Determination of food web support and trophic position of the mummichog,Fundulus heteroclitus, in New Jersey smooth cordgrass (Spartina alterniflora), common reed (Phragmites australis), and restored salt marshes

The invasion ofPhragmites australis into tidal marshes formerly dominated bySpartina alterniflora has resulted in considerable interest in the consequences of this invasion for the ecological functions of marsh habitat. We examined the provision of trophic support for a resident marsh fish,Fundulus heteroclitus, in marshes dominated byP. australis, byS. alterniflora, and in restored marshes, using multiple stable isotope analysis. We first evaluated our ability to distinguish among potential primary producers using the multiple stable isotope approach. Within a tidal creek system we found significant marsh and elevation effects on microalgal isotope values, and sufficient variability and overlap in primary producer isotope values to create some difficulty in identifying unique end members. The food webs supportingF. heteroclitus production were examined using dual isotope plots. At both sites, the δ¹³C values ofF. heteroclitus were clustered over values for benthic microalgae (BMI) and approximately midway between δ¹³C values ofSpartina andPhragmites. Based on comparisons of fish and primary producer δ¹³C, δ¹⁵N, and δ³⁴S values, and consideration ofF. heteroclitus feeding habits, we conclude that BMI were a significant component of the food web supportingF. heteroclitus in these brackish marshes, especially recently-hatched fish occupying pools on the marsh surface. A 2‰ difference in δ¹³C betweenFundulus occupying nearly adjacentSpartina andPhragmites marshes may be indicative of relatively less reliance on BMI and greater reliance onPhragmites production inPhragmites-dominated marshes, a conclusion consistent with the reduced BMI biomass found inPhragmites marshes. The mean δ¹³C value ofF. heteroclitus from restored marshes was intermediate between values of fish from naturally occurringSpartina marshes and areas invaded byPhragmites. We also examined the isotopic evidence for ontogenetic changes in the trophic position of larval and juvenileF. heteroclitus. We found significant positive relationships betweenF. heteroclitus δ¹⁵N values and total length, reflective of an increase in trophic position as fish grow.F. heteroclitus δ¹⁵N values indicate that these fish are feeding approximately two trophic levels above primary producers.     

Impact of common reed,<Emphasis Type="Italic">Phragmites australis</Emphasis>, on essential fish habitat: Influence on reproduction,embryological development,and larval abundance of mummichog (<Emphasis Type="Italic">Fundulus heteroclitus</Emphasis>)

The invasion ofSpartina marshes by the common reed,Phragmites australis, along the east coast of the United States over the last several decades has been well documented, although we know little about the impact of this invasion on the fish fauna and the few published papers seem contradictory. During 1999–2000 (May–September) we evaluated the fish response to vegetation type (Phragmites australis veersusSpartina alterniflora) by monitoring several aspects of fish early life history (egg deposition, embryonic development, hatching success, and larval and juvenile abundance) in low salinity marshes in the Mullica River in southern New Jersey. The dominant fish species using the marsh surface,Fundulus heteroclitus (93% of total catch, n=996 individuals), reproduced in both vegetation types with eggs deposited in leaf axils near the base of the plant inSpartina and in broken stems ofPhragmites during both years. These eggs also undergo successful embryonic development to hatching in both vegetation types. Larval and juvenile (5–75 mm total length, but 95% < 34 mm TL) abundance of this species is much reduced onPhragmites-dominated (mean CUPE=0.02, n=7 ind) marsh surface relative toSpartina (mean CPUE=2.31). These findings, and similar results for fish abundance in 1997 and 1998, indicate that theSpartima marsh surface is likely essential fish habitat for this species because it provides habitat for larvae and small juveniles, whilePhragmites does not. ThePhragmites invasion in brackish marshes may be having deleterious effects on fish populations and possibly on predators that prey uponF. heteroclitus, and as a result, marsh secondary production.     

