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Studies in Avian Biology No. 32:155–175

IMPACTS OF MARSH MANAGEMENT ON COASTAL-MARSH HABITATS

LAURA R. MITCHELL, STEVEN GABREY, PETER P. MARRA, AND R. MICHAEL ERWIN

Abstract. The effects of habitat-management practices in coastal marshes have been poorly evaluated. We summarize the extant literature concerning whether these manipulations achieve their goals and the effects of these manipulations on target (i.e., waterfowl and waterfowl food ) and non-target organisms (particularly coastal-marsh endemics). Although we focus on the effects of marsh manage- ment on , we also summarize the scant literature concerning the impacts of marsh manipulations on wildlife such as small mammals and . We address three common forms of anthro- pogenic marsh disturbance: prescribed fi re, structural marsh management, and open-marsh water management. We also address marsh perturbations by native and introduced vertebrates.

Key Words: Disturbance, impoundment, marsh endemic, marsh management, mosquito control, open- marsh water management, prescribed fi re, structural marsh management.

IMPACTOS DEL MANEJO DE MARISMA EN HABITATS DE AVES DE COSTA-MARISMA Resumen. Los efectos por las prácticas de manejo del hábitat en marismas de costa han sido pobre- mente evaluados. Resumimos la literatura existente que concierne a que ya sea si estas manipula- ciones alcanzan sus metas, y los efectos de estas manipulaciones en organismos blanco (ej. Gallinas de agua y plantas de alimento de gallinas de agua) y en organismos no-blanco (particularmente en endémicos de marisma de costa). A pesar de que nos enfocamos en los efectos del manejo de marisma en aves, también resumimos la escasa literatura que concierne a los impactos de la manipulación de marisma en la vida silvestre, tales como pequeños mamíferos e invertebrados. Dirigimos tres for- mas comunes de perturbación antropogénica de marisma: quemas prescritas, manejo estructural de marisma, y manejo de agua en marisma-abierto. También dirigimos perturbaciones del marisma por vertebrados nativos e introducidos.

Nearly three-quarters of the 2,500,000 ha of Much less is known about the impacts of marsh coastal marshes of the US are located along the manipulations on small mammals, , southeast Atlantic (North Carolina–Florida) and and invertebrates. In this review, we address northern Gulf of (Florida–Texas) coasts three common forms of anthropogenic marsh (Alexander et al. 1986, Chabreck 1988). The exten- disturbance—prescribed fi re, structural marsh sive gulf and Atlantic marshes of the Southeast management, open-marsh water management. are intensely managed by federal and state land Additionally, we address marsh perturbations management agencies, conservation organiza- by native and introduced vertebrates. Several tions, and private landowners. Managers dis- other marsh-management actions we consid- turb these ecosystems, often yearly, through ered beyond the scope of this review including: prescribed burns, herbicide applications, ditch- insecticides targeting mosquitoes, herbicides ing, and shallow pond construction. The ratio- for exotic or control, salt-hay nale for these manipulations fall broadly under cropping, and cattle grazing. These processes, categories of: (1) wildlife enhancement, (2) fl ood particularly insecticide and herbicide use, often control, (3) mosquito control, and (4) erosion create impacts on nearly all of our marsh lands, mitigation (Daiber 1987, Chabreck 1988, Nyman but have been so little studied in the wildlife and Chabreck 1995). These manipulations have arena that little can be concluded, despite their occurred since historical times, but have been clearly major impacts. poorly, or only recently, evaluated in terms of their impact on wildlife. EFFECTS OF PRESCRIBED FIRE ON In this chapter we address two primary ques- COASTAL-MARSH BIRDS tions: (1) are these manipulations achieving their wildlife management goals, and (2) what are Prescribed fi re is widely used to manipulate the effects of these manipulations on non-target marsh vegetation. Although prescribed fi re tra- organisms, particularly coastal-marsh endem- ditionally is used in gulf and southeast Atlantic ics? The majority of available data focuses on Coast marshes, its application has spread the effects of marsh management on birds. throughout much of the eastern seaboard. For

