Ascophyllum Nodosum) and Associated Epifaunal Communities Following Cutter Rake Harvesting in Maine
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Evaluation of short-term changes in rockweed (Ascophyllum nodosum) and associated epifaunal communities following cutter rake harvesting in Maine Thomas J. Trott Friedman Field Station, Suffolk University, Edmunds, Maine 04628 Department of Biology, Suffolk University, Boston, Massachusetts 02114 and Peter F. Larsen Bigelow Laboratory, PO Box 475, 180 McKown Point Rd. West Boothbay, Maine 04575 Report to Department of Marine Resources Marine Resources Laboratory P.O. Box 8 194 McKown Point Rd. W. Boothbay Harbor, Maine 04575-0008 INTRODUCTION North Atlantic intertidal areas are often dominated by the rockweed Ascophyllum nodosum which can grow to greatest length and mass on wave protected shores. Rockweed is a long-lived canopy alga. In some areas it can achieve near mono-specific cover. The holdfasts and three-dimensional canopy of rockweed beds provide habitat for intertidal invertebrates and fish, and offer a place for attachment of their eggs and epiphytic algae. Rockweed is harvested for use in cosmetics, processed foods, domestic animal feeds and fertilizers. It has been harvested commercially for decades in the Gulf of Maine. Concerns about harvesting have generated numerous publications and workshops (e.g., Rangeley and Davies, 2000). While it is well documented that harvesting rockweed stimulates growth (Lazo and Chapman, 1996; Ugarte, et al., 2006), less attention has been directed towards disturbance effects at the community level. If associated animal communities are affected by rockweed harvesting, then the way that harvesting is done with a cutter-rake would produce a patchwork of disturbance. Harvesting with a cutter-rake removes rockweed in patches over large areas on an incoming tide. Once in a rockweed bed, a harvester targets the largest or bulkier plants in the water that can be reached with his rake. The harvester’s boat moves with the wind and tide to a random position where the bulkiest floating plants are targeted. As a result, the cut plant is chosen by the harvester but position in the bed is not. The goal is to harvest maximum biomass, although the lengths that plants are cut can vary from changes in visibility, the harvester’s stroke and depth perception. Harvesting is conducted in accordance to regulations requiring a 16” length minimum of plant along with its holdfast remain. Potential effects on fauna associated with rockweed at the scale of a rockweed bed would be linked to the patches where targeted plants were removed. The degree of disturbance might fluctuate according to the varying amount of plant removed. A recent increase in harvesting rockweed has raised public concern in Maine and re- stimulated interest to evaluate some basic potential impacts. This study was undertaken to examine the effectiveness of current regulations on cutter-rake harvesting for preventing and/or minimizing impact on rockweed and associated macroinvertebrates. Short-term recovery, two months following a single harvest, was the time course for evaluating disturbance effects. 1 METHODS A field experiment was conducted using a modified Before-After, Control-Impact (BACI) experimental design with stratified random destructive sampling to evaluate some potential effects of harvesting. The study site was located in Shackford Head State Park, Cobscook Bay, Maine to minimize uncontrolled disturbance from anthropogenic activities (Figure 1). In July 2008, rockweed biomass and associated epifaunal species on rockweed thalli and holdfasts with contiguous substrate were assessed in adjacent control and experimental plots (Figure2). In each plot, four quadrats were sampled per tidal stratum (high, mid, low intertidal). Positions of randomly selected 1/16-m2 quadrats were recorded with GPS. Within each quadrat, plants were sheared just above their holdfasts (<1cm) and transported to the laboratory where they were washed with freshwater under pressure above a 0.5 mm screen to collect mobile epifauna. Material retained on the sieve was preserved in 10% formalin and stained with Rose Bengal. Rockweed samples were drained of excess water and weighed. Individual thalli were examined for attached and remaining mobile epifauna, and eggs. All mobile fauna was saved with sieved material. In the field, epifauna within each quadrat exposed after removing rockweed was collected with forceps. Surfaces were scraped to collect rockweed holdfasts and remaining epifauna (Figure 3). Samples were washed and material retained on a 0.5mm sieve was preserved in 10% formalin and stained with Rose Bengal. Animals were sorted, identified to the lowest taxon possible, and counted. Species names were checked in the World Register of Marine Species database (http://www.marinespecies.org/) to insure use of current valid names. All formalin preserved samples were transferred to ethanol for archiving at the Maine Department of Marine Resources. The impact of harvesting as it is done commercially was a priority so treatments could not be randomly assigned to quadrats within each plot with plants cut at uniform heights. That would not match how rockweed is commercially harvested. Instead, immediately after sampling, rockweed was harvested in the experimental plot by a professional harvester from Acadian Seaplants Limited, New Brunswick, using a cutter rake (Figure 4). The harvester followed Maine Department of Marine Resources harvesting regulations and their activities observed and 2 documented (Figure 5). During harvesting, plants were not cut below 16 inches or the first branch of the primary thalli. Approximately 1.25 wet tons of rockweed was harvested by one person in 1.5 h and an estimated 17% of the biomass of rockweed in the study area was removed. Both plots were re-sampled in September leaving two months of recovery for the harvested experimental plot. Re-sampling the same quadrat was avoided since the GPS location of each quadrat sampled in July was known. Making a priority on assessing the effects of harvesting rockweed commercially rather than through experimentally consistent, controlled hand removal from much smaller patches placed conditions on how data were analyzed. Although each plot had 12 sample quadrats, these were not independent replicates. This meant that parametric analyses were not valid. Keeping this in mind, analyses performed were for summarizing results observed and not for statistical significance. When statistical significance is given, it is for presentation purposes only. None- the- less, these results are clearly evident in graphical illustrations even without statistics. Another constraint on the experimental design was financial. This dictated the number of plots that could be studied. In keeping with the priority of commercial harvesting effects, having more than one experimental plot was not financially feasible. These caveats need to be held in mind by the reader throughout the presentation of data, results and conclusions. Rockweed biomass and epifauna data were analyzed with a paired BACI design where the differences in measurements before and after in control and experimental plots were tested for using a two-sample t-test. This procedure tests for the significance of impact (Smith et al. 1993; Stewart-Oaten et. al., 1986; Stewart-Oaten and Bence, 2001) which in this case is harvesting. Specifically, before harvest measurements from experimental plots are subtracted from control plot measurements and the mean difference compared using a two-sample t-test with that derived using the same method but with measurements taken after harvesting. Additional tests for differences in epifauna distribution according to habitat were conducted using ANOVA. When warranted, multiple comparison tests were performed using the Student Newman-Keuls Test to identify significant differences among groups. Rockweed biomass was analyzed using simple linear regression to examine relationships. Cluster analysis and non-parametric multidimensional scaling were performed on epifauna data using PRIMER ver. 6 (Plymouth Routines in Multivariate Ecological Research) 3 (Clarke and Warwick, 2001). Species counts were filtered according to statistical test protