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2015
Nordstrom, MC, Demopoulos AWJ, Whitcraft CR, Rismondo A, McMillan P, Gonzalez JP, Levin LA.  2015.  Food web heterogeneity and succession in created saltmarshes. Journal of Applied Ecology. 52:1343-1354.   10.1111/1365-2664.12473   AbstractWebsite

Ecological restoration must achieve functional as well as structural recovery. Functional metrics for re-establishment of trophic interactions can be used to complement traditional monitoring of structural attributes. In addition, topographic effects on food web structure provide added information within a restoration context; often, created sites may require spatial heterogeneity to effectively match structure and function of natural habitats. We addressed both of these issues in our study of successional development of benthic food web structure, with focus on bottom-up-driven changes in macroinvertebrate consumer assemblages in the saltmarshes of the Venice Lagoon, Italy. We combined quantified estimates of the changing community composition with stable isotope data (C-13:C-12 and N-15:N-14) to compare the general trophic structure between created (2-14years) marshes and reference sites and along topographic elevation gradients within saltmarshes. Macrofaunal invertebrate consumers exhibited local, habitat-specific trophic patterns. Stable isotope-based trophic structure changed with increasing marsh age, in particular with regard to mid-elevation (Salicornia) habitats. In young marshes, the mid-elevation consumer signatures resembled those of unvegetated ponds. The mid-elevation of older and natural marshes had a more distinct Salicornia zone food web, occasionally resembling that of the highest (Sarcocornia-dominated) elevation. In summary, this indicates that primary producers and availability of vascular plant detritus structure consumer trophic interactions and the flow of carbon. Functionally different consumers, subsurface-feeding detritivores (Oligochaeta) and surface grazers (Hydrobia sp.), showed distinct but converging trajectories of isotopic change over time, indicating that successional development may be asymmetric between brown' (detrital) guilds and green' (grazing) guilds in the food web.Synthesis and applications. Created marsh food webs converged into a natural state over about a decade, with successional shifts seen in both consumer community composition and stable isotope space. Strong spatial effects were noted, highlighting the utility of stable isotopes to evaluate functional equivalence in spatially heterogeneous systems. Understanding the recovery of functional properties such as food web support, and their inherent spatial variability, is key to planning and managing successful habitat restoration. Created marsh food webs converged into a natural state over about a decade, with successional shifts seen in both consumer community composition and stable isotope space. Strong spatial effects were noted, highlighting the utility of stable isotopes to evaluate functional equivalence in spatially heterogeneous systems. Understanding the recovery of functional properties such as food web support, and their inherent spatial variability, is key to planning and managing successful habitat restoration.

2002
Levin, LA, Talley TS.  2002.  Natural and manipulated sources of heterogeneity controlling early faunal development of a salt marsh. Ecological Applications. 12:1785-1802.   10.2307/3099938   AbstractWebsite

Ecosystem recovery following wetland restoration offers exceptional opportunities to study system structure, function, and successional processes in salt marshes. This study used observations of natural variation and large-scale manipulative experiments to test the influence of vascular vegetation and soil organic matter on the rate and trajectory of macrofaunal recovery in a southern California created salt marsh, the Crown Point Mitigation Site. During the first three years following marsh establishment, macrofaunal density and species richness recovered rapidly within the Spartina foliosa (cordgrass) zone; densities in the created marsh were 50% of those in the natural marsh after 16 mo and 97% after 28 mo. However, the early successional assemblage had a lower proportion of tubificid and enchytraeid oligochaetes, and a higher proportion of chironomids and other insect larvae than did the mature natural marsh. Most of the colonizers arrived by rafting on sea grass and algae rather than by larval dispersal. Initial planting of S. foliosa had no influence on macrofaunal recovery, perhaps because of variable transplant survival. However, subsequently, both positive and negative correlations were observed between densities of some macrofaunal taxa and shoot densities of S. foliosa or Salicornia spp. (pickleweed). Salinity and measures of soil organics (belowground biomass, combustible organic matter, and chlorophyll a) also were correlated with macrofaunal densities and taxon richness. Of foul added soil amendments (kelp, alfalfa, peat, and Milorganite), Milorganite (a sewage product) and kelp both promoted macrofaunal colonization during year 1, but effects were short lived. The most significant sources of heterogeneity in the recovering marsh were associated with site history and climate variation. Faunal recovery was most rapid in highly localized, organic-rich marsh sediments that were remnants of the historical wetland. Elevated sea level during the 1998 El Nino corresponded with similarity of macrofaunal communities in the created and natural marshes. The large spatial scale and multi-year duration of this study revealed that natural sources of spatial and temporal heterogeneity may exert stronger influence on faunal succession in California wetlands than manipulation of vegetation or soil properties.

