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2001
Levin, LA, Boesch DF, Covich A, Dahm C, Erseus C, Ewel KC, Kneib RT, Moldenke A, Palmer MA, Snelgrove P, Strayer D, Weslawski JM.  2001.  The function of marine critical transition zones and the importance of sediment biodiversity. Ecosystems. 4:430-451.   10.1007/s10021-001-0021-4   AbstractWebsite

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.

Levin, LA, Etter RJ, Rex MA, Gooday AJ, Smith CR, Pineda J, Stuart CT, Hessler RR, Pawson D.  2001.  Environmental influences on regional deep-sea species diversity. Annual Review of Ecology and Systematics. 32:51-93.   10.1146/annurev.ecolsys.32.081501.114002   AbstractWebsite

Most of our knowledge of biodiversity and its causes in the deep-sea benthos derives from regional-scale sampling studies of the macrofauna. Improved sampling methods and the expansion of investigations into a wide variety of habitats have revolutionized our understanding of the deep sea. Local species diversity shows clear geographic variation on spatial scales of 100-1000 km. Recent sampling programs have revealed unexpected complexity in community structure at the landscape level that is associated with large-scale oceanographic processes and their environmental consequences. We review the relationships between variation in local species diversity and the regional-scale phenomena of boundary constraints, gradients of productivity, sediment heterogeneity, oxygen availability, hydrodynamic regimes, and catastrophic physical disturbance. We present a conceptual model of how these interdependent environmental factors shape regional-scale variation in local diversity. Local communities in the deep sea may be composed of species that exist as metapopulations whose regional distribution depends on a balance among global-scale, landscape-scale, and small-scale dynamics. Environmental gradients may form geographic patterns of diversity by influencing local processes such as predation, resource partitioning, competitive exclusion, and facilitation that determine species coexistence. The measurement of deep-sea species diversity remains a vital issue in comparing geographic patterns and evaluating their potential causes. Recent assessments of diversity using species accumulation curves with randomly pooled samples confirm the often-disputed claim that the deep sea supports higher diversity than the continental shelf. However, more intensive quantitative sampling is required to fully characterize the diversity of deep-sea sediments, the most extensive habitat on Earth. Once considered to be constant, spatially uniform, and isolated, deep-sea sediments are now recognized as a dynamic, richly textured environment that is inextricably linked to the global biosphere. Regional studies of the last two decades provide the empirical background necessary to formulate and test specific hypotheses of causality by controlled sampling designs and experimental approaches.

1997
Levin, LA, Edesa S.  1997.  The ecology of cirratulid mudballs on the Oman margin, northwest Arabian Sea. Marine Biology. 128:671-678.   10.1007/s002270050134   AbstractWebsite

Mudball-building cirratulid polychaetes have been described previously only from the southern California margin. During a study of oxygen minimum-zone benthos in fall 1994, we observed dense aggregations of agglutinated mudballs at 840 to 875 m on the Oman margin in the northwest Arabian Sea. These were inhabited, and probably constructed, by a cirratulid polychaete species in the genus Monticellina. The mudballs were cigar-shaped, 4.5 to 25 mm long, and positioned vertically so as to protrude several millimeters above the sediment-water interface. Total mudball densities were similar to 16000 m(-2). Occupied mudballs occurred at densities of 2112 m(-2); 89% were in the uppermost 2 cm of sediment, and no occupied mudballs were found below 10 cm. Organisms other than the cirratulid were present on 1.7% of the mudballs examined, and included epizoic polychaetes, agglutinated and calcareous Foraminifera. Various polychaetes, a nemertean and nematodes were found inside tests. Mudball abundance exhibited positive associations with densities of several paraonid polychaete species, and with densities of burrowing and subsurface-deposit-feeding polychaetes. Negative associations were observed between mudballs and three tube-building taxa (two polychaetes and an amphipod). Mudball-inhabiting cirratulids are abundant in at least two low-oxygen, margin settings. We expect further sampling of bathyal environments to yield additional systems in which cirratulid mudballs are common. Such observations are valuable because mudballs appear to represent a significant source of heterogeneity that can influence macrofaunal community structure in deep-sea sediments.

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.

