Niche Écologique

Le concept de niche écologique
Importance en écologie et conservation
Pr. Frédéric Médail

frederic.medail@imbe.fr
1
Pourquoi le concept de niche écologique
est-il important?
Niche écologique
Plan
1. Le concept de niche : définitions
2. Evolution et conservatisme de niche
3. Changements climatiques et évolution de niche
4. Importance de la micro-échelle
3
1. Le concept de niche : définitions
4
La niche selon Grinnel
Grinnel (1917)
Zone où la somme des facteurs limitants, abiotiques, permet de maintenir un taux de
croissance positif pour l’espèce
Niche de l’espèce A
+ : présence de l’espèce ; o : absence de l’espèce
Pulliam H.R. 2000. Ecol. Lett., 3.

Définitions
Hutchinson (1957)
• Niche fondamentale
Somme des facteurs limitants pour maintenir un taux de croissance positif,
ignorant les facteurs biotiques
• Niche réalisée
Portion de la niche fondamentale dans laquelle une espèce peut avoir
un taux de croissance positif, malgré les pressions biotiques (compétition,
prédation)
Construction de niche
Elton (1927)
Interaction de l’espèce avec les autres espèces
Dynamiques ressource-consommateur
La niche selon Hutchinson
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Niche réalisée d’un compétiteur B

qui exclut l’espèce A de sites favorables
Niche fondamentale
de l’espèce A
Niche réalisée
de l’espèce A

La niche est un volume

multidimensionnel qui décrit
l’écologie d’une espèce dans un
hyper-espace à n-dimensions
Hutchinson 1957 Cold Spring Harbor Symposia on quantitative Biology

Un modèle qui n’est pas parfait : dynamique source-puits
Si les ressources sont

abondantes (parcelles sources) :
Niche réalisée?
- Production de plus
d’individus que nécessaires
- Dispersion vers d’autres
parcelles (puits) où le taux de
croissance reste positif
uniquement grâce à
l’immigration

Un modèle qui n’est pas parfait : cas d’une dispersion limitée
Une espèce peut être absente

dans des stations favorables si sa
capacité de dispersion est réduite

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Le concept de dualité
-> Décrit la correspondance réciproque entre l’espace de la niche et l’espace physique

(”le biotope”)
• Le biotope constitue
l’environnement physique
de l’espèce
• L’espace de niche: partie

de hyperspace de n-
dimensions dans laquelle
la niche peut être définie
• Chaque observation
d’espèce dans le biotope a
une équivalance dans
l’espace de la niche
• Par contre, l’intégralité de

la niche écologique n’est
pas forcément représentée
dans un biotope donné
Biotope 1
Distribution réalisée de S1
Distribution réalisée de S2

Imaginons un nouveau biotope, où pouvons nous prédire S1 et S2 dans le biotope?
Biotope 2
Temperature Algal food size

Imaginons un nouveau biotiope, où pouvons nous prédire S1 et S2 dans le biotope?
Biotope 2
S2
S1
Temperature Algal food size

Chevauchements de niches entre espèces
Dans tous ces cas :

- Si l’espèce S2 est supérieure, l’espèce S1 est
exclue du chevauchement
- Si l’espèce S1 est supérieure, l’espèce S2 est

exclue du chevauchement
http://www.zo.utexas.edu/courses/bio301/chapters/Chapter13/Chapter13.html
Niche réalisée et niche fondamentale
Modèle théorique de niche fondamentale

(partie pleine et hachurée) et de niche
réalisée (partie hachurée) pour une espèce
G après exclusion compétitive due à six
espèces plus compétitives (A, B, C, D, E, F)
Phénomène de compression de niche
Hypothèse de compression (McArthur et Wilson 1967, The Theory of Island Biogeography):

Elle prédit que la proportion d’habitat utilisé par chaque espèce dans un biotope décroit
lorsque l’intensité de la compétition augmente
Niche réalisée, abondance et aire de répartition
Cas rare :
Abondances faibles sur les marges de l’aire
et forte au centre de la niche réalisée / aire
de répartition
Cas plus commun :

Abondances faibles sur les marges de l’aire,
et à la fois faibles et fortes au centre de la
niche réalisée / aire de répartition
Gaston 2003, The structure and dynamics of geographic ranges

Relation entre la niche de l’espèce (ou du clade)
Discussion
et la niche des populations (ou des taxons du clade)
Plusieurs populations d’une

même espèce peuvent avoir
des niches qui se superposent
à des degrés divers
Figure 6-2. Relationship between population niche and species niche.

Several populations of a single species might have niches that overlap in different extent. Reproduced from Holt (2009).
Niche climatique des populations
au centre ou en périphérie de
l’aire de distribution
Merendera filifolia Narcissus dubius
Polygala rupestris Viola arborescens
Climatic niche of 11 Mediterranean plants on

the first two axes of a Principal Component
Analysis of the Mediterranean climate in the
Western Mediterranean basin.
The blue line represents the climatic limits of the
western
Figure Mediterranean
1. Distribution maps of thebasin.
11 studied species in the western Medit
survey
Theinredthe polygon
central partrepresents
of a species distribution,
the climatic black stars are
niche of peripheral
species from de Bolòs and Vigo (2015).
central populations.
The
soil, blackmoss,
lichen, polygon represents
herbaceous the climatic
and woody niche
litter, and of Eute
living
peripheral
plants. populations.
Each contacted plant species was identified. When of w
several
Bluecomponents
area symbolizewere touched at a given point,
the intersection betweenwe con-
the rated
strained the value of the contact point to 1, so that the total ues i
twopertypes
cover of niches
quadrat could not exceed 100%. For each quadrat Wate
we calculated species richness (the number of ‘contacted’ spe- lost
cies), the cover of each biological type (Raunkiaer 1934) and tion
the Hill number associated with Papuga G., 2016. PhD
the Shannon Thesis index
diversity Univ. Sassari.
of w
Définir la niche d’une espèce : dimensions temporelle et spatiale
Difficulté de définir la niche d’une espèce dans le temps et l’espace
Prise en compte de la dimension temporelle
Prise en compte de la dimension spatiale sur de vastes territoires

Dimension temporelle à long terme de la niche
Zelkova
Cas de Zelkova sicula:
sicula (Ulmaceae) an emblematic relict...
Site 2 – Contrada Ciranna
Site 1 - Bosco Pisano
0 100 m
0 100 m
• Current distribution: 2 single, isolated
populations, some hundred trees in total
• Area: 0.5 ha (site 1), 0.8 ha (site 2)
• Habit: shrub to small tree
• Habitat: sparse, disturbed sclerophyllous
forests
• Conservation: critical G. Garfi et al., ined, IUCN Symosium Malte, nov. 2018
Extinct from
La communauté continental
d’espèces Europe
forestières associées at il31000
à Zelkova y a 100 000BP
ans
Valle di Castiglione EXTINCTION

Fraxinus
Age Corylus
ka Carpinus Tilia Ulmus Quercus Lake of Pergusa (Sicily)
Fagus Acer Hedera deciduous
3.5
50
80
100
130
150
ZELKOVA
200
(Sadori et al., 2008)

(Follieri et al., 1988)
Sitedeselection
Prise en compte criteria
la niche temporelle for
de Zelkova danstranslocation
des opérations de translocation
Madonie Mts. – Bosco Pomieri (1341 m a.s.l.) Nebrodi Mts. – Bosco Tassita (1309 m a.s.l.)
• Paleobotanical data (past floristic
assemblages)
• Biogeography of the genus Zelkova
• Elevation (500 to 1340 m a.s.l.)
• Micro-topography (streamsides,
northern face)
• Protected areas
• Public property
SUPRA-MEDITERRANEAN BELT SUPRA-MEDITERRANEAN BELT
Quercus deciduous-Fagus forests Fagus-Acer-Taxus forests
Sicani Mts. – Bosco Ficuzza (1014 m a.s.l.) Iblei Mts. – Bosco Pisano-Manco (507 m a.s.l.)
Bosco Ficuzza – Alpe Cucco (1014 m a.s.l.)
(MESO-)SUPRA-MEDITERRANEAN BELT THERMO-MEDITERRANEAN BELT

Quercus deciduous/evergreen-Acer forests Quercus evergreen forests
G. Garfi et al., ined, IUCN Symosium Malte, nov. 2018
Dimension temporelle à court terme de la niche réalisée
Fortes (extrêmes) fluctuations inter-annuellles dans

le cycle de vie d’une plante à bulbe (géophyte) :
Colchicum filifolium dans le Sud de la France
Photos D. Pavon / IMBE
Dimension temporelle de la niche réalisée
Plant persistence along biogeographical gradients

Patterns induced by several selective force that shape the
evolution of plant reproductive traits such as resprouting
ability and propagule persistence in soil
The ca. 1500 geophytes of the Mediterranean Basin

are mostly rare and narrow endemic species; their
persistence by longevity through seeds and storage
organs often reduces their local extinction.
Representation of multiple demographic strategies

of persistence and regeneration of a long-lived species,
and the biological traits promoting them
L = Longevity
VR = Vegetative Reproduction
S = Seeding
Acis nicaeensis (Maritime Alps)
Garcia & Zamora, 2003. Journal of Vegetation Science, 14.
Dimension spatiale de la niche réalisée :
migrations transcontinentales des vertébrés
Nous ne pouvons pas afficher cette image pour l’instant.
Source :
National Geographic
novembre 2010
Dimension spatiale de la niche réalisée : migration régionale

Dimension spatiale de la niche réalisée : migration transcontinentale
Cas du puffin cendré
Suivis Argos durant la période de migration automnale (août 2011 à janvier 2012) (projet CEFE).
Dimension spatiale de la niche réalisée : échelle locale
Cas des fauvettes méditerranéennes
Document de Jacques Blondel
F. à lunettes
F. pitchou
F. mélanocéphale
F. passerinette
F. orphée
F. à tête noire
Comment étudier la niche écologique d’une espèce en pratique ?
Echelles
macro-
écologiques
Echelles
micro-
écologiques
o Estimer l’aire totale de distribution de l’espèce (données actuelles et données historiques)
o Analyser l’analyse écologique la plus précise possible de l’habitat de l’espèce :