1999
Talley, TS, Levin LA.  1999.  Macrofaunal succession and community structure in Salicornia marshes of southern California. Estuarine Coastal and Shelf Science. 49:713-731.   10.1006/ecss.1999.0553   AbstractWebsite

Lack of basic understanding of ecosystem structure and function forms a major impediment to successful conservation of coastal ecosystems. This paper provides a description of the fauna and examines faunal succession in Salicornia-vegetated sediments of southern California. Environmental attributes (vegetation and sediment properties) and macrofaunal (animals greater than or equal to 0.3 mm) community structure were examined in sediments of five natural, southern California Salicornia spp. marshes (Tijuana Estuary, San Diego Bay, Mission Bay, Upper Newport Bay and Anaheim Bay) and in created Salicornia marshes 16 months to 10 years in age, located within four of the bays. Oligochaetes and insects were the dominant taxa in both natural (71 to 98% of total fauna) and created (91 to 97%) marshes. In San Diego, Newport and Anaheim Bays, macrofaunal densities were generally higher in the created marshes (88 000 to 290 000 ind m(-2)) than in their natural counterparts (26 000 to 50 000 ind m(-2)). In the youngest system, Mission Bay, the reverse was true (natural: 113 000 vs created: 28 000 ind m-2). Similar species numbers were recorded from the created and adjacent natural marshes. Insects, especially chironomids, dolichopodids, and heleids, as well as the naidid oligochaete, Paranais litoralis, characterize early successional stages. Enchytraeid and tubificid oligochaetes reflect later succession evident in natural and older created marshes. Sediment organic matter (both combustible and below-ground plant biomass) was the environmental variable most commonly associated with densities of various macrofaunal taxa. These relationships were generally negative in the natural marshes and positive in the created marshes. Within-bay comparisons of macrofauna from natural Salicornia- vs Spartina-vegetated habitat in San Diego and Mission Bays revealed lower macrofaunal density (San Diego Bay only), proportionally fewer oligochaetes and more insects, and no differences in species richness in the Salicornia habitat. The oldest created Salicornia marsh (San Diego Bay) exhibited an assemblage intermediate in composition between those of the natural Salicornia- and Spartina-vegetated marshes. These results suggest: (a) faunal recovery following Salicornia marsh creation can require 10 or more years, (b) high macrofaunal variability among bays requires marsh creation reference site selection from within the same bay, and (c) Spartina-based research should not be used for Salicornia marsh management decisions. (C) 1999 Academic Press.

1996
Levin, LA, Talley D, Thayer G.  1996.  Succession of macrobenthos in a created salt marsh. Marine Ecology-Progress Series. 141:67-82.   10.3354/meps141067   AbstractWebsite

Early succession of macrofauna was examined over several years in a created Spartina alterniflora marsh located on the Newport River Estuary, North Carolina, USA. Epifauna and infaunal community structure and composition were compared at 2 elevations in plots planted with S. alterniflora, plots left bare of vegetation and vegetated plots in a nearby natural S, alterniflora marsh. No significant successional differences were observed between vegetated and unvegetated sediments in the created marsh. The earliest stages of colonization involved recruitment by opportunistic estuarine polychaetes: Streblospio benedicti, Capitella spp, and Polydora cornuta. Capitella spp. dominated the macrofauna a month after marsh creation, but thereafter S. benedicti was the most abundant species. During the first few years, the artificial marsh retained early successional characteristics, with S, benedicti, Capitella spp. and turbellarians accounting for 75 to 95% of the total macrofauna. Fiddler crabs were common epifaunal colonists. After 4 yr, species richness increased and dominance by the early colonists diminished. Taxa lacking planktonic larvae and swimming adults were particularly slow to recover in the created marsh, but accounted for over 25% of the infauna by Year 4. Oligochaetes, which comprised over 50% of the fauna in the natural marsh, remained absent or rare in the artificial system throughout the study. Infaunal recovery appears to be more rapid in lower than upper marsh elevations. Although macrofaunal densities and species richness of sediments in the lower created marsh came to resemble those of the natural marsh within 6 mo, species composition and faunal feeding modes did not. These observations suggest there may be significant functional differences between young artificial marshes and older natural marshes. Consideration of the timing of marsh creation, marsh configuration, continuity with natural marshes, seeding of taxa with poor dispersal, and attention to species habitat requirements are recommended to accelerate infaunal colonization of created Spartina marshes.