Use of drift nets to monitor production and limiting factors in recruitment of larval Pacific lamprey proposal 200706300

FY07-09 proposal 200706300

Section 1. Administrative

Proposal title	Use of drift nets to monitor production and limiting factors in recruitment of larval Pacific lamprey
Proposal ID	200706300
Organization	Oregon State University
Short description	The primary purpose of this proposal is to assess a larval drift protocol for general application throughout the CRB, describe local spawning stock - larval recruitment relationships, and quantify factors limiting early recruitment of Pacific lamprey.
Information transfer	In addition to inclusion in annual reports, information gained from this proposal will be presented annually at Oregon Chapter of AFS and other meetings, submitted to peer reviewed journals, and made available to the Columbia River Basin Lamprey Technical Workgroup for circulation to fisheries managers.
Proposal contact person or principal investigator

Contacts

Contact	Organization	Email
Form submitter
Abel Brumo	Oregon State University	abel.brumo@oregonstate.edu
All assigned contacts
Abel Brumo	Oregon State University	abel.brumo@oregonstate.edu
Abel Brumo	Oregon State University	abel.brumo@oregonstate.edu
Doug Markle	Oregon State University	douglas.markle@oregonstate.edu

Section 2. Locations

Province / subbasin: Mainstem/Systemwide / Systemwide

Latitude	Longitude	Waterbody	Description
		Blue River	Likely location for Objective 1. Potential location for Objectives 2 & 3.
		Deer Creek	Likely location for Objective 1. Potential location for Objectives 2 & 3.
		Horse Creek	Likely location for Objective 1. Potential location for Objectives 2 & 3.
		Lost Creek	Likely location for Objective 1. Potential location for Objectives 2 & 3.
		North Fork Gate Creek	Likely location for Objective 1. Potential location for Objectives 2 & 3.
		Quartz Creek	Likely location for Objective 1. Potential location for Objectives 2 & 3.
		South Fork Gate Creek	Likely location for Objective 1. Potential location for Objectives 2 & 3.
		South Fork McKenzie River	Likely location for Objective 1.
		South Fork McKenzie River	Likely location for Objectives 2 and 3. Directly below Cougar Reservoir. ODFW personnel have observed a spawning population of Pacific lamprey at this site. Easy access exists.

Section 3. Focal species

primary: Pacific Lamprey
secondary: Western Brook Lamprey

Section 4. Past accomplishments

Year	Accomplishments

Section 5. Relationships to other projects

Funding source	Related ID	Related title	Relationship
Other: BPA-07-09	200716500	Relative abundance, distribution, and population structure of lampreys in the Columbia River Basin	This project refers to a 2007-2009 BPA proposal being submitted by USGS and CRITFIC. We participated in the initial planning meetings, reviewed a draft proposal, discussed our project plans with the principal investigators, and are aware of potential collaboration possibilities. For example, a key objective of proposal 200716500 is to develop standardized protocols for sampling lampreys in all life stages and implement them throughout the entire CRB. The information on sampling adult and emergent larvae gained from our project will compliment the above project—aiding in interpretation of their results throughout the basin. We will encourage the principal investigators to use the McKenzie basin as a focal basin for their project, assist with logistical and field support to the maximum extent possible, and potentially take the lead on standardized early ammocoete surveys. Our presence/absence of older ammocoete surveys (Work Element 1.7), will be coordinated for consistency with this project. Lastly, since both projects propose publication of sampling manuals for respective life stages, cooperative effort in publishing a multi-life stage manual is probable.
BPA	200201600	Determine lamprey species composition, larval distribution, and adult abundance in the Deschutes River subbasin	We have met with project leaders to discuss potential collaboration and use of our larval drift methodologies in the Deschutes River. The Deschutes River may be a good candidate for application of our larval sampling protocol if logistic and analytical problems are addressed. Because of this collaboration, data from our project will have a direct bearing on interpretation of Deschutes lamprey monitoring data. Researchers working on this project have expertise in ammocoete presence/absence sampling and will be consulted for assistance with our sampling designs. Continued coordination of efforts between the two projects will occur.
BPA	200002800	Evaluate the status of Pacific lamprey in the Clearwater River drainage, Idaho	This project employs ammocoete survey protocols that could be implemented in Work Element 1.7. Project leaders will be consulted with on sampling methods and informed of our results. This project presents evidence for lack of larval recruitment in the S.F. Clearwater River subbasin (Cochnauer and Claire 2003). Data from our proposed project is vital for understanding this trend.

