1 temporal niche expansion in mammals from a nocturnal ...€¦ · maor et al. evolution of...
TRANSCRIPT
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
Maor et al. Evolution of activity patterns in mammals
2
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
Maor et al. Evolution of activity patterns in mammals
3
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
Maor et al. Evolution of activity patterns in mammals
4
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
Maor et al. Evolution of activity patterns in mammals
5
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
Maor et al. Evolution of activity patterns in mammals
6
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
Maor et al. Evolution of activity patterns in mammals
7
= 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
Maor et al. Evolution of activity patterns in mammals
8
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
Maor et al. Evolution of activity patterns in mammals
9
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
Maor et al. Evolution of activity patterns in mammals
10
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
Maor et al. Evolution of activity patterns in mammals
11
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
Maor et al. Evolution of activity patterns in mammals
12
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
Maor et al. Evolution of activity patterns in mammals
13
‘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
Maor et al. Evolution of activity patterns in mammals
14
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
Maor et al. Evolution of activity patterns in mammals
15
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
Maor et al. Evolution of activity patterns in mammals
16
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
Maor et al. Evolution of activity patterns in mammals
17
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
Maor et al. Evolution of activity patterns in mammals
18
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
Maor et al. Evolution of activity patterns in mammals
19
References 420
1 Aronson, B. D. et al. Circadian rhythms. Brain Research Reviews 18, 315-333, 421
doi:10.1016/0165-0173(93)90015-R (1993). 422
2 Kronfeld-Schor, N. & Dayan, T. Partitioning of Time as an Ecological Resource. 423
Annual Review of Ecology, Evolution, and Systematics 34, 153-181, 424
doi:10.1146/annurev.ecolsys.34.011802.132435 (2003). 425
3 DeCoursey, P. J. Diversity of Function of SCN Pacemakers in Behavior and Ecology 426
of Three Species of Sciurid Rodents. Biological Rhythm Research 35, 13-33, 427
doi:10.1080/09291010412331313214 (2004). 428
4 Hut, R. A., Kronfeld-Schor, N., van der Vinne, V. & De la Iglesia, H. in Progress in 429
Brain Research: The Neurobiology of Circadian Timing Vol. 199 (eds A. Kalsbeek, 430
M. Merrow, T. Roenneberg, & R.G. Foster) Ch. 17, 281-304 (Elsevier, 2012). 431
5 Joffe, B., Peichl, L., Hendrickson, A., Leonhardt, H. & Solovei, I. Diurnality and 432
Nocturnality in Primates: An Analysis from the Rod Photoreceptor Nuclei Perspective. 433
Evolutionary Biology 41, 1-11, doi:10.1007/s11692-013-9240-9 (2014). 434
6 Melin, A. D., Matsushita, Y., Moritz, G. L., Dominy, N. J. & Kawamura, S. Inferred 435
L/M cone opsin polymorphism of ancestral tarsiers sheds dim light on the origin of 436