The influence of<Emphasis Type="Italic">Phragmites</Emphasis> (common reed) on the distribution,abundance, and potential prey of a resident marsh fish in the Hackensack Meadowlands,New Jersey

This study investigates the influence ofPhragmites australis (common reed) invasion on the habitat of the resident marsh fish,Fundulus heteroclitus (mummichog) in the Hackensack Meadowlands, New Jersey. These abundant fish play an important role in the transfer of energy from the marsh surface to adjacent subtidal waters and thus estuarine food webs. The objectives of this 2-yr study (1999 and 2000) were to compare the distribution and abundance of the eggs, larvae, juveniles, and adults of mummichog and their invertebrate prey inhabitingSpartina alterniflora-dominated marshes withPhragmites-dominated marshes, and to experimentally investigate the influence of marsh surface microtoprography on larval fish abundance withinPhragmites-dominated marshes. In 2000, we verified that egg deposition does occur inPhragmites-dominated marshes. In both years, the abundance of larvae and small juveniles (4–20 mm TL) inS. alterniflora was significantly greater than inPhragmites-dominated marshes, while larger juveniles and adults (>20 mm TL) were similarly abundant in both habitat types. The overall abundance of larvae and small juveniles was significantly greater in experimentalPhragmites plots in which microtopography was manipulated to resemble that ofSpartina marshes than inPhragmites control plots. Major groups of invertebrate taxa differed between marsh types with potential prey for larval fish being significantly more abundant inS. alterniflora marshes.Phragmites-dominated marshes may not provide the most suitable habitat for the early life-history stages of the mummichog. The low abundance of larvae and small juveniles inPhragmites marshes is likely due to inadequate larval habitat and perhaps decreased prey availability for these early life history stages.     

Mechanisms of marsh habitat alteration due to<Emphasis Type="Italic">Phragmites</Emphasis>: Response of young-of-the-year mummichog (<Emphasis Type="Italic">Fundulus heteroclitus</Emphasis>) to treatment for<Emphasis Type="Italic">Phragmites</Emphasis> removal

In recent decades, marshes naturally dominated bySpartina spp. have been replaced byPhragmites australis throughout the northeastern United States. We suggest that early in this invasion there was little effect on the fish fauna. As the invasion proceeds, the marsh surface habitat became more altered (i.e., elevated, flattened, reduced water-filled depressions, and reduced standing water), which resulted in a reduction of feeding, reproduction, and nursery function for fishes, especiallyFundulus spp. These potential changes in marsh habitat and function have resulted in numerous attempts to removePhragmites and restoreSpartina spp. To evaluate the response of marsh surface fishes toPhragmites treatment, we examined fish use in the brackish water reaches of Alloway Creek in the Delaware Bay estuary. ReferencePhragmites habitats were compared with referenceSpartina alterniflora-dominated habitats and sites treated (1996–1998) to removePhragmites to restore former vegetation (i.e., restored, now comprised of 100%Spartina). Fish were sampled with an array (n=9 at each site) of shallow pit traps (rectangular glass dishes, 27.5×17.5×3.7 cm). Small individuals (mean=17.5, 5–45 mm TL) dominated all pit trap collections. Fish abundance was highest at the restored (catch per unit effort [CPUE]=2.16) andSpartina (CPUE=0.81) sites with significantly lower values atPhragmites (CPUE=0.05) habitats. Samples were dominated by young-of-the-year mummichog,Fundulus heteroclitus (98% of total fish, n=631). The only other fish species collected was spotfin killifish,Fundulus luciae (2% of total catch, n=14), which was only present in restored andSpartina habitats. These observations suggest that the restored marsh is providing habitat (water-filled depressions on the marsh surface) for young-of-the-yearFundulus spp. These marshes are responding favorably to the restoration based on the much greater abundance of fish in restored versusPhragmites habitats and the overall similarity between restored andSpartina habitats.     

Tidal hydrology and habitat utilization by resident nekton in<Emphasis Type="Italic">Pragmites</Emphasis> and Non-<Emphasis Type="Italic">Phragmites</Emphasis> Marshes

