Opening keynote at the Colombian Congress on Evolutionary Biology, 17 August 2017

1. Twenty million years of extinction and
survival in the Caribbean
Liliana M. Dávalos
ColEvol
17 August 2017

2. Biodiversity:
Equilibrium
• Spp. richness ~ resource
abundance
• Long-term spp. richness
stable
• Diversity dynamics follow
predictable course after
perturbation

3. Biodiversity:
Disequilibrium
• Communities are
unsaturated
• Disturbance prevents
communities from
reaching equilibria
• Dispersal limitation slows
reaching equilibrium

4. Most tests limited to static
prediction of dynamics
Morgan & Woods 1986 Biol. J.
Linnean Soc.

5. Dynamic equilibrium
needs phylogenies
• Test of MW evolutionary
equilibrium dynamics
using phylogenies
Ricklefs & Bermingham 2001 Science

6. Rojas, Warsi, Dávalos
2016 Syst. Biol.

7. Sphaeronycteris
Ariteus
Ardops
Pygoderma
Stenoderma
Ametrida
Phyllops falcatus
† Cubanycteris silvai
Centurio
† Phyllops silvai
Artibeus (Artibeus
† Phyllops vetus
Million years before present
Figu
0.05.010.015.020.025.0
Posterior
probability
1.00
0.48
Oligocene Miocene Pliocene
Short-faced bats
Stenodermatina
Most of the fossils are in
phylogeny too
Tavares, Dávalos et al. in revision

8. Data on Greater
Antillean bats
Valente, Etienne, Dávalos 2017 Nature
Ecol. & Evol.

9. Equilibrium dynamics!
Age
Ma
λc μ γ λa
20 0.22 0.26 0.017 0.27
45 0.26 0.35 0.021 0.27
Valente, Etienne, Dávalos 2017 Nature
Ecol. & Evol.

10. What about extinct
species?
Valente, Etienne, Dávalos 2017 Nature
Ecol. & Evol.

11. Fossils make a
difference
Age
Ma
λc μ γ λa
20 0.28 0.33 0.030 0.19
45 0.32 0.41 0.036 0.16
Age
Ma
λc μ γ λa
20 0.22 0.26 0.017 0.27
45 0.26 0.35 0.021 0.27
Valente, Etienne, Dávalos 2017 Nature
Ecol. & Evol.

12. 0 10 20 30 40
020406080
Numberofendemicspecies
0 10 20 30 40
01020304050
Numberofendemicspecies
Time (Ma)Time (Ma)
BA
How to restore species?
Wait for 8 Ma
Valente, Etienne, Dávalos 2017 Nature
Ecol. & Evol.

13. Highlights
• Longstanding equilibrium
upended by recent
extinctions
• diversity is predictable
• time to recovery long
• phylogenies require
fossils to discover
patterns
Photo by Jon Flanders

15. What killed the mammals
of the Caribbean?
Deglaciation!
© Adrian Tejedor

16. Islands of the Caribbean:
The West Indies
Dávalos,& Turvey 2012 Bones,
Clones, and Biomes

17. Dávalos & Russell 2012 Ecol. Evol.

18. Trouble is we didn’t have
the dates
Soto-Centeno & Steadman 2015 Sci. Rep.

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

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

23. 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

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

25. 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.

26. 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.

27. A few traits
• Bat extinction
• Larger species
• Herbivores
• Among non-bats
• Larger
• Smaller?!
Cooke, Dávalos et al. 2017 Annu. Rev.
Ecol. Evol. Syst.

28. Mammals of the Caribbean
© Yomangani/Creative Commons © Eladio Fernandez
© Jon Flanders

29. 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.

30. Modeling extinction/
survival
Unpublished

31. Modeling extinction/
survival
Unpublished
Spp. traits
Island characteristics
Mass is quadratic
Many island characteristics
can be added

32. Intercept on probability of survival
−2000 −1000 0 1000 2000
Standard deviation
Mean
Zygodontomys_sp
Xenothrix_mcgregori
Xaymaca_fulvopulvis
Tainotherium_valei
Solenodon_paradoxus
Solenodon_marcanoi
Solenodon_cubanus
Solenodon_arredondoi
Sheep_Pen_femur_unidentified_rodent_taxon
Rhizoplagiodontia_lemkei
Quemisia_gravis
Plagiodontia_velozi
Plagiodontia_spelaeum
Plagiodontia_aedium
Pennatomys_nivalis
Parocnus_serus
Parocnus_browni
Paralouatta_varonai
Oryzomys_antillarum
Oryzomyini_undescribed_species_small_Montserrat_taxon
Oryzomyini_undescribed_species_small_Grenada_taxon
Oryzomyini_undescribed_species_large_Montserrat_taxon
Oryzomyini_undescribed_species_large_Grenada_taxon
Oryzomyini_undescribed_species_St._Martin_taxon
Oryzomyini_undescribed_species_Saba_taxon
Oryzomyini_undescribed_species_Anguilla_taxon
Oligoryzomys_victus
Nesophontes_zamicrus
Nesophontes_sp2
Nesophontes_sp1
Nesophontes_paramicrus
Nesophontes_micrus
Nesophontes_hypomicrus
Nesophontes_edithae
Neocnus_toupiti
Neocnus_gliriformis
Neocnus_dousman
Neocnus_comes
Mysateles_prehensilis
Mesocapromys_sanfelipensis
Mesocapromys_nanus
Mesocapromys_melanurus
Mesocapromys_kraglievichi
Mesocapromys_auritus
Mesocapromys_angelcabrerai
Megalomys_luciae
Megalomys_georginae
Megalomys_desmarestii
Megalomys_audreyae
Megalocnus_zile
Megalocnus_rodens
Macrocapromys_latus
Macrocapromys_acevedo
Isolobodon_portoricensis
Isolobodon_montanus
Insulacebus_toussaintiana
Hyperplagiodontia_araeum
Hexolobodon_phenax
Heteropsomys_insulans
Geocapromys_thoracatus
Geocapromys_sp2
Geocapromys_sp1
Geocapromys_ingrahami
Geocapromys_columbianus
Geocapromys_brownii
Elasmodontomys_obliquus
Clidomys_osborni
Capromys_sp1
Capromys_pilorides
Brotomys_voratus
Brotomys_contractus
Boromys_torrei
Boromys_offella
Antillothrix_bernensis
Antillomys_rayi
Acratocnus_ye
Acratocnus_simorhynchus
Acratocnus_odontrigonus
Acratocnus_antillensis
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
Initial results species
• Species vary in
probability of survival
• Some appear doomed

33. Initial results islands
• Multiple species were
extinction traps

34. Initial results
predictors
• Island area weakly
related to survival
• Elevation strong
predictor extinction
• Intuition about large
and small spp. correct!

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

36. 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

37. Larger mammals probably hunted early on
© Ghedoghedo Wilson 1989 J Field Archaeol.

38. Many extinction events recorded after arrival of Old
World invasive competitors/predators
Turvey et al. 2012 Mamm. Biol.

39. Agriculture likely important for certain species
© Adrian Tejedor ©Pierpont Morgan Library

40. 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

41. Why is this important? The world is made of islands

42. This is happening now Ray et al. 2015 Mongabay

43. What happens when the
frontier closes?
1869

44. Frontier closes, forest
vanishes
Stanton 2014 Biol. Cons.

45. 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.
Fractionforest

46. The Andean frontier A general model
Fractionforest
coca beachhead
Time ->
mixed agriculture
properties
pastures!

47. 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

48. Thanks!