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The importance of models —dynamics of extinction and their drivers from the Miocene to the last 40 years
Liliana M. Dávalos
National Science Foundation15 July 2016
!
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Who am I? Evolution & Conservation
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Our research mission
Biological diversityDiversification Extinction
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Focus first part
Biological diversity
Colonization, speciation Extinctionincrease decrease
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The debate: equilibrium
• Spp. richness ~ resource abundance
• Long-term spp. richness stable
• Diversity dynamics follow predictable course after perturbation
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The debate: disequilibrium
• Communities are unsaturated
• Disturbance prevents communities from reaching equilibria
• Dispersal limitation slows reaching equilibrium
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Elements of the equilibrium model MacArthur & Wilson 1963 Evolution
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Most common application
Morgan & Woods 1986 Biol. J. Linnean Soc.
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Extended application Dávalos & Russell 2012 Ecol. Evol.
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Change in habitat ~ change in spp. Dávalos & Russell 2012 Ecol. Evol.
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But larger Greater Antilles < spp. than expected Dávalos & Russell 2012 Ecol. Evol.
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But what about the dynamics over time? Rabosky & Hurlbert 2015 Am. Nat.
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Phylogenies and equilibrium
• Test of MW evolutionary equilibrium dynamics using phylogenies
Ricklefs & Bermingham 2001 Science
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A model including speciation Valente et al. 2015 Ecol. Lett.
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Key findings from model comparison
• Disequilibrium in most bird groups
• Tendency to equilibrium in Darwin’s finches
• Diversity-dependent
• Together, community dynamics are non-equilibrial
Valente et al. 2015 Ecol. Lett.
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Rojas et al. 2016 Syst. Biol.
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Rojas et al. 2016 Syst. Biol.
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Figure 2
Ametrida Sphaeronycteris Centurio Pygoderma Ardops Ariteus Phyllops
† Phyllops silvai† Phyllops vetus
Stenoderma † Cubanycteris silvai
Artibeus (Artibeus)
† Artibeus anthonyiArtibeus (Koopmania)
Artibeus (Dermanura)
Ectophylla Enchisthenes Chiroderma Vampyressa (Vampyriscus)
Vampyressa (Metavampyressa)
Mesophylla
Vampyressa (Vampyressa)
Platyrrhinus Vampyrodes Uroderma
12 10 8 6 4 2 0
Mil l ion years before present
S A
M2.83
1.780.28
0.12
2.60
1.02
Founder-event dispersal(20% of cladogenetic events)
S A
M
1.13
0.32
6.465.04 0.68
0.73
Anagenetic dispersal(33% of cladogenetic events)
Miocene Pliocene
Tavares et al. in prep.
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How old are the Greater Antilles? 35 Ma?
Iturralde-Vinent & MacPhee 1999 Bull. Am. Mus. Nat. Hist
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Input data on Greater Antillean bats Valente, Etienne, Dávalos in review
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Equilibrium dynamics! Valente, Etienne, Dávalos in review
Age Ma
λc μ γ λa
20 0.28 0.33 0.030 0.1945 0.32 0.41 0.036 0.16
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What happens if no fossils? Valente, Etienne, Dávalos in review
Age Ma
λc μ γ λa
20 0.22 0.26 0.017 0.2745 0.26 0.35 0.021 0.27
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Why are so many bats extinct? Climate? Humans?
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It’s looking more and more like humans Stoetzel et al. 2016 Quat. Sci. Rev.
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0 10 20 30 40
020
4060
80
Num
ber o
f end
emic
spe
cies
0 10 20 30 40
010
2030
4050
Num
ber o
f end
emic
spe
cies
Time (Ma)Time (Ma)
BA
How long to restore the species? Valente, Etienne, Dávalos in review
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Conclusions part I
• First proposed in the 1960s, some evolutionary aspects of equilibrium theory can only be tested now
• Using phylogenies
• Longstanding questions remain to be answered, e.g.,
• is diversity predictable?
• relative contribution biotic vs. abiotic factors
Photo by Jon Flanders
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More conclusions part I
• Need to include extinct species
• Underestimate extinction rate (obvious)
• Underestimate immigration rate (less obvious)
• Human effects large
• Long time to recovery
• Habitat change probably important Dávalos & Russell 2012 Ecol. Evol.