anthropoid primates. Proceedings of the Royal Society B: Biological Sciences 280, 437
doi:10.1098/rspb.2013.0189 (2013). 438
7 Gutman, R. & Dayan, T. Temoral partitioning: A experiment with two species of 439
spiny mice. Ecology 86, 164-173, doi:10.1890/03-0369 (2005). 440
8 Jones, K. E. et al. PanTHERIA: a species-level database of life history, ecology, and 441
geography of extant and recently extinct mammals. Ecology 90, 2648-2648, 442
doi:10.1890/08-1494.1 (2009). 443
Maor et al. Evolution of activity patterns in mammals
20
9 Refinetti, R. The diversity of temporal niches in mammals. Biological Rhythm 444
Research 39, 173-192, doi:10.1080/09291010701682690 (2008). 445
10 Heesy, C. P. & Hall, M. I. The Nocturnal Bottleneck and the Evolution of Mammalian 446
Vision. Brain, Behavior and Evolution 75, 195-203 (2010). 447
11 Walls, G. L. The Vertebrate Eye and its Adaptive Radiation. (Cranbrook Institute of 448
Science, 1942). 449
12 Davies, W. I. L., Collin, S. P. & Hunt, D. M. Molecular ecology and adaptation of 450
visual photopigments in craniates. Molecular Ecology 21, 3121-3158, 451
doi:10.1111/j.1365-294X.2012.05617.x (2012). 452
13 Gerkema, M. P., Davies, W. I. L., Foster, R. G., Menaker, M. & Hut, R. A. The 453
nocturnal bottleneck and the evolution of activity patterns in mammals. Proceedings 454
of the Royal Society B: Biological Sciences 280, doi:10.1098/rspb.2013.0508 (2013). 455
14 Peichl, L. Diversity of mammalian photoreceptor properties: Adaptations to habitat 456
and lifestyle? The Anatomical Record Part A: Discoveries in Molecular, Cellular, and 457
Evolutionary Biology 287A, 1001-1012, doi:10.1002/ar.a.20262 (2005). 458
15 Hayden, S. et al. Ecological adaptation determines functional mammalian olfactory 459
subgenomes. Genome Research 20, 1-9, doi:10.1101/gr.099416.109 (2010). 460
16 Coleman, M. N. & Boyer, D. M. Inner Ear Evolution in Primates Through the 461
Cenozoic: Implications for the Evolution of Hearing. The Anatomical Record: 462
Advances in Integrative Anatomy and Evolutionary Biology 295, 615-631, 463
doi:10.1002/ar.22422 (2012). 464
17 Diamond, M. E., von Heimendahl, M., Knutsen, P. M., Kleinfeld, D. & Ahissar, E. 465
'Where' and 'what' in the whisker sensorimotor system. Nat Rev Neurosci 9, 601-612, 466
doi:10.1038/nrn2411 (2008). 467
Maor et al. Evolution of activity patterns in mammals
21
18 Crompton, A. W., Taylor, C. R. & Jagger, J. A. Evolution of homeothermy in 468
mammals. Nature 272, 333-336, doi:10.1038/272333a0 (1978). 469
19 Barnosky, A. D. et al. Has the Earth/'s sixth mass extinction already arrived? Nature 470
471, 51-57, doi:10.1038/nature09678 (2011). 471
20 Brusatte, S. L. et al. The extinction of the dinosaurs. Biological Reviews 90, 628-642, 472
doi:10.1111/brv.12128 (2015). 473
21 Angielczyk, K. D. & Schmitz, L. Nocturnality in synapsids predates the origin of 474
mammals by over 100 million years. Proceedings of the Royal Society B: Biological 475
Sciences 281, doi:10.1098/rspb.2014.1642 (2014). 476
22 Schmitz, L. & Motani, R. Nocturnality in Dinosaurs Inferred from Scleral Ring and 477
Orbit Morphology. Science, doi:10.1126/science.1200043 (2011). 478
23 Hall, M. I., Kamilar, J. M. & Kirk, E. C. Eye shape and the nocturnal bottleneck of 479
mammals. Proceedings of the Royal Society B: Biological Sciences 279, 4962-4968, 480
doi:10.1098/rspb.2012.2258 (2012). 481