- dans les situations de présence actuelle / de présence historique où l’espèce est
non revue ou éteinte,
- dans les situations écologiques a priori similaires aux conditions actuelles mais où
l’espèce est absente
o Dans le cas de vastes distributions ou d’espèces fréquentes, opérer des sélections

aléatoires de mailles qui feront l’objet des analyses écologiques précises pour estimer
l’habitat de l’espèce
o Estimer (si possible !) la magnitude des interactions entre espèces à l’échelle de l’aire de
distribution (notamment entre populations centrales vs. marginales)
o Réaliser un modèle de distribution de l’espèce (Species Distribution Model, SDM)

Distribution du triton (Triturus vulgaris) en Europe

● occurrences post-1970 ❍ occurrences pré-1970
Taxonomy
Comment
Kingdom
étudier
Phylum
la niche
Class
écologique
Order
d’une
Family
espèce en pratique ?
The IUCN Red List of Threatened Species™
Plantae Tracheophyta Liliopsida
ISSN 2307-8235 (online) Asparagales Asparagaceae
IUCN 2008: T15418037A15418040
Scope: Global & Mediterranean
Language: English
Taxon Name: Prospero hierae C.Brullo, Brullo, Giusso, Pavone & Salmeri
Taxonomic Source(s):
The Plant List. 2017. The Plant List. Version 1.1. Available at: http://www.theplantlist.org/.
Prospero hierae
Assessment Information
Assessment by: Lansdown, R.V.
?
Red List Category & Criteria: Vulnerable D2 ver 3.1
Year Published: 2018
Date Assessed: July 29, 2015
Justification:
This species endemic to the small island of Marettimo in Sicily has a restricted distribution, with an Area
of Occupancy (AOO) and Extent of Occurrence (EOO) both estimated at 8 km2. It has been described as
rare, and is threatened due to grazing byView non-native ungulates, however there is no evidence that there
on www.iucnredlist.org
is a current population decline. Threats acting on this species are very likely to impact the entire
population and therefore the number of locations has been estimated at one. Based on this, the species
qualifies as Vulnerable (VU D2) because of its restricted distribution and likelihood that the ongoing
threats may drive the taxon to be listed as CR or EX in a very short time. The species is close to qualifying
forCitation:
Critically
Lansdown,Endangered, however
R.V. 2018. Prospero hierae. The IUCNat
Red present theSpecies
List of Threatened population
2018: is considered to be stable. Urgent
e.T15418037A15418040. http://dx.doi.org/10.2305/IUCN.UK.2018-
population monitoring
1.RLTS.T15418037A15418040.en and habitat status assessment are needed in order to design a conservation plan
forCopyright:
this species.
© 2018 International Union for Conservation of Nature and Natural Resources
Population
Reproduction of this publication for educational or other non-commercial purposes is authorized without prior written
Geographic Range
permission from the copyright holder provided the source is fully acknowledged.
Reproduction of this publication for resale, reposting or other commercial purposes is prohibited without prior written
There
permissionisfrom
nothedetailed
copyright holder.information at present
For further details see Terms of Use. on population trends for this species, however there is no
Range
evidence
The IUCN RedDescription:
that
List there
of Threatened is a
Species™ current
is produced andpopulation
managed by the IUCN decline.
Global Species Programme, the IUCN
Species Survival Commission (SSC) and The IUCN Red List Partnership. The IUCN Red List Partners are: Arizona State
Current
This Botanic Population
Royal species Gardens,isKew; Sapienza Trend:
endemic to the
University Stable
University; BirdLife International; Botanic Gardens Conservation International; Conservation International; NatureServe;
island
of Rome; Texas A&Mof Marettimo
University; and Zoologicalin the
Society Egadi Archipelago, off the west coast of Sicily
of London.
(Brullo eterrors
If you see any al. or2009), Italy.or suggestions on what is shown in this document, please provide us with
have any questions
feedback so that we can correct or extend the information provided.
La flora
flora vascolare
vascolare
dell’Isola dell’Isola di(Arcipelago
di Marettimo Marettimodelle
(Arcipelago delle Egadi,
Egadi, Sicilia Sicilia occidentale):
occidentale): … … 361 361
La flora
La flora vascolare vascolare
dell’Isola dell’Isola di(Arcipelago
di Marettimo Marettimodelle
(Arcipelago delle Egadi,
Egadi, Sicilia Sicilia occidentale):
occidentale): … … 361 361
lare dell’Isola di Marettimo (Arcipelago delle Egadi, Sicilia occidentale): …

361

La flora vascolare dell’Isola di Marettimo (Arcipelago delle Egadi, Sicilia occidentale): … 361
o

o

Downloaded by [University of Toronto Libraries] at 10:55 03 December 2014
Prospero hierae
(oct.-nov. 2019)
Prospero hierae
(oct.-nov. 2019)


Prospero hierae
(oct.-nov. 2019) Prospero hierae
(oct.-nov. 2019)

Fig. 1 – Ubicazione e principali località dell’Isola di Marettimo.