We compared nekton densities over a range of measured flooding conditions and locations withinPhragmites australis andSpartina alterniflora (salt marsh cordgrass) at the Charles Wheeler Salt Marsh, located on the lower Housatonic River estuary in southwestern Connecticut. Nekton were sampled on nine spring high tide events from May to October 2000 using bottomless lift nets positioned between 0–5 and 10–20 m from the creek edge. Flooding depth, duration, and frequency were measured from each vegetation type during each sampling month. Benthic macroinvertebrate density was also measured within each vegetation type in May, July, and September. Frequency of flooding was 52% lower and flooding depth and duration were also significantly reduced inP. australis relative toS. alterniflora. A total of 4,197 individuals representing 7 species, mostlyPalaemonetes pugio (dagger-blade grass shrimp) andFundulus heteroclitus (common mummichog), were captured.P. pugio densities were significantly greater inS. alterniflora as were benthic macroinvertebrate density and taxa richness during May, but not during June or October. Total fish density was not significantly different betweenP. australis andS. alterniflora and was independent of location on the marsh. Significantly more juvenileF. heteroclitus were collected withinS. alterniflora relative toP. australis in June and July, suggesting that recruitment of this species may be lower inP. australis habitat. Fish density generally did not vary predictably across the range of flooding depth and duration; there was a positive relationship between flooding depth and fish density inS. alterniflora. The measured reduction in flooding frequency (52%) withinP. australis at the Housatonic site would result in an average total monthly fish use, expressed as density, of 447 ind m⁻² forP. australis and 947 ind m⁻² forS. alterniflora. WhenP. australis expansion results in reduction of flooding frequency and duration, nekton community composition can change, access to the marsh surface is reduced twofold, and nursery habitat function may be impaired.     

Diet and movement of the killifish,Fundulus heteroclitus, in a Maine salt marsh assessed using gut contents and stable isotope analyses

Killifish are ecologically important components of salt marsh ecosystems, but no studies have determined the importance of locally produced versus allochthonous food sources on a scale of less than multiple kilometers. The goal of our study was to examine diet and movement of the killifish,Fundulus heteroclitus, collected from a Maine salt marsh to assess the importance of locally produced versus allochthonous food sources on a scale of several hundred meters. We compared the gut contents and stable isotope signatures ofF. heteroclitus from four regions along the central river of a Maine salt marsh to the distinct food sources and isotopic signatures of the region of the marsh in which they were caught.F. heteroclitus were relying on locally produced food sources even on the scale of several hundred meters. They fed daily in a small area less than 6 ha and maintained relatively strong site fidelities over the course of several months. Phytoplankton and salt marsh detritus both contributed to the high production ofF. heteroclitus; terrestrial plant detritus was not an important component of their diet. The diet and feeding patterns ofF. heteroclitus from this small Maine salt marsh were similar to the patterns found in much larger salt marshes, suggesting that locally produced organic matter is essential to the production of these ecologically important fish.     

Effects of common reed (Phragmites australis) invasion on marsh surface macrofauna: Response of fishes and decapod crustaceans

The tidally inundated marsh surface is an importnat site for energy exchanges for many resident and transient species. In many areas along the East Coast of the U.S. the dominant vegetation,Spartina alterniflora, has been replaced by the common reed (Phragmites australis). This shift has caused concern about the impact ofPhragmites on marsh fauna but research in this area has been limited. During 1997 and 1998, we examined the effect ofPhragmites on fish and decapod crustacean use of the marsh surface in the brackish water reaches of the Mullica River, in southern New Jersey, U.S. Fish and decapod crustaceans were sampled with an array of shallow pit traps (rectangular glass dishes, 27.5×17.5×3.7 cm) and with flumes (1.3 m wide×10 m long of 3.2-mm mesh). Fish (2–60 mm TL) dominated pit trap collections withFundulus heteroclitus andFundulus luciae significantly more abundant atSpartina sites.Fundulus heteroclitus was also the dominant fish (15–275 mm TL) collected in flumes but collections with this gear, including a number of species not collected in pit traps, showed no distinct preferences for different marsh vegetation types. Decapod crustaceans (1–48 mm CW) collected in pit traps were generally less abundant than fishes withCallinectes sapidus andPalaemonetes spp. most abundant inSpartina, whileRhithropanopeus harrisii was most abundant inPhragmites. The same decapod crustacean species (2–186 mm CW) dominanted the flume collections and, similar to the pattern of fish collected by the flumes, there were no distinct habitat preferences for different marsh vegetation types. As a result of these observations, with different sampling techniques, it appears there is an overall negative effect ofPhragmites on larval and small juvenile fish but less or no effect on larger fish and decapods crustaceans.     