Section 6. Biological objectives

Biological objectives	Full description	Associated subbasin plan	Strategy
Limitations to larval recruitment	Describe and quantify the chief factors limiting Pacific lamprey larval recruitment in focal spawning areas.	Willamette	"Strategy 2: Monitor habitat capacity, focal species survival and productivity, and biotic processes in selected watersheds within the Willamette Basin."
Spawning stock - larval recruitment	Investigate the relationship between Pacific lamprey spawning stock and recruitment to larval phase.	Willamette	"...species recovery efforts must be placed within the context of complete life cycles." (Conservation Guidelines)
Use of larval drift nets to evaluate status	Establish and assess a monitoring protocol that employs larval drift sampling to gauge Pacific lamprey distribution, status, and life history in large river basins where little information exists.	Willamette	"Strategy 7: Standardize monitoring designs, assessment protocols, and methods to manage, analyze data, and report findings."

Section 7. Work elements (coming back to this)

Work element name	Work element title	Description	Start date	End date	Est budget
Analyze/Interpret Data	Describe and/or quantify and model key variables explaining Pacific lamprey larval recruitment.	Multiple linear or quantile regression techniques will be employed to analyze and model potential factors influencing larval recruitment. A statistician at Oregon State University will be consulted both before and after sampling to aid in selecting robust models.	8/1/2008	12/1/2009	$10,000
Biological objectives Limitations to larval recruitment			Metrics
Analyze/Interpret Data	Evaluate relationships between spawning adults, redds, and emergent larval Pacific lamprey in focal spawning areas.	Employing the temperature-development relationship in Work Element 1.2, the date offspring from each adult observation (Work Element 2.2) are expected emerge into the drift will be predicted. To investigate the relationship between stock and larval recruitment, each spawning observation will be linked with the larval drift rate observed at the expected emergence date. Various stock-recruitment models will be used to describe observed correlations at each focal site. Analyses inspecting differences between sites and among sites within and between years will be carried out. Although stock-recruitment is typically evaluated on an annual basis, the greater number of data points obtained by using weekly cohorts as described here is advantageous when long term data sets are not available.	8/1/2007	12/1/2009	$10,000
Biological objectives Spawning stock - larval recruitment			Metrics
Collect/Generate/Validate Field and Lab Data	Apply standardized ammocoete presence/absence survey for comparison with results from larval drift nets.	Using a standardized sampling protocol likely developed by the Lamprey Technical Workgroup, older ammocoete surveys will be carried out in tributaries sampled in Work Element 1.1. This protocol will likely involve use of an AbP-2 backpack electrofisher (Engineering Technical Services, University of Wisconsin, Madison, Wisconsin). The purpose of this task is to compare presence/absence of young of the year (drift nets) with presence/absence of older ammocoetes. These surveys will take place after spawning season in the fall 2007, when likelihood of detecting ammocoetes is predicted to be maximized.	8/1/2007	12/1/2008	$25,000
Biological objectives Use of larval drift nets to evaluate status			Metrics
Collect/Generate/Validate Field and Lab Data	Determine seasonal timing of Pacific lamprey spawning in eight McKenzie Basin tributaries above Leaburg Dam.	Dates and duration of spawning in each tributary will be calculated using drift net samples described above in conjunction with a temperature-development relationship and in situ water temperature data. The temperature-development equation was generated using a power function fit to USGS laboratory data on development time of Pacific lamprey reared at 10, 14, 18, and 22ºC (Brumo and Markle, 2005, unpublished; Meeuwig et al. 2004). This relationship was verified in the field in 2005 using in situ fertilization and incubation of Pacific lamprey eggs (Brumo and Markle, 2005, unpublished data). Onset Optic Stowaway Temp Logger® will be placed at each net site from January until August in 2007, taking readings every 30 minutes. Mean water temperature during the 30 days prior to larval emergence will be used in the power equation to predict spawning date for each group of larvae, resulting in continuous picture of spawning activity and larval production over