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 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 g laciation and succession) –fire has return rate of 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 FireWind Clear-cutting Total Biomass Time (White Mountains, NH; Bormann and Likens 1979)

80 20 Do We Really Know Habitat Needs? Spring Males Females Important % of Each Age/Sex In Group 10 Fall Adults Hatch Year 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 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 – years, 200 species

The HCP Process 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 validityhere

NCEAS evaluation of HCPs Kareiva et al. online here 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 Designation Pacific Salmon and Steelhead (2005) –19 Species or “ESUs” –13 in WA, OR, ID, 7 in CA Southern Resident Killer Whales Presentation by Donna Darm, NOAA Fisheries

Relevance of Critical Habitat Section 7(a)(2): all federal agencies shall ensure actions they fund, permit or carry out are not likely to:Section 7(a)(2): all federal agencies shall ensure actions they fund, permit or carry out are not likely to: –jeopardize species’ continued existence –destroy or adversely modify critical habitat

What is critical habitat? Occupied areas with physical or biological featuresOccupied areas with physical or biological features –essential to conservation –may require special management Unoccupied areas essential for conservation ESA §3(5)Unoccupied areas essential for conservation ESA §3(5)

Step 1: Identify “Critical Habitat” Map actual fish distributionMap actual fish distribution Verify features and special managementVerify features and special management Group by watershed into “specific areas”Group by watershed into “specific areas” Identify unoccupied areas essential for conservationIdentify unoccupied areas essential for conservation

How does it get designated? Best scientific dataBest scientific data Consider economic, national security, other relevant impactsConsider economic, national security, other relevant impacts May exclude if benefits of exclusion outweigh benefits of designationMay exclude if benefits of exclusion outweigh benefits of designation (Unless exclusion results in extinction)(Unless exclusion results in extinction) ESA §4(b)(2) ESA §4(b)(2) Benefits of Designation Benefits of Exclusion Conservation Avoid Impacts

Step 2: Consider Impacts, Balance Benefits Benefit of designation is the protection of section 7 (no adverse modification by federal agencies)Benefit of designation is the protection of section 7 (no adverse modification by federal agencies) Designation also gives notice of areas important to speciesDesignation also gives notice of areas important to species Benefits of exclusion (i.e., avoiding impacts) may be economic, national security, or “other”Benefits of exclusion (i.e., avoiding impacts) may be economic, national security, or “other”

Balancing Benefits (Economic) Consider for exclusion areas with a relatively high economic impact and a relatively low conservation valueConsider for exclusion areas with a relatively high economic impact and a relatively low conservation value Made sense with salmon because:Made sense with salmon because: –Large number of habitat areas –Not all areas are equally important for conservation Conservation threshold – don’t exclude if it significantly impedes conservationConservation threshold – don’t exclude if it significantly impedes conservation

Relative Conservation Value of Habitat Areas

Example Exclusion Scenario

Balancing Benefits (Other) National Security (military readiness during global war on terror)National Security (military readiness during global war on terror) Conservation partnerships on private land (HCPs) = net conservation benefitConservation partnerships on private land (HCPs) = net conservation benefit Tribal sovereignty and self-determination (conservation partnership also relevant)Tribal sovereignty and self-determination (conservation partnership also relevant)

Net Occupied Habitat Areas = 23,447 miles - Dept. of Defense Sites - Indian Lands - 4 HCP Holders - Economic Impacts Net Designated Habitat Areas = 20,630 miles August Final Critical Habitat for 12 ESUs of Salmon and Steelhead

Southern Resident Killer Whales Very different species, very different habitatVery different species, very different habitat Marine versus fresh waterMarine versus fresh water Habitat less affected at small scales by human actionsHabitat less affected at small scales by human actions

Geographical Area Occupied by the Species Pacific coast from SF Bay to Queen Charlotte IslandsPacific coast from SF Bay to Queen Charlotte Islands Inland waters of WashingtonInland waters of Washington Georgia Basin in CanadaGeorgia Basin in Canada

Geographical Area Occupied by the Species: Data and Issues Extensive record of sightings compiled by Whale MuseumExtensive record of sightings compiled by Whale Museum No sightings in Hood Canal within past 20 yearsNo sightings in Hood Canal within past 20 years Shallow water and tidesShallow water and tides In ocean, only 28 sightings over 30 yrsIn ocean, only 28 sightings over 30 yrs

Specific Areas Area 1- core summer area (feeding, resting, socializing)Area 1- core summer area (feeding, resting, socializing) Area 2- Puget Sound area (fall feeding)Area 2- Puget Sound area (fall feeding) Area 3- Strait of Juan de Fuca area (mainly migration)Area 3- Strait of Juan de Fuca area (mainly migration)

Particular Areas Same as specific areasSame as specific areas Plus military areasPlus military areas

Exclusions No exclusions based on economic impactsNo exclusions based on economic impacts No overlap with tribal lands (waters deeper than 20 feet)No overlap with tribal lands (waters deeper than 20 feet) Exclusions based on impacts to national securityExclusions based on impacts to national security

References Minett, M. and T. Cullinan A citizen’s guide to HCPs. National Audubon Society. Washington DC. USFWS Kaiser, J When a habitat is not a home. Science 276: Bormann, FH. And GE Likens Catastrophic disturbance and the steady state in northern hardwood forests. Am. Scientist 67: Doncaster, CP, Clobert, J, Doligez, B, Gustafsson, L, and E. Danchin Balanced dispersal between spatially varying local populations: an alternative to the source-sink model. Am. Nat. 150:

More References Levins, R Some demographic and genetic consequences of environmental heterogeneity for environmental control. Bull. Entomol. Soc. Am. 15: Stacey, PB. And M. Taper Environmental variation and the persistence of small populations. Ecol. Appl. 2: Pulliam, HR Sources, sinks, and population regulation. Am. Natural. 132: Doak, DF and LS Mills A useful role for theory in conservation. Ecology 75: Botkin, DB. And MJ. Sobel Stability in time- varying ecosystems. Am. Nat. 109:

More References Yong, W., Finch, DM, Moore, FR, and JF Kelly Stopover ecology and habitat use of migratory Wilson’s Warblers. Auk 115: Van Horne, B Density as a misleading indicator of habitat quality. JWM 47:

References Crandall, KA, Bininda-Emonds, ORP, Mace, GM, and RK Wayne Considering evolutionary processes in conservation biology. Trends in Ecology and Evolution. 15: Gardenfors, U Classifying threatened species at national versus global levels. Trends in Ecology and Evolution. 16: National Research Council Science and the endangered species act. National Academy Press. Washington, DC. Moritz, C Defining “evolutionary significant units” for conservation. Trends in Ecology and Evolution. 9: Ryder, OA Species conservation and systmatics: the dilemma of subspecies. Trends in Ecology and Evolution. 1:9-10. Waples, RA Pacific salmon, Oncorhynchus spp., and the definition of “species” under the endangered species act. Marine Fisheries Review. 53: Wiley, E Phylogenetics: the theory and practice of phylogenetic systematics. New York. John Wiley & Sons Wilcove, D.S, M. McMillan, and K. C. Winston What exactly is an endangered species? An analysis of the U.S. Endangered Species List: Conservation Biology 7:87-93.