But, What Do We Mean by “Species”?

• ESA and CITES define species as “any subspecies of fish or wildlife or plants, and any distinct population segment of any species of vertebrate fish or wildlife which interbreeds when mature”

– plant populations do not get special consideration

– NMFS often lists fish stocks (local non-interbreeding populations)

• concept of Evolutionary Significant Unit (requires reproductive isolation)

What is Actually Listed?• Wilcove et al.’s Analysis suggested that most listed

species were full species– only 20% of listed species were subspecies or

populations, but this varied by taxonomic group• birds---80% of listed “species” were subspecies or

populations• mammals--70% of listed “species” were subspecies or

populations• Mollusks--5%of listed “species” were subspecies or

populations• Plants--14% of listed “species” were subspecies or

populations

Biological vs. Evolutionary Species Concepts

• ESA uses a biological species concept because it emphasizes that groups to be listed are reproductively isolated from other such groups

• But, if the goal is to preserve biodiversity, then what we really want to preserve is unique genetic material, thus the evolutionary species concept may be more appropriate

Biological vs. Evolutionary Species Concepts

• Evolutionary species are those lineages that maintain their “own evolutionary tendencies and historical fates” (Wiley 1981)

• The National Research Council (NRC) review of the ESA discusses the EU and suggests it may be an especially valuable way to view species for listing– Evolutionary Unit-- “group of organisms that

represents a segment of biological diversity that shares evolutionary lineage and contains the potential for a unique evolutionary future”

Evolutionary Significant Units

•Others have defined EUs, or more generally, ESUs•Reviewed in Crandall et al. 2000

•Ryder 1986: populations that actually represent significant adaptive variation based on concordance between sets of data derived by different techniques

•Waples 1991: populations that are reproductively separate from other populations and have unique or different adaptations

•Moritz 1994: populations that are reciprocally monophyletic for mtDNA alleles and show significant divergence of allele frequences at nuclear loci

Ecological and Genetic Exchangeability

•Ecological exchangeability: the factors that define the fundamental niche and the limits of spread of new genetic variants through genetic drift and natural selection

•rejected with evidence for population differentiation owing to genetic drift or natural selection

•Differences in life histories, morphology, habitat, allozymes under selection (preferably heritable ones)

•Genetic exchangeability: the factors that define the limits and spread of new genetic variants through gene flow

•Rejected when there is evidence of restricted gene flow between populations

•Differences in microsatellites, nucleotide sequences, (mtDNA, cpDNA, nDNA) and allozymes

(Crandall et al. 2000)

Assessing Ecological and

Genetic Exchangeability

(Crandall et al. 2000)

Why Conserve Habitat?

• Critical to species’ survival

• Protection applies to more than just the species of interest

• Know more about habitat hot spots and distribution than about species distributions

• Know habitat loss and degradation are major reasons for endangerment

Metapopulation Review

• Subpopulations connected by dispersal (Levins 1969)

• Good way to describe structure and dynamics of populations scattered across a landscape in spatially isolated patches

– common in managed landscapes

• Some sub-populations may be sinks and some may be sources, but this is only a special case of general metapopulation model– core-satellite or simultaneous sink-source may be

more common (Doak and Mills 1994; Doncaster et al. 1997)

Key Messages for Endangered Species Management

• Extinction of subpopulations in metapopulation is to be expected

• Subpopulation dynamics may be controlled by dynamics of other subpopulations– rescue by dispersal– need to ID sources or cores

• Functioning metapopulation may be necessary for species to remain extant– Acorn Woodpeckers in New Mexico

• (Stacey and Taper 1992)

Another Key: Habitat is Not Constant in Space or Time

• It is a “shifting mosaic” (Bormann and Likens 1979, Botkin and Sobel 1975)

– habitat composition in landscape changes naturally• usually slowly

• BWCA (continual change at replacement rate every 2-4

centuries from glaciation and succession)– fire has return rate of 20-200 years

– GPP may ~ Respiration at ecosystem scale (steady state), but individual stands change frequently

Management Implications of Shifting Mosaics

• Land management usually decreases time between disturbances

• may also affect spatial arrangement by increasing edge

• Endangered species may need change or may need specific disturbance state– Kirtland’s Warbler and Red-cockaded Woodpecker

Fire Wind

Clear-cutting

Tot

al B

i om

ass

Time (White Mountains, NH; Bormann and Likens 1979)

80

20

Do We Really Know Habitat Needs?

