Répartition

géographique
Gabriel Blouin-Demers

Professeur titulaire – Département de biologie

BIO2529 – Écologie
 Facteurs les plus importants
pour la répartition des espèces

• Évolution

• Dispersion

• Persistance / Extinction
Évolution
Pianka & Vitt (2003)
Pianka & Vitt (2003)
 Molecular Phylogenetics and Evolution 94 (2016) 537–547

Contents lists available at ScienceDirect

Molecular Phylogenetics and Evolution

journal homepage: www.elsevier.com/locate/ympev

Combining phylogenomic and supermatrix approaches, and a

time-calibrated phylogeny for squamate reptiles (lizards and snakes)
based on 52 genes and 4162 species q
Yuchi Zheng a,b, John J. Wiens b,⇑
a
Department of Herpetology, Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu 610041, China
b
Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, AZ 85721-088, USA

a r t i c l e i n f o a b s t r a c t

Article history: Two common approaches for estimating phylogenies in species-rich groups are to: (i) sample many loci
Received 26 May 2015 for few species (e.g. phylogenomic approach), or (ii) sample many species for fewer loci (e.g. supermatrix
Revised 30 September 2015 approach). In theory, these approaches can be combined to simultaneously resolve both higher-level rela-
Accepted 8 October 2015
tionships (with many genes) and species-level relationships (with many taxa). However, fundamental
Available online 22 October 2015
questions remain unanswered about this combined approach. First, will higher-level relationships more
closely resemble those estimated from many genes or those from many taxa? Second, will branch sup-
Keywords:
port increase for higher-level relationships (relative to the estimate from many taxa)? Here, we address
Missing data
Phylogenomic
these questions in squamate reptiles. We combined two recently published datasets, one based on 44
Phylogeny genes for 161 species, and one based on 12 genes for 4161 species. The likelihood-based tree from the
Squamata combined matrix (52 genes, 4162 species) shared more higher-level clades with the 44-gene tree (90%
Supermatrix vs. 77% shared). Branch support for higher level-relationships was marginally higher than in the
12-gene tree, but lower than in the 44-gene tree. Relationships were apparently not obscured by the
abundant missing data (92% overall). We provide a time-calibrated phylogeny based on extensive
sampling of genes and taxa as a resource for comparative studies.
! 2015 Elsevier Inc. All rights reserved.
 Chelydra serpentina
133.7
Podocnemis expansa
264.5 Gallus gallus
106.2
Dromaius novaehollandiae
3 239.0
Crocodylus porosus
33.0
Alligator mississippiensis
1 299.8 Sphenodon punctatus
Dibamidae
Carphodactylidae
72.4
Pygopodidae
76.1
Gekkota Diplodactylidae
2 277.6 4 137.4
Eublepharidae
133.5 Sphaerodactylidae
119.5 Phyllodactylidae
110.3
Gekkonidae
Xantusiidae Cricosaurinae
205.1 98.3 Xantusiinae
53.0
Lepidophyminae
5 157.1 Gerrhosauridae Gerrhosaurinae
59.7
Zonosaurinae

Scincidae
Scincoidea 87.1 Cordylidae Platysaurinae1
170.9 59.7 Platysaurus pungweensis
43.4
Cordylinae
Acontiinae
202.1 Scincidae
13 97.3 Scincinae1

Clade ancien
Scincinae2
84.2
82.3 Ateuchosaurus pellopleurus
72.0
Lygosominae1
75.3
Lygosominae2
Teiidae Tupinambinae
85.1
Teiinae
94.0
6 Alopoglossinae

Lacertoidea
89.4 Ecpleopinae
Gymnophthalmidae 76.3
Cercosaurinae
79.4
Bachiinae
74.7 Rhachisaurinae
64.9
179.6 Gymnophthalminae
196.9 Rhineuridae
146.6 Bipedidae
91.0 Cadeidae
75.0
Blanidae
87.0
7 153.8 Trogonophiidae
80.5
Amphisbaenidae
Lacertidae Gallotiinae
85.1
Lacertinae
99.6
Helodermatidae
12 Xenosauridae
10 94.4 Diploglossinae
68.1 Anniellidae
Anguimorpha
114.1 61.0 Anguinae
43.3
Gerrhonotinae
Shinisauridae
188.3 9 104.5 Lanthanotidae
64.4
Varanidae
182.0 Chamaeleonidae Brookesiinae
62.0
Acrodonta Chamaeleoninae
126.1
Uromastycinae
119.8
Leiolepidinae
Agamidae Hydrosaurinae
116.2 95.8
Amphibolurinae
11 168.2 105.6
Iguania Agaminae
100.7
Draconinae
Leiocephalidae
Pleurodonta Iguanidae
91.3 Hoplocercidae
81.8
87.2
Corytophanidae
76.5
Crotaphytidae
184.6 Tropiduridae
85.7
Opluridae
71.3 Leiosauridae Enyaliinae
84.7 34.3
80.1 Leiosaurinae
Liolaemidae
82.7
Phrynosomatidae
81.4
Polychrotidae
78.1
Dactyloidae
Leptotyphlopidae
122.7 Gerrhopilidae
89.0
Xenotyphlopidae
85.4
Typhlopidae
Serpentes Anomalepididae
128.1 Aniliidae
79.8
Tropidophiidae
Cylindrophis ruffus
124.7 29.2

