Nutrient loading by anadromous alewife (Alosa
pseudoharengus): contemporary patterns and
predictions for restoration efforts
Derek C. West, Annika W. Walters, Stephen Gephard, and David M. Post
Can. J. Fish. Aquat. Sci. Downloaded from www.nrcresearchpress.com by CLEMSON UNIVERSITY on 11/10/14

Abstract: Anadromous alewives (Alosa pseudoharengus) have the potential to alter the nutrient budgets of coastal lakes
as they migrate into freshwater as adults and to sea as juveniles. Alewife runs are generally a source of nutrients to the
freshwater lakes in which they spawn, but juveniles may export more nutrients than adults import in newly restored popu-
lations. A healthy run of alewives in Connecticut imports substantial quantities of phosphorus; mortality of alewives con-
tributes 0.68 g Pfish–1, while surviving fish add 0.18 g P, 67% of which is excretion. Currently, alewives contribute 23%
of the annual phosphorus load to Bride Lake, but this input was much greater historically, with larger runs of bigger fish
contributing 2.5 times more phosphorus in the 1960s. A mesocosm experiment in a nearby lake showed that juvenile ale-
wife growth is strongly density dependent, but early survival may be too low for juvenile outmigration to balance adult in-
puts. In eutrophic systems where nutrients are a concern, managers can limit nutrient loading by capping adult returns at a
level where juvenile populations would not be suppressed.
Résumé : Les gaspareaux (Alosa pseudoharengus) anadromes peuvent potentiellement modifier les bilans de nutriments
des lacs côtiers pendant leurs migrations adultes vers les eaux douces et leurs migrations juvéniles vers la mer. Les mon-
taisons de gaspareaux sont généralement des sources de nutriments pour les lacs d’eau douce dans lesquels ils fraient,
mais, dans les populations nouvellement rétablies, les jeunes peuvent exporter plus de nutriments que les adultes n’en im-
For personal use only.

portent. Une montaison saine au Connecticut importe des quantités importantes de phosphore; la mortalité des gaspareaux
fournit 0,68 g Ppoisson–1, alors que les poissons survivants apportent 0,18 g P, dont 67 % est de l’excrétion. Actuellement,
les gaspareaux fournissent 23 % de la charge annuelle de phosphore du lac Bride, mais cet apport était beaucoup plus élevé
dans le passé lorsque, durant les années 1960, des montaisons plus importantes de poissons plus grands fournissaient 2,5
fois plus de phosphore. Une expérience en mésocosme dans un lac adjacent montre que la croissance des gaspareaux est
fortement reliée à la densité, mais que la survie au début du cycle est peut-être trop faible pour que l’émigration des jeunes
contrebalance les apports des adultes. Dans les systèmes eutrophes dans lesquels les gestionnaires se préoccupent des nutri-
ments, ceux-ci peuvent contrôler la charge de nutriments en limitant les retours d’adultes au niveau nécessaire pour que
les populations de jeunes ne soient pas réduites.
[Traduit par la Rédaction]

Introduction are essential components of both the freshwater and marine
systems in which they reside; however, many diadromous
In coastal watersheds across New England, diadromous fish species are in decline (Limburg and Waldman 2009).
fish are regaining access to inland waters from which they While species such as Atlantic salmon (Salmo salar) and
have been barred for a century or more. The removal of American shad (Alosa sapidissima) have garnered the pre-
dams and the installation of fishways have already reopened ponderance of public attention, ecological interactions may
250 miles of stream to diadromous fish in the state of Con- be most strongly influenced by less glamorous, but histori-
necticut, USA alone. Restoration of the spatial connection cally more abundant fish such as alewife (Alosa pseudohar-
between the ocean and freshwater ecosystems is a priority engus), and blueback herring (Alosa aestivalis) (collectively
for diadromous fish management and conservation (Atlantic called river herring).
States Marine Fisheries Commission 2009). Diadromous fish Alewives are an important component of the marine and

Received 5 October 2009. Accepted 15 May 2010. Published on the NRC Research Press Web site at cjfas.nrc.ca on 19 July 2010.
Paper handled by Associate Editor Yves Prairie.
D.C. West,1 A.W. Walters,2 and D.M. Post. Department of Ecology and Evolutionary Biology, Yale University, 165 Prospect Street,
New Haven, CT 06511, USA.
S. Gephard. Connecticut Department Environmental Protection, Inland Fisheries Division, P.O. Box 719, Old Lyme, CT 06371, USA.
1Corresponding author (e-mail: derek.west@yale.edu).
2Presentaddress: National Marine Fisheries Service, Northwest Fisheries Science Center, 2725 Montlake Boulevard East, Seattle, WA
98112, USA.

