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Analysis Plan for the Endangered Species Act Section 7(a)(2) Consultation on the U.S. 

Environmental Protection Agency’s Vessels General Permit and Small Vessels General Permit

Nation Marine Fisheries Service and United States Fish and Wildlife Service (Services)

2 August 2012

Overview of the Services’ Assessment Framework

Section 7(a)(2) of the Endangered Species Act of 1973, as amended, requires federal Agencies, in consultation with 

and with the assistance of the Services, to insure that any action they authorize, fund, or carry out is not likely to 

jeopardize the continued existence of endangered species, threatened species, or destroy or adversely modify critical 

habitat that has been designated for these species (16 U.S.C. 1539). During consultations on specific actions, the 

Services fulfill their obligations by using an assessment framework that begins by identifying the physical, chemical, 

or biotic components of proposed actions that are likely to have individual, interactive, or aggregate direct and 

indirect effect on the environment (we use the term “potential stressors” for these components of an action); we then 

determine whether listed species or designated critical habitat are likely to be exposed to those potential stressors; 

we then estimate how listed species or designated critical habitat are likely to respond to any exposure; then we 

conclude by estimating the risks those responses pose to the individuals, populations, and species or designated 

critical habitat that are likely to be exposed. 

Federal Agency general permits apply to activities over large geographic areas over long periods of time, with substantial uncertainty about the number, location, timing, frequency, and intensity of specific activities general permits

would authorize, fund, or carry out. Traditional section 7 consultations are small-scale, proximal and focus on the 

specific effects of a specific proposal. However, those consultations are not designed to deal with the spatial and 

temporal scales and level of uncertainty that are typical of consultations on general permits. 

Instead of trying to adapt traditional consultation approaches to general permit consultations, we have developed an 

assessment framework that specifically allows us to help Federal Agencies insure that their programs comply with 

the requirements of section 7(a)(2) of the ESA as described in the Endangered Species Consultation Handbook 

(USFWS and NMFS 1998). Our assessment framework for general permits first assesses whether the general 

permits are likely to adversely affect listed species or critical habitats by estimating exposure and response, as 

described above for traditional consultations. If listed species and critical habitats are likely to be adversely affected 

we then assess how the permit is structured to comply with section 7(a)(2). Specifically, our general permit

consultations examine the decision-making processes that are integrated into Federal Agency programs to determine 

whether they are likely to insure that specific actions the Agency authorizes, funds, or carries out through the 

general permit comply with the requirements of section 7(a)(2). That is, during general permit consultations we ask 

whether or to what degree the agency has structured its proposed permit to (1) collect the information necessary to 

know or reliably estimate the probable individual and aggregate consequences of the action to listed resources; (2) 

evaluate the information to assess how its actions have affected listed resources; and (3) use its authorities to bring 

those activities into compliance with the requirements of section 7(a)(2) of the ESA, if the information it collects 
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suggests that it is necessary. Here, the “structure of permit” or “permit structure” refers to the decision-making 

processes, applications of standards and criteria (including standards of information and treatment of uncertainty), 

feedback loops and internal audits, and controls (including permit conditions) that the agencies rely upon to fulfill 

their obligations under the ESA.

Figure 1 displays a normative model of a decision-making process that includes these program elements and 

feedback loops. The process trigger on the left-hand side of the figure (Box D1) might represent an application from 

a prospective permittee or licensee, a request for prospective funding, or a prospective proposal that would be 

undertaken by a Federal Agency. These process triggers are typically subjected to two screening processes (Box 

D2): 

1. an initial screening process that is designed to insure that proposals minimally comply with statutory, 

regulatory, or policy requirements that are applicable to requests for permits, licenses, or funding; and,

2. secondary screening processes that are designed to insure that an Agency satisfies the statutory, regulatory, 

or policy requirements or criteria that must be met before an Agency can issue a permit, license, or funding.