Pythonidae
Cylindrophis maculatus
24.7
44.4 Anomochilidae
Uropeltidae
Xenopeltidae
8 67.7
52.4
92.7 Loxocemidae
33.4

Clade récent
Pythonidae
62.9 Xenophidiidae
48.5
Bolyeriidae
60.4 Calabariidae
45.4
87.5 Sanziniinae
44.1
Boidae Ungaliophiinae
41.3
Candoiinae
40.4
Erycinae
39.6
Boinae
Acrochordidae
Xenodermatidae
80.6 Pareatidae
75.2 Viperinae
Viperidae 42.8
Azemiopinae
37.2
64.2 Crotalinae
Homalopsidae
61.4
Prosymninae
45.4
Psammophiinae
46.8 Atractaspidinae
36.1
Aparallactinae
44.5
Pseudaspidinae
54.8 47.9 43.5
Lamprophiinae
42.3
Pseudoxyrhophiinae
Micrelaps bicoloratus
46.7
Oxyrhabdium leporinum
44.2
53.0 Elapidae
Sibynophiinae

Molecular Phylogenetics and

47.1
Grayiinae
42.1
44.9
Colubrinae1
Colubrinae2
48.7
Colubridae Calamariinae
43.4
Pseudoxenodontinae

Evolution (2016) 94: 537-547

47.1
47.7
Natricinae
Dipsadinae
 Scincidae (171 ma)
• Amérique du nord

• Amérique du sud

• Europe

• Asie

• Afrique

• Australie
 Pythonidae (33 ma)

• Asie

• Afrique

• Australie
Impacts of the Cretaceous Terrestrial (BP
S4).
men
Revolution and KPg Extinction on to S
~30

Mammal Diversification disa

mam
1 2 1 3
carn
Robert W. Meredith, * Jan E. Janečka, * John Gatesy, Oliver A. Ryder, clus
2 4 4 5
Colleen A. Fisher, Emma C. Teeling, Alisha Goodbla, Eduardo Eizirik, on t
5 6 7 8
Taiz L. L. Simão, Tanja Stadler, Daniel L. Rabosky, Rodney L. Honeycutt, upo
9,10 9 3 11
John J. Flynn, Colleen M. Ingram, Cynthia Steiner, Tiffani L. Williams, acte
12 1,13 14
Terence J. Robinson, Angela Burk-Herrick, Michael Westerman, earl
1,15 1 2
Nadia A. Ayoub, Mark S. Springer, †‡ William J. Murphy †‡ R
(18,
Previous analyses of relations, divergence times, and diversification patterns among extant sup
mammalian families have relied on supertree methods and local molecular clocks. We constructed to re
a molecular supermatrix for mammalian families and analyzed these data with likelihood-based niot
methods and relaxed molecular clocks. Phylogenetic analyses resulted in a robust phylogeny with to S
better resolution than phylogenies from supertree methods. Relaxed clock analyses support the crep
long-fuse model of diversification and highlight the importance of including multiple fossil the s
calibrations that are spread across the tree. Molecular time trees and diversification analyses line
suggest important roles for the Cretaceous Terrestrial Revolution and Cretaceous-Paleogene (KPg) and
mass extinction in opening up ecospace that promoted interordinal and intraordinal diversification, met
respectively. By contrast, diversification analyses provide no support for the hypothesis concerning whe
the delayed rise of present-day mammals during the Eocene Period. ofte
Science (2011) 334: 521-524attra
 Soricidae
Clade ancien

Downloaded from http://science.sciencemag.org/ on July 11, 2018

Muridae
Clade récent

Lemuridae
Clade récent

Science (2011)
334: 521-524
 Soricidae (70 ma)

• Amérique du nord

• Amérique du sud

• Europe

• Asie

• Afrique
 Lemuridae (30 ma)

• Madagascar
 Muridae (25 ma)
• Amérique du nord

• Amérique du sud

• Europe

• Asie

• Afrique

• Australie
Dispersion
Ecological Monographs, 59(2), 1989, pp. 59-123
© 1989 by the Ecological Society of America

PLANT RECOLONIZATION AND VEGETATION

SUCCESSION ON THE KRAKATAU ISLANDS, INDONESIA'
R. J. WHITTAKER
School of Geography, University of Oxford, Mansfield Road, Oxford, OXJ 3TB, United Kingdom
M. B. BusH 2
Department of Geography, University of Hull, Cottingham Road, Hull, HU6 7RX. United Kingdom
AND