Tidal utilization of nekton in Delaware Bay restored and reference intertidal salt marsh creeks

Large-scale marsh restoration efforts were conducted to restore normal salt marsh structure and function to degraded marshes (i.e., former salt hay farms) in the mesohaline lower Delaware Bay. While nekton response has been previously evaluated for the marsh surface and subtidal creeks in these marshes, little effort has been focused on intertidal creeks. Nekton response in intertidal creeks was evaluated by sampling with seines to determine if restored (i.e., former salt hay farms restored in 1996) and reference (i.e., natural or relatively undisturbed) salt marshes were utilized by intertidal nekton in a similar manner. The overall nekton assemblage during June–October 2004–2005 was generally comprised of the same species in both the restored and reference marshes. Intertidal creek catches in both marsh types consisted primarily ofFundulus heteroclitus andMenidia menidia, with varying numbers of less abundant transient species present. Transient nekton were more abundant at restored marshes than reference marshes, but in insufficient numbers to cause differences in nekton assemblages. In both marsh types, low tide stages were characterized by resident nekton, dominated byF. heteroclitus, while high tide stages were characterized by a variable mix of transient and resident nekton. Assemblage level analyses indicated that intertidal creeks in restored and reference marshes were generally utilized in a similar manner by a similar nekton assemblage, so restoration efforts were deemed successful. This is in agreement with multiple comparative studies from the ame marshes examining fish, invertebrates, and vegetation in different marsh habitats.     

Responses of resident marsh fishes to stages ofPhragmites australis invasion in three mid Atlantic estuaries

Modification of brackish marshes by nonindigenousPhragmites australis has occurred across a broad geographical area in eastern North America. Among its effects on marsh processes,Phragmites may be increasingly unfavorable to marsh surface fishes as its invasion progresses within an estuary. We assessed the effect of thePhragmites invasion on resident marsh surface fishes by examining the population response ofFundulus heteroclitus (mummichog, 5–48 mm TL) andF. luciae (spotfin killifish, 5–41 mm TL) to four distinct invasion stages in three estuaries of the U.S. mid Atlantic region (New Jersey, Delaware, and Maryland). We documented precipitous declines in mean catch per unit effort ofF. heteroclitus in pit traps from natural marsh (51.6), through initial (33.8), early (12.3), and late invasion stages (2.4) across all sites. A similar pattern was documented forF. luciae, with mean catch per unit effort in pit traps declining from natural marsh (48.9), through initial (39.1), early (9.3), and late invasion stages (2.7). Population structure of both species also changed somewhat across invasion stages such that we collected a narrower size range of individuals of both species from late invasion stages. Patterns suggest that as thePhragmites invasion progresses, there is a decline in habitat function for larval and juvenileF. heteroclitus and an increased risk of extirpation ofF. luciae from brackish marshes along the east coast of the U.S.     

Comparison of fish and macroinvertebrate use of<Emphasis Type="Italic">Typha angustifolia,Phragmites australis</Emphasis>, and treated<Emphasis Type="Italic">Phragmites</Emphasis> marshes along the lower Connecticut River

Since 1965 large areas of lower Connecticut River tidelands have been converted from high diversity brackish meadow andTypha angustifolia marsh to near monocultures ofPhragmites australis. This study addresses the impact ofPhragmites invasion on fish and crustacean use of oligohaline high marsh. During spring tides from early June through early September 2000, fishes and crustaceans leaving flooded marsh along 3 km of the Lieutenant River, a lower Connecticut River tributary, were captured with Breder traps at 90 sites, equally distributed amongPhragmites, Typha, and treated (herbicide and mowing)Phragmites areas. Pit traps, 18 per vegetation type in 2000 and 30 each inPhragmites andTypha in 2001, caught larvae and juveniles at distances of up to 30 m into the marsh interior. There were no significant differences in fish species compositions or abundances among the vegetation types. Size distributions, size specific biomasses, and diets ofFundulus heteroclitus, the numerically dominant fish, were also similar. The shrimpPalaemonetes pugio was more abundant inPhragmites than in other types of vegetation, whereas the fiddler crabUca minax was least numerous inPhragmites. Mean numbers ofF. heteroclitus andP. pugio caught per site event were positively correlated with increasing site hydroperiod. Significantly moreF. heteroclitus were captured along the upper reach of the river where marsh elevations were lower than farther downstream. MoreF. heteroclitus and fewerP. pugio andU. minax were captured during the day than at night. A relatively small number of larval and juvenileFundulus sp. were captured in pit traps, but consistently fewer inPhragmites than inTypha, suggesting thatTypha and brackish meadow marshes may provide better nursery habitat. Vegetation was sampled along a 30 m transect at each trap site in 2000. Plant species diversity was greatest in treatedPhragmites areas and lowest inPhragmites sites.     