155 156 STUDIES IN AVIAN BIOLOGY NO. 32 example, in 2002, the USDI and Wildlife wetlands remains weak; few analytical papers Service (USFWS), National Wildlife Refuge have documented fi re’s effects on marsh wild- System burned approximately 9,500 ha of life. Many coastal researchers consider marsh coastal marsh along the Texas coast; 9,300 ha of burning to be simply a “long-standing cultural tidal and freshwater marshes along the North practice…apparently done because of tradition Carolina–Florida coast; 4,000 ha of coastal or with poorly planned objectives” (Nyman marsh in Louisiana; and 2,000 ha of salt and and Chabreck 1995:134). We summarize extant brackish tidal marshes in the Chesapeake Bay information on the effi cacy of prescribed burn- watershed (Dave Brownlie, USFWS Region 4, ing on improving waterfowl habitat and popu- pers. comm.; Mark Kaib, USFWS Region 2 pers. lations and examine possible indirect effects of comm.; Roger Boykin, USFWS Region 4 pers. burning on non-target marsh birds. comm.; Allen Carter, USFWS Region 5, pers. comm.). These fi gures exclude widespread EFFECTS OF PRESCRIBED BURNING ON WATERFOWL prescribed burning by state agencies, National AND THEIR HABITAT Park Service, or private individuals. Prescribed fi re in marshes gained initial Lynch (1941) advocated prescribed fi res support as a management tool for improv- to enhance waterfowl wintering habitat. He ing wintering waterfowl habitat in the 1930s reported that experimental burning on federal and 1940s along the Gulf Coast (Lynch 1941, refuges in Louisiana attracted >500,000 geese Nyman and Chabreck 1995). In the following and thousands of ducks to marshes that had decades, wildlife managers from the East and previously held few waterfowl. Lynch (1941) Gulf coasts embraced the recommendations of states that these burns removed dense veg- authors who advocated prescribed burning as etation that interfered with growth of preferred a tool for conditioning marsh habitats (Lynch waterfowl foods, increased nutritional quality 1941, Hoffpauir 1961, Givens 1962, Hoffpauir of forage for cattle and geese, and increased 1968, Perkins 1968). Such conditioning includes waterfowl access to seeds and rhizomes. No removal of litter and dead vegetation, or veg- details regarding counting methods, use of etation considered to be of little or no value control marshes, or historical occurrence of to gamebirds (e.g. cattails [Typha spp.] and fi res or waterfowl in that area were provided. cordgrasses [ spp.]), and reduction of Interestingly, Lynch (1941) recommended that shrub cover. These burns were also purported prescribed fi res be used only on Gulf Coast to stimulate growth or seed production of food marshes until experiments had been conducted plants eaten by waterfowl such as bulrushes in other coastal regions. (Schoenoplectus spp. formerly Scirpus spp.), For 20 yr after Lynch, burning in coastal bristle grasses (Setaria spp.) and Echinichloa marshes continued without critical evaluation. spp. Other purported benefi ts of marsh burn- In the 1960s, several authors began assess- ing include: (1) maintaining a mixture of ing marsh-burning effi cacy, mostly based on open-water and vegetated cover for resting, observational and anecdotal data (Givens 1962, loafi ng, and breeding activities by waterfowl Hoffpauir 1968, Perkins 1968). Those authors and other water birds, (2) facilitating trapping focused on benefi ts to waterfowl by the favor- (primarily for muskrats [Ondatra zibethicus] and ing of preferred forage plants and maintaining American alligators [Alligator mississippiensis]), shrub-free and otherwise open-marsh habitat. (3) recycling dead material and increasing As an example of the anecdotal nature of primary productivity through nutrient release, the evidence presented, Hoffpauir (1968:135) in and (4) reducing the risk of unpredictable or coastal Louisiana, noted cover or wet burning uncontrollable fi res, or fi res that would dam- 2–3 wk prior to arrival of Snow Geese (Chen age the marsh system, e.g., peat fi res (Nyman caerulescens) provided fresh green vegetation and Chabreck 1995, Foote 1996). Despite the and increased access to below-ground vegeta- long history of fi re, and ongoing federal expen- tion; however, geese used these areas for only ditures for prescribed fi re programs, critical 3–4 wk. Certain dabbling ducks appeared to evaluations of the effects of fi re on target and use the burned areas extensively, feeding in non-target species have been scarce. potholes left behind by the activity of the Snow In reviewing the available literature on fi re Geese. However, no information regarding use in North American wetland ecosystems in 1988, of controls, numbers of burned areas, or quan- Kirby et al. (1988:iii) declared that the science titative data were provided. of using fi re in natural and anthropogenic Despite these claims concerning waterfowl wetlands to perpetuate wildlife and plant com- habitat improvement, we found only one study munities was still in its infancy. Over a decade in which investigators surveyed waterfowl later, the predictive science of prescribed fi re in response to prescribed burns in coastal-marsh IMPACTS OF MARSH MANAGEMENT—Mitchell et al. 157 habitats using a standard methodology and intermediate marshes dominated by salt comparing burned areas to controls (Gabrey et meadow cordgrass. Burns increased total live al. 1999). The authors conducted aerial surveys above-ground biomass but failed to increase from December–February immediately follow- bulrush species. Post-burn, species composition ing 14 prescribed burns on a 30,700 ha state did not change, and post-burn fl owering and wildlife refuge in coastal Louisiana. Gabrey et seed production were nearly nonexistent, there- al. (1999) reported that 10 fl ocks of white geese fore, post-burn growth appeared to be from (Snow Goose and Ross’s Goose [Chen rossii]), below ground rhizomes and roots of the burned ranging in size between 300 and 17,500 individ- plants (Gabrey and Afton 2001; S. W. Gabrey, uals, used recently burned marsh areas exclu- pers. obs). Smooth cordgrass biomass in burned sively during the December–February period. plots was lower compared to unburned plots; However, the authors collected no behavioral or burning had no effect on inland saltgrass bio- dietary data to assess goose activity or possible mass (Gabrey and Afton 2001). The most nota- goose attractants in burned areas. ble and longest lasting effect of these burns was Habitat enhancement burns are intended to the dramatic reduction in dead above ground increase biomass and seed production of marsh biomass, which remained below unburned lev- plants preferred by migrating or wintering els for at least 3 yr post-burn. waterfowl (Lynch 1941), while reducing compe- Flores and Bounds (2001) studied six rep- tition from less preferred plants such as inland licate marsh sites in the Chesapeake Bay of saltgrass (Distichlis spicata), smooth cordgrass Maryland. All plots were burned in winter (Spartina alternifl ora), or salt meadow cordgrass 1998 and treatment plots were burned again (S. patens) (Lynch 1941, Nyman and Chabreck in winter 1999. Vegetation samples collected in 1995). DeSzalay and Resh (1997) evaluated late the fall of 1999 (following treatments) showed summer burns in inland saltgrass dominated that live above-ground biomass of inland salt- coastal marshes in and found that grass, chairmaker’s bulrush, and saltmeadow percent cover of inland saltgrass was reduced, cordgrass was greater in sites burned in 1999 while that of goosefoots (Chenopodium spp.) and than in those left unburned. Total biomass did purslanes (Sesuvium spp.) important in dab- not differ between treatments. Sites burned bling duck diets was increased, in burn treat- in 1999 had signifi cantly higher mean stem ments versus controls. densities than those left unburned. At 6 mo In brackish marshes in Chesapeake Bay, win- post-burn no signifi cant difference was found ter burning increased culm density and above in live aboveground biomass of black needle- ground biomass of live chairmaker’s bulrush rush (Juncus roemerianus) or smooth cordgrass (Schoenoplectus americanus) in burned plots 1 yr between burned and unburned treatments. The post-fi re, compared to plots that had not been researchers report an overall increase in plant burned for 2–3 yrs (Pendleton and Stevenson community stem density, but lack of increase in 1983, Stevenson et al. 2001). Biomass of dead overall plant community biomass, in response bulrush was greater in unburned plots than to burning. Although burning increased bio- burned plots. Burning did not affect biomass mass of bulrush, it did not reduce biomass of of plants other than bulrush. Pendleton and either cordgrass or saltgrass. Stevenson (1983) concluded that the greater Some researchers report that burning bulruish biomass produced following burning coastal marshes enhances primary productiv- was a consequence of increased stem density ity. Spring, summer, and winter burns in Texas rather than increased biomass of individual each increased live gulf cordgrass (Spartina stems. Standing-dead material limited the total spartinae) standing crop and the percentage of number of living culms in the unburned stands, fl owering plants by the end of the fi rst grow- and shaded new culms, therefore delaying the ing season post-burn. The greatest growth onset of spring growth. response resulted from spring treatment, pos- Turner (1987) found that late-winter burn- sibly because of post-burn rainfall (McAtee et ing in smooth cordgrass marshes in Georgia al. 1979). Winter cover burns on the Mississippi reduced net aboveground primary production coast increased net primary production (NPP) by 35%. Burning signifi cantly reduced mean of above ground plant material by 56% and 49% dry biomass of live rhizomes of smooth cord- in black needlerush and big cordgrass (Spartina grass in the top 10 cm of sediment. Burned cynosuroides) marsh communities, respectively plots exhibited a denser growth of smaller, fi ner (Hackney and de la Cruz 1981). smooth cordgrass plants than control plots. Season of burn and frequency of burn may Gabrey and Afton (2001) evaluated the effects explain in part the variability of vegetation of winter burning in 14 burned/unburned response. In a greenhouse study using small plot pairs in Louisiana saline, brackish, and buckets to simulate marshes, Chabreck (1981) 158 STUDIES IN AVIAN BIOLOGY NO. 32 showed that varying the season of burn altered 1981, DeSzalay and Resh 1997). Komarek (1984: the post-fi re plant community. October burns 6) reported that following a single prescribed appeared the most successful at promoting burn during the winter in a Juncus-Spartina the growth of bulrush species. while burns marsh at St. George Island, Florida, three spe- between December and February promoted salt cies of snail appeared to be more abundant in meadow cordgrass growth. In addition, O’Neil the burned section of the marsh. He observed (1949) recommended 3–4 yr of repeated burn- higher populations of fi ddler (Uca ing followed by periodic burning at 4-yr inter- spp.) in burned coastal marshes compared to vals to convert salt meadow cordgrass-inland unburned areas. However, because no data saltgrass dominated marsh to sturdy bulrush were provided to support these comments, it (Schoenoplectus robustus)-saltmeadow cordgrass is unclear if such reported increases actually marsh in Louisiana. However, numerous other refl ect increased abundance, per- environmental variables, such as air or water haps in response to greater nutrient availabil- temperature, salinity, pre-fi re vegetation com- ity, or greater invertebrate visibility in burned munity, likely infl uence the composition of the areas versus unburned sites. post-burn plant community. A few studies demonstrate that various Another popular objective of habitat- invertebrate taxa may respond differently to fi re. enhancement burns in coastal marshes is to On marsh islands in Virginia, Matta and Clouse increase the nutritive quality of available (1972) collected invertebrates in sweep nets at 2- plant foods. McAtee et al. (1979) report that wk post-burn intervals from six sites representa- digestible energy and crude protein content tive of four burn treatments. The occurrence of of gulf cordgrass was signifi cantly increased most adult forms was not signifi cantly affected on Texas coastal prairie in response to burn- by burning, although the principal insect herbi- ing. Smith et al. (1984) conducted fall burning vore, a meadow katydid (Conocephalus sp.) did in a Utah alkali marsh; protein increased in show signifi cant differences among sites, with inland saltgrass, tule bulrush (Schoenoplectus fewer numbers at recently burned sites. Turner acutus), and cattail (Typha spp.), but not in alkali (1987) found that abundance of the periwinkle bulrush (Bolboschoenus maritimus). Schmalzer snail (Littorarea irrorata), an important winter and Hinkle (1993) evaluated black needlerush food for American Black Ducks (Anas rubripes), and sand cordgrass (Spartina bakeri) marshes was reduced by burning in smooth cordgrass in burned in December, at Merritt Island, Florida, Georgia marshes. In the most extensive study and found that plant-tissue nutrient concentra- to date, DeSzalay and Resh (1997) found den- tions generally declined post-fi re. One year sities of many invertebrates important in the after burning, nitrogen (N) content in live diets of dabbling ducks in California wetlands vegetation was lower than pre-burn content for (for example, Chironomus spp. and Trichocorixa. all plant species. Phosphorous (P) concentra- spp.) to be greater in burned compared to tions increased in sand cordgrass, decreased unburned control marshes. However, densi- in bulltongue arrowhead (Sagittaria lancifolia) ties of other invertebrates, such as copepods and black needlerush in the black needlerush and oligochaetes, were lower in open sections marsh, and were unchanged in other species. of burned marshes compared to unburned However, the P:N ratio increased in all live bio- marshes. The researchers attributed lower den- mass types. Potassium (K) concentrations in live sities and biomass of these invertebrates in burn tissues declined or did not change signifi cantly areas to mortality due to vegetation removal, in all species whereas calcium (Ca) concentra- desiccation, or elevated soil temperatures. tions increased in black needlerush and sand The existing evidence supports the long- cordgrass. Magnesium (Mg) concentrations standing assumption that winter burning in decreased in live and dead black needlerush coastal marshes does attract waterfowl; the but increased in live bulltongue arrowhead and evidence is strongest for geese. However, the cordgrass species. Overall, biomass and nutri- mechanism for the attraction and the benefi ts ent content in these marshes did not return to accrued to the waterfowl populations remain pre-burn levels at 1 yr post-burn. unclear. Winter burning removes undesirable A potentially important, but poorly studied plants species and promotes growth of pre- effect of prescribed fi re is the possible impact ferred waterfowl plant foods under some con- on coastal-marsh invertebrates important in ditions (O’Neil 1949, Chabreck 1981, Pendleton waterfowl diets. Some researchers have specu- and Stevenson 1983, Turner 1987, DeSzalay and lated that burning may reduce invertebrate Resh 1997, Stevenson et al. 2001) but not oth- populations in the short term by altering marsh ers (Flores and Bounds 2001, Gabrey and Afton surface temperature or exposing to 2001). Effects of burning on the nutritional greater predation risk (Hackney and de la Cruz quality of marsh vegetation appear ambiguous IMPACTS OF MARSH MANAGEMENT—Mitchell et al. 159

(McAtee et al. 1979, Schmalzer and Hinkle the fi rst year post-burn, but increased 3–4 yr 1993). The scant studies on marsh invertebrate post-burn. At one location, breeding densities community response are also inconclusive declined during the fi fth breeding season, post- (Matta and Close 1972, Turner 1987, DeSzalay burn. Werner (1975) suggests that sparrows and Resh 1997). Plant and invertebrate com- decline in numbers immediately post-burn, then munity changes to burning are variable and increase in density 3–4 yr after a fi re, and may likely depend on environmental factors such as abandon a site after the sixth year after a fi re as season of burn, fi re intensity, water depth and vegetation density increases. He speculates that salinity, and post-burn rainfall. Although stud- optimum sparrow habitat could be maintained ies of vegetative productivity, plant nutritional if marshes are burned every 4–5 yr. Werner quality, and invertebrate abundance are impor- (1975) based his conclusions on a very small tant, it is also necessary to determine if such sample size without control sites. Of additional changes indicate a change in habitat quality interest in this study was the direct observation and benefi t the organisms such as birds that for- of individually marked sparrows fl eeing the age on the vegetation or invertebrates. Habitat fl aming front of a winter wildfi re into adjacent quality might be assessed by quantifying in unburned areas and fl ying in circles in areas of these improved areas the: (1) activity or energy smoke and fl ames. Although sparrow density in budgets, (2) foraging effort and behavior, (3) the burned area returned to pre-burn levels by physiological indices such as suffi cient energy the next breeding season, none of the marked stores for migration or breeding activities, (4) sparrows returned. or movement among burned and unburned Taylor (1983) censused Cape Sable Seaside patches. We are unaware of rigorous fi re stud- Sparrows at Taylor Slough, Florida. The study ies that have been designed to answer these design was a randomized-block design, with questions. three different prescribed burn treatments (annual, 3-yr rotation, 5-yr rotation) allocated EFFECTS OF PRESCRIBED BURNING ON OTHER to a set of three, 20-ha plots; a set of plots was MARSH BIRDS located at each of three different marsh loca- tions (blocks). In addition, a single control site Given the lack of information on waterfowl had not been burned for 10 yr, and was not response to burning in coastal marshes, it is not burned during the study. However, the season surprising that few quantitative studies address of prescribed fi res differed between blocks, the effects of fi re on other (non-game) birds e.g., burns at one marsh were applied only (Rotenberry et al. 1995). Herein we summarize in December (annually, every 3 yr, and every the few quantitative studies and include results 5 yr), while at another marsh area, all burns of qualitative observational work on the effects were conducted in July. Therefore, the study of coastal-marsh fi re on breeding and wintering had no true replication of treatments. coastal-marsh birds, including and Taylor (1983) reported that on burned sites raptors. with deeper soils (>20 cm), vegetation recov- ery was more rapid and sparrow populations Emberizidae (sparrows) recovered and peaked earlier than on sites with shallower soils. The former populations re-colo- The Cape Sable Seaside Sparrow nized rapidly and began to decline 4 yr post- (Ammodramus maritimus mirabilis) is an endan- fi re. In burned sites with shallow soils, plant gered whose relationship to fi re has biomass recovery was slower and sparrows did come under scrutiny due to recent population not even re-colonize these areas until about 4 yr declines. Although now restricted to inland sub- post-fi re and densities remained low for up to tropical marshes and seasonally fl ooded prairies 10 yr. In addition, post-fi re breeding territories of southern Florida (Werner 1975), this subspe- were clumped, presumably because birds were cies of a primarily coastal-marsh species is one forced to use marginal areas following large of the best researched passerines with respect to fi res (Taylor 1983). Fires created long edges in the effects of habitat burning, so we will review which birds concentrated during the fi rst breed- these studies in some detail. Werner (1975) ing season post-fi re and created a mosaic of tracked sparrow populations in two locations unburned patches in which birds nested. Taylor at Everglades National Park, Florida, for which (1983) concluded that fi re regimes shorter than historical fi re data indicated that these areas had 8–10 yr could be detrimental to Cape Sable experienced wildfi res in 1969 and 1972. A fi re in Seaside Sparrow populations. 1974 also burned one of the two locations. The Curnutt et al. (1998) provides the most com- author reports that at each of these sites breed- prehensive analysis of fi re’s effects on Cape ing densities of sparrows were sparse during Sable Seaside Sparrows. The authors analyzed 160 STUDIES IN AVIAN BIOLOGY NO. 32

