

Rhinolophus darlingi

Rhinolophus denti

Rhinolophus ferrum

equinumR

hinolophus hildebrandtiiR

hinolophus hipposiderosR

hinolophus megaphyllus

Rhinolophus philippinensis

Rhinolophus sim

ulatorR

hinolophus swinnyi

Rhinolophus landeri

Rhinonicteris aurantia

Triaenops furculusTriaenops rufus

Hipposideros ater

Hipposideros caffer

Hipposideros speoris

Hipposideros cervinus

Cloeotis percivali

Hipposideros com

mersoni

Thyroptera discifera

Thyroptera tricolor

Natalus stram

ineus

Mystacina tuberculata

Noctilio albiventris

Noctilio leporinus

Morm

oops megalophylla

Pteronotus davyi

Pteronotus personatus

Pteronotus m

acleayii

Desm

odus rotundus

Macrophyllum

macrophyllum

Lonchorhina aurita

Macrotus californicus

Macrotus w

aterhousiiLam

pronycteris brachyotisM

icronycteris megalotis

Vam

pyrum spectrum

Lophostoma silvicolum

Phyllostom

us discolorP

hyllostomus hastatus

Bra

chyp

hylla

cav

erna

rum

Bra

chyp

hylla

nan

aP

hyllo

nyct

eris

aph

ylla

Lonc

hoph

ylla

mor

dax

Lonc

hoph

ylla

rob

usta

Lept

onyc

teris

cur

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eLe

pton

ycte

ris y

erba

buen

aeLe

pton

ycte

ris n

ival

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loss

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omm

issa

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Glo

ssop

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sor

icin

aA

nour

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ltrat

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a ge

offr

oyi

Cho

eron

ycte

ris m

exic

ana

Rhi

noph

ylla

pum

ilio

Car

ollia

cas

tane

aC

arol

lia p

ersp

icill

ata

Stu

rnira

liliu

mS

turn

ira e

ryth

rom

os

Cen

turio

sen

exS

phae

rony

cter

is to

xoph

yllu

m

Phy

llops

falc

atus

Arit

eus

flave

scen

s

Art

ibeu

s ja

mai

cens

isA

rtib

eus

litur

atus

Vam

pyre

ssa

pusi

llaV

ampy

ress

a bi

dens

Vam

pyro

des

cara

ccio

liP

laty

rrhi

nus

helle

ri

Che

irom

eles

torq

uatu

sM

yopt

erus

dau

bent

onii

Cyn

omop

s gr

eenh

alli

Cyn

omop

s pl

aniro

stris

Mor

mop

teru

s be

ccar

iiM

orm

opte

rus

norf

olke

nsis

Mor

mop

teru

s pl

anic

eps

Tada

rida

bras

ilien

sis

Tada

rida

fulm

inan

s

Nyc

tinom

ops

fem

oros

accu

sN

yctin

omop

s la

ticau

datu

sN

yctin

omop

s m

acro

tis

Mol

ossu

s si

nalo

ae

Mol

ossu

s cu

rren

tium

Mol

ossu

s m

olos

sus

Mol

ossu

s ru

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Mol

ossu

s pr

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sus

Eum

ops

pero

tis

Eum

ops

glau

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umop

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dulu

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Min

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sch

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inio

pter

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Lasi

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Nyc

ticei

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umer

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Sco

tean

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Oto

nyct

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hem

pric

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Ant

rozo

us p

allid

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espe

rtili

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urin

usN

eoro

mic

ia n

anus

His

tiotu

s m

onta

nus

Sco

tore

pens

gre

yii

Sco

tore

pens

san

born

i

Hyp

sugo

bod

enhe

imer

iV

espa

delu

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lturn

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azor

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Cha

linol

obus

tube

rcul

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Cha

linol

obus

dw

yeri

Cha

linol

obus

gou

ldii

Cha

linol

obus

mor

io

Cha

linol

obus

nig

rogr

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Lasi

urus

egr

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Lasi

urus

ega

Lasi

urus

inte

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Lasi

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cin

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Lasi

urus

bor

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Lasi

urus

sem

inol

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Nyc

toph

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arnh

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sis

Nyc

toph

ilus

geof

froy

iN

ycto

philu

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uldi

Nyc

toph

ilus

timor

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is

Bar

bast

ella

bar

bast

ellu

s

Eud

erm

a m

acul

atum

Idio

nyct

eris

phy

llotis

Cor

ynor

hinu

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fines

quii

Cor

ynor

hinu

s to

wns

endi

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leco

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aurit

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Sco

toph

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borb

onic

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coto

philu

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ridis

Sco

toph

ilus

robu

stus

Pip

istr

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s ku

hlii

Pip

istr

ellu

s ru

stic

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ipis

trel

lus

pipi

stre

llus

Pip

istr

ellu

s te

nuis

Ept

esic

us n

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Ept

esic

us b

rasi

liens

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ptes

icus

bot

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Ept

esic

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rinal

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ptes

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fusc

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ptes

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hot

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Myo

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lbes

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Myo

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ustr

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yotis

gris

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yotis

levi

sM

yotis

luci

fugu

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yotis

oxy

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Myo

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elife

rM

yotis

vol

ans

Myo

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uman

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yotis

viv

esi

Myo

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ipar

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Myo

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dver

sus

Myo

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ocag

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yotis

lani

ger

Myo

tis d

aube

nton

ii

Myo

tis w

elw

itsch

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Myo

tis e

votis

Myo

tis k

eeni

iM

yotis

thys

anod

es

Myo

tis c

alifo

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usM

yotis

leib

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Myo

tis m

ysta

cinu

s

Myo

tis k

eays

i

Myo

tis n

igric

ans

Myo

tis s

odal

is

Talpa davidiana

Talpa stankovici

Talpa romana

Talpa occidentalis

Talpa europaea

Talpa caeca

Talpa levantis

Mogera w

ogura

Mogera tokudae

Mogera im

aizumii

Euroscaptor mizura

Mogera uchidai

Condylura cristata

Parascalops breweri

Scalopus aquaticus

Scapanus latimanus

Scapanus townsendii

Scapanus orariusN

eurotrichus gibbsii

Dym

ecodon pilirostris

Urotrichus talpoides

Desm

ana moschata

Galem

ys pyrenaicus

Echinosorex gymnura

Podogymnura aureospinula

Podogymnura truei

Hylom

ys suillus

Neotetracus sinensis

Neohylom

ys hainanensis

Paraechinus hypomelas

Paraechinus aethiopicus

Paraechinus micropus

Hem

iechinus auritus

Hem

iechinus collaris

Mesechinus dauuricus

Mesechinus hughi

Erinaceus concolor

Erinaceus am

urensis

Erinaceus europaeus

Atelerix frontalis

Atelerix algirus

Atelerix albiventris

Atelerix sclateriC

rocidura cyanea

Crocidura flavescens

Crocidura fuscom

urina

Crocidura hirta

Crocidura leucodon

Crocidura luna

Crocidura m

ariquensis

Crocidura olivieri

Crocidura suaveolens

Diplom

esodon pulchellum

Myosorex cafer

Myosorex longicaudatus

Myosorex varius

Scutisorex som

ereni

Suncus etruscus

Suncus infinitesim

us

Suncus m

urinus

Suncus varilla

Sorex arcticus

Sorex coronatus

Sorex araneus

Sorex unguiculatus

Sorex m

inutus

Sorex cinereus

Sorex hoyi

Sorex fum

eusS

orex longirostris

Sorex ornatus

Sorex bairdi

Sorex palustris

Sorex m

onticolus

Sorex vagrans

Sorex pacificus

Sorex trow

bridgii

Sorex arizonae

Blarina carolinensis

Blarina brevicauda

Cryptotis goodw

ini

Cryptotis m

agna

Cryptotis m

eridensis

Cryptotis nigrescens

Cryptotis parva

Cryptotis thom

asi

Cryptotis gracilis

Cryptotis endersi

Cryptotis hondurensis

Cryptotis m

exicana

Cryptotis goldm

ani

Cryptotis squam

ipes

Cryptotis m

ontivaga

Neom

ys anomalus

Neom

ys fodiens

Neom

ys teres

Chim

arrogale platycephalus

Chim

arrogale himalayica

Notiosorex craw

fordi

Solenodon cubanus

Solenodon paradoxus

Choloepus didactylus