time. This technique assumes age-0 larvae are newly emerged. Our work on the Coquille River showed older and larger age-0 larvae were caught with increasing frequency as the spawning season progressed. In order to address this problem, newly-emerged Pacific lamprey ammocoetes were defined as 8-9mm based on length frequency of early season samples and data from USGS laboratory studies (Brumo and Markle, 2005 unpublished data; M. Meeuwig, personal communication). In the case river system-specific size differences exist, the size range used to distinguish emergent larvae from older individuals will be adjusted based on length frequencies of early season samples from each tributary. Pacific lamprey will be distinguished from western brook using the methods described in Work element 1.6.	1/10/2007	8/1/2009	$25,000
Biological objectives Use of larval drift nets to evaluate status			Metrics
Collect/Generate/Validate Field and Lab Data	Estimate relative drift rates of newly emerged ammocoetes from each focal spawning area throughout spawning seasons.	Using methods described in Work Element 1.1 age-0 ammocoetes will be collected from roughly 30 meters downstream of the lower boundary of each focal spawning area. Beginning about 20 days after initial spawning activity, three drift nets will be set twice a week at each site for 10-20 minutes, 2-4 hours after sunset. Emergent Pacific lamprey larval drift rates for each sample will be calculated following Work Element 1.3. Sampling will continue until emergent larvae are not present in the drift. This task will occur in 2008 and 2009 spawning seasons.	3/1/2008	8/1/2009	$40,000
Biological objectives Spawning stock - larval recruitment			Metrics
Collect/Generate/Validate Field and Lab Data	Evaluate temporal trends in age-0 ammocoete production on eight McKenzie Basin tributaries.	Using data and analyses from Work Elements 1.1, 1.2 and 1.3, ammocoete drift rates for study tributaries will be assessed within and between spawning seasons. In spring 2007, all eight study tributaries will be evaluated to determine if most larval production results from early or later spawners. Tributaries containing the three focal populations chosen for Objectives 2 and 3 will be monitored in 2008 and 2009 to investigate annual differences in tributary larval production and within-season drift patterns.	2/1/2007	8/1/2009	$25,000
Biological objectives Use of larval drift nets to evaluate status			Metrics
Collect/Generate/Validate Field and Lab Data	Identify lamprey larvae to species using size, temporal differences in spawning, anatomy, and molecular techniques.	Although reliable techniques for distinguishing older Pacific lamprey larvae from western brook lamprey do not currently exist (Meeuwig et al. 2004), it is likely that they will by the time this project begins (D. Goodman, personal communication). Because L. richardsoni may be present in our study sites, a series of strategies will be employed to minimize species misidentification. Pacific lamprey have larger eggs and emergent larvae than brook lamprey. Based on USGS laboratory data and our previous research, all larvae under 7.2 mm will be considered L. richardsoni (Brumo and Markle, 2005, unpublished data; M. Meeuwig, personal communication). Because L. richardsoni typically spawn later than L. tridentata, early-season emergent larvae will be primarily Pacific lamprey. In previous work, we found a pulse of smaller larvae later in the season when spawning L. richardsoni were present. Larval pigmentation differences also have potential (Meeuwig et al. 2004) and we have noticed some easily observed anatomical differences in internal organs (unpublished observations). These differences do not exist after metamorphosis so corroboration will be established using previously designed diagnostic mitochondrial loci (Docker et al. 1999). Specimens for molecular analyses will be collected once a week from drift nets and preserved in 95% ethanol.	8/1/2007	12/1/2009	$20,000
Biological objectives Use of larval drift nets to evaluate status			Metrics
