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The last ones standing: why mammalogists should
care about Caribbean bats
Liliana M. Dávalos
Ecology and Evolution, SUNY at Stony Brook

Danny Rojas
Department of Biology and Centre for Environmental and Marine Studies,
University of Aveiro

ASM Meeting

Jacksonville, 15 June 2015
Caribbean mammal
fauna
• Depauperate
compared to mainland
• Shaped by extinction
• Native extant
mammals are bats
• Rest is rounding error
• Except for present
company
Drawing	
  by	
  A.	
  Tejedor
However, many
endemics = irreplaceable
Myers et al. 2000 Nature
This is not new
The number of peculiar
genera in all classes of animals
is so great in proportion to
those in common with the
adjacent mainland, as to lead
us to conclude that,
subsequent to the original
separation from the Mexican
area, a very large tract of land
existed, calculated to support
a rich and varied fauna, and,
by the interaction of
competing types, give rise to
peculiar and specially modified
organisms…
Wallace 1876 The Geographical Distribution
of Animals
How can we measure
this?
• Phylogenetic diversity (PD) = sum of
branch length between root and tips for
a community
• Identifies evolutionary information
added by each species
• Mean pairwise distance (MPD) = mean
distance between all pairs of species
• Reflects structuring across the entire
tree
• Mean nearest taxon distance (MNTD) =
mean of branch lengths from each
species to closest relative
• Reflects phylogenetic structure of tips
Winter et al. 2013 TREE
Hypothesis &
prediction
• Richness alone
severely
underestimates
Caribbean biodiversity
• Measures of
evolutionary diversity
(ED) will be
comparable to
communities on
mainland
Data criteria
• Published or collected
ourselves
• Local scale
• Community sampling
• Floor on richness
• ≥5 spp.
Photo by Jon Flanders
Island caves = 61
Mainland caves = 21
Mainland forests = 18
Photo	
  @	
  GEPOG
Photo	
  @	
  Guia	
  Cavernas	
  
Santander
Artibeus anderseni
Artibeus cinereus
Artibeus phaeotis
Artibeus watsoni
Artibeus bogotensisArtibeus glaucusArtibeus gnomusArtibeus amplusArtibeus planirostris
Artibeus jamaicensis
Artibeus obscurus
Artibeus lituratus
Artibeus concolor
Sphaeronycteris toxophyllum
Ametrida centurio
Centurio senex
Phyllops falcatus
Stenoderma rufum
Ariteus flavescens
Ectophylla
alba
Enchisthenes hartii
Platyrrhinus
lineatus
Platyrrhinus
helleri
Platyrrhinus
brachycephalus
Platyrrhinus
infuscus
Vam
pyrodes
caraccioli
Vam
pyressa
m
elissa
Vampyressapusilla
Mesophyllamacconnelli
Chirodermavillosum
Chirodermatrinitatum
Chirodermasalvini
Vampyriscusnymphaea
Vampyriscusbrocki
Vampyriscusbidens
Urodermabilobatum
Urodermamagnirostrum
Sturniratildae
Sturniraludovici
Sturniramagna
Sturniralilium
Sturniraluisi
Rhinophyllafischerae
Rhinophyllapumilio
Glyphonycterisdaviesi
Glyphonycterissylvestris
Trinycterisnicefori
Carolliabrevicauda
Carolliaperspicillata
Carolliacastanea
Lionycterisspurrelli
Lonchophylladekeyseri
Lonchophyllarobusta
Lonchophyllamordax
Hsunycteristhomasi
Phyllonycterispoeyiobtusa
Phyllonycterispoeyipoeyi
Phyllonycterisaphylla
Erophyllabombifrons
Erophyllasezekorni
Brachyphyllacavernarum
Brachyphyllapumila
Brachyphyllanana
Glossophagasoricina
Glossophaga
com
m
issarisi
M
onophyllusredm
ani