Fig. 1 – Ubicazione e principali località dell’Isola di Marettimo.
Figure 1.Fig.
Localization on Marettimo
1 – Ubicazione e principali island
località of the eight
Figure
dell’Isola Prospero
di1.Fig.
Localization
Marettimo. hierae populations
on Marettimo
1 – Ubicazione e principali island
observed
località of the eight
dell’Isola
in Prospero
this study
di Marettimo.
(fullpopulations
hierae dots) and observed in this study (full dots) and
of the population of Semaforo quoted by Brullo of theetpopulation
al. (2009)of(empty
Semaforocircle);
quoted thebyProspero
Brullo et sp. population
al. (2009) (emptyofcircle);
Monte the Prospero sp. population of Monte
Falcone that has a dubious taxonomical identity is alsothat
Falcone quoted (empty taxonomical
has a dubious square). identity is also quoted (empty square).
aratteristiche geologiche, bioclimatiche e fitocenotiche Caratteristiche geologiche,(Rock
affiorante bioclimatiche
Outcrop), e fitocenotiche
litosuoli (Lithic affiorante
Xerorthents) (Rock Outcrop), litosuoli (Lithic Xerorthents)
Populations Main location Populations
Caratteristiche
Micro-habitats Main location
geologiche, bioclimatiche
Populations’ Micro-habitats
e fitocenotiche
Estimated affiorante Putative (Rock Populations’
Outcrop), litosuoli Estimated Putative
(Lithic Xerorthents)
Oltre cheCaratteristiche geologiche,
con le caratteristiche bioclimaticheOltre
geomorfologiche e che
e fitocenoticheeconterrele rosse
caratteristiche
affiorante
(Lithic geomorfologiche
(Rock Outcrop),
Rhodoxeralfs). elitosuoli
e terre(Lithicrosse (Lithic Rhodoxeralfs).
Xerorthents)
Oltre on the islandfloristica anddi relevés Laperformed extent number of threats
pografiche Oltre on the islandfloristica
che con lel’originalità
del territorio, caratteristiche topografiche
and
di relevés
geomorfologiche eche
del territorio,
Laperformed
con lel’originalità
caratteristiche
e terreorografica,
dorsale rosse (Lithic geomorfologiche
extent
Rhodoxeralfs).
posta secondo number la
e dorsale
ofe terreorografica,
direttrice
rosse (Lithic
threats Rhodoxeralfs).
posta secondo la direttrice
individuals
Marettimotopografiche
va messa indel territorio, proprio
correlazione l’originalità con floristica
le individuals
NO/SE, di è dominata La dorsale orografica,
da Pizzo Falconeposta(686 secondo
m s.l.m.),lail direttrice
arettimotopografiche
va messa indel territorio, proprio
correlazione l’originalitàcon
complesse floristica
le NO/SE,
1.Marettimo
Between di
vicende Punta è
va La
dominata
messa
paleogeograficheWesterndorsale
in da
part orografica,
Pizzo
correlazione
che hanno Falcone
Rocky posta
(686
sub-halophilous
proprio
inte- con
quale secondo
m s.l.m.),
grassland,
lesi ergeNO/SE, la
il direttrice
on
nell’ambito è dominata 1000-5000
da PizzodiFalcone
di un sistema (686 _m s.l.m.), il
vette superan-
mplesse1.Marettimo
Between
vicende Punta va messa Western
paleogeografiche part
in correlazioneFig. Rocky
che hanno ressato
sub-halophilous
1 –proprio
inte-
Ubicazione
ildiCanale coneand
quale
Libecci
complesse di
grassland,
lesiSicilia
erge
principali
CalaNO/SE,
vicende nell’ambito
località
nel
on
corsoè dominatadi un
dell’Isola
paleogeografiche delle da
di
chePizzo
sistema
small
ultime Marettimo.
pockets
ere
hanno
1000-5000
diFalcone
vette
tiof
spesso
inte- (686
terrasuperan-
rosa _m s.l.m.), il
between
i quale
400 metri
si erge dinell’ambito
quota. I versanti
di un declinano
sistema di ripi-
vette superan-
– Ubicazione
di Libecci eandprincipali
Cala località dell’Isolasmall di Marettimo.
pockets
Sparmaturiof terra rosa between the blocks, alt. 5-10 m (R1-R12)
sato il Canale
complesse di Sicilia
vicendenelpaleogeografiche
corso delle ultime che erehanno
geologiche. ti inte-
ressato spesso
il m i quale
Canale400dimetri siSicilia
erge dinel
quota.corsoI delle
nell’ambito versanti un declinano
di ultime sistema
damenteere di tiripi-
vette
versospesso superan-
la costa,
i 400interrotti da numerose
metri di quota. I versantiincisio-
declinano ripi-
Sparmaturi the blocks, 2. alt.
Punta 5-10
Bassano, (R1-R12) Southern part Rocky halophilous grassland, 100-500 _
ologiche.ressato il Canale di Sicilia nel corso delle L’isolaultime damente
rappresenta
geologiche. ere laverso tiporzione
spessola costa, 400interrotti
iemersa metri
più occiden- da numerose
di quota. I versanti
ni incisio-
torrentizie, declinano
damente il cuiverso ripi-
ruscellamento impetuoso
la costa, interrotti durante
da numerose incisio-
near Puntagrassland,
di Cala between the pebbles and the_
L’isola2.rappresenta
Punta Bassano,
geologiche. Southern
la porzione emersapart più occiden-
Rocky halophilous
tale della Catena
Marino
ni torrentizie,
L’isoladelle Egadi,
damente
rappresenta ilchecui
la aruscellamento
sua volta
verso
porzione la fa parte
costa,
emersa
blocks,
impetuoso
interrotti
più
alt.
occiden-
5 m
100-500
le piogge
da
(R18-R20)
durante
determina
numerose
ni torrentizie, una ilnotevole
incisio- cui erosione e ilimpetuoso
ruscellamento denuda- durante
near Punta di Cala between
Caratteristiche the pebbles
geologiche, and the
bioclimatiche e fitocenotiche affiorante (RockdeterminaOutcrop),
e della Catena L’isoladelle Egadi, che
rappresenta a sua
la porzione volta della
fa parte
emersa più
più ampia
3.tale ledella
occiden- “Thrust
piogge CatenaniBelt”
determinadelle individuata
torrentizie, una
Egadi, che
partil cuinelle
notevole
aVery
sua aree
erosione famento
voltagrasslands
ruscellamento deilesubstrati
e ilimpetuoso
parte denuda- piogge rocciosi,
durante unalitosuoli
processi notevole (Lithic
che probabilmen-
erosione Xerorthents)
50-100 eTrampling
il denuda-
iche Marino Oltre blocks,
chearee alt.
con 5Punta
lem Bassano,
(R18-R20)
caratteristiche Southern
geomorfologiche open
eche e130terre between
rosse rocks
(Lithic Rhodoxeralfs).
lla piùgeologiche,
ampia bioclimatiche
“Thrust Belt” e fitocenotiche
individuata marine
nelle contermini,
della affiorante
mentopiù ampia formatasi (Rock
“Thrust
deilesubstrati nel Outcrop),
corso
Belt”
rocciosi, del Miocene
individuata
processi litosuoli
nelle (Lithic
teprobabilmen-
si
areesono mento Xerorthents)
accentuati dei a seguito
substrati dei disboscamenti
rocciosi, processi cheeprobabilmen-
dei
3.tale della Catena delle Egadi, che a sua volta fa parte piogge determina
upper ridge oruna notevole
pebbles, alt. erosione
m (R13-R14) e il denuda-
Punta Bassano, Southern part
topograficheVery open
superiore del grasslands
territorio,
(Catalano between
et al., 1985; rocks
l’originalità Catalano, floristica
nel1986; di Miocene
tagli La50-100 dorsale Trampling orografica, posta secondo la direttrice
he con
arine le ridge
della
upper
caratteristiche
contermini, formatasi
più ampia nel
“Thrust geomorfologiche
corso
Belt” del Miocene
individuata
or pebbles,
e4.nelle
marine e130
NEtePunta
alt.Essa
terre
si
areesono
m
rosse
contermini, accentuati
mento
(R13-R14)
(Lithic
formatasi
Southern
deiquattroapart
seguito
substrati deiAr-
Rhodoxeralfs).
corso
Open
rocciosi, del
disboscamenti
sub-halophilous
processi
operati sino
teeprobabilmen-
sidei
grassland,
che sono alla metà delaXX
accentuati secolo.
seguito dei Parte
50-100
delle
disboscamenti e dei
Landslide?
Marettimo gnani, va 1987).
messa
superiore
Bassano, in
near è(Catalano
costituita
correlazione
Praia etdaalla
al., proprio
1985; unità contetto-
Catalano,
between 1986;precipitazioni
lepebbles,NO/SE,
Ar-alt 5 mtagli è la
(R15) meteoriche
dominata
operati sinoviene
daalla invece
Pizzo incanalata
metàFalcone
del XX (686 pene-
secolo. mParte
s.l.m.),
delle il
he del
periore territorio,
(Catalano
marine
4. NE Punta
et l’originalità
al.,
contermini, 1985; Catalano,
formatasi
Southerncomplesse
part
floristica
nel 1986;
corso
niche,
Open
Ar-di
del tagli
Miocene
derivanti
sub-halophilous dalla
La
operati dorsale
te sino
si
deformazioni
grassland, sono orografica,
metà
accentuati
di substrati
del a XXposta
seguito
meso-
secolo. secondo
dei
trando
Parte delle
disboscamenti
50-100lesirocce
direttrice
e dei
vicende gnani,
i Nacchi paleogeografiche
1987). Essameteoriche che
è costituita hanno
da quattro inte- unità quale
tetto- fratturate:
ergeLandslide?
nell’ambito
precipitazioni sull’isola
di unaffiorano
meteoriche sistema
viene infatti
invece di- superan-
diincanalata
vette pene-
oani,va1987).
messa Essa
superiore in è costituita
correlazione
near(Catalano da quattro
proprio
et al., ressato unità
1985; Catalano, tetto-
con le 1986; precipitazioni
NO/SE,
Ar- è
tagli dominata
operati viene
da
sinoultime invece
Pizzo
alla incanalata
Falcone (686 pene- m s.l.m.), il
Bassano, Praia zoici,
ilbetween
Canalecostituiti
pebbles,
5. Near
niche, prevalentemente
di Sicilia alt 5 m
Ficaredda
derivanti nel (R15)corso da delle
North-Eastern
dalla dolomie,
deformazioni di metà
marne
Rocky eree del
xeric
substrati tiXX
verse spesso
grassland,
meso- secolo.
sorgenti i 400
ontrando
small Parte
naturali
lemetri delle
rocce(Cusimano
di quota.
fratturate: etsull’isola
al., 1985).
I versanti 10-50 declinano
affiorano infatti ripi-
Trampling di-
che, derivanti
vicende gnani, dalla
1987). deformazioni
paleogeograficheEssa è costituita di substrati
che hanno
da quattro meso-
inte-
calcari unità
del trando
Trias quale
tetto-
medio lesi(?)-Lias
rocce
erge fratturate:
nell’ambito
precipitazioni inferioremeteorichesull’isola
(Abate dietun affiorano
ofsistema
viene
al., Uninfatti
invece di-assai
diincanalata
habitatvette superan-
pene-
i Nacchi geologiche. zoici, costituiti part
prevalentemente da pockets
dolomie, terra
marne rosa
damente e between verso
verse the lapeculiare
sorgenti costa,
naturaliè rappresentato
interrotti
(Cusimano dal
al.,siste-
da etnumerose1985). incisio-
ici, costituiti
Canale5.niche, Near prevalentemente
di Sicilia
Ficaredda
derivantineldalla
corso da dolomie,
delleL’isola
North-Eastern
deformazioni marne
1982,
ultime
Rocky ere e
1998).
xeric
dirappresenta
substrati verse
ti la
grassland,
calcarimeso- sorgenti
spesso ontrando
delporzione
Trias small naturali
imedio
400 lemetri
emersa
(Cusimano
roccepiù
(?)-Lias di quota.
fratturate:
occiden-
inferiore
et
blocks,(Abate
alt.al.,
I290 1985).
ma
versanti
sull’isola rupicolo,
m al.,
ni
et (R17)
10-50
affiorano
torrentizie, con
declinano leil imponenti
Trampling
Un infatti
habitat cuiripi-
di- pareti rocciose
ruscellamento
assai peculiare di Pizzo dal
impetuoso
è rappresentato durante
siste-
.cari delzoici,
Trias costituiti
medio (?)-Lias partinferiore
prevalentemente tale (Abate
della et
da pockets
dolomie,
Catenaal.,of6.1982,
L’associazionemarne
delle
Rumurale
Un
Near Canale
terra dietipi
rosa
damente
1998).
Egadi,habitat
pedologici
between
verseche assai
North-Eastern
verso the
sorgenti
part
a sua peculiare
lapiù diffusa
costa,
naturali
volta è(Fierot-
rappresentato
Rocky
interrotti
fa
and very
(Cusimano
parte
grassland, on
opendal
Falcone,
da
le etnumerose
piogge
pebbles,
al., siste-
Pizzo
xeric
ma
alt.
1985). del incisio-
rupicolo,
determina
250 m
Capraro,
conunale Pizzo
imponentiLisandro,
notevole pareti
100-500
erosione
nonché Trampling
rocciose di Pizzo
e il denuda-
82, 1998). ti et al.,alt.
1988; ma Fierotti,
rupicolo, 1997) conètipi
rappresentata
leilpedologici
imponenti da roccia
pareti delle aree del Libbano, Bassano, Orru Chiàppara, ecc. nonché
rappresenta calcari ladelporzione
Trias medio emersa(?)-Lias più
della blocks,
occiden-
inferiore
più ampia (Abate 290 ni
et
“Thrust m al.,
(R17)
L’associazione
torrentizie,
Belt” Un di habitat
individuata cui assai più
ruscellamento
nelle aree rocciose
diffusa
peculiare (Fierot-
è mento didei
impetuoso
rappresentato Pizzo
Falcone, dal
substrati Pizzo
durante
siste-del Capraro,
rocciosi, processiPizzoche Lisandro,
probabilmen-
L’associazione
6. Near Canale
1982, di tipi pedologici
1998). più
North-Eastern diffusacontermini,
marine (Fierot-
Rocky and ti veryetFalcone,
open
al., 1988;
formatasi Pizzo
xeric Fierotti,
ma nel delcorso
rupicolo, Capraro,
1997) è una
del
con Pizzo
rappresentata
Miocene
le imponenti Lisandro,
da roccia
te 100-500
si sono
pareti nonché Trampling
delleaccentuati
rocciose aree didel PizzoLibbano,
a seguito Bassano, Orru Chiàppara,
dei1000-5000
disboscamenti eecc.
dei
Catena delle
Rumurale
Egadi, chepart
a sua volta fa parte
grassland,
7. Isthmusle
on delle
piogge
pebbles,
of Punta determina
North-Eastern notevole
Rocky erosione
sub-halophilous grassland,e il denuda-
on Trampling
et al., 1988; Fierotti, 1997)
L’associazione di ètipi
rappresentata
superiore
pedologici da
più roccia
(Catalano
diffusa Troia et
(Fierot- areealt.
al., del
1985; 250
Falcone,
m
Libbano,
Catalano,
part Pizzo Bassano,
1986;
del Orru
Capraro,
small Ar-
sandy Chiàppara,
tagli
Pizzo
pockets operati
Lisandro,
between ecc. the sino nonchéalla metà del XX secolo. Parte delle
ampia “Thrust Belt” individuata nelle aree mento dei substrati rocciosi, processi che probabilmen-
gnani, 1987). Essa pebbles, alt. 5 m
ntermini, 7.tiIsthmus
et al., 1988;
formatasi Fierotti,
nel 1997)
corso è rappresentata
del Miocene da èroccia
tecostituita
sislopesono delle daaree quattro unità
del Libbano, tetto-
Bassano, precipitazioni
Orru Chiàppara, meteoriche dei viene invece10-50
eecc. incanalata pene-
of Punta North-Eastern Rocky sub-halophilous
niche, derivanti 8.dalla
Eastern deformazioni of accentuati
grassland, Mainon di island's a seguito
substratiPockets meso-
dei1000-5000
of terra disboscamenti
rosa
trando between
le rocceTrampling
the fratturate: sull’isola affiorano _ infatti di-
(Catalano Troia et al., 1985;part Catalano, 1986;
small Ar-sandy Pizzo
zoici, costituiti prevalentemente
tagli
pockets
Craparo, operati
between
near mountainthesino ridge alla metà
blocks,del alt. 505XX m (R16)secolo.
da dolomie, marne e verse sorgenti naturali (Cusimano et al., 1985).
Parte delle
87). Essa è costituita da quattro unità pebbles, Craparizza
alt. 5 m
calcari del tetto-
Trias medio precipitazioni
(?)-Lias inferiore meteoriche (Abate et vieneal., invece Unincanalata
habitat assai pene-peculiare è rappresentato dal siste-
ivanti8.dalla Eastern slope of
deformazioni Maindi island's
substrati
1982, 1998).
Pockets
meso- of terra
Table rosa
1.trando
Synoptic between
letable
rocce the
of the fratturate:
main environmental sull’isola 10-50 _ infatti
maaffiorano
and demographic
rupicolo, con di- of thepareti
characteristics
le imponenti 8 populations
roccioseofdiProspero
Pizzo
Pizzo Craparo, near mountain ridge blocks, alt. 505 m (R16)
ituiti prevalentemente
Craparizza
da dolomie, marnehierae
L’associazione e di currently
verse present on
sorgenti
tipi pedologici thediffusa
naturali
più Marettimo(Cusimano
(Fierot-Island. Falcone, et al., 1985). Pizzo del Capraro, Pizzo Lisandro, nonché
Trias medio (?)-Lias inferioreti (Abate et al., 1988;et al., Fierotti, 1997) Un habitat è rappresentata assai peculiare da rocciaè rappresentato delle aree del Libbano, dal siste-Bassano, Orru Chiàppara, ecc.
2. Evolution et conservatisme de niche
38
Conservatisme de niche
Conservatisme de niche [niche conservatism (NC)]

