Resilience and Innovation Resilience and Innovation

in Complex Systemsin Complex Systems

ResilienceResilience

Resilience is the ability of an system to remain within a state.

Resilience is a measure of the amount of perturbation required to cause a system to change states

Resilience is a cross-scale measure. Ecological resilience depends upon:

– Control of disturbance

– Regulation of renewal

Overlapping Function Within a ScaleOverlapping Function Within a Scale

At the same scale species from different functional groups specialize in the use of different resources, but each guild can use other resources at lower efficiencies.

Use of different resources at same scale

Guild C

Guild B

Guild A

Guild D

Functional Reinforcement Across ScalesFunctional Reinforcement Across Scales

Scale (species body mass)

Guild A

Species from the same guild can utilize the same resource at different scales. Resources that are more dispersed can be utilized be a large animal, but with a decrease in efficiency. Species are performing similar functions at different scales.

Birds and BudwormBirds and Budworm

Use of different resources at same scale

Scale (species body mass)

NectivoreCarnivoreGranivoreInsectivores

As budworm populations increase and occur larger aggregation both larger birds and birds that would not normally consume budworm switch to the use of budworm.This process provides robust control of budworm populations over a wide range of budworm densities.

predation of budworm at different aggregations

small birds large birds

Cross-Scale ResilienceCross-Scale Resilience

Use of different resources at same scale

Scale (species body mass)

Guild DGuild CGuild BGuild A

Overlapping function within scales and functional reinforcement across scales.

At the same scale species from different guilds specialize in the use of different resources, but each guild can use other resources at lower efficiencies. Species in a guild utilize the same resource, but at different scales.

Ecological ScalingEcological Scaling

Scale is the spatial and temporal frequency of a process or structure.

A scale domain is bounded by the grain size of processes detected and the extent or span of processes attended.

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Vegetative ScalesVegetative Scales

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regionBoreal forest, forExample, is patterned across a range of scales.

Larger slower structures usually constrain the behavior of faster smaller scales.

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DisturbanceProcesses

Vegetation

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food choice

home range

dispersal & migration

moosebeaver

deer mouse

Herbivorous Mammals

Different species interact with the landscape at different scales.

An animal makes different decisions at different scales.

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Animal body mass patterns reflect the scale-specific domains of key structuring process.

Textural Discontinuity Hypothesis

Vertebrate body masses are discontinuous

Increasing body mass

A.

B.

C.

Cross-Scale ResilienceCross-Scale Resilience

Use of different resources at same scale

Scale (species body mass)

Guild DGuild CGuild BGuild A

Overlapping function within scales and functional reinforcement across scales.

At the same scale species from different guilds specialize in the use of different resources, but each guild can use other resources at lower efficiencies. Species in a guild utilize the same resource, but at different scales.

© Photohome.com

MethodsMethods

U.S. Census data set incorporating the U.S. Census data set incorporating the Urbanized Area definitionUrbanized Area definition

Bureau of Economic Analysis (BEA) Bureau of Economic Analysis (BEA) regionsregions

Data from the southwestern region of the Data from the southwestern region of the United StatesUnited States

MethodsMethods

Gap Rarity Index (GRI)Gap Rarity Index (GRI) Split moving-window (SMW) Split moving-window (SMW)

boundary analysis boundary analysis Hierarchical cluster analysis Hierarchical cluster analysis Aggregation = 3 or more cities Aggregation = 3 or more cities 48 cities in 1890 to 161 cities in 199048 cities in 1890 to 161 cities in 1990

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3.40 3.90 4.40 4.90 5.40 5.90 6.40

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Log10 (City Size)1 2 3 4 5 6 7

Year Largest City1890 Dallas

1900 San Antonio

1910 San Antonio

1920 San Antonio

1930 Houston

1940 Houston

1950 Houston

1960 Houston

1970 Houston

1980 Dallas-Fort Worth

1990 Dallas-Fort Worth

City size distributions for the southwestern region of the United States were discontinuous

Distinct aggregations of cities were identified in each decade, by all methods of analysis.