Rates, patterns, and impacts ofPhragmites australis expansion and effects of experimentalPhragmites control on vegetation, macroinvertebrates, and fish within tidelands of the lower Connecticut River

Phragmites expansion rates (linear at 1–3% yr⁻¹) and impacts of this expansion on high marsh macroinvertebrates, aboveground production, and litter decomposition fromPhragmites and other marsh graminoids were studied along a polyhaline to oligohaline gradient. These parameters, and fish use of creeks and high marsh, were also studied inPhragmites control sites (herbicide, mowing, and combined herbicide/mow treatments).Phragmites clones established without obvious site preferences on oligohaline marshes, expanding radially. At higher salinities,Phragmites preferentially colonized creekbank levees and disturbed upland borders, then expanded into the central marsh. Hydroperiods, but not salinities or water table, distinguishedPhragmites-dominated transects. Pooled samples ofPhragmites leaves, stems, and flowers decompose more slowly than other marsh angiosperms;Phragmites leaves alone decompose as or more rapidly than those of cattail. AbovegroundPhragmites production was 1,300 to 2,400 g m⁻² (about 23% of this as leaves), versus 600–800 g m⁻² for polyhaline to mesohaline meadow and 1,300 g m⁻² for oligohaline cattail-sedge marsh. Macroinvertebrates appear largely unaffected byPhragmites expansion or control efforts; distribution and densities are unrelated to elevation or hydroperiod, but densities are positively related to litter cover. Dominant fish captured leaving flooded marsh wereFundulus heteroclitus andAnguilla rostrata; both preyed heavily on marsh macroinvertebrates.A. rostrata andMorone americana tended to be more common inPhragmites, but otherwise there were no major differences in use patterns betweenPhragmites and brackish meadow vegetation. SAV and macroalgal cover were markedly lower within aPhragmites-dominated creek versus one withSpartina-dominated banks. The same fish species assemblage was trapped in both plus a third within the herbicide/mow treatment. Fish biomass was greatest from theSpartina creek and lowest from thePhragmites creek, reflecting abundances ofF. heteroclitus. Mowing depressedPhragmites aboveground production and increased stem density, but was ineffective for control.Phragmites, Spartina patens, andJuncus gerardii frequencies after herbicide-only treatment were 0.53-0.21; total live cover was <8% with a heavy litter and dense standing dead stems. After two growing seasonsAgrostis stolonifera/S. patens/J. gerardii brackish meadow characterized most of the herbicide/mow treatment area;Phragmites frequency here was 0.53, contributing 3% cover. Both values more than doubled after four years; a single treatment is ineffective for long-termPhragmites control.     

Impact of Armored Shorelines on Shore-Zone Fish Density in a Mid-Atlantic,USA, Estuary: Modulation by Hypoxia and Temperature

Anthropogenic modifications of estuarine environments, including shoreline hardening and corresponding alteration of water quality, are accelerating worldwide as human population increases in coastal regions. Estuarine fish species inhabiting temperate ecosystems are adapted to extreme variations in environmental conditions including water temperature, salinity, and dissolved oxygen across seasonal, daily, and hourly time scales. The present research utilized quantitative sampling to examine the spatiotemporal distribution of shore-zone estuarine fish species in association with four unique shoreline types across a range of water temperature and dissolved oxygen conditions. Fish were collected from the intertidal and shallow subtidal region of four shoreline types, Spartina alterniflora marsh, Phragmites australis marsh, riprap, and bulkhead, in the summer and fall of 2009 and 2010. Analyses were performed to (1) compare mean fish density among shoreline types across all water conditions and (2) explore relationships of the complete fish assemblage, three functional species groupings, and two fish species (Fundulus heteroclitus and Menidia menidia) to unique shoreline/water conditions. Significantly greater mean fish densities were found along S. alterniflora shorelines than armored shorelines. Several metrics including fish density, species richness, and occurrence rates suggest S. alterniflora shorelines may serve as a form of refuge habitat during periods of low dissolved oxygen and high temperatures for various species, particularly littoral-demersal species including F. heteroclitus. Potential mechanisms that could contribute to a habitat providing refuge during adverse water quality conditions include tempering of the adverse condition (decreased temperatures, increased dissolved oxygen), predation protection, and increased foraging opportunities.     