Choloepus hoffmanni

Bradypus torquatus

Bradypus tridactylus

Bradypus variegatus

Tamandua tetradactyla

Tamandua m

exicana

Myrm

ecophaga tridactyla

Cyclopes didactylus

Chlamyphorus truncatus

Euphractus sexcinctus

Zaedyus pichiy

Chaetophractus nationi

Chaetophractus vellerosus

Chaetophractus villosus

Priodontes maxim

us

Tolypeutes matacus

Cabassous centralis

Cabassous chacoensis

Cabassous tatouay

Cabassous unicinctus

Dasypus kappleri

Dasypus novem

cinctus

Chrysospalax trevelyani

Chrysochloris asiatica

Chrysochloris stuhlmanni

Amblysom

us hottentotus

Neamblysom

us gunningi

Neamblysom

us julianae

Limnogale m

ergulus

Geogale aurita

Oryzorictes tetradactylus

Oryzorictes hova

Microgale brevicaudata

Microgale longicaudata

Potamogale velox

Micropotam

ogale lamottei

Micropotam

ogale ruwenzorii

Hemicentetes sem

ispinosus

Hemicentetes nigriceps

Tenrec ecaudatus

Setifer setosus

Echinops telfairi

Elephantulus brachyrhynchus

Elephantulus fuscipes

Elephantulus fuscus

Elephantulus intufi

Elephantulus rupestris

Elephantulus revoili

Elephantulus rufescens

Elephantulus edwardii

Elephantulus myurus

Elephantulus rozeti

Macroscelides proboscideus

Petrodromus tetradactylus

Rhynchocyon chrysopygus

Rhynchocyon cirnei

Rhynchocyon petersi

Orycteropus aferDugong dugon

Trichechus inunguis

Trichechus manatus

Dendrohyrax arboreus

Dendrohyrax dorsalis

Heterohyrax brucei

Procavia capensis

Elephas maxim

us

Loxodonta africana

Loxodonta cyclotis

Acrobates pygmaeus

Distoechurus pennatus

Tarsipes rostratus

Dactylopsila megalura

Dactylopsila palpator

Dactylopsila tatei

Dactylopsila trivirgata

Gymnobelideus leadbeateri

Petaurus abidi

Petaurus australis

Petaurus breviceps

Petaurus gracilis

Petaurus norfolcensis

Hemibelideus lemuroides

Petauroides volans

Petropseudes dahli

Pseudochirops albertisii

Pseudochirops coronatus

Pseudochirops archeri

Pseudochirops cupreus

Pseudochirops corinnae

Pseudochirulus cinereus

Pseudochirulus herbertensis

Pseudocheirus peregrinus

Ailurops ursinus

Phalanger carmelitae

Phalanger vestitus

Phalanger orientalis

Phalanger intercastellanus

Phalanger sericeus

Phalanger lullulae

Strigocuscus pelengensis

Strigocuscus celebensis

Phalanger gymnotis

Spilocuscus maculatus

Spilocuscus kraemeri

Spilocuscus rufoniger

Phalanger matanim

Phalanger ornatusTrichosurus arnhemensis

Trichosurus vulpecula

Trichosurus caninus

Wyulda squamicaudata

Burramys parvus

Cercartetus caudatus

Cercartetus concinnus

Cercartetus lepidus

Cercartetus nanus

Aepyprymnus rufescens

Bettongia gaimardi

Bettongia lesueur

Bettongia penicillata

Bettongia tropicaDendrolagus bennettianus

Dendrolagus dorianus

Dendrolagus scottae

Dendrolagus goodfellowi

Dendrolagus matschiei

Dendrolagus spadix

Dendrolagus ursinus

Dendrolagus inustus

Dendrolagus lumholtzi

Lagorchestes conspicillatus

Lagorchestes hirsutus

Macropus agilis

Macropus eugenii

Macropus dorsalis

Macropus parm

a

Macropus irm

a

Macropus parryi

Macropus rufogriseus

Macropus antilopinus

Macropus rufus

Macropus robustus

Macropus bernardus

Macropus fuliginosus

Macropus giganteus

Wallabia bicolor

Onychogalea fraenata

Onychogalea unguifera

Setonix brachyurus

Petrogale assimilis

Petrogale godmani

Petrogale inornata

Petrogale penicillata

Petrogale lateralis

Petrogale persephone

Petrogale rothschildi

Petrogale xanthopus

Petrogale burbidgei

Petrogale concinna

Thylogale billardierii

Thylogale stigmatica

Thylogale thetis

Dorcopsis atrata

Dorcopsis hageni

Dorcopsis luctuosa

Dorcopsulus macleayi

Dorcopsulus vanheurni

Lagostrophus fasciatus

Potorous longipes

Potorous tridactylus

Potorous gilbertii

Hypsiprymnodon moschatus

Lasiorhinus krefftii

Lasiorhinus latifrons

Vombatus ursinus

Phascolarctos cinereus

Dromiciops gliroides

Antechinus bellus

Antechinus flavipes

Antechinus leo

Antechinus stuartii

Antechinus godmani

Antechinus minimus

Antechinus swainsonii

Murexechinus melanurus

Micromurexia habbema

Phascomurexia naso

Murexia longicaudata

Paramurexia rothschildi

Phascogale calura

Phascogale tapoatafa

Dasyuroides byrnei

Dasycercus cristicauda

Dasyurus albopunctatus

Dasyurus geoffroii

Dasyurus spartacus

Dasyurus viverrinus

Dasyurus maculatus

Dasyurus hallucatus

Sarcophilus harrisii

Neophascogale lorentzi

Phascolosorex doriae

Phascolosorex dorsalis

Pseudantechinus bilarni

Pseudantechinus macdonnellensis

Pseudantechinus woolleyae

Dasykaluta rosamondae

Parantechinus apicalis

Myoictis melas

Pseudantechinus ningbing

Ningaui ridei

Ningaui yvonnae

Ningaui timealeyi

Sminthopsis aitkeni

Sminthopsis griseoventerSminthopsis archeri

Sminthopsis dolichura

Sminthopsis gilberti

Sminthopsis leucopus

Sminthopsis murina

Sminthopsis hirtipesSminthopsis ooldea

Sminthopsis youngsoni

Sminthopsis psammophila

Sminthopsis longicaudata

Sminthopsis granulipes

Sminthopsis butleri

Sminthopsis douglasi

Sminthopsis virginiae

Sminthopsis macroura

Sminthopsis crassicaudata

Antechinomys laniger

Planigale gilesi

Planigale ingrami

Planigale novaeguineae

Planigale tenuirostris

Planigale maculata

Myrmecobius fasciatus

Notoryctes typhlops

Echymipera clara

Echymipera kalubu

Echymipera rufescens

Microperoryctes papuensis

Peroryctes broadbenti

Peroryctes raffrayana

Isoodon auratus

Isoodon obesulus

Isoodon macrourus

Perameles bougainville

Perameles gunnii

Perameles nasuta

Macrotis lagotis

Caenolestes caniventer

Caenolestes convelatus

Caenolestes fuliginosus

Lestoros inca

Rhyncholestes raphanurus

Caluromys derbianus

Caluromys lanatus

Caluromys philander

Caluromysiops irrupta

Glironia venusta

Chironectes minimus

Didelphis albiventrisDidelphis aurita

Didelphis marsupialis

Didelphis virginiana

Philander andersoni

Philander opossum

Lutreolina crassicaudata

Metachirus nudicaudatus

Gracilinanus aceramarcae

Gracilinanus agilis

Gracilinanus microtarsus

Gracilinanus emiliae

Gracilinanus marica

Gracilinanus dryas

Marmosops cracens

Marmosops dorotheaMarmosops fuscatus

Marmosops parvidens

Marmosops handleyi

Marmosops impavidus

Marmosops incanus

Marmosops invictus

Marmosops noctivagus

Lestodelphys halliThylamys elegans

Thylamys pallidior

Thylamys macrurus

Thylamys pusillus

Marmosa andersoni

Tlacuatzin canescens

Marmosa lepida

Marmosa mexicana

Marmosa murina

Marmosa robinsoni

Marmosa rubra

Marmosa tyleriana

Marmosa xerophila

Micoureus alstoni

Micoureus constantiaeMicoureus demerarae

Micoureus paraguayanus

Micoureus regina

Monodelphis adusta

Monodelphis americana

Monodelphis brevicaudata

Monodelphis dimidiata

Monodelphis domestica

Monodelphis emiliae

Monodelphis iheringi

Monodelphis kunsi

Monodelphis maraxina

Monodelphis osgoodi

Monodelphis rubida

Monodelphis scalops

Monodelphis sorex

Monodelphis theresa

Monodelphis unistriata

160 120 80 40 0 Mya

a

Abrocoma bennettii

Ailu

rus

fulg

ens

Ant

iloca

pra

amer

ican

a

Aplodontia rufa

Crase

onyc

teris

thon

glong

yai

Cyclopes didactylus

Daubentonia madagascariensis

Dinomys branickii

Dromiciops gliroides

Dugong dugon

Esc

hric

htiu

s ro

bust

us

Hypsiprymnodon moschatus

Inia

geo

ffren

sis

Macrotis lagotis

Myocastor coypus

Myrmecobius fasciatus

Mystacina tuberculata

Nandinia binotata

Natalus stra

mineus

Notoryctes typhlops

Odo

benu

s ro

smar

us

Ornithorhynchus anatinus

Orycteropus afer

Ped

etes

cap

ensi

s

Petromus typicus

Phascolarctos cinereus

Ptilocercus lowii

Tarsipes rostratus

Acrobates pygmaeus

Distoechurus pennatus

Aepyprymnus rufescens

Bettongia gaimardi

Bettongia lesueur

Bettongia penicillata

Bettongia tropica

Potorous longipes

Potorous tridactylus

Potorous gilbertii

Cuniculus paca

Cuniculus taczanowskii

Urs

us a

rcto

sU

rsus

mar

itim

usU

rsus

thib

etan

usH

elar

ctos

mal

ayan

usM

elur

sus

ursi

nus

Urs

us a

mer

ican

usTr

emar

ctos

orn

atus

Ailu

ropo

da m

elan

oleu

ca

Chrysospalax trevelyani

Chrysochloris asiatica

Chrysochloris stuhlm

anni

Amblysom

us hottentotus

Neam

blysomus gunningi

Neam

blysomus julianae

Ano

mal

urus

bee

crof

ti

Ano

mal

urus

der

bian

us

Ano

mal

urus

pel

ii

Ano

mal

urus

pus

illus

Idiu

rus

mac

rotis

Idiu

rus

zenk

eri

Arc

toce

phal

us a

ustr

alis

Arc

toce

phal

us fo

rste

riA

rcto

ceph

alus

gal

apag

oens

isA

rcto

ceph

alus

gaz

ella

Arc

toce

phal

us p

usill

usN

eoph

oca

cine

rea

Eum

etop

ias

juba

tus

Zal

ophu

s ca

lifor

nian

usC

allo

rhin

us u

rsin

us

Desmodus rotundusMacrophyllum macrophyllum

Lonchorhina aurita

Macrotus californicus

Macrotus waterhousii

Lampronycteris brachyotis

Micronycteris megalotisVampyrum spectrum

Lophostoma silvicolumPhyllostomus discolor

Phyllostomus hastatusBrachyphylla cavernarum

Brachyphylla nana

Phyllonycteris aphyllaLonchophylla mordax

Lonchophylla robusta

Leptonycteris curasoae

Leptonycteris yerbabuenae

Leptonycteris nivalis

Glossophaga commissarisi

Glossophaga soricina

Anoura cultrata

Anoura geoffroyi

Choeronycteris mexicana

Rhinophylla pumilioCarollia castanea

Carollia perspicillataSturnira lilium

Sturnira erythromosCenturio senex

Sphaeronycteris toxophyllumPhyllops falcatus

Ariteus flavescensArtibeus jamaicensis

Artibeus lituratus

Vampyressa pusilla

Vampyressa bidens

Vampyrodes caraccioli

Platyrrhinus helleri

Alouatta belzebulAlouatta caraya

Alouatta coibensisAlouatta palliata

Alouatta guaribaAlouatta pigra

Alouatta saraAlouatta seniculus

Ateles belzebuth

Ateles chamekAteles marginatusAteles fuscicepsAteles geoffroyiAteles paniscusBrachyteles arachnoidesBrachyteles hypoxanthusOreonax flavicaudaLagothrix lagotricha