For example, the screening process the EPA applies to general permits includes reviews for completeness; analyses 

for compliance with Clean Water Act (CWA) guidelines; compliance with State water quality standards; compliance 

with toxic effluent standards; and compliance with the requirements of section 7(a)(2) of the ESA. 

Agency screening processes typically produce recommendations to Agency decision-makers, who have the authority 

to make final decisions on Agency Actions (Figure 1, Box D3). Following those decisions, the Action Agency, 

permittee, licensee, or funding recipient undertakes the action, including any terms or conditions the Action Agency 

has attached (Figure 1, Box O1). The action produces a set of direct, indirect, and aggregate effects on the environment and any living organisms that occur in or rely on the environment that is affected by the action (Figure 1, Box 

O2) and the condition of the environment changes in response to those effects (Figure 1, Box O3). 

Figure 1 also displays a program that contains an audit function represented by a monitoring component (Figure 1, 

Lines M1 to M4), a feedback component (Figure 1, Lines F1 to F4), and an information gathering and evaluation 

component that informs a screening process (Figure 1, Box D2.1). The monitoring component would collect 

empirical information on individual actions or a sample of individual actions to (a) identify what action actually 

occurred for comparison with the action that had been proposed and approved (implementation monitoring; Figure 

1, Line M1); (b) identify which terms and conditions, if any, were satisfied, including any mitigation measures that 

were required (implementation monitoring; Figure 1, Line M1); (c) gather empirical information on the action’s 

direct and indirect effects on the environment, including the effectiveness of any mitigation measures that had been 

required (validation and compliance monitoring; Figure 1, Line M2); (d) gather the empirical evidence to determine 

whether or to what degree the environment changed in response to those effects (Figure 1, Line M3); and (e) gather 

the empirical evidence sufficient to determine whether a proposal contributed to environmental conditions that do 

not meet program purposes and standards (compliance monitoring; Figure 1, Line M4). The feedback component 

evaluates empirical data collected by monitoring and incorporates those data into Agency decisions about prior or 

subsequent actions (Figure 1, Lines F1 through F4).

Five components are critical to our assessment of whether a general permit would be expected to insure that 

individual actions authorized, funded, or carried out by the program comply with the requirements of section 7(a)(2) 

of the ESA. 
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Figure 1. A normative model of a decision-making process that includes these program elements and feedback loops. See text for further explanation
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The first critical component is the screening process an Agency applies to specific actions authorized, funded, or 

carried out by a program. When we examine this component of an Agency’s decision-making process, questions we 

ask include: What standards apply to the screening process? How rigorous are those standards? How rigorously 

does the Action Agency apply those standards? Are proposals assumed to comply with an Agency’s statute barring 

evidence of non-compliance or vice versa? 

The second critical component is the information that forms the foundation for the Agency’s screening process 

(Figure 1, Boxes D2.1, and D2.2). When we examine this component, questions we ask include: Does the Agency

assess the individual and cumulative impacts of specific proposals? Does the Agency’s methodology consider all of 

the variables that would have to be considered to determine whether a specific proposal is likely to have adverse 

consequence for endangered or threatened species and designated critical habitat? Do assessments employ data

acquisition procedures that are likely to identify, gather, and analyze all of the information that would be relevant to 

identify the presence or absence of consequence for endangered or threatened species and designated critical 

habitat? Does the assessment process incorporate quality assurance and quality control procedures? Are those 

procedures designed to prevent Type I decision error (falsely concluding that a proposal had an adverse impact), 

Type II decision error (falsely concluding that a proposal had no adverse impact), or both?

The third critical component is an Action Agency’s decision-making process, which includes the information and 

variables that inform the Agency’s decision on whether or not to authorize, fund, or implement an action and any 

conditions or terms the Agency attaches to its decision. 

