The influence of leaf litter on zooplankton in floodplain wetlands: changes resulting from river regulation

1. The loss of input of leaf litter through clearing of riparian vegetation may result in significant changes to aquatic ecosystems. River red gums (Eucalyptus camaldulensis) surrounding floodplain wetlands in the Murray–Darling Basin, Australia, contribute large quantities of leaf litter, but the quality of this resource may change depending on the timing of inundation. 2. We used experimental mesocosms to test the hypotheses that zooplankton would have a greater abundance with an input of leaf litter and that fewer zooplankton would emerge from egg banks in cleared than forested wetlands. The experiment was carried out in summer/autumn and in spring to test a third hypothesis that zooplankton would respond to changes in the timing of wetland inundation as a result of river regulation. 3. In summer/autumn, leaf litter reduced zooplankton abundance by 89% at the beginning of the experiment through its influence on water quality. Only a few taxa (Polyarthra spp., Colurella spp. and the cladoceran Family Moinidae) responded positively to leaf litter when water quality improved later in the experiment, indicating a switch in the role of leaf litter from a non‐trophic to a trophic pathway. 4. In spring, microcrustaceans emerged in smaller numbers from sediment sourced from cleared compared to forested wetlands, reflecting different communities in these two wetland types and/or disturbances to the sediment that interfere with emergence. 5. Although leaf litter appears not to be an important resource for zooplankton in floodplain wetlands, riparian clearing may have lasting effects on future emerging zooplankton communities. Additionally, river regulation may have considerable impacts on the influence of leaf litter on zooplankton, which has implications for the management of floodplain river systems.     

Benthic diatom composition in isolated forested wetlands subject to drying: Implications for monitoring and assessment

The development of bioindicators for wetlands, especially ephemerally hydrated depressional and isolated wetlands, can be problematic because of seasonal changes in hydrology and target indicator organism biology. To determine if benthic diatoms could be used as a multi-season biological indicator of wetland condition in isolated forested wetlands of Florida, USA, 11 wetlands were sampled twice during a 5-month period, once when dry, then again when hydrated. Sites sampled when dry had significantly higher diatom taxa richness at genus and species levels. Non-metric multidimensional scaling and multiple response permutation process analyses resulted in no obvious or significant wet/dry grouping of species or genus level abundance data. Five of seven diatom metrics of the Florida Wetland Condition Index (FWCI) for depressional forested wetlands were significantly linearly correlated (p < 0.05), while only one of seven metrics (a dissolved oxygen indicator) had a significantly different mean in paired t-test analyses. The final FWCI was significantly correlated (Pearson's r = 0.85, p < 0.001) between wet and dry sites, and no difference was found in mean FWCI score between wet and dry sites (t = −1.98, p = 0.076), suggesting that with additional research, benthic diatoms may be used to monitor and assess wetland condition regardless of season or site hydrologic conditions.     

Assessment of Elk Highway Permeability by Using Global Positioning System Telemetry

Abstract: Highways have significant direct and indirect impact on natural ecosystems, including wildlife barrier and fragmentation effects, resulting in diminished habitat connectivity and highway permeability. We used Global Positioning System (GPS) telemetry to assess Rocky Mountain elk (Cervus elaphus nelsoni) permeability across a 30-km stretch of highway in central Arizona, USA, currently being reconstructed with 11 wildlife underpasses, 6 bridges, and associated ungulate-proof fencing. The highway was reconstructed in phases, allowing for comparison of highway crossing and passage rates during various stages of reconstruction. We instrumented 33 elk (25 F, 8 M) with GPS receiver collars May 2002 to April 2004. Our collars accrued 101,506 GPS fixes with 45% occurring within 1 km of the highway. Nearly 2 times the proportion of fixes occurred within 1 km of the highway compared with random. We think elk were attracted to the highway corridor by riparian—meadow foraging habitats that were 7 times more concentrated within the 1-km zone around the highway compared with the mean proportion within elk use areas encompassing all GPS fixes. Elk crossed the highway 3,057 times; crossing frequency and distribution along the highway were aggregated compared with random. Crossing frequency within 0.16-km highway segments was negatively associated with the distance to riparian—meadow habitats (r_s = -0.714, n = 190, P < 0.001). Mean observed crossing frequency (92.6 ± 23.5 [SE] crossings/elk) was lower than random (149.6 ± 27.6 crossings/elk). Females crossed 4.5 times as frequently as males. Highway permeability among reconstruction classes was assessed using passage rates (ratio of highway crossings to approaches); our overall mean passage rate was 0.67 ± 0.08 crossings per approach. The mean passage rate for elk crossing the highway section where reconstruction was completed (0.43 ± 0.15 crossings/approach) was half that of sections under reconstruction and control sections combined (0.86 ± 0.09 crossings/approach). Permeability was jointly influenced by the size of the widened highway and associated vehicular traffic on all lanes. Crossing frequency was used to delineate where ungulate-proof fencing yielded maximum benefit in intercepting and funneling crossing elk toward underpasses, promoting highway safety. Use of passage rates provides a quantitative measure to assess permeability, conduct future pre- and postconstruction comparisons, and to develop mitigation strategies to minimize highway impacts to wildlife.     