Addax

nas

omac

ulat

us

Ory

x ga

zella

Ory

x be

isa

Ory

x le

ucor

yx

Hippo

tragu

s eq

uinu

s

Hippo

tragu

s ni

ger

Alcela

phus

bus

elap

hus

Alcela

phus

licht

enst

eini

i

Conno

chae

tes

gnou

Conno

chae

tes

taur

inus

Damal

iscus

luna

tus

Damal

iscus

pyg

argu

s

Amm

otra

gus l

ervia

Budor

cas t

axico

lor

Ovis a

mm

on

Ovis a

ries

Ovis d

alliCap

ra ca

ucas

ica

Capra

falco

neri

Capra

hirc

us

Capra

pyre

naica

Capra

sibir

ica

Hemitr

agus

hyloc

rius

Hemitr

agus

jaya

kari

Hemitr

agus

jem

lahicu

s

Pseud

ois n

ayau

r

Pseud

ois sc

haef

eri

Rupica

pra

pyre

naica

Rupica

pra

rupic

apra

Capric

ornis

crisp

us

Capric

ornis

miln

eedw

ards

ii

Ovibos

mos

chat

us

Oream

nos a

mer

icanu

s

Aepy

cero

s m

elam

pus

Antid

orca

s m

arsu

pial

is

Antil

ope

cerv

icap

ra

Gaz

ella

ben

netti

i

Gaz

ella

sub

guttu

rosa

Gaz

ella

dor

cas

Gaz

ella

gaz

ella

Nange

r dam

a

Nange

r soe

mm

errin

gii

Nange

r gra

nti

Eudo

rcas

thom

soni

i

Lito

cran

ius

walle

ri

Saig

a ta

taric

aCepha

loph

us c

allip

ygus

Cepha

loph

us o

gilb

yi

Cepha

loph

us d

orsa

lis

Cepha

loph

us s

ilvicu

ltor

Cepha

loph

us s

padi

x

Cepha

loph

us je

ntin

ki

Cepha

loph

us n

atal

ensis

Cepha

loph

us n

igrif

rons

Cepha

loph

us ru

filatu

s

Cepha

loph

us le

ucog

aste

r

Cepha

loph

us z

ebra

Sylvi

capr

a gr

imm

ia

Phila

ntom

ba m

axwel

lii

Phila

ntom

ba m

ontic

ola

Dor

catra

gus

meg

alot

is

Rap

hice

rus

cam

pest

ris

Rap

hice

rus

shar

pei

Rap

hice

rus

mel

anot

is

Kobu

s el

lipsi

prym

nus

Kobu

s le

che

Red

unca

aru

ndin

um

Red

unca

redu

nca

Red

unca

fulvo

rufu

la

Pele

a ca

preo

lus

Mad

oqua

gue

nthe

ri

Mad

oqua

kirk

ii

Mad

oqua

pia

cent

inii

Mad

oqua

sal

tiana

Neo

tragu

s ba

tesi

Neo

tragu

s m

osch

atus

Neo

tragu

s py

gmae

us

Ore

otra

gus

oreo

tragu

s

Our

ebia

our

ebi

Proc

apra

gut

turo

sa

Biso

n bi

son

Biso

n bo

nasu

s

Bos

grun

nien

s

Bos

sauv

eli

Bos

front

alis

Bos

java

nicu

s

Buba

lus

buba

lis

Buba

lus

depr

essi

corn

is

Buba

lus

min

dore

nsis

Buba

lus

quar

lesi

Sync

erus

caf

fer

Bose

laph

us tr

agoc

amel

us

Tetra

ceru

s qu

adric

orni

s

Taur

otra

gus

oryx

Trag

elap

hus

stre

psic

eros

Trag

elap

hus

anga

sii

Trag

elap

hus

eury

ceru

s

Trag

elap

hus

spek

ii

Trag

elap

hus

scrip

tus

Trag

elap

hus

imbe

rbis

Bradypus torquatus

Bradypus tridactylus

Bradypus variegatus

Caenolestes caniventer

Caenolestes convelatus

Caenolestes fuliginosus

Lestoros inca

Rhyncholestes raphanurus

Callimico goeldii

Callithrix argentataCallithrix emiliaeCallithrix intermedia

Callithrix leucippeCallithrix humeraliferaCallithrix chrysoleuca

Callithrix pygmaea

Callithrix auritaCallithrix flaviceps

Callithrix geoffroyiCallithrix kuhliiCallithrix jacchus

Callithrix penicillataLeontopithecus caissara

Leontopithecus chrysopygusLeontopithecus rosalia

Leontopithecus chrysomelas

Saguinus bicolorSaguinus midas

Saguinus geoffroyiSaguinus oedipusSaguinus leucopusSaguinus imperator

Saguinus labiatusSaguinus mystax

Saguinus fuscicollisSaguinus tripartitusSaguinus nigricollisSaguinus inustusCebus albifronsCebus capucinusCebus olivaceusCebus apellaCebus xanthosternos

Saimiri boliviensisSaimiri vanzolinii

Saimiri oerstediiSaimiri sciureusSaimiri ustus

Caluromys derbianus

Caluromys lanatus

Caluromys philander

Caluromysiops irrupta

Glironia venusta

Chironectes minimusDidelphis albiventris

Didelphis aurita

Didelphis marsupialis

Didelphis virginiana

Philander andersoni

Philander opossum

Lutreolina crassicaudata

Metachirus nudicaudatus

Gracilinanus aceramarcae

Gracilinanus agilis

Gracilinanus microtarsus

Gracilinanus emiliae

Gracilinanus marica

Gracilinanus dryas

Marmosops cracens

Marmosops dorotheaMarmosops fuscatus

Marmosops parvidens

Marmosops handleyi

Marmosops impavidus

Marmosops incanus

Marmosops invictus

Marmosops noctivagus

Lestodelphys halliThylamys elegans

Thylamys pallidior

Thylamys macrurus

Thylamys pusillus

Marmosa andersoni

Tlacuatzin canescens

Marmosa lepida

Marmosa mexicana

Marmosa murina

Marmosa robinsoni

Marmosa rubra

Marmosa tyleriana

Marmosa xerophila

Micoureus alstoni

Micoureus constantiaeMicoureus demerarae

Micoureus paraguayanus

Micoureus regina

Monodelphis adusta

Monodelphis americana

Monodelphis brevicaudata

Monodelphis dimidiata

Monodelphis domestica

Monodelphis emiliae

Monodelphis iheringi

Monodelphis kunsi

Monodelphis maraxina

Monodelphis osgoodi

Monodelphis rubida

Monodelphis scalops

Monodelphis sorex

Monodelphis theresa

Monodelphis unistriata

Can

is lu

pus

Can

is la

tran

sC

anis

sim

ensi

sC

anis

adu

stus

Can

is a

ureu

sLy

calo

pex

culp

aeus

Lyca

lope

x gr

iseu

sLy

calo

pex

gym

noce

rcus

Lyca

lope

x se

chur

aeLy

calo

pex

vetu

lus

Ate

locy

nus

mic

rotis

Cer

docy

on th

ous

Chr

ysoc

yon

brac

hyur

usC

uon

alpi

nus

Lyca

on p

ictu

sS

peot

hos

vena

ticus

Nyc

tere

utes

pro

cyon

oide

s

Vul

pes

cors

ac

Vul

pes

ruep

pelli

iV

ulpe

s vu

lpes

Vulpes lagopus

Vul

pes

velo

x Vulpes macrotis

Vulpes cana

Vulpes zerda

Vulpes bengalensis

Vulpes cham

aV

ulpes pallidaU

rocyon cinereoargenteusU

rocyon littoralisO

tocyon megalotis

Capromys pilorides

Geocaprom

ys brownii

Geocaprom

ys ingrahami

Mesocaprom

ys melanurus

Plagiodontia aedium

Castor canadensis

Castor fiber

Hydrochoerus hydrochaeris

Kerodon rupestris

Cavia porcellus

Cavia aperea

Microcavia australis

Galea musteloides

Galea spixii

Galea flavidens

Dolichotis patagonum

Dolichotis salinicola

Cer

atot

heriu

m s

imum

Dic

eros

bic

orni

s

Dic

eror

hinu

s su

mat

rens

is

Rhi

noce

ros

unic

orni

s

Burramys parvus

Cercartetus caudatus

Cercartetus concinnus

Cercartetus lepidus

Cercartetus nanus

Alc

es a

lces

Bla

stoc

erus

dic

hoto

mus

Hip

poca

mel

us a

ntis

ensi

s

Hip

poca

mel

us b

isul

cus

Ozo

toce

ros

bezo

artic

us

Maz

ama

amer

ican

a

Maz

ama

nana

Odo

coile

us h

emio

nus

Odo

coile

us v

irgin

ianu

sP

udu

puda

Ran

gife

r tar

andu

s

Axi

s ax

is

Axi

s ca

lam

iane

nsis

Axi

s ku

hlii

Axi

s po

rcin

us

Ruc

ervu

s du

vauc

elii

Cer

vus

elap

hus

Ruc

ervu

s el

dii

Cer

vus

nipp

on

Rus

a tim

oren

sis

Rus

a un

icol

orM

untia

cus

athe

rode

s

Mun

tiacu

s m

untja

k

Lagostomus m

aximus

Chinchilla lanigera

Chinchilla chinchilla

Lagidium peruanum

Lagidium viscacia

Lagidium wolffsohni

Choloepus didactylus

Choloepus hoffmanni

Abrothrix longipilisAbrothrix sanborni

Akodon affinisAkodon albiventerAkodon azaraeAkodon cursorAkodon iniscatusAbrothrix olivaceusNecromys urichiAkodon boliviensis

Akodon dayiAkodon molinaeAkodon varius

Thaptomys nigritaAbrothrix andinusAbrothrix jelskii

Auliscomys boliviensisLoxodontomys micropusAuliscomys pictusAuliscomys sublimis

Phyllotis caprinusPhyllotis darwiniPhyllotis magisterPhyllotis osgoodiPhyllotis osilaePhyllotis xanthopygus

Graomys griseoflavus

Andinomys edax

Kunsia frontoKunsia tomentosusScapteromys tumidus

Necromys lasiurusNecromys obscurus

Calomys hummelinckiChelemys macronyxChinchillula sahamae

Delomys dorsalisDelomys sublineatus

Eligmodontia typusGeoxus valdivianus

Holochilus brasiliensisHolochilus chacariusLundomys molitorHolochilus sciureus

Irenomys tarsalisMelanomys caliginosus

Neacomys guianaeNeacomys pictusNeacomys spinosusNeacomys tenuipes

Nectomys palmipesNectomys rattusNectomys squamipes

Neotomys ebriosusOecomys bicolor

Oligoryzomys chacoensisOligoryzomys delticolaOligoryzomys destructorOligoryzomys flavescensOligoryzomys fulvescensOligoryzomys microtisOligoryzomys nigripes

Oryzomys alfaroiOryzomys bolivarisOryzomys angouyaOryzomys megacephalusOryzomys chapmaniOryzomys couesiOryzomys russatusOryzomys macconnelliOryzomys melanotisOryzomys nitidusOryzomys palustrisOryzomys rostratusOryzomys subflavusOryzomys talamancae

Oxymycterus nasutusOxymycterus rufus

Punomys lemminusReithrodon auritus

Rhipidomys austrinusRhipidomys caucensisRhipidomys couesiRhipidomys fulviventerRhipidomys latimanusRhipidomys leucodactylusRhipidomys macconnelliRhipidomys mastacalisRhipidomys nitelaRhipidomys ochrogasterRhipidomys venezuelaeRhipidomys venustusRhipidomys wetzeli