K. RICHARDS
Robertson Research International, Llandudno, Gwynedd, LL30 JSA, United Kingdom

Abstract. The development of the vegetation and floras of the Krakatau Islands in the Sunda Straits,
Indonesia, since their "sterilization" in 1883 is described. Key features of the post-1883 environment, such as
pedogenesis, geomorphology (coastal change), human influence, and recent volcanic activity are detailed, and
their possible influence on spatial and temporal patterns in both vegetation and flora is discussed.
Field work conducted in 1979, 1983, and 1984 has enabled an assessment of the present state of vegetation
development, aided by plot-based sampling and analysis of the arboreal component, employing numerical
classification (by TWINSPAN) and ordination (by DCA). The coastal communities were established early and
have remained little changed, although the distribution of the various components has changed according to
the influence of a dynamic coastal geomorphology. Fifty years after forest closure, the forests of the interiors
remain species-poor and composed of typical early-sera! species. Rakata was dominated inland by Neonauclea
ú =550 m altitude, above which a mossy forest of Ficus spp. and Schejjlera
 Colonisation
400

Ncumulatif
Depuis l’irruption
300
Nombre d’espèces (N)

200

Nrecensées
Lors de l’échantillonnage
100

0
1850 1875 1900 1925 1950 1975 2000
Ecological Monographs (1989) 59: 59-123
 Colonisation

• Immigration (I) : arrivée de nouvelles espèces sur l’île

• Extirpation (E) : disparition d’espèces de l’île

• Nc, t2 = Nc, t1 + I

• Nr, t2 = Nr, t1 + I - E
 Colonisation
400

Changement : ΔN
Taux de changement : ΔN / ΔT
300
Nombre d’espèces (N)

ΔN

200 ΔT

100

0
1850 1875 1900 1925 1950 1975 2000
Ecological Monographs (1989) 59: 59-123
 Devoir
N N
Année ΔT I E I / ΔT E / ΔT
recensées cumulatif

1883 0 0

1886 17 17

1897 39 47

1921 165 179

1931 173 209

1991 236 324

 Devoir
• Complétez le tableau

• Faites des graphiques des taux d’immigration et des taux

d’extirpation en fonction du temps

• Proposez deux hypothèses plausibles pour expliquer (1)

la variation du taux d’immigration en fonction du temps et
(2) la variation du taux d’extirpation en fonction du temps

• À remettre le
Niche écologique
 Niche d’habitat

• Joseph Grinnell (1917)

• Tout ce qui conditionne

l'existence d'une espèce à un
endroit donné, incluant les
facteurs abiotiques (température,
humidité, précipitations) et
biotiques (présence de nourriture,
compétiteurs, prédateurs, abris,
etc.)
 Niche fonctionnelle

• Charles Elton (1927)

• Place occupée par une espèce

dans les chaînes trophiques
(carnivore, herbivore, etc.)
Niche multi-dimensionnelle

• Evelyn Hutchinson (1957)

• Hypervolume où chaque
dimension de l'espace représente
une ressource (alimentaire, en
matériaux, spatiale, offre en
cachette, substrats ou perchoirs,
etc.) ou une condition
(température, précipitations,
acidité, etc) de l’environnement
Niche multi-dimensionnelle

• Hutchinson fait la distinction entre la niche fondamentale

et la niche réalisée

• Niche fondamentale : tous les composants et toutes les

conditions environnementales nécessaires à l'existence
d'un organisme

• Niche réalisée : comprise dans la niche fondamentale,

mais réduite à l'espace que l’espèce est contrainte
d’occuper du fait de la compétition avec les autres
espèces (ou à l’incapacité à se disperser)
 Principales dimensions de
la niche écologique
• Espace

• Temps

• Nourriture

• Température (?)

• Précipitation, pH, abri, site de ponte, site d’hibernation,

etc.
Dimensions de la niche
Densité

Dimension
Dimensions de la niche

Dimension 1 Dimension 2
Dimension 2

Dimension 1
Dimensions de la niche

Dimension 1 Dimension 2 Dimension 3

3
i on
n s
e
Dimension 2

im
D

Dimension 1
Jusqu’à n dimensions
Généralistes et spécialistes
Densité

Dimension
Généralistes et spécialistes
Dimension 2

Dimension 1
Généralistes et spécialistes
Dimension 2

Dimension 1
Généralistes et spécialistes
Dimension 2

Dimension 1
Pourquoi la niche réalisée
≠ niche fondamentale?
Barrières à la dispersion
 Compétition