Can. J. Fish. Aquat. Sci. 67: 1211–1220 (2010) doi:10.1139/F10-059 Published by NRC Research Press

wives could exacerbate water quality problems if alewives ing strong changes in zooplankton community structure and increase net nutrient loading into the freshwater ecosystems causing trophic cascades in their natal rearing lakes (Brooks where they spawn. 1969. This has caused concern among some lo- and Dodson 1965. Daily Load studies for nutrient abatement). Marine. and direct restoration efforts (Havey 1961). (1979) studied the movement of tant to the alewife nutrient loading model. alewives are iteroparous. 2009) and lakes in which they spawn. First. Dramatic declines have been budget. Net loading into freshwater ecosystems is fully de- will readily ascend fishways. Sci. 2000) primary water quality concern (e. Materials and methods tions on fish passes (Davis and Schultz 2009). The one load to that in the 1960s when alewives were more numer- available estimate of nutrient loading to lakes suggests that ous and larger. Can. the run is dominated alewife population that we used to both evaluate contempo- by smaller age 3 and 4 fish (Davis and Schultz 2009). we evaluate the patterns of net nutrient loading by For personal use only. J. gametes. (Schindler 1978). Naiman et al. 2010 freshwater ecosystems in which they reside. Bride Lake to Long Island Sound (see Dalton et al. ga. Despite restoration efforts. Aquat. 2009). than many Pacific salmon runs (Durbin et al. In the fall. on an areal the net nutrient loading for a newly restored population basis. Juvenile ale. Although the migration of obvious and much better studied (Moore and Schindler adult alewives is obstructed by even low-head dams. the ale. Vol. although inputs are typically more their natal systems to spawn. Aquat. at low densities and import nutrients when at high densities derived nutrients into freshwater ecosystems (Donaldson (Scheuerell et al.. Connecticut. the body size of alewife Study site spawners has decreased substantially. Finally. 1998). We then compare the current alewife nutrient (Garman and Macko 1998. import large quantities of nutrients as adult returns (escape- tic salmon. fish can export nutrients from freshwater ecosystems when cused on the role of Pacific salmon in loading marine. Historically.nrcresearchpress. 2008). and is linked to Long Island Sound productive. Coastal ecosystems throughout southern New by Bride Brook. 1212 Can. we estimate net alewife nutrient Walters et al. rary and historic net nutrient loading by an alewife popula- While Pacific salmon typically spawn in nutrient-poor tion and parameterize our general alewife nutrient loading lakes and streams where marine-derived nutrients are con. Juvenile alewives are cal stakeholders and has the potential to delay or prevent re- important prey for largemouth bass (Micropterus salmoides) storation efforts in watersheds where nutrient loading is a and pickerel (Esox spp. and excretion make anadromous alewives a poten. mental Protection Agency has mandated Total Maximum orous birds (Hartman 2003. While an enormous amount of research has fo. 1978). In addition to the decline in population size. Bride Lake is the site of extensive historical (Marcy sidered ecologically beneficial (Stockner 1987. model. which flows 3. 2005). We concentrate metes. Bilby et al. not 18 ha.3 km from the outlet of England are naturally mesotrophic and are typically im.g. 1979). where the US Environ- and adults are important prey for costal predators and pisciv.com by CLEMSON UNIVERSITY on 11/10/14 juvenile anadromous alewives migrate to the ocean where Anadromous fish move nutrients both into and out of they feed and grow for several years before returning to freshwater ecosystems.) (Cooper 1961. anadromous land waters. Like Atlan. Post and Walters 2009). consisted of fish that Pond in Connecticut. Downloaded from www. In this context. J. as it is generally the nutrient that limits pro- tially sizable source of marine-derived nutrients for the duction in the lake ecosystems in which alewives spawn streams through which they migrate (Post and Walters 2009. Post et al. Fish. Next. cally density dependent (Jonsson et al. we predict trogen and phosphorus to freshwater ecosystems. 2009). alewives over a range of population sizes. as previously reported Published by NRC Research Press . Connecticut. to the coastal ocean is the reestablishment of the movement Since juvenile growth and survival in freshwater are typi- of nutrients by diadromous fish between the ocean and in. Kissil 1974) and current research on alewife (Davis 2000. 67. across a range of adult escapement. but currently. driv. adults surviving spawning to return to sea (Cooper 1961. alewives transport nutrients into and Schultz 2009. alewife loading term in the context of the lake’s nutrient Limburg and Waldman 2009). Sci. Inputs from carcasses. since little is known about the actual anadromous alewives may contribute considerably more ni. Here. excretion of nutrients minus the removal of nutrients from One consequence of reconnecting freshwater ecosystems freshwater ecosystems by juvenile fish when they emigrate. the restoration of anadromous ale- wives are extremely effective size-selective predators. seen from 1950 to 2000 in both commercial fisheries land- ings and the passage of river herring through monitoring sta. Naiman et al. Gresh et al. with about half of ment) increase. loading and parameterize an alewife nutrient loading model derived organic matter from alewives has been shown to be using data from an existing run of anadromous alewives in incorporated into stream invertebrates and piscivores Bride Lake. Our research was conducted at Bride Lake and Linsley wife run at Bride Lake. 2009 bedded within a human-dominated landscape where cultural for map). 2009). much less is omous alewives to export nutrients from freshwater ecosys- known about the role of other anadromous fish as nutrient tems early in restoration when adult densities are low and to vectors. Yako et al. Fish. Dalton et al. This sets up the potential for anadr- 1969. making them a good target for scribed as inputs due to adult mortality. alewife an export coefficient model to estimate the landscape phos- populations are in decline throughout much of their native phorus loading to Bride Lake and Linsley Pond to place the range (Atlantic States Marine Fisheries Commission 2009. Kissil 1974. Bride Lake contains an anadromous were age 5 or greater. 2002). on phosphorus. It is located in and out of freshwater ecosystems that are typically quite East Lyme. Bride Lake is 28. we use nutrients by alewives. patterns of nutrient loading during restoration. Dalton et al. they 2004). 2002). Walters et al. particularly those in the Atlantic Ocean. We use a local sensitiv- Alewife populations have changed considerably since the ity analysis to determine which parameters are most impor- 1970s when Durbin et al. 1996.7 ha in area (based on a high-res- eutrophication is a persistent management concern (Brugam olution areal photograph.