Tendance à ce que plusieurs traits d’histoire de vie d’une espèce
restent identiques à travers une longue période de temps (> 10 000 ans)
- Importance dans la prévision de la réponse des espèces face aux

changements d’environnements passés ou actuels
- Importance dans le domaine de la paléoécologie (principe

d’actualisme)
- Importance en biologie de la conservation

However, NC can also be viewed
Conservatisme as a process, if this pattern
de niche
of ecological similarity helps create other patterns (e.g.
Ecology Letters, (2010) 13: 1310–1324 doi: 10.1111/j.1461-0248.2010.01515.x
REVIEW AND
SYNTHESIS Niche conservatism as an emerging principle
Ecology Letters, (2010) 13: 1310–1324
doi: 10.1111/j.1461-0248.2010.01515.x
REVIEW AND
in ecology and conservation biology
SYNTHESIS Niche conservatism as an emerging principle
Abstract and conservation biology
in ecology
John J. Wiens,1* David D. The diversity of life is ultimately generated by evolution, and much attention has focused
Ackerly,2 Andrew P. Allen,3 on the rapid evolution of ecological traits. Yet, the tendency for many ecological traits to
Brian L. Anacker, 4
Lauren B. Abstract
instead remain similar over time [niche conservatism (NC)] has many consequences for
Resource or environment 1 Resource or environment 2
1
John J. Wiens,
5 * David D. 4 The diversity of life is ultimately generated by evolution, and much attention has focused
Buckley, Howard V. Cornell,
2
Ackerly, Andrew P. Allen,
6
3
on the the
rapidfundamental patterns
evolution of ecological andYet,processes
traits. the tendency studied
for many inecological
ecologytraits
andtoconservation biology.
Ellen I. Anacker,
Brian L. Damschen, 4
Lauren T. B.JonathaninsteadHere,
remainwe describe
similar the
over time mounting
[niche evidence
conservatism (NC)]for
hasthe
many importance
consequences offor
NC to major topics in
7,8 9
Buckley,5 Howard
Davies, John-Arvid 4
Grytnes,
V. Cornell, the fundamental patterns and processes studied in ecology and conservation biology. (e.g. climate change,
Ellen I. P.
Damschen, 6
T.4Jonathan
ecology (e.g. species richness, ecosystem function) and conservation
Susan Harrison, Bradford A.Here, we describe the mounting evidence for the importance of NC to major topics in
Davies,7,8 John-Arvid Grytnes,9 11
Hawkins,10 Robert D. Holt,
invasiveSpecies
ecology (e.g. species richness,1
species). We also reviewSpecies
ecosystem function)
other areas2whereSpecies
and conservation 3
it may be important
(e.g. climate change, Species 4
but has generally Species 5 Species 6
Susan P. Harrison,4 Bradford A. been overlooked, in both ecology (e.g. food webs, disease ecology, mutualistic
Christy 12 invasive species). We also review other areas where it may be important but has generally
Hawkins,M. 10 McCain
Robert D. Holt, and
11
Patrick
Christy M. R.McCain
Stephens12
and13 interactions)
been overlooked, in and
both conservation
ecology (e.g. food (e.g. habitat modification).
webs, disease We summarize methods for
ecology, mutualistic
Patrick R. Stephens13 interactions)
testingandforconservation
NC, and (e.g. habitat
suggest modification).
that a commonly We used
summarizeand methods
advocated for method (involving a
testing for NC, and suggest that a commonly used and advocated method (involving a
test for phylogenetic signal) is potentially problematic, and describe alternative
test for phylogenetic signal) is potentially problematic, and describe alternative
approaches.
approaches. We suggest Wethat
suggest that NC:
considering considering
(1) focusesNC: (1) focuses
attention attention on the within-
on the within-
speciesspecies
processesprocesses
that cause that
traits cause traits toover
to be conserved be conserved over time,
time, (2) emphasizes (2) emphasizes connections
connections
between questionsquestions
between and researchandareas that are areas
research not obviously
that arerelated (e.g. invasives,
not obviously global (e.g. invasives, global
related
warming, tropical richness), and (3) suggests new areas for research (e.g. why are some
warming, tropical richness), and (3) suggests new areas for research (e.g. why are some
clades largely nocturnal? why do related species share diseases?).
clades largely nocturnal? why do related species share diseases?).
Keywords
ClimateKeywords
change, community assembly, conservation, disease ecology, food webs, habitat
destruction, invasive species, niche conservatism, phylogeny, species richness.
Climate change, community assembly, conservation, disease ecology, food webs, habitat
Ecology destruction, invasive
Letters (2010) 13: species,
niche conservatism, phylogeny, species richness.
1310–1324
Ecology Letters (2010) possess.

13: 1310–1324
Many biologists are enthralled by spectacular
INTRODUCTION
examples of the rapid evolution of species and ecological
Evolution is the ultimate cause of the diversity of life, from
the origin of species to the variety of ecological, physiolog-
traits (e.g. Darwin!s finches, Rift-lake cichlids) and con-
cerned about evolutionary
possess. Manyresponses to human impacts
biologists (e.g.
are enthralled
by spectacular
Species 4
Iical,
N T morphological
R O D U C T I Oand
N behavioural traits that those species reduced body sizes in fisheries). Yet, there may also be many
examples of the rapid evolution of species and ecological
Evolution
1
Department is
of the ultimate
Ecology causeStony
& Evolution, of the
Brookdiversity of life, 8from
University, traits
Department of (e.g.
Biology, Darwin!s
McGill University,finches, Rift-lake cichlids) and con-
Quebec, Canada
the
Stonyorigin
Brook, of
NY species to the variety of ecological, physiolog-
11794, USA H3A 1B1 cerned about evolutionary responses to human impacts (e.g.
2 9
Department of Integrative Biology, University of California, Department of Biology, University of Bergen, N-5020 Bergen,
ical, morphological and behavioural traits that those species reduced body sizes in fisheries). Yet, there may also be many
Berkeley, CA 94720, USA Norway
3 10
Department of Biological Sciences, Macquarie University, Department of Ecology & Evolutionary Biology, University of
1 8
Department
Sydney, of Ecology
NSW 2109, Australia& Evolution, Stony Brook University, Department
California, Irvine, of Biology, McGill University, Quebec, Canada
CA 92696, USA
4 11
Stony Brook,
Department of NY 11794, USA
Environmental Science & Policy, University of Department H3A 1B1 University of Florida, Gainesville,
of Biology,
2California, Davis, CA 95616, USA FL 32611, USA 9Department of Biology, University of Bergen, N-5020 Bergen,
5
Department of Integrative Biology, University of California,
12
Department of Biology, University of North Carolina, Chapel Department of Ecology & Evolutionary Biology, University of
Berkeley, CA 94720, USA Norway
Hill, NC 27599, USA Colorado, and University of Colorado Natural History Museum,
3 10
Conservatisme de niche
Int. J. Plant Sci. 164(3 Suppl.):S165–S184. 2003.
! 2003 by The University of Chicago. All rights reserved.
1058-5893/2003/16403S-0012$15.00
COMMUNITY ASSEMBLY, NICHE CONSERVATISM, AND ADAPTIVE EVOLUTION