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Focus second part
Biological diversity
Colonization, speciation
Land use changeincrease decrease
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When the UN comes knocking The Special Session of 2016
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But illicit crops are everywherePrevious focus on
Colombia
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Why tropical deforestation? I
• 2 models
• Immiserization:
• poor growers bring marginal land into production
• external shocks (e.g., debt crisis)
• internal dynamics (e.g., demographic growth)
Rudel & Roper 1997 World Dev.
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Why tropical deforestation? II
• Frontier:
• investors/government build roads
• roads bring colonization
• colonists clear land
• frontier stabilizes (development),
• or recedes (hollow frontier)
Rudel & Roper 1997 World Dev.
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1400
1600
1800
2000
2200●
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3035404550
Forest
NP
1500
2000
2500
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28 30 32 34 36 38 40
Pasture
150200250300350400
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0.51.01.52.02.5
Coca
500
550
600
650
700 ●●
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3035404550
PA
(ha)
150
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250
300
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28 30 32 34 36 38 40
2468101214
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0.51.01.52.02.5
90
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3035404550PLAND
EN
N (m
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95100105110115120
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28 30 32 34 36 38 40PLAND
300
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0.51.01.52.02.5PLAND
Figure 2
BA
G
C
E FD
H I
253035404550
2001 2004 2007 201025
30
35
40
2001 2004 2007 2010
1
2
3
2001 2004 2007 2010
PLA
ND
K L M
The closing of the forest frontier
• An example from Guaviare, Colombia
• Forest declines, pastures increase
• Forest fragments become smaller and more isolated
• New stable equilibrium loses forest almost completely
Dávalos et al. 2014 Biol. Cons.
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Where do illicit crops grow?
• In 2000s Colombia
• Correspond to forest frontier
• Interface between settled land and forests
• Spatially clustered
Dávalos et al. 2011 Environ. Sci. Technol.
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What happened in the 1960s? Etter et al. 2006 J. Environ. Manage.
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Opening the frontier Denevan 1966 Geogr. Rev.
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Roads and colonization projects
Brücher 1977 Geogr. Z. Schuurman 1978 Tijdschr. Econ. Soc. Ge.
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Brazil
Bolivia
Peru
Colombia
Ecuador
Coca cultivation
Government-sponsored
before 1979
1990/1992
Projects
Coca cultivation
High
Low
2014
Percent forest Probability of coca
a b c
A remarkable spatial pattern
Dávalos, Sanchez, Armenteras in revision
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Brazil
Bolivia
Peru
Colombia
Ecuador
Coca cultivation
Government-sponsored
before 1979
1990/1992
Projects
Coca cultivation
Percent forest
a b
Location of projects ~ C emissions Harris et al. 2012 Science
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Coca cultivation
High
Low
2014
Percent forestHigh
Low
Probability of coca
b c
Is coca spatially clustered around projects? Dávalos, Sanchez, Armenteras in revision
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High
Low
Probability of coca
c
~40 years later, Amazonia coca remains clustered Dávalos, Sanchez, Armenteras in revision
Parameter 2.5 Mean 97.5Intercept -1.2671 -1.1540 -1.0410Distance to nearest project
-0.6096 -0.5613 -0.5129
Precision municipality intercept
1.116 1.4582 1.910
Precision spatial
0.243 0.5211 1.079
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Coca is spatially associated with the frontier UNODC 2016
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Can coca explain some deforestation?
• Three independent studies say no
• Dávalos et al. 2011 Environ. Sci. Technol. no effect after including social factors
• Armenteras et al. 2013 Reg. Environ. Change no effect of coca in Amazon
• Sánchez-Cuervo et al. 2013 Ecosystems no effect any time or place Photo by Ana Luz Porzecanski
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No relationship coca cultivation ~ deforestation
ratesUnpublished
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Conclusions part II
• Actions from 4 decades ago have large effects
• Brought colonists to the Amazon
• Farmers turned to coca
• Landscapes transform following roads
• End point often minimal forest
Photo by OSITRAN, Peru
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More conclusions part II
• Spatial patterns misleading without history
• History illuminates the opening of the frontier
• Frontier deforestation can be connected to poverty
• But actually requires investment, e.g.,
• Roads
• Development projects
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Thanks!