Spring

MalesFemales

Important

% o

f E

ach

Age

/Sex

In

Gro

up

10

Fall

AdultsHatchYear

1020

80

AG CN CR CS SS WI

AG CN CR CS SS WI

• Van Horne (1983)– abundance quality

• Yong et al. (1998)– Wilson’s Warblers in New

Mexico

– Habitat needs differ from spring to fall (breeeding to migration)

• cottonwood not used in spring

– Habitat needs differ from adults to subadults

• ag for juveniles, willow for adults

Habitat Conservation Plans

• The way habitat is protected on non-federal lands (rather than designation of critical habitat which applies to Fed lands)

• Allows non-federal landowners to get incidental take permit (Sect 10(a))– implementation of HCP “will, to the maximum

extent practicable, minimize and mitigate the impacts of such taking” and “not appreciably reduce the likelihood of survival and recovery of the species in the wild”

HCPs as a Solution to a Problem

• Services view HCPs as a way to balance a citizen’s right to use their property with the nation’s interest in conserving rare and endangered species

• Goal is to create “creative partnerships” between landowners wanting to develop their land and our natural heritage

Increase in HCPs

020406080

100120140160180200

No. of HCPs

90 91 92 93 94 95 96

• San Bruno Mtn. Cal (1983)• Over 200 in 1997, 200 more

in preparation• Range in size

– 1/2 acre lot (Fl. Scrub Jay)– 170,000 acres

• Plum Creek Timber– 100 years, 285listed and unlisted

species

– 1.6 million acres• WA DNR

– 70-100 years, 200 species

The HCP Process (USFWS 1998)

• Plan Development– permit application ($25)

– the plan

– document of compliance with NEPA

– implementation agreement

• Review– service

– public (published in Federal Register)

• Monitoring– service monitors compliance with HCP

Contents of HCP (USFWS 1998)

• Species covered (listed and non-listed)• Assessment of impacts of take• How take will be monitored, minimized, and mitigated• Plan for funding the proposed monitoring and

mitigation • Alternatives to take and why they are not being

adopted• Argument that taking will not reduce the species’

survival and recovery

Recent HCP Evaluation

• The National Center for Ecological Analysis and Synthesis reviewed many HCPs and their results echo those previously mentioned

• View their report here to better understand HCPs and evaluate their scientific validity

NCEAS evaluation of HCPs

• Kareiva et al. online here

• 43 focal plans, from 1997 and earlier

Types of Mitigation

How Well Was Each

Step Analyzed?

HCPs are not Recovery Plans• Another criticism is that HCPs often do little for the

listed species– Requirement is that plan MINIMIZES and MITIGATES take

• they do not have to contribute to RECOVERY• alternatives easily dismissed

– Rota’s proposed HCP would take 1/2 of Mariana Crow’s habitat!

– Balcones Canyonlands HCP (Texas) provided 12,000 ha, but science report called for 53,000 ha

• black-capped vireo is likely to go locally extinct

Limited Public Participation• A serious criticism from environmental

organizations– Years of negotiation between service and landowner

prior to review– Service does not have to use public comments

obtained during review when making their final decision

– Too much invested in negotiations to change after public comments

– Environmental organizations are out of loop and don’t like it

Making HCPs Better (Kaiser 1997)

• Require plan to boost, not reduce, populations of listed species

• Initial plan developed by scientists with no vested interests in planning area

• Wait for recovery plan before HCP is approved

– allows range-wide coordination of efforts

• Allow for adjustment even with “no surprise”– public funding for surprises– good monitoring and adaptive response

Critical Habitat DesignationCritical Habitat DesignationPacific Salmon and Steelhead (2005)Pacific Salmon and Steelhead (2005)– 19 Species or “ESUs” 19 Species or “ESUs”