The mass of phosphorus loaded per fish by gametes Root SR-1601 fish counter (2005–2008). Can. Downloaded from www. so all nutrients excreted during this time are of marine origin where ma is the adult in-lake mortality rate (per year). Lake in 2003 and 2006 (J. Davis. The phosphorus content of unspawned adults ([Pa]) adults returning to Bride Lake to spawn using a blocking was taken from Durbin et al. masse is the mass deep and 35. of each egg. the 1 – ma term was by YOY or juvenile fish as they emigrate from the ecosys. which corresponds to the median stay in freshwater (Pmort) was modeled as for alewives at the time when migration peaks (Kissil 1969). the general each adult that dies and the 1 – ma term was included to model estimates net phosphorus flux from alewives as avoid double counting Pgametes from adults that die. tion in TP inputs. The massa from the 1960s was determined spawning population and young-of-the-year (YOY) in the using the length of fish found by Kissil (1969) and the lake.7 m (Kissil of adult immigration and emigration taken at the Bride 1974). Fish. has a maximum depth of 10.9 km to the Branford Supply Ponds Dam and ken at the entrance of Bride Lake in early May of 2004 and then another 1. The na was used as a dynamic variable to look at pat. The mass concentration of 150 mg PL–1. 1213 in Kissil (1974)).01) derived from data in Kissil Branford Supply Ponds Dam in the fall of 2005. the Connecticut Department of Environ. Current massa was estimated Bride Lake anadromous alewife population was studied in directly from 120 sampled fish entering Bride Lake in 2004. The ovaries of 10 anadromous alewives ta- which runs 3. J. The Fa was Sound.64. In 2005. the lake had a spring TP Marlborough. r2 = to develop a Total Maximum Daily Load for the watershed 0. in mass of the testes was calculated as the mass of the testes tered the supply ponds. The change year that the fishway was opened. The initial mass of testes was estimated as a reach Linsley Pond in the coming years. and had an average summer total phosphorus (TP) Lake outlet in 1966 and 1967 by Kissil (1974) and in 2005 concentration of 21. With an annual time step. Excessive nutrient function of lengtha and based on the relationship masst = inputs from development and industry have led the CTDEP 0.43.3 mg PL–1 from 2005 to 2007. t = 13.563 (n = 289. The densities. Pmort includes Pgametes for for anadromous fish. The mass of phosphorus loaded through direct excretion where na is the number of adults. In 2005. and Pexcretion ¼ ð1  ma Þ  massa  Ea  ta Pexcretion is the phosphorus loaded into the lake through di. r2 = 0. Dmasst is the 13. PYOY is export of phosphorus freshwater by spawning adults. of spawned testes was taken to be 5. of their phosphorus to the lake. West et al.18 g (Durbin et al. Connecticut.45 m long and encircles an area of 100 m2. 3123 adult alewives en- For personal use only. CT 06447. A fishway was built at the t = 4. 209 Hebron Road. Thus. The 1 – ma term was included because we assume Alewife nutrient loading model that all of the phosphorus contained in a fish remains in the Here.01) for fish captured at the entrance to Bride in 2005.04. p < 0. Linsley Pond flows into Pisgah Brook.4 m located on the border of Branford and North Bran- change in mass of the testes. The Ea was calculated with alewives entering Bride (grams phosphorus per gram wet weight). The seine was 4. Eastern District Headquarters. In the first (1969) and from fish sampled in 2004 and 2005. included because we assume that fish that die contribute all tem. massa (Cooper 1961). 1979).P. Pmort ¼ ma  massa  ½Pa  Adults do not feed between migration and spawning. The by the CTDEP to estimate ma. Excretion is only accounted terns of net nutrient loading across a range of adult for in fish that survive spawning and leave the system.87 m where Fa is the fecundity of each female. The analysis found that Linsley Pond currently re. we present a fully described nutrient loading model lake when that fish dies. time that adult alewives spend in freshwater was set as The mass of phosphorus loaded per fish by adult mortality 14 days. Again. and [Pa] is the concentration of phosphorus in adults short. they are expected to spawned fish. and [Pt] is the concentration of ford. Sci.157 (gmm–1)  lengtha – 28. p <0. unpublished data). The river linkage between Linsley Pond and Long estimated as a function of adult length (lengtha) based on the Island Sound was severed in 1900 by the construction of relationship Fa = 3596 (mm–1)  lengtha – 766186 (n = 24. Pgametes is the phos- phorus loaded into the lake by inputs of eggs and sperm. We as- Net P flux ¼ na  ðPmort þ Pgametes þ Pexcretion Þ  PYOY sumed a 1:1 sex ratio. length–weight regression for current adult alewives in Bride mental Protection (CTDEP) has monitored the number of Lake.nrcresearchpress.18 mm mesh.com by CLEMSON UNIVERSITY on 11/10/14 weir (2003–2005. the Branford Supply Ponds Dam. Published by NRC Research Press . the fish (Pgametes) was modeled as counter was also used to estimate the number of adults leav- ing Bride Lake. the 1960s by Kissil (1974) to evaluate characteristics of the 2005. where Ea is the excretion rate of adults (grams phosphorus rect excretion of phosphorus by adults during their residence per gram wet weight per day) and ta is the time spent in in the freshwater ecosystem. and although no alewives have been of unspawned alewives minus the mass of the testes of confirmed in Linsley Pond to date. Division. We used counts Lake (Post and Walters 2009). and 2008. USA. CTDEP Inland Fisheries ceives ~100 kg P annually and recommended a 43% reduc.6 km to Branford Harbor and Long Island April of 2005 were measured for masse and [Pe].39. Since 2003. Adults may feed in the lake between spawn- is the average mass of the adults (wet weight) entering the ing and returning to the ocean but this time is generally lake. [Pe] is the concentration of phosphorus in each Linsley Pond is a 9. P in the testes.4 ha pond with a maximum depth of egg (grams phosphorus per gram wet weight). (1979).  ðFa  masse  ½Pe  þ Dmasst  ½Pt Þ gic purse seine with 3. Davis and Schultz 2009) and a Smith. Pmort is the phosphorus (Pexcretion) was modeled as loaded into the lake by adult mortality. YOY alewives in Bride Lake were sampled Pgametes ¼ ð1  ma Þ  0:5 monthly from June to August from 2004 to 2006 in a pela. Aquat.