IN CHANGING ENVIRONMENTS
David D. Ackerly1
ACKERLY—COMMUNITY ASSEMBLY
Department of Biological Sciences, AND
Stanford University, Stanford, ADAPTIVE
California 94305, U.S.A. EVOLUTION S171
The widespread correspondence between phenotypic variation and environmental conditions, the “fit” of
organisms to their environment, reflects the adaptive value of plant functional traits. Several processes con-
tribute to these patterns: plasticity, ecological sorting, and adaptive evolution. This article addresses the im-
portance of ecological sorting processes (community assembly, migration, habitat tracking, etc.) as primary
causes of functional trait distributions at the local and landscape level. In relatively saturated communities,
Ceanothus. (Rhamnaceae)
plants will establish and regenerate in environments to which they are well adapted, so their distributions,
and the distributions of associated functional traits, will reflect the distribution of optimal or near-optimal
environmental conditions in space and time. The predicted evolutionary corollary of this process is that traits
related to habitat occupancy, e.g., environmental tolerances, will be under stabilizing selection. This process
Phylogeny of some Ceanothus species,
contributes to the widely observed pattern of phylogenetic niche conservatism, i.e., ecological and phenotypic
similarities of closely related species. Evidence for niche conservatism in plants is reviewed. Based on Jackson
illustrating divergence in regeneration
and Overpeck’s concept of the realized environment, I propose three scenarios in which a species’ distributional
responses to environmental conditions will lead to a “mismatch” between its environmental tolerances and strategy and several functional traits.
the environments it occupies, thus creating opportunities for adaptive evolution: (1) the colonization of “en-
vironmental islands” (habitats that are discontinuous in niche space) that require large adaptive shifts in
tolerance of one or more environmental factors; (2) the persistence of “trailing-edge” populations in species
tracking changing climate, if barriers to dispersal of competitors prevent competitive exclusion in the dete-
riorating conditions; and (3) responses to changes in the realized environment in multidimensional niche space,
A: Specific leaf area
in which species are predicted to track environmental factors for which they exhibit narrow tolerances and
exhibit adaptive evolutionary response along axes where they exhibit greater niche breadth. These three
scenarios provide a conceptual framework that emphasizes the role of ecological sorting processes and sta- B: Expression of an experimental heat
bilizing selection as the context for understanding opportunities for adaptive evolution in heterogeneous and
changing environments. stress (heat shock proteins )
Keywords: adaptation, California flora, climate change, community assembly, environmental tolerances, niche,
phylogeny, specific leaf area, stabilizing selection. C: Drought tolerance
1. Introduction ation and environmental conditions (i.e., the state of adapt-
(water potential at which stems exhibit 50%
Plant functional traits are directly responsible for the ac-
edness) reflects the adaptive value of functional traits. For ex-
ample, leaf size commonly increases with water availability,
loss of hydraulic conductivity due to
quisition of resources required for growth (light, water, nu-
trients, CO2) and the regulation of conditions that influence
and this pattern may be observed among individuals (Sultan embolism).
and Bazzaz 1993), species (Cunningham et al. 1999), and com-
metabolism (e.g., temperature, turgor pressure). Functional munity assemblages (Dolph and Dilcher 1980; Fonseca et al.
traits vary across a wide range of spatial and temporal scales
and among cells, leaves, shoots, individuals, populations, and
2000). Depending on the scale of analysis, these patterns reflect
one or more of three interacting processes: (1) phenotypic plas-
In C, the lines connect sympatric
ecosystems. Functional traits are considered adaptive if the
phenotype occurring in a particular environment enhances per-
ticity: the modification of the phenotype during development
in response to the environment (Sultan 1987); (2) ecological
species pairs along an altitudinal
formance in that environment (e.g., improved resource acqui-
sition, growth, survival and/or reproduction) relative to alter-
sorting: the differential success, due to their functional char-
acteristics, of populations or species in contrasting environ- gradient.
native phenotypic states. Familiar examples include carbon ments (Weiher and Keddy 1995); (3) adaptation by natural
gain of shade versus sun leaves in contrasting light environ- selection: heritable changes in the phenotype in populations
ments, contrasting allocation strategies related to resource occupying different environments, as a result of the adaptive
availability, and variation in leaf size and angle with respect
value of the trait (Cody and Mooney 1978).
to radiation and water availability.
Mécanisme du conservatisme de niche
- La température
change au cours du
biotope
temps
- Les niches
écologiques ne
changent pas mais
leurs expressions
dans le biotope
Espace environnemental
changent en fonction
du climat
- Chaque espèce
”tente de suivre” sa
niche écologique en
modifiant son aire de
répartition
Colwell & Rangel 2009, PNAS

Mécanisme du conservatisme de niche
- La température
- La température change
biotope
change au cours du
temps au cours du temps
- Les niches - Les espèces s’adaptent

écologiques ne aux changements
changent pas mais environnementaux et
leurs expressions peuvent conserver
dans le biotope leurs aires de
Espace environnemental
changent en fonction répartition
du climat
- Ce sont les niches
- Chaque espèce écologiques qui
”tente de suivre” sa changent car les
niche écologique en espèces doivent
modifiant son aire de modifier leurs
répartition exigences écologiques
Conservatisme et évolution de niches écologiques fait toujours référence à la niche fondamentale

Conservatisme de niche ou adaptation évolutive ?
Cas de la convergence vs. non-convergence fonctionnelle

des végétaux ligneux entre les 5 écorégions méditerranéennes du globe
La convergence de traits d’histoire de vie peut s’expliquer :
By similar environmental factors:

- Climate and notably the intensity of summer drought
- Soil nutrient status fertility
- Fire regimes
- Rainfall reliability
- Landscape age :
• Old, Climatically-Buffered Infertile Landscapes (OCBILs)
Predominance in SW Australia FR, Greater Cape FR of S Africa
• Young, Often-Disturbed Fertile Landscapes (YODELs)
Predominance in Mediterranean Basin, California, Chile
By similar evolutionary history (phylogenetic composition of lineages) induced by a

common historical biogeography
Conservatisme de niche ou adaptation évolutive ?
Two processes can explain the observed convergences of plant traits:
(i) Evolutionary adaptation, i.e. the production of new phenotypes by

the action of natural selection.
(ii) Niche conservatism, with a relative stasis in trait evolution, induced

by the ecological match between organisms and their environment
caused by the spatial and temporal sorting of existing lineages.
Similarity between traits of Mediterranean woody plants could be due

more to phylogenetical inertia than to common adaptive strategies under
similar Mediterranean climates.
The absence of deep morphological changes in leave size and specific

leaf area (SLA) suggests that most of the ancestors of shrubland taxa had
already acquired plant life-history traits that contributed to their success
under mediterranean climates.
Subtropical “phantoms” pre-date the onset of the mediterraneity
during the mid-Pliocene (C.M. Herrera 1992, Amer. Nat.).
Existence de 2 grandes lignées fonctionnelles
chez les végétaux ligneux des régions méditerranéennes
A tropical-like lineage that evolved during the Tertiary,

before the Mediterranean climates appeared
- Sclerophyllous or lauriphyllous
- Cauliflorous trees, small flowers
- Dioecy and wind- pollination
- Fleshy-fruited and large seeds
- Vertebrate-dispersed
- Regeneration by resprouting
Broad eco-morphological
similarities among the
Quercus ilex
different Mediterranean
vegetations of the world
Erica arborea
Existence de 2 grandes lignées fonctionnelles
chez les végétaux ligneux des régions méditerranéennes
A true Mediterranean lineage related to Quaternary taxa,
with higher plant diversity within genera
- Non-sclerophyllous
- Wind-dispersed or autochorous seeds
- Dry-fruited, numerous and small seeded plants
- Flowers coloured, large, often hermaphrodites
- Pollination by insects
- Regeneration by seeds
Cistus albidus
Cistus monspeliensis Genista dorycnifolia Echium valentinum

However, as
Given information onwith
the food webs andrequirements
ecological disease ecology,nec-
the
Conservatisme
essary for persistence of a species
remain understudied.
deinniche,intact vs.phylogénie
ecological mechanisms that underlie NC in these cases
modified et impacts humains
habitats (e.g. microclimates,
Finally, a recent studyfood
(Gómez resources),
et al. 2010) foundstudying
evidence
for phylogenetic conservatism in
habitat modification from a NC perspective can help all types of interspecific
elucidate the interactions
potential (host–parasite, predator–prey, mutualism) in
for those requirements to evolve,
116 clades (genera) across the Tree of Life. These authors
including generalists and specialists and viruses, bacteria,
fungi, plants and animals.
Figure 4 Niche conservatism, phylogeny and anthropogenic
Conservation and habitat modification change. Six species belong to two clades (black vs. white circles)
and occur in three communities (squares; where circles represent
Habitat destruction is often considered the most important species from each clade). In the top row of communities, the
current threat to biodiversity (e.g. Dirzo & Raven 2003), and distribution of species is associated with phylogenetically conserved
can also be seen from the framework of NC: habitat is tolerances to a range of conditions along a natural environmental
modified faster than a species can adapt to these changes gradient. In the bottom set of communities, species composition
(Holt & Gomulkiewicz 2004). To some, this may seem has now changed due to conserved tolerances to anthropogenic
trivial; little evolutionary perspective seems necessary to changes, with the loss of one of the clades from these communities
understand why cutting down a forest leads to extinction of and a reduction in phylogenetic diversity. Empirical studies have
endemic, forest-dwelling species. But threats from habitat now demonstrated changes in the phylogenetic composition of
modification will depend on the interaction between the communities in response to climate change, pollution, invasive
species, agriculture, urbanization, and other human modifications
type of modification (e.g. clearcutting, selective logging,
(Knapp et al. 2008; Willis et al. 2008; Dinnage 2009; Helmus et al.
agriculture, pollution), tolerances of species to that modi- 2010), and in some cases have shown the specific biological traits
fication (e.g. ability to withstand heat, low pH), and whether that are conserved and seemingly underlie these responses (e.g.
those tolerances will evolve rapidly or be conserved. flowering time and climate change; Willis et al. 2008). Anthropo-
Recent studies have shown that habitat modification can genic changes have also been shown to lead to a loss of
have non-random phylogenetic effects on impacted com- phylogenetic diversity (e.g. Helmus et al. 2010), seemingly through
munities (e.g. Knapp et al. 2008; Dinnage 2009; Helmus conservatism of niche-related traits.
! 2010 Blackwell Publishing Ltd/CNRS

Figure 4 Niche conservatism, phylogeny and anthropogenic
change. Six species belong to two clades (black vs. white circles)
and occur in three communities (squares; where circles represent
species from each clade). In the top row of communities, the
distribution of species is associated with phylogenetically conserved
tolerances to a range of conditions along a natural environmental
gradient. In the bottom set of communities, species composition
3. Changements climatiques
et évolutions de niche
49
Evidences de conservation de niche ou de déplacement d’espèces
Araschnia levana en Scandinavie
2000 2005 2010

Dans ce cas les optimums

altitudinaux des espèces
ont augmenté,
potentiellement à cause
du réchauffement
climatique
Felde et al 2012, Ecography

Les niches réalisées

d’arbres juvéniles (▴) se
situent en général plus
en altitude que les
niches réalisées des
arbres adultes (●)
Lenoir et al 2009, Ecography

Modélisation de niche écologique (ENM)
La modélisation de niche écologique (Ecological

