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Susceptibility of Nereocystis Luetkeana (Laminariales, Ochrophyta) and Eualaria Fistulosa (Laminariales, Ochrophyta) Spores to Sedimentation

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Susceptibility of Nereocystis Luetkeana (Laminariales, Ochrophyta) and Eualaria Fistulosa (Laminariales, Ochrophyta) Spores to Sedimentation Research Article Algae 2012, 27(2): 115-123 http://dx.doi.org/10.4490/algae.2012.27.2.115 Open Access Susceptibility of Nereocystis luetkeana (Laminariales, Ochrophyta) and Eualaria fistulosa (Laminariales, Ochrophyta) spores to sedimentation Melissa Deiman1,a,*, Katrin Iken1 and Brenda Konar1 1School of Fisheries and Ocean Sciences, University of Alaska Fairbanks, P.O. Box 757220, Fairbanks, AK 99775, USA The establishment of algal spores plays an essential role in adult kelp distribution and abundance patterns. Sedimen- tation is a key variable regulating algal spore settlement and success, possibly controlling species-specific dominance in situ. Laboratory experiments were used to determine spore attachment and survival rates of two Alaskan canopy-forming kelps, Nereocystis luetkeana (K. Mertens) Postels & Ruprecht and Eualaria fistulosa (Postels & Ruprecht) M. J. Wynne, to various types of sediment loading. Spore attachment for both species was significantly and similarly affected by three sediment treatments: suspended particles; settled sediment covering the substratum; and smothering of attached spores by settling sediment. Spore attachment decreased by approximately 90% at 420 mg sediment L-1, the highest sediment load tested here, under all three treatments for both species. These results suggest that increases in sedimentation may constrain the success of the spore stages, but sediment does not seem to be a likely factor explaining species-specific distribution patterns. However, while sedimentation affected spores of both species similarly, timing of spore release in relation to times of maximum sediment load in the water might differ for different species, possibly explaining kelp spe- cies distribution patterns. Key Words: Alaska; Eualaria; kelp; Nereocystis; sediment; spore INTRODUCTION Survival, successful development, and distribution of (Amsler et al. 1992, 1999). Typically, kelp spores are viable kelp are influenced by environmental factors (Dayton immediately upon release and may settle on the seafloor 1985), which can affect the microscopic stages (spores, within minutes (Santelices 1990, Reed et al. 1992). The gametophyte, and male and female gametes) and the successful settlment and attachment of spores plays an macroscopic stage (sporophyte). All life stages exhibit important role in determining patterns of adult kelp dis- varying tolerances to environmental factors (Lüning tribution and abundance (Dayton 1985, Schiel and Foster 1980, Deysher and Dean 1986, Amsler and Neushul 1991, 1986, Santelices 1990). Sedimentation has been shown Dring et al. 1996, Edwards 2000), making some species to be a particularly influential parameter affecting spore and life stages more or less adapted to changing condi- survival and attachment success (Devinny and Volse tions. In particular, spores will preferentially settle onto 1978, Airoldi 2003). suitable substrata in appropriate microenvironments Increasing temperatures have led to accelerated glacial This is an Open Access article distributed under the terms of the Received February 15, 2012, Accepted May 2, 2012 Creative Commons Attribution Non-Commercial License (http://cre- Corresponding Author ativecommons.org/licenses/by-nc/3.0/) which permits unrestricted * non-commercial use, distribution, and reproduction in any medium, E-mail: [email protected] provided the original work is properly cited. Tel: +1-907-299-3296, Fax: +1-907-581-1572 aPresent address: Alaska Department of Fish and Game, Dutch Harbor, AK 99692, USA Copyright © The Korean Society of Phycology 115 http://e-algae.kr pISSN: 1226-2617 eISSN: 2093-0860 Algae 2012, 27(2): 115-123 melting in Alaska (VanLooy et al. 2006), which probably It is important to separately analyze the effects of sedi- increases the amount of glacially-discharged sediments mentation on Nereocystis and Eualaria spore attachment that enters the nearshore environment. Spores within and survival in an effort to tease apart how this variable the water column can attach to suspended particles dis- affects each species in a controlled laboratory setting. charged by glacial and terrestrial runoff, preventing them Understanding the effects of sediments on individual from attaching to bottom substrate (Arakawa and Mat- species may help us discern what drives the spatial and suike 1992, Arakawa 2005). Sediment can also occlude temporal variability of these species. The objective of this suitable surfaces preventing settlement, and sediments study was, therefore, to determine spore attachment suc- can smother settled spores, leading to mortality (Devinny cess