Growth and Movements of Mummichogs (<Emphasis Type="Italic">Fundulus heteroclitus</Emphasis>) Along Armored and Vegetated Estuarine Shorelines

Alteration of estuarine shorelines associated with increased urbanization can significantly impact biota and food webs. This study determined the impact of shoreline alteration on growth and movement of the estuarine fish Fundulus heteroclitus in a tributary of the Delaware Coastal Bays. Fundulus heteroclitus is abundant along the east coast of the USA, and is an important trophic link between marsh and subtidal estuary. The restricted home range of F. heteroclitus allowed discrete sampling, and fish growth comparisons, along 35–65-m long stretches of fringing Spartina alterniflora and Phragmites australis marsh, riprap, and bulkhead. Fundulus heteroclitus were tagged with decimal Coded Wire Tags. Of 725 tagged F. heteroclitus, 89 were recaptured 30–63 days later. Mean growth rate (0.06–0.15 mm day^?1 across all shoreline types) was greatest at riprap, lowest at Spartina and Phragmites, and intermediate at bulkhead, where growth was not significantly different from any other shoreline. This suggests that discernible environments exist along different shoreline types, even at the scale of tens of meters. No difference in movement distance was detected at different shoreline types; most individuals displayed a high degree of site fidelity. Forty-seven percent were recaptured within 5 m of their tagging location, although alongshore movements up to 475 m were recorded. Estimates of relative F. heteroclitus productivity, using relative density data from a concurrent study, were highest along Spartina and Phragmites, intermediate at riprap, and lowest at bulkhead. Therefore, despite greater growth rates along riprap than at vegetated shores, armoring reduces abundance sufficiently to negatively impact localized productivity of F. heteroclitus.     

Blue crab,<Emphasis Type="Italic">Callinectes sapidus</Emphasis>, response to the invasive common reed,<Emphasis Type="Italic">Phragmites australis</Emphasis>: Abundance,size, sex ratio,and molting frequency

Much effort has been directed recently at restoring marshes, by the removal of the invasive common reed,Phragmites australis, yet it is not clear how fish and invertebrates have responded either to the invasion ofPhragmites or to marsh restoration. The blue crab,Callinectes sapidus, uses marsh habitats during much of its benthic life. We investigated the response of blue crabs toPhragmites invasion and restoration efforts by comparing crab abundance (catch per unit effort), mean size and size frequency distribution, sex ratio, and molting of crabs in three physically similar areas differing in marsh vegetation;Spartina-dominated,Phragmites-dominated, and a treated area (Phragmites removed and now dominated bySpartina) in one marsh in the upper portion of Delaware Bay. Field sampling occurred monthly (April to November) from 1999 to 2001 using replicate daytime otter trawls in large marsh creeks. Crabs were categorized by carapace width into recruits (<30 mm), juveniles (30–115 mm), and adults (>115 mm). Juveniles dominated the system, representing 69.4% of all crabs. Similar monthly increases in mean size and molting patterns during the growing season (May–August) occurred inSpartina (natural and treated sites) andPhragmites sites suggesting that, subtidal habitats, used for molting, in these areas do not differ. More juveniles in the feeding molt stage (i.e., intermolt) than in other molt stages and more recruits predominantly in the feeding molt stage than adults were inSpartina, suggesting differences in the marsh surfaces used as feeding habitats withSpartina being preferred. Sex ratios of each life history stage were skewed towards males, but this was related to the low salinity of Alloway Creek, rather than marsh surface vegetation. Our results suggest that marsh surface vegetation influences the way blue crabs use marsh surface habitats, thus restoration efforts focusing on changing vegetation type may have a positive influence on blue crabs.     