Niche Modelling: ENM) est fréquemment utilisée
pour estimer les occurrences potentielles des
espèces en fonction des variables
environnementales (en général le climat) dans le
cas de différents scénarios de changements
climatiques passés ou futurs.
L’ENM permet d’inférer les dynamiques de

changements passés des aires de distribution
d’espèces clés au cours du temps :
- Processus de migration
- Processus d’extinction
- Processus de persistance ?
Prise en compte de la seule niche climatique
Source : INRA Nancy - IFN, Programme CARBOFOR

Pertinence de la seule enveloppe climatique ?
« Although the complexity of the natural

system presents fundamental limits to
predictive modelling, the bioclimate envelope
approach can provide a useful first
approximation as to the potentially dramatic
impact of climate change on biodiversity »
Pertinence de la seule enveloppe climatique ?
Implicit is the assumption that all

individuals within a species are
adapted to the same climate
enveloppe - an envelope
determined from the current range
of the species
Nouveaux climats et interactions biotiques différentes
Liens théoriques entre les niches écologiques de 4

espèces (Sp.) et les changements d’enveloppes
climatiques globales entre le XXe et le XXe siècles
• Modification de distribution et désagrégation
de la communauté (sp. 1 et 3) Woodwardia radicans
• Nouvelles interaction et communauté (sp. 2 et 3)
• Extinction (sp. 4)
D’après Williams. et al., 2007. PNAS, 104.
Utilisation du concept de niche écologique pour modéliser
la distribution géographique d’une espèce
Comment connaître la niche écologique d’une espèce ?
Algorithme de
modélisation
La niche écologique est modélisée à

partir d’observations de présence et
d’absence sur le terrain
C’est donc la niche réalisée qui est

modélisée et projetée
Modélisation de niche écologique (ENM): cas du chêne liège
Quaternary Science Reviews 119 (2015) 85e93
Contents lists available at ScienceDirect
Quaternary Science Reviews

journal homepage: www.elsevier.com/locate/quascirev
Quercus suber range dynamics by ecological niche modelling: from the

Last Interglacial to present time
Federico Vessella*, 1, Marco Cosimo Simeone, Bartolomeo Schirone
Dipartimento Ambiente, Foreste, Natura ed Energia (D.A.F.N.E.), Universit!
a degli Studi della Tuscia, 01100 Viterbo, Italy
F. Vessella et al. / Quaternary Science Reviews 119 (2015) 85e93
a r t i c l e i n f o a b s t r a c t
Table 1
Article history:
Received 2 February 2015
Ecological Niche Modelling (ENM) is widely used to depict species potential occurrence according to
environmental variables under different climatic scenarios. We tested the ENM approach to infer past retrieved calculating the
List of environmental layers
implicationsused inwiththis
a betterstudy. All variables
and conservation are raster data at 30 arc-
Received in revised form range dynamics of cork oak, a keystone species of the Mediterranean Biome, from 130 ka to the present
20 April 2015
Accepted 23 April 2015
time. Hindcasting
management for the future.
would deal species risk assessment
matic surfaces and pre-in
second resolution.
Available online
We modelled present and past occurrence of cork oak using seven ENM algorithms, starting from
Keywords:
Climate change
63,733 spatially unique presence points at 30 arc-second resolution. Fourteen environmental variables
were used and four time slices were considered (Last Interglacial, Last Glacial Maximum, mid-Holocene
PMIP3, then added to Wo
downscaled 30 arc-second
and present time). A threshold-independent evaluation of the goodness-of-fit of the models was eval-
openModeller
Variable typeuated by means of ROC curve and fossil or historical evidences were used to validate the results.
Palaeodistribution modelling Source
Putative refugia Four weighted average maps depicted the dynamics of area suitability for cork oak in the last 130 ka.
works considered in this w

Quercus suber The derived species autoecology allowed its long-term occurrence in the Mediterranean without striking
Annual Mean Temperature (! C) WorldClim

Range dynamics range reduction or shifting. Fossil and historical post-processing validation support the modelled past
spatial extension and a neglected species presence at Levantine until the recent time.
Despite the severe climatic oscillation since the Last Glacial Maximum, cork oak potential distribution
Mean Diurnal Range (! C) area experienced limited range changes, confirming its strong link with the Mediterranean Basin. The
ecological amplitude of Quercus suber could be therefore adopted as a reference to trace the Mediter-
WorldClim
Temperature Seasonality (SD " 100) (! C)ranean bioclimate area. A better knowledge of the past events of Mediterranean vegetation, a wider
range of study species and environmental determinants are essential to inform us about its current state,
WorldClim
Max Temperature of Warmest Month (! C)
its sensitivity to human impact and the potential responses to future changes.
© 2015 Elsevier Ltd. All rights reserved. WorldClim 2.3. Ecological niche model
Min Temperature of Coldest Month (! C) WorldClim
Temperature Annual Range (! C) WorldClim Cork oak potential distr
1. Introduction It is widely accepted that the present geographic distribution of
Annual Precipitation (mm)a species is the result of environmental driving forces which acted WorldClim
The factors shaping the distribution patterns of living organisms
on Earth are thePrecipitation of Wettest
key issues of a recent multidisciplinary
in the Quaternary period, i.e. the last 2.6 Ma (Hampe and Jump,
perspective Month (mm)produced range contraction, especially of
2011), and cyclically WorldClim
spatially unique points d
that includes ecology, ecophysiology, population biology, biogeog-
Precipitation of Driest Month
raphy, evolutionary studies, genetics, phylogeography, and history
warm-adapted taxa, during glaciations and expansion in the warm-
(mm)
interglacial periods (Hewitt, 2003; Stewart et al., 2010; Beatty and
WorldClim layers as set of georeferen
as well (Novicic et al., 2012). The main challenge is to understand Provan, 2013), together with human influence as a landscape
Precipitation Seasonality
how species interact with biotic and abiotic factors, and react to
changes in the ecosystem where they live (Gonza "lez-Salazar et al., (Coeff. of Variation)
modeller especially in species with commercial relevance
(Valbuena-Carabana et al., 2010). Such range fluctuations mostly
WorldClim 1.1 was employed to run w
tions of climate Digital Elevation
change on biodiversity Model
under future scenarios (m)
2013). This is worthy of interest especially for the over implica- reflect the behaviour of a species to track its favourite conditions by
shifting the geographic range (niche conservatism), or to respond ASTER GDEM algorithms set up based on
(Lepetz et al., 2009). to environmental changes by adjusting the niche through an
Slope (degrees) Derived from DEM
adaptive process (niche evolution). Stability of realized niche, in
particular, played a key role in recent biogeographic and ecological 2011). Seven algorithms w
Aspect (degrees) modelling studies to forecast distribution under several climate Derived from DEM
sent potential distribution
* Corresponding author. Tel.: þ390761357391; fax: þ390761357250.
E-mail address: vessella@unitus.it (F. Vessella).
change scenarios in tree species, which have dominated terrestrial
1
Emberger Q Index
Skype: federico.vessella. ecosystems for over 370 Ma (Ruiz-Labourdette et al., 2013).
This work
http://dx.doi.org/10.1016/j.quascirev.2015.04.018
0277-3791/© 2015 Elsevier Ltd. All rights reserved.
Climate Space Model (CSM
Entropy (MaxEnt) and Support Vector Machine (SVM) (Mun
~ oz
et al., 2011 and reference therein).
Modélisation de niche écologique (ENM): cas du chêne liège
Changements climatiques et modification de la distribution des espèces
Modeling late 21st century (2080) distributions (50 x 50 km UTM) for 1350 common
European plant species under 7 climate change scenarios, using a niche-based model
(BIOMOD), and two contrasting assumptions (no migration vs. universal migration)
N = 2294 plantes d’Europe Þ 1350 plantes (> 20 pixels)

A1 scenario
[CO2] = 800 ppm
Tm Ý 3.6°C
A2 scenario
[CO2] = 700 ppm
Tm Ý 2.8°C
B1 scenario
[CO2] = 520 ppm
Tm Ý 1.8°C
B2 scenario
[CO2] = 550 ppm
Tm Ý 2.1°C
Under the worse scenario (A1 HadCM3, no migration):

2% extinction by 2080, 22% species critically endangered
Projections régionales des pertes en espèces
végétales / modifications de GDD et d’humidité
atmosphérique
Perte attendue en végétaux très

variable à l’horizon 2080 :
- selon les scénarios climatiques
(27-42%)
- selon les régions (2,5-86%)
• Excès de perte d’espèces (ca. 60%)

: régions de moyenne montagne
(Alpes, Pyrénées, centre Espagne,
Cévennes, Balkans, Carpathes)
• Moins de perte en espèces :

régions méridionales d’Europe
méditerranéenne, des îles et de la
région pannonienne
Modélisation de la répartition potentielle de
la grenouille-taureau invasive en Amérique
du Sud (Lithobates catesbeianus) en utilisant
les données empiriques de l’aire native de
October 2011 | Volume 6 | Issue 10 | e25718
l’espèce (Amérique du Nord)
4. Importance de la micro-échelle
67
Des micro-niches différentes au sein d’un même écosystème
Les niches réalisées des espèces

s’observent à échelles très fines
https://lamaisondalzaz.wordpress.com/2010/09/01/la-niche-ecologique/
Des micro-niches différentes au sein d’un même écosystème
https://lamaisondalzaz.wordpress.com/2010/09/01/la-niche-ecologique/
Existence de micro-niches climatiques différentes
sur un même versant de montagne (Alpes suisses)
Les différences microclimatiques

(températures) sur de très courtes
distances peuvent excéder de 5 fois les
prévisions issues des scénarios du GIEC.
Source : C. Körner, Univ. Bâle (Suisse), 2005

Ecography 39: 001–011, 2016
doi: 10.1111/ecog.02494
Importance de considérer la micro-échelle