1995
Levin, LA, Dibacco C.  1995.  Influence of sediment transport on short-term recolonization by seamount infauna. Marine Ecology-Progress Series. 123:163-175.   10.3354/meps123163   AbstractWebsite

Rates and mechanisms of infaunal recolonization in contrasting sediment transport regimes were examined by deploying hydrodynamically unbiased colonization trays at 2 sites similar to 2 km apart on the flat summit plain of Fieberling Guyot in the eastern Pacific Ocean. Both study sites experienced strong bottom currents and high shear velocity (u* exceeding 1.0 cm s(-1) daily). Macrofaunal recolonization of defaunated sediments on Fieberling Guyot was slow relative to observations in shallow-water sediments, but rapid compared to other unenriched deep-sea treatments. Microbial colonization was slower but macrofaunal colonization was faster at White Sand Swale (WSS, 585 m), where rippled foraminiferal sands migrate daily, than at Sea Pen Rim (SPR, 635 m), where the basaltic sands move infrequently. Total densities of macrofaunal colonizers at WSS were 31 and 75% of ambient after 7 wk and 6.4 mo, respectively; at SPR they were 6 and 49% of ambient, respectively. Over 3/4 of the colonists were polychaetes (predominantly hesionids and dorvilleids) and aplacophoran molluscs. Species richness of colonizers was comparable at SPR and WSS and did not differ substantially from ambient. Most of the species (91%) and individuals (95%) recovered in colonization trays were taxa present in background cores. However, only 25% of the taxa colonizing tray sediments occurred in trays at both WSS and SPR. Sessile species, carnivores and surface feeders were initially slow to appear in colonization trays, but after 6.4 mo, colonizer feeding modes, life habits and mobility patterns mirrored those in ambient sediments at WSS and SPR. Defaunated sediments were colonized by larvae, juveniles and adults at both sites. These experiments provide the first observations of infaunal colonization on seamounts, and in deep, high-energy settings. Passive bedload transport appears to be a dominant colonization mechanism in unstable foraminiferal sands at WSS. Based on the rapid recovery of infauna in trays and low diversity at WSS, we infer that disturbance is a natural feature of this site and that the ambient fauna of WSS retains features of early succession. Infaunal colonization is slower in the stable substrate at SPR, where physical disturbance may occur much less frequently.

1994
Levin, LA, Leithold EL, Gross TF, Huggett CL, Dibacco C.  1994.  Contrasting effects of substrate mobility on infaunal assemblages inhabiting two high-energy settings on Fieberling Guyot. Journal of Marine Research. 52:489-522.   10.1357/0022240943077028   AbstractWebsite

The influence of seamount-intensified flows on the structure of infaunal assemblages was examined at two sand-covered sites located 2.3 km apart atop the summit plain of Fieberling Guyot (32-degrees 27.6'N 127-degrees 48.0'W). Both sites experience strong, tidal bottom currents with flows exceeding 20 cm/ s on a daily basis (4 mab). Estimates of shear velocity (u*) did not differ significantly between the two sites. However, differences in sediment composition and density produced different sediment transport regimes at the two sites. At Sea Pen Rim (SPR), located on the NW perimeter (635 m), sedimentary particles were composed primarily of basaltic sands that experienced negligible transport during the study period. At White Sand Swale (WSS, 580 m), a narrow valley enclosed on three sides by basalt outcrops, sediments were composed almost entirely of foraminiferal sands that moved daily. Sediment organic content and microbial abundances were similar at the two sites. Infauna (> 300 mum) had higher densities at WSS (1870/m2) than SPR (1489/m2), but lower expected species richness. Although the 2 sites shared nearly 50% of identified species, peracarid crustaceans, echinoderms, sponges, and bryozoans were proportionally more important in the stable substrates of SPR, while turbellarians, bivalves, and aplacophorans were better represented in the shifting sands of WSS. The infauna of WSS lived deeper in the sediment column (> 50% below 2 cm) than that of SPR (> 50% in the upper 1 cm), at least partly because the majority (83%) at WSS were subsurface burrowers with motile lifestyles. Tube-building and epifaunal lifestyles were more common at SPR than WSS, as were surface-deposit and filter-feeding modes. Fences and weirs were deployed at the study sites for 6.5-wk and 6-mo periods to manipulate bottom stress. Changes in faunal patterns within weirs at WSS reinforced our conjecture that contrasting sediment transport regimes explain between-site differences in community structure. Fence effects varied with deployment period and site. Topographic features on Fieberling Guyot produce heterogeneous sedimentary settings characterized by different transport regimes. Our results suggest that substrate mobility exerts primary control over infaunal community structure at the two high-energy sites.