The fourth critical component is an audit function. Does the Action Agency audit the results of its actions? Are the 

monitoring and feedback loops (Figure 1, lines M1 to M4 and F1 to F4) designed to allow the Agency to (a) collect 

empirical information that allows them to insure that specific actions they authorize, fund, or carry out are 

undertaken as designed (including any terms, conditions, or mitigation measures associated with the proposal), (b) 

assess the actual effects of those actions, (c) determine whether the program is fulfilling its mandate, purposes, and 

goals with respect to the ESA.

The final critical component is the Agency’s authority to modify its prior and subsequent decisions when new 

information (particularly information provided by the audit function) reveals that particular authorizations have not 

satisfied applicable environmental mandates, standards, criteria, and program purposes (the applicable 

environmental mandates includes compliance with section 7(a)(2) of the ESA).

We organize our general permit consultations using a sequence of questions, listed below, that focus on the 

Agency’s decision-making process in general and the five critical components we just described. Those questions 

focus on whether and to what degree an Agency has structured a program so that the Agency is in a position to know 

or reliably estimate whether endangered or threatened species or designated critical habitat are likely to be: (a) 

exposed to stressors associated with specific actions a program would authorize, fund, or carry out; (b) respond to 

that exposure; and (c) experience individual-level, population-level, or species-level risks as a result of those 

responses (or similar reductions in the features and value of critical habitat). Further, we ask whether or to what 

degree an Agency actively gathers that information, whether or to what degree an Agency incorporates that 

information into its decision-making processes about specific actions, and whether or to what degree an Agency

changes the decisions it makes about specific actions based on that information.
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Application of this Approach in this Consultation

As we have already discussed, we treat the issuance of the proposed VGPs as “programs” that would authorize 

discharges of pollutants over a five-year period. As we described earlier, during general permit consultations we ask 

whether or to what degree the programs: 

Collect or will have access to the information necessary to allow it to know how the actions it permits 

affect listed resources; 

Evaluate that information to assess how its actions have affected endangered species, threatened species, 

and designated critical habitat; and 

Uses its authorities to bring those activities into compliance.

Specific questions we ask about the proposed VGPs are:

1. Scope

Has the general permit been structured to reliably estimate the probable number, location and timing of the 

discharges that would be authorized by the program?

2. Stressors

Has the general permit been structured to reliably estimate the physical, chemical, or biotic stressors that are likely 

to be produced as a direct or indirect result of the discharges that would be authorized (that is, the stressors produced 

by the actual discharges to waters of the U.S.)?

3. Overlap

Has the general permit been structured to reliably estimate whether or to what degree specific endangered or 

threatened species or designated critical habitat are likely to be exposed to potentially harmful impacts that the 

proposed permit would authorize?

4. Monitoring/Feedback

Has the general permit been structured to identify, collect, and analyze information about authorized actions that 

may have exposed endangered or threatened species or designated critical habitat to stressors at concentrations, 

intensities, durations, or frequencies that are known or suspected to produce physical, physiological, behavioral, or 

ecological responses that have potential individual or cumulative adverse consequences for individual organisms or 

constituent elements of critical habitat?

5. Responses of Listed Resources

Does the general permit have an analytical methodology that considers: 

a) the status and trends of endangered or threatened species or designated critical habitat; 

b) the demographic and ecological status of populations and individuals of those species given their exposure 

to pre-existing stressors in different drainages and watersheds; 

c) the direct and indirect pathways by which endangered or threatened species or designated critical habitat 

might be exposed to the discharges to waters of the United States; and 
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d) the physical, physiological, behavior, sociobiological, and ecological consequences of exposing endangered 

or threatened species or designated critical habitat to stressors from discharges at concentrations, 

intensities, durations, or frequencies that could produce physical, physiological, behavioral, or ecological 

responses, given their pre-existing demographic and ecological condition?

6. Compliance 

Does the general permit have a mechanism to reliably determine whether or to what degree operators have complied 

with the conditions, restrictions or mitigation measures the proposed permit requires when they discharge to waters 

of the United States?