mate of the response of the net nutrient flux: tion of a fish in the middle of the experiment. the atmosphere on top. One of the 0. 4.32 rameter by ±10% and recording the centered distance esti- fishm–2 treatments was omitted due to the inadvertent addi.30 mg Pm–3day–1. and 10. Vol. massa. early survival is likely density dependent and survival 2 m in diameter with 6 m deep cylindrical sides of solid could be considerably higher at densities lower than that polyethylene. Each mesocosm was ever.26 g (SD = 0. square metre) of YOY alewives. 67. which was 30 June in 2005 and 19 June in 2006. Fish were censused ures the percent change in the model output for a 1% change on 26 July and 16 August and a final count was made on in a parameter.1. They were closed on the bottom and open to seen in Bride Lake. Using the number of spawners and fecundity.27. Aquat. verted to the number of spawners (na) by fitting an exponen- eled as a function with an exponential rise-to-maximum as tial decline between the time when eggs were spawned on 1 the number of YOY individuals increases. 2008). and pi is the YOY produced was fit with an asymptotic exponential func. Fish.55. Sci. caught in Bride Lake in the falls of 2004 and 2005. 2006). and Ea by sequentially adjusting each pa- to be measured for length and mass. with densities of 0. we ment performed in Linsley Pond in the summer of 2005 (see can estimate the number of eggs spawned and measure the below). Phosphorus per mesocosm) replicated three times. a is the maximum biomass density (asymptote). we matched a to the biomass den- relates the biomass density of YOY alewives to the number sity of YOY alewives observed in midsummer in Bride Lake of adult spawners. [Pt]. Aquat. Yð1:1  pi Þ  Yð0:9  pi Þ We fit a negative exponential model to the growth rate of si ¼ YOY alewives across stocking densities and a linear model 0:2  Yðpi Þ to mortality rate across stocking densities. 1. Phosphorus content of fish (adults. 2010 The mass of phosphorus exported from the lake by YOY YOY alewives at stocking. so we expected to obtain greater densities of YOY alewives in the Linsley Pond mesocosms than in PYOY ¼ nYOY  massYOY  ½PYOY   area Bride Lake. Linsley Pond is more productive alewives (PYOY) was modeled as than Bride Lake. ma.1 mm) and an average mass of 0. in the estimate of potential For personal use only. which was 1. gametes. Therefore. The sides of the mesocosms were to the estimation of net nutrient flux.09 g). content of eggs and YOY was estimated by ashing dried ments. The survival of alewives in periment in Linsley Pond to estimate the density-dependent Bride Lake provides a best estimate of early survival. cosms. The [PYOY] was measured in 13 YOY alewives seine. 5. Astoria 2 autoanalyzer (Higgins et al. The initial density x of YOY alewives was con- from the lake was calculated on an areal basis and was mod. Y( ) is between initial fish density and final biomass density of the net nutrient flux for the given parameter set. The other five treat. We varied na. 30 August when all fish were removed from the mesocosms Dmasst. [PYOY] is the concentration of phosphorus for 2004 and 2005.64. This density-dependent relationship reflects a de. tion: We analyzed the sensitivity of the full model to the esti- mated biomass density of YOY fish (nYOY  massYOY) with nYOY  massYOY ¼ a  ð1  ebx Þ the same method. parameter being adjusted (Ellner and Guckenheimer 2006). b is the slope of the relationship be. how- growth and survival of YOY alewives. 8. Downloaded from www. We stocked YOY creases 10-fold from that seen in Bride Lake. Fish stocked into mesocosms had an aver. ta. Mesocosms were lowered to around nutrient export from Linsley Pond. The mesocosm experiment provides a good estimate of the density-dependent growth of alewives larger than Density-dependent growth and survival of YOY alewives 30 mm in Linsley Pond. anadromous alewives from Bride Lake into the mesocosms on 12 July 2005 at densities that spanned the range of den. age length of 32 mm (SD = 3. Fish. were first able to estimate the population size using a purse creases. Watershed loading model tween density and biomass. (1979).60 gm–2 (Post et al. J. [Pe]. and analysis of liberated soluble reactive phosphorus on an cated two times. in each YOY alewife (grams phosphorus per gram wet The year 2006 was excluded because an early emigration weight). The sensitivity meas- scrubbed clean of periphyton weekly. and area is the lake area in hectares. and x is the initial density of To understand the impact that alewives are having on Published by NRC Research Press . 2. were repli. adjusting the asymptote and slope para- where nYOY  massYOY is the biomass density (grams per meters by ±10% and recording the change to the full model. Fa. survival of smaller alewives. masse. We Phosphorus content of unspawned adult alewives and ale- had one treatment with a density of 0. The relationship where si is the sensitivity of the model to parameter i. To translate the biomass from the experiment where nYOY  massYOY is a density-dependent function that to biomass in Bride Lake. we conducted a mesocosm ex. We conducted a local sensitivity analysis of our alewife Nitrogen and phosphorus were added to each mesocosm at a nutrient loading model for the current alewife run in Bride 25:1 molar ratio based on a phosphorus loading rate of Lake to determine the relative importance of each parameter 0. case where the survival of YOY alewives to 30 mm in- proximately 19 m3 of water from the lake. and YOY) sities observed in local lakes with anadromous alewives. we also examine the 7 m depth and raised through the water column isolating ap. J. 1214 Can. but we do not know the growth or In the summer of 2005. The form of May and 12 July. slope of the exponential decline to the time at which we crease in growth and survival as the number of juveniles in.32 fishm–2 (one fish wife testes was taken from Durbin et al.19 samples in a muffle furnace followed by digestion with HCl fishm–2 (2. 16. and 32 fish per mesocosm). when fish were stocked into the meso- nYOY  massYOY was estimated from the mesocosm experi. The biomass event of YOY alewives greatly reduced their midsummer Can. Sci.nrcresearchpress. [Pa].com by CLEMSON UNIVERSITY on 11/10/14 density of YOY alewives (nYOY  massYOY) emigrating density. Fish were stocked at this size because Sensitivity analysis fish <25–30 mm are difficult to keep alive during transport.

