COMMENTARY
COMMENTARY
Present-day drivers do not explain biodiversity
patterns in mammals
Richard T. Corletta,b,1
Mammals are an obvious choice for analyses of global
biodiversity patterns. They are not too diverse, dispro-
portionately well studied, and even nonspecialists will
be interested in the results. They are also fairly good
indicators of overall vertebrate diversity (1). Moreover,
the limited ability of most mammals to cross oceanic
barriers and the lack of direct land connections between
the Neotropics (South and Central America), Africa,
Madagascar, Asia, and Australasia (Australia and New
Guinea) provide an opportunity to compare more or
less independent evolutionary responses to similar trop-
ical and subtropical environments (2). Such comparisons
are complicated, however, by the widespread impacts
of climate change and human activities over the last
100,000 y, from the Late Pleistocene megafaunal ex-
tinctions to the ongoing consequences of recent human
population growth and economic development across
the tropics (3). Understanding the reasons for the simi-
larities and differences within and between regions is
of more than just theoretical interest, since it may also
have practical importance for conservation manage-
ment. In PNAS, Rowan et al. (4) address this issue and
attempt to determine the relative importance of differ-
ent factors in driving patterns of mammalian biodiver-
sity across the tropics and subtropics.
They excluded smaller species (<500 g) from their
analyses, greatly reducing the scale of the task, since
most mammals are small rodents or bats, while greatly
increasing the quality of the data available, since
larger species are easier to find and study. They were
then able to assemble an impressive dataset consist-
ing of 515 checklists of medium and large mammals
from across four zoogeographic realms: Afrotropical,
Indomalayan, Malagasy (Madagascar), and Neotropical.
They excluded tropical Australasia, presumably be-
cause there were too few complete checklists from this
realm. The final dataset included 852 species, many of
which are now under threat from hunting and/or habit
loss. For each community, they quantified both phylo-
genetic structure, using two measures of relatedness
among the component species, and functional structure,
a
Center for Integrative Conservation, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Mengla 666303, Yunnan,
China; and bCenter of Conservation Biology, Core Botanical Gardens, Chinese Academy of Sciences, Mengla 666303, Yunnan, China
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Author contributions: R.T.C. wrote the paper.
The author declares no competing interest.
Published under the PNAS license.
See companion article 10.1073/pnas.1910489116.
1
Email: corlett@xtbg.org.cn.
www.pnas.org/cgi/doi/10.1073/pnas.1921257117
using the range of body masses and diet types within
the community, and the evenness with which the spe-
cies are spaced along these trait axes.
They investigated four factors that potentially
influenced these current patterns—present and past
climates and recent and prehistoric human impacts.
Each of these has been studied before, but not to-
gether, not across all four realms, and not with such a
large database. To assess the potential importance
of past climates they used changes in temperature
and precipitation since the Last Glacial Maximum,
20,000 y ago. As a proxy for recent human impacts,
they used changes in the extent of anthropogenic
biomes (anthromes, such as rangelands, croplands,
and settlements) mapped for AD 1700 and 2000 (5).
Prehistoric human impacts were estimated by sub-
tracting present-day mammalian species richness from
estimates of what this richness would have been if
prehistoric human-driven extinctions and extirpations
since the Late Pleistocene had not occurred. The rela-
tionships between each community diversity metric and
the four potential drivers were then analyzed separately
for each realm using simultaneous autoregressive spa-
tial error models.
The relative importance of each factor, and even
the directions in which they acted, varied greatly across
realms (Table 1). Present-day drivers alone were no-
where sufficient to explain observed patterns in commu-
nity structure, although modern climate—temperature
and rainfall—was the most important single factor for
Africa. Modern climate—but temperature more than
rainfall—was also important in the Indomalayan and
Neotropical realms. Past climates also had an influ-
ence in Africa, but the effects of both recent and pre-
historic human impacts were weak. This is consistent
with the idea that the gradual evolution of humans in
Africa allowed the regional fauna to adapt, while the
other tropical regions experienced a sudden influx of
a novel intelligent, social, tool-using predator. Human
influences on community structure were strongest in
the Neotropics, followed by the Indomalayan realm,
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 Table 1. The influence of present and past climates and recent
and prehistoric human impacts on mammal community
structures in four zoogeographical realms in the tropics
and subtropics
Factors Neotropics Malagasy Afrotropics Indomalayan
Modern climate Moderate (T) Weak Strong (T, P) Strong (T)
Past climates Weak None Strong Moderate
Historical human Strong None Weak Moderate
impacts
Prehistoric human Strong None Weak Moderate
impacts
Modern climates are represented by mean annual temperature (T) and annual
precipitation (P) and past climates by the changes since the Last Glacial Maximum.
Historical human impacts are the changes in the extent of human-modified
“anthromes” from AD 1700 to 2000. Prehistoric human impacts are the differences
between the expected species richness if human-driven late-Quaternary extinc-
tions and extirpations had not occurred and present-day richness.
with the lower impact in the latter perhaps reflecting the presence
of Homo erectus for more than 1 million y before the arrival of
full-fledged modern humans. Past climates had a stronger im-
pact in Indomalaya than in the Neotropics, where any effects
may have been masked by the overwhelming human impacts.
Finally, Malagasy mammal communities were poorly predicted
by any of the factors included in the analyses, perhaps because
of their low phylogenetic diversity, since few clades were able to
colonize across oceanic barriers, and the massive anthropogenic
extinctions of the last 1,000 y (6).