7. Adequacy of Controls

Does the general permit have a mechanism to prevent or minimize endangered or threatened species or designated 

critical habitat from being exposed to stressors from discharges: 

a) at concentrations, durations, or frequencies that are potentially harmful to individual listed organisms, 

populations, or the species, or; 

b) to ecological consequences that are potentially harmful to individual listed organisms, populations, the 

species or Primary Constituent Elements of designated critical habitat? 

As with all of our consultations, we conduct our section 7 analyses through a series of steps. These include: a

examination of the action; identification of the action area; exposure analyses; response analyses; analyses of 

aggregate impacts; and risk analyses. These steps are described below.

PROPOSED ACTION

In reviewing the proposed action and biological evaluation, the Services will examine the activities that would be 

authorized by the proposed VGPs. This step of our analyses identifies spatial and temporal patterns associated with 

each category of activity; specifically (a) the geographic distribution of the different activities; (b) the number 

activities; (c) the amounts of pollutants that are likely to be discharged; and (d) the rate of discharges.

Our analysis will evaluate the effects of all discharges of biological and chemical pollutants to waters of the US 

incidental to the normal operation of all eligible vessels that would be authorized by the VGPs. In addition, the 

Services must consider the effects of interrelated and interdependent actions of the proposed action. Interdependent 

actions are actions having no independent utility apart from the proposed action (50 CFR 402-021). They are 

typically a consequence of the proposed action. For example, if our consultation were evaluating the effects of 

building a road, an interdependent action would be the planned construction of homes and other structures that 

would not be accessible without the presence of that road. Interrelated actions are actions that are part of a larger 

action and depend on the larger action for their justification (50 CFR 402-023). They are actions that are typically 

associated with the proposed action.

In this consultation, NMFS would address the potential effects of the proposed action for marine, coastal, estuarine, 

or anadromous endangered species, threatened species, and critical habitat that has been designated for those species 

 

1

See: http://cfr.vlex.com/vid/402-02-definitions-19895209
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in those ecosystems under its jurisdiction; FWS would address the potential effects of the proposed action for 

terrestrial, freshwater, and resident estuarine endangered species, threatened species, and critical habitat that has 

been designated for those species under FWS’ jurisdiction in those ecosystems. 

ACTION AREA

The Services will determine the degree of geographic and temporal overlap between the activities that would be 

authorized by the proposed VGPs and endangered and threatened species and designated critical habitat. The Action 

Area for this consultation consists of all waters of the United States in the United States, its territories, and its 

possessions (which includes American Samoa, Baker Island, Guam, Howland Island, Jarvis Island, Johnston Atoll, 

Midway Islands, Navassa Island, the Commonwealth of the Northern Mariana Islands, Palmyra Atoll, Puerto Rico, 

the U.S. Virgin Islands, and Wake Island) where discharges incidental to the normal operation of all nonrecreational and non-military vessels are to be authorized by the EPA’s VGPs. Waters of the U.S. (as defined in 40 

CFR 122.22) are:

1. All waters which are currently used, were used in the past, or may be susceptible to use in interstate or 

foreign commerce, including all waters which are subject to the ebb and flow of the tide;

2. All interstate waters, including interstate “wetlands;”

3. All other waters such as intrastate lakes, rivers, streams (including intermittent streams), mudflats, 

sandflats, “wetlands,” sloughs, prairie potholes, wet meadows, playa lakes, or natural ponds the use, 

degradation, or destruction of which would affect or could affect interstate or foreign commerce including 

any such waters:

a. Which are or could be used by interstate or foreign travelers for recreational or other purposes;

b. From which fish or shellfish are or could be taken and sold in interstate or foreign commerce; or

c. Which are used or could be used for industrial purposes by industries in interstate commerce;

4. All impoundments of waters otherwise defined as waters of the United States under this definition;

5. Tributaries of those waters described above;

6. The territorial sea; and

7. “Wetlands” adjacent to waters (other than waters that are themselves wetlands).