Monitoring food web changes in tide-restored salt marshes: A carbon stable isotope approach

Primary producer (angiosperms, macroalgae, submerged aquatic vegetation), suspended particulate matter, andFundulus heteroclitus isotope values (δ¹³C, δ¹⁵N, δ³⁴S) were examined to assess their use as indicators for changes in food web support functions in tidally-restored salt marshes. Study sites, located throughout the southern New England region (USA), ranged fromSpartina alterniflora-dominated reference marshes, marshes under various regimes and histories of tide restoration, and a severely tide-restrictedPhragmites australis marsh.Fundulus δ¹³C values were greater for fish from referenceSpartina marshes than for fish from adjacent tide-restricted or tide-restored marshes where higher percent cover of C₃ plants, lower water column salinities, and more negative dissolved inorganic δ¹³C values were observed. The difference inFundulus δ¹³C values between a tide-restrictedPhragmites marsh and an adjacent referenceSpartina marsh was great compared to the difference between marshes at various stages of tide restoration and their respective reference marshes, suggesting that food web support functions are restored as the degree of tidal restriction is lessened. While a multiple isotopic approach can provide valuable information for determining specific food sources to consumers, this study demonstrates that monitoringFundulus δ¹³C values alone may be useful to evaluate the trajectory of ecological change for marshes undergoing tidal restoration.     

Effects of Life History Strategy on Fish Distribution and Use of Estuarine Salt Marsh and Shallow-Water Flat Habitats

To assess the potential for habitat isolation effects on estuarine nekton, we used two species with different dispersal abilities and life history strategies, mummichog (Fundulus heteroclitus) and pinfish (Lagodon rhomboides) to examine: (1) distribution trends among estuarine shallow-water flat and various intertidal salt marsh habitats and (2) the influence of salt marsh habitat size and isolation. Collections were conducted using baited minnow traps set within nonisolated interior marshes (interior), nonisolated fringing marshes (nonisolated), isolated island marshes (isolated), and shallow-water flat habitats (flat) that were adjacent to isolated and nonisolated marshes. Size range of individuals collected included juvenile and adult F. heteroclitus (20–82-mm standard length) and L. rhomboides (22–151-mm standard length). During high tide, F. heteroclitus exclusively used marsh habitats, particularly high marsh, whereas L. rhomboides used marshes and flats. F. heteroclitus abundance followed an interior > nonisolated > isolated pattern. L. rhomboides abundance patterns were less consistent but followed a nonisolated > isolated > interior pattern. A size-dependent water depth relationship was observed for both species and suggests size class partitioning of marsh and flat habitats during high tide. Minimum water depth (~31 cm) restricted L. rhomboides populations in marshes, while maximum water depth (~69 cm) restricted F. heteroclitus population use of marshes and movement between marsh habitats. Disparities in F. heteroclitus young of year contribution between isolated compared to nonisolated and interior marsh types suggests isolated marshes acted as population sinks and were dependent on adult emigrants. Resident and transient salt marsh nekton species utilize estuarine habitats in different ways and these fundamental differences can translate into how estuarine landscape might affect nekton.     

Salt-marsh tide pools as winter refuges for the mummichog,Fundulus heteroclitus,in New Jersey

Mummichog,Fundulus heteroclitus, were collected weekly from a southern New Jersey high-salinity salt marsh from October 1988 to June 1989 and from September 1989 to June 1990 to determine the overwintering habitat. Major habitat types sampled within the salt marsh were subtidal creek, intertidal creeks, and salt-marsh pools. Few individuals were collected in the intertidal creek or the subtidal creek from the end of October through the beginning of May for both years, when creek water temperatures were low. Both young-of-the-year and adults of both sexes were abundant in the salt-marsh pools (total lengths ranged from 29 mm to 125 mm) throughout the winter. In the spring, catch per unit effort (CPUE) within the tidal creek increased with increasing water temperature, while CPUE in marsh pools decreased with increases in estuarine water temperature. These collection patterns indicate that the majority ofF. heteroclitus may move from subtidal and intertidal creeks into salt-marsh pools in the late fall and leave in the spring. This seasonal movement could explain how fish survive winter environmental conditions because daily average water temperatures of salt-marsh pools were warmer than subtidal creek temperatures for much of the winter.     