the slope (decay constant) of the exponential decline of the TPin YOY population in Bride Lake was found to be –0. Connecticut (42 km to the When one outlier was removed. 11 Phelps Way.39 YOY alewives surviving to 12 July (start of the mesocosm experiment) per adult spawner. 1215 their natal systems. 2c).89. adult spawners and YOY biomass export of phorus to Bride Lake. Predicted At the current escapement of 90 000 adults.26  e–0. With the months.001) (Fig. Each survival is the same as in Bride Lake. Pond mesocosms was strongly density dependent (Fig. AECOM the density-dependent growth of the YOY alewives. CT 06279.10 = 36. tional Land Cover Database to determine areal land uses Growth rate declined with increasing initial stocking den- (Homer et al.46 g fish less than 30 mm increased by an order of magnitude so of phosphorus to the system. The extent of the watershed of Bride Lake YOY growth and survival was determined using the National Hydrography Dataset Growth and survival of YOY alewives in the Linsley and overlaid on land cover information from the 2001 Na. The Pgametes is split nearly evenly between males (Johnes 1996). r2 = 0.01) sonable. phosphorus added to Bride Lake in the 1960s was 2.42 gm–2  ington. adult alewives total nutrient load from the landscape (Reckhow and Simp. 2a). Downloaded from www. if early the nutrient loading model are provided (Table 1). p < 0.6 kg. 3). result- Global Environment. nYOY  massYOY ¼ 2:67 kg  ð1  e0:000061na Þ (Fig. p < 0. where Y is the daily indi- nutrient inputs from the approximately 50 blue heron (Ardea vidual mortality rate and x is the initial density of YOY ale- herodias) individuals nesting in the watershed were included wives (n = 11. Will. and it showed a greater than it is today (Fig. USA. 2004). from 1982 to 1996. known as the phosphorus days 14 and 35 of the experiment. July. Export coefficient models are excretion (Pexcretion) with gametes (Pgametes) contributing the widely used to estimate the nutrient loads from watersheds remainder. entering Brides Lake were larger than present-day alewives son 1980). The average population that flows overland (runoff) and subsurface (baseflow) and returning to Bride Lake from 2003 to 2008 was nearly produces an estimate for the nutrient yield of a watershed 90 000 fish. In the 1960s. but the net input would have been much higher with 100.84x. tween initial stocking density and daily mortality rate was ues of loading estimates for each land use were chosen and found with Y = 0. land uses. Wagner. The proportion of phosphorus that makes it was a treatment with two fish in which one died between through the watershed to the lake.67 kg of phosphorus.0004. adult population of 162 000 found in 1960s.7 ha). (1 – e–2. The outlier in the model. p < 0. where Y is growth rate (grams per day) and x has changed little due to the presence of two large prisons is the initial density of YOY alewives stocked on 12 July in the watershed. an and contributed 39% more phosphorus per fish (mean of export coefficient model that was developed for use in 0. phorus loading to lake phosphorus concentration.77. aged over 160 000 fish. Although we do not have land-use in. Median val. the resulting relationship would be nYOY  massYOY ¼ with each carcass contributing 0. where x is the initial stocking density of YOY To ensure that the estimate of phosphorus loading was rea. Aquat. West et al. we estimated background phosphorus that survive each add 0. alewife in the mesocosms (n = 12.0006x – 0. which estimates the lake phosphorus concentration (TPlake After multiplying the density of fish by the area of the lake (micrograms per litre)) using the inflow phosphorus (TPin (28. r2 = 0.001) (Fig.13 day–1 TPlake ¼ 1 þ 1:12ðyear0:47 Þ  t 0:47 w to give an estimate of 6.18 g of phosphorus. This estimate is reason- Published by NRC Research Press . After adjusting the asymptote for Bride Lake. of which 67% is loading into Bride Lake. 1).73x). 1). These models use an estimate of the nutrient and females.8 kg of Results phosphorus imported to Bride Lake. Box 506.0112 + formation from the 1960s. r2 = 0. personal communication (2009)). 2). 2b). Alewife nutrient loading model For Linsley Pond.46. the Bride Lake watershed likely 0. The values for all parameters used in nYOY  massYOY ¼ 2:96 kg  ð1  e0:000061na Þ. Annual precipitation was calculated from monthly precip. the resulting relationship between spawner density (micrograms per litre)) and the water residence time (tw and the mass of YOY alewives in Bride Lake was (years)). and basin characteristics. Fish.66 kg and the net phos- in the pelagic zone of Bride Lake during the summer phorus input to Bride Lake by alewives is 38. As a result. Survival declined with increasing initial stocking density. while in 1966 and 1967. close match to other estimates of nutrient loading (Stahl and Bolton 2005). the model output was entered into the equation (Fig. Sci. The difference in nutrient loading between southern New England (AECOM Technology Corporation the present and the 1960s is magnified when the size of the Can. Adults 2:96 kg  ð1  e0:00061na Þ (Fig.5 times mum Daily Load analysis of Linsley Pond.9 YOY alewives surviving to 12 (Pmort) make up 83% of the total alewife phosphorus input.com by CLEMSON UNIVERSITY on 11/10/14 2009). F2. ing in the relationship nYOY  massYOY = 5. was considered to be moderate due to the An asymptotic exponential function provided a good fit to presence of a wetland in the basin (K. itation records from Middletown. basin attenuation. a linear relationship be- For personal use only.68 g of phosphorus. YOY export is expected to have been similar at 2. northwest of Bride Lake). Brett and Benjamin (2008) found this relationship to perform better than several older models relating phos. The Lake Loading Response Model considers water populations is taken into account. as the higher phosphorus content of testes com- yield for each land-use type in a watershed to calculate the pensates for their lower mass. sities as a negative exponential in the form of Y = 0. we estimate a relationship between Adult alewives contribute considerable amounts of phos. Adults that die in the system each spawner resulted in 63. the phospho- TPlake was compared with measured values of TPlake taken rus export from YOY alewives is 2.64 g Pfish–1). 2005 (n = 12. We used the Lake Loading Response Model. the total amount of The model has been used by the CTDEP for Total Maxi.57. If the survival rate of adult entering Bride Lake currently adds a mean of 0.nrcresearchpress. J. the population aver- based on precipitation. P.O.

