Publications

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2016
Burkett, AM, Rathburn AE, Perez ME, Levin LA, Martin JB.  2016.  Colonization of over a thousand Cibicidoides wuellerstorfi (foraminifera: Schwager, 1866) on artificial substrates in seep and adjacent off-seep locations in dysoxic, deep-sea environments. Deep-Sea Research Part I-Oceanographic Research Papers. 117:39-50.   10.1016/j.dsr.2016.08.011   AbstractWebsite

After-1 yr on the seafloor at water depths of similar to 700 m on Hydrate Ridge in the Pacific, eight colonization experiments composed primarily of a plastic mesh cube (from here on refered to as SEA(3), for Seafloor Epibenthic Attachment Cubes) were colonized by 1076 Cibicidoides wuellerstorfi on similar to 1841 cm(2) of experimental substrate. This species is typically considered an indicator of well-oxygenated conditions, and recruitment of such large numbers in bottom waters with low dissolved oxygen availability (0.24-0.37 mL/L) indicate that this taxon may not be as limited by oxygen as previously thought. Clues about substrate preferences were evident from the distribution, or lack thereof, of individuals among plastic mesh, coated steel frame, wooden dowels and reflective tape. Abundance, individual size distributions within cage populations and isotopic biogeochemistry of living foraminifera colonizing experimental substrates were compared between active seep and adjacent off seep experiment locations, revealing potential differences between these environments. Few studies have examined foraminiferal colonization of hard substrates in the deep-sea and to our knowledge no previous study has compared foraminiferal colonization of active seep and off-seep substrates from the same region. This study provides initial results of recruitment, colonization, geochemical and morphological aspects of the paleoceanographically significant species, C. wuellerstorfi, from dynamic deep-sea environments. Further experimental deployments of SEA(3)s will provide a means to assess relatively unknown ecologic dynamics of important foraminiferal deep-sea species.

2014
Cook, GS, Parnell PE, Levin LA.  2014.  Population connectivity shifts at high frequency within an open-coast marine protected area network. Plos One. 9   10.1371/journal.pone.0103654   AbstractWebsite

A complete understanding of population connectivity via larval dispersal is of great value to the effective design and management of marine protected areas (MPA). However empirical estimates of larval dispersal distance, self-recruitment, and within season variability of population connectivity patterns and their influence on metapopulation structure remain rare. We used high-resolution otolith microchemistry data from the temperate reef fish Hypsypops rubicundus to explore biweekly, seasonal, and annual connectivity patterns in an open-coast MPA network. The three MPAs, spanning 46 km along the southern California coastline were connected by larval dispersal, but the magnitude and direction of connections reversed between 2008 and 2009. Self-recruitment, i.e. spawning, dispersal, and settlement to the same location, was observed at two locations, one of which is a MPA. Self-recruitment to this MPA ranged from 50-84%; within the entire 60 km study region, self-recruitment accounted for 45% of all individuals settling to study reefs. On biweekly time scales we observed directional variability in alongshore current data and larval dispersal trajectories; if viewed in isolation these data suggest the system behaves as a source-sink metapopulation. However aggregate biweekly data over two years reveal a reef network in which H. rubicundus behaves more like a well-mixed metapopulation. As one of the few empirical studies of population connectivity within a temperate open coast reef network, this work can inform the MPA design process, implementation of ecosystem based management plans, and facilitate conservation decisions.

2011
Fodrie, FJ, Becker BJ, Levin LA, Gruenthal K, McMillan PA.  2011.  Connectivity clues from short-term variability in settlement and geochemical tags of mytilid mussels. Journal of Sea Research. 65:141-150.   10.1016/j.seares.2010.09.001   AbstractWebsite