227 sites (as surveyed on a 1-km grid within demonstrated by direct evidence of immedi- Everglades National Park) that contained Cape ate, negative effects of burning on sparrow Sable Seaside Sparrows between 1992–1996, and populations, and the reported role of fi re in for which dates and spatial extent of fi res from periodically maintaining open habitats attrac- 1982–1996 were known. Sites had experienced tive to the sparrows (Werner 1975, Taylor 1983, fi res caused by lightning strikes, unplanned Werner and Woolfenden 1983, Curnutt et al. human ignitions, or prescribed fi re activities. 1998). Walters et al. (2000) analyzed population The analysis did not control for likely differ- census data in Everglades National Park from ences beside fi re frequency and time since last 1981–1998 and concluded that two northeastern fi re between sites (Walters et al. 2000). populations appear to have declined due to For each site and for each sparrow survey, abnormally high fi re frequencies, and that dry- Curnutt et al. (1998) determined the frequency season fi res pose greater threats to breeding of fi res, the number of days since the most birds than wet-season fi res. They cite evidence recent fi re, and whether the most recent fi re that increased fi re frequency has been a direct occurred during the wet (1 June–31 October) result of anthropogenic water diversions, sub- or dry (1 November–May) season. They found sequent reduced hydroperiods, and exposure that sparrow densities were lowest at sites that to human-caused dry-season fi re. The authors had a dry-season fi re as their most recent fi re speculate that occasional fi re is necessary for occurrence. In contrast to Werner (1975) and continued occupancy of a marsh by Cape Sable Taylor (1983), Curnutt et al. (1998) found no Seaside Sparrows because it inhibits invasion evidence that sparrows abandon a site imme- by woody shrubs, including non-natives such diately post-burn and suggest that sparrows as paper barked tea tree (Melaleuca quinque- are able to occupy marsh sites immediately nervia; Curnutt et al. 1998), which can eliminate following a burn due to the patchy nature sparrow nesting habitat. Finally, Walters et al. of natural fi re in the Everglades. Curnutt et (2000) stress the need to incorporate prescribed al. (1998) also found that sparrow popula- burning into rigorous experimental studies, tions increase in density with no evidence of including studies of the dispersal patterns of eventual declines for up to 10 yr following a the birds through telemetry, to determine the fi re event. For those sites that held sparrows direct effects of fi re frequency on habitat and over the entire period of record, fi re frequency sparrow populations. ranged from one–seven fi res/10 yr, with a Effects of fi re on other coastal sparrow mean of 2.97 fi res/10 yr. Sparrow densities populations, such as the Louisiana Seaside were highest where there had been one–two Sparrow (Ammodramus maritimus fi sheri) and fi res over the previous 10-yr period, lower Nelson’s Sharp-tailed Sparrow (Ammodramus where fi re frequencies were greater or equal to nelsoni), have received recent attention (Gabrey three–six fi res/10 yr, and absent from sites that et al 1999, Gabrey and Afton 2000, Gabrey et were burned more than seven times in 10 yr. al. 2001). Gabrey et al. (1999) surveyed bird- The authors’ fi ndings support those of Taylor species composition and abundance and veg- (1983)—sparrows will use sites that had burned etation structure on 14 pairs of winter-burned 10–12 yr previously—and contradict Werner’s and unburned marshes in Louisiana. Winter (1975) suggestions that sparrows decline in bird surveys were conducted immediately fol- numbers and will abandon a site after the sixth lowing burns and again one full year post-fi re. year after a fi re. The primary conclusion from Immediately following burn treatment, plant this study is that frequent fi res are harmful to community visual obstruction and percent Cape Sable Seaside Sparrows and may be the cover were lower in burned plots; at 1 yr post- cause of declines in populations on the north- burn, vegetation structure was similar between eastern edge of the sparrow’s range. Curnutt treatment and control plots. Wintering Seaside et al. (1998) also suggested that that the artifi - Sparrows were absent immediately following cially drained nature of the coastal prairies in burns, but were present in unburned marshes; this region increased the fl ammability of these Seaside Sparrows were present in burn-treat- habitats, amplifying negative effects on Cape ment plots 1 yr post-burn. Nelson’s Sharp-tailed Sable Seaside Sparrows. Sparrows, a migratory species that winters In summarizing past studies on the Cape exclusively in coastal marshes (Greenlaw and Sable Seaside Sparrow, Walters et al. (2000: Rising 1994), were present in burned marshes 1104) state that catastrophic sparrow population during the fi rst winter but only in scattered declines in the past decade cannot be directly patches of unburned vegetation; however, they attributed to fi re. Nevertheless, authors of that were recorded frequently in unburned plots paper concluded that fi re has affected sparrow during both survey periods and in burn treat- populations by altering habitat suitability, as ment plots 1 yr post-burn. The authors conclude IMPACTS OF MARSH MANAGEMENT—Mitchell et al. 161 that winter burning reduces the suitability of population density, interfering with pair bonds, the marsh as winter habitat for these marsh- and delaying territory establishment and nest- dependent sparrows, but only for a few months ing activities. immediately following the burn. In other fi re studies in the Chenier Coastal Most studies of fi re effects on birds rely on Plain in Louisiana, Gabrey et al. (2001) found relative abundance as the response variable; that total abundance of sparrows (primar- rarely are demographic parameters such as ily Seaside Sparrows) did not differ between nest success or survival addressed. Gabrey et burned and unburned marshes during the al. (2002) used artifi cial nests to investigate fi rst or third summers, post-burn, but were effects of winter marsh burning on nest suc- two times greater in burned than unburned cess of two coastal-marsh endemics—Louisiana marshes during the second summer post-burn. Seaside Sparrows and Mottled Ducks (Anas The peak in sparrow abundance coincided with fuligula). They recorded apparent nest success the recovery of dead vegetation cover to pre- of nests containing quail eggs (to simulate spar- burn levels. Gabrey et al. (2001) concluded from row nests) and chicken eggs (to simulate duck both wintering and breeding season studies that nests) in four pairs of burned and unburned periodic but infrequent fi res that remove dense, marshes, during the breeding season prior to dead vegetation benefi t sparrow populations on and following experimental burns. They found the Chenier Coastal Plain. no difference in vegetation structure or success Baker (1973) reported that two wildfi res in of either type of artifi cial nest in the post-burn December 1972 and January 1973 burned about breeding season. Although no effect of winter 690 ha at St. Johns National Wildlife Refuge in burns on artifi cial-nest success was detected, Florida, leaving few patches of unburned sand the authors caution that their study involved cordgrass. Immediately following these fi res, only four marshes and that the timing of burn- color-banded individuals of the now-extinct ing may affect success of those birds that nest Dusky Seaside Sparrows (Ammodramus mari- early in the season before suffi cient vegetation timus nigrescens) were displaced from burned re-growth has occurred (Gabrey et al. 2002). areas. In early May, however, banded males Gabrey and Afton (2000) examined effects reappeared and defended territories in burned of winter marsh burning on Louisiana Seaside areas. Baker (1973) speculates that rather than Sparrows nesting activity. Measurements were occupying small, unburned cordgrass patches made during the breeding season (April–July) within burn areas, the birds moved to nearby, prior to experimental burns and during two unburned cover. Three birds banded on the area breeding seasons post-burn. Male sparrows prior to burns were recaptured immediately were absent from burned marshes during the after the burn in unburned cover, 900 m from start of the fi rst breeding season after burns, but their original locations. had reached abundances comparable to control Walters (1992) reported that in 1975, a fi re marshes by June of that season. During the intentionally set on private land escaped con- second breeding season post-burn, numbers of trol lines and burned nearly 850 ha of Dusky male sparrows were greater in burned marshes Seaside Sparrow habitat on the St. Johns than in unburned marshes. Nesting activity National Wildlife Refuge. Thirty-six male spar- indicators showed a similar but non-signifi cant rows had occupied this area pre-fi re; however, pattern in response to burning. The authors only seven were recorded post-fi re. The refuge linked sparrow abundance and nesting activ- reported that six Dusky Seaside Sparrows ity to dead-vegetation cover, which was lower escaped the fi re to an adjacent private land in burn plots during the fi rst breeding season area, which was subsequently burned by its post-burn but recovered to pre-burn levels by owner. The sparrows then disappeared from the second breeding season post-burn. Gabrey the site. and Afton (2000) speculated that reduced veg- Although diffi cult to quantify, fi re may also etation cover might provide less invertebrate have more direct effects on the survival of spar- prey and nest material for Louisiana Seaside rows. Legare et al. (2000) captured and banded Sparrows. During the study, the researchers fi ve Swamp Sparrows ( georgiana) recaptured birds banded as adults in unburned on the St. Johns National Wildlife Refuge, in marshes during subsequent breeding seasons, Florida. A sparrow banded on 20 March 1994 but failed to recapture birds banded in burned was recovered dead in burned sand cordgrass marshes. The authors suggest that the sparrows within 50 m of the original banding location on move to nearby unburned marsh following a 5 January 1995, following a prescribed fi re on fi re and that such displacement could affect the refuge. Although the authors report that the short-term reproductive success by forcing bird had most of its feathers burned, a conclu- dispersal into lesser quality habitats, increasing sive cause of death was not reported. 162 STUDIES IN AVIAN BIOLOGY NO. 32