Scolomys melanopsScolomys ucayalensis

Sigmodontomys alfari

Wiedomys pyrrhorhinos

Wilfredomys oenaxZygodontomys brevicauda

Ichthyomys pittieriIchthyomys stolzmanni

Ichthyomys tweediiNeusticomys monticolus

Rheomys raptorSigmodon alstoniSigmodon arizonae

Sigmodon hispidusSigmodon toltecusSigmodon ochrognathus

Nyctomys sumichrasti

Otonyctomys hatti

Ototylomys phyllotis

Tylomys miraeTylomys nudicaudus

Tylomys tumbalensis

Tylomys watsoni

Baiomys musculus

Baiomys taylori

Scotinomys teguina

Ochrotomys nuttalli

Habromys lophurus

Neotomodon alstoni

Podomys floridanus

Osgoodomys banderanus

Peromyscus attwateri

Peromyscus aztecus

Peromyscus boylii

Peromyscus bullatus

Peromyscus californicus

Peromyscus caniceps

Peromyscus crinitus

Peromyscus dickeyi

Peromyscus difficilis

Peromyscus eremicus

Peromyscus eva

Peromyscus furvus

Peromyscus gossypinus

Peromyscus grandis

Peromyscus gratus

Peromyscus guardia

Peromyscus guatemalensis

Peromyscus gymnotis

Peromyscus hooperi

Peromyscus interparietalis

Peromyscus leucopus

Peromyscus levipes

Peromyscus madrensis

Peromyscus maniculatus

Peromyscus mayensis

Peromyscus megalops

Peromyscus mekisturus

Peromyscus melanocarpus

Peromyscus melanophrys

Peromyscus melanotis

Peromyscus melanurus

Peromyscus merriami

Peromyscus mexicanus

Peromyscus nasutus

Peromyscus ochraventer

Peromyscus keeni

Peromyscus pectoralis

Peromyscus perfulvus

Peromyscus polionotus

Peromyscus polius

Peromyscus pseudocrinitus

Peromyscus sejugis

Peromyscus simulus

Peromyscus slevini

Peromyscus spicilegus

Peromyscus stephani

Peromyscus stirtoni

Peromyscus truei

Peromyscus winkelmanni

Peromyscus yucatanicus

Peromyscus zarhynchusOnychomys leucogaster

Onychomys torridus

Isthmomys flavidus

Isthmomys pirrensis

Reithrodontomys brevirostris

Reithrodontomys darienensis

Reithrodontomys gracilis

Reithrodontomys mexicanus

Reithrodontomys paradoxus

Reithrodontomys spectabilis

Reithrodontomys creper

Reithrodontomys microdon

Reithrodontomys rodriguezi

Reithrodontomys tenuirostrisReithrodontomys fulve

scens

Reithrodontomys hirsutus

Reithrodontomys burti

Reithrodontomys chrysopsis

Reithrodontomys humulis

Reithrodontomys megalotis

Reithrodontomys montanus

Reithrodontomys raviventris

Reithrodontomys sumichrasti

Reithrodontomys zacatecae

Xenomys nelsoni

Neotoma albigula

Neotoma bryanti

Neotoma devia

Neotoma floridana

Neotoma magisterNeotoma lepida

Neotoma martinensis

Neotoma mexicana

Neotoma micropus

Neotoma phenax

Neotoma cinerea

Neotoma fuscipes

Cricetus c

ricetus

Cricetulus m

igratoriu

s

Mesocri

cetus a

uratus

Mesocri

cetus n

ewtoni

Phodopus campbelli

Phodopus roborov

skii

Phodopus sungoru

s

Alticola argentatus

Alticola st

oliczka

nus

Alticola st

relzowi

Myodes c

alifornicu

s

Myodes g

apperi

Myodes g

lareolus

Myodes r

ufocanus

Myodes r

utilus

Dinaromys bogdanovi

Arborimus a

lbipes

Arborimus l

ongicaudus

Arborimus p

omo

Phenacomys

interm

edius

Phenacomys

ungavaArvico

la sapidus

Arvico

la amphibius

Chionomys niva

lis

Dicrosto

nyx groenlandicu

s

Dicrosto

nyx ric

hardsoni

Prometheomys sc

haposchniko

wi

Lagurus l

agurus

Lasiopodomys

brandtii

Lasiopodomys

fusc

us

Lasiopodomys

mandarin

us

Lemmiscus c

urtatus

Lemmus lemmus

Myopus s

chist

icolor

Synaptomys

borealis

Synaptomys

cooperi

Microtus a

grestis

Microtus c

hrotorrhinus

Microtus r

ichardso

ni

Microtus x

anthognathus

Microtus g

uatemalensis

Microtus lo

ngicaudus

Microtus m

exicanus

Microtus a

rvalis

Microtus californicus

Microtus montanus

Microtus breweri

Microtus pennsylvanicus

Microtus townsendii

Microtus o

economus

Microtus o

chrogaste

r

Microtus p

inetorum

Microtus a

bbreviatus

Microtus m

iurus

Microtus multip

lex

Microtus duodecimcostatus

Microtus s

avii

Microtus bavaricus

Microtus subterraneus

Neofiber a

lleni

Ondatra zi

bethicusLo

phiom

ys im

haus

i

Crocuta crocuta

Hyaena hyaena

Hyaena brunnea

Proteles cristata

Massoutiera mzabi

Pectinator spekei

Ctenodactylus gundi

Ctenodactylus valiFelovia vae

Ctenomys haigi

Ctenomys opim

us

Ctenomys torquatus

Ctenomys talarum

Galeopterus variegates

Cynocephalus volans

Myoprocta acouchy

Dasyprocta punctata

Dasyprocta leporina

Dasyprocta azarae

Dasyprocta coibae

Dasyprocta cristata

Dasyprocta fuliginosa

Dasyprocta guamara

Dasyprocta kalinowskii

Dasyprocta mexicana

Dasyprocta prymnolopha

Dasyprocta ruatanica

Chlamyphorus truncatus

Euphractus sexcinctus

Zaedyus pichiy

Chaetophractus nationi

Chaetophractus vellerosus

Chaetophractus villosus

Priodontes maxim

us

Tolypeutes matacus

Cabassous centralis

Cabassous chacoensis

Cabassous tatouay

Cabassous unicinctus

Dasypus kappleri

Dasypus novemcinctus

Antechinus bellus

Antechinus flavipes

Antechinus leo

Antechinus stuartii

Antechinus godmani

Antechinus minimus

Antechinus swainsonii

Murexechinus melanurus

Micromurexia habbema

Phascomurexia naso

Murexia longicaudata

Paramurexia rothschildi

Phascogale calura

Phascogale tapoatafa

Dasyuroides byrnei

Dasycercus cristicauda

Dasyurus albopunctatus

Dasyurus geoffroii

Dasyurus spartacus

Dasyurus viverrinus

Dasyurus maculatus

Dasyurus hallucatus

Sarcophilus harrisii

Neophascogale lorentzi

Phascolosorex doriae

Phascolosorex dorsalis

Pseudantechinus bilarni

Pseudantechinus macdonnellensis

Pseudantechinus woolleyae

Dasykaluta rosamondae

Parantechinus apicalis

Myoictis melas

Pseudantechinus ningbing

Ningaui ridei

Ningaui yvonnae

Ningaui timealeyi

Sminthopsis aitkeni

Sminthopsis griseoventerSminthopsis archeri

Sminthopsis dolichura

Sminthopsis gilberti

Sminthopsis leucopus

Sminthopsis murina

Sminthopsis hirtipesSminthopsis ooldea

Sminthopsis youngsoni

Sminthopsis psammophila

Sminthopsis longicaudata

Sminthopsis granulipes

Sminthopsis butleri

Sminthopsis douglasi

Sminthopsis virginiae

Sminthopsis macroura

Sminthopsis crassicaudata

Antechinomys laniger

Planigale gilesi

Planigale ingrami

Planigale novaeguineae

Planigale tenuirostris

Planigale maculata

Per

ogna

thus

alti

colu

sP

erog

nath

us a

mpl

usP

erog

nath

us fa

scia

tus

Per

ogna

thus

flav

esce

nsP

erog

nath

us fl

avus

Per

ogna

thus

inor

natu

sP

erog

nath

us lo

ngim

embr

isP

erog

nath

us m

erria

mi

Per

ogna

thus

par

vus

Cha

etod

ipus

form

osus

Cha

etod

ipus

pen

icill

atus

Cha

etod

ipus

inte

rmed

ius

Cha

etod

ipus

nel

soni

Cha

etod

ipus

line

atus

Cha

etod

ipus

spi

natu

s

Cha

etod

ipus

falla

xC

haet

odip

us c

alifo

rnic

us

Cha

etod

ipus

bai

leyi

Cha

etod

ipus

his

pidu

s

Dip

odom

ys o

rdii

Dip

odom

ys g

ravi

pes

Dip

odom

ys in

gens

Dip

odom

ys c

ompa

ctus

Dip

odom

ys p

anam

intin

usD

ipod

omys

hee

rman

ni

Dip

odom

ys s

teph

ensi

Dip

odom

ys v

enus

tus

Dip

odom

ys a

gilis

Dip

odom

ys s

imul

ans

Dip

odom

ys m

icro

ps

Dip

odom

ys e

lato

r

Dip

odom

ys p

hilli

psii

Dip

odom

ys c

alifo

rnic

us

Dip

odom

ys m

erria

mi

Dip

odom

ys n

itrat

oide

s

Dip

odom

ys d

eser

tiD

ipod

omys

spe

ctab

ilis

Dip

odom

ys n

elso

ni

Mic

rodi

podo

ps m

egac

epha

lus

Mic

rodi

podo

ps p

allid

us

Het

erom

ys d

esm

ares

tianu

sH

eter

omys

ore

ster

usH

eter

omys

nel

soni

Liom

ys a

dspe

rsus

Liom

ys s

alvi

niLi

omys

pic

tus

Liom

ys ir

rora

tus

Het

erom

ys a

ustr

alis

Het

erom

ys a

nom

alus

Het

erom

ys g

aum

eri

Limnogale m

ergulus

Geogale aurita

Oryzorictes tetradactylus

Oryzorictes hova

Microgale brevicaudata

Microgale longicaudata

Potamogale velox

Micropotam

ogale lamottei

Micropotam

ogale ruwenzorii

Hem

icentetes semispinosus

Hem

icentetes nigriceps

Tenrec ecaudatus

Setifer setosus

Echinops telfairi

Echymipera clara

Echymipera kalubu

Echymipera rufescens

Microperoryctes papuensis

Peroryctes broadbenti

Peroryctes raffrayana

Isoodon auratus

Isoodon obesulus

Isoodon macrourus

Perameles bougainville

Perameles gunnii

Perameles nasuta

Elephantulus brachyrhynchus

Elephantulus fuscipes

Elephantulus fuscus

Elephantulus intufi

Elephantulus rupestris

Elephantulus revoili

Elephantulus rufescens

Elephantulus edwardii

Elephantulus myurus

Elephantulus rozeti

Macroscelides proboscideus

Petrodromus tetradactylus

Rhynchocyon chrysopygus

Rhynchocyon cirnei

Rhynchocyon petersiElephas m

aximus

Loxodonta africana

Loxodonta cyclotis

Saccolaimus peli

Saccolaimus flaviventris

Saccolaimus mixtus

Saccolaimus saccolaimusTaphozous hamiltoni

Taphozous nudiventris

Taphozous theobaldiTaphozous hilliTaphozous australis

Taphozous georgianus

Taphozous kapalgensis

Taphozous hildegardeae

Taphozous longimanus

Taphozous mauritianus

Taphozous perforatus

Taphozous melanopogonMosia nigrescens

Emballonura atrata

Emballonura beccarii

Emballonura dianae

Emballonura furax

Emballonura raffrayana

Emballonura semicaudata

Coleura afra

Coleura seychellensis

Emballonura alecto

Emballonura monticolaRhynchonycteris nasoCyttarops alecto

Diclidurus isabellus

Diclidurus scutatusDiclidurus albus

Diclidurus ingensCentronycteris maximiliani

Cormura brevirostrisSaccopteryx bilineata

Saccopteryx leptura

Saccopteryx gymnura

Saccopteryx canescens

Balantiopteryx io

Balantiopteryx infusca

Balantiopteryx plicata

Peropteryx leucopteraPeropteryx kappleri

Peropteryx macrotis

Equ

us a

sinu

sE

quus

zeb

ra

Equ

us h

emio

nus

Equ

us c

abal

lus

Chaetomys subspinosus

Erethizon dorsatum

Sphiggurus insidiosus

Sphiggurus villosus

Sphiggurus mexicanus

Sphiggurus spinosus

Sphiggurus vestitus

Coendou bicolor

Coendou prehensilis

Coendou rothschildi

Echinosorex gymnura

Podogymnura aureospinula

Podogymnura truei

Hylom

ys suillus

Neotetracus sinensis

Neohylom

ys hainanensis

Paraechinus hypomelas

Paraechinus aethiopicus

Paraechinus micropus

Hem

iechinus auritus

Hem

iechinus collaris

Mesechinus dauuricus

Mesechinus hughi

Erinaceus concolor

Erinaceus amurensis

Erinaceus europaeus

Atelerix frontalis

Atelerix algirus

Atelerix albiventris

Atelerix sclateri

Panthera leo

Panthera pardus

Panthera onca

Panthera tigris

Uncia uncia

Neofelis nebulosa

Pardofelis m

armorata

Lynx canadensisLynx lynxLynx pardinusLynx rufusC

atopuma tem

minckii

Profelis aurata

Leopardus tigrinusLeopardus geoffroyiLeopardus guignaLeopardus colocoloLeopardus jacobitus