and [Pt]) had little impact on the net nu. Adult phosphorus concentration had a sensitivity (83%) of the input from alewives is from alewives that die of 0.0031 g Pg egg–1 1 D testes wet mass (Dmasst) 7.17 mg Pg–1h–1 6 Adult time in system (ta) 14 days 2 YOY P concentration ([PYOY]) 0. Fig.03 each. Comparing the phosphorus concen- tration of Bride Lake with the Brett and Benjamin (2008) model shows that our loading estimate is reasonable but is likely an overestimate of the true phosphorus load to the lake.0042 g Pg fish–1 4 Adult fecundity (Fa) 3596  lengtha – 766 187 1.07. which affect years. with sensitivities of 1. Of this. Pgametes. Fish. 1216 Can.3 kgyear–1. 67. 2 Adult P concentration ([Pa]) 0. as Havey (1973) found an average of 366 YOY ale. 11. 222. Landscape loading model The parameters used for the lake loading response model For personal use only. with a peak of 58% of the phosphorus budget for the input from Pmort.6 kg input from ale- wives brings our total estimated phosphorus load to Bride Lake to 169. Alewives currently import 23% low following the restoration of an alewife population in of the total phosphorus input to Bride Lake.4 g (1960s) 1. Aquat. which For the Bride Lake nutrient loading model. Fish. supply 3%–28% of lake phosphorus budgets in normal tion size and adult body size (na and massa). Vol. greater in the 1960s than at present because of both greater masse. 4. Davis (CTDEP Inland Fisheries Division. with sensitivity values of just 0.5 kg was from waterfowl. 1998). J.nrcresearchpress. The amount of phosphorus added annually to Bride Lake from external sources (excluding alewives) was 130.12. shows that alewives are loading substantial quantities of wife emigrants per adult spawner when adult densities were phosphorus to Bride Lake. 2. 2 Egg P concentration ([Pe]) 0. Trait Value Reference(s) Adult numbers (na) Variable 1. adult densities seen in Bride Lake. run size and larger body size in the 1960s than at present. Sci. increases to 44% for estimates from the 1960s. The Brett and Benjamin (2008) model predicted a lake phosphorus concentration of 23. The Although fish were only 38% heavier and 80% more numer- adult time in the lake and excretion rate (ta and Ea) each ous in the 1960s.6 mgL–1. Eastern District Headquarters. Kissil (1974). Discussion The full model for nutrient loading by an iteroparous fish able. We do not know the size of the alewife run Published by NRC Research Press .56year–1 2. (2009). Maine.00012 g 1.65.0088 g Pg testes–1 3 Adult excretion rate (Ea) 2. CT 06447. Aquat. This study.com by CLEMSON UNIVERSITY on 11/10/14 Egg wet mass (masse) 0. 11% greater than the observed average summer phosphorus concentration. The 38.6 kg was directly deposited on the lake from the atmosphere. [Pe]. eter (b) had nearly no effect on the overall model at the doharengus) and landscape sources across a range of adult densi. and this number Love Lake. Dalton et al.5 times had a sensitivity of 0. adult popula.08 g (present). unpublished data).0058 g Pg YOY–1 1 Note: 1. Downloaded from www. 209 Hebron Road.P. 5. J. the majority spectively.5 g (present). The nutrient input from alewives was much with a sensitivity of 0. J. were the most Lake Iliamna in a peak year for sockeye returns (Stockner sensitive parameters. 1987. Post and Walters (2009). The parameters for Pgametes (Fa. 2 Adult mortality rates (ma) 0.88. the combination resulted in 2.07 and 1. The asymptote of juvenile biomass greater phosphorus input by alewives in the 1960s than in density (a) had a sensitivity of 0. Marlborough. Gross et al. Phosphorus flux to Bride Lake from alewives (Alosa pseu. 2 Can. and 4. while the slope param. USA. 5 Testes P concentration ([Pt]) 0. Parameters and sources for the alewife (Alosa pseudoharengus) nutrient loading model. while adult mortality was slightly less important in the system. Average current (2003–2008) and 1960s (1966–1967) adult alewife escapement indicated by vertical bars. 2 Adult length (lengtha) 260 mm (present).12. (1979). ties. export coefficient model are listed in Table 2. Sci. 2010 Table 1. trient flux. 289 mm (1960s) 1. 6. 1. Not surprisingly. the early 2000s. Durbin et al. re.6 kg of phosphorus was from the watershed. 3 Adult wet mass (massa) 161.60 g (1960s) 3. Dmasst. and Pexcretion. 1). 8.7 kg (Fig. These num- bers are relatively high compared to the phosphorus loading Sensitivity analysis reported for sockeye salmon (Oncorhynchus nerka). 3. 117.

Although they have not yet reached Linsley Pond.nrcresearchpress. growth and survival current levels of adult escapement.com by CLEMSON UNIVERSITY on 11/10/14 be a net exporter of phosphorus when adult populations are small. prohibiting emigration. The triangle in Fig. early in the recovery of an alewife popu- lation. In the Bride Lake model. The growth of YOY alewives in managing flow regimes to allow alewife emigration through- mesocosms was strongly density dependent. parameters related to of juveniles smaller than 30 mm are strongly density de. Can.9 YOY per spawner surviving to 12 July (gray dot–dashed line). Linsley Pond. with good early survival. If these fish had moved into Linsley Pond rather than remaining in the artifi- cial supply ponds. then it is possible that alewife populations could to nutrient loading. 1217 Fig. however. If 63. Aquat. we believe that alewives are rarely nutrient cific competition throughout the growing season. phosphorus input by adult mortality are the most important pendent. YOY alewives can migrate an even more important part of the nutrient budget of to the ocean throughout the summer. unpublished data). Yako et al. the outlet streams often go dry for most of the were likely considerably less (Brugam 1978) and alewife summer months. 3.39 per adult estimated from early survival in Bride Lake. more than 3100 fish ascended the Branford fishway into the Branford Supply Ponds the first year that it opened (D. Note that axes scales differ from those on Fig. we predict they would have imported 0. As a result. Sci. ley Pond. West et al. possibly due to an increase in intraspe- for Bride Lake. sensitivity analysis shows that at trient inputs of the parents. we estimate that the population will be a net exporter of phosphorus until the population reaches 6500 adults (690ha–1). YOY export pre- Linsley Pond mesocosm experiment.39 YOY per excluded from the analysis of density-dependent mortality. If. 2. One possible source of bias in our estimates of nutrient ex- port by YOY alewives is that juveniles can leave their natal lakes at different times throughout the year. but landscape nutrient inputs outlet streams. Thus. For Bride Lake.9 juveniles per adult survive to 12 July in Lins- For personal use only. In- put by adults is for current fish (black dashed line). rather than the 6.8 kg of phosphorus to Linsley Pond and. but the early out the summer may both benefit alewife populations and in- survival rates that we measured in Bride Lake were too low crease nutrient export from eutrophic systems.M. Estimated phosphorus flux due to alewives (Alosa pseudo- (c) biomass production for alewives (Alosa pseudoharengus) in the harengus) in Linsley Pond following restoration. The loss of some individuals early in the year may release the remaining fish from density dependence and increase the total biomass of YOY fish that emigrate from the system. exporters for a watershed. (2002) found coastal lakes in precolonial New England. Patterns of density-dependent (a) growth. Post. In wet years and in run size considerably larger. net loading is likely to be small or even slightly neg- ative (net export) where YOY survival and growth are high. suggesting that alewives were lakes with larger outlet streams. they could have exported greater than 1 kg of phosphorus. 2b was dicted from juvenile growth in mesocosms with 6. that alewives in lakes where they cannot emigrate throughout Based on the in-lake-survival of YOY alewives observed the year were smaller. 1. to produce a sufficient number of young to exceed the nu. Downloaded from www. J. (b) mortality. and Fig. Fish. and many other coastal lakes with low-gradient before European settlement. spawner surviving to 12 July (black dot–dashed line) and 63. Adult escapement and adult body size Published by NRC Research Press .