Limits and effects of invasion by the nonindigenous wetland plant <Emphasis Type="Italic">Lythrum salicaria</Emphasis> (purple loosestrife): a seed bank analysis

We used seed bank analyses to investigate the role of dispersal in limiting invasion by Eurasian Lythrum salicaria within and among North American wetlands, and the changes in seed bank diversity associated with this invader. We compared the number and species composition of seedlings emerging from soil sampled in 11 uninvaded wetlands and paired uninvaded and invaded sites within 10 invaded wetlands under both seedling competition and noncompetitive conditions. Almost no L. salicaria emerged in samples from uninvaded wetlands, indicating dispersal limitation despite prodigious seed production in nearby wetlands. However L. salicaria emerged in all samples from uninvaded sites in invaded wetlands, suggesting environmental limits on establishment within invaded wetlands. Conditions that provided opportunities for seedlings to compete reduced survival of Typha spp. but not L. salicaria seedlings. However, this was due to species-specific differences in post-emergence mortality rather than response to competition. Competition did reduce seedling mass, but this effect did not differ among species. Species richness of emerging seedlings was lower for invaded than uninvaded wetlands. Lower seed bank richness may be a cause or consequence of L. salicaria invasion. Efforts to reduce seed dispersal to uninvaded wetlands would likely slow the spread of this invader.     

Re-establishment of epibiotic communities in reforested mangroves of Gazi Bay, Kenya

Recolonization of epibiotic flora and fauna in two fringing Sonneratia alba reforestation plots was investigated and compared to a natural mangrove stand and a denuded site in Gazi Bay, Kenya. The reforested sites differed with respect to land history and planting density. Habitat availability in the form of pneumatophore surface differed among forested sites (P<0.001), and between landward and seaward zones (P<0.05). Eighteen algal species were found in the natural area compared to 23 and 10 in replanted sites. Only one species was encountered in the denuded area. SIMPER analysis distinguished Enteromorpha ramulosa, Polysiphonia sp., Hypnea sp. and Caloglossa leprieuri as the main algal species responsible for differences between sites. Algal biomass was positively correlated to pneumatophores area (P<0.001). Total algal biomass differed markedly between forested sites: 1.4 (matrix replantation), 28.6 (natural stand) and 44.3 g m⁻² (integrated replantation) in the seaward zones. The matrix replantation showed strong differences in algal community assemblages compared to the other forested sites, and this site also had significantly lower biomass of sessile benthic fauna (P<0.001). Statistical differences in algal (P<0.01) and sponge (P<0.05) community composition between landward and seaward zones were observed in all sites and trunk fouling fauna was distinctly different between sites. Reasons for the above patterns are discussed and it is suggested that zonation patterns affecting pneumatophore surface and inundation time, in combination with proximity of sites to natural seeding areas, are the most likely explanations for observed patterns of epibiotic community distribution in this study.     