24 Emerling, C. A., Huynh, H. T., Nguyen, M. A., Meredith, R. W. & Springer, M. S. 482
Spectral shifts of mammalian ultraviolet-sensitive pigments (short wavelength-483
sensitive opsin 1) are associated with eye length and photic niche evolution. 484
Proceedings of the royal Society B: Biological Sciences 282, 20151817, 485
doi:10.1098/rspb.2015.1817 (2015). 486
25 Reppert, S. M. & Weaver, D. R. Molecular Analysis of Mammalian Circadian 487
Rhythms. Annual Review of Physiology 63, 647-676, 488
doi:10.1146/annurev.physiol.63.1.647 (2001). 489
26 Griffin, R. H., Matthews, L. J. & Nunn, C. L. Evolutionary disequilibrium and 490
activity period in primates: A bayesian phylogenetic approach. American Journal of 491
Physical Anthropology 147, 409-416, doi:10.1002/ajpa.22008 (2012). 492
Maor et al. Evolution of activity patterns in mammals
22
27 Santini, L., Rojas, D. & Donati, G. Evolving through day and night: origin and 493
diversification of activity pattern in modern primates. Behavioral Ecology, 494
doi:10.1093/beheco/arv012 (2015). 495
28 Heesy, C. P. & Ross, C. F. Evolution of activity patterns and chromatic vision in 496
primates: morphometrics, genetics and cladistics. J Hum Evol 40, 111-149, 497
doi:10.1006/jhev.2000.0447 (2001). 498
29 Roll, U., Dayan, T. & Kronfeld-Schor, N. On the role of phylogeny in determining 499
activity patterns of rodents. Evolutionary Ecology 20, 479-490, doi:10.1007/s10682-500
006-0015-y (2006). 501
30 Meredith, R. W. et al. Impacts of the Cretaceous Terrestrial Revolution and KPg 502
Extinction on Mammal Diversification. Science 334, 521-524, 503
doi:10.1126/science.1211028 (2011). 504
31 Bininda-Emonds, O. R. P. et al. The delayed rise of present-day mammals. Nature 505
446, 507-512, doi:10.1038/nature05634 (2007). 506
32 Fritz, S. A., Bininda-Emonds, O. R. P. & Purvis, A. Geographical variation in 507
predictors of mammalian extinction risk: big is bad, but only in the tropics. Ecology 508
Letters 12, 538-549, doi:10.1111/j.1461-0248.2009.01307.x (2009). 509
33 Meade, A. & Pagel, M. BayesTraits v3 (2017). 510
34 Melin, A. D. et al. Euarchontan Opsin Variation Brings New Focus to Primate 511
Origins. Molecular Biology and Evolution 33, 1029-1041, 512
doi:10.1093/molbev/msv346 (2016). 513
35 Sakamoto, M., Benton, M. J. & Venditti, C. Dinosaurs in decline tens of millions of 514
years before their final extinction. Proceedings of the National Academy of Sciences 515
113, 5036-5040, doi:10.1073/pnas.1521478113 (2016). 516
Maor et al. Evolution of activity patterns in mammals
23
36 Close, Roger A., Friedman, M., Lloyd, Graeme T. & Benson, Roger B. J. Evidence 517
for a Mid-Jurassic Adaptive Radiation in Mammals. Current Biology 25, 2137-2142, 518
doi:10.1016/j.cub.2015.06.047 (2015). 519
37 Lee, Michael S. Y. & Beck, Robin M. D. Mammalian Evolution: A Jurassic Spark. 520
Current Biology 25, R759-R761, doi:10.1016/j.cub.2015.07.008 (2015). 521
38 Wilson, G. P. et al. Adaptive radiation of multituberculate mammals before the 522
extinction of dinosaurs. Nature 483, 457-460, doi:10.1038/nature10880 (2012). 523
39 Krause, D. W. et al. First cranial remains of a gondwanatherian mammal reveal 524
remarkable mosaicism. Nature 515, 512-517, doi:10.1038/nature13922 (2014). 525
40 Ross, C. F. Into the Light: The Origin of Anthropoidea. Annual Review of 526