– 13 in WA, OR, ID, 7 in CA13 in WA, OR, ID, 7 in CA

Southern Resident Killer WhalesSouthern Resident Killer Whales

Presentation by Donna Darm, NOAA Presentation by Donna Darm, NOAA FisheriesFisheries

Relevance of Critical Habitat Relevance of Critical Habitat

• Section 7(a)(2): all Section 7(a)(2): all federal agencies shall federal agencies shall ensure actions they ensure actions they fund, permit or carry fund, permit or carry out are not likely to:out are not likely to:

– jeopardize species’ jeopardize species’ continued existencecontinued existence

– destroy or adversely destroy or adversely modify critical habitat modify critical habitat

What is critical habitat?What is critical habitat?

• Occupied areas with Occupied areas with physical or biological physical or biological featuresfeatures– essential to essential to

conservation conservation – may require special may require special

managementmanagement• Unoccupied areas Unoccupied areas

essential for essential for conservationconservation ESA ESA §§3(5)3(5)

Step 1: Identify “Critical Habitat”Step 1: Identify “Critical Habitat”

• Map actual fish Map actual fish distribution distribution

• Verify features and Verify features and special managementspecial management

• Group by watershed into Group by watershed into “specific areas”“specific areas”

• Identify unoccupied areas Identify unoccupied areas essential for conservationessential for conservation

How does it get designated? How does it get designated?

• Best scientific dataBest scientific data• Consider economic, Consider economic,

national security, other national security, other relevant impactsrelevant impacts

• May exclude if benefits May exclude if benefits of exclusion outweigh of exclusion outweigh benefits of designationbenefits of designation

• (Unless exclusion (Unless exclusion results in extinction)results in extinction)

ESA ESA §4§4(b)(b)(2)(2)

Benefits of Designation

Benefits of Exclusion

ConservationAvoid

Impacts

Step 2: Consider Impacts, Balance Step 2: Consider Impacts, Balance BenefitsBenefits

• Benefit of designation is the protection of Benefit of designation is the protection of section 7 (no adverse modification by section 7 (no adverse modification by federal agencies)federal agencies)

• Designation also gives notice of areas Designation also gives notice of areas important to speciesimportant to species

• Benefits of exclusion (i.e., avoiding Benefits of exclusion (i.e., avoiding impacts) may be economic, national impacts) may be economic, national security, or “other”security, or “other”

Balancing Benefits (Economic)Balancing Benefits (Economic)

• Consider for exclusion areas with a relatively Consider for exclusion areas with a relatively high economic impact and a relatively low high economic impact and a relatively low conservation valueconservation value

• Made sense with salmon because:Made sense with salmon because:– Large number of habitat areasLarge number of habitat areas– Not all areas are equally important for conservationNot all areas are equally important for conservation

• Conservation threshold – don’t exclude if it Conservation threshold – don’t exclude if it significantly impedes conservationsignificantly impedes conservation

Relative Conservation Value of Relative Conservation Value of Habitat AreasHabitat Areas

Example Exclusion ScenarioExample Exclusion Scenario

Balancing Benefits (Other)Balancing Benefits (Other)

• National Security (military readiness National Security (military readiness during global war on terror)during global war on terror)

• Conservation partnerships on private land Conservation partnerships on private land (HCPs) = net conservation benefit(HCPs) = net conservation benefit

• Tribal sovereignty and self-determination Tribal sovereignty and self-determination (conservation partnership also relevant)(conservation partnership also relevant)

Net Occupied Habitat Areas = 23,447 miles - Dept. of Defense Sites - Indian Lands - 4 HCP Holders - Economic ImpactsNet Designated Habitat Areas = 20,630 miles

August 2005 - Final Critical Habitat for 12 ESUs of Salmon and Steelhead

Southern Resident Killer Southern Resident Killer WhalesWhales

• Very different species, very different Very different species, very different habitathabitat

• Marine versus fresh waterMarine versus fresh water

• Habitat less affected at small scales by Habitat less affected at small scales by human actionshuman actions

Geographical Area Occupied by the Geographical Area Occupied by the SpeciesSpecies

• Pacific coast from SF Pacific coast from SF Bay to Queen Bay to Queen Charlotte IslandsCharlotte Islands