Dimension 2
Dimension 2

Allopatrie

Dimension 1 Dimension 1
Dimension 2

Sympatrie

Dimension 1

REVIEW AND
SYNTHESIS
Miguel B. Arau !jo,1,2,3,*a Francisco
~ez,1a Francisco Bozinovic,4
Ferri-Ya!n
Pablo A. Marquet,4,5,6 Fernando
Valladares1 and Steven L. Chown7
INTRODUCTION
Ecology Letters, (2013) 16: 1206–1219 doi: 10.1111/ele.12155
Heat freezes niche evolution
Abstract
Climate change is altering phenology and distributions of many species and further changes are projected.
Can species physiologically adapt to climate warming? We analyse thermal tolerances of a large number of
terrestrial ectotherm (n = 697), endotherm (n = 227) and plant (n = 1816) species worldwide, and show
that tolerance to heat is largely conserved across lineages, while tolerance to cold varies between and within
species. This pattern, previously documented for ectotherms, is apparent for this group and for endotherms
and plants, challenging the longstanding view that physiological tolerances of species change continuously
across climatic gradients. An alternative view is proposed in which the thermal component of climatic
niches would overlap across species more than expected. We argue that hard physiological boundaries exist
that constrain evolution of tolerances of terrestrial organisms to high temperatures. In contrast, evolution
of tolerances to cold should be more frequent. One consequence of conservatism of upper thermal toler-
ances is that estimated niches for cold-adapted species will tend to underestimate their upper thermal limits,
thereby potentially inflating assessments of risk from climate change. In contrast, species whose climatic
preferences are close to their upper thermal limits will unlikely evolve physiological tolerances to increased
heat, thereby being predictably more affected by warming.
Keywords
Bioclimatic envelope models, biological invasions, climate change, CTmax, CTmin, evolutionary rates, lower
thermal tolerance, niche conservatism, species distributions, thermal adaptation, upper thermal tolerance.
Ecology Letters (2013) 16: 1206–1219
differences among various traits in the scope of their variation (e.g.
Kellermann et al. 2012a,b).
 (a)

(b)

(c)

Figure 7 (a) Response curves estimated with bioclimatic envelope models (logistic regression) relating species distributions of three species of Liolaemus lizards in central
Chile (see panel c) against mean annual air environmental temperatures (from Hijmans et al. 2005) (i.e. characterization of the realized niche for mean annual
temperature); (b) empirical performance curves for the same species of Liolaemus (from F. Ferri-Y!a~
nez, unpublished data) measured with sprint speed in laboratory
conditions against body temperature (i.e. characterization of the fundamental niche for body temperature); and (c) geographical distributions of the three species of
Liolaemus (from P.A. Marquet and C. Gar!ın, unpublished data) overlaid on mean annual temperature.
Ecology Letters (2013) 16: 1206-1219
Oecologia (2017) 183:337–345
DOI 10.1007/s00442-016-3762-7

CONCEPTS, REVIEWS AND SYNTHESES

Do ectotherms partition thermal resources? We still do not know

James E. Paterson1 · Gabriel Blouin-Demers1

Received: 19 November 2015 / Accepted: 26 October 2016 / Published online: 15 November 2016
© Springer-Verlag Berlin Heidelberg 2016

Abstract Partitioning of the niche space is a mechanism of the thermal resource partitioning hypothesis are required
used to explain the coexistence of similar species. Ecto- before we can assess whether it is widespread in communi-
 Oecologia (2017) 183:337–345

or
s
ces

Oecologia (2017) 183: 337-345

 Annu. Rev. Ecol. Evol. Syst. 2004. 35:113–47
doi: 10.1146/annurev.ecolsys.35.021103.105725
Copyright ⃝ c 2004 by Annual Reviews. All rights reserved
First published online as a Review in Advance on June 10, 2004

ECOLOGICAL IMPACTS OF DEER OVERABUNDANCE

Steeve D. Côté,1 Thomas P. Rooney,2 Jean-Pierre Tremblay,1
Christian Dussault,1 and Donald M. Waller2
1
Chaire de Recherche Industrielle CRSNG-Produits forestiers Anticosti, Département de
Biologie and Centre d’études nordiques, Université Laval, Québec G1K 7P4, Canada;
email: steeve.cote@bio.ulaval.ca, jean-pierre.tremblay@bio.ulaval.ca,
christian.dussault@fapaq.gouv.qc.ca
/18. For personal use only.

2
Department of Botany, University of Wisconsin, Madison, Wisconsin 53706;
email: tprooney@facstaff.wisc.edu, dmwaller@wisc.edu

Key Words browsing, Cervidae, forest regeneration, herbivory, plant-herbivore

interactions
■ Abstract Deer have expanded their range and increased dramatically in abun-
Espèces exotiques
envahissantes
 eat-
s of
hys-
eas- BIOGEOGRAPHY
23).
pre-
pen-
s of
The dispersal of alien species
sen- redefines biogeography in
the Anthropocene
per-
for
im-
and
César Capinha,1,2* Franz Essl,3 Hanno Seebens,4
hing
Dietmar Moser,3 Henrique Miguel Pereira1,5,6
con-
nvi-
It has been argued that globalization in human-mediated dispersal of species breaks down
more
biogeographic boundaries, yet empirical tests are still missing. We used data on native and
ting
alien ranges of terrestrial gastropods to analyze dissimilarities in species composition among
56 globally distributed regions. We found that native ranges confirm the traditional
biogeographic realms, reflecting natural dispersal limitations. However, the distributions of
gastropods after human transport are primarily explained by the prevailing climate and, to
a smaller extent, by distance and trade relationships. Our findings show that human-mediated
25 dispersal is causing a breakdown of biogeographic barriers, and that climate and to some
extent socioeconomic relationships will define biogeography in an era of global change.
4

, he reduced similarity in species composi- range of natural dispersal (4). Notwithstanding,

tion between distant locations is one of human trade and travel have been transgressing
t,
the most noticeable patterns in nature (1–3). natural barriers to dispersal(2015)
Science (5), and348:
increasing
1248-1251
LETTER doi:10.1038/nature14910