Anthropology 29, 147-194, doi:10.1146/annurev.anthro.29.1.147 (2000). 527
41 dos Reis, M., Donoghue, P. C. J. & Yang, Z. Neither phylogenomic nor 528
palaeontological data support a Palaeogene origin of placental mammals. Biology 529
Letters 10, doi:10.1098/rsbl.2013.1003 (2014). 530
42 Foley, N. M., Springer, M. S. & Teeling, E. C. Mammal madness: is the mammal tree 531
of life not yet resolved? Philosophical Transactions of the Royal Society B: Biological 532
Sciences 371, doi:10.1098/rstb.2015.0140 (2016). 533
43 Tarver, J. E. et al. The Interrelationships of Placental Mammals and the Limits of 534
Phylogenetic Inference. Genome Biology and Evolution, doi:10.1093/gbe/evv261 535
(2016). 536
44 Springer, M. S. et al. Waking the undead: Implications of a soft explosive model for 537
the timing of placental mammal diversification. Molecular Phylogenetics and 538
Evolution 106, 86-102, doi:10.1016/j.ympev.2016.09.017 (2017). 539
45 Donati, G. & Borgognini-Tarli, S. M. From darkness to daylight: cathemeral activity 540
in primates. Journal of Anthropological Sciences 84, 7-32 (2006). 541
Maor et al. Evolution of activity patterns in mammals
24
46 Fullard, J. H. & Napoleone, N. Diel flight periodicity and the evolution of auditory 542
defences in the Macrolepidoptera. Animal Behaviour 62, 349-368, 543
doi:10.1006/anbe.2001.1753 (2001). 544
47 O'Leary, M. A. et al. The Placental Mammal Ancestor and the Post–K-Pg Radiation 545
of Placentals. Science 339, 662-667, doi:10.1126/science.1229237 (2013). 546
48 Wilson, D. E. & Reeder, D. A. Mammal Species of the World. (John Hopkins 547
University Press 2005). 548
49 Price, S. A., Bininda-Emonds, O. R. P. & Gittleman, J. L. A complete phylogeny of 549
the whales, dolphins and even-toed hoofed mammals (Cetartiodactyla). Biological 550
Reviews 80, 445-473, doi:10.1017/S1464793105006743 (2005). 551
50 O'Leary, M. A. & Gatesy, J. Impact of increased character sampling on the phylogeny 552
of Cetartiodactyla (Mammalia): combined analysis including fossils. Cladistics 24, 553
397-442, doi:10.1111/j.1096-0031.2007.00187.x (2008). 554
51 Springer, M. S., Meredith, R. W., Teeling, E. C. & Murphy, W. J. Technical 555
Comment on “The Placental Mammal Ancestor and the Post–K-Pg Radiation of 556
Placentals”. Science 341, 613-613, doi:10.1126/science.1238025 (2013). 557
52 Paradis, E., Claude, J. & Strimmer, K. APE: analyses of phylogenetics and evolution 558
in R language. Bioinformatics 20, 289-290, doi:10.1093/bioinformatics/btg412 (2004). 559
53 Schliep, K. P. phangorn: phylogenetic analysis in R. Bioinformatics 27, 592-593, 560
doi:10.1093/bioinformatics/btq706 (2011). 561
54 R: A language and environment for statistical computing (R Foundation for Statistical 562
Computing, Vienna, Austria, 2015). 563
55 Pagel, M. & Meade, A. Bayesian analysis of correlated evolution of discrete 564
characters by reversible-jump Markov chain Monte Carlo. The American Naturalist 565
167, 808-825, doi:10.1086/503444 (2006). 566
Maor et al. Evolution of activity patterns in mammals
25
56 Kirk, E. C. Effects of activity pattern on eye size and orbital aperture size in primates. 567
J Hum Evol 51, 159-170, doi:10.1016/j.jhevol.2006.02.004 (2006). 568
57 Xie, W., Lewis, P. O., Fan, Y., Kuo, L. & Chen, M.-H. Improving Marginal 569
Likelihood Estimation for Bayesian Phylogenetic Model Selection. Systematic 570
