Twenty million years of extinction and survival in the Caribbean

Liliana M. Dávalos

ColEvol17 August 2017

Biodiversity: Equilibrium

• Spp. richness ~ resource abundance

• Long-term spp. richness stable

• Diversity dynamics follow predictable course after perturbation

Biodiversity: Disequilibrium

• Communities are unsaturated

• Disturbance prevents communities from reaching equilibria

• Dispersal limitation slows reaching equilibrium

Most tests limited to static prediction of dynamics

Morgan & Woods 1986 Biol. J. Linnean Soc.

Dynamic equilibrium needs phylogenies

• Test of MW evolutionary equilibrium dynamics using phylogenies

Ricklefs & Bermingham 2001 Science

Rojas, Warsi, Dávalos 2016 Syst. Biol.

Sphaeronycteris

Vampyressa (Metavampyressa)

Ectophylla

Rhinophylla

† Artibeus anthonyi

AriteusArdopsPygoderma

Stenoderma

Artibeus (Dermanura)

Uroderma

Ametrida

Carollia Phyllostomus

Chiroderma

Phyllops falcatus

† Cubanycteris silvai

Vampyressa (Vampyressa)

Centurio

Enchisthenes

† Phyllops silvai

Sturnira

Artibeus (Artibeus)

Artibeus (Koopmania)

Vampyrodes

Mesophylla

Platyrrhinus

† Phyllops vetus

Vampyressa (Vampyriscus)

Mill ion years before present

Figure 1

0.05.010.015.020.025.0

Posteriorprobability

1.00

0.48

Oligocene Miocene Pliocene

Short-faced batsStenodermatina

Stenodermatinae

Stenodermatini

Most of the fossils are in phylogeny too Tavares, Dávalos et al. in revision

Data on Greater Antillean bats

Valente, Etienne, Dávalos 2017 Nature Ecol. & Evol.

Equilibrium dynamics!

Age Ma

λc μ γ λa

20 0.22 0.26 0.017 0.2745 0.26 0.35 0.021 0.27

Valente, Etienne, Dávalos 2017 Nature Ecol. & Evol.

What about extinct species?

Valente, Etienne, Dávalos 2017 Nature Ecol. & Evol.

Fossils make a difference

Age Ma

λc μ γ λa

20 0.28 0.33 0.030 0.1945 0.32 0.41 0.036 0.16

Age Ma

λc μ γ λa

20 0.22 0.26 0.017 0.2745 0.26 0.35 0.021 0.27

Valente, Etienne, Dávalos 2017 Nature Ecol. & Evol.

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 to restore species? Wait for 8 Ma

Valente, Etienne, Dávalos 2017 Nature Ecol. & Evol.

Highlights

• Longstanding equilibrium upended by recent extinctions

• diversity is predictable

• time to recovery long

• phylogenies require fossils to discover patterns

Photo by Jon Flanders

What killed the mammals of the Caribbean? Deglaciation!

© Adrian Tejedor

Islands of the Caribbean: The West Indies

Dávalos,& Turvey 2012 Bones, Clones, and Biomes

Dávalos & Russell 2012 Ecol. Evol.

Trouble is we didn’t have the dates

Soto-Centeno & Steadman 2015 Sci. Rep.

Timing is everything Cooke, Dávalos et al. 2017 Annu. Rev. Ecol. Evol. Syst.

Large islands: Two extinction waves

• Larger islands show some pre-human extinctions• Not all extinction

caused by humans• But another cluster of

extinction after human arrival• Humans responsible

for some lossesCooke, Dávalos et al. 2017 Annu. Rev. Ecol. Evol. Syst.

Lesser Antilles: One extinction wave

• Many smaller mammals survived human use• Found in middens

and were presumably eaten

• But could not recover after European arrival• Cats, rats, mice,

goats, introduced

Lessons from Marie Galante Stoetzel et al. 2016 Quat. Sci. Rev.

Many species lack dates

• Some groups/islands well studied• Many missing

• Rodents & bats• Even in the Greater

Antilles • We know too little

about most species• But we know one

thing…Cooke, Dávalos et al. 2017 Annu. Rev. Ecol. Evol. Syst.

Phylogeny!

• Some Caribbean groups distinct• e.g., Solenodon

• Others less distinct• Some hutias &

Short-faced bats• Still, too little life

history

Cooke, Dávalos et al. 2017 Annu. Rev. Ecol. Evol. Syst.

A few traits

• Bat extinction• Larger species• Herbivores

• Among non-bats• Larger• Smaller?!

Cooke, Dávalos et al. 2017 Annu. Rev. Ecol. Evol. Syst.

Mammals of the Caribbean

© Yomangani/Creative Commons © Eladio Fernandez

© Jon Flanders

Survivors are on the brink of extinction

• Among bats• 9 of 60 threatened• 3 known from 1

cave each• Among non-bats• 8 of 12 threatened• How to predict

extinction or survival?