The use of geochemical tags in calcified structures of fish and invertebrates is an exciting tool for investigating larval population connectivity. Tag evaluation over relatively short intervals (weeks) may detect environmental and ecological variability at a temporal scale highly relevant to larval transport and settlement. We collected newly settled mussels (Mytilus californianus and M. galloprovincialis) weekly during winter/spring of 2002 along the coast of San Diego, CA, USA, at sites on the exposed coast (SIO) and in a protected coastal bay (HI), to investigate temporal patterns of geochemical tags in mussel shells. Analyses of post-settlement shell via LA-ICP-MS revealed statistically significant temporal variability for all elements we examined (Mg, Mn, Cu, Sr, Cd, Ba, Pb and U). Despite this, our ability to distinguish multielemental signatures between sites was largely conserved. Throughout our 13-week study, SIO and HI mussels could be chemically distinguished from one another in 78-87% of all cases. Settlement varied between 2 and 27 settlers grambyssus(-1) week(-1) at 510 and HI, and both sites were characterized by 2-3 weeks with "high" settlement. Geochemical tags recorded in early larval shell of newly settled mussels differed between "high" and "low" settlement weeks at both sites (MANOVA), driven by Mg and Sr at SIO (p = 0.013) and Sr, Cd, Ba and Pb at HI (p < 0.001). These data imply that shifts in larval sources or transport corridors were responsible for observed settlement variation, rather than increased larval production. In particular, increased settlement at HI was observed concurrent with the appearance of geochemical tags (e.g., elevated Cd), suggesting that those larvae were retained in upwelled water near the mouth of the bay. Such shifts may reflect short-term changes in connectivity among sites due to altered transport corridors, and influence the demography of local populations. (C) 2010 Elsevier B.V. All rights reserved.

2009
Fodrie, FJ, Levin LA, Lucas AJ.  2009.  Use of population fitness to evaluate the nursery function of juvenile habitats. Marine Ecology-Progress Series. 385:39-49.   10.3354/meps08069   AbstractWebsite

Juveniles of many fish and invertebrate species are able to select among a diverse portfolio of nursery habitat alternatives. Environmental heterogeneity among these habitats generates variation in the vital rates of young individuals that may influence overall population dynamics. Therefore, understanding how these habitat options affect population fitness is crucial for identifying habitats that widen bottlenecks in early life histories and promote population persistence. We used cohort analyses and demographic models to explore the population-level consequences of habitat selection by juvenile California halibut Paralichthys californicus in southern California, focusing on population growth rate (lambda) as a measure of fitness. Although alternative juvenile habitats (exposed coast and coastal embayments) could contribute an approximately equal number of recruits to the adult stock, positive overall population growth (lambda > 1) depended critically on the subpopulations of juveniles that utilized coastal embayments (bays, lagoons, and estuaries). Conversely, the juvenile subpopulation along the exposed coast contributed negatively to overall population growth (lambda < 1) in 3 of the 4 years we conducted this study, due to elevated local mortality in that habitat. Life table response experiments confirmed that juvenile growth and survivorship were responsible for differences in lambda, and that nursery habitat choice could be a key contributor toward overall population fitness. Considering nurseries in a demographic source-sink context could aid conservation efforts by allowing identification or prioritization of the juvenile habitats most critical for population persistence.

2008
Fodrie, FJ, Levin LA.  2008.  Linking juvenile habitat utilization to population dynamics of California halibut. Limnology and Oceanography. 53:799-812.   10.4319/lo.2008.53.2.0799   AbstractWebsite

We investigated the nursery role of four coastal ecosystems for the California halibut (Paralichthys californicus) using the following metrics: (1) contribution in producing the fish that advance to older age classes, (2) connectivity of coastal systems resulting from migration of fish from juvenile to subadult habitats, and (3) effect of nursery habitat usage and availability on subadult population size, specifically evaluating the concentration hypothesis. Potential nurseries were grouped using a robust classification scheme that segregated exposed, bay, lagoon, and estuarine environments. Assignment of nursery origins for individual subadult fish via elemental fingerprinting indicated that exposed coasts, bays, lagoons, and estuaries contributed 31%, 65%, 1%, and 3% of advancing juvenile halibut during 2003, versus 49%, 33%, 16%, and 2% during 2004, respectively. These results were remarkably similar to "expected'' nursery contributions derived from field surveys, suggesting that in this system juvenile distributions were a good indicator of unit-area productivity of juvenile habitats and that density-dependent mechanisms during the juvenile phase did not regulate recruitment pulses. Elemental fingerprinting also demonstrated that individuals egressing from bays did not migrate far from their nursery origins (, 10 km), resulting in reduced connectivity along the 110-km study region over the timescale of approximately one generation. Consequently, we observed considerably higher subadult densities at sites near large bays, while populations distant from large bays appeared to be more influenced by nursery habitat limitation. Over large (similar to 100 km) scales, the location and availability of nursery habitat alternatives had significant effects on the population dynamics of an important member of the ichthyofaunal community of southern California.