Biology 60, 150-160, doi:10.1093/sysbio/syq085 (2011). 571
58 Revell, L. J. phytools: an R package for phylogenetic comparative biology (and other 572
things). Methods in Ecology and Evolution 3, 217-223, doi:10.1111/j.2041-573
210X.2011.00169.x (2012). 574
59 Maddison, W. P. Confounding asymmetries in evolutionary diverification and 575
character change. Evolution 60, 1743-1746, doi:10.1554/05-666.1 (2006). 576
60 FitzJohn, R. G. Diversitree: comparative phylogenetic analyses of diversification in R. 577
Methods in Ecology and Evolution 3, 1084-1092, doi:10.1111/j.2041-578
210X.2012.00234.x (2012). 579
61 Kuhn, T. S., Mooers, A. Ø. & Thomas, G. H. A simple polytomy resolver for dated 580
phylogenies. Methods in Ecology and Evolution 2, 427-436, doi:10.1111/j.2041-581
210X.2011.00103.x (2011). 582
583
584
Maor et al. Evolution of activity patterns in mammals
26
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
Maor et al. Evolution of activity patterns in mammals
27
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
Maor et al. Evolution of activity patterns in mammals
28
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
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
Tachyglossus aculeatus
Zaglossus bruijni
Ornithorhynchus anatinus
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
Ped
etes
cap
ensi
s
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
Acomys
cahirin
us
Acomys
russ
atus
Acomys
spinosis
simus
Acomys
subsp
inosus
Lophuromys
flavo
punctatus
Lophuromys
melanony
x
Lophuromys
sika
pusi
Uranomys
ruddi
Aetho
mys
chry
soph
ilus
Aetho
mys
inep
tus
Aetho
mys
kaise
ri
Mica
elam
ys n
amaq
uens
is
Aetho
mys
nyik
ae
Chirur
omys
lamia
Cross
omys
mon
ckton
i
Hyomys
golia
th
Lorentzi
mys nouhuys
i
Pseud
ohyd
romys
eller
mani
Parale
ptomys
rufila
tus
Macru
romys
majo
r
Mallom
ys ro
thsch
ildi
Apode
mus
mys
tacin
us
Apode
mus
with
erby
i
Apode
mus
flavic
ollis
Apode
mus
sylva
ticus
Apomys abrae
Apomys datae
Apomys hy
locetes
Apomys in
signis
Apomys lit
toralis
Apomys m
icrodon
Apomys m
usculus
Apomys sa
cobianus
Archboldomys
luzo
nensis
Rhynch
omys is
arogensis
Arvi
cant
his
nilo
ticus
Bandi
cota
ben
gale
nsis
Band
icota
indi
ca
Band
icota
sav
ilei
Cra
tero
mys
sch
aden
berg
i
Chiro
podo
mys
glir
oide
s
Hapal
omys
del
acou
ri
Hapal
omys
long
icaud
atus
Col
omys
gos
lingi
Conilurus penicil
latus
Leporillus c
onditor
Mesembrio
mys gouldii
Mesembrio
mys macru
rus
Melomys burto
ni
Melomys ca
pensis
Melomys ce
rvinipes
Paramelomys platyo
ps
Melomys rubico
la
Uromys ca
udimacu
latus
Zyzomys
argurus
Zyzomys
woodwardi
Leggadina forre
sti
Leggadina lake
downensis
Notomys alexis
Notomys aquilo
Notomys ce
rvinus
Notomys fu
scus
Notomys m
itchellii
Pseudomys albocinereus
Pseudomys australis
Pseudomys bolami
Pseudomys delicatulus
Mastacomys fuscus
Pseudomys gracilicaudatus
Pseudomys hermannsburgensis
Pseudomys higginsi
Pseudomys nanus
Pseudomys novaehollandiae
Pseudomys occidentalis
Pseudomys pilligaensis
Pseudomys
shortri
dgei
Das
ymys
inco
mtu
s
Das
ymys
rufu
lus
Depho
mys de
fua
Depho
mys eb
urne
ae
Hybom
ys pl
anifro
ns
Hybom
ys tr
ivirg
atus
Hybom
ys un
ivitta
tus
Hydro
mys ch
ryso
gaste
r
Xerom
ys m
yoide
s
Echio
thrix
leuc
ura
Mel
asm
othr
ix na
so
Tate
omys
mac
roce
rcus
Tate
omys
rhino
grad
oides
Gram
mom
ys co
met
es
Gram
mom
ys d
olich
urus
Gram
mom
ys p
oens
is
Thallo
mys