Collect/Generate/Validate Field and Lab Data	Measure relative larval production of Pacific lamprey in eight McKenzie Basin tributaries above Leaburg Dam.	Comparisons of relative lamprey larval production from each study tributary will be made in 2007 using data from drift net collections described in Work Element 1.1. To control for differences in stream discharge between samples and streams, all comparisons will be based on a standardized drift rate (individuals per minute). Drift rate will be calculated by multiplying mean drift density (individuals per m3 H2O filtered) by estimated discharge (m3 H2O per minute) at each tributary. Data from all three nets (described in Work Element 1.1) will be combined. Discharge will be estimated using a hand held flow meter. To control for diel differences in drift rate, the order tributaries are sampled will be alternated each night. An unknown variable is the distance 8-9mm larvae will drift from spawning redds. White and Harvey (2003) found ammocoetes drifted at similar rates longitudinally within two northern California basins. For small tributaries such as these, we assume drift rates at sample sites will be representative of the stream.	2/1/2007	8/1/2009	$23,376
Biological objectives Use of larval drift nets to evaluate status			Metrics
Collect/Generate/Validate Field and Lab Data	Monitor Pacific lamprey redds in three focal spawning areas throughout spawning seasons.	All Pacific lamprey redds in each focal area will be counted on a weekly basis (Brumo and Markle 2004). During each count, observers will systematically wade through the focal area, counting and marking each redd. Only redds estimated to be built within the previous week, based on redd integrity and color will be counted. In 2008 and 2009, consistent methods will be used for redd counts from March through August or until spawning ceases.	3/1/2008	8/1/2009	$30,000
Biological objectives Spawning stock - larval recruitment			Metrics
Collect/Generate/Validate Field and Lab Data	Observe and document abiotic variables affecting larval recruitment in focal spawning areas.	Water temperature, discharge, water chemistry, and other relevant abiotic variables will be monitored in terms of factor limiting factor analysis.	1/10/2008	8/1/2009	$30,000
Biological objectives Limitations to larval recruitment			Metrics
Collect/Generate/Validate Field and Lab Data	Observe and document biotic variables affecting larval recruitment in focal spawning areas.	During intensive monitoring of the three focal areas used in Objective 2, relevant biotic variables influencing larval recruitment at each site will be observed and recorded. A combination of systematic wading, snorkeling surveys, and video observations will be carried out at least twice a week throughout 2008 and 2009 spawning seasons. In 2007, quantitative and or qualitative observation protocols will be developed. Based on previous research, we foresee egg predation to be the primary biotic factor, but will also describe the influence of other factors such as adult predation and redd superimposition (Brumo and Markle 2004; Pletcher 1963).	3/1/2008	8/1/2009	$30,000
Biological objectives Limitations to larval recruitment			Metrics
Collect/Generate/Validate Field and Lab Data	Quantify production of emergent larval Pacific lamprey as a surrogate for presence/absence of spawners in eight McKenzie River Basin tributaries above Leaburg Dam.	Presence and absence of spawning lamprey will be evaluated during the spring spawning season using nighttime drift samples taken in eight wadeable McKenzie Basin tributaries above Leaburg Dam. Focal tributaries are N. Fork Gate Cr., S. Fork Gate Cr., Quartz Cr., Blue River, S. Fork McKenzie, Horse Creek, Lost Creek, and Deer Creek (near Trail Bridge Reservoir). Nets will be set at locations with easy access and proximity to the mouth of each tributary. Samples will be collected between 2-4 hrs after civil twilight with 500µm mesh zooplankton-drift nets [30 x 45 cm opening] equipped with a flow meter. Diel drift samples from previous research demonstrated peak larval drift occurs during this period (Brumo and Markle 2004; Brumo and Markle, 2005, unpublished data; White and Harvey 2003). Three nets will be set concurrently at even intervals across the wetted-width to describe spatial differences in drift density. In 2007 drift nets will be set from March to August, or until larvae are not longer caught, a period predicted to encompass the entire period of age-0 lamprey redd emergence. Depending on flow conditions, nets will be fished 10-20 minutes for each tributary. Each of the eight streams will be sampled twice per week and four streams will be fished per sampling night, resulting in four sampling nights and 16 sampling events per week. A wash-down pump will be used to rinse fish larvae into net cod-end and contents will be stained with Rose Bengal dye to facilitate sorting. Samples will be preserved in 5% formalin for subsequent sorting, counting, identification, and measuring. All non-lamprey juvenile and adult fish will be identified to species, enumerated, and released unharmed. Previous work with similar sized drift nets resulted in minimal non-larval by-catch. All age-one and older ammocoetes