EXPOSURE ANALYSES

Before assessing an agency’s decision-making process (as described above), the Services will first establish whether 

the proposed action could expose listed species and critical habitat to potentially harmful stressors and whether 

 

2

See: http://ecfr.gpoaccess.gov/cgi/t/text/textidx?type=simple;c=ecfr;cc=ecfr;idno=40;region=DIV1;q1=122.2;rgn=div8;sid=eefc0a55c51938afa484bbfe4d9ea5a2;view=text;

node=40%3A22.0.1.1.12.1.6.2
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listed species and critical habitats are likely to respond adversely. If exposure to stressors and adverse responses are 

not likely to occur, we do not assess the agency’s decision-making process.

Our assessment will focus on whether and to what degree the EPA structured the VGPs in ways that would prevent 

or minimize endangered or threatened species or critical habitat that has been designated for those species from 

being exposed to harmful discharges of pollutants into waters of the United States and other harmful activities 

because such exposures commonly trigger responses that are difficult to prevent. For example, once individual 

plants and animals are exposed to a discharge containing pollutants, their responses to the exposure is controlled by 

the concentration, duration, and frequency associated with the exposure, their sensitivity to the discharged materials, 

other physical, chemical, or biotic stressors they are exposed to in the same time interval, their pre-existing 

physiological state, and their constitutional endowment. Because it is so difficult to prevent free-ranging organisms 

from responding to anthropogenic stressors once they have been exposed, the most effective management measures 

are designed to influence the exposure itself. For that reason, our assessment focuses on whether and to what degree 

the VGPs prevent or minimize endangered and threatened species and designated critical habitat from being exposed 

to harmful discharges and other harmful activities that would be authorized by the proposed VGPs. Rather than 

discuss the literature for each species, it would be only be necessary to discuss the risks of exposing species groups 

(for example, Pacific Salmon; Sturgeon; Sea Turtles; etc.).

RESPONSE ANALYSES

The Services will conduct a detailed review of the literature available on the physical, physiological, behavioral, 

social, and ecological responses of endangered or threatened species or constituent elements of critical habitat given 

exposure to harmful discharges into waters of the United States or to the effects of those discharges and the activities 

associated with those activities on the ecology of the watersheds in which they occur (that is, effects resulting from 

changes in populations of prey, predators, competitors, symbionts, etc.). Rather than discuss the literature for each 

species, we organize the data using species groups (for example, Pacific Salmon; Sturgeon; Sea Turtles; etc.). We 

summarize the probable consequences of the identified responses for populations of endangered and threatened 

species and designated critical habitat. 

TREATMENT OF AGGREGATE IMPACTS 

To address the question of whether the activities that would be authorized by the proposed VGPs have direct and 

indirect effects on listed resources that are sufficiently small both individually and in aggregate, we explicitly 

consider the aggregate impacts of the proposed permits in the Effects of the Action chapter of our Opinion. This 

usage of “aggregate impacts” is distinct from the term “cumulative effects” which the section 7 regulations defines 

as “those effects of future State or private activities, not involving Federal activities, that are reasonably certain to 

occur within the action area of the Federal action subject to consultation” (50 CFR 402.02). 

Aggregate impacts include: (1) time-crowded perturbations or perturbations that are so close in time that the effects 

of one perturbation do not dissipate before a subsequent perturbation occurs; (2) space-crowded perturbations or 

perturbations that are so close in space that their effects overlap; (3) interactions or perturbations that have 

qualitatively and quantitatively different consequences for the ecosystems, ecological communities, populations, or 

individuals exposed to them because of synergism (when stressors produce fundamentally different effects in 
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combination than they do individually), additivity, magnification (when a combination of stressors have effects that 

are more than additive), or antagonism (when two or more stressors have less effect in combination than they do 

individually); and (4) nibbling or incremental and decremental effects are often, but not always, involved in each of

the preceding three categories (NRC 1986).