Global exchange and accumulation of

non-native plants
Mark van Kleunen1, Wayne Dawson1, Franz Essl2, Jan Pergl3, Marten Winter4, Ewald Weber5, Holger Kreft6, Patrick Weigelt6,
John Kartesz7, Misako Nishino7, Liubov A. Antonova8, Julie F. Barcelona9, Francisco J. Cabezas10, Dairon Cárdenas11,
Juliana Cárdenas-Toro12,13, Nicolás Castaño11, Eduardo Chacón2,14, Cyrille Chatelain15, Aleksandr L. Ebel16, Estrela Figueiredo17,18,
Nicol Fuentes19, Quentin J. Groom20, Lesley Henderson21, Inderjit22, Andrey Kupriyanov23, Silvana Masciadri24,25, Jan Meerman26,
Olga Morozova27, Dietmar Moser2, Daniel L. Nickrent28, Annette Patzelt29, Pieter B. Pelser9, Marı́a P. Baptiste12, Manop Poopath30,
Maria Schulze31, Hanno Seebens32, Wen-sheng Shu33, Jacob Thomas34, Mauricio Velayos10, Jan J. Wieringa35,36 & Petr Pyšek3,37,38

All around the globe, humans have greatly altered the abiotic and
biotic environment with ever-increasing speed. One defining fea-
ture of the Anthropocene epoch1,2 is the erosion of biogeographical
barriers by human-mediated dispersal of species into new regions,
where they can naturalize and cause ecological, economic and
social damage3. So far, no comprehensive analysis of the global
accumulation and exchange of alien plant species between conti-
nents has been performed, primarily because of a lack of data. Here
we bridge this knowledge gap by using a unique global database on
the occurrences of naturalized alien plant species in 481 mainland
and 362 island regions. In total, 13,168 plant species, correspond-
ing to 3.9% of the extant global vascular flora, or approximately the
size of the native European flora, have become naturalized some-
where on the globe as a result of human activity. North America has
accumulated the largest number of naturalized species, whereas the
Pacific Islands show the fastest increase in species numbers with
respect to their land area. Continents in the Northern Hemisphere
have been the major donors of naturalized alien species to all other
continents. Our results quantify for the first time the extent of
plant naturalizations worldwide, and illustrate the urgent need
for globally integrated efforts to control, manage and understand
the spread of alien species.
The magnitude of impacts caused by alien species on native biota
and human societies is increasing rapidly3. However, our knowledge of
the global spread and distribution of naturalized species (that is, alien
4,5
superiority6,7. It has also been suggested that islands have more alien
species than mainland areas, among others because of unfilled niche
space on islands7,8 or, as shown for birds, a higher introduction effort9.
Although these hypotheses have been tested for some parts of the
world9,10, global tests are still lacking.
Scientific and societal concerns about alien species have led to
improved documentation of their distributions, and inventories have
become available for many regions11. Many of these inventories are
still incomplete, especially for megadiverse taxonomic groups that are
difficult to survey, such as invertebrates and microorganisms, and for
less well-surveyed regions. However, vascular plants are well docu-
mented because of long histories of exploration. Recently, there have
been several major efforts to combine inventories of alien species for
large geographical regions (for example, Delivering Alien Invasive
Species Inventories for Europe (DAISIE; http://www.europe-aliens.
org/)) and for those considered to be the most problematic invaders
globally12. However, a global database of the distribution of all natur-
alized alien plant species had not yet been built. Such data are essential
for understanding global naturalization patterns and their underlying
processes, reporting biodiversity status in terms of essential biodiver-
sity variables13, and informing environmental managers across polit-
ical borders via early warning systems.
Here, we present an analysis of naturalized vascular plant species in
843 non-overlapping regions (countries, federal states, islands) cover-
ing ,83% of the Earth’s land surfaceNature (Fig. 1).(2015)
We used a525: 100-103
novel data-
 a

Pacific Island

b 6,0
0 1,753
Number of naturalized species

Figure 1 | Naturalized vascular plant species in the 843 regions covered by 5,0

Cumulative number of species

the GloNAF database. The heat-map colours correspond to the number of
naturalized species in each of the regions (including 362 island regions). Areas
4,0
permanently covered by ice sheets are indicated in hatched cyan. Grey areas
indicate regions lacking naturalized plant data. To allow comparisons between
the sizes of the GloNAF regions, we used a Mollweide equal-area projection. 3,0
However, to increase the visibility of small islands and island groups on the
map, they are represented by circles.
2,0
We found that at least 13,168 vascular plant species have become
Nature (2015) 525: 100-103
WALDO SWIEGERS/BLOOMBERG VIA GETT

Fall armyworm caterpillars are devastating crops in Africa.

AG RICULTU RE

Invasive pest hits Afric

Fall armyworm caterpillars are devastating crops in Africa.