Leopardus pardalisLeopardus w

iediiFelis m

argaritaFelis nigripesFelis silvestrisFelis bietiFelis chausFelis m

anulC

aracal caracalLeptailurus servalP

rionailurus bengalensis

Prionailurus iriom

otensis

Prionailurus viverrinus

Prionailurus rubiginosus

Prionailurus planiceps

Pum

a yagouaroundi

Pum

a concolorA

cinonyx jubatus

Galidictis fasciata

Galidictis grandidieri

Galidia elegans

Mungotictis decem

lineataS

alanoia concolorE

upleres goudotiiFossa fossanaC

ryptoprocta ferox

Genetta abyssinica

Genetta angolensis

Genetta m

aculataG

enetta genettaG

enetta tigrinaG

enetta servalinaG

enetta cristataP

rionodon linsangP

rionodon pardicolorP

oiana richardsoniiG

enetta piscivoraV

iverra civettinaV

iverra megaspila

Viverra tangalunga

Viverra zibetha

Viverricula indica

Civettictis civetta

Paradoxurus jerdoni

Paradoxurus zeylonensis

Paradoxurus herm

aphroditus

Arctictis binturong

Pagum

a larvataM

acrogalidia musschenbroekii

Arctogalidia trivirgata

Chrotogale ow

stoniD

iplogale hoseiH

emigalus derbyanus

Cynogale bennettii

Thom

omys bottae

Thom

omys um

brinusT

homom

ys talpoides

Geom

ys personatus

Geom

ys bursariusG

eomys attw

ateriG

eomys breviceps

Geom

ys arenarius

Geom

ys pinetis

Cratogeom

ys neglectus Ort

hoge

omys

het

erod

usO

rtho

geom

ys h

ispi

dus

Ort

hoge

omys

che

rrie

iO

rtho

geom

ys u

nder

woo

diO

rtho

geom

ys d

arie

nsis

Ort

hoge

omys

cav

ator

Ort

hoge

omys

cun

icul

usOrthogeom

ys laniusO

rthogeomys thaeleri

Dryom

ys laniger

Dryom

ys nitedula

Chaetocauda sichuanensis

Eliomys m

elanurus

Eliomys quercinus

Selevinia betpakdalaensis

Muscardinus avellanarius

Glis glis

Glirulus japonicus

Graphiurus christyi

Graphiurus kelleni

Graphiurus lorraineus

Graphiurus m

urinus

Graphiurus ocularis

Graphiurus platyops

Graphiurus surdus

Graphiurus crassicaudatus

Graphiurus nagtglasii

Heterocephalus glaber

Heliophobius argenteocinereus

Cryptomys damarensis

Cryptomys hottentotus

Cryptomys mechowi

Georychus capensis

Bathyergus suillus

Dendrohyrax arboreus

Dendrohyrax dorsalis

Heterohyrax brucei

Procavia capensis

Hex

apro

todo

n lib

erie

nsis

Hip

popo

tam

us a

mph

ibiu

s

Rhinon

icter

is au

rant

ia

Triae

nops

furcu

lus

Triae

nops

rufu

sHipp

oside

ros a

ter

Hippos

idero

s caff

er

Hippos

idero

s spe

oris

Hippos

idero

s cer

vinus

Cloeoti

s per

civali

Hippos

idero

s com

merso

ni

Gorilla gorilla

Homo sapiens

Pan paniscusPan troglodytes

Pongo pygmaeusPongo abelii

Trinomys dim

idiatus

Hoplom

ys gymnurus

Proechimys quadruplicatus

Proechimys sim

onsi

Euryzygomatom

ys spinosus

Lonchothrix emiliae

Proechimys longicaudatus

Proechimys trinitatus

Carterodon sulcidens

Proechimys brevicauda

Proechimys canicollis

Proechimys guyannensis

Proechimys cuvieri

Proechimys sem

ispinosus

Proechimys guairae

Proechimys hoplom

yoides

Proechimys m

agdalenae

Proechimys oconnelli

Proechimys poliopus

Proechimys steerei

Makalata didelphoides

Echimys chrysurus

Phyllomys blainvillii

Phyllomys brasiliensis

Phyllomys dasythrix

Makalata grandis

Phyllomys lam

arum

Phyllomys nigrispinus

Echimys sem

ivillosus

Phyllomys thom

asi

Isothrix bistriata

Isothrix pagurus

Diplom

ys caniceps

Diplom

ys labilis

Diplom

ys rufodorsalis

Kannabateomys am

blyonyx

Dactylom

ys dactylinus

Dactylom

ys boliviensis

Olallam

ys edax

Mesom

ys hispidus

Mesom

ys leniceps

Makalata obscura

Mesom

ys stimulax

Trichys fasciculata

Hystrix africaeaustralis

Hystrix cristata

Hystrix brachyura

Hystrix indica

Atherurus macrourus

Atherurus africanus

Par

adip

us c

teno

dact

ylus

Sic

ista

bet

ulin

a

Sic

ista

sub

tilis

Dip

us s

agitt

a

Jacu

lus

jacu

lus

Jacu

lus

blan

ford

i

Jacu

lus

orie

ntal

is

Sty

lodi

pus

andr

ewsi

Sty

lodi

pus

sung

orus

Sty

lodi

pus

telu

m

Nap

aeoz

apus

insi

gnis

Zap

us h

udso

nius

Zap

us p

rince

ps

Zap

us tr

inot

atus

Car

dioc

rani

us p

arad

oxus

Sal

ping

otul

us m

icha

elis

Pyg

eret

mus

pum

ilio

Pyg

eret

mus

pla

tyur

us

Pyg

eret

mus

shi

tkov

i

Cam

elus

dro

med

ariu

s

Lam

a gl

ama

Vic

ugna

vic

ugna

Eulemur coronatus

Eulemur fulvusEulemur albocollaris

Eulemur collarisEulemur rufus

Eulemur sanfordiEulemur macacoEulemur mongoz

Eulemur rubriventer

Hapalemur aureus

Hapalemur griseusHapalemur alaotrensis

Hapalemur occidentalis

Prolemur simus

Lemur cattaVarecia variegata

Varecia rubra

Nesolagus netscheri

Sylvilagus aquaticus

Sylvilagus brasiliensis

Sylvilagus palustris

Sylvilagus audubonii

Sylvilagus bachmani

Caprolagus hispidus

Poelagus marjorita

Lepus alleni

Lepus callotis

Lepus flavigularis

Lepus californicus

Lepus americanus

Lepus arcticus

Lepus timidus

Lepus townsendii

Lepus brachyurus

Lepus europaeus

Lepus comus

Lepus capensis

Lepus saxatilis

Lepus microtis

Lepus hainanus

Lepus nigricollis

Oryctolagus cuniculusBunolagus monticularis

Romerolagus diazi

Pentalagus furnessi

Pronolagus crassicaudatus

Pronolagus rupestris

Pronolagus randensis

Allenopithecus nigroviridis

Cercopithecus ascanius

Cercopithecus cephus

Cercopithecus erythrogaster

Cercopithecus erythrotis

Cercopithecus petaurista

Cercopithecus sclateri

Cercopithecus mitis

Cercopithecus albogularis

Cercopithecus nictitansCercopithecus campbelli

Cercopithecus mona

Cercopithecus pogonias

Cercopithecus wolfi

Cercopithecus dentiCercopithecus diana

Cercopithecus dryas

Cercopithecus neglectus

Cercopithecus hamlyniCercopithecus lhoesti

Cercopithecus preussi

Cercopithecus solatus

Chlorocebus aethiops

Chlorocebus pygerythrus

Chlorocebus sabaeus

Chlorocebus tantalusErythrocebus patas

Miopithecus talapoin

Cercocebus agilis

Cercocebus galeritus

Cercocebus torquatus

Cercocebus atys

Mandrillus leucophaeus

Mandrillus sphinx

Lophocebus albigena

Lophocebus aterrimus

Papio hamadryas

Papio anubis

Papio cynocephalusPapio papio

Papio ursinus

Theropithecus gelada

Macaca arctoides

Macaca assamensis

Macaca thibetana

Macaca radiata

Macaca sinica

Macaca cyclopis

Macaca mulatta

Macaca fuscata

Macaca fascicularis

Macaca maura

Macaca nigra

Macaca nigrescens

Macaca ochreata

Macaca tonkeana

Macaca hecki

Macaca nemestrina

Macaca leonina

Macaca pagensis

Macaca silenus

Macaca sylvanus

Colobus angolensis

Colobus guereza

Colobus polykomosColobus vellerosusColobus satanas

Piliocolobus badius

Piliocolobus preussiPiliocolobus kirkii

Piliocolobus pennantii

Piliocolobus rufomitratusProcolobus verus

Simias concolorNasalis larvatus

Presbytis comataPresbytis hosei

Presbytis thomasi

Presbytis femoralis

Presbytis siamensis

Presbytis melalophos

Presbytis frontata

Presbytis potenziani

Presbytis rubicunda

Semnopithecus entellus

Trachypithecus auratus

Trachypithecus cristatus

Trachypithecus francoisi

Trachypithecus delacouri

Trachypithecus laotum

Trachypithecus poliocephalus

Trachypithecus geei

Trachypithecus pileatusTrachypithecus obscurus

Trachypithecus phayrei

Trachypithecus johnii

Trachypithecus vetulusRhinopithecus avunculus

Rhinopithecus bieti

Rhinopithecus brelichi

Rhinopithecus roxellanaPygathrix nemaeus

Dendrolagus bennettianus

Dendrolagus dorianus

Dendrolagus scottae

Dendrolagus goodfellowi

Dendrolagus matschiei

Dendrolagus spadix

Dendrolagus ursinus

Dendrolagus inustus

Dendrolagus lumholtzi

Lagorchestes conspicillatus

Lagorchestes hirsutus

Macropus agilis

Macropus eugenii

Macropus dorsalis

Macropus parm

a

Macropus irm

a

Macropus parryi

Macropus rufogriseus

Macropus antilopinus

Macropus rufus

Macropus robustus

Macropus bernardus

Macropus fuliginosus

Macropus giganteus

Wallabia bicolor

Onychogalea fraenata

Onychogalea unguifera

Setonix brachyurus

Petrogale assimilis

