Ronald T. Kneib

The function of marine critical transition zones and the importance of sediment biodiversity

Estuaries and coastal wetlands are critical transition zones (CTZs) that link land, freshwater habitats, and the sea. CTZs provide essential ecological functions, including decomposition, nutrient cycling, and nutrient production, as well as regulation of fluxes of nutrients, water, particles, and organisms to and from land, rivers, and the ocean. Sediment-associated biota are integral to these functions. Functional groups considered essential to CTZ processes include heterotrophic bacteria and fungi, as well as many benthic invertebrates. Key invertebrate functions include shredding, which breaks down and recycles organic matter; suspension feeding, which collects and transports sediments across the sediment–water interface; and bioturbating, which moves sediment into or out of the seabed. In addition, macrophytes regulate many aspects of nutrient, particle, and organism dynamics above- and belowground. Animals moving within or through CTZs are vectors that transport nutrients and organic matter across terrestrial, freshwater, and marine interfaces. Significant threats to biodiversity within CTZs are posed by anthropogenic influences; eutrophication, nonnutrient pollutants, species invasions, overfishing, habitat alteration, and climate change affect species richness or composition in many coastal environments. Because biotic diversity in marine CTZ sediments is inherently low whereas their functional significance is great, shifts in diversity are likely to be particularly important. Species introductions (from invasion) or loss (from overfishing or habitat alteration) provide evidence that single-species changes can have overt, sweeping effects on CTZ structure and function. Certain species may be critically important to the maintenance of ecosystem functions in CTZs even though at present there is limited empirical evidence that the number of species in CTZ sediments is critical. We hypothesized that diversity is indeed important to ecosystem function in marine CTZs because high diversity maintains positive interactions among species (facilitation and mutualism), promoting stability and resistance to invasion or other forms of disturbance. The complexity of interactions among species and feedbacks with ecosystem functions suggests that comparative (mensurative) and manipulative approaches will be required to elucidate the role of diversity in sustaining CTZ functions.

Patterns of Invertebrate Distribution and Abundance in the Intertidal Salt Marsh: Causes and Questions

The intertidal marsh community comprises both benthic and natant faunal components. The benthic components are primarily small invertebrates residing within or on the soft sediments of the vegetated marsh surface. The natant components include larger, fully aquatic organisms (e.g., fish and shrimp) that inhabit the shallow waters adjacent to the marsh at low tide but interact with the benthic components of the community when the marsh is tidally inundated. In this structurally complex and often expansive intertidal environment, patterns of invertebrate distribution and abundance are not apparent to the casual observer. Benthic core samples taken along an intertidal marsh transect on Sapelo Island, Georgia, USA show that many of the inconspicuous infaunal organisms, which numerically dominate the macrofaunal elements of this soft-substrate community, exhibit zonal distribution patterns along a tidal gradient. Patterns of invertebrate distribution in the intertidal salt marsh are often attributed to the activities of aquatic predators. The results of most predator exclusion experiments have left little doubt that predation/disturbance can be an important determinant of invertebrate abundance in soft-substrate communities; but a growing number of experiments, in both freshwater and marine environments, have produced results that apparently conflict with this, general tenet. Dismissed by some as “failed” experiments, these investigations have exposed our lack of knowledge about the effects of specific predators and the importance of complex interactions which involve more than two trophic levels. Although the importance of predation has been stressed in many recent experimental investigations, there are many other factors that, alone or in combination, may also influence the structure of salt marsh invertebrate assemblages. Included among these are: (1) various density-dependent processes (e.g., adult-larval interactions, agonistic behavior, interspecific competition), (2) selective larval settlement or mortality, (3) the influence of physical factors expressed through habitat preferences, and (4) unpredictable or cyclic physical disturbances. Many questions concerning the spatial and temporal patterns of invertebrate distribution and abundance in the salt marsh are unresolved and remain as challenges to our understanding of soft-substrate community dynamics.

Early Life Stages of Resident Nekton in Intertidal Marshes

Variability in early life stages of species that are permanent residents of the estuarine nekton is poorly understood, especially in systems with extensive areas of emergent vegetation (e.g., salt marshes and mangroves). Sampling small mobile nekton in these shallow intertidal habitats presents a difficult methodological challenge. Simulated aquatic microhabitats (SAMs) were used to collect the early life stages of resident nekton that remained on the emergent marsh surface after it was exposed by the tide and could not be adequately sampled by traditional methods. Where the intertidal is a prominent areal component of the estuary, a large portion of young nekton could be overlooked using other common survey methods (e.g., plankton tows or block nets). Populations of young fishes and natant crustaceans were monitored for a year at 3-d to 6-d intervals from both low and high intertidal elevations within each of two marsh sites on Sapelo Island, Georgia, USA. Three species accounted for >99% of the 41,023 individuals collected. These were the killifishesFundulus heteroclitus (57.0%) andF. luciae (4.0%), and the daggerblade grass shrimp,Palaemonetes pugio (38.4%). YoungF. heteroclitus were used in field enclosure experiments to relate abundance data to actual areal densities. Average annual estimated density of young nekton on the surface of the intertidal marsh at low tide was 7.2 individuals m−2. Early life stages of estuarine resident species, particularly those with demersal young, are not affected by the same physical processes influencing larval supply and recruitment variability in marine-spawned species. In salt marshes, biotic factors (e.g., adult reproductive activity, predation, and food limitation) may be more important as proximate causes of variation during the early life histories of resident nekton.