We observed 4 to 7 aggregations in each decadal dataset

Maintenance of similar discontinuous structure over time suggests this structure is conserved and resilient.

How does the discontinuous distribution of body masses, or other complex system components (e.g., cities)relate to the emergence of innovation and novelty?

The South Florida The South Florida terrestrial terrestrial vertebrate vertebrate faunafauna

NATIVE DECLINE N.I.S.

MAMMALS 35 10(29%) 10(22%)

BIRDS 120 22(18%) 32(21%)

HERPS 49 7(14%) 21(30%)

Historic body mass Historic body mass structure,structure,South Florida vertebratesSouth Florida vertebrates

mam

mal

sm

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als

herp

sbi

rds

Increasing body mass

Tests of selected Tests of selected hypotheses in South hypotheses in South Florida:Florida: invasive species don't tend to be smaller than native speciesinvasive species don't tend to be smaller than native species endangered species don't tend to be larger than non-endangered species don't tend to be larger than non-

endangered native speciesendangered native species no gross differences in trophic level between endangered no gross differences in trophic level between endangered

and invasive speciesand invasive species no niche replacementno niche replacement no phylogenetic influence on endangermentno phylogenetic influence on endangerment

Taxonomic groups as replicates; Mann-Whitney Utests, Fisher's exact test, chi-square goodness-of-fit.

Random

In lumps

In gaps

Within asingle scale

At lump edges

Possible patterns of invasive or declining species in terms of lump structure

Null hypothesis.No relationship withbody mass structure.

Hypothesissupported

Gaps are repellant.

Morphologicaloverdispersion -competitivestructuring.

Scale-specific effect or

.Turbulencebetween rangesof scale.

phylogeny

Historic body mass Historic body mass structure,structure,South Florida vertebratesSouth Florida vertebrates

mam

mal

sm

amm

als

herp

sbi

rds

Increasing body mass

Species Rank

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Everglades Mammals

INVASIVE SPECIES AND ENDANGERED INVASIVE SPECIES AND ENDANGERED SPECIES ARE NON-RANDOMLY SPECIES ARE NON-RANDOMLY DISTRIBUTED IN RELATION TO THE DISTRIBUTED IN RELATION TO THE BODY MASS DISTRIBUTIONS OF BODY MASS DISTRIBUTIONS OF EVERGLADES ECOSYSTEM VERTEBRATESEVERGLADES ECOSYSTEM VERTEBRATES Invasive species are Invasive species are

successful at the edge of successful at the edge of aggregationsaggregations

Endangered species occur Endangered species occur at the edge of aggregationsat the edge of aggregations

Results are similar for all Results are similar for all taxonomic groups analyzedtaxonomic groups analyzed

RUSSIACANADA

UNITED STATES

MEXICO

CHINA

MONGOLIAKAZAKHSTAN

INDIA

UZBEKISTAN

PAKISTAN

AFGHANISTAN

TURKMENISTAN

KYRGYZSTAN

TAJIKISTAN

NEPALBHUTAN

BANGLADESH

BURMAVIETNAM

THAILAND

LAOS

CAMBODIA

MALAYSIA

INDONESIAPAPUA NEW GUINEA

AUSTRALIA

NEW ZEALAND

GUATEMALA

BELIZEHONDURAS

EL SALVADOR

COSTA RICA

PANAMA

DOMINICAN REPUBLIC

CUBAHAITI

VENEZUELA

COLOMBIA

ECUADOR

PERU

BOLIVIA

GUYANASURINAME

FRENCH GUIANA

BRAZIL

ARGENTINA

CHILE

PARAGUAY

URUGUAY

JAPAN

NORTH KOREA

SOUTH KOREA

PHILIPPINES

TAIWAN

WHY?WHY? NOT due to morphological overdispersion. NOT due to morphological overdispersion.

With the addition of exotics, the result is not a With the addition of exotics, the result is not a continuous distribution; exotics aren't "filling in the continuous distribution; exotics aren't "filling in the gaps".gaps".

Gaps may be analogous to zones of phase transition, Gaps may be analogous to zones of phase transition, and represent zones of crisis and opportunity for and represent zones of crisis and opportunity for species exploiting them.species exploiting them.