Petrogale godmani

Petrogale inornata

Petrogale penicillata

Petrogale lateralis

Petrogale persephone

Petrogale rothschildi

Petrogale xanthopus

Petrogale burbidgei

Petrogale concinna

Thylogale billardierii

Thylogale stigmatica

Thylogale thetis

Dorcopsis atrata

Dorcopsis hageni

Dorcopsis luctuosa

Dorcopsulus macleayi

Dorcopsulus vanheurni

Lagostrophus fasciatus

Manis crassicaudata

Manis pentadactyla

Manis tem

minckii

Cardio

derm

a co

r

Lavia

fron

s

Mac

rode

rma

gigas

Mus

tela

lutre

ola

Mus

tela

lutr

eolin

a

Mus

tela

nud

ipes

Mus

tela

sib

irica

Mus

tela

eve

rsm

anii

Mus

tela

nig

ripes

Mus

tela

put

oriu

sM

uste

la a

ltaic

aM

uste

la e

rmin

eaM

uste

la fr

enat

aM

uste

la n

ival

isM

uste

la a

fric

ana

Neo

viso

n m

acro

don

Neo

viso

n vi

son

Mar

tes

amer

ican

a

Mar

tes

mel

ampu

sM

arte

s zi

belli

naM

arte

s m

arte

sM

arte

s fo

ina

Mar

tes

flavi

gula

Mar

tes

gwat

kins

iiM

arte

s pe

nnan

tiG

ulo

gulo

Eira

bar

bara

Gal

ictis

cuj

aG

alic

tis v

ittat

a

Lync

odon

pat

agon

icus

Icto

nyx

libyc

aIc

tony

x st

riatu

sVo

rmel

a pe

regu

sna

Poe

cilo

gale

alb

inuc

haM

elliv

ora

cape

nsis

Arc

tony

x co

llaris

Mel

es m

eles

Mel

es a

naku

ma

Mel

ogal

e ev

eret

tiM

elog

ale

mos

chat

aM

elog

ale

orie

ntal

isM

elog

ale

pers

onat

a

Lont

ra fe

lina

Lont

ra p

rovo

cax

Lont

ra lo

ngic

audi

sLo

ntra

can

aden

sis

Lutr

a lu

tra

Aon

yx c

apen

sis

Aon

yx c

iner

ea

Hyd

rictis

mac

ulic

ollis

Lutr

ogal

e pe

rspi

cilla

taP

tero

nura

bra

silie

nsis

Enh

ydra

lutr

is

Taxi

dea

taxu

s

Myd

aus

java

nens

isM

ydau

s m

arch

ei

Con

epat

us c

hing

aC

onep

atus

leuc

onot

usC

onep

atus

hum

bold

tiiC

onep

atus

sem

istr

iatu

s

Mep

hitis

mac

rour

aM

ephi

tis m

ephi

tis

Spi

loga

le p

utor

ius

Spi

loga

le g

raci

lisS

pilo

gale

pyg

mae

a

Allocebus trichotisMirza coquereli

Microcebus murinusMicrocebus griseorufusMicrocebus myoxinusMicrocebus rufus

Cheirogaleus majorCheirogaleus crossleyi

Cheirogaleus mediusPhaner furcifer

Miniopterus schreibersii

Miniopterus australis

Lasionycteris noctivagans

Nycticeius humeralis

Scoteanax rueppellii

Otonycteris hemprichii

Antrozous pallid

us

Vespertilio murin

us

Neoromicia nanus

Histiotus montanus

Scotorepens greyii

Scotorepens sanborni

Hypsugo bodenheimeri

Vespadelus vulturnus

Nyctalus azoreum

Chalinolobus tuberculatusChalinolobus dwyeri

Chalinolobus gouldii

Chalinolobus morio

Chalinolobus nigrogriseus

Lasiurus egregiusLasiurus ega

Lasiurus intermedius

Lasiurus cinereus

Lasiurus borealis

Lasiurus seminolus

Nyctophilus arnhemensis

Nyctophilus geoffroyi

Nyctophilus gouldi

Nyctophilus timoriensis

Barbastella barbastellus

Euderma maculatum

Idionycteris phyllotis

Corynorhinus rafinesquii

Corynorhinus townsendii

Plecotus auritus

Scotophilus borbonicus

Scotophilus viridis

Scotophilus robustus

Pipistrellus kuhlii

Pipistrellus rusticus

Pipistrellus pipistrellus

Pipistrellus tenuis

Eptesicus nilssoniiEptesicus brasiliensis

Eptesicus bottae

Eptesicus furinalis

Eptesicus fuscus

Eptesicus hottentotus

Myotis albescens

Myotis austroriparius

Myotis grisescens

Myotis levis

Myotis lucifugus

Myotis oxyotus

Myotis velifer

Myotis volans

Myotis yumanensis

Myotis vivesi

Myotis riparius

Myotis adversus

Myotis bocagii

Myotis laniger

Myotis daubentoniiMyotis welwitschii

Myotis evotis

Myotis keenii

Myotis thysanodesMyotis californicus

Myotis leibii

Myotis mystacinus

Myotis keaysi

Myotis nigricans

Myotis sodalis

Mos

chus

ber

ezov

skii

Mos

chus

chr

ysog

aste

r

Mos

chus

fusc

us

Mos

chus

mos

chife

rus

Acom

ys c

ahiri

nus

Acom

ys ru

ssat

us

Acom

ys s

pino

sissim

us

Acom

ys s

ubsp

inos

us

Loph

urom

ys fl

avop

unct

atus

Loph

urom

ys m

elan

onyx

Loph

urom

ys s

ikapu

si

Urano

mys

rudd

i

Aeth

omys

chr

ysop

hilu

s

Aeth

omys

inep

tus

Aeth

omys

kai

seri

Mic

aela

mys

nam

aque

nsis

Aeth

omys

nyi

kae

Chiru

rom

ys la

mia

Cross

omys

mon

ckto

ni

Hyom

ys g

olia

thLo

rent

zimys

nou

huys

i

Pseu

dohy

drom

ys e

llerm

ani

Para

lept

omys

rufila

tus

Mac

ruro

mys

maj

or

Mal

lom

ys ro

thsc

hild

i

Apod

emus

mys

taci

nus

Apod

emus

with

erby

i

Apod

emus

flav

icol

lis

Apod

emus

syl

vatic

us

Apom

ys a

brae

Apom

ys d

atae

Apom

ys h

yloce

tes

Apom

ys in

signi

s

Apom

ys lit

tora

lis

Apom

ys m

icrod

on

Apom

ys m

uscu

lus

Apom

ys s

acob

ianu

s

Archb

oldo

mys

luzo

nens

is

Rhync

hom

ys is

arog

ensis

Arv

ican

this

nilo

ticus

Ban

dico

ta b

enga

lens

is

Ban

dico

ta in

dica

Ban

dico

ta s

avile

i

Cra

tero

mys

sch

aden

berg

i

Chi

ropo

dom

ys g

liroi

des

Hap

alom

ys d

elac

ouri

Hap

alom

ys lo

ngic

auda

tus

Col

omys

gos

lingi

Conilu

rus p

enici

llatu

s

Lepo

rillus

cond

itor

Mes

embr

iomys

gou

ldii

Mes

embr

iomys

mac

ruru

s

Melomys

burto

ni

Melomys

cape

nsis

Melo

mys ce

rvini

pes

Param

elomys

plat

yops

Melo

mys ru

bicola

Uromys

caud

imac

ulatu

s

Zyzom

ys a

rgur

us

Zyzom

ys w

oodw

ardi

Legg

adina

forre

sti

Legg

adina

lake

downe

nsis

Notom

ys a

lexis

Notom

ys a

quilo

Notom

ys ce

rvinu

s

Notom

ys fu

scus

Notom

ys m

itche

llii

Pseud

omys

albo

ciner

eus

Pseud

omys

austr

alis

Pseud

omys

bolam

i

Pseud

omys

delic

atulus

Mastac

omys

fusc

us

Pseud

omys

grac

ilicau

datus

Pseud

omys

herm

anns

burg

ensis

Pseud

omys

higg

insi

Pseud

omys

nanu

s

Pseud

omys

nova

eholl

andia

e

Pseud

omys

occid

ental

is

Pseud

omys

pillig

aens

is

Pseud

omys

shor

tridg

ei

Das

ymys

inco

mtu

s

Das

ymys

rufu

lus

Dep

hom

ys d

efua

Dep

hom

ys e

burn

eae

Hybom

ys p

lani

frons

Hybom

ys tr

ivirg

atus

Hybom

ys u

nivit

tatu

s

Hyd

rom

ys c

hrys

ogas

ter

Xero

mys

myo

ides

Echi

othr

ix le

ucur

a

Mel

asm

othr

ix n

aso

Tate

omys

mac

roce

rcus

Tate

omys

rhin

ogra

doid

esG

ram

mom

ys c

omet

es

Gra

mm

omys

dol

ichu

rus

Gra

mm

omys

poe

nsis

Thal

lom

ys n

igric

auda

Thal

lom

ys p

aedu

lcus

Hei

mys

cus

fum

osus

Hyl

omys

cus

aeta

Hyl

omys

cus

alle

ni

Hyl

omys

cus

parv

us

Hyl

omys

cus

stel

la

Oen

omys

hyp

oxan

thus

Oen

omys

orn

atus

Tham

nom

ys v

enus

tus

Lem

nisc

omys

bar

baru

s

Lem

nisc

omys

mitt

endo

rfi

Lem

nisc

omys

rosa

lia

Lem

nisc

omys

stri

atus

Leno

thrix

can

us

Leop

olda

mys

edw

ards

i

Leop

olda

mys

sab

anus

Mal

acom

ys c

ansd

alei

Mal

acom

ys e

dwar

dsi

Mal

acom

ys lo

ngip

es

Pra

omys

luko

lela

e

Mas

tom

ys n

atal

ensi

s

Mas

tom

ys s

hort

ridge

i

Max

omys

inas

Max

omys

raja

h

Max

omys

sur

ifer

Max

omys

whi

tehe

adi

Mic

rom

ys m

inut

us

Mus

mus

culu

s

Mus

spr

etus

Mus

indu

tus

Mus

min

utoi

des

Mus

set

ulos

us

Mus

trito

n

Myl

omys

dyb

owsk

ii

Pelo

mys

falla

x

Pra

omys

dal

toni

Myo

mys

cus

verr

eaux

ii

Ste

noce

phal

emys

alb

ocau

data

Nes

okia

indi

ca

Niv

iven

ter c

onfu

cian

us

Niv

iven

ter c

rem

oriv

ente

r

Niv

iven

ter f

ulve

scen

s

Oto

mys

ang

onie

nsis

Oto

mys

irro

ratu

s

Myo

tom

ys s

logg

etti

Oto

mys

typu

s

Myo

tom

ys u

nisu

lcat

us

Paro

tom

ys b

rant

sii

Paro

tom

ys li

ttled

alei

Phl

oeom

ys c

umin

gi

Phl

oeom

ys p

allid

us

Pogo

nom

ys c

ham

pion

i

Pogo

nom

ys lo

riae

Pogo

nom

ys m

acro

urus

Pogo

nom

ys s

ylve

stris

Pra

omys

del

ecto

rum

Pra

omys

tullb

ergi

Rattu

s coll

etti

Rattu