Hylonycteris
underwoodi
Lichonycteris
obscura
C
hoeroniscus
godm
ani
Choeroniscus
m
inor
Anoura
caudifer
Anoura
geoffroyi
Scleronycteris ega
Lonchorhina sp
Lonchorhina aurita
Phyllostomus elongatus
Phyllostomus hastatus
Phyllostomus latifolius
Phyllostomus discolor
Phylloderma stenops
Gardnerycteris crenulatum
Lophostoma brasiliense
Lophostoma carrikeri
Lophostoma schulzi
Lophostoma evotis
Lophostoma silvicolumTonatia bidensTonatia saurophilaMacrophyllum macrophyllum
Trachops cirrhosus
Chrotopterus auritus
Vampyrum spectrum
Mimon bennettii
Mimon cozumelae
Desmodus rotundus
Diaemus youngi
Diphylla ecaudata
Micronycteris megalotis
Micronycteris microtis
Micronycteris brosseti
Micronycteris hirsuta
Micronycteris homezi
Micronycteris minuta
Micronycteris sanborni
Micronycteris schmidtorum
Lampronycteris brachyotis
Macrotus waterhousii
Pteronotus macleayii
Pteronotus quadridens
Pteronotus davyi
Pteronotus gym
nonotus
Pteronotus
parnellii
M
orm
oops
blainvillei
M
orm
oops
m
egalophylla
Noctilio
albiventris
Noctilio
leporinus
Furipterushorrens
Thyroptera
discifera
Thyropteratricolor
Peropteryxmacrotis
Peropteryxkappleri
Cormurabrevirostris
Peropteryxleucoptera
Diclidurusalbus
Diclidurusscutatus
Cyttaropsalecto
Saccopteryxcanescens
Saccopteryxgymnura
Saccopteryxbilineata
Saccopteryxleptura
CentronycterismaximilianiCentronycteriscentralisRhynchonycterisnaso
Cynomopsparanus
Cynomopsabrasus
Cynomopsgreenhalli
Cynomopsplanirostris
Neoplatymopsmattogrossensis
Molossopstemminckii
MolossopsneglectusNyctinomopslaticaudatusNyctinomopsmacrotisEumopsglaucinusEumopstrumbulli
Eumopsauripendulus
Eumopshansae
Molossusrufus
Molossuscoibensis
Molossussinaloae
Molossuscurrentium
Molossusbarnesi
Molossusmolossus
Promopscentralis
Tadaridabrasiliensis
Eptesicusfurinalis
Eptesicusbrasiliensis
Eptesicus
dim
inutus
Eptesicus
chiriquinus
Eptesicus
lynni
Eptesicus
fuscus
Rhogeessa
tum
ida
Bauerus dubiaquercus
Pipistrellus subflavus
Lasiurus blossevillii
Lasiurus atratus
Lasiurus ega
Myotis riparius
Myotis elegans
Myotis simus
Myotis keaysi
Myotis albescens
Myotis nigricans
Myotis fortidens
Natalus major
Natalus jamaicensis
Natalus primus
Natalus stramineus
Natalus tumidirostris
Chilonatalus micropus
Chilonatalus macer
Nyctiellus lepidus
We need a phylogeny to
calculate ED
Shi & Rabosky 2015 Evolution

Rojas et al. Accepted Syst Biol
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500
1000
island caves mainland caves mainland forests
Geographic location
Faith'sphylogeneticdiversity
Mainland forests win PD,
caves are alike, though
PD = evo info added by species
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20 40 60 80
Species richness
Faith'sphylogeneticdiversity
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island caves
mainland caves
mainland forests
PD strongly depends on
richness
PD = evo info added by species
−4
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−1
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1
island caves mainland caves mainland forests
Geographic location
StandardizedeffectsizeofMPDvs.nullcommunities
Island caves win MPD Standardized to richness
−3
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−1
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1
2
island caves mainland caves mainland forests