Other passerines Blackbirds, and Boat-tailed Grackles were the dominant species in these marshes. Winter In studies of wintering bird populations burns dramatically lowered Seaside Sparrow in coastal Louisiana marshes, Gabrey et al. abundance but increased blackbird and grackle (1999) found that several species of sparrows abundance in the fi rst breeding season post- and wrens avoided recently burned marshes burn. During the second breeding season but reappeared one winter later. Common post-burn, sparrow abundance increased and Yellowthroats (Geothlypis trichas) and Sedge blackbird and grackle abundance decreased in Wrens (Cistothorus platensis) were absent from burn treatment plots to the point where each recently burned marshes, during the fi rst win- variable was similar to pre-burn conditions ter, but present in unburned marshes. One year (Gabrey and Afton 2004). Bird community post-fi re, Marsh Wrens (Cistothorus palustris) changes were strongly correlated with percent were found more frequently in unburned versus cover of dead vegetation and live salt meadow burned marshes. The authors concluded that cordgrass—plots with greater percent cover winter habitat for several passerine species was had greater sparrow densities and lower black- reduced during the winter in which the burns bird and grackle densities. occurred, particularly if a high proportion of the plot burned. In contrast, Boat-tailed Grackles Raptors (Quiscalus major) and Red-winged Blackbirds (Agelaius phoenecius) preferred recently burned Some research suggests that raptors use plots, possibly because burns reduced visual smoke and fi re as a foraging cue, suggesting obstruction and increased visual contact with that raptors feed opportunistically upon prey conspecifi cs, and reduced ground cover, facili- either chased from cover by fi re or left without tating foraging for aquatic prey. cover by the burn (Baker 1940, Komarek 1969, Gabrey et al. (2001) evaluated relative abun- Tewes 1984). Anecdotal evidence suggests that dance of birds during the breeding season this occurs in marsh burns as well. Following immediately following winter burns and for two burns in gulf cordgrass communities at two consecutive breeding seasons thereafter. Aransas National Wildlife Refuge (ANWR), Structural vegetation characteristics (visual White-tailed (Buteo albicaudatus) report- obstruction and percent cover) did not differ edly dived through smoke to capture cotton rats between burned and unburned plots by the fi rst (Sigmodon hispidus), pocket mice (Perognathus summer post-burn. Neither treatment affected spp.), and grasshoppers (Acrididae) (Stevenson bird species richness or species composition. Of and Meitzen 1946). Tewes (1984) reported simi- the 10 most abundant bird species, only Sedge lar behavior during a 40-ha prescribed burn in Wrens were absent from burned marshes but gulf cordgrass at ANWR, when 14 White-tailed present in unburned marshes during the fi rst Hawks appeared near the fi re, hovering near post-burn breeding season. Sedge Wrens were the ground and grasping prey in the ash. Other present in burned marshes by the second breed- raptors noted soaring in the smoke column ing season post-burn. Total birds/survey for all and hunting in the burned area included two species combined and for sparrows (primar- Northern Harriers (Circus cyanneus), a White- ily Seaside Sparrows) did not differ between tailed Kite (Elanus leucurus), an American burned and unburned marshes during the fi rst Kestrel (Falco sparverius), and a Short-eared or third summers post-burn, but were two times (Asio fl ammeus). No raptors were noted during greater in burned than unburned marshes dur- post-fi re strip-transect counts, suggesting that ing the second summer post-burn, coinciding the enhanced foraging opportunities afforded with the recovery of dead vegetation cover to the raptors was extremely short lived. Tewes pre-burn levels. The researchers concluded that (1984) speculated this could be due to extensive managed burns for winter waterfowl foods and complete removal of vegetative cover forc- appear compatible with maintaining popula- ing small mammals, , and other prey spe- tions of certain other marsh birds, provided that cies to abandon the site. large contiguous marsh areas are not burned in any single winter, and >2 yr are allowed THE POSSIBLE ROLE OF LIGHTNING FIRES between burns. Gabrey and Afton (2004) conducted multi- Lightning-ignited fi res are a common occur- variate analyses of breeding bird abundance rence in coastal marshes, especially on the Gulf in four pairs of burned and unburned marshes Coast and southeast Atlantic Coast. Such fi res in the breeding season prior to experimental likely would have little detrimental impact on burns and in two breeding seasons post burn. bird species endemic to these areas. Some evi- Louisiana Seaside Sparrows, Red-winged dence exists to support the idea that Seaside IMPACTS OF MARSH MANAGEMENT—Mitchell et al. 163

Sparrow habitat, for example, in the Gulf Coast been done based upon documented fi re effects and southeast Atlantic Coast depends on peri- on coastal-marsh vegetation and known odic but relatively infrequent fi res (Taylor 1983, breeding or wintering habitat requirements Gabrey and Afton 2000); we are unaware of of coastal-marsh birds. Prescribed burns may published studies that address effects of burn- indirectly affect bird populations through a ing on Seaside Sparrows in the northern part variety of pathways. Some of the more obvious of their range—habitats which naturally expe- mechanisms include direct or indirect effects rience a lower frequency of lightning-ignited on vegetation structure, changes in amount fi res. In southern marshes in the absence of and distribution of open water, or changes in fi re, vegetation density increases to a point at availability and quality of plant or food which the marsh is no longer suitable to Seaside items. A summary of potential mechanisms is Sparrows. Immediately post-fi re, it appears that presented in Table 1. We emphasize that these while numbers of breeding Seaside Sparrows are possible short-term impacts, based on the and Marsh Wrens and wintering Nelson’s few quantitative studies that have been pub- Sharp-tailed Sparrows are reduced, these spe- lished. Long-term impacts have not yet been cies may subsequently show a positive response investigated. For example, Seaside Sparrow for one or more years following the immediate numbers may be temporarily reduced imme- post-burn season. However, as fi re frequency diately following a fi re but may increase for increases (i.e., to every year), fi res suppress a period afterwards. In addition, many other vegetation density, rendering both breeding variables such as water depth, salinity, and pre- and wintering habitat unsuitable for several cipitation could infl uence vegetation responses passerines (Common Yellowthroats and Sedge to fi re. Wrens) including species dependent upon coastal-marsh habitats (Seaside and Nelson’s EFFECTS OF STRUCTURAL MARSH Sharp-tailed sparrows). Frequent fi res would MANAGEMENT ON COASTAL-MARSH likely also increase habitat availability for wide- BIRDS spread habitat-generalist species such as black- birds and grackles at the expense of habitat for In addition to prescribed burns, marsh man- endemic Seaside Sparrows or Nelson’s Sharp- agers frequently alter marsh habitat by inter- tailed Sparrows (Gabrey et al 1999, Gabrey and rupting normal tidal cycles and manipulating Afton 2004). Therefore, periodic but infrequent the timing, depth, and duration of fl ooding, and fi res (Gabrey et al. 2001), possibly mimicking salinity. Structural marsh management (SMM; the historic fi re regimes of these coastal habi- Chabreck 1988, U.S. Environmental Protection tats, are probably most likely to benefi t spar- Agency 1998) involves the use of weirs, dams, row and other passerine populations on the tide gates, canals, or other structures that alter southeast coast. Whether such patterns occur the hydrology of coastal marshes. These struc- in coastal marshes outside of the Southeast is tures allow managers to manipulate water unknown. Few studies have addressed effects depth, timing of fl ooding or drying, and salin- on demographic parameters. ity, to achieve the following objectives: No studies to date have adequately 1. Prevent encroaching isohaline lines from addressed the likely effects of fi re, either natu- changing the distribution of marsh types. ral or prescribed, on other marsh-bird groups, 2. Encourage production of preferred water- such as raptors or colonial waterbirds. Research fowl and muskrat foods while discourag- on population responses of these species to con- ing growth of plants with less waterfowl trolled fi res in marsh habitats using standard value (primarily cordgrass species). methodologies and sound statistical design 3. Create or maintain shallow water or open is needed to increase our understanding of water areas. the effects of prescribed burning on the entire 4. Reduce loss of existing marshes to erosion, coastal-marsh avian community. sea-level rise, and saltwater intrusion. 5. Create new emergent wetlands from pre- MECHANISMS OF CHANGE IN COASTAL MARSH-BIRD viously inundated areas. COMMUNITIES IN RESPONSE TO FIRE 6. Provide for ingress and egress of selected estuarine organisms (e.g., shrimp and lar- Because few scientifi c studies have focused val fi sh). on the effects of prescribed burns on marsh 7. Control biting insect populations (mos- birds, the best we can do at present is to specu- quitoes). late about the potential effects of fi re on various Although the scientifi c and management species and recommend that these potential communities have begun to evaluate the effec- relationships be investigated fully. This has tiveness of SMM on coastal-marsh ecosystems, 164 STUDIES IN AVIAN BIOLOGY NO. 32 ). ),

), Swamp )? Ceryle ammeus spp.). Ammodramus Asio fl sher ( sher Sterna forsteri ) and Marsh ), Pied-billed ), Black Tern ), blackbirds ), Northern Harriers ), American Bittern Aimophila ), waterfowl, waders, ), Red-winged Blackbird Falco peregrinus ). ), Common Yellowthroat Ammodramus maritimus ), Seaside Sparrow ( Fulica americana ), Forster’s Tern ( ), Short-eared owl ( Ixobrychus exilis Chen caerulescens Cistothorus platensis Laterallus jamaicensis Melospiza georgiana ), other sparrows ( Podilymbus podiceps Cistothorus palustris ), Peregrine Falcon ( Botaurus lentiginosus Chlidonias niger Circus cyaneus Agelaius phoeniceus Geothlypis trichas alcyon maritimus . ats for foraging Grebes ( AVIFAUNA

MARSH

ng shorebirds. ON

IMPACTS

POSSIBLE

AND

STRUCTURE

VEGETATION

ON

BURNS

PRESCRIBED

OF

res inhibit Lack of shrubby cover, perches − Species nesting in or foraging Colonial waterbirds, Belted Kingfi FFECTS 1. E ABLE remove emergent vegetation Frequent fi temporarily reduced due to loss of standing vegetation visual surface feeders (Icteridae). Effect of prescribed burn Cover burns reduce or Marsh appearance Vegetation height and density + Impact Waterfowl, gregarious, Guild(s) potentially affected Representative species Snow Goose ( T woody vegetation establishment (live or dead) absent, standing dead vegetation cover removed, from strong perches; species nesting in shrubs or otherwise Sparrow ( water level, cover for small of previous year’s grasses; Least Bittern ( Cover burns reduce or remove surface litter Marsh surface exposed, no decumbent vegetation above − Species that nest in/on this canopy removed material or that nest on mats Black Rail ( ( require woody structure ( vegetation Peat or root burns Open water more abundant; mammals and tunnels removed − Species using emergent species that prey on rodents ( Sedge Wren ( submerged vegetation increases; + marsh or mudfl Species that forage on ground Seaside Sparrow ( no longer inhibited by dense inhibits re-vegetation re-vegetation expose underlying mineral soil inhibits emergent vegetation replaced by shallow to moderately deep vegetation in unbroken marsh; Wren ( species that nest or forage on ponds; long-term inundation ( Ponds remain as open water, + ground Species using broken, hemi- American Coot ( ( Fire suppression obscured Dense emergent vegetation and ? or pond bottoms exposed due to tides or drought litter accumulation, woody shrubs or loafi colonize; small open-water ponds IMPACTS OF MARSH MANAGEMENT—Mitchell et al. 165 few quantitative studies have been published. areas of open water suitable for dabbling duck The greatest extent of SMM application to foraging (Gordon et al. 1989). Finally, Gabrey et coastal marshes are in Louisiana and South al. (1999) conducted fi ve aerial counts of white Carolina (Day et al. 1990, U.S. Environmental geese (Snow Goose and Ross’ Goose) wintering Protection Agency 1998); consequently, most in managed marshes in southwestern Louisiana published studies of impacts on wetland veg- from December 1995 to February 1996 and etation and wildlife comes from these two found several fl ocks present in recently burned, states. We summarize below the current state unimpounded marshes or recently burned, of knowledge regarding effects of hydrology impounded marshes; however, no description manipulations on coastal-marsh birds. of goose behavior (e.g., foraging was reported). No geese were observed in unburned, unim- BIRD USE OF IMPOUNDMENTS DURING WINTER pounded marshes.