but tle effect on the overall nutrient budget.50 0. water basin attenuation 0. which may vary considerably transport much of the nutrients away from the lake as allo- between lakes and could have a strong effect on the amount cation to growth and reproduction or as feces. Fish. Although Bride Lake is many times number of adults will not have a large effect on the biomass larger than the stream. bringing the total contribution from populations for one of two goals: minimizing nutrient inputs excretion to as much as 26% of the total alewife phosphorus while allowing some alewives to spawn or maximizing ale. the YOY because although an increase in adult mortality increases the biomass exported would be expected to be greater than 75% phosphorus input from mortality.004 Meadow 25. the retention of car- population is near the asymptote of YOY biomass density.10 0. Since ju.50 0. park. casses in the lake should be high.010 Upland 377.1 ha lot commercial Institutional. Runoff Baseflow Runoff P export Baseflow P export Land use Area (ha) fraction fraction (kgha–1year–1) (kgha–1year–1) Notes Low-density residential 40.010 <0. which is body size. for exam. input.com by CLEMSON UNIVERSITY on 11/10/14 or recreational Agricultural.87 0.4 ha lot Medium-density residential or 8.05 0. and adult mortality are expected to vary consider.10 0. Downloaded from www.9 kg of phosphorus could be excreted in the lake by coastal ponds will allow for adaptive management of alewife fish that eventually die. they remain on the lake for the majority of the day.60 0.3 1. as excretion from fish is. which drains Bride Lake. Nislow et al. ably from system to system. The net nutrient input gametes and excretion.8.010 >0.28 0. J. One exception is the time of tality in Bride Lake (Dalton et al. 2010 Table 2. came from excretion. In Linsley al. and P basin attenuation 0.40 0. (1979) who found that 18% of alewife phosphorus input Pond.26 0. this where water flow necessitates an explicit consideration of parameter is very important to the overall nutrient budget.1–0. Since the YOY production is a small frac. it is important that they are ac. likely because it is not important to the net ers could constrain the number of fish that are allowed into energy budget of semelparous species.25 1. Aside from mobile predators.25 1. J.15 0.010 Agricultural 36. Sci. The nu- tion of the net nutrient flux at the current escapement levels trients excreted by cormorants are not available for direct for Bride Lake.010 Can. managers can control the number of ale.3 0.6.3 kg of phosphorus. manag. the YOY biomass density asymptote has lit. uptake by primary producers. Since adult mortality is likely not instantaneous. for example. row crops 1.65 0.010 0. unlike in lotic systems In lakes that have smaller adult populations. 66.3 0. Sci.4 ha lot highway corridors High-density residential or 1. since the 2002). Since the model is not very sensitive some of the nutrients from carcasses can be removed from to any of the factors affecting the phosphorus load from ga.004 Wetland/lake 55. however.15 2. cemetery. Once bolic processes from anadromous fish has received relatively a run of alewives becomes established in the pond.004 Note: Other model parameters: precipitation 1.5 kg Pyear–1 (50 birds). For personal use only. adult for this many fish would be 8. little attention. Vol. We estimate that 11% of the phosphorus input from ale- ple.45 0. carcass retention (Garman 1992. 1218 Can. In the projected nutrient budget for Linsley Pond. although considerably less than population size or body size capping the adult run at 23 000 fish (2450ha–1). Adult mortality also has a very strong impact. Although the birds adult residence in freshwater.10 0. The nutrient loading estimate from adult mortality that we curately measured to get a good estimate of net phosphorus provide is an upper estimate for the nutrient load to lakes. Aquat. YOY export term. In the Bride Lake feces are mineralized by microbes to available forms (Vanni model. although the lake is considerably longer than in the stream. it decreases the input from of the maximum capacity for the lake. If early survival of input terms.05 2.23 0. changing the (Walters et al. 2009). an addi- Knowing the nature of the nutrient flux from alewives to tional 5. 67. as they are important for all the number of juveniles will be affected.05 0. 2009).00 0.91 0.3 0. has shown that alewife ex- at a point where YOY export is nearly maximized.50 0.4 0.26 0. Aquat. Since adult population size.40 0. as loading from alewives.3 myear–1 (Middletown. Connecticut).20 0. waterfowl 4.4 0. they excrete a of excretion and the overall model if differences are on the substantial proportion of their diet back into the lake. have the highest sensitivities. the residence time for alewives in of juveniles that survive to emigrate from the pond. less than 10% of the current phosphorus load to the pond. the slope parameter has virtually no effect.3 0. 2004). the model is very sensitive to the slope parameter of the wives is due to excretion of alewives that survive spawning. At present.nrcresearchpress. the system by mobile predators.3 0. alewives in Linsley Pond is similar to that in Bride Lake. these parameters are less important to wives. The importance of inputs from meta- wives that ascend the Branford Supply Ponds fishway. since order of weeks.20 0. cretion is an important source of nutrients for the stream venile growth is strongly density dependent. Fish. This estimate closely approximates that of Durbin et wife returns while limiting nutrient loading.40 0. Parameters for the Bride Lake export coefficient model. responsible for as much as 48% of the alewife mor- measure in different systems. Recent work in Bride the pond to a point where there is nearly zero net loading or Brook. cormorants metes or excretion and these factors are not expected to vary (Phalacrocorax auritus) are a major predator of adult ale- greatly between lakes. resulting Published by NRC Research Press .