Geographic location
StandardizedeffectsizeofMNTDvs.nullcommunities
Mainland caves win
MNTD
Standardized to richness
Extinction
experiments
• Estimate effect of
losing 1 species
• Remove species
from phylogeny =
extinct
• Remove species
from community =
extirpate
Photo by Jon Flanders
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island caves mainland caves mainland forests
Geographic location
DifferenceinmeanFaith'sphylogeneticdiversity
betweenthemodifiedcommunitiesandtheoriginalcommunities
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−3
−2
−1
0
island caves mainland caves mainland forests
Geographic location
DifferenceinmeanFaith'sphylogeneticdiversity
betweenthemodifiedcommunitiesandtheoriginalcommunities
Extirpation affects PD of mainland forests the most,
island caves next
Photo by Jon Flanders
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−0.1
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island caves mainland caves mainland forests
Geographic location
DifferenceinmeanMPD
betweenthemodifiedcommunitiesandtheoriginalcommunities
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−0.075
−0.050
−0.025
0.000
0.025
island caves mainland caves mainland forests
Geographic location
DifferenceinmeanMPD
betweenthemodifiedcommunitiesandtheoriginalcommunities
Extirpation affects MPD of mainland caves the most,
mainland forests next
Photo by Laurel Yohe
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island caves mainland caves mainland forests
Geographic location
DifferenceinmeanMNTD
betweenthemodifiedcommunitiesandtheoriginalcommunities
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−0.025
0.000
0.025
0.050
island caves mainland caves mainland forests
Geographic location
DifferenceinmeanMNTD
betweenthemodifiedcommunitiesandtheoriginalcommunities
Extirpation affects MNTD of mainland forests the most,
island caves next
Photo by Laurel Yohe
To summarize
• Island caves
• Richness = mainland caves
• MPD > mainland
• MNTD > mainland forests
• Extirpation on island caves
• Reduces MPD more than
for mainland caves
• Increases MNTD as for
mainland forests Photo by Jon Flanders
Were we right?
• Richness underestimates
island biodiversity
• Yes, richness also
determines PD
• ED comparable to mainland
communities
• Yes, MPD and MNTD
• Effect of loss comparable to
mainland or even greater
• Endemicity
• Functional diversity (?)
Photo by Jon Flanders
Thanks for inviting me!

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The last ones standing: why mammalogists should care about Caribbean bats

  • 1. The last ones standing: why mammalogists should care about Caribbean bats Liliana M. Dávalos Ecology and Evolution, SUNY at Stony Brook Danny Rojas Department of Biology and Centre for Environmental and Marine Studies, University of Aveiro ASM Meeting Jacksonville, 15 June 2015
  • 2. Caribbean mammal fauna • Depauperate compared to mainland • Shaped by extinction • Native extant mammals are bats • Rest is rounding error • Except for present company Drawing  by  A.  Tejedor
  • 3. However, many endemics = irreplaceable Myers et al. 2000 Nature