Waterfowl Other bird species

As with prescribed burns, habitat manage- Most assessments of the value of structural ment for waterfowl, particularly wintering marsh management evaluate relative abun- habitat, has been a major justifi cation for SMM. dance of birds during winter or migration SMM became a common practice in the 1950s periods, and focus upon birds associated with and the fi rst evaluation of its effectiveness was ponds, mudfl ats, or open water, i.e., waterfowl, presented by Chabreck (1960). He reported that shorebirds, herons, egrets, gulls, and terns. prior to construction of impoundments in 1954, Habitat within impounded marshes may sup- about 75,000 waterfowl wintered on Rockefeller plement natural habitat in unmanaged marshes State Wildlife Refuge in southwest Louisiana. In or provide protection from oil spills or other post-construction surveys, however, >320,000 coastal catastrophes. Weber and Haig (1996) waterfowl wintered in the new impoundments, counted shorebirds and waterfowl in man- with another 120,000 in surrounding areas aged and unmanaged coastal marshes in South within the refuge. He attributed the dramatic Carolina. Managed marshes were drawn down increase in numbers to increased food produc- through April then re-fl ooded in June, July, or tion and constant shallow water. Chabreck et August. They found that throughout the winter al. (1974) later compared duck use of impound- and spring seasons (January–May), shorebird ments with that of control areas—unimpounded density at high tide (when natural marshes marshes and marshes that had been drained were fl ooded) was greater in managed exposed and converted to pasture. They reported that in mudfl ats than in natural marshes. Even dur- general duck usage was highest in freshwater ing low tides, shorebird density was generally impoundments; numbers varied with vegeta- greater in managed than in natural marshes. tion type, water depth, and time of year. They concluded that managed marshes pro- Gordon et al. (1998) compared relative duck vide alternative or supplementary feeding or abundance between abandoned rice fi elds roosting habitats during high tides or adverse that were diked and managed for waterfowl weather. Differences in shorebird density were and adjacent tidal (unimpounded) wetlands attributed to consistently shallower water depth in South Carolina. Winter use of managed and greater invertebrate occurrence in managed wetlands by seven dabbling duck species was marshes. greater than expected; winter use of unman- Impoundments in South Carolina are typi- aged tidal marshes was less than expected cally managed for production of wigeongrass for six of the seven species, American Black (Ruppia maritima), spikerushes (Eleocharis spp.), Duck (Anas rubripes) being the exception. The bulrushes., and other waterfowl foods through authors attributed these fi ndings to differ- spring drawdowns and summer re-fl ooding ences in hydrology of the two types of marshes. (Epstein and Joyner 1988). Consequently, vege- Tidal marshes are fl ooded and drained daily; tation composition differs between impounded hence, availability of open water for foraging is marshes and natural tidally infl uenced marshes, unreliable. In addition, the intertidal period, in which are dominated by big cordgrass (Epstein which the marsh surface is exposed, allows for and Joyner 1988). Epstein and Joyner (1988) denser vegetation growth that inhibits water- compared relative abundance of waterbirds fowl access. In managed marshes, however, in six managed (impounded) and two unman- water level may remain relatively constant at aged (unimpounded) South Carolina marshes. a depth suitable for waterfowl foraging and Except for Clapper Rails (Rallus longirostris) the continuous fl ooding may prevent dense and Northern Harriers, most bird species vegetation growth while maintaining large groups (particularly shorebirds, waterfowl, and 166 STUDIES IN AVIAN BIOLOGY NO. 32

waders) were more abundant in managed than Other bird species in unmanaged marshes. The authors felt that the greater number of species and of individu- Bird use of impounded and unimpounded als in managed marshes was due to moist soil marshes during the breeding season has conditions that increase access to invertebrates received little attention. Brawley et al. (1998) and seeds and to fi sh prey concentrated in pro- compared breeding bird abundance in two gressively smaller ponds. restored (formerly impounded) marshes with The above studies have addressed to some that of three reference sites in Connecticut. They extent the question of whether waterbird use found that marsh specialists—those species that differs between impounded and natural or breed only in coastal marsh (Seaside Sparrow, unimpounded marshes. However, birds that Willet [Catoptrophorus semipalmatus], Marsh do not use open water or mudfl at habitats but Wren, and Saltmarsh Sharp-tailed Sparrow nest or forage in the emergent vegetation (e.g., [Ammodramus caudacutus])—were more abun- passerines, rails, some herons, and egrets) have dant in the restored marshes than in the refer- received less attention. Gabrey et al. (1999) ence marshes. Three of these four species are addressed the issue of wintering passerines listed as threatened (Willet) or of special con- in impounded versus unimpounded coastal cern (both sparrows), in the state. The authors marshes in Louisiana. They found that some state that these species were absent from the species (Seaside and Nelson’s Sharp-tailed restored marshes prior to the re-establishment sparrows) were found almost exclusively in of tidal activity. Brawley et al. (1998) suggest unimpounded marshes, possibly because of a that the frequent tidal inundation and expo- preference for shorter vegetation and because sure maintained the low-marsh community ground-foraging behavior required exposed dominated by short-form smooth cordgrass in marsh surfaces. However, impoundment effects which Seaside and sharp-tailed sparrows prefer were confounded with salinity effects. This to nest. Marsh areas in the high-marsh zone are raises the question of the importance of vegeta- not fl ooded frequently enough or of suffi cient tion variables in habitat selection. While most duration to allow for establishment of short- avian ecologists agree that vegetation structure form smooth cordgrass. Marsh areas below the is an important criterion, other factors such as low-marsh zone are permanently fl ooded and invertebrate abundance, salinity, competitors, so also do not support smooth cordgrass. or predators may infl uence bird community Brawley et al.’s (1998) fi ndings, that marsh composition differently in managed compared impoundment benefi ts a diversity of bird spe- to unmanaged marshes. cies but limits habitat availability for marsh- It is interesting to note that three of the specialist species, supports results from other species listed above as being more abun- regions. In New Jersey, Burger et al. (1982) dant in unimpounded, natural marshes are found greater biomass and diversity of birds coastal-marsh endemics (Clapper Rail, Seaside in impounded marshes compared to ditched or Sparrow, and Nelson’s Sharp-tailed Sparrow). unimpounded marshes. However, species that Consequently, although impounded marshes nest exclusively in coastal marshes (Seaside and benefi t a large suite of species, conservation of sharp-tailed sparrows, and Clapper Rails) were unimpounded coastal marshes is necessary for recorded only in unimpounded marshes. They coastal-marsh endemics. (Burger et al. 1982) stated that while generalist or relatively abundant species used impounded BIRD USE OF IMPOUNDMENTS DURING THE BREEDING marshes, maintaining natural unimpounded SEASON coastal marsh was necessary for the conserva- tion of coastal-marsh specialists. Waterfowl Gabrey et al. (2001) detected different bird communities present in impounded Several waterfowl species breed in coastal and unimpounded marshes in southwest- marshes of the northeast Atlantic coast (e.g., ern Louisiana. Red-winged Blackbirds and Mallard [Anas platyrhynchos], American Black Boat-tailed Grackles were more abundant in Duck, Blue-winged Teal [A. discors], and impounded than in unimpounded marshes, Gadwall [A. strepera] although most such popu- whereas Seaside Sparrows were more abundant lations are small (Bellrose 1976, Sauer et al. 2004). in unimpounded than in impounded marshes. In southern marshes Mottled Ducks nest in large The authors attributed these differences to veg- numbers (Moorman and Gray 1994). However, etation structure and hydrology. Vegetation of we are unaware of any published studies that impounded marshes included patches of - address effects of marsh impoundment on any tails (Typha spp.) and common reed (Phragmites waterfowl nesting in coastal marshes. australis); blackbirds and grackles readily IMPACTS OF MARSH MANAGEMENT—Mitchell et al. 167 nested in these patches of tall vegetation. Chabreck 1988). Ditches approximately 2–4 m Unimpounded marshes, on the other hand, wide and 1–2 m deep were dug in parallel fash- were dominated by low-growing salt meadow ion every 50 m in high-marsh areas from the cordgrass and inland saltgrass. These two plant upland-marsh ecotone bayward. The amount species form a low, densely interwoven canopy of Atlantic Coast marsh altered by this method of relatively uniform height (<1 m), which pre- has been estimated at about 90% and extends sumably provides protection from predators from Massachusetts to Florida (Tiner 1984, The for the ground-foraging Seaside Sparrow. In Conservation Foundation 1988). addition, the surface of impounded marshes is With the increasing awareness of the high often continually fl ooded. The marsh surface of productivity of coastal marshes in estuaries unimpounded marshes is exposed during part (e.g., the rise of the Odum school in ecology of the tidal cycle; hence sparrows are able to for- during the late 1950s and 1960s) and recogni- age on the ground. tion of the importance of natural tidal fl ood- ing and hydrology to the integrity of marsh MECHANISMS OF CHANGE IN COASTAL-MARSH BIRD systems, improvements in marsh management COMMUNITIES IN RESPONSE TO SMM were attempted. Experimentation began in New Jersey with a method that became known Structural marsh management infl uences as open-marsh water management (OMWM) coastal-marsh bird communities through its (Cottam 1938, Ferrigno and Jobbins 1968). This effects on open water or mudfl at availability, method substitutes biological control of mos- timing and frequency of fl ooding, modifi cation quito larvae using predatory fi sh, and by alter- of the plant community, and salinity (Table ing mosquito egg-laying habitat, instead of 2). In general, SMM appears to benefi t water- drainage and pesticide applications. In short, fowl and other species such as herons and mosquito depressions in the marsh not con- blackbirds that are attracted to open water, nected to existing ditches are either connected exposed mudfl ats, lower salinity, or tall, dense to ditches using new spurs or, if the depres- vegetation. This likely is due to reduced diur- sions are very dense, a pond is constructed. nal variability in fl ooding due to the exclusion The ponds originally were small (<0.05 ha), of tides. Thus, impoundments that are drawn deep (often >60 cm), and had a deeper area or down to moist soil conditions maintain those sump added to enable mummichog (Fundulus conditions until managers fl ood the impound- spp.) to survive during summer droughts. The ment. In contrast, unimpounded marshes fl ood material dredged to create the new ditches and at daily high tides; mudfl ats are then exposed ponds was spread thinly over the marsh sur- for only about half a day. Disruption of tidal face to reduce the prospects that common reed hydrology often increases the area of open (Phragmites australis) or woody vegetation such water and decreases the amount of grass and as marsh elder (Iva frutescens) might invade. short herbaceous vegetation. Consequently, Later, in other regions such as , the although SMM provides habitat for a diversity practice expanded but some modifi cations of bird species, certain species such as Seaside were added, such as adding sills to the ends of Sparrows, sharp-tailed sparrows, and Clapper large ditches to retain ground water (Meredith Rails, that require grassland-like conditions et al. 1987). In spite of the popularity of the and alternating cycles of inundation and expo- method in New Jersey and Delaware and its sure of the marsh surface, likely do not benefi t expansion to other states, little research on from impoundments. effects on wildlife has been performed and published in the peer-reviewed literature EFFECTS OF MOSQUITO CONTROL AND (Erwin et al. 1994 and references therein). In OPEN-MARSH WATER MANAGEMENT ON addition, Wolfe (1996) provided a summary of COASTAL-MARSH BIRDS the effects of OMWM on birds, fi sh, mammals and other tidal resources in the Atlantic region. The history of coastal-marsh alteration to The practice remains somewhat controversial control the mosquito as a human pest and dis- among wetland ecologists and federal and ease vector, or for agriculture (livestock grazing state resource managers because of concerns and salt-hay farming), goes back centuries in the for converting and altering pristine marsh US (see Daiber 1987 for a review). During the (Table 3). early part of the 20th Century, the Old World Post (1974) was one of the earliest to dem- notion of draining much of the high marsh was onstrate the behavioral and ecological effects popular, and thus began an ambitious campaign of ditching on marsh birds, specifi cally Seaside of ditching both by hand and with horse or Sparrows, revealing that ditches could alter mule during the 1930s and 1940s (Daiber 1987, the shape and sizes of territorial boundaries. 168 STUDIES IN AVIAN BIOLOGY NO. 32 ), ), s. ), Marsh ). ). ), Common Agelaius phoeniceus ). Quiscalus major Ammodramus maritimus Geothlypis trichas Cistothorus platensis . Cistothorus palustris AVIFAUNA Ammodramus nelsoni