under increasing levels of sedimentation, specifi- and Volse 1978, Airoldi 2003). For example, the effects of cally the effects of suspended sediment loads, of different sediments on Macrocystis pyrifera (Linnaeus) C. Agardh quantities of settled sediments, and of sediment smoth- spores appear to be immediate and significantly hindered ering of attached spores by known sedimentation rates. spore attachment in the laboratory (Devinny and Volse Due to Nereocystis’ current dominance in Kachemak Bay, 1978). By determining how sedimentation influences the which corresponds with the increase in sediment load- establishment and survival of various canopy-forming ing from glacial melt, it was hypothesized that Nereocystis kelp species, we may be able to predict how changes in spores would have higher survival and attachment rates this climate-change related variable may impact the kelp under increased levels of sedimentation compared to composition and distribution of nearshore kelp forests. Eualaria spores. Within the nearshore of much of Alaska’s rocky coast- line, the two surface canopy-forming kelp species, Nereocystis luetkeana (K. Mertens) Postels & Ruprecht MATERIALS AND METHODS and Eualaria fistulosa (Postels & Ruprecht) M. J. Wynne (formerly Alaria fistulosa; hereafter referred to as Nereo- Laboratory experiments using fertile sori from two cystis and Eualaria), may form mono-specific or mixed canopy forming kelp species collected at five sites in species kelp forests that provide very different structural Kachemak Bay, Alaska, tested the effects of suspended habitats for associated communities (Efird and Konar and deposited sediments on spore attachment, and unpublished data). The long, slender stipe of Nereocystis sedimentation on spore survival. Kachemak Bay, located provides much less midwater structure than the exten- north of the Gulf of Alaska at 59° N and 151° W, was cho- sive, submerged blade structure of Eualaria. The many sen as the study area because of its presence of both kelp apical blades of Nereocystis create considerably denser canopy-forming species, Nereocystis and Eualaria. Sorus surface structure than the single Eualaria blade. Hence, collection sites were a representative selection of the ma- the abundance or dominance of one canopy kelp species jor kelp beds along the southern shore of Kachemak Bay over the other may have habitat implications for associ- (Fig. 1); multiple collection sites were used to exclude any ated communities. In addition, the density and distribu- site-specific fitness patterns of the parent populations. tion of these species are temporally and spatially vari- All laboratory experiments were conducted in the sum- able (Schoch 2001). This has been well documented in mer of 2009 at the Kasitsna Bay Laboratory located on the Kachemak Bay, Alaska, where Eualaria was the dominant southern shore of Kachemak Bay. Sedimentation rates in canopy-forming species in the 1970s (Lees 1976), while the summer within the bay range from 1 to 5 mg cm-2 day-1 Nereocystis was the dominant species in the early 2000s (Spurkland and Iken 2011). (Chenelot 2003, Schoch and Chenelot 2004, Deiman un- published data). Currently, Nereocystis still dominates Sorus collection and spore release but Eualaria is being observed with increasing frequency (personal observation). Kachemak Bay is an estuarine Sorus collections were made from individual algae fjord under strong influence of glacial sediment dis- growing at a water depth of 6.1 ± 1.5 m in rocky habitats at charge (VanLooy et al. 2006, Spurkland and Iken 2011). the various sites (Fig. 1). Approximately 15-20 fertile sori This high sedimentation may be a variable that prevents were collected from at least five mature individuals per one or both canopy-forming species from settling and species at each site. Sori of both species were collected establishing in areas of high sedimentation (Schoch and in close proximity so that all sori at a site originated from Chenelot 2004), an effect that may increase with increas- equivalent environmental conditions. For Nereocystis, ing glacial discharge. sori were gathered from floating blades at the surface http://dx.doi.org/10.4490/algae.2012.27.2.115 116 Deiman et al. Kelp Spore Susceptibility to Sedimentation Fig. 1. Map of Kachemak Bay indicating the locations of Nereocystis luetkeana and Eualaria fistulosa sorus collection sites. Sori of both species were collected at each site. Sites: OB, Outside Beach; HI, Hesketh Island; YI, Yukon Island; IC, Inner Cohen Island; OC, Outer Cohen Island. and placed into coolers filled with ambient seawater. Eualaria sori were not incubated damp but immediately Eualaria sori were gathered from the base of the mature placed into beakers with 1 L of 0.2 μm filtered seawater. sporophytes using scuba. The sori from each individual The Eualaria sorus solution became dark after approxi- alga were placed in separate mesh bags and brought
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