Schaff, TR, Levin LA.  1994.  Spatial heterogeneity of benthos associated with biogenic structures on the North Carolina continental slope. Deep-Sea Research Part Ii-Topical Studies in Oceanography. 41:901-&.   10.1016/0967-0645(94)90053-1   AbstractWebsite

The objective of this study was to determine if biogenic features such as mounds, pits and tubes produce small-scale (0.1-100 m) spatial heterogeneity in macrofaunal community structure on the continental slope off North Carolina at 850 m. Macrofaunal and microbial communities associated with sediment mounds, pits and level areas were compared off Cape Lookout, North Carolina. No significant differences were found in sediment microbial counts or total macrofaunal distributions. One paraonid polychaete (Levensenia gracilis) was more abundant in pits than in the other samples, and infaunal anemones exhibited depressed densities in sediment mounds. At a second site, off Cape Hatteras, North Carolina, infaunal heterogeneity associated with the tube-building foraminiferan Bathysiphon filiformis was examined by comparing an area with high tube densities (93.8 m(-2)) with an area 100 m away without tubes. No significant differences were found in the distribution and abundances of bacteria between the two areas. The only significant difference found in infaunal densities was the presence of high numbers of reproductive oligochaetes in the 5-10 cm fraction beneath tube beds. One terebellid polychaete species (Nicolea sp.), which lives exclusively on B. filiformis tubes, was absent in the non-tube area. With a few exceptions, the biogenic structures examined at these two sites appeared to exert only minor influence on macrofaunal or microbial community structure. Within each site, slope assemblages examined in this study appeared to be homogeneous on the small scales examined.

1992
Gooday, AJ, Levin LA, Thomas CL, Hecker B.  1992.  The distribution and ecology of Bathysiphon filiformis sars and B. major de folin (Protista, Foraminiferida) on the continental slope off North Carolina. Journal of Foraminiferal Research. 22:129-146. AbstractWebsite

Two large species of the agglutinated foraminifera genus Bathysiphon are common in samples and photographs from bathyal depths on the North Carolina continental slope: B. filiformis off Cape Hatteras (588-930 m bathymetric depth) and B. major off Cape Lookout (850-1950 m depth). The sampling area, and particularly the 850 m station where B. filiformis is abundant (mean densities of 59-154 per m2), is believed to receive large inputs of organic material from various sources. This is consistent with the previously observed occurrence of large Bathysiphon species in regions of high food supply. Ten camera sled transects across the eastern U.S. continental slope between 32-degrees-N and 41-degrees-N emphasize the abundance of B. filiformis in the Cape Hatteras area compared with its rarity or absence elsewhere along the continental slope. Box cores, bottom photographs, and direct submersible observations indicate that B. filiformis tubes project above the sediment in an arcuate curve with only the lower 1 cm or so buried. Bathysiphon major adopts a similar orientation but has a greater proportion (50-80%) of the tube buried. The voluminous, dense, granular protoplasm of both species contains biogenic particles (including diatoms, in B. filiformis only), dinoflagellate cysts, fungal remains, pollen grains, tintinnid loricae, polychaete jaws and setae, benthic foraminiferal tests, and fish tooth fragments), suggesting that they feed mainly on material derived from the sediment surface. Submersible observations indicate that B. filiformis is patchily distributed at 100 m scales. Smaller scale dispersion patterns (analyzed from photographs) are generally random but with a tendency to be aggregated at lower densities and uniform at higher densities. A variety of metazoans and foraminifers live epifaunally on the outer surfaces of B. filiformis tubes. The most frequently occurring metazoans were larvae and juveniles of an unidentified gastropod and a tubiculous terebellid polychaete Nicolea sp. The most common epifaunal foraminifers were Tritaxis conica and Trochammina sp. Tubes of B. major, however, are virtually devoid of epifauna. Our results support the view that large, agglutinated rhizopod tests may influence the structure of deep-water benthic communities. However, in the case of Bathysiphon on the North Carolina continental slope, the effect appears limited to taxa directly associated with the foraminiferal tubes.