The impact of small chironomid grazers on epiphytic algal abundance and dispersion

• 1 A combination of field and laboratory experiments was used to assess the impact of chironomid grazers on taxonomic composition, abundance and dispersion of epiphytic algal assemblages.
• 2 In the laboratory, Psectrodadius sp. reduced the biovolume of algal species preferred as food and increased the degree of clumping of non-preferred species. Thienemanniella cf. fusca had both positive and negative effects (depending on the algal species) on the biovolumes of algal species preferred as food and increased the degree of clumping of non-preferred species.
• 3 In field exclosures, no effect of removal of chironomid larvae from the grazer assemblage could be detected in autumn or winter experiments. A third, longer removal experiment, conducted in summer, resulted in increased biovolumes of edible Cosmarium spp. and Aphanocapsa spp., preferred foods of chironomid larvae. Biovolumes of Lyngbya sp., Eulbochaete spp. and Oedogortium spp., filamentous taxa used extensively in larval case construction, also increased. Chironomid larvae had no effect on total algal biovolume or biovolume of large unicellular algae.
• 4 Chironomid larvae can influence epiphytic algal assemblages through selective grazing by reducing the biovolumes of preferred foods and through case-building activity by reducing the biovolumes of construction materials.

     

Patterns of Distribution and Environmental Correlates of Macroalgal Assemblages and Sediment Chlorophyll A in Oregon Tidal Wetlands

Algae have important functional roles in estuarine wetlands. We quantified differences in macroalgal abundance, composition and diversity, and sediment chl a and pheophytin a (pheo a) among three National Wetlands Inventory (NWI) emergent marsh classes in four Oregon estuaries spanning a range of riverine to marine dominance. We also assessed the strength of macroalgal‐vascular plant associations and the degree to which environmental variables correlated with algal community metrics in marsh and woody wetlands. The frequency of occurrence of most macroalgal genera, total benthic macroalgal cover, macroalgal diversity, and sediment chl a content were several times higher in low emergent marsh than in high marsh or palustrine tidal marsh. Conversely, pheo a: chl a ratios were highest in high and palustrine marsh. Attached macroalgae (Fucus and Vaucheria) were strongly associated with plants common at lower tidal elevations such as Sarcocornia perennis and Jaumea carnosa; Ulva (an unattached alga) was not strongly associated with any common low marsh plants. In structural equation models, intertidal elevation was the most influential predictor of macroalgal cover and richness and chl a; light availability and soil salinity played secondary roles. Although common taxa such as Ulva spp. occurred across a broad range of salinities, wetlands with oligohaline soils (salinity < 5) had the lowest macroalgal diversity and lower sediment chl a. These types of baseline data on algal distributions are critical for evaluating the structural and functional impacts of future changes to coastal estuaries including sea‐level rise (SLR), altered salinity dynamics, and habitat modification.     

Lake Kinneret phytoplankton: Response to N and P enrichments in experiments and in nature

A series of water samples from Lake Kinneret was supplemented with 100 µM N (as NH₄ or as NO₃ and/or 10 µM orthophosphate-P. The yield of phytoplankton both as chlorophyll and in cell numbers of major species was determined after a two-week incubation. During these experiments, some of the algae present initially never multiplied (e.g. Peridinium and Peridiniopsis spp. Cryptomonas spp., Rhodomonas spp. and Crysochromulina parva); others e.g. Anomoeoneis exilis, Synedra sp., Chlamydomonas sp., Elakatothrix gelatinosa), undetected in the original sample, grew out during the incubation. Chlorophyte species (the majority of commonly observed forms in the lake) responded most readily to added nutrients.The results of these enrichment experiments were related to the long-term record of phytoplankton populations observed in the lake and suggest that through summer and fall, when ambient levels of both P and N are minimal, P was generally, but not always, the most limiting nutrient for algal growth. In the spring, after the decline of the Peridinium bloom, P appeared to be limiting the growth of Chlorophyta. Although most algal species grew equally well on NH₄ or NO₃, some species appeared to respond preferentially either to the former (Coelastrum, Chodatella) or to the latter (Chroococcus, Anomoeoneis) source of N.     

Pelagic Ciliates in a Large Mesotrophic Lake: Seasonal Succession and Taxon‐Specific Bacterivory in Lake Constance

The taxonomic composition of the ciliate assemblage and their taxon‐specific bacterial grazing rates in Lake Constance were investigated over the course of one year. Bacterial grazing rates were measured using natural fluorescently labelled bacteria (FLB) and compared to bacterial production. Small species such as Balanion planctonicum/Urotricha furcata and Rimostrombidium spp./Halteria sp. were the most numerous ciliates on the annual average. Larger ciliates such as Rimostrombidium lacustris and Limnostrombidium spp. contributed significantly to total ciliate biomass, but were relatively unimportant as bacterial grazers. Per capita ingestion rates ranged from 0–194 bacteria ciliate⁻¹ h⁻¹ and changed seasonally up to a hundredfold within a given taxon. Approximately 1% of the bacterial production were removed by the ciliate community on the annual average. (© 2004 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Seed viability and seed dormancy of non-native Phragmites australis in suburbanized and forested watersheds of the Chesapeake Bay, USA