s fus

cipes

Rattu

s lut

reolu

s

Rattu

s sor

didus

Rattu

s tun

neyi

Rattu

s villo

sissim

us

Rattu

s le

ucop

us

Rattu

s an

nand

alei

Rattu

s ex

ulan

sRat

tus a

rgen

tiven

ter

Rattu

s eve

retti

Rattu

s rat

tus

Rattu

s and

aman

ensis

Rattu

s tio

man

icus

Rattu

s no

rveg

icus

Rha

bdom

ys p

umili

o

Rat

tus

vere

cund

us

Sto

chom

ys lo

ngic

auda

tus

Sun

dam

ys m

uelle

ri

Vand

eleu

ria n

olth

enii

Vand

eleu

ria o

lera

cea

Zelo

tom

ys w

oosn

ami

Mer

ione

s cr

assu

s

Mer

ione

s lib

ycus

Mer

ione

s sa

cram

enti

Mer

ione

s tr

istra

mi

Mer

ione

s un

guic

ulat

us

Sek

eeta

mys

cal

urus

Psa

mm

omys

obe

sus

Psa

mm

omys

vex

illar

is

Rho

mbo

mys

opi

mus

Des

mod

illis

cus

brau

eri

Des

mod

illus

aur

icul

aris

Ger

billu

rus

paeb

a

Ger

billu

rus

tyto

nis

Ger

billu

rus

valli

nus

Ger

billu

s an

ders

oni

Dip

odill

us c

ampe

stris

Ger

billu

s ch

eesm

ani

Dip

odill

us d

asyu

rus

Ger

billu

s ge

rbill

us

Ger

billu

s he

nley

i

Ger

billu

s na

nus

Ger

billu

s py

ram

idum

Mic

rodi

llus

peel

i

Ger

billi

scus

boe

hmi

Ger

billi

scus

bra

ntsi

i

Ger

billi

scus

incl

usus

Ger

billi

scus

leuc

ogas

ter

Tamandua tetradactyla

Tamandua m

exicana

Myrm

ecophaga tridactyla

Noctilio albiventris

Noctilio leporinus

Hylobates agilisHylobates larHylobates moloch

Hylobates muelleriHylobates pileatusHylobates klossii

Bunopithecus hoolock

Symphalangus syndactylusNomascus concolor

Nomascus leucogenysNomascus gabriellae

Nycteris thebaicaNycteris grandis

Nycteris hispida

Arctocebus aureus

Arctocebus calabarensis

Perodicticus potto

Loris tardigradus

Loris lydekkerianus

Nycticebus coucang

Nycticebus pygmaeus

Nyctim

ene robinso

ni

Cynopteru

s sphinx

Megaloglossus w

oermanni

Eidolon helvum

Eidolon dupreanum

Myony

cteris

torquata

Rousettu

s aegyp

tiacu

s

Rousettu

s lesc

henaultii

Rousettu

s madagasc

ariensis

Scotonyc

teris ze

nkeri

Nanonycteris

veldka

mpii

Hypsig

nathus monstr

osus

Epomops franqueti

Micropteropus p

usillus

Epomophorus wahlbergi

Epomophorus angolensis

Epomophorus gambianus

Macrogloss

us minim

us

Syconyc

teris austr

alis

Dobsonia m

oluccensis

Pteropus melanotus

Pteropus hyp

omelanus

Pteropus alecto

Pteropus dasy

mallus

Pteropus livin

gstonii

Pteropus consp

icillatus

Pteropus samoensis

Pteropus tonganus

Pteropus polio

cephalus

Pteropus moloss

inus

Pteropus giganteus

Pteropus vampyru

s

Pteropus sca

pulatus

Pteropus insu

laris

Ochotona collaris

Ochotona princeps

Ochotona koslowi

Ochotona rufescens

Ochotona macrotis

Ochotona roylei

Ochotona curzoniae

Octodon bridgesi

Octodon degus

Tympanoctom

ys barrerae

Octom

ys mim

ax

Spalacopus cyanus

Octodontom

ys gliroides

Aconaemys sagei

Aconaemys fuscus

Gira

ffa c

amel

opar

dalis

Oka

pia

john

ston

i

Euoticus elegantulusEuoticus pallidus

Galago gallarumGalago matschiei

Galago moholiGalago senegalensis

Otolemur crassicaudatusOtolemur garnettii

Galago alleniGalago demidoff

Galago zanzibaricus

Cat

agon

us w

agne

ri

Taya

ssu

peca

riPe

cari

taja

cu

Dactylopsila megalura

Dactylopsila palpator

Dactylopsila tatei

Dactylopsila trivirgata

Gymnobelideus leadbeateri

Petaurus abidi

Petaurus australis

Petaurus breviceps

Petaurus gracilis

Petaurus norfolcensis

Bea

mys

hin

dei

Cric

etom

ys e

min

i

Cric

etom

ys g

ambi

anus

Sac

cost

omus

cam

pest

ris

Eliu

rus

min

or

Eliu

rus

myo

xinu

s

Bra

chyu

rom

ys b

etsi

leoe

nsis

Bra

chyu

rom

ys ra

miro

hitra

Nes

omys

ruf

us

Hyp

ogeo

mys

ant

imen

a

Mac

rota

rsom

ys b

asta

rdi

Mac

rota

rsom

ys in

gens

Del

anym

ys b

rook

siP

etro

mys

cus

barb

ouri

Pet

rom

yscu

s co

llinu

s

Pet

rom

yscu

s m

ontic

ular

is

Pet

rom

yscu

s sh

ortr

idge

i

Den

drom

us m

elan

otis

Den

drom

us m

esom

elas

Den

drom

us m

ysta

calis

Ste

atom

ys c

aurin

us

Ste

atom

ys c

uppe

dius

Ste

atom

ys ja

ckso

ni

Ste

atom

ys k

rebs

ii

Ste

atom

ys p

arvu

s

Ste

atom

ys p

rate

nsis

Ste

atom

ys o

pim

us

Mal

acot

hrix

typi

ca

Meg

aden

drom

us n

ikol

ausi

Mys

trom

ys a

lbic

auda

tus

Prio

nom

ys b

ates

i

Pus

a si

biric

a

Pho

ca la

rgha

Pho

ca v

itulin

a

Hyd

rurg

a le

pton

yxLo

bodo

n ca

rcin

opha

gaLe

pton

ycho

tes

wed

delli

iM

iroun

ga a

ngus

tiros

tris

Miro

unga

leon

ina

Mon

achu

s sc

haui

nsla

ndi

Mon

achu

s m

onac

hus

Pro

cyon

can

criv

orus

Pro

cyon

loto

rP

rocy

on p

ygm

aeus

Nas

ua n

aric

aN

asua

nas

uaN

asue

lla o

livac

eaB

assa

riscu

s as

tutu

sB

assa

riscu

s su

mic

hras

tiB

assa

ricyo

n be

ddar

diB

assa

ricyo

n ga

bbii

Bas

saric

yon

lasi

usB

assa

ricyo

n pa

uli

Pot

os fl

avus

Avahi lanigerAvahi occidentalis

Propithecus diademaPropithecus edwardsi

Propithecus perrieri

Propithecus tattersalli

Propithecus verreauxiPropithecus coquereli

Indri indri

Hemibelideus lemuroides

Petauroides volans

Petropseudes dahli

Pseudochirops albertisii

Pseudochirops coronatus

Pseudochirops archeri

Pseudochirops cupreus

Pseudochirops corinnae

Pseudochirulus cinereus

Pseudochirulus herbertensis

Pseudocheirus peregrinus

Mormoops megalophyllaPteronotus davyi

Pteronotus personatus

Pteronotus macleayii

Rhinolophus blasii

Rhinolophus capensis

Rhinolophus cliv

osus

Rhinolophus darlin

gi

Rhinolo

phus

denti

Rhinolo

phus

ferru

mequin

um

Rhinolo

phus

hilde

bran

dtii

Rhinolo

phus

hipp

oside

ros

Rhinolo

phus

meg

aphy

llus

Rhinolo

phus

philip

pinen

sis

Rhinolo

phus

simula

tor

Rhinolo

phus

swinn

yi

Rhinolo

phus

land

eri

Marm

ota menzbieri

Marm

ota caudata

Marm

ota himalayana

Marm

ota marm

ota

Marm

ota monax

Marm

ota flaviventrisM

armota olym

pus

Marm

ota caligata

Marm

ota vancouverensis

Sperm

ophilus columbianus

Sperm

ophilus parryiiS

permophilus richardsonii

Sperm

ophilus armatus

Sperm

ophilus tridecemlineatus

Sperm

ophilus franklinii

Sperm

ophilus washingtoni

Sperm

ophilus spilosoma

Sperm

ophilus beecheyi

Sperm

ophilus citellus

Sperm

ophilus suslicus

Sperm

ophilus beldingi

Sperm

ophilus adocetus

Sperm

ophilus mollis

Sperm

ophilus brunneus

Sperm

ophilus townsendii

Sperm

ophilus tereticaudus

Sperm

ophilus mohavensis

Sperm

ophilus saturatus

Sperm

ophilus mexicanus

Sperm

ophilus elegans

Sperm

ophilus lateralis

Sperm

ophilus variegatus

Cynom

ys gunnisoni

Cynom

ys leucurus

Cynom

ys parvidens

Cynom

ys ludovicianus

Cynom

ys mexicanus

Am

mosperm

ophilus leucurus

Am

mosperm

ophilus nelsoni

Am

mosperm

ophilus interpres

Am

mosperm

ophilus harrisiiTamias striatus

Tamias sibiricus

Tamias am

oenus

Tamias quadrim

aculatus

Tamias canipes

Tamias rufus

Tamias quadrivittatus

Tamias dorsalis

Tamias palm

eri

Tamias um

brinus

Tamias tow

nsendii

Tamias senex

Tamias m

erriami

Tamias obscurus

Tamias ruficaudus

Tamias ochrogenys

Tamias speciosus

Tamias m

inimus

Tamias alpinus

Tamias panam

intinusMyosciurus pum

ilioH

eliosciurus ruwenzorii

Heliosciurus gam

bianus

Heliosciurus m

utabilis

Heliosciurus punctatus

Heliosciurus rufobrachium

Epixerus ebii

Protoxerus stangeri

Funisciurus pyrropus

Funisciurus anerythrus

Funisciurus congicus

Funisciurus isabella

Funisciurus lemniscatus

Paraxerus cepapi

Paraxerus flavovittis

Paraxerus lucifer

Paraxerus palliatus

Paraxerus poensis

Sperm

ophilopsis leptodactylus

Atlantoxerus getulus

Xerus rutilus

Xerus erythropus

Xerus inauris

Xerus princeps

Tamiasciurus hudsonicus

Tamiasciurus douglasii

Sciurus carolinensis

Sciurus niger

Sciurus granatensisS