nigr
icaud
a
Thallo
mys
pae
dulcu
s
Hei
mys
cus
fum
osus
Hylom
yscu
s ae
ta
Hylom
yscu
s al
leni
Hylom
yscu
s pa
rvus
Hylom
yscu
s st
ella
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
Praom
ys lu
kole
lae
Mas
tom
ys n
atal
ensi
s
Mas
tom
ys s
hortr
idge
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
culus
Mus
spre
tus
Mus
indu
tus
Mus
minu
toide
s
Mus
setu
losus
Mus
trito
n
Myl
omys
dyb
owsk
ii
Pelo
mys
falla
x
Prao
mys
dal
toni
Myo
mys
cus
verre
auxii
Sten
ocep
hale
mys
alb
ocau
dataN
esok
ia in
dica
Nivive
nter
con
fucia
nus
Niv
ivent
er c
rem
orive
nter
Niv
ivent
er fu
lvesc
ens
Otomys
ango
niens
is
Otomys
irror
atus
Myo
tomys
slog
getti
Otomys
typu
s
Myo
tomys
unis
ulcat
us
Parot
omys
bra
ntsii
Parot
omys
little
dalei
Phlo
eom
ys c
umin
gi
Phlo
eom
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
ylves
tris
Prao
mys
del
ecto
rum
Prao
mys
tullb
ergi
Rattus c
olletti
Rattus f
uscipes
Rattus l
utreolus
Rattus s
ordidus
Rattus t
unneyi
Rattus v
illosis
simus
Rattus l
eucopus
Rattus a
nnandalei
Rattus e
xulans
Rattus a
rgentiventer
Rattus e
veretti
Rattus r
attus
Rattus a
ndamanensis
Rattus t
iomanicus
Rattus n
orvegicu
s
Rha
bdom
ys p
umilio
Rat
tus
vere
cund
us
Stoc
hom
ys lo
ngic
auda
tus
Sund
amys
mue
lleri
Vand
eleu
ria n
olth
enii
Vand
eleu
ria o
lera
cea
Zelo
tom
ys w
oosn
ami
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
Cric
etus
cric
etus
Cric
etul
us m
igra
toriu
sM
esoc
ricet
us a
urat
us
Mes
ocric
etus
new
toni
Pho
dopu
s ca
mpb
elli
Pho
dopu
s ro
boro
vski
i
Pho
dopu
s su
ngor
us
Alti
cola
arg
enta
tus
Alti
cola
sto
liczk
anus
Alti
cola
stre
lzow
iM
yode
s ca
lifor
nicu
s
Myo
des
gapp
eri
Myo
des
glar
eolu
s
Myo
des
rufo
canu
s
Myo
des
rutil
us
Din
arom
ys b
ogda
novi
Arb
orim
us a
lbip
es
Arb
orim
us lo
ngic
audu
s
Arb
orim
us p
omo
Phe
naco
mys
inte
rmed
ius
Phe
naco
mys
ung
ava
Arv
icol
a sa
pidu
s
Arv
icol
a am
phib
ius
Chi
onom
ys n
ival
is
Dic
rost
onyx
gro
enla
ndic
us
Dic
rost
onyx
rich
ards
oni
Pro
met
heom
ys s
chap
osch
niko
wi
Lagu
rus
lagu
rus
Lasi
opod
omys
bra
ndtii
Lasi
opod
omys
fusc
us
Lasi
opod
omys
man
darin
us
Lem
mis
cus
curt
atus
Lem
mus
lem
mus
Myo
pus
schi
stic
olor
Syn
apto
mys
bor
ealis
Syn
apto
mys
coo
peri
Mic
rotu
s ag
rest
is
Mic
rotu
s ch
roto
rrhin
us
Mic
rotu
s ric
hard
soni
Mic
rotu
s xa
ntho
gnat
hus
Mic
rotu
s gu
atem
alen
sis
Mic
rotu
s lo
ngic
audu
s
Mic
rotu
s m
exic
anus
Mic
rotu
s ar
valis
Mic
rotu
s ca
lifor
nicu
s
Mic
rotu
s m
onta
nus
Mic
rotu
s br
ewer
i
Mic
rotu
s pe
nnsy
lvan
icus
Mic
rotu
s to
wns
endi
iMic
rotu
s oe
cono
mus
Mic
rotu
s oc
hrog
aste
r
Mic
rotu
s pi
neto
rum
Mic
rotu
s ab
brev
iatu
s
Mic
rotu
s m
iuru
sM
icro
tus
mul
tiple
x
Mic
rotu
s du
odec
imco
stat
us
Mic
rotu
s sa
vii
Mic
rotu
s ba
varic
us
Mic
rotu
s su
bter
rane
us
Neo
fiber
alle
ni
Ond
atra
zib
ethi
cus
Mer
ione
s cr
assu
s
Mer
ione
s lib
ycus
Mer
ione
s sa
cram
enti
Mer
ione
s tri
stra
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
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
sB
rach
yuro
mys
bet
sile
oens
is
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
Rhi
zom
ys s
umat
rens
is
Tach
yory
ctes
mac
roce
phal
us
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
Loph
iom
ys im
haus
i
Mal
acot
hrix
typi
ca
Meg
aden
drom
us n
ikol
ausi
Mys
trom
ys a
lbic
auda
tus
Spa
lax
ehre
nber
gi
Spa
lax
leuc
odon
Spa
lax