2002
Davis, JLD, Levin LA, Walther SM.  2002.  Artificial armored shorelines: sites for open-coast species in a southern California bay. Marine Biology. 140:1249-1262.   10.1007/s00227-002-0779-8   AbstractWebsite

Artificial hard substrates have been used to stabilize naturally soft bay shorelines for centuries. Despite the loss of over half of the natural shoreline in many bays, little attention has been paid to the communities inhabiting armored shorelines and to the ecological implications of armoring. The goal of the present study was to examine factors affecting spatial and temporal variation of intertidal, hard-substrate biota (emergent species and fishes), with emphases on the influence of exposure, distance from the open ocean, and similarity to open-coast, hard-substrate communities. We examined community composition at eight San Diego Bay (California, USA) sites (an exposed and a protected site at four bay locations) in June and November 2000 and two open-coast sites in August 2000. At all bay sites, the shore was armored with granite boulders, a form of shoreline stabilization referred to as "riprap." Community structure was more variable spatially than temporally on the scales we studied, affected more by distance from the bay mouth and exposure to wave energy than by differences between June and November. Exposed sites near the bay mouth were more similar to natural open-coast sites, sharing about 45% of their species, than protected sites and sites farther from the mouth, which shared as few as 8%. Species richness was generally higher in exposed than protected bay sites. Species tended to occur higher in the intertidal zone at exposed than protected sites, and higher in November, when sea level was higher, than in June. Such results will be useful to shoreline managers who examine the ecological implications of hardening long stretches of coastline and may suggest ways to incorporate artificial structures into ecosystems in a more meaningful way.

2000
Dibacco, C, Levin LA.  2000.  Development and application of elemental fingerprinting to track the dispersal of marine invertebrate larvae. Limnology and Oceanography. 45:871-880. AbstractWebsite

The early life history of many marine benthic invertebrate and fish species involves a planktonic larval stage that allows exchange of individuals among separated adult populations. Here, we demonstrate how natural and anthropogenic trace elements can be used to determine larval origins and assess bay-ocean exchange of invertebrate larvae. Trace elements can be effective site markers for estuaries because run-off and pollutant loading often impart distinct elemental signatures to bay habitats relative to nearshore coastal environments. Crab larvae originating from San Diego Bay (SDB) were distinguished from those originating in neighboring embayments and exposed coastal habitats by comparing multiple trace-element concentrations ("fingerprints") in individuals. Discriminant function analysis (DFA) was used to characterize stage I zoeae of the striped shore crab, Pachygrapsus crassipes, of known origin (reference larvae) via trace-elemental composition (i.e., Cu, Zn, Mn, Sr, Ca). Linear discriminant functions were used to identify the origin and characterize the exchange of stage I P. crassipes zoeae between SDB and the nearshore coastal environment during one spring tidal cycle. Elemental fingerprinting revealed that most (87%) of the stage I larvae collected at the bay entrance during the flood tide were larvae of SDB origin that were reentering the bay. Nearly one third of zoeae sampled (32%) at the entrance during ebb tide were coastal larvae leaving the bay and returning to open water. The observed bidirectional exchange contrasts with the unidirectional transport of zoeae out of the bay predicted from stage I vertical migratory behavior. Because P. crassipes zoeal survivorship is lower in SDB than in coastal waters, bay-ocean exchange has significant implications for the dynamics of P. crassipes populations. Trace-elemental fingerprinting of invertebrate larvae promises to facilitate investigations of many previously intractable questions about larval transport and dynamics.

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.