RISK ANALYSES 

The Services help Action Agencies determine whether or to what degree they have complied with the requirements 

of section 7(a)(2) of the ESA by assessing whether and to what degree an Agency has structured a program so that 

the Agency is in a position to know or reliably estimate: (a) whether endangered or threatened species are likely to 

be placed at increased risk of extinction, or (b) if those species avoid extinction, whether they are likely to 

experience increased risk of failing to recover from having been endangered or threatened because of the actions the 

program authorizes, funds, or carries out. 

Our consideration of how well an Agency’s program manages risks to endangered and threatened species reflects 

the relationship between a listed species, the populations it comprises, and the individuals those populations 

comprise. The continued existence of a species is determined by the fate of the populations it comprises and the 

continued existence of a population is determined by the fate of the individuals they comprise. Populations grow or 

decline as the individuals they comprise live, die, grow, mature, migrate, and reproduce, or fail to do so. When we

assess whether or to what degree an Agency’s program is likely to eliminate or avoid risks to endangered or 

threatened species, we are mindful of the distinction between species, the populations they comprise, and the 

individuals comprised by those populations.

When we assess whether or to what degree an Agency’s program is likely to eliminate or avoid risks to individual 

members of endangered or threatened species, we think in terms of the individuals’ fitness, which integrates an 

individuals’ longevity with its current and future reproductive success. In particular, we examine the scientific and 

commercial data available to determine if an individual’s probable response to stressors produced by an Action 

would reasonably be expected to reduce the individual’s current or expected future reproductive success by 

increasing an individual’s likelihood of dying prematurely, increasing the age at which it becomes reproductively 

mature, reducing the age at which it stops reproducing, reducing the number of live births it produces during any 

reproductive bout, reducing the number of reproductive bouts it engages in over its reproductive lifespan (in animals 

that reproduce multiple times), or causing the individual’s progeny to experience any of these phenomena (Brommer 

2000, Brommer et al. 1998, 2002; Clutton-Brock 1998, Coulson et al. 2006, Kotiaho et al. 2005, McGraw and 

Caswell 1996, Newton and Rothery 1997, Oli and Dobson 2003, Roff 2002, Stearns 1992, Turchin 2003).

We evaluate whether individual members of an endangered or threatened species experience reductions in their 

current or expected future reproductive success or experience reductions in the rates at which they grow, mature, or 

become reproductively active. Then we determine if those reductions also reduce the abundance, reproduction rates, 

and growth rates (or increase variance in one or more of these rates) of the populations those individuals represent 

(see Stearns 1992). Similarly, we determine if those population-level reductions reduce the viability of the species 

those population(s) comprise. Our general permit assessments focus on whether or to what degree an Agency’s 

program is likely to insure that the direct or indirect effects of actions the program would authorize are not likely to 

reduce the fitness of listed individuals to a degree that would be sufficient to reduce the viability of the population(s) 

those individuals represent and jeopardize the survival and recovery of the species.
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Our consideration of how well an Agency’s program manages risks to designated critical habitat focuses on the 

value of the physical, chemical, or biotic phenomena of the critical habitat for the conservation of the endangered 

and threatened species for which the critical habitat was designated. In this step of our assessment, we consider 

information about the contribution of constituent elements of critical habitat (or of the physical, chemical, or biotic 

features that give the designated area value for the conservation of listed species, particularly for older critical 

habitat designations that have no constituent elements) to the conservation value of those areas of critical habitat that 

occur in the action area. Then we consider the contribution of the conservation value of those areas to the 

conservation value of the entire critical habitat designation. Our general permit assessments focus on whether or to 

what degree an Agency’s program is likely to insure that the direct or indirect effects of actions the program would 

authorize are not likely to reduce the value of the physical, chemical, or biotic phenomena of the critical habitat 

areas affected or are not likely to reduce that value of the physical, chemical, or biotic phenomena to a degree that 

would be sufficient to reduce the value of the entire critical habitat for the conservation of the endangered and 

threatened species for which the critical habitat was designated.