  • 4. This is not new The number of peculiar genera in all classes of animals is so great in proportion to those in common with the adjacent mainland, as to lead us to conclude that, subsequent to the original separation from the Mexican area, a very large tract of land existed, calculated to support a rich and varied fauna, and, by the interaction of competing types, give rise to peculiar and specially modified organisms… Wallace 1876 The Geographical Distribution of Animals
  • 5. How can we measure this? • Phylogenetic diversity (PD) = sum of branch length between root and tips for a community • Identifies evolutionary information added by each species • Mean pairwise distance (MPD) = mean distance between all pairs of species • Reflects structuring across the entire tree • Mean nearest taxon distance (MNTD) = mean of branch lengths from each species to closest relative • Reflects phylogenetic structure of tips Winter et al. 2013 TREE
  • 6. Hypothesis & prediction • Richness alone severely underestimates Caribbean biodiversity • Measures of evolutionary diversity (ED) will be comparable to communities on mainland
  • 7. Data criteria • Published or collected ourselves • Local scale • Community sampling • Floor on richness • ≥5 spp. Photo by Jon Flanders
  • 8. Island caves = 61 Mainland caves = 21 Mainland forests = 18 Photo  @  GEPOG Photo  @  Guia  Cavernas   Santander
  • 9. Artibeus anderseni Artibeus cinereus Artibeus phaeotis Artibeus watsoni Artibeus bogotensisArtibeus glaucusArtibeus gnomusArtibeus amplusArtibeus planirostris Artibeus jamaicensis Artibeus obscurus Artibeus lituratus Artibeus concolor Sphaeronycteris toxophyllum Ametrida centurio Centurio senex Phyllops falcatus Stenoderma rufum Ariteus flavescens Ectophylla alba Enchisthenes hartii Platyrrhinus lineatus Platyrrhinus helleri Platyrrhinus brachycephalus Platyrrhinus infuscus Vam pyrodes caraccioli Vam pyressa m elissa Vampyressapusilla Mesophyllamacconnelli Chirodermavillosum Chirodermatrinitatum Chirodermasalvini Vampyriscusnymphaea Vampyriscusbrocki Vampyriscusbidens Urodermabilobatum Urodermamagnirostrum Sturniratildae Sturniraludovici Sturniramagna Sturniralilium Sturniraluisi Rhinophyllafischerae Rhinophyllapumilio Glyphonycterisdaviesi Glyphonycterissylvestris Trinycterisnicefori Carolliabrevicauda Carolliaperspicillata Carolliacastanea Lionycterisspurrelli Lonchophylladekeyseri Lonchophyllarobusta Lonchophyllamordax Hsunycteristhomasi Phyllonycterispoeyiobtusa Phyllonycterispoeyipoeyi Phyllonycterisaphylla Erophyllabombifrons Erophyllasezekorni Brachyphyllacavernarum Brachyphyllapumila Brachyphyllanana Glossophagasoricina Glossophaga com m issarisi M onophyllusredm ani Hylonycteris underwoodi Lichonycteris obscura C hoeroniscus godm ani Choeroniscus m inor Anoura caudifer Anoura geoffroyi Scleronycteris ega Lonchorhina sp Lonchorhina aurita Phyllostomus elongatus Phyllostomus hastatus Phyllostomus latifolius Phyllostomus discolor Phylloderma stenops Gardnerycteris crenulatum Lophostoma brasiliense Lophostoma carrikeri Lophostoma schulzi Lophostoma evotis Lophostoma silvicolumTonatia bidensTonatia saurophilaMacrophyllum macrophyllum Trachops cirrhosus Chrotopterus auritus Vampyrum spectrum Mimon bennettii Mimon cozumelae Desmodus rotundus Diaemus youngi Diphylla ecaudata Micronycteris megalotis Micronycteris microtis Micronycteris brosseti Micronycteris hirsuta Micronycteris