mic
roph
thal
mus
Prio
nom
ys b
ates
i
Castor canadensis
Castor fiber
Per
ogna
thus
alti
colu
s
Per
ogna
thus
am
plus
Per
ogna
thus
fasc
iatu
s
Per
ogna
thus
flav
esce
ns
Per
ogna
thus
flav
usP
erog
nath
us in
orna
tus
Per
ogna
thus
long
imem
bris
Per
ogna
thus
mer
riam
i
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
x
Cha
etod
ipus
cal
iforn
icus
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
Tho
mom
ys b
otta
e
Thom
omys um
brinusT
homom
ys talpoides Geo
mys
per
sona
tus
Geo
mys
bur
sariu
sG
eom
ys a
ttwat
eri
Geo
mys
bre
vice
psG
eom
ys a
rena
rius
Geo
mys
pin
etis
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
Cra
toge
omys
neg
lect
usO
rtho
geom
ys h
eter
odus
Ort
hoge
omys
his
pidu
sO
rtho
geom
ys c
herr
iei
Ort
hoge
omys
und
erw
oodi
Ort
hoge
omys
dar
iens
isO
rtho
geom
ys c
avat
orO
rtho
geom
ys c
unic
ulus
Ort
hoge
omys
lani
usO
rtho
geom
ys th
aele
ri
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
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
lerv
ia
Budor
cas t
axico
lor
Ovis a
mm
on
Ovis a
ries
Ovis d
alli
Capra
cauc
asica
Capra
falco
neri
Capra
hirc
us
Capra
pyr
enaic
a
Capra
sibir
ica
Hemitr
agus
hylo
crius
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
rupi
capr
a
Capric
orni
s cr
ispus
Capric
orni
s m
ilnee
dwar
dsii
Ovib
os m
osch
atus
Ore
amno
s am
erica
nus
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
Nan
ger d
ama
Nan
ger s
oem
mer
ringi
i
Nan
ger g
rant
i
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
Bos
elap
hus
trago
cam
elus
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
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
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
asoa
eLe
pton
ycte
ris y
erba
buen
aeLe
pton
ycte
ris n
ival
isG
loss
opha
ga c
omm
issa
risi
Glo
ssop
haga
sor
icin
aA
nour
a cu
ltrat
aA
nour
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
fus
Mol
ossu
s pr
etio
sus
Eum
ops
pero
tis
Eum
ops
glau
cinu
sE
umop
s au
ripen
dulu
sC
haer
epho
n ni
geria
e
Min
iopt
erus
sch
reib
ersi
iM
inio
pter
us a
ustr
alis
Lasi
onyc
teris
noc
tivag
ans
Nyc
ticei
us h
umer
alis
Sco
tean
ax r
uepp
ellii
Oto
nyct
eris
hem
pric
hii
Ant
rozo
us p
allid
usV
espe
rtili
o m
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
s vu
lturn
usN
ycta
lus
azor
eum
Cha
linol
obus
tube
rcul
atus
Cha
linol
obus
dw
yeri
Cha
linol
obus
gou
ldii
Cha
linol
obus
mor
io
Cha
linol
obus
nig
rogr
iseu
s
Lasi
urus
egr
egiu
s
Lasi
urus
ega
Lasi
urus
inte
rmed
ius
Lasi
urus
cin
ereu
s
Lasi
urus
bor
ealis
Lasi
urus
sem
inol
us
Nyc
toph
ilus
arnh
emen
sis
Nyc
toph
ilus
geof
froy
iN
ycto
philu
s go
uldi
Nyc
toph
ilus
timor
iens
is
Bar
bast
ella
bar
bast
ellu
s
Eud
erm
a m
acul
atum
Idio
nyct
eris
phy
llotis
Cor
ynor
hinu
s ra
fines
quii
Cor
ynor
hinu
s to
wns
endi
iP
leco
tus
aurit
us
Sco
toph
ilus
borb
onic
usS
coto
philu
s vi
ridis
Sco
toph
ilus
robu
stus
Pip
istr
ellu
s ku
hlii
Pip
istr
ellu
s ru
stic
usP
ipis
trel
lus
pipi
stre
llus
Pip
istr
ellu
s te
nuis
Ept
esic
us n
ilsso
nii
Ept
esic
us b
rasi
liens
isE
ptes
icus
bot
tae
Ept
esic
us fu
rinal
isE
ptes
icus
fusc
usE
ptes
icus
hot
tent
otus
Myo
tis a
lbes
cens
Myo
tis a
ustr
orip
ariu
sM
yotis
gris
esce
nsM
yotis
levi
sM
yotis
luci
fugu
sM
yotis
oxy
otus
Myo
tis v
elife
rM
yotis
vol
ans
Myo
tis y
uman
ensi
sM
yotis
viv
esi
Myo
tis r
ipar
ius
Myo
tis a
dver
sus
Myo