Evidence Available for the Consultation

For all of the above analyses, the Services will rely on two bodies of evidence for this consultation. We will use the 

first body of evidence to determine whether or to what degree the proposed general permits can insure that vessel 

discharges on, over, or near waters of the United States are not likely to jeopardize the continued existence of 

endangered or threatened species or result in the destruction or adverse modification of critical habitat that has been 

designated for those species. To build this body of evidence, we will search for, gather, and analyze published and 

unpublished sources that examine the effectiveness previous programs, and whether or to what degree the program 

has had: (1) adverse consequences for natural flora and fauna that have some dependency on the quality of waters 

of the United States, or (2) adverse consequences for species that have been listed as endangered or threatened. In 

particular, we will consider information contained in biological assessments, program reviews of effectiveness, 

compliance monitoring databases, risk assessments and ESA Section 7 consultations on similar actions, species 

status reviews, listing documents, recovery plans, reports on the status and trends of water quality, past and current 

research and population dynamics modeling. We will supplement this information by searching for compliance data

from available databases that track the number of inspections and enforcement actions over time. 

We will use a second body of evidence to assess the probable consequences of authorizing discharges of pollutants 

on, over, or near waters of the United States to endangered or threatened species or critical habitat that has been 

designated for those species. To assemble this body of information, we shall search peer-reviewed scientific 

literature, master’s theses and doctoral dissertations, government reports and studies and reports from commercial 

vendors. For example, the National Library of Medicine’s Hazardous Substance Data Bank, TOXNET, Toxline, 

EXTOXNET pesticide information profiles provide information on the toxicity of the pollutants. These searches 

will focus on identifying recent information on the biology, ecology, distribution, status, and trends of the threatened 

and endangered species considered in this consultation. We will consider the results of these searches based on the 

quality of their study design, sample sizes and study results. Studies that relied on large sample sizes with small 

variances will be generally ranked higher than studies that relied on small sample sizes or large variances.
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Literature Cited

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Brommer, J. E., J. Merila, and H. Kokko. 2002. Reproductive timing and individual fitness. Ecology Letters 5:802-

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Clutton-Brock, T. H., I. R. Stevenson, P. Marrow, A. D. MacColl, A. I. Houston, and J. M. McNamara. 1996. 

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Coulson, T., T. G. Benton, P. Lundberg, S. R. X. Dall, B. E. Kendall, and J. M. Gaillard. 2006. Estimating 

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Kotiaho, J.S., V. Kaitala, A. Komonen and J. Paivinen. 2005. Predicting the risk of extinction from shared 

ecological characteristics. Proceedings of the National Academy of Sciences of the United States of America 

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McGraw, J. B., and H. Caswell. 1996. Estimation of individual fitness from life-history data. The American 

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National Research Council. 1986. The special problem of cumulative effects. Pages 93-136 In: Ecological 

knowledge and environmental problem-solving. National Academy Press, Washington, D.C. 

Newton, I. and P. Rothery. 1997. Senescence and reproductive value in sparrowhawks. Ecology 78:1000-1008.

Oli, M.K. and F.S. Dobson. 2003. The relative importance of life-history variables to population growth rate in 

mammals: Cole's prediction revisited. The American Naturalist 161(3):422-440.

Roff, D.A. 2002. Life history evolution. Sinauer Associates, Inc.; Sunderland, Massachusetts.

Stearns, S. C. 1992, The evolution of life histories. New York, New York, Oxford University Press.

Turchin, P. 2003. Complex population dynamics: a theoretical/empirical synthesis. Princeton University Press; 

Princeton, New Jersey.