homezi Micronycteris minuta Micronycteris sanborni Micronycteris schmidtorum Lampronycteris brachyotis Macrotus waterhousii Pteronotus macleayii Pteronotus quadridens Pteronotus davyi Pteronotus gym nonotus Pteronotus parnellii M orm oops blainvillei M orm oops m egalophylla Noctilio albiventris Noctilio leporinus Furipterushorrens Thyroptera discifera Thyropteratricolor Peropteryxmacrotis Peropteryxkappleri Cormurabrevirostris Peropteryxleucoptera Diclidurusalbus Diclidurusscutatus Cyttaropsalecto Saccopteryxcanescens Saccopteryxgymnura Saccopteryxbilineata Saccopteryxleptura CentronycterismaximilianiCentronycteriscentralisRhynchonycterisnaso Cynomopsparanus Cynomopsabrasus Cynomopsgreenhalli Cynomopsplanirostris Neoplatymopsmattogrossensis Molossopstemminckii MolossopsneglectusNyctinomopslaticaudatusNyctinomopsmacrotisEumopsglaucinusEumopstrumbulli Eumopsauripendulus Eumopshansae Molossusrufus Molossuscoibensis Molossussinaloae Molossuscurrentium Molossusbarnesi Molossusmolossus Promopscentralis Tadaridabrasiliensis Eptesicusfurinalis Eptesicusbrasiliensis Eptesicus dim inutus Eptesicus chiriquinus Eptesicus lynni Eptesicus fuscus Rhogeessa tum ida Bauerus dubiaquercus Pipistrellus subflavus Lasiurus blossevillii Lasiurus atratus Lasiurus ega Myotis riparius Myotis elegans Myotis simus Myotis keaysi Myotis albescens Myotis nigricans Myotis fortidens Natalus major Natalus jamaicensis Natalus primus Natalus stramineus Natalus tumidirostris Chilonatalus micropus Chilonatalus macer Nyctiellus lepidus We need a phylogeny to calculate ED Shi & Rabosky 2015 Evolution Rojas et al. Accepted Syst Biol
  • 10. ● ● ● ● ● ● 500 1000 island caves mainland caves mainland forests Geographic location Faith'sphylogeneticdiversity Mainland forests win PD, caves are alike, though PD = evo info added by species
  • 12. −4 −3 −2 −1 0 1 island caves mainland caves mainland forests Geographic location StandardizedeffectsizeofMPDvs.nullcommunities Island caves win MPD Standardized to richness
  • 13. −3 −2 −1 0 1 2 island caves mainland caves mainland forests Geographic location StandardizedeffectsizeofMNTDvs.nullcommunities Mainland caves win MNTD Standardized to richness
  • 14. Extinction experiments • Estimate effect of losing 1 species • Remove species from phylogeny = extinct • Remove species from community = extirpate Photo by Jon Flanders
  • 15. ● ●● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ●● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ●● ● ● ● ● ●● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● −20 −10 0 island caves mainland caves mainland forests Geographic location DifferenceinmeanFaith'sphylogeneticdiversity betweenthemodifiedcommunitiesandtheoriginalcommunities ●● ● ●●● ●●● ● ● ●● ● ●●●●●●●● ● ● ●●●●●●●●●●●●● ● ●●●● ● ● ●● ●●●● ● ●●●●●● ● ● ● ● ● ●●●●●●●●● ● ● ●● ●●● ● ●● ● ● ●●●●● ●● ● ● ●● ● ●●● ●●●● ●●●●● ● ●●●●● ●●●●●●●● ●●●● ● ●●● ● ●●●●●●●●●●●●●● ● ●● ● ● ● ● ● ●●●●●●● ●● ●●●● ● ●●●●● ● ● ●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●● ● ●●●●●●●●●●●●●●●●●●●● ● ●●●●● ● ●●●●●●●●●●●● ●●●●●●●●●●●●●●●●●●●●● ● ●●●●● ● ●●● ● ● ● ●●●● ● ●●● ● ● ● ● ● ●● ●● ● ●●●●● ●● ● ● ● ● ●●● ●● ● ● ● ●●●● ● ●● ● ●●●●●●●●●●●●●●● ● ●● ● ●●●●●●●●● ● ●●●● ●●● ● ●● ● ● ●● ● ●●●●● ● ● ● ●●● ● ●● ●● ●● ● ● ●●●●●●●● ● ●●● ● ● ●● ● ● ●● ● ● ● ●●●● ● ●● ● ● ●●● ● ●● ● ● ● ● ● ● ● ● ● ● ●●● ●● ● ● ●●● ● ● ●● ● ● ● ●●●●● ● ● ●● ● ●●● ● ● ● ●●●●●● ●●● ● ● ●●●●●●● ●● ●● ●●● ●● ●●●●●●●●●●● ● ● ● ● ● ● ● ●● ● ●●● ● ● ● ●● −3 −2 −1 0 island caves mainland caves mainland forests Geographic location DifferenceinmeanFaith'sphylogeneticdiversity betweenthemodifiedcommunitiesandtheoriginalcommunities Extirpation affects PD of mainland forests the most, island caves next Photo by Jon Flanders