The non-native, invasive haplotype of Phragmites australis is rapidly invading tidal and non-tidal wetlands across North America. Phragmites has the potential to spread by seeds and rhizomes. Seed viability and dormancy differences were quantified among 18 patches of non-native Phragmites in subestuarine wetlands in developed (i.e., suburbanized) vs. forested watersheds of the Chesapeake Bay. We used tetrazolium and germination assays to assess seed viability and compared germination percentages and rate of germination among fresh seeds, cold–moist treated seeds, and warm–dry treated seeds to evaluate seed dormancy. Seed viability was <1% in most patches but a few patches produced abundant viable seeds (5–21%). Seed viability, however, did not differ significantly between wetlands in forested vs. developed watersheds. Contrary to studies of Phragmites seed dormancy in European populations, some Phragmites seeds were dormant at maturity; cold–moist treated seeds germinated faster and to higher percentages than fresh seeds or warm–dry treated seeds.     

The Effects of pH on a Periphyton Community in an Acidic Wetland, USA

Despite the importance of peatlands, the algal ecology of peatlands and the periphyton communities which are abundant in these habitats are relatively understudied. We performed an in situ manipulation of pH in an intermediate fen in northern lower Michigan in order to examine how hydrogen ion concentrations structure an epiphytic algal community. Levels of pH were manipulated in enclosures from the control level (pH = 5) to an acid treatment (pH = 4) by adding H₂SO₄ and a neutral treatment (pH = 7) by adding NaOH. Algal communities growing on sections of Chamaedaphne calyculata (L.) Moench stems were examined after 22 days of colonization. Chlorophyll a concentration was significantly greater only in the acid treatment (~5.5 mg m⁻²) relative to the control (~3.5 mg m⁻²). Taxa richness was lower in the acid treatment. The algal assemblages were dominated by filamentous green algae and a filamentous taxon, Mougeotia spp., was significantly greater in the acid treatment relative to the control. Increases in Zygnemataceae and Oedogonium spp. most likely account for the higher chlorophyll a in the acid treatment. Most treatment differences were detected in the neutral treatment, including increased abundances of Closterium polystichum Nygaard, Cosmarium sp., Peridinium inconspicuum Lemmermann, and Synedra acus Kütz. Unexpectedly, there was no strong response of the desmid community. These data can be informative in the development of algal monitoring programs in peatlands when assessment of acidification is desired.     

The invasive green alga Avrainvillea sp. transforms native epifauna and algal communities on a tropical hard substrate reef

Some introduced species compete directly with native species for resources and their spread can alter communities, while others do not proliferate and remain benign. This study compares community structure and diversity in adjacent areas dominated by the introduced alga Avrainvillea sp. or native algal species on a hard substrate reef. The biomass and species composition of 15 paired plots (30 in total, plot type based on dominance of Avrainvillea sp. or native species) were quantified. Plots dominated by Avrainvillea sp. had a significantly different assemblage of species characterized by lower algal diversity, mostly Dictyota spp. and Laurencia sp., and a higher abundance and diversity of invertebrates, such as small arthropods, polychaetes, and brittlestars. These results suggest that as Avrainvillea sp. becomes more abundant on hard substrate reefs, it will engineer a different community composed of algal epiphytes and an invertebrate assemblage more typically associated with algae in soft sediments.     