MARSH

ON

IMPACTS

POSSIBLE

AND

STRUCTURE

) + Species that nest in tall dense Red-winged Blackbird ( VEGETATION

ON spp., − Species associated with Seaside Sparrow, Nelson’s Sharp-tailed Sparrow Phragmites (SMM) Distichlis ) or reeds ( ats may be available + Shorebirds, waterfowl, gulls. Typha MANAGEMENT

MARSH

STRUCTURAL

OF

FFECTS 2. E ABLE salt meadow cordgrass) grassland-like habitats ( vegetation increases Effect of SMM Marsh ponds not subject to tides Ponds retain water year-round; vegetation winter + Marsh appearance limited? Species that feed or loaf on Waterfowl, coots, herons, egrets, tern Timing of drawdowns ratio of open water to emergent controllable, not dependent on droughts; Impact Open mudfl wetland manager actively Guild(s) potentially affected summer drawdowns may conducts seasonal draw enhance production of seed- vegetation increases, submerged downs Representative species producing annual plants; open shallow to moderately deep during spring or fall migration; throughout Marsh surface inundated year- round; access to invertebrates ponds ponds decreases − Ground-foraging or ground- Seaside Sparrow ( water may be available nesting species Flood-tolerant vegetation dominates; wetland Cattail ( manager favors out deep- water management regime Marsh surface may be inundated distribution increases year-round; access to invertebrates Sedge Wren ( may be limited? − distribution Availability of open water habitat may be limited during drawdowns Ground-foraging or ground- − nesting species Seaside Sparrow, Sedge Wren? Species that feed or loaf on shallow to moderately deep Waterfowl, coots, herons, egrets, terns. vegetation Short vegetation ( Boat-tailed Grackle ( Wren ( Yellowthroat ( T IMPACTS OF MARSH MANAGEMENT—Mitchell et al. 169

TABLE 3. POTENTIAL ECOLOGICAL EFFECTS OF OPEN-MARSH WATER MANAGEMENT ON COASTAL EMERGENT MARSHES OF THE ATLANTIC COAST.

Negative effects Positive effects Loss of salt meadow cordgrass habitat for Reduction of mosquito breeding sites. Seaside (Ammodramus maritimus) and sharp-tailed sparrows (Ammodramus nelsoni and A. caudacutus); loss of short-form smooth cordgrass (Spartina alternifl ora), Fragmentation of inner marsh with pools and Increased forage fi sh populations and enhanced waterbird radials; exacerbation of erosion and marsh (wading birds, shorebirds) feeding habitats. loss in the face of sea-level rise Compaction of emergent marsh due to Restoration of hydrology (with ditch plugging). operation of heavy equipment on marsh surface Invasion of shrubs, (Iva spp., Baccharis spp.), Augmentation of perches and nesting substrates for and reeds (Phragmites australis) due to passerines (marsh sparrows and wrens), wading birds. slight elevation changes; change in vegetation community structure

For larger waterbirds, more recent studies in study by Meredith and Saveikis (1987) revealed California have revealed that under certain that waterfowl use of OMWM ponds was only circumstances, waterfowl use of marshes origi- about one half that of natural ponds. The con- nally ditched and then diked for mosquito clusions of that study are problematic however, control can achieve positive results for both because natural ponds were larger than were objectives (Batzer and Resh 1992). Along the OMWM ponds. Walbeck (1989) conducted a Atlantic Coast, studies in New England demon- study with limited information (only conducted strated that draining of high marshes reduced for 1 yr) on the Eastern Shore of Maryland their use by waterbirds because of the loss of where large impoundment use by waterfowl ponds and pannes (Clarke et al. 1984, Brush et was greater than use of OMWM ponds. al. 1986, Wilson et al. 1987). Several studies conducted in the mid- Other, early studies often examined marsh- Atlantic region, examining many waterbird alteration effects only at one local site and only species, revealed that sizes of ponds and the on one or a few species, such as Herring Gulls water/marsh ratio of the study site were the (Larus argentatus; Burger and Shisler 1978) or most important determinant of use. Burger et Clapper Rails (Shisler and Schultze 1976). A al. (1982) examined six different marsh sites more comprehensive analysis of OMWM effects in New Jersey and found high use of larger on waterfowl at fi ve New Jersey sites from ponds by some shorebird and wading bird 1959–1984 was attempted (Shisler and Ferrigno species; however, they cautioned that adding 1987); however, counting techniques and per- ponds to the high marsh could adversely affect sonnel changes rendered interpretation of the breeding Clapper Rails. Erwin et al. (1991) results diffi cult. found among nine marsh sites in three states In a study of effects of OMWM on waterbirds that the water/marsh ratio was positively cor- in New Jersey, Erwin et al. (1994) determined related with use by waterfowl, and separately, year-round relative abundance of waterfowl, American Black Ducks, but pond number was shorebirds, waders, gulls, and terns in ponds not. Larger ponds (>0.25 ha) tended to be used in OMWM-managed marshes, unmanaged tidal more than smaller ponds by most bird species, ponds, and managed impoundments (>400 ha). but no treatment effect (OMWM vs. natural They found that spring and summer densities pond) was found. In a later experimental of American Black Ducks were greatest in the study at Forsythe National Wildlife Refuge two large impoundments when compared to in New Jersey, Erwin et al. (1994) compared OMWM and tidal ponds. use by larger waterbirds of OMWM, small In New England, several authors monitoring natural ponds, and nearby impoundments. shorebirds, wading birds, and waterfowl have They reported results that varied by guild concluded that use of marsh sites treated with and season. Higher densities were not always OMWM resulted in little difference when com- found in larger ponds for waterfowl, but this pared with sites with natural ponds. However, did seem to be the case for spring-summer the method in New England did not include cre- shorebirds. When comparing small pond use ation of new ponds (Clarke et al. 1984, Brush et (both OMWM and natural) with impoundment al. 1986, Wilson et al. 1987). In Delaware, a 2-yr use, however, American Black Ducks and 170 STUDIES IN AVIAN BIOLOGY NO. 32 other waterfowl used impoundments in higher densities for both fall and winter feeding and

nesting than they did small marsh ponds. They e to recommended that a smaller number of larger ponds be created in the high marsh if mosquito control is deemed necessary, and that ponds have shallow and sloping sides to accommo- ). of birds.

date shorebird, wading bird, and rail use. The TATES authors also concluded that in areas near large S impoundments, small water bodies would add NITED little waterbird habitat value. numbers U THE ANIMALS AS MARSH ARCHITECTS/ IN MANAGERS (AND THEIR MANAGEMENT)

Although wildlife managers tend to think b of marsh management as a strictly human PRIMARILY endeavor, many animal species have demon- ( strated quite remarkable abilities to manipulate

the structure, and hence functions, of marshes MARSHES

to differing degrees (Table 4). In some regions, - for rails, marsh sparrows, waterfowl (feeding). as their populations have increased, some of + for waterfowl, open-water species,

these species have created conditions con- COASTAL

sidered undesirable from the perspective of OF resource managers. Thus, managing the animal managers has become simultaneously a chal- lenge and an ethical paradox, i.e., managing the

STRUCTURE Schoenoplectus ) marsh environment for human values is accept-

able but for other animals to do so requires THE

corrective (often lethal) measures (Table 5). We ), + for shorebirds and wading birds. will explore and summarize some of the major aspects of animal architect activities in the US in ora MODIFYING the following sections. IN

BIRDS ); fragmentation of eat-outs - for rails, marsh sparrows. . Schoenoplectus americanus The effect of marsh grazing by the Snow Spartina alternifl ARCHITECTS Goose can be signifi cant in coastal marshes, because the birds typically pull up the above- MARSH ground stems to gain access to the rhizomes AS

(Belanger and Bedard 1994, Jefferies and Rockwell 2002). In brackish marshes, geese ACT tend to uproot primarily bulrush species while

in saltmarshes along the Atlantic, the princi- THAT pal plant affected is smooth cordgrass. Larger patches of denuded marsh were referred to as SPECIES eat-outs. Effects on marsh (at high densities) Effects on birds

In the early years of study (1940s–1970s) of a

the Snow Goose, such goose eat-outs in win- ANIMAL ter were believed to be benefi cial to wildlife, LA, TX, MS, MD Destruction of bulrushes ( as apparently they opened up parts of the ) in US; nearly global (cattle) and invertebrates, destroys nests reduced food. MAJOR ) (