Despite the huge dataset and sophisticated analyses, the lack
of consistent patterns across the tropics—and of any patterns in
Malagasy—may concern ecologists, who are wary of case-by-case
storytelling, even when data driven. Also, several potentially signif-
icant drivers were left out of the analyses. Rowan et al. (4) highlight
the potential importance of the “deep time” factors responsible for
the distinctiveness of the Malagasy fauna and, to a lesser extent, the
Neotropics, but there is no obvious way of quantifying these effects.
Other potential limitations are the use of only mean annual temper-
ature and precipitation to represent climate, ignoring seasonality,
and the use of changes in the extent of anthromes to represent all
types of recent human impacts, including hunting. Robust season-
ality estimates are unavailable for paleoclimates, however, at least at
a global scale, and there is currently no better global proxy for
recent human impacts, so it is unlikely that this study will be super-
seded in the near future.
It is interesting to speculate about where Australia and New
Guinea’s marsupial-dominated tropical mammal faunas would
have fitted in the analysis if they had been included. Previous
studies have shown strong trait convergence at the species and
assemblage level between Australia and areas with similar climates
in other realms (7). Like the Neotropics, Australasia experienced
massive Late Pleistocene megafaunal extinctions, but there was
no equivalent of the Great American Biotic Interchange, which
transformed the Neotropical mammal fauna relatively recently
(8). Despite this difference, the major drivers in Australasia seem
likely to be similar to those in the Neotropics, with both human
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1 C. N. Jenkins, S. L. Pimm, L. N. Joppa, Global patterns of terrestrial vertebrate diversity and conservation. Proc. Natl. Acad. Sci. U.S.A. 110, E2602–E2610 (2013).
2 R. T. Corlett, R. B. Primack, Tropical Rain Forests: An Ecological and Biogeographical Comparison (Wiley-Blackwell, Chichester, UK), ed. 2, 2011).
3 F. A. Smith et al., The accelerating influence of humans on mammalian macroecological patterns over the late Quaternary. Quat. Sci. Rev. 211, 1–16 (2019).
4 J. Rowan et al., Geographically divergent evolutionary and ecological legacies shape mammal biodiversity in the global tropics and subtropics. Proc. Natl. Acad.
Sci. U.S.A., 10.1073/pnas.1910489116 (2019).
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and climatic factors significant, but, like the Afrotropics, with
rainfall at least as important as temperature. Could the same
approach be applied to birds, which have similar patterns of
species diversity to mammals (1) and a lot of high-quality data?
Prehistoric bird extinctions were as size-selective as those of
mammals and thus involved fewer species, but there are no
obvious reasons to expect that the other factors—recent human
impacts and the influence of past and present climates—would
have acted differently on birds. The deep-time evolutionary his-
tory of birds was very different from that of mammals, however,
and they are more capable of crossing oceanic barriers: both of
which may have influenced present-day community structures.
Unfortunately, there is probably no other group of organisms for
which the available data are good enough.
What are the implications for conservation? We can’t change
the legacy effects of past climates and human activities, but we
can potentially use them as a guide to future actions (9). First,
Rowan et al. (4) once again highlight the unique position of
Africa as the last remaining area that still possesses more or less
intact, climatically determined, mammal communities. Africa
shows what nature can do if given the opportunity. These mam-
mal communities are now under massive threats and the African
countries that host them need all of the help they can get in
meeting human needs while safeguarding their globally impor-
tant biotas (10). A similar priority should be given to the few
remaining sites in Asia with near-intact communities. Where
species in Africa and Asia have been locally extirpated in histor-
ical times, but still survive elsewhere, reintroduction may be an
option in the future. No such option exists where species are
globally extinct, as with most of the larger species lost from the
Neotropics and Malagasy, but these two realms also still support
hyperdiverse mammalian communities, with most species en-
demic, so their conservation continues to be both worthwhile
and urgent.
Finally, the analyses of Rowan et al. (4) have mixed messages in
regard to the impacts of future, anthropogenic, climate change.
Past climate change had a broad impact on community structure
everywhere except Malagasy. This shows that large-bodied endo-
therms are not, as might have been expected, less sensitive to
changes in temperature and rainfall than other taxonomic groups
and also that the relatively smaller climatic fluctuations experienced
at low latitudes than at high latitudes can still have an impact. This
suggests that tropical mammal faunas may be more vulnerable to
future climate change than has sometimes been assumed so far. On
the other hand, human impacts from hunting and land-use change
appear to have at least partly obscured the effects of past climates
in Indomalaya and the Neotropics, underlining the fact that mam-
mal species need secure habitats and protection from overhunting
before climate change becomes the biggest concern. We live in a
time of twin climatic and biodiversity crises, and they both need to
be addressed urgently.
Acknowledgments
R.T.C.’s research is supported by the Chinese Academy of Sciences (Grants
Y6ZK121B01 and Y9ZK011B08).
Corlett
 5 E. C. Ellis et al., Anthropogenic transformation of the biomes, 1700 to 2000. Glob. Ecol. Biogeogr. 19, 589–606 (2010).
6 K. Douglass et al., A critical review of radiocarbon dates clarifies the human settlement of Madagascar. Quat. Sci. Rev. 221, 105878 (2019).
7 F. Mazel et al., The geography of ecological niche evolution in mammals. Curr. Biol. 27, 1369–1374 (2017).
8 C. D. Bacon et al., Biological evidence supports an early and complex emergence of the Isthmus of Panama. Proc. Natl. Acad. Sci. U.S.A. 112, 6110–6115 (2015).
9 E. Polaina, M. González-Suárez, E. Revilla, The legacy of past human land use in current patterns of mammal distribution. Ecography 42, 1623–1635 (2019).
10 P. A. Lindsey et al., More than $1 billion needed annually to secure Africa’s protected areas with lions. Proc. Natl. Acad. Sci. U.S.A. 115, E10788–E10796 (2018).
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