Other ecosystems and faunas should reveal the same association between population status (invasive, declining)and body-mass lump edges.

PREDICTION:

TEST:Mexican cave bat community (declining/vulnerable species)Mediterranean-climate faunas (invasive)

Mexican cave bats: - relatively large proportion of vulnerable species- well studied: status not politically derived

Mediterranean -climate faunas:- well studied- relatively large number of invasions

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Mexican Cave Bats. Association with scale breaks

Dist. to Edge: Median, End. = 0.00, Median safe = 0.02, M-W P = 0.014

Location N Obs. Exp. PMAMMALS

California 2 2 0.38Chile 11 5 1.63Greece 4 2 0.58South Africa 8 1 1.11Southwestern Australia 8 2 1.14Southcentral Australia 10 5 1.61 0.0001

BIRDS California 18 3 1.59Chile 7 2 0.85South Africa 8 5 0.89Spain 4 1 0.47Turkey 3 0 0.23Southwestern Australia 9 4 1.02 <0.0001

Successful introductions should have body masses closer to body-mass lump edges than unsuccessful introductions.

PREDICTION:

TEST:

Avian introductions in the Everglades sub-ecoregion:

- Well documented- Large number in both categories- Sufficient information to differentiate between species withand without a sufficient propagule

Successful v. unsuccessful introductions in south FloridaLogistic regression

35 successful introductions45 unsuccessful introductions

Variables: body mass, distance to nearest neighbor (in terms of body mass), year of introduction, presence of congeners, distribution in a body mass aggregation or gap, and distance to body-mass aggregation edge (in terms of body mass).

* Successful introductions have body masses that place them closer to body-mass lump edges.

Other variable phenomena reflecting species turnover should be associated with the edge of body mass aggregations.

PREDICTION:

TESTS:

Nomadic birds in South Central Australia:

- Well documented- Large number in both categories

Migrant birds in South Florida, South Carolina and Costa Rica:- Well documented

Significant predictors ofnomadism:(model selection based on AIC)

Parameter WaldVariable estimate Chi-Square Probability

Mass 1.31 14.97 0.0001

Edge -1.21 6.42 0.0113

MedianClass edge distance N T P

Migratory 0.00133 10

Permanent residents 0.0108 96 374.0 0.042

Migratory birds in the south Florida everglades ecosystem:

Migratory birds body masses place them 10 times closer, onaverage, to body mass aggregation edges.

Log Body Mass

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Mammals

Herpetofauna

Avifauna

Historic - Future comparisons of the South Florida fauna

Invasions and extinctions represent turnover in species composition(25% declining in SFL, a further 25% are non-native)

The generation of innovation in complex systems? Entrainment

Hierarchically Structured Landscape

Discontinuous Animal Size Distributions

Species Pool, Continuous Distribution ?

Self Organization

SCALE-SPECIFIC STRUCTURING PROCESSES

Stochastic Climatic and Edaphic Patterns

Vegetation Architecture

Forest Stands Patch

Structure

Forest Patch

Structure

Clump 1 Clump 2 Clump 3

Character Displacement and/or Species Assortment

Over-Dispersed Size Distribution Within Clump

Gap Gap

Competition?

Inter-specific Interactions

Increasing Size

The generation of innovation in complex systems?

Increasing Size

Aggregations Discontinuities

Increasing Size

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Variability and innovation between ranges of scales?

The profile

-Nomadic birds in SC Australia.-Migrant birds, Everglades and South Carolina.-Invasive herpetofauna, birds, and mammals,Everglades.-Invasive species, global mediterranean-climate systems.-Declining herpetofauna, birds, and mammals, Everglades.-Declining cave bats, Mexico.

Variability between ranges of scales?The significance

-Understanding of the link between landscape change and the composition of species communities.

-Understanding complex phenomena such as invasions, migration and nomadism.

-Understanding how resilience is generated – turnover at scalebreaks does not alter gross system structure but propagatesinnovation.

- Insight into the evolution of complex behaviors and species.