) ( monotypic marshes and allowed access for ) spp OF feeding by a variety of other birds and fur ) (MD–GA), southern cover and destroys nests - for nesting gulls, and total bearers (Lynch et al. 1947, Chabreck 1988). In t for the target species/group, whereas a - sign indicates negative effect. fact, small patches of eat-outs in the cordgrass XAMPLES marshes of New Jersey and Delaware, which make small fi shes and invertebrates more 4. E Ondatra zibethicus Myocastor coypus Equus caballus Bos taurus/Ovis aries available to predators, can attract over six spe- Chen caerulescens ABLE + indicates a benefi State names are abbreviated. Muskrat ( ( US wide Cattle/sheep Atlantic and Gulf coasts Eat outs especially Olney threesquare Trampling reduces both annual grasses + for waterfowl (roosting), open-water species, - for wintering waterfowl and shorebirds, rails du Horses ( Southeast US islands Reduced structure, trampling reduces + for total species richness of birds; Nutria ( Species Area cies of feeding spring migrant shorebirds, as T Snow Goose ( East and Gulf coasts Large eat outs of smooth cordgrass a - for breeding rails, marsh sparrows, terns b IMPACTS OF MARSH MANAGEMENT—Mitchell et al. 171

TABLE 5. MANAGEMENT METHODS EMPLOYED TO CONTROL DENSITIES OF SPECIES THAT ACT AS MARSH ARCHITECTS IN MODIFYING THE STRUCTURE OF COASTAL MARSHES.