Planktonic food chains of a highly humic lake

The development and metabolism of the plankton of a highly humic lake were followed over the vernal primary production maximum. The study was made in a mesocosm in which large filter feeders, typical of this lake in summer, were absent. During the rising phase of phytoplankton, the community was predominantly autotrophic. The most important constituents in the algal biomass were a dinoflagellate, Gymnodinium sp. (40–50%), and a prasinophycean, Scourfieldia cordiformis (7%). The biomasses of Chlamydomonas spp. and Chrysococcus spp. reached their maxima a few days later and Cryptomonas sp. became most abundant at the end of the experiment. After the phytoplankton maximum, about one week from the beginning ofthe experiment, grazing of algae by phagotrophic protozoans and phosphate depletion led to a rapid decrease of algal biomass and the community became predominantly heterotrophic. In spite of a large variation in algal biomass and primary production, the biomass of bacteria remained of the same order of magnitude as in algae both before and after the algal maximum. Bacteria were mostly responsible for the plankton respiration, which also showed no dependence on primary production. Since exudation by phytoplankton was also low, the nutrition of bacterioplankton was probably mainly based on allochthonous dissolved organic matter rather than or primary production. Thus the production of bacteria was an additional food source for higher trophic levels along with phytoplankton. Because filter feeding zooplankton was absent in the experiment, protozoans were the only grazers utilizing algae and bacteria. Essentially all growth of bacteria was used by bacterivores.     

小兴安岭天然森林沼泽湿地生态系统碳源/汇

采用静态箱-气相色谱法与相对生长方程法,同步测定小兴安岭7种天然沼泽湿地(草丛沼泽-C、灌丛沼泽-G、毛赤杨沼泽-M、白桦沼泽-B、落叶松苔草沼泽-L-T、落叶松藓类沼泽-L-X、落叶松泥炭藓沼泽-L-N)的土壤呼吸(CO_2、CH_4)净碳排放量、植被年净固碳量,并依据生态系统净碳收支平衡,揭示温带天然沼泽湿地的碳源/汇作用规律。结果表明:(1)7种天然沼泽CH_4年通量(0.006—7.756 mg m~(-2)h~(-1))呈M(高于其他类型1.0—1291.7倍,P0.05)C、G、B(高于针叶林沼泽17.7—649.0倍,P0.05)针叶林沼泽变化趋势,其季节动态存在3种类型(C、G单峰型、M、B多峰型及针叶林沼泽排放与吸收交替型);(2)其CO_2年通量(157.40—231.06 mg m~(-2)h~(-1))呈G(高于森林沼泽28.7%—46.8%,P0.05)C(高于森林沼泽7.4%—22.5%,P0.05)森林沼泽的变化趋势,其季节动态存在2种类型(C、G、L-X和L-N双峰型和M、B、L-T单峰型);(3)C、G、M、B、L-N CH_4排放仅受0—40 cm不同土壤层温度所控制;7种天然沼泽土壤CO_2排放均受气温及0—40 cm不同土壤层温度所控制,但B、L-X、L-N受温度与水位综合控制;(4)其植被年净固碳量((2.05±0.09)—(6.75±0.27)t C hm-2a-1)呈C(高于其他类型65.4%—229.3%,P0.05)G、B、L-T、L-X、L-N(高于M 80.0%—99.0%,P0.05)M变化趋势;(5)7种天然沼泽的碳源/汇(-2.32—2.09 t C hm-2a-1)作用不同,C、B和L-N为碳吸收汇(C强汇、B和L-N弱汇),M、G、L-T和L-X则为碳排放源(M、G强源、L-T和L-X弱源)。因此,温带小兴安岭草丛沼泽为碳强汇、灌丛沼泽为碳强源、森林沼泽基本维持碳平衡(除M外)。     

Macroalgal decomposition: Laboratory studies with particular regard to microorganisms and meiofauna

The microbial degradation of North Sea macroalgae was studied in laboratory microcosms, containing autoclaved seawater and a mixture of equal parts of air-driedDelesseria sanguinea, Ulva lactuca, andLaminaria saccharina (red, green and brown algae, respectively). To determine the influence of different organisms on the decomposition rate (expressed in terms of algal dry weight loss relative to the material present at time zero) and their development during decomposition processes, yeast, flagellates, ciliates, nematodes and a harpacticoid copepod species were introduced to the microcosms. Results show that microbial degradation compared to the controls was enhanced in the presence of non-axenic nematodes (Monhystera sp.) and protozoans, including bacterivorous ciliates (Euplotes sp. and aUronema-like sp.) and flagellates. No enhancement occurred with yeast (Debaryomyces hansenii) or with the harpacticoid copepodTisbe holothuriae. The most rapid algal dry weight loss (78.7% after 14 d at 18°C) occurred with the addition of raw seawater sampled near benthic algal vegetation and containing only the natural microorganisms present. These consisted mainly of bacteria with different morphological properties, whereby their numbers alone (viable counts) could not be correlated with algal dry weight loss. Although no single dominant species could be determined, lemon yellow pigmented colonies were frequently found. During decomposition in all microcosms the formation of algal particles 40–400 μm was observed, which were rapidly colonized by the other organisms present.     