AG R I C U LT U R E

Invasive pest hits Africa

As hungry caterpillar eats its way through 12 countries, researchers be
As hungry caterpillar eats its way through 12 countries, researchers begin to fight back.
BBYYS ASRAAR
H AWH
I L DW I L D response.
response. Sixteen countries Sixteen
agreed to urgentcountries agreed
January 2016, tosince
and has urgent
moved toJanuary
at least 2016
planscapacity
plans to boost the region’s to boost the region’s
to manage Nature
capacity
12 countries on theto (2017)
manage
continent, 543: 7 of13-14
12 countries
reaching
Ecology (1987) 68: 660-668
Perte totale : 10 oiseaux
endémiques en 15 ans

Ecology (1987) 68: 660-668

Science (2013) 342: 1166-1167
PNAS (2012) 109: 2418-2422
PNAS (2012) 109: 2418-2422
PNAS (2012) 109: 2418-2422
management of invasive predators on islands should be a global
conservation priority. Understanding and mitigating the impact of
invasive mammalian predators is essential for reducing the rate of
global biodiversity loss.
extinction | feral cat | island | invasive mammal | trophic cascade
of either 0.25 (secondary cause of species decline), 0.75 (primary
cause of species decline), or 1.0 (species extinction attributed to
the predator), and we weighted these values by the strength of
evidence available, drawing on a total of 996 supporting references
(Methods). The severity of predator impacts and the strength of
evidence supporting them [the inverse of the width of confidence
intervals (CIs)] was higher for bird and mammal species compared

ECOLOGY
IInvasive predators and global biodiversity loss
nvasive mammalian predators (“invasive predators” hereafter)
are arguably the most damaging group of alien animal species
with reptile species (Fig. 1).
Rodents are linked to the extinction of 75 species (52 bird, 21
for global biodiversity (1–3). Species such as cats (Felis catus), mammal, and 2 reptile species; 30% of all extinctions) and cats
a,b,1
rats
Tim(Rattus rattus),
S. Doherty mongoose Glenc, Dale
, Alistair S.(Herpestes G. Nimmod, Euan
auropunctatus), and G.toRitchie a
, and Chris
63 extinctions (40,R. 21,
Dickman e
and 2 species, respectively; 26%)
stoats
a
(Mustela erminea) threaten biodiversity through predation whereas red foxes, dogs (Canis familiaris), pigs (Sus scrofa), and
Centre for Integrative Ecology, School of Life and Environmental Sciences, Deakin University, Geelong, VIC 3216, Australia; bCentre for Ecosystem
(4,Management,
5), competition (6), disease transmission (7), and facilitation
School of Natural Sciences, Edith Cowan University, Joondalup, WA small Indian
6027, mongoose
Australia; c
Landcare (H. auropunctatus)
Research, are implicated
Auckland 1072, New Zealand; in 9–11
with
d otherforinvasive
Institute species
Land, Water (8). The
and Society, decline
School and extinction
of Environmental Science,of extinctions
Charles eachAlbury,
Sturt University, (Fig. NSW
2). For2640,all threatened
Australia; e
and extinct
and Desert Ecology species
native
Researchspecies
Group,due to of
School invasive
Life and predators
Environmental canSciences,
have impacts
Universitythat
of Sydney, Sydney, NSW
combined, cats2006,
andAustralia
rodents threaten similar numbers of species
cascade throughout entire ecosystems (9). For example, pre- (430 and 420 species, respectively), followed by dogs (156 spe-
Edited by Daniel S. Simberloff, The University of Tennessee, Knoxville, TN, and approved July 20, 2016 (received for review February 12, 2016)
dation by feral cats and red foxes (Vulpes vulpes) has led to the cies), pigs (140 species), mongoose (83 species), red foxes (48
decline
Invasive or species
extinction of twobiodiversity
threaten thirds of Australia’s
globally, and digging mammal
invasive mam- include three
species), stoatscanids, seven (Fig.
(30 species) mustelids,
2), andfive
therodents,
remaining twopredators
procyo-
species
malianover the past
predators 200 y (10,damaging,
are particularly 11). Reduced havingdisturbance
contributed to to nids, three viverrids, two primates, two marsupials, two mon-
topsoil in the absence
considerable species of digging
decline mammals
and has We
extinction. led to impoverished
provide a global gooses, and single representatives from four other families, with
Significance
landscapes
metaanalysis where little impacts
of these organicand matter
revealaccumulates and rates
their full extent. of
Invasive 60% from the order Carnivora (Table S1). The 738 impacted
seed germination
predators are low (10).
are implicated in 87Inbird,
the 45Aleutian
mammal, archipelago, pre-
and 10 reptile species consist of 400 bird species from 78 families, 189 mammal
Invasive mammalian predators are arguably the most damag-
dation
species of extinctions—58%
seabirds by introduced Arctic
of these foxes contemporary
groups’ (Alopex lagopus) has
extinc- species from 45 families, and 149 reptile species from 26 families
ing group of alien animal species for global biodiversity. Thirty
lowered nutrient input
tions worldwide. Theseand soil fertility,
figures are likelyultimately
underestimatedcausingbecause
vege- (Dataset S1). Invasive mammalian predators emerge as causal
23 critically endangered species that we assessed are classed as
species of invasive predator are implicated in the extinction or
tation to transform from grasslands to dwarf shrub/forb-dominated factors in the extinction of 87 bird, 45 mammal, and 10 reptile
extinct.” Invasive mammalian predators endanger a fur- endangerment of 738 vertebrate species—collectively contrib-
systems
“possibly (12). species, which equates to 58% of modern bird, mammal, and reptile
ther 596 species at risk of extinction, with cats, rodents, pred-
dogs, uting to 58% of all bird, mammal, and reptile extinctions. Cats,
Mitigating the negative impacts of invasive mammalian species extinctions globally (including those species classed as “ex-
and ispigs threatening rodents, dogs, and pigs have the most pervasive impacts, and
ators a primary goal the most species agencies
of conservation overall. Species most(1,
worldwide at 13,
risk tinct in the wild”). Invasive predators also threaten 596 species
endemic island faunas are most vulnerable to invasive preda-
from
14). predators there
Regardless, have highremainsevolutionary
no globaldistinctiveness
synthesis of the and role
inhabit
of classed as “vulnerable” (217 species), “endangered” (223), or “criti-
insular environments. Invasivedeclines
mammalian tors. That most impacted species are insular indicates that
invasive predators in species andpredators
extinctions are therefore
(but see cally endangered” (156), of which 23 are classed as “possibly extinct.”
management of invasive predators on islands should be a
important
refs. 3 and 15). drivers
Here, ofwe irreversible
quantify the lossnumber
of phylogenetic diversity
of bird, mammal, To assess the comparative severity of predator impacts, we
worldwide. That most impacted species are insular indicates that global conservation priority. Understanding and mitigating the
and reptile species threatened by, or thought to have become assigned each of 1,439 predator-threatened species cases a value
management of invasive predators on islands should be a global impact of invasive mammalian predators is essential for re-
extinct (since AD 1500) due to, invasive mammalian predators. of either 0.25 (secondary cause of species decline), 0.75 (primary
conservation priority. Understanding and mitigating the impact of ducing the rate of global biodiversity loss.
We use metaanalysis to examine taxonomic and geographic cause of species decline), or 1.0 (species extinction attributed to
invasive mammalian predators is essential for reducing the rate of the predator), and we weighted these values by the strength of
trends in these impacts and show how the severity of predator Author contributions: T.S.D., A.S.G., D.G.N., E.G.R., and C.R.D. designed research; T.S.D.
global biodiversity loss. evidence available, drawing
impacts varies according to species endemicity and evolutionary and A.S.G. performed research; T.S.D.on a total
analyzed ofand
data; 996T.S.D.,
supporting references
A.S.G., D.G.N., E.G.R., and
distinctiveness. (Methods). The
C.R.D. wrote the paper. severity of predator impacts and the strength of
extinction | feral cat | island | invasive mammal | trophic cascade
evidence
The supporting
authors declare them
no conflict [the inverse of the width of confidence
of interest.
Results and Discussion intervals
article is(CIs)]
a PNASwas higher for bird and mammal species compared