tis b
ocag
iiM
yotis
lani
ger
Myo
tis d
aube
nton
ii
Myo
tis w
elw
itsch
ii
Myo
tis e
votis
Myo
tis k
eeni
iM
yotis
thys
anod
es
Myo
tis c
alifo
rnic
usM
yotis
leib
ii
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
0.0 0.2 0.4 0.6 0.8
020
4060
80
Sam
ple
dens
ity
0.74
0.24
0.02 ●
●
●
NocturnalCathemeralDiurnal
a
0.0 0.2 0.4 0.6 0.8
020
4060
80
Posterior Probability
Sam
ple
dens
ity
0.590.1
0.31
b
MonotremataMonotremata
RodentiaRodentia
Rodentia
RodentiaRodentia
RodentiaRodentia
RodentiaRodentiaRodentiaRodentiaRodentiaRodentia
RodentiaRodentia
RodentiaRodentia
LagomorphaLagomorpha
PrimatesPrimatesPrimates
Primates
PrimatesPrimates
Primates
DermopteraScandentia
ArtiodactylaArtiodactyla
CetaceaCetaceaArtiodactyla
ArtiodactylaArtiodactylaPerissodactylaPerissodactylaPerissodactylaCarnivoraCarnivoraCarnivoraCarnivoraCarnivoraPholidota
ChiropteraChiropteraChiropteraChiropteraChiropteraChiropteraChiropteraChiropteraChiropteraChiropteraChiropteraChiropteraChiropteraChiroptera
Soricomorpha
ErinaceomorphaErinaceomorpha
ErinaceomorphaSoricomorphaSoricomorpha
Soricomorpha
Pilosa
PilosaPilosa
Afrosoricida
AfrosoricidaAfrosoricida
Afrosoricida
MacroscelideaMacroscelideaTubulidentata
SireniaSirenia
HyracoideaProboscidea
DiprotodontiaDiprotodontia
DiprotodontiaMicrobiotheriaDasyuromorphiaNotoryctemorphiaPeramelemorphiaPaucituberculata
Didelphimorphia
DidelphimorphiaDidelphimorphia
DidelphimorphiaDidelphimorphia
166.2 140 120 100 80 60 40 20 0
Jura
ssic
Cre
tace
ous
K−Pg Mya
Rodentia
Primates
Chiroptera
Cetartiodacty
laCarn
ivora
Soricomorp
haAfro
theria
Marsupialia
Rodentia
Monotremata
a
02
520
5020
0
No.
of l
inea
ges
bNocturnalCathemeralDiurnal
Rodentia
Carnivora
Rodentia
Chiroptera
Pilosa
Primates
Microbiotheria
Sirenia
Cetacea
Diprotodontia
PeramelemorphiaDasyuromorphia
Chiroptera
Carnivora
Chiroptera
Notoryctemorphia
Tubulidentata
Rodentia
Scandentia
AfrosoricidaAfrosoricidaAfrosoricida
RodentiaRodentia
Chiroptera
Artiodactyla
Pilosa
PaucituberculataPaucituberculata
Didelphimorphia
DidelphimorphiaDidelphimorphia
DidelphimorphiaDidelphimorphia
Carnivora
RodentiaRodentia
Perissodactyla
Rodentia
RodentiaRodentia
DermopteraDermoptera
CingulataCingulata
Rodentia
AfrosoricidaAfrosoricida
Afrosoricida
MacroscelideaMacroscelidea
ProboscideaProboscidea
ChiropteraChiroptera
PerissodactylaPerissodactyla
ErinaceomorphaErinaceomorpha
Erinaceomorpha
RodentiaRodentiaRodentia
Rodentia
RodentiaRodentia
Rodentia
HyracoideaHyracoideaHyracoideaHyracoidea
Artiodactyla
Rodentia
RodentiaRodentiaRodentia
ArtiodactylaArtiodactyla
LagomorphaLagomorpha
Primates
PholidotaPholidota
Chiroptera
Primates
Chiroptera
Chiroptera
ChiropteraChiroptera
ChiropteraChiroptera
Chiroptera
ChiropteraChiroptera
ChiropteraChiroptera
LagomorphaLagomorpha
Primates
Chiroptera
RodentiaRodentia
SoricomorphaSoricomorpha
Soricomorpha
SoricomorphaSoricomorphaSoricomorphaSoricomorpha
Rodentia
Artiodactyla
Monotremata
Chiroptera
SoricomorphaSoricomorpha
Soricomorpha
SoricomorphaSoricomorpha
SoricomorphaSoricomorpha
Soricomorpha
Perissodactyla
PrimatesPrimates
Chiroptera
Scandentia
ScandentiaScandentia
217.8 200 180 160 140 120 100 80 60 40 20 0
Cre
tace
ous
Jura
ssic
Tria
ssic
K−Pg Mya
Rodentia
Primates
Chiroptera
Cetartiodacty
laCarn
ivora
Soricomorp
ha
Afrotheria
Marsupialia
Rodentia
Monotremata
a
02
520
5020
0
No.
of l
inea
ges
bNocturnalCathemeralDiurnal