Biological control of the wetlands weed purple loosestrife (Lythrum salicaria) in the Pacific northwestern United States

Purple loosestrife (Lythrum salicaria) is an Eurasian perennial hydrophyte that has become naturalized in wetlands and in and along waterways throughout temperate North America. The ecological integrity of such areas is threatened by rapidly forming monotypic infestations that displace valued flora and diminish critical fish and wildlife habitat. The inability of physical, cultural, and chemical methods to provide adequate control of the weed has led to the development of an insect-based biological control program. The first field releases of the bud and leaf feeding beetles, Galerucella calmariensis and G. pusilla, and a root-mining weevil, Hylobius transversovittatus, were made in the United States and Canada in 1992. A total of 4740 Galerucella spp. adults were released in central Washington during 1992 and 1993 at eight sites and 471 H. transversovittatus egg inoculations were made in 1993 at three locations. Establishment of both Galerucella spp. was confirmed and Hylobius colonization was achieved.     

Algal richness and life‐history strategies are influenced by hydrology and phosphorus in two major subtropical wetlands

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The permeability of highway in Gorski kotar (Croatia) for large mammals

The highway from Zagreb to Rijeka stretches 68.5 km through a wildlife core area in Gorski kotar (Croatia). It has 43 viaducts and tunnels, and one specifically constructed (100 m wide) green bridge (Dedin). One quarter of the total highway length consists of possible crossing structures. At Dedin green bridge, a total of 12,519 crossings have been recorded during 793 different days of active infrared monitors being in operation, or 15.8 crossings per day. Two monitored tunnel overpasses had 11.2 and 37.0 crossings per day, respectively, whilst 4.3 crossings occurred per day under one monitored viaduct. Of those crossings, 83.2% were by ungulates and 14.6% by large carnivores. Radio-tracked large carnivores, brown bear (Ursus arctos), grey wolf (Canis lupus) and Eurasian lynx (Lynx lynx), expressed strong positive selection for tunnels and viaducts, whilst avoiding small underpasses or bridges. Selection for the use of Dedin green bridge was equal to its availability. We conclude that this green bridge, constructed as a measure to mitigate the negative effects of the studied highway, served its purpose acceptably. Territorial and dispersing radio-tracked large carnivores crossed the highway 41 times, using both sides of the highway as parts of their home ranges. Overall, the highway in Gorski kotar does not seem to be a barrier. This demonstrates that it is possible to maintain habitat connectivity during the process of planning the highway route.     

Consideration of Geography and Wetland Geomorphic Type in the Development of Great Lakes Coastal Wetland Bird Indicators

We examined how geographic distribution of birds and their affinities to three geomorphic wetland types would affect the scale at which we developed indicators based on breeding bird communities for Great Lakes coastal wetlands. We completed 385 breeding bird surveys on 222 wetlands in the US portion of the basin in 2002 and 2003. Analyses showed that wetlands within two ecoprovinces (Laurentian Mixed Forest and Eastern Broadleaf Forest) had different bird communities. Bird communities were also significantly different among five lakes (Superior, Michigan, Huron, Erie, and Ontario) and among three wetland types (lacustrine, riverine, barrier-protected). Indicator values illustrated bird species with high affinities for each group (ecoprovince, lake, wetland type). Species with restricted geographic ranges, such as Alder and Willow Flycatchers (Empidonax alnorum and E. traillii), had significant affinities for ecoprovince. Ten bird species had significant affinities for lacustrine wetlands. Analyses on avian guild metrics showed that Lake Ontario wetlands had fewer long-distant migrants and warblers than other lakes. Numbers of short-distant migrants and total individuals in wetlands were higher in the Eastern Broadleaf Forest ecoprovince. Number of flycatchers and wetland obligate birds were not different among provinces, lakes, or wetland type. One potential indicator for wetland condition in Great Lakes wetlands, proportion of obligate wetland birds, responded negatively to proportion of developed land within 1 km of the wetland. We conclude that, although a guild approach to indicator development ameliorates species-specific geographic differences in distribution, individual species responses to disturbance scale will need to be considered in future indicator development with this approach.