© 2016 The Authors. This is an Online Open article
Subject Editor: Nathalie Butt. Editor-in-Chief: Miguel Araújo. Accepted 19 July 2016
Fine-grain, large-domain climate models based on climate station

and comprehensive topographic information improve microrefugia
detection
Area where the temperature
Ecography 39: 001–011, 2016
doi: 10.1111/ecog.02494 logger dataset was collected
© 2016 The Authors. This is an Online Open article
Eric MeineriSubject
and Kristoffer HylanderButt. Editor-in-Chief: Miguel Araújo. Accepted 19 July 2016
Editor: Nathalie
E. Meineri (meineri.eric@gmail.com) and K. Hylander, Dept of Ecology, Environment and Plant Sciences, Stockholm Univ., Stockholm, Sweden.
Large-domain species distribution models (SDMs) fail to identify microrefugia, as they are based on climate estimates
-domain climate models based on climate station

that are either too coarse or that ignore relevant topographic climate-forcing factors. Climate station data are considered
inadequate to produce such estimates, a viewpoint we challenge here.
Using climate stations and topographic data, we developed three sets of large-domain (450 000 km²), fine-grain (50 m)
ive topographic
Digital elevationinformation improve microrefugia
temperature grids accounting for different levels of topographic complexity. Using these fine-grain grids and the Worldclim
data, we fitted SDMs for 78 alpine species over Sweden, and assessed over- versus underestimations of local extinction and
model of Sweden
area of microrefugia by comparing modelled distributions at species’ rear edges. Accounting for well-known topographic
climate-forcing factors improved our ability to model fine-scale climate, despite using only climate station data. This
approach captured the effect of cool air pooling, distance to sea, and relative humidity on local-scale temperature, but the
and subset network

effect of solar radiation could not be accurately accounted for. Predicted extinction rate decreased with increasing spatial
resolution of the climate models and with increasing number of topographic climate-forcing factors accounted for. About
half of the microrefugia detected in the most topographically complete models were not detected in the coarser SDMs and
ferof climate stations

in the models calibrated from climate variables extracted from elevation only.
Hylander Although major limitations remain, climate station data can potentially be used to produce fine-grain topoclimate
grids, opening up the opportunity to model local-scale ecological processes over large domains. Accounting for the topo-
m) used
and K. to validate the
graphic complexity encountered within landscapes permits the detection of microrefugia that would otherwise remain
Hylander,
undetected.Dept of Ecology,
Topographic Environment
heterogeneity is likely toand
have aPlant
massiveSciences,
impact onStockholm Univ.,
species persistence, andStockholm, Sweden.
should be included in
temperature studies on the effects of climate change.
stribution models (SDMs)

Microrefugia are local fail to identify
patches microrefugia,
where species persist as they are based
Correlative specieson climate models
distribution estimates(hereafter SDM)
or thatwhen the relevant
ignore regional climate becomesclimate-forcing
topographic unsuitable (Keppel are Climate
factors. commonlystation
used to predict
data arespecies ’ range shifts, extinction
considered
et al. 2012, 2015). For example, during the Last Glacial risks, and biodiversity loss due to climate change. However,
ch estimates, a viewpoint we challenge here.
Area where species
Maximum, marginal populations of pine and spruce most SDMs fail to identify microrefugia (Ashcroft 2010,
and topographic data, we developed
remained in microrefugia far norththree
of thesets of large-domain
species ’ suit- Dobrowski (450 2011,
000 km ²), fine-grain
Hannah et al. 2014).(50 m)The reasons for distribution models of
ting forable
diffregional
erent levels
climate of attopographic
the time, fromcomplexity.
which theyUsing later these fine-grain
this are two-fold;grids andspatial
first, the the Worldclim
resolution of the climatic
78 alpine species over
re-expanded their Sweden,
distributionsand(Parducci
assessed et over- versus underestimations
al. 2012). of local SDMs
predictors used to calibrate extinction and not been
have generally 78 alpine vascular
Likewise,
omparing in thedistributions
modelled current context of climate
at species ’ rear change, sufficiently fifor
edges. Accounting ne-grained
well-known to capture microrefugia (hereafter
topographic
microrefugia may provide safe havens for many organisms the grain-size issue; Randin et al. 2009, Franklin et al. 2013), plants were projected
mproved our ability to model fine-scale climate, despite using only climate station data. This
(Ashcroft 2010, Dobrowski 2011, Hannah et al. 2014). An and second, topographic climate-forcing factors other than
ffect of improved
cool air ability
pooling, distance
to predict to sea, of
the location and relative humidity
microrefugia on local-scale
has altitude that are known temperature,
to be relevantbut mustthebe included in at current and future
ould not be accurately
repeatedly been sought accounted
(Ashcroftfor.
2010,Predicted
Dobrowski extinction
2011, the ratemodelling
decreased with
of the increasing
climate predictors spatial
used to calibrate time periods
modelsHannah
and withet al.increasing
2014). Here number of topographic
we demonstrate a methodclimate-forcing factors accounted
to the SDMs (hereafter the topoclimate for. issue;
About Ashcroft 2010,
etecteddownscale
in the mostlocaltopographically
climate variables complete
across largemodels
domains, were Dobrowski
not detected 2011,inHannah et al. 2014).
the coarser SDMs and
and show how such models can improve the detection of Regarding the grain-size issue, Randin et al. (2009)
from climate variables extracted from elevation only.
microrefugia. found that potential microrefugia for alpine plants were
tations remain, climate station data can potentially be used tocaptured
better producewithin
fine-grain
fine-grain topoclimate
SDMs (few decametres),
portunity to model local-scale ecological processes over large and domains.
that Accounting
coarse-grain SDMs for the topo-
(several kilometres) overes-
This is an open access article under the terms of the Creative
untered withinAttribution
Commons landscapes permits
License, which the detection
permits of microrefugia
use, distribution thatextinctions
timated local would otherwise
because local remain
landscape patches
heterogeneity is likely
and reproduction to have
in any medium, a massive impact
provided the onwork
original species
is with colder conditions
persistence, and should remained undetected.inAn additional
be included
0
Ecography 39: 001–011, 2016
699000
1200
110
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doi: 10.1111/ecog.02494 1000 1000 11
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00
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© 2016 The Authors. This is an Online Open article 1000
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120
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Subject Editor: Nathalie Butt. Editor-in-Chief: Miguel Araújo. Accepted 19 July 2016 10
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Ecography 39: 001–011, 2016 1000
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Eric Meineri and Kristoffer Hylander doi: 10.1111/ecog.02494
© 2016 The Authors. This is an Online Open article 1320000 1340000 1360000
E. Meineri (meineri.eric@gmail.com) and K. Hylander, Dept of Ecology, Environment and Plant Sciences, Stockholm Univ., Stockholm, Sweden.
Subject Editor: Nathalie Butt. Editor-in-Chief: Miguel Araújo. Accepted 19 July 2016
Rhododendron lapponicum (b) Angelica archangelica
(a)
700 700 800
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Large-domain700 species70distribution models (SDMs) fail to identify microrefugia, as they are700based
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1000 800
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Using climate stations and topographic data, we developed three sets of large-domain (450 00050km
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7010000
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temperature grids accounting for different levels of topographic complexity. Using these fine-grain grids and the Worldclim
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data, we fitted SDMs for 78 alpine species over Sweden, and assessed over- versus8underestimations of local extinction and 800
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110
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d K. Hylander, Dept of Ecology, Environment and Plant Sciences, Stockholm Univ., Stockholm, Sweden. 00 00
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Microrefugia are local 90 patches where species persist Correlative species
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Mapthe of distributions predicted by the TopoClim SDMs for a high- and a low-alpine species (a and b, respectively) a
(b)Map of distributions
models can improvepredicted
for. Predicted extinctionbyrate the TopoClim SDMs for a high-et al.and a low-alpine species (arefer
andto b,
therespectively) according to theabsent at both
and show how such theAngelica
detection ofarchangelica
Regarding the grain-size issue, Randin (2009)
not bemicrorefugia.
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increasing
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spatial RCP4.5.
plants were
Numbers in the legend following categories: 1) species predicted
els and with CO2 increasingemissionnumber
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800 refer
(2085)About
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time
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2) habitat predicted
decametres), 1)between
to be lost species nowpredicted
and 2085; 3) absent at both
habitat predicted to be gained between now
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4) microrefugia (stable range)

overes- predicted by all methods; 5) microrefugia undetected in the ElevOnly SDMs; 6) microrefugia un
800
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Thismost topographically
current accessandarticlefuturecomplete
00 termsmodels
11the (2085) of thetimewere notand
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microrefugia undetected in both the ElevOnly and Worldclim SDMs. The gained
between now and 2085; 3) habitat predicted to be
that coarse-grain SDMs (several kilometres)
0
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is an open under Creative 70 0
climateCommons
variables extracted
Attribution License,from whichelevation distribution800 timated local 0extinctions
only.
permits use, 0 thebecause
Worldclim SDMs; 7)
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between
and reproduction
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with colder(stable
to produce
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fi
0 00 range)
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remained 5 00
predicted
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area topoclimate
projected within by SDMs
the all methods;
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a description of each type of SDM. C
010000
1bias
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1200
nity to model local-scale ecological processes over large domains. Accounting for the topo-
1000
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120
0
80 100 600
10
0
five individuals per population. For each diversity index, mean values O’Connell, Brown, Guin
Importance de considérer la micro-échelle across 1,000 replicates were used in correlation analyses (see below). Topographic variables
(a) (b)
Areas: A B C D
Received: 13 November 2017 | Revised: 17 July 2018 | Accepted: 28 July 2018
DOI: 10.1111/ddi.12836
BIODIVERSITY RESEARCH
Topography explains the distribution of genetic diversity in one

of the most fragile European hotspots
José Luis Blanco-Pastor1,2 | Mario Fernández-Mazuecos1 | Alberto J. Coello1,3 |

Julia Pastor1 | Pablo Vargas1
L. glacialis populations
1
Real Jardín Botánico de Madrid (RJB-CSIC),
Madrid, Spain Abstract (c)
2
INRA, Centre Nouvelle-Aquitaine-Poitiers, Aim: To investigate factors that explain the spatial pattern of genetic diversity in
UR4 (URP3F), Lusignan, France
three closely related species (Linaria glacialis, Linaria nevadensis and Chaenorhinum
3
Universidad Rey Juan Carlos, Madrid, Spain
glareosum) endemic to a fragile high mountain ecosystem.
Correspondence Location: The alpine belt of Sierra Nevada, Spain.
José Luis Blanco-Pastor, INRA, Centre
Methods: We analysed the spatial pattern of cpDNA diversity of the three species.
11
10
Nouvelle-Aquitaine-Poitiers, UR4 (URP3F), 6
Lusignan, France. To explain the distribution of genetic diversity, we investigated the effect of topo- 12 4 9
Email: jose-luis.blanco-pastor@inra.fr
graphic features and the evolutionary history of the species (demography, habitat 3 7 8
2 5
To investigate factors that explain
Funding information availability and colonization dynamics).
The Spanish Ministry for Environment, and 1 13
Results: Genetic diversity was heterogeneous across the landscape. We found mod-
Rural and Marine Affairs, Grant/Award
Number: 005/2008; Spanish National erate positive correlation values between genetic diversity indices of the two Linaria
the spatial pattern of genetic