ciurus flamm

iferS

ciurus variegatoides

Sciurus anom

alus

Sciurus stram

ineus

Sciurus lis

Sciurus igniventris

Sciurus spadiceus

Sciurus arizonensis

Sciurus oculatusS

ciurus griseusS

ciurus alleniS

ciurus richmondi

Sciurus colliaei

Sciurus sanborni

Sciurus aureogaster

Sciurus pyrrhinus

Sciurus aestuansS

ciurus vulgaris

Sciurus ignitus

Sciurus yucatanensis

Sciurus gilvigularis

Sciurus pucheranii

Sciurus nayaritensis

Sciurus deppei

Microsciurus flaviventer

Microsciurus m

imulus

Microsciurus santanderensis

Microsciurus alfari

Syntheosciurus brochus

Pterom

ys volans

Pterom

ys mom

onga

Petaurista petaurista

Petaurista philippensis

Petaurista alborufus

Petaurista nobilis

Petaurista m

agnificus

Petaurista elegans

Petaurista xanthotis

Trogopterus xanthipes

Pterom

yscus pulverulentus

Aerom

ys tephromelas

Aerom

ys thomasi

Eupetaurus cinereus

Aeretes m

elanopterus

Glaucom

ys volans

Glaucom

ys sabrinus

Eoglaucom

ys fimbriatus

Hylopetes lepidus

Petinom

ys setosus

Petinom

ys crinitus

Petinom

ys genibarbis

Petinom

ys hageni

Petinom

ys sagitta

Petinom

ys vordermanni

Hylopetes alboniger

Hylopetes nigripes

Hylopetes spadiceus

Petaurillus kinlochii

Petaurillus hosei

Iomys horsfieldii

Ratufa bicolor

Ratufa affinis

Ratufa indica

Ratufa m

acroura

Callosciurus prevostii

Callosciurus caniceps

Callosciurus finlaysonii

Callosciurus orestes

Callosciurus adam

si

Callosciurus notatus

Callosciurus inornatus

Callosciurus baluensis

Callosciurus phayrei

Callosciurus m

elanogaster

Callosciurus quinquestriatus

Lariscus insignis

Lariscus obscurus

Rhinosciurus laticaudatusF

unambulus palm

arum

Funam

bulus pennantii

Tamiops m

cclellandii

Tamiops m

aritimus

Tamiops rodolphii

Tamiops sw

inhoei

Drem

omys rufigenis

Drem

omys lokriah

Nannosciurus m

elanotis

Sundasciurus hippurus

Sundasciurus low

ii

Sundasciurus tenuis

Callosciurus nigrovittatus

Callosciurus pygerythrus

Sciurillus pusillus

Solenodon cubanus

Solenodon paradoxus

Crocidura cyanea

Crocidura flavescens

Crocidura fuscom

urina

Crocidura hirta

Crocidura leucodon

Crocidura luna

Crocidura m

ariquensis

Crocidura olivieri

Crocidura suaveolens

Diplom

esodon pulchellum

Myosorex cafer

Myosorex longicaudatus

Myosorex varius

Scutisorex som

ereni

Suncus etruscus

Suncus infinitesim

us

Suncus m

urinus

Suncus varilla

Sorex arcticus

Sorex coronatus

Sorex araneus

Sorex unguiculatus

Sorex m

inutus

Sorex cinereus

Sorex hoyi

Sorex fum

eusS

orex longirostris

Sorex ornatus

Sorex bairdi

Sorex palustris

Sorex m

onticolusS

orex vagransS

orex pacificus

Sorex trow

bridgii

Sorex arizonae

Blarina carolinensis

Blarina brevicauda

Cryptotis goodw

ini

Cryptotis m

agna

Cryptotis m

eridensis

Cryptotis nigrescens

Cryptotis parva

Cryptotis thom

asi

Cryptotis gracilis

Cryptotis endersi

Cryptotis hondurensis

Cryptotis m

exicana

Cryptotis goldm

ani

Cryptotis squam

ipes

Cryptotis m

ontivaga

Neom

ys anomalus

Neom

ys fodiens

Neom

ys teres

Chim

arrogale platycephalus

Chim

arrogale himalayica

Notiosorex craw

fordi

Rhi

zom

ys s

umat

rens

is

Tach

yory

ctes

mac

roce

phal

usS

pala

x eh

renb

ergi

Spa

lax

leuc

odon

Spa

lax

mic

roph

thal

mus

Helogale hirtula

Helogale parvula

Mungos gam

bianusM

ungos mungo

Crossarchus alexandri

Crossarchus ansorgei

Crossarchus obscurus

Ichneumia albicauda

Suricata suricatta

Bdeogale crassicauda

Bdeogale jacksoni

Bdeogale nigripes

Liberiictis kuhniC

ynictis penicillataP

aracynictis selousiH

erpestes edwardsi

Herpestes javanicus

Herpestes brachyurus

Herpestes fuscus

Herpestes urva

Herpestes ichneum

onH

erpestes nasoH

erpestes semitorquatus

Herpestes sm

ithiiH

erpestes vitticollisG

alerella pulverulentaG

alerella sanguineaA

tilax paludinosusD

ologale dybowskii

Rhynchogale m

elleri

Bab

yrou

sa b

abyr

ussa

Hyl

ocho

erus

mei

nert

zhag

eni

Pota

moc

hoer

us la

rvat

us

Pota

moc

hoer

us p

orcu

s

Sus

bar

batu

s

Sus

ceb

ifron

s

Sus

cel

eben

sis

Sus

phi

lippe

nsis

Sus

sal

vani

usS

us s

crof

a

Pha

coch

oeru

s ae

thio

picu

s

Pha

coch

oeru

s af

rican

us

Tachyglossus aculeatus

Zaglossus bruijni

Cheiromeles t

orquatus

Myopterus d

aubentonii

Cynomops g

reenhalli

Cynomops p

lanirostr

is

Mormopterus b

eccarii

Mormopterus n

orfolke

nsis

Mormopterus p

lanicepsTa

darida brasili

ensis

Tadarid

a fulm

inans

Nyctinomops fe

morosaccu

s

Nyctinomops la

ticaudatus

Nyctinomops m

acrotisMolossus sinaloae

Molossus currentium

Molossus molossus

Molossus rufus

Molossus pretiosus

Eumops perotis

Eumops glaucinus

Eumops auripendulus

Chaerephon nigeriae

Talpa davidiana

Talpa stankovici

Talpa romana

Talpa occidentalis

Talpa europaea

Talpa caeca

Talpa levantis

Mogera w

ogura

Mogera tokudae

Mogera im

aizumii

Euroscaptor m

izura

Mogera uchidai

Condylura cristata

Parascalops breweri

Scalopus aquaticus

Scapanus latim

anus

Scapanus tow

nsendii

Scapanus orarius

Neurotrichus gibbsii

Dym

ecodon pilirostris

Urotrichus talpoides

Desm

ana moschata

Galem

ys pyrenaicus

Tapi

rus

baird

ii

Tapi

rus

pinc

haqu

e

Tapi

rus

terr

estr

isTa

piru

s in

dicu

s

Tarsius bancanusTarsius dentatusTarsius pumilusTarsius tarsierTarsius syrichta

Thryonomys swinderianus

Thryonomys gregorianus

Thyroptera discifera

Thyroptera tricolor

Hye

mos

chus

aqu

atic

us

Mos

chio

la m

emin

na

Trag

ulus

java

nicu

s

Trag

ulus

nap

u

Trichechus inunguis

Trichechus manatus

Ailurops ursinus

Phalanger carmelitae

Phalanger vestitus

Phalanger orientalis

Phalanger intercastellanus

Phalanger sericeus

Phalanger lullulae

Strigocuscus pelengensis

Strigocuscus celebensis

Phalanger gymnotis

Spilocuscus maculatus

Spilocuscus kraemeri

Spilocuscus rufoniger

Phalanger matanim

Phalanger ornatusTrichosurus arnhemensis

Trichosurus vulpecula

Trichosurus caninus

Wyulda squamicaudata

Anathana ellioti

Dendrogale melanura

Dendrogale murina

Tupaia belangeri

Tupaia dorsalis

Tupaia glis

Tupaia gracilis

Tupaia javanica

Tupaia longipes

Tupaia minor

Tupaia montana

Tupaia nicobarica

Tupaia palawanensis

Tupaia picta

Tupaia splendidula

Tupaia tana

Cep

halo

rhyn

chus

hec

tori

Ste

nella

coe

rule

oalb

a

Sot

alia

fluv

iatil

is

Ste

nella

long

irost

ris

Lasiorhinus krefftii

Lasiorhinus latifrons

Vombatus ursinus

200 160 120 80 40 0 Mya

b

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0.0 0.2 0.4 0.6 0.8

020

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0.74

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a

0.0 0.2 0.4 0.6 0.8

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Sam

ple

dens

ity

0.590.1

0.31

b

Page 32: 1 Temporal niche expansion in mammals from a nocturnal ...€¦ · Maor et al. Evolution of activity patterns in mammals 5 91 before the extinction of dinosaurs, calling for a wider

MonotremataMonotremata

RodentiaRodentia

Rodentia

RodentiaRodentia

RodentiaRodentia

RodentiaRodentiaRodentiaRodentiaRodentiaRodentia

RodentiaRodentia

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PrimatesPrimates

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166.2 140 120 100 80 60 40 20 0

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Rodentia

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DermopteraDermoptera

CingulataCingulata

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AfrosoricidaAfrosoricida

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MacroscelideaMacroscelidea

ProboscideaProboscidea

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PerissodactylaPerissodactyla

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PholidotaPholidota

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