• Inland waters of Inland waters of WashingtonWashington

• Georgia Basin in Georgia Basin in CanadaCanada

Geographical Area Occupied by the Geographical Area Occupied by the Species: Data and IssuesSpecies: Data and Issues

• Extensive record of Extensive record of sightings compiled by sightings compiled by Whale Museum Whale Museum

• No sightings in Hood No sightings in Hood Canal within past 20 Canal within past 20 yearsyears

• Shallow water and Shallow water and tidestides

• In ocean, only 28 In ocean, only 28 sightings over 30 yrssightings over 30 yrs

Specific AreasSpecific Areas

• Area 1- core summer Area 1- core summer area (feeding, resting, area (feeding, resting, socializing)socializing)

• Area 2- Puget Sound Area 2- Puget Sound area (fall feeding)area (fall feeding)

• Area 3- Strait of Juan Area 3- Strait of Juan de Fuca area (mainly de Fuca area (mainly migration)migration)

Particular AreasParticular Areas

• Same as specific Same as specific areasareas

• Plus military areasPlus military areas

ExclusionsExclusions

• No exclusions based on economic impactsNo exclusions based on economic impacts

• No overlap with tribal lands (waters deeper No overlap with tribal lands (waters deeper than 20 feet)than 20 feet)

• Exclusions based on impacts to national Exclusions based on impacts to national securitysecurity

References• Minett, M. and T. Cullinan.1997. A citizen’s guide to HCPs.

National Audubon Society. Washington DC.• USFWS. 1998. Www.fws.gov/r9endspp/hcpplan.html• Kaiser, J. 1997. When a habitat is not a home. Science

276:1636-1638.• Bormann, FH. And GE Likens. 1979. Catastrophic

disturbance and the steady state in northern hardwood forests. Am. Scientist 67:660-669.

• Doncaster, CP, Clobert, J, Doligez, B, Gustafsson, L, and E. Danchin. 1997. Balanced dispersal between spatially varying local populations: an alternative to the source-sink model. Am. Nat. 150:425-445.

More References

• Levins, R. 1969. Some demographic and genetic consequences of environmental heterogeneity for environmental control. Bull. Entomol. Soc. Am. 15:237-240.

• Stacey, PB. And M. Taper. 1992. Environmental variation and the persistence of small populations. Ecol. Appl. 2:18-29.

• Pulliam, HR. 1988. Sources, sinks, and population regulation. Am. Natural. 132:652-661.

• Doak, DF and LS Mills. 1994. A useful role for theory in conservation. Ecology 75:615-626.

• Botkin, DB. And MJ. Sobel. 1975. Stability in time-varying ecosystems. Am. Nat. 109:625-646.

More References

• Yong, W., Finch, DM, Moore, FR, and JF Kelly. 1998. Stopover ecology and habitat use of migratory Wilson’s Warblers. Auk 115:829-842.

• Van Horne, B. 1983. Density as a misleading indicator of habitat quality. JWM 47:893-901.

References• Crandall, KA, Bininda-Emonds, ORP, Mace, GM, and RK Wayne. 2000. Considering

evolutionary processes in conservation biology. Trends in Ecology and Evolution. 15:290-295.

• Gardenfors, U. 2001. Classifying threatened species at national versus global levels. Trends in Ecology and Evolution. 16:511-516.

• National Research Council. 1995. Science and the endangered species act. National Academy Press. Washington, DC.

• Moritz, C. 1994. Defining “evolutionary significant units” for conservation. Trends in Ecology and Evolution. 9:373-375.

• Ryder, OA. 1986. Species conservation and systmatics: the dilemma of subspecies. Trends in Ecology and Evolution. 1:9-10.

• Waples, RA. 1991. Pacific salmon, Oncorhynchus spp., and the definition of “species” under the endangered species act. Marine Fisheries Review. 53:11-22.

• Wiley, E. 1981. Phylogenetics: the theory and practice of phylogenetic systematics. New York. John Wiley & Sons

• Wilcove, D.S, M. McMillan, and K. C. Winston. 1993. What exactly is an endangered species? An analysis of the U.S. Endangered Species List: 1985-1991. Conservation Biology 7:87-93.