Research Council, CSIC; Spanish Ministry of
Education, Grant/Award Number: AP2007-
01841 and FPU16/05681; Spanish Ministry
species. We also observed moderate negative correlation values between genetic
diversity indices of C. glareosum and those of L. glacialis and L. nevadensis. Topographic
(d) L. nevadensis populations
diversity in three closely related
of Economy and Competitivity, Grant/Award
variables correlated positively with genetic diversity of the Linaria species and nega-
Number: IJCI-2015-23459
tively with genetic diversity of C. glareosum. Bayesian skyline plots (BSPs) displayed a
Editor: Aibin Zhan
shared demographic pattern with a population size stabilization/increase since the
species (Linaria glacialis, Linaria LGM (the last 21 kyr) in all three species. Discrete phylogeographical analyses
showed similar patterns of westward diffusion for L. nevadensis and C. glareosum.
6 4
nevadensis and Chaenorhinum Species distribution models pointed to similar range dynamics in all three species,
with a reduction in range size since the LGM. 5 11
12 2
7
14
9
Main conclusions: Different dispersal abilities, demographic trends and colonization 1
glareosum) endemic to a fragile patterns can hardly explain the differences in spatial patterns of genetic diversity
between the Linaria species and C. glareosum. In contrast, topographic features seem
13
3 8 10
high mountain ecosystem to be an important factor to explain the distribution of genetic diversity in the alpine
belt of Sierra Nevada. We point to a relevant role of microniche partitioning in deter-
mining patterns of genetic diversity distribution in alpine Mediterranean ecosystems.
Furthermore, we highlight the role of microhabitat heterogeneity in the maintenance
of distinct lineages, species and genetic diversity in high mountain biodiversity
(e) C. glareosum populations
hotspots.
KEYWORDS
comparative phylogeography, genetic diversity, high mountain, niche partitioning, Quaternary, 7
topography 11 2
4 3 10
1 8 9
5
|
Diversity and Distributions. 2018;1–16. wileyonlinelibrary.com/journal/ddi © 2018 John Wiley & Sons Ltd 1 6
cialis (cpDNA) and L. nevadensis. We found significant positive cor- the growth-decline model. The distributions for L. nevadensis and
relation values between topographic variables and diversity indices of C. glareosum fitted equally well to the constant size and the growth–
the two Linaria species (0.03 < r < 0.1; p < 0.05). In contrast, we found decline models.
9
(a) L. glacialis
outgroup
2
4 3
4
10 5 5
8
6
7
2 8
3
7 9
10
6 11
11
(b) L. nevadensis
1 + outgroup
5 2
3
4
6 5
4 6
F I G U R E 2 CpDNA haplotype 2 3
networks for Linaria glacialis, Linaria
nevadensis and Chaenorhinum glareosum,
and spatial distribution of haplotypes
in the Sierra Nevada National Park.
Each haplotype is represented by both (c) C. glareosum
10 7
a number and a colour. Areas of circles 1 + outgroup
in haplotype networks are proportional 2
3
to the number of individuals displaying 9 4
each haplotype. In the L. glacialis network, 11 5
haplotype 1 (red) represents the outgroup 6
2 7
species (L. verticillata). In the other two
4 6 8 8
networks, haplotype 1 is shared between 9
the outgroup (L. amoi for L. nevadensis and 5 10
C. villosum for C. glareosum) and the study 11
1 2.5 0 2.5 5 7.5 10 km
species 3
C. glareosum
(a) L. glacialis
Importance
(c) de considérer
C. glareosum la micro-échelle
7.0
3.1 11.6
75.4 3.5
29.9 13.1
3.3
3.7 52.8
3.8
00 10,000 15,000 20,000 25,000 30,000 35,000
(b) L. nevadensis
Time (kya)
Areas: A B C D
13.4
F I G U R E 5 Results of Bayesian discrete phylogeographical
5.2
17.7 4.3 analyses (DPAs). Analyses are based on cpDNA sequences of Linaria
glacialis, Linaria nevadensis and Chaenorhinum glareosum. Arrows
represent spread routes supported by Bayes factors (BF > 3).
Bayes factor values are shown. Pie charts represent probabilities of
00 10,000 15,000 20,000 25,000 30,000 35,000
ancestral location
cene (c) Würm glaciation
C. glareosum
PMIP3 (CHELSA) layers recovered no potential distribution of these two
Bayesian skyline plots (BSPs) representing the species, while PMIP2 (WorldClim) layers recovered relatively broad dis-
11.6
story of Linaria glacialis, Linaria nevadensis and tributions. For L. nevadensis, projections displayed little variation in the
13.1
glareosum in the past c.28, c.10 and c.35 thousand extent of potential distribution through time in WorldClim-based analy-
3.3
vely, in the alpine areas of Sierra
3.7 Nevada (Spain).
52.8 The
ses. In the CHELSA-based analysis, projection to the LGM under PMIP3
ts time; the Y-axis represents effective3.8 population
from cpDNA data sets. The bottom graph represents data displayed a widespread distribution, even including lowlands sur-
riation measured from the Vostok (Antarctica) ice rounding Sierra Nevada. To visualize the effect of single variables on the
Areas: A B C D
from Petit et al. (1999)
L. glacialis L. nevadensis C. glareosum
Last
interglacial
(WorldClim)
The observed spatial
Last
structure of cpDNA diversity
glacial is linked to plant
maximum
(WorldClim,
CMIP5)
colonization success.
Last
glacial Topography is a key
maximum
(WorldClim,
PMIP2)
determinant of climatic
variation at small spatial
Last
glacial scales, especially in treeless
maximum
(CHELSA, areas such as alpine
PMIP3)
regions.
Mid-
Holocene
(WorldClim, Association between
CMIP5)
topographic microniche
variation and the spatial
Present
(WorldClim) distribution of genetic
diversity of plant species in
this heterogeneous alpine
Present
(CHELSA)
landscape.
F I G U R E 6 Results of species distribution modelling for Linaria glacialis, Linaria nevadensis and Chaenorhinum glareosum using the
maximum entropy algorithm, as implemented in maxent. Analyses are based on WorldClim and CHELSA variables, and projected to past
conditions: last interglacial (LIG, c. 120–140 kya), last glacial maximum (LGM, c. 21 kya) and mid-Holocene (MH, c. 6 kya). The presence/
absence was estimated using the maximum training sensitivity plus specificity logistic threshold
• Modélisation des aires de répartition
potentielles de six espèces.
• Résolution spatiale 150 m
àPuis visites des sites avec des forts potentiels de
présence pour trouver de nouvelles populations
16 nouvelles populations trouvées !
77

Niche Écologique

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Le concept de niche écologique

Importance en écologie et conservation

Pr. Frédéric Médail

1. Le concept de niche : définitions

2. Evolution et conservatisme de niche

3. Changements climatiques et évolution de niche

+ : présence de l’espèce ; o : absence de l’espèce

Pulliam H.R. 2000. Ecol. Lett., 3.

Niche réalisée d’un compétiteur B

+ : présence de l’espèce ; o : absence de l’espèce

La niche est un volume

Hutchinson 1957 Cold Spring Harbor Symposia on quantitative Biology

Si les ressources sont

+ : présence de l’espèce ; o : absence de l’espèce

Une espèce peut être absente

+ : présence de l’espèce ; o : absence de l’espèce

-> Décrit la correspondance réciproque entre l’espace de la niche et l’espace physique

• L’espace de niche: partie

• Par contre, l’intégralité de

Hutchinson 1957 Cold Spring Harbor Symposia on quantitative Biology

Imaginons un nouveau biotope, où pouvons nous prédire S1 et S2 dans le biotope?

Temperature Algal food size

Imaginons un nouveau biotiope, où pouvons nous prédire S1 et S2 dans le biotope?

Temperature Algal food size

Dans tous ces cas :

- Si l’espèce S1 est supérieure, l’espèce S2 est

Modèle théorique de niche fondamentale

Hypothèse de compression (McArthur et Wilson 1967, The Theory of Island Biogeography):

Cas plus commun :

Gaston 2003, The structure and dynamics of geographic ranges

Plusieurs populations d’une

Figure 6-2. Relationship between population niche and species niche.

Polygala rupestris Viola arborescens

Climatic niche of 11 Mediterranean plants on

Difficulté de définir la niche d’une espèce dans le temps et l’espace

Prise en compte de la dimension temporelle

Prise en compte de la dimension spatiale sur de vastes territoires

Site 1 - Bosco Pisano

Valle di Castiglione EXTINCTION

(Sadori et al., 2008)

(MESO-)SUPRA-MEDITERRANEAN BELT THERMO-MEDITERRANEAN BELT

Fortes (extrêmes) fluctuations inter-annuellles dans

Plant persistence along biogeographical gradients

The ca. 1500 geophytes of the Mediterranean Basin

Representation of multiple demographic strategies

Nous ne pouvons pas afficher cette image pour l’instant.

Nous ne pouvons pas afficher cette image pour l’instant.

Nous ne pouvons pas afficher cette image pour l’instant.

o Estimer l’aire totale de distribution de l’espèce (données actuelles et données historiques)

o Analyser l’analyse écologique la plus précise possible de l’habitat de l’espèce :

o Dans le cas de vastes distributions ou d’espèces fréquentes, opérer des sélections

o Réaliser un modèle de distribution de l’espèce (Species Distribution Model, SDM)

Distribution du triton (Triturus vulgaris) en Europe

Year Published: 2018

Date Assessed: July 29, 2015

lare dell’Isola di Marettimo (Arcipelago delle Egadi, Sicilia occidentale): …

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Fig. 1 – Ubicazione e principali località dell’Isola di Marettimo.

Conservatisme de niche [niche conservatism (NC)]

- Importance dans la prévision de la réponse des espèces face aux

- Importance dans le domaine de la paléoécologie (principe

- Importance en biologie de la conservation

Ecology Letters (2010) possess.

COMMUNITY ASSEMBLY, NICHE CONSERVATISM, AND ADAPTIVE EVOLUTION