Habitat use patterns of the mummichog,Fundulus heteroclitus, in New England. I. Intramarsh variation

Although studies of intertidal habitats have contributed greatly to the field of ecology, the processes governing the use of these areas by highly mobile animals such as fish remain poorly understood. In particular, although large-scale patterns of estuarine fish abundances are well known, fine-scale patterns of habitat use have been largely overlooked. Here, I examine among and within habitat use patterns of the mummichog,Fundulus heteroclitus, in a New England salt marsh. Using minnow traps I sampled changes in mummichog habitat use among closely spaced sites within creeks, mudflat, and channel over 2 yr. The general pattern of mummichog captures was consistent among years, showing peaks in summer months and lows in winter. Use of specific habitat types was also seasonally dependent. For most of the year (fall, winter, spring) mudflat capture rates were lower than creek capture rates in the summer; however, densities in the mudflat equal or exceed those in creeks. In most months channel habitats did not differ significantly in use from either creek or mudflat habitat. Consistent patterns also occur within induividual marsh habitats.F. heteroclitus concentrate in the uppermost portion of creeks. Additionally, mudflat capture rates (August 1990) declined with increasing water depth. Diurnal habitat use is heavier than nocturnal use for all habitats. These findings demonstrate that mummichogs are restricted to areas representing a very small proportion of total available habitat and that their patterns of habitat use are strongly seasonal. Only careful experimental work can determine the relative advantages of these habitats for estuarine fish. Further comparative work is required to reveal how biological and physical parameters affecting habitat use may vary geographically.     

Life history strategies of estuarine nekton: The role of marsh macrophytes,benthic microalgae,and phytoplankton in the trophic spectrum

The stable isotope signatures of marine transient and resident nekton were used to investigate trophic linkages between primary producers, marsh macrophytes, phytoplankton, benthic microalgae, and consumers within the Delaware Bay. A whole estuary approach was used to compare the flux of nutrients from primary producers to juvenile weakfish (Cynoscion regalis), bay anchovy (Anchoa mitchilli), and white perch (Morone americana) in open waters of the lower and upper Bay and adjacent salt marshes dominated by eitherSpartina alterniflora orPhragmites australis. Our results suggest that trophic linkages vary significantly along the salinity gradient, reflecting the transition fromSpartina toPhragmites-dominated marshes, and secondarily, in a marsh to open water (offshore) direction at a given salinity. Superimposed on this pattern was a gradient in the proximate use of organic matter that depended on life history traits of each species ranging from pelagic to benthic in the order bay anchovy > weakfish > white perch.     

Habitat use and movement of the mummichog (<Emphasis Type="Italic">Fundulus heteroclitus</Emphasis>) in a restored salt marsh

The mummichog,Fundulus heteroclitus, is one of the most abundant macrofaunal components of salt marsh ecosystems along the east coast of the United States. During April–November 1998, we determined the habitat use and movement patterns of young-of-the-year (YOY) and adult mummichogs in a restored marsh, formerly a salt hay farm, and an adjacent creek in order to expand our understanding of the ecology of the species and evaluate the success of the restoration. Four major fish habitat types (large first-order natural creek, second-order created creek, linear drainage ditch, and marsh surface) were identified within the study site. Patterns of relative abundance and mark and recapture using coded wire tags were used to determine the habitat use, tidal movements, home range, and site fidelity of the species within these habitat types. A total of 14,784 fish, ranging from 20–100 mm SL, were captured with wire mesh traps and tagged, and 1,521 (10.3%) fish were recaptured. A variety of gears were used to attempt to recapture fish across all habitat types, including wire mesh traps, push nets, and otter trawls. Based on abundance and recaptures of tagged fish, the YOY and adults primarily used the shallow subtidal and intertidal areas of the created creek, the intertidal drainage ditches, and the marsh surface of the restored marsh but not the larger, first-order natural creek. At low tide, large numbers were found in the subtidal areas of the created creek; these then moved onto the marsh surface on the flooding tide. Elevation, and thus hydroperiod, appeared to influence the microscale use of the marsh surface. We estimated the home range of adults and large YOY (20–100 mm SL) to be 15 ha at high tide, which was much larger than previously quantified. There was strong site fidelity to the created creek at low tide. The habitat use and movement patterns of the mummichog appeared similar to that reported for natural marshes. Coupled with the results of other studies on the feeding, growth, and production of this species in this restoreh, the species appeared to have responded well to the restoration.