. Higgins. Fish. N. Ecosyst.CO. dromous alewife (Alosa pseudoharengus) in south-central portant but often overlooked piece of the nutrient budget for Connecticut.2. Early life history and spawning migration of Can.0.00365. of lake phosphorus retention and the nutrient loading concept. CT.. Kingston.R..x. the Interstate Fishery Management Plan for shad and river herring. University of 2009). composition of plankton. come addition to lakes that are already eutrophic. M.E. Wash. 2009. Fisheries US Environmental Protection Agency STAR Graduate Fel- (Bethesda). Contribution of marine-de- Acknowledgements rived organic matter to an Atlantic coast. 01862. 052[0917:PSIAAT]2. for the lake loading response model.C. Excretion from surviving Brett. and life history of an anadromous alewife population in Connec- tional Marine Fisheries Service 2006). D. doi:10. doi:10. 2002. phorus samples.org/Pages/ Havey. Fransen. Willson. 1996. Although Garman.x.B. Science. 1992. 1973. 1978. Bilby.J. J. P. Washington County. Princeton. 17(3): 277–285. Alosa pseudoharengus. 2006. Ph.150.CO. University of Washington. Biol.M. Trans.I. nity to restore fish populations and strengthen the overall Donaldson.2007. Manag. AE.. M.A. doi:10. A. and J. PMID:17829740. Maine. P. anadromous alewife population. and L. Detritivory Fish. may be coming from excretion before senescence. 25(1): 15–21. Ecology.. M. restoration would be a de. alewife populations can be adaptively managed to minimize Trans. Havey.A. anadromous sockeye salmon in the nutrient loading and produc- tivity of Redfish Lake.1. The impact of dou- cretion is in an inorganic form that can be readily used by ble-crested cormorant (Phalacrocorax auritus) predation on ana- primary producers. West et al.1577/1548-8659(1973) nitrogen and carbon from spawning coho salmon into the trophic 102<434:POJAAP>2. doi:10. Project No. C.1139/F08-198. tidal We thank E.easthamptonct. S.2.1365-2427. stream by anadromous clupeid fishes. freshwater. 1965. 53(1): 194–211.P.C. Sci. For most diadromous fish as related to the cyclic abundance of sockeye salmon. G. M. and Bisson. COM Technology Corporation. Pond. 53(1): 164–173. The claim of Johnston et al. Brugam. ing anadromous fish carcasses in an Atlantic coastal river. Sci. Alosa pseudoharengus.28. doi:10.. body size. and the National Science Foun- Gross. Ecology. and González. 121(3): 390–394.3. 2000.1577/C08-003. 60(1): 8–17. AECOM Technology Corporation..1577/1548-8659(1998)127<0001:TROASS>2. ial photographs. Princeton University Press. Benthol.. Dalton. Soc.P. J. a freshwater systems of the Pacific Northwest.R.. Willington.M. Garman. Wagner provided valuable support Pacific salmon in aquatic and terrestrial ecosystems.. Ecol. Sci. H. Maine. D. J. doi:10. 66(2): 177–186. Ratliff for running phos. system of small streams: evidence from stable isotopes. Am.A. 1979. 0717265). Effects of the the demolition of dams or the installation of fish ladders. R. Downloaded from www. Production of juvenile alewives.P.2307/1468331..1139/cjfas-53-1-164.. Fish. 1998. J. 8659(1992)121<0390:FAPSOP>2. Nixon. Use of the lake loading Hartman. J. CT.: Dynamics Manag. The additional nutrients seen in the natural system Rhode Island.0.3692. do not excrete phosphorus when they are not Brooks..CO. W.1365-2400. Wurtsbaugh. doi:10. Davis for providing alewife Gende. the alewife. R. lowship Award to A.M. Soc. Lichatowich. K. Trans. on Alewives. and feeding is not relevant to breeding individuals. R. fisheries managers have the opportu. Population-level consumption by Atlantic response model (LLRM) in TMDL development for Lake Poco.pdf. Soc.S. and Schultz. and Wipfli. 127(1): 1– References 18. at Love Lake.2307/ For personal use only..A.1577/1548- the extent of potentially harmful nutrient loading.2307/1936629. By providing passage ticut. J. et al. North Am. thesis. Atlantic States Marine Fisheries Commission. Temporal shifts in demography the listing of alewife as a species of special concern (Na. spawning migration of the alewife. 1998. rhynchus spp. Am. anadromous fish (Post and Walters 2009).. ence in nitrogen and phosphorus dynamics between natural Cooper.C. Edwards.2.A. doi:10. and Dodson. coastal striped bass and the influence of population recovery topaug. Can. 150(3692): 28–35. 2008. J. Ellis. We thank three anonymous reviewers for Gresh. C. E. T. sis. Fish.P. (2004) that Onco. and Guckenheimer. 1961. Available from www. doi:10.0. Ellner.Sc. S. 2009. Can. D. A review and reassessment alewives alone equals 3% of the external load of the phos.. 2003.. Soc.A. 2006. the- whose populations are declining. 90–106. Am. 1(1): past existing barriers. S. The role of dation (DEB No.G. D. Coast. A recent meta-analysis shows a striking differ. Seattle. sirable outcome and environmental groups would rally for Durbin.J. embedded in the densely populated coast of eastern North 1936461. G. Sci. East Hampton.1046/j.. Butman assisted 52(10): 917–928.. 1969.2. M.0. M. The decline in the size of existing alewife runs has led to Davis. ley Pond mesocosm experiment. and Post. B. doi:10. America where the nutrients that they import are an unwel. however. An estima- helpful comments on this manuscript. S. 1219 in more nutrients being excreted.nrcresearchpress. doi:10. Palkovacs for field assistance in the Lins. Fish. Fish. 12095-005.W. and the stoichiometry of nutrient cycling by a dominant fish Published by NRC Research Press . Phosphorus budget of Iliama Lake. Aquat.T.D. USA. Vanni. Washington. ology changes dramatically as they prepare to spawn (Gende 1126/science. This research was tion of historic and current levels of salmon production in the northeast Pacific ecosystem: evidence of a nutrient deficit in the supported by the Connecticut Institute of Water Resource. phorus to the lake at present and would have been 5% in Freshw.L.I. 10(5): 281–288. whose physi. Soc. R. K. K. Incorporation of Am. and Luecke. harengus... 1961. Amendment 2 to 1577/1548-8659(1961)90[281:ROAAAL]2. and Macko. Alaska.J. Bioscience.1111/j. Mar. Since ex.F.M. doi:10. face the additional obstacle of being freshwater ecosystems. Fish.W. K.T. Fish. 90(3): 281–286. Idaho.CO. Predation. 2009. Human disturbance and historical development of Linsley Pond. C.com by CLEMSON UNIVERSITY on 11/10/14 salmon runs and artificial carcass additions (Janetski et al. 59(1): 19–36. doi:10. Bonneau assisted with aer. doi:10. J. Aquat. K.A. S. Trans.2. M.J. 102(2): 434–437. and Oviatt. T. Aquat.CO. Restoration of anadromous alewives at Long LakePocotopaugLLRMAug09.2003. Alosa pseudo- Atlantic States Marine Fisheries Commission. upon prey communities.1641/0006-3568(2002) with GIS methodologies. it appears that excretion may be an im. and Schoonmaker. J. the 1960s. Dynamic models in biology. 2009. Fish. Quality Assurance Project Plan.A.T. and Benjamin. physiological data. R. 2002).W. Fate and potential significance of postspawn- this makes alewife restoration projects more challenging.

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