will be separated from samples on site, measured, identified, and released unharmed. Age-0 ammocoetes and other larval fish will be sorted and tallied in the laboratory using a microscope. As described in Work Element 1.6 (Project Narrative), all, or a random subsample of, age-0 ammocoetes will be identified to species to distinguish western brook lamprey (L. richardsoni). A random subsample of 100 age-0 ammocoetes will be measured to the nearest 0.1mm using an ocular micrometer. The relative densities of emergent Pacific lamprey will be a surrogate for evidence of spawning. Absence of emergents might indicate a poor year class in a stream that otherwise has spawners or be evidence that the stream is unsuited for lamprey spawning. To detect an interannual effect and illustrate pros and cons of larval drift nets, an alternative presence/absence methodology for older ammocoetes will be used (Task 1.7).	2/1/2007	8/1/2007	$25,000
Biological objectives Use of larval drift nets to evaluate status			Metrics
Collect/Generate/Validate Field and Lab Data	Survey adult Pacific lamprey in three focal spawning areas throughout spawning seasons.	Beginning in March, counts of sexually mature Pacific lamprey and western brook lamprey will be carried out at least twice each week and be consistently made in late afternoon. Adult surveys will be based on visual counts from systematic wading through each focal area (Brumo and Markle 2004). The number of adult Pacific lamprey will serve as a relative spawning activity index to be used in subsequent analysis of stock-recruitment and larval survival. Adult monitoring will take place in 2008 and 2009 from March through August or until spawning ceases.	3/1/2008	8/1/2009	$30,000
Biological objectives Spawning stock - larval recruitment			Metrics
Develop RM&E Methods and Designs	Identify and delineate at least three significant and discrete Pacific lamprey spawning areas.	Sites for intensive monitoring of Pacific lamprey spawning populations will be identified using 2007 results from Objective 1, plus periodic snorkeling and pontoon boat surveys (Brumo and Markle 2004). Sites will have single, discrete spawning areas that can be easily monitored for Pacific lamprey adults, redds, and emergent larvae. Ideally, sites will have easy access, be wadeable and clear, and have different densities of spawners. If possible, sites will have minimal upstream influence by being located below a barrier to anadromy, being located a large distance downstream from other spawning areas, or being located below a series of significant rearing habits where drifting larvae settle out. One such site has been identified immediately below Cougar Dam on the S. Fork McKenzie River. Once other sites are located, the spawning area will be demarcated and mapped on a grid to assist with collecting data on adults, redds, and predators.	2/1/2007	2/1/2008	$10,000
Biological objectives Spawning stock - larval recruitment			Metrics
Disseminate Raw/Summary Data and Results	Communicate objective results of at Oregon AFS meetings, McKenzie Watershed Council meetings, and in peer reviewed journals.	Results will be presented at annual Oregon AFS meetings, McKenzie Watershed Council meetings, and in relevant peer reviewed journals such as Transactions of the American Fisheries Society.	2/1/2008	3/1/2010	$5,000
Biological objectives Spawning stock - larval recruitment			Metrics
Disseminate Raw/Summary Data and Results	Communicate results of Objective 3 at Oregon AFS meetings, McKenzie Watershed Council Meetings, and in peer reviewed journals.	Results will be presented at annual Oregon AFS meetings, McKenzie Watershed Council meetings, and in relevant peer reviewed journals such as Transactions of the American Fisheries Society.	2/1/2008	3/1/2010	$5,000
Biological objectives Limitations to larval recruitment			Metrics
Disseminate Raw/Summary Data and Results	Publish and circulate a manual on field, laboratory, and analytical techniques for monitoring emergent larval lamprey.	Work Elements 1.1-1.4 will serve as a proof-of-concept for larval drift net methodologies. A field manual, similar in concept to “Standard techniques for pelagic fish egg and larva surveys” (Smith and Richardson 1977) will be completed in 2008. Key elements are field operations, laboratory procedures, data summarization, statistics, and theoretical considerations associated with detection of spawning. The first draft of the manual will be made available online and, with collaborators, potentially expanded to include all life stages.	10/1/2007	12/1/2008	$30,000
Biological objectives Use of larval drift nets to evaluate status			Metrics