Managing Critical Transition Zones

Ecosystems that function as critical transition zones (CTZs) among terrestrial, freshwater, and marine habitats are closely connected to the ecosystems adjacent to them and are characterized by a rapid flux of materials and organisms. CTZs play various roles, including mediating water flows, accumulating sediments and organic matter, processing nutrients, and providing opportunities for recreation. They are particularly difficult to manage because they tend to be small, albeit important, components of large watersheds, and managers may not have control over the entire landscape. Moreover, they are often the focus of intensive human activity. Consequently, CTZs are critically important zones, and their preservation and protection are likely to require unique collaboration among scientists, managers, and stakeholders. Scientists can learn a great deal from the study of these ecosystems, taking advantage of small size and the importance of fluxes, but a good understanding of adaptive management strategies is needed to establish a dialogue with managers and stakeholders on technical and management issues. An understanding of risk analysis is also important to help set meaningful goals and establish logical strategies that include all of the interested parties. Successful restoration of a CTZ is the best test of the quality of knowledge about its structure and function. Much has already been learned about coastal CTZs through restoration projects, and the large number of such projects involving riparian CTZs in particular suggests that there is considerable opportunity for fruitful collaborations between scientists and managers.

Efficacy of minnow traps for sampling mummichogs in tidal marshes

We evaluated two types of wire-mesh minnow traps for sampling mummichogs (Fundulus heteroclitus) from intertidal rivulets on the surface of a salt marsh. In field trials, mean catch (no. of fish trap−1) did not differ significantly between trap types nor across soak times ranging from 30 to 240 min. Catch rate (no. of fish trap−1 min−1) was significantly (K-W ANOVA on ranked data, H4df=24.79; p<0.001) greater at shorter soak times, and larger size classes (>45 mm TL) of fish were relatively more abundant (Kolmogorov-Smirnov test, p<0.0001) in collections from double-funnel cylindrical traps (volume=10,330 cm3) compared to rectangular traps having about twice the volume and a single funnel cylindrical traps (volume=10,330 cm3) compared to rectangular traps having about twice the volume and a single funnel opening. We estimated entry and escape rates by adding marked fish to traps at initial densities of 30, 60, and 120 trap−1 and measuring changes in the numbers of marked and unmarked fish in each trap type at soak times ranging from 15 to 120 min during the flood stage of the tidal cycle. Escape rate was almost, twice as great from the two-funnel cylindrical traps (0.77 fish min−1) than from the single-funnel rectangular type (0.42 fish min−1) but entry rates did not differ significantly (K-W ANOVA, H4df=0.06; p=0.813) between trap types. Escape rate increased and entry rate decreased as fish density within traps was increased. Escape rates were highest immediately following immersion of the traps, but approached a constant value after 30 min. Trap-specific rates of entry and escape can be affected by a variety of factors including physical characteristics of different trap types, method and timing of trap deployment, and fish behavior. Catch data from minnow traps are unlikely to be useful in measuring relative abundance of mummichogs among habitats unless very short soak times (≤60 min) are used and calibration tests are performed. Use of longer soak times virtually ignores the dynamics involved in determining catch, and is unlikely to provide accurate information on either abundance or size distribution of mummichogs from tidal marsh habitats.

Intertidal distribution and feeding habits of the Mud Crab, Eurytium limosum

Average summer densities of the xanthid crab,Eurytium limosum, in an intertidal salt marsh on Sapelo Island, Georgia were in the range of 7.5 to 80.0 individuals m−2. Crab densities were lowest in wet, lowlying marsh and highest in well-drained creekbank and mussel mound habitats. An analysis of crab stomach contents indicated that feeding occurred mostly around high tide, especially at night. Although the diet included some plant material,Eurytium limosum is primarily predatory and consumed crabs, polychaetes, ostracods, bivalves, and snails. In the laboratory, under simulated low-tide field conditions, both small (11–15 mm carapace width) and large (20–37 mm CW)Eurytium could capture and consume young killifish (Fundulus heteroclitus). Large crabs consumed the entire size range (7–19 mm total length) of larval/juvenile fish offered, but small crabs did not prey upon fish >11.5 mm TL. The potential importance ofE. limosum as a predator on young killifish may not be realized in the field because alternative prey are available and the crabs feed primarily at high tide, when young killifish are dispersed in the water column and are less vulnerable to benthic predators.