ECOLOGY
I
This Direct Submission.
nvasive
In total, 596mammalian
threatenedpredators (“invasive
and 142 extinct predators”
species hereafter)
(total 738) have 1with reptile species (Fig. 1).
To whom correspondence should be addressed. Email: tim.doherty.0@gmail.com.
are arguably the most damaging group of
suffered negative impacts from 30 species of invasive mammalian alien animal species Rodents are linked to the extinction of 75 species (52 bird, 21
This article contains supporting information online at www.pnas.org/lookup/suppl/doi:10.
for globalfrom biodiversity (1–3).andSpecies
eightsuch as cats (Felisspecies
catus), 1073/pnas.1602480113/-/DCSupplemental.
mammal, and 2 reptile species; 30% of all extinctions) and cats
to 63 extinctions (40,PNAS 21, and (2016)
2 species, 113: 11261-11265
predators 13 families orders. These
rats (Rattus rattus), mongoose (Herpestes auropunctatus), and respectively; 26%)
Zealand (15), and Hawaii (11), with the remaining regions having
0–7 species extinctions each (Fig. 4). The taxonomy of impacted
species varied among regions, with the highest numbers of im-
pacted mammal species occurring in Australia and Central
America, and most of the impacted reptile species occurring in
Micro-/Mela-/Polynesia and Central America (Fig. 4). Most im-
pacted bird species are in Micro-/Mela-/Polynesia, New Zealand,
the Madagascar region, Central America, and Hawaii (Fig. 4).
Insular endemics accounted for 87% of extinct species (124
species) and 81% of the sum of all threatened/extinct species
(601 species). The proportions of total threatened/extinct species
that were insular endemics varied between taxonomic classes
species (23), and the red fox, along with the feral cat, is an im-
portant driver of Australian mammal species extinctions (11).
Fewer reptile species were negatively affected by invasive
mammalian predators, compared with bird and mammal species.
Reptiles also had a lower average impact score, which may be
because reptiles are less studied than birds and mammals (9),
with only 40% of the world’s reptiles having been assessed for the
Red List thus far (compared with ∼99% for birds and mammals)
(24). Further insights will likely emerge once the conservation
status of most reptiles has been determined. Detailed studies
from individual regions nonetheless demonstrate that invasive
predators can have severe impacts on local reptile assemblages