Section 8. Budgets

Itemized estimated budget

Item	Note	FY07	FY08	FY09
Personnel	Faculty Research Assistant	$30,000	$31,000	$32,000
Personnel	Field Technician	$12,000	$12,000	$12,000
Personnel	Undergraduate Lab Workers	$2,000	$3,000	$3,000
Fringe Benefits	For Faculty Research Assistant	$19,500	$20,150	$20,800
Fringe Benefits	For Field Technician	$8,760	$8,760	$8,760
Fringe Benefits	For Undergraduate Lab Workers	$160	$240	$240
Supplies	Laboratory and Project Supplies	$2,000	$2,000	$2,000
Supplies	Molecular Genetics Supplies	$4,000	$2,000	$2,000
Supplies	Field supplies (nets, pontoon boat, waders, boots, headlamps, washdown pump, temperature loggers, dry ice, ice)	$2,500	$1,500	$1,500
Other	Publication Charges	$0	$1,000	$1,000
Travel	Vehicle, Housing, and Per Diem	$5,500	$5,500	$5,500
Travel	Oregon AFS Meeting	$0	$750	$750
Overhead	OSU Indirect Costs @41.5%	$35,864	$36,479	$37,163
Totals		$122,284	$124,379	$126,713

Total estimated FY 2007-2009 budgets

Total itemized budget:	$373,376
Total work element budget:	$373,376

Cost sharing

Funding source/org	Item or service provided	FY 07 est value ($)	FY 08 est value ($)	FY 09 est value ($)	Cash or in-kind?	Status
Oregon State University	Markle, 0.1 FTE	$14,449	$15,170	$15,928	In-Kind	Confirmed
Totals		$14,449	$15,170	$15,928

Section 9. Project future

FY 2010 estimated budget: $0
FY 2011 estimated budget: $0

Comments:

Future O&M costs:

Termination date: 3/10/2010
Comments: Project fieldwork will be completed by September 2009. Final laboratory work, data analysis, and publication will occur after that time.

Final deliverables: 1. Annual and Final reports to BPA 2. Larval drift sampling protocol manual 3. Publication of results in peer-reviewed journals 4. Findings made available to CRB researchers through Columbia River Lamprey Technical Workgroup

Section 10. Narrative and other documents

Reviews and recommendations

FY07 budget	FY08 budget	FY09 budget	Total budget	Type	Category	Recommendation
NPCC FINAL FUNDING RECOMMENDATIONS (Oct 23, 2006) [full Council recs]
$0	$0	$0	$0	Expense	Basinwide	Do Not Fund
NPCC DRAFT FUNDING RECOMMENDATIONS (Sep 15, 2006) [full Council recs]
$0	$0	$0	$0		Basinwide

ISRP PRELIMINARY REVIEW (Jun 2, 2006)

Recommendation: Fundable in part

NPCC comments: Fundable in part – Objective # 1 only. This proposal addresses the problem of accurately assessing the abundance of early life stage lamprey larvae and developing spawner-larval recruitment relationships. The sponsors objectively discuss the advantages and disadvantages of sampling techniques including their own and conclude that the technique they propose would be better for sampling early life stages of larvae. Data on lamprey abundance are identified as a priority in the Willamette subbasin plan. The proponents have done substantial networking with other lamprey researchers in the Columbia River Basin. The latter are not using drift methodology, so the proponents could have a unique methodology that could be integrated with studies elsewhere. The proponents have laid out a measured and defensible plan to assess the drift methodology with a proof of concept approach. Comments on Objective 1: “Establish and assess a monitoring protocol that employs larval drift sampling to gauge Pacific lamprey distribution, status, and life history in large river basins where little information exists.” The proponents should assess sampling efficiency of the drift nets. As well, cross channel differences in lampreys could affect abundance estimates depending on where the three nets were deployed. It is likely that habitat impacts vary between the eight subbasins to be investigated. And it would be useful to tie this work into habitat planning as much as possible as data on flows, substrates etc could be used by others studying the Willamette basin. This would enhance the benefits of the study. Comments on “not fundable” objective 2: “Investigate the relationship between Pacific lamprey spawning stock and recruitment to larval phase.” The design for objective 2 does not mimic the natural situation that will occur in most rivers. Downstream drifting emergents may come from several spawning areas upstream, mortality would occur as they drift downstream, and some would have settled out before reaching the sampling. It would be virtually impossible to predict the number of spawners that produced the larvae captured in the drift nets. The investigators do not discuss how corrections will be made for net efficiency. Nor do they discuss the kinds of stock-recruitment models that would be appropriate. To estimate abundance of emergents it would seem simpler to just cap nests and determine the number of eggs that survive to emergence. The types of analyses that will be conducted are not given. Comments on “not fundable” objective 3: “Describe and quantify the chief factors limiting Pacific lamprey larval recruitment in focal spawning areas.” The sponsors do not explain how mortality from egg predation will be quantified. How will the affect of abiotic variables be analyzed taking into account differences in fecundity, which will not be measured? Adult predation and redd superimposition were other factors mentioned but no details on how they would be assessed were given.

ISRP FINAL REVIEW (Aug 31, 2006)

Recommendation: Fundable in part