  • 16. ● ● ● ● ● ● ● ● ● ● ●● ● ● ● ● ● ● ● ● ● ● ● −0.2 −0.1 0.0 0.1 0.2 island caves mainland caves mainland forests Geographic location DifferenceinmeanMPD betweenthemodifiedcommunitiesandtheoriginalcommunities ● ● ●●●●● ● ● ●● ● ● ● ● ● ● ● ● ● ● ● ●●●● ●●● ● ● ● ●● ● ● ● ● ● ● ● ● ● ● ● ● ● ●● ● ● ● ● ●● ● ● ●● ● ● ● ●●●●●● ● ●●● ●●●●●●● ● ●●●● ● ● ● ● ●● ●● ● ●●●● ● ● ● ●● ● ● ● ● ● ● ● ● ● ● ● ●● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ●● ● ●●●●●●● ● ● ● ● ● ● ● ● ● ● ●● ● ●● ● ● ● ● ● ●● ●●● ●● ●● ● ● ●● ● ● ●● ● ● ● ● ● ● ● ●● ● ●● ● ● ●● ● ● ● ● ● ● ● ● ● ●● ● ● ● ● ●●●● ● ● ● ●● −0.075 −0.050 −0.025 0.000 0.025 island caves mainland caves mainland forests Geographic location DifferenceinmeanMPD betweenthemodifiedcommunitiesandtheoriginalcommunities Extirpation affects MPD of mainland caves the most, mainland forests next Photo by Laurel Yohe
  • 17. ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ●● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ●● ● ● ● ● ●● ● ●●● −0.2 0.0 0.2 island caves mainland caves mainland forests Geographic location DifferenceinmeanMNTD betweenthemodifiedcommunitiesandtheoriginalcommunities ●● ● ● ●● ● ● ● ● ● ●●● ● ● ●● ● ●●●●●●●● ● ●●● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ●● ● ● ● ● ● ●● ● ●● ●●● ● ●● ● ●● ● ● ● ●● ● ●●●● ● ● ● ● ● ● ●● ● ● ● ● ● ● ● ● ● ● ●●● ● ● ● ●●● ● ● ●●● ●● ●● ● ● ●● ● ● ● ● ●● ● ● ● ● ● ● ● ● ●● ●● ● ● ●●●● ● ● ● ● ●● ●● ● ●● ● ● ● ● ● ● ●● ● ● ●● ●●● ● ● ●● ● ● ● ● ● ●● ● ● ●● ● ● ●●● ● ●●● ●● ●● ● ● ● ● ● ● ● ● ● ●●● ● ●● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ●●● ● ● ● ● ● ● ● ●● ● ● ●● ●●● ● ● ● ●● ● ● ●● ● ● ● ● ● ●● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ●● ● ● ●● ● ● ● ● ● ● ● ● ● ●● ● ● ● ● ●●● ● ● ● ● ● ● ● ● ●● ● ●● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ●● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ●● ● ● ●●●● ●● ● ● ● ● ● ● ● ● ● ●● ●● ● ● ● ● ●●● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ●● ● ● ●● ● −0.025 0.000 0.025 0.050 island caves mainland caves mainland forests Geographic location DifferenceinmeanMNTD betweenthemodifiedcommunitiesandtheoriginalcommunities Extirpation affects MNTD of mainland forests the most, island caves next Photo by Laurel Yohe
  • 18. To summarize • Island caves • Richness = mainland caves • MPD > mainland • MNTD > mainland forests • Extirpation on island caves • Reduces MPD more than for mainland caves • Increases MNTD as for mainland forests Photo by Jon Flanders
  • 19. Were we right? • Richness underestimates island biodiversity • Yes, richness also determines PD • ED comparable to mainland communities • Yes, MPD and MNTD • Effect of loss comparable to mainland or even greater • Endemicity • Functional diversity (?) Photo by Jon Flanders