300
and threatened species
Number of extinct

200

100

0
B M R B M R B M R B M R B M R B M R B M R

Fig. 2. Numbers of threatened and extinct bird (B), mammal (M), and reptile (R) species negatively affected by invasive mammalian predators. Gray bars are
the total number of extinct and threatened species, and red bars are extinct species (including those classed as extinct in the wild). Predators affecting <15
species are not shown here. Predators (L to R) are the cat, rodents, dog, pig, small Indian mongoose, red fox, and stoat.

11262 | www.pnas.org/cgi/doi/10.1073/pnas.1602480113 Doherty et al.

PNAS (2016) 113: 11261-11265

Applications
PNAS (2014) 111: 10233-10238
Fig. 1. Projections of the native (red points) and Australian (blue points) ranges of R. marina onto the first two axes of a PCA. Red and blue contours
represent the climatic conditions available in the two ranges, whereas the gray contour represents climates within the species’ native fundamental niche.
Maps illustrate how R. marina has filled its geographically projected fundamental niche (gray shading) over time. The correlation circle in the middle of the
figure indicates the relative importance of each variable on the PCA axes. See Materials and Methods for variable descriptions.

10234 | www.pnas.org/cgi/doi/10.1073/pnas.1405766111 PNAS (2014) 111: 10233-10238 Tingley et al.

 org
A
in
management of range shifts and, t izat
development of more effective ma tion
(Qu
face of climate change and increasin Par
For example, detecting realized nich
Fig. 3. Prediction of the fundamental niche (potential number of breeding
months per year) of R. marina according to a mechanistic model. Predictions
are

are depicted in 10 equal potential

interval classes,biocontrol
with the highest agents for9–invas
class depicting Clim
fying fundamental niche shifts
12 breeding months per year and the white area representing no breeding
months per year. White dots represent occurrence records of R. marina,
cou
dat
199
ability
whereas black dots represent of records
occurrence native of aspecies
congener (R.to adapt totemc
schneideri).
approach
dently assigned to either species.
can be applied to any
Gray dots in Central Brazil lie within a hybrid zone and could not be confi-
taxo
war
of
thus holds great promise for iden qua
proximate causes of niche
Afrotropical, and Indo-Malayan realms than did the correlative
shifts thro
rect
the
model based on native-range data (Fig. S5). Collectively, these results wit
provide evidence for aMaterials and Methods
shift in the realized, as opposed to the fun-
Mea
damental, niche between the species’Data.
Occurrence nativeData
and Australian ranges.distrib
on the native R. m
Why does R. marina fail to fill its fundamental niche in its
squares, 10’ in size) and R. schneideri (n wit =
native range? One possibility is that the presence of a closely com
our own in
related species (R. schneideri) surveys,
cooler HerpNET (www.herpnet.
and drier regions of PCA
formation Facility (www.gbif.org),
Southern Brazil may be preventing R. marina from colonizing speciesLink
sen
suitable environmentsdiversity
south ofand its present range. Indeed,
Environmental ResourcetheseDatabet
S
two species hybridizebz),at the den
thesouthern
World range edge of R.
Biodiversity marina
Information N
ran
(30), and even low rates of interspecific hybridization
remib_ingles/doctos/remibnodosdb.html), can en-
tha
force stable parapatric range boundaries (31). In addition, the
ranges of R. marina andMuseu de História
R. schneideri Natural fill
collectively de the
Sãonative
Paulo. Toovea
sim
fundamental niche ofwe R. excluded
marina (Fig.occurrence records
3), and the thatoc-
climates wereove
w
cupied by R. marina deri and R. (where
in Australia marinathe (30)species
and only included
is closer to rec
rea
filling its fundamental niche)ranges
native are similar
of R.toschneideri
those occupied
and R.bymarinwh
R. schneideri in Southrange
America (Fig. 2A and from
Fig. S3). ove
maps available theHowever,
Global Amph nich
all niche models, whether correlative or mechanistic, should be
org/initiatives/amphibians).
viewed as tools for generating hypotheses and elucidating knowl- ran
grid

edge gaps (32, 33). The Australian

fact that therange data model
mechanistic for R. marina
and the (as100o
correlative model basedin size)
on thewere sourced
Australian fromniche
realized researchers
both a
T
predicted suitable climatic conditions
izations: south of the
FrogWatch, observed native
Department of Envi
nich
range of R. marina suggests
tional thatParksinterspecific hybridization
and Wildlife Service,may Fores
pro
be important; however, future studies could usefully test this wh
(Queensland Environmental Protection not A
hypothesis, using laboratory or field experiments.
PNAS
Parks
Our results demonstrate and
that(2014)
Wildlife 111:
Service.
contrasting 10233-10238
Most ofand
fundamental these the
da