Species Methods adopted Outcome Snow Goose Special early season hunts, scare Scaring and fall-winter hunts mostly ineffective; (Chen caerulescens) decoys and noisemakers, shooting recent spring hunts in under evaluation. on the breeding grounds (Canada) Muskrat Trapping Ineffective currently since market value is so low. (Ondatra zibethicus) Nutria Trapping; shooting, poisoning Ineffective to date with low market values; (Myocastor coypus) shooting effective in some winters in Maryland. Horse Reducing size of herd by culling; Sterilants costly and time consuming; exclosures (Equus caballus) use of exclosures, and sterilants only for local control. Roundups may be most effective (e.g., annual pony roundup in Chincoteague, Virginia). Cattle/sheep (Bos Reducing size of herd by culling, Pasture rotation (seasonal) and annual cull and taurus/Ovis aries) exclosures, and pasture rotation sale most effective. well as summering egrets, herons, and Glossy MAMMALS Ibis (Plegadis falcinellus; R. M. Erwin, unpubl. data). In the past few decades, however, Snow Muskrats Goose populations have exploded, resulting in major marsh damage on the breeding grounds The muskrat is a native species that, like especially near St. James Bay, Canada, on the Snow Goose, has evolved in the marshes of staging areas along the St. Lawrence River, . The role of muskrats and their and on wintering areas from New Jersey to management in marshes remains one of the Maryland (see Batt 1998 for a summary of classics in North American wildlife literature the goose problem). Intense grazing by large (Errington 1961). Without muskrats, fresh and numbers of these social birds has resulted in brackish marshes may often become domi- rather large eat-outs that may require a decade nated by cattail although moderate muskrat or more for the vegetation to recover (Smith densities control the cattail and keep the marsh and Odum 1981, Young 1987). In some cases open. Waterfowl managers speak of an ideal if the bare areas were extensive, they have hemi-marsh with 40–50% open water in which lost their organic composition and became muskrats are dense enough to control cattails hypersaline; the marsh may have shifted to an and keep some open water, but are in turn kept alternative stable state (Jefferies and Rockwell under control by regular trapping (O’Neil 1949, 2002). Early attempts to remedy the goose Bishop et al. 1979). In coastal marshes along the problem relied on using hazing techniques Atlantic and Gulf coasts, the species that may and special extended season hunts during benefi t most from muskrat foraging activities fall and winter on national wildlife refuges, and tunneling include migrant and winter- initially at Forsythe National Wildlife Refuge, ing Blue-winged Teal, Green-winged Teal New Jersey (M. C. Perry, USGS, pers. comm.), (Anas crecca), Mallard, American Widgeon (A. Bombay Hook, and Prime Hook national wild- americana), and American Black Duck. During life refuges, Delaware (P. Daly, USFWS, pers. the breeding season, Coastal Plain Swamp comm.). These proved largely unsuccessful Sparrows appear to achieve their highest densi- however in reducing regional populations, and ties in association with intense muskrat work- in recent years, the Canadian Wildlife Service ings (B. Olsen and R. Greenberg, pers. comm.). is directing a special large-scale spring breed- On occasion, muskrat population densi- ing season harvest (Batt 1998; Table 5). ties and associated tunneling activities may The manager’s challenge concerning the result in confl icts with wildlife management in Snow Goose becomes one of partial suppres- marshes (Lynch et al. 1947). Examples include sion of a native species that is an important eroding the earthen plugs that marsh manag- game species, a popular species among bird ers use in constructing OMWM sill ditches in watchers and photographers, a charismatic Delaware (W. Meredith, Delaware Division species that precludes some types of draconian of Fish and Wildlife, pers. comm.) and plug- control methods (e.g., poisoning), and a species ging old tidal ditches in New England (C. T. that has had a long co-evolutionary history with Roman, National Park Service, pers. comm.). marsh-vegetation dynamics. Although poisons have been used on occasion 172 STUDIES IN AVIAN BIOLOGY NO. 32 as control, regular trapping remains the most of patches of short and tall grasses (Menard et widely acceptable method to control popula- al. 2002). This suggests potential indirect com- tions (Table 5); in recent decades however, with petition between horses and dabbling ducks declining fur prices, reduced trapping has ren- (Menard et al. 2002). In the mid-Atlantic region dered population control ineffective (Chapman of the US, horse grazing was thought to reduce and Feldhammer 1982). the density of smooth cordgrass (Furbish and Albano 1994). In North Carolina, marshes Nutria subjected to moderate grazing by feral horses supported a higher diversity of foraging water- The nutria (Myocastor coypu), a native of South birds, a higher density of crabs, but had less America, was released in the Louisiana marshes vegetation and a lower diversity and density in 1938 as part of a fur-bearing animal experi- of fi shes than ungrazed marshes (Levin et al. ment and rapidly expanded throughout the 2002). Horse trampling of bird nests has occa- Gulf Coast brackish marshes (Kinler et al. 1987, sionally been detected (I. Ailes, USFWS, pers. Chabreck 1988). Along the East Coast, nutria comm.) but is probably a minor factor in most are found sporadically from Georgia north to locations. the Blackwater National Wildlife Refuge in the The primary method of controlling feral Chesapeake Bay, Maryland, where they have horse numbers is simply reducing herd sizes created much controversy because of signifi cant and alternating the use of pasturage and wet- marsh losses on refuge lands (Chapman and land areas. On Assateague Island National Feldhammer 1982). As with the Snow Goose, Seashore and the Chincoteague National small and localized nutria populations did not Wildlife Refuge in eastern Virginia, a fi xed damage marshes, and it had been claimed that percentage of the annual foal production is only for giant cordgrass (Spartina cyanosuroides) removed from the herd during a July drive and did nutria have any major impact (Harris and managed roundup and are auctioned to the Webert 1962). In moderate numbers, nutria were public in what has been a major tourist event felt to benefi t some waterfowl, because the ani- (Keiper 1985). Experimentation with steriliza- mals created open patches in otherwise dense tion of horses has also been tried at several marsh grass (Chabreck 1988). However, like island locations along the Atlantic Coast, but the muskrat, the fur-trade decline has resulted with limited success (J. Schroer, USFWS, pers. in fewer trappers, and hence less control of comm.). Sterilization of dominant stallions local and region populations by trapping. As without other control measures is unlikely to a result, large populations of these herbivores control feral horse populations. have caused very extensive eat-outs, resulting in marshes reverting to open water pools and Cattle and sheep lakes. In Maryland, the state natural resource agency is attempting to eradicate the species As with horses, light-to-moderate numbers by trapping and shooting on all public lands of livestock (0.5–1.0 animal/ha) probably are (B. Eyler, Maryland Department of Natural not deleterious to marsh vegetation or to the Resources, pers. comm.). associated bird life (Chabreck 1988). Cattle graze forbs and shrubs and may retard the Horses invasion of woody vegetation into emergent marshes (Menard et al. 2002). In Europe, cattle In a relatively small number of regions today grazing has been cited as benefi ting grazing (e.g., southern France, Spain, and southeastern waterfowl as well as a common nesting shore- US), domestic or feral horses (Equus caballus) bird (Redshank [Tringa tetanus]) by maintaining occur in coastal marshes, at times in high den- early successional stages and a diverse array sities (Keiper 1985, Menard et al. 2002). As in of halophytic plant species, (Norris et al. 1997, previous examples, light to moderate grazing Esselink et al. 2000). Along the US Gulf Coast, probably has little effect, but with more intense Chabreck (1968) mentioned that moderate cattle grazing impacts accumulate. In Georgia, signifi - grazing in marshes might benefi t the Snow cantly more periwinkle snails (Littorea irrorata), Goose (Chabrek (1968) and, more interestingly, a potential waterbird prey, were found inside the Yellow Rail (Coturnicops novaboracensis; compared to outside of exclosures, and tram- Mizell 1999). On the other hand, overgrazing pling by horses reduced above ground biomass by cattle reduces biomass of many annual seed- of vegetation by 20–55% (Turner 1987). In south- producing grasses and sedges, reducing food ern Europe, horses reduced plants more than availability for wintering waterfowl, especially did cattle (Bos taurus), removed more vegetation ducks (Chabreck 1968). In addition, in Germany per unit body mass, and maintained a mosaic an experiment conducted using three levels of IMPACTS OF MARSH MANAGEMENT—Mitchell et al. 173 grazing (0.5–2.0 animals/ha) over 9 yr demon- salinity, ambient or water temperature, humid- strated that grazing could depress population ity, fuel load, fi re intensity, and season of burn densities, species richness, and community likely strongly infl uence vegetation responses diversity of invertebrates (Andresen et al. 1990); but have not been investigated. In particular, hence, many shorebird species could potentially comparisons between biological responses to be affected. winter management burns and summer light- Sheep (Ovis aries) grazing in wetlands is ning fi res could improve our understanding most common in Europe. In general, as wet- of the pre-management-era role of fi re in these lands revert to upland pasture for sheep and systems, and possible marsh community altera- cattle by drainage or diking into polders, poten- tions caused by human-imposed fi re regimes. tial wetland-dependent birds suffer habitat loss; Similarly, effects of prescribed fi res on inverte- such has occurred in The Netherlands (Hotker brate foods are unclear. 1992) and elsewhere in Europe (Finlayson et Gulf Coast marshes in which Seaside al. 1992). In England, some attempts have been Sparrows breed are prone to lightning-ignited made to reduce the potential confl ict between fi res; thus, these birds have likely evolved sheep grazing and wintering waterfowl use by behavioral or other responses that allow their imposing seasonal restrictions for sheep grazing persistence in a frequently disturbed habitat. from April–October in designated wet pastures Prescribed fi res appear benefi cial to breeding (Cadwalladr and Morley 1973). sparrow populations, presumably because veg- Management of potential deleterious effects etation that inhibits the birds’ movements along of cattle and sheep involves reducing the herd the ground is removed. Wrens and other small periodically or alternating pasturage areas. passerines apparently avoid recently burned Also, where signifi cant waterfowl populations marshes for about 1 yr, likely due to loss of arrive in fall and winter, seasonal closures of vegetative cover. Burning marshes during the some marshes from October through March fall and winter reduces winter habitat qual- may be appropriate to reduce disturbance. ity for migratory species such as Sedge Wrens and Nelson’s Sharp-tailed Sparrows, which SUMMARY AND FUTURE DIRECTIONS winter almost exclusively in coastal marshes. Widespread and abundant species such as Red- Coastal marshes are subject to lightning- winged Blackbirds and Boat-tailed Grackles ignited fi res that typically occur during the seem to prefer recently burned marshes. summer when thunderstorms are most fre- Observations of fi re effects on raptors and quent, and vegetation is actively growing. On waterbirds are far too limited to make any sig- the other hand, marsh managers typically burn nifi cant inferences. Although these species do marshes during late fall or winter, the time when not necessarily nest in the marsh itself, they are migratory or wintering waterfowl are present, important components of the marsh system as and vegetation, at least at higher latitudes, is predators and vehicles of nutrient cycles; their generally dormant. Observational data provide responses should be investigated further. limited evidence that these management burns Impounded marshes appear to attract water- attract some species of waterfowl (wintering birds in greater numbers than do neighboring Snow Goose in particular), at least occasion- unmanaged, tidally infl uenced marshes and ally. Unfortunately, lack of comparisons with may contribute signifi cantly to shorebird con- unburned or control marshes limit inferences servation because they provide supplemental that can be made from these observations. feeding and roosting areas, particularly when We only can speculate as to what feature(s) of natural marshes are inundated by high tide. these burned marshes are attractive (e.g., food However, passerines and other species that availability and nutritional content of vegeta- do not frequent large open-water ponds or tion, changes in predator communities, social mudfl ats may be negatively affected by conver- interaction, and/or altered vegetation structure sion of tidal marshes into non-tidal marshes. facilitating animal movement), or under what Impounded marsh habitat differs suffi ciently other environmental conditions waterfowl will from unimpounded marsh habitat in that use burned marshes (e.g., availability of food in distinctive bird groups use one but generally the surrounding landscape). avoid the other. Thus, managers are faced with Results of studies of vegetation responses a diffi cult task of integrating and improving indicate that prescribed burns sometimes, but management of impounded marshes with the not always, produce the desired results (i.e., management and preservation (in as natural changes in plant community composition, bio- a state as possible) of unimpounded marshes. mass, or seed production). Numerous environ- Areas in which information appears to be lack- mental or other factors, including water depth or ing for coastal impounded marshes include 174 STUDIES IN AVIAN BIOLOGY NO. 32 effects of timing and duration of drawdown on ponds may be used by more birds than smaller wildlife use and invertebrate communities ones, no treatment effect was detected (i.e., cre- Management values of impounded marshes ated versus natural pond use); also, at least for during the breeding season are similar to those waterfowl, nearby large impoundments (100s of during winter and migration. Birds that benefi t hectares in size) harbored both a larger number are typically those associated with open water and higher density of birds than did the cre- ponds or mudfl ats; the specifi c nature of ben- ated ponds in fall and winter. Thus, the entire efi ts for even these species have not been rigor- landscape surrounding the treatment areas of ously evaluated. At the same time, water levels the marsh must be considered when addressing within impoundments often are too deep to be habitat use. Recent improvements in the design suitable for ground-foraging passerines. These of small OMWM ponds include using very shal- species appear to depend on periodic exposure low, sloping perimeters to maximize shorebird of the marsh surface, possibly to facilitate forag- use, and creating larger ponds unless dredged ing or because invertebrate prey are more vul- material deposition precludes that option. nerable at low tides. Habitat structure may also Many animals other than humans have been play a role in the distribution and abundance marsh managers for years; however a limited of bird species because salinities and plant amount of research has been conducted to communities differ between impounded and evaluate effects of such activity. In general, unimpounded marshes. Invertebrate commu- removing animals (annual sales or culls) or nities and availability may also differ between rotating pasture lands have been effective in managed and unmanaged marshes. preventing overgrazing. In some cases, perma- Mosquito-control ditches drastically alter nent fencing of selected areas may be necessary the hydrology, hence vegetation communities, where critical species require increased protec- of the marshes and set the stage for more dra- tion (e.g., Piping Plovers [Charadrius melodus]) matic marsh transformations. Since the 1960s in the beach-marsh complexes of Chincoteague in the mid-Atlantic region, OMWM has been National Wildlife Refuge where feral horses are developed to facilitate the biological control of managed). Additional work is needed to assess mosquitoes. OMWM attempts to enhance fi sh the level of grazing and trampling that can be populations while decreasing ovipositon sites sustained by the local soil invertebrates and for mosquitoes by creating high-marsh pools native grasses and sedges before community and radial ditches isolated from daily tides. In dynamics are altered. spite of the appeal of depending upon biological Ironically, in light of the species’ importance rather than chemical means to control mosqui- as an impetus of coastal-marsh management, toes, the practice has proven controversial with recent increases in the Snow Goose in much of some marsh ecologists remaining concerned North America have been a major concern for about the mechanical alteration of marshes. state and federal wildlife managers and coastal The effect of OMWM on waterbirds has wetland managers because of their potential been studied in several locations, but relatively to damage marshes and nearby crops. Special little research has been done on a larger suite of hunts have been used to attempt to reduce these potential ecological effects that might accrue due populations; however, the effectiveness of these to OMWM treatment. Some of these potential measures is unclear. Additional research and effects are being addressed presently through monitoring are necessary to determine the effec- research projects on six national wildlife ref- tiveness of different levels of control in altering uges from Maine to Delaware (James-Pirri et al. goose populations. 2004). Additional work is needed in other areas Medium-sized fur-bearing mammals also as well over longer time frames to determine the modify marshes considerably. The native immediate versus longer-term effects of altering muskrat, however is less cause for concern in its the hydrology and structure of the marsh. With marsh plant consumption and tunneling than is the onset of sea-level rise, any additional inte- the exotic nutria. Where population levels are rior fragmentation of marshes may prove inimi- moderate for each species, the opening of small cal to a healthy marsh ecosystem. pockets in the monoculture of marsh grasses In general, larger OMWM ponds (>0.1 ha) may benefi t waterfowl, rails, and other species. and pools attract more shorebirds and water- However, nutria have caused extensive marsh fowl than do small ones, although densities fragmentation and loss, especially in Louisiana may not be greater. Several studies attempting and Maryland. Trapping no longer is viable eco- to assess bird use of ponds were compromised nomically nor is it effective in population con- due to either insuffi cient controls, or inappro- trol. An extermination program is underway in priate survey methods. One experimental study Maryland and Louisiana, and research efforts in New Jersey indicated that, although larger are underway to evaluate how population IMPACTS OF MARSH MANAGEMENT—Mitchell et al. 175 reduction rates are affecting declines and demo- and historic disturbances (fi re). We suggest that graphic aspects of the Maryland population. scaling back the use of prescribed burning by Although some evidence suggests that we reducing extent and frequency, particularly in can improve marshes for waterfowl, herons, areas in which fi re is historically not a frequent and, possibly in some cases, passerines, using disturbance, is certainly advisable, given the certain marsh-management activities, success levels of uncertainty. In a similar vein, taking is often hit or miss. Additionally, the effects on a go-slow approach on OMWM, especially in non-target organisms, particularly those that relatively pristine, unaltered coastal marshes is depend on coastal marshes for at least part of recommended. Coastal-marsh restoration, such their life cycle (e.g., endemic sparrows, rails, as ditch plugging in the Northeast and opening small mammals, snakes, and fi sh) are at best up diked marshes (Cape Cod, Delaware Bay ambiguous and at worst harmful. As a result, marshes; salt ponds; Merritt many generally abundant and widespread spe- Island, Florida) should be encouraged. cies may benefi t, whereas the few coastal-marsh A precautionary approach that uses adap- specialists probably do not. tive resource management and attempts several Nearly all studies of avian responses to experiments simultaneously to compare and coastal-marsh management document simple evaluate model parameters is well advised. We abundance or density measures that may not encourage researchers and managers to work best refl ect habitat quality (Van Horne 1983). together to monitor and evaluate management Unknown are the effects of actions on biological activities while emphasizing an experimental parameters closely related to fi tness, such as sur- approach (Ratti and Garton 1996). Such col- vival, nesting success, and physiological condi- laborations should emphasize well-designed tion, or shifts in intrinsic (e.g., foraging behavior long-term studies that document meaningful and social organization) or environmental fac- ecological responses (e.g., avian productiv- tors (food availability and predator populations) ity or nutrient cycling). Only by treating each that lead to changes in these parameters. In addi- management activity, when possible, as a fi eld tion, most studies have attempted to relate bird experiment, complete with suitable control responses local habitat features alone. Landscape treatments and true replication, can signifi cant scale variables such as area and extent of pre- advances in the science of coastal-wetland man- scribed burns, proximity of other foraging areas, agement be made. Information gleaned from food sources, open water, or emergent vegeta- these sound practices can be used to justify or tion, and habitat diversity and juxtaposition have alter coastal-marsh management activities with also been largely ignored. Longer-term effects of greater confi dence. changes in ecosystem processes (vertical accre- tion, compaction, sedimentation, and nutrient ACKNOWLEDGMENTS cycling) have also received comparatively little attention. We would like to thank the USDI Fish and Finally, given the variable nature of coastal Wildlife Service, National Wildlife Refuge marshes, we should consider the merits of con- System Region 5, Northwestern State University, tinuing to manage these habitats as we have Smithsonian Environmental Research Center, historically (occasionally achieving some objec- USGS Patuxent Wildlife Research Center, and tives) while risking potential irreversible ecosys- the University of Virginia. We also wish to tem effects, such as the loss of a coastal-marsh thank S. Droege, R. Greenberg, and two anony- endemic species. An alternative is to revise man- mous reviewers for their discerning comments agement goals and procedures to emphasize res- on previous drafts of this manuscript. toration of natural marsh processes (hydrology) Studies in Avian Biology No. 32:300–339

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