Illustration by Adrián Tejedor

Cooke, Dávalos et al. 2017 Annu. Rev. Ecol. Evol. Syst.

Modeling extinction/survival Unpublished

Modeling extinction/survival Unpublished

Spp. traits

Island characteristics

Mass is quadratic

Many island characteristics can be added

Intercept on probability of survival

−2000 −1000 0 1000 2000

Standard deviationMean

Zygodontomys_spXenothrix_mcgregoriXaymaca_fulvopulvis

Tainotherium_valeiSolenodon_paradoxusSolenodon_marcanoiSolenodon_cubanus

Solenodon_arredondoiSheep_Pen_femur_unidentified_rodent_taxon

Rhizoplagiodontia_lemkeiQuemisia_gravis

Plagiodontia_veloziPlagiodontia_spelaeum

Plagiodontia_aediumPennatomys_nivalis

Parocnus_serusParocnus_browni

Paralouatta_varonaiOryzomys_antillarum

Oryzomyini_undescribed_species_small_Montserrat_taxonOryzomyini_undescribed_species_small_Grenada_taxon

Oryzomyini_undescribed_species_large_Montserrat_taxonOryzomyini_undescribed_species_large_Grenada_taxon

Oryzomyini_undescribed_species_St._Martin_taxonOryzomyini_undescribed_species_Saba_taxon

Oryzomyini_undescribed_species_Anguilla_taxonOligoryzomys_victus

Nesophontes_zamicrusNesophontes_sp2Nesophontes_sp1

Nesophontes_paramicrusNesophontes_micrus

Nesophontes_hypomicrusNesophontes_edithae

Neocnus_toupitiNeocnus_gliriformisNeocnus_dousman

Neocnus_comesMysateles_prehensilis

Mesocapromys_sanfelipensisMesocapromys_nanus

Mesocapromys_melanurusMesocapromys_kraglievichi

Mesocapromys_auritusMesocapromys_angelcabrerai

Megalomys_luciaeMegalomys_georginae

Megalomys_desmarestiiMegalomys_audreyae

Megalocnus_zileMegalocnus_rodens

Macrocapromys_latusMacrocapromys_acevedoIsolobodon_portoricensis

Isolobodon_montanusInsulacebus_toussaintianaHyperplagiodontia_araeum

Hexolobodon_phenaxHeteropsomys_insulans

Geocapromys_thoracatusGeocapromys_sp2Geocapromys_sp1

Geocapromys_ingrahamiGeocapromys_columbianus

Geocapromys_browniiElasmodontomys_obliquus

Clidomys_osborniCapromys_sp1

Capromys_piloridesBrotomys_voratus

Brotomys_contractusBoromys_torrei

Boromys_offellaAntillothrix_bernensis

Antillomys_rayiAcratocnus_ye

Acratocnus_simorhynchusAcratocnus_odontrigonus

Acratocnus_antillensis

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Initial results species

• Species vary in probability of survival

• Some appear doomed

Initial results islands

• Multiple species were extinction traps

Initial results predictors

• Island area weakly related to survival

• Elevation strong predictor extinction

• Intuition about large and small spp. correct!

A history of extinction Cooke, Dávalos et al. 2017 Annu. Rev. Ecol. Evol. Syst.

A history of people

• Humans arrived in four waves• First three waves

from New World• Introduced dogs

and agriculture• Last wave from Old

World• Introduced

invasive spp.

©Theodor de Bry/National Library of Medicine

Larger mammals probably hunted early on

© Ghedoghedo Wilson 1989 J Field Archaeol.

Many extinction events recorded after arrival of Old World invasive competitors/predators

Turvey et al. 2012 Mamm. Biol.

Agriculture likely important for certain species

© Adrian Tejedor ©Pierpont Morgan Library

Highlights

• Multiple extinction drivers

• some before humans

• most after humans

• both small and large terrestrial species went extinct (not just hunting)

• agriculture possibly important for bats, others

© Yomangani/Creative Commons

Why is this important? The world is made of islands

This is happening now Ray et al. 2015 Mongabay

What happens when the frontier closes? 1869

Frontier closes, forest vanishes Stanton 2014 Biol. Cons.

The frontier

• Border between settled lands and natural habitats (nominally belongs to the state)• Forest->property• Final state = no forest• Already happened in other

regions• Most of Andes• western Caquetá• western Putumayo

• Currently unfolding in parts of Amazonia, most of Chocó

Etter et al. 2006 J. Environ. Manage.

Frac

tion

fore

st

The Andean frontier A general model

Frac

tion

fore

stcoca beachhead

Time ->

mixed agriculture

properties

pastures!

The sixth extinction is (partially) avoidable

• The species-area relationship is real• so is species loss with

fragmentation• We have the tools to

conserve and connect ecosystems

• We have roughly 30 years before the Amazon is transformed into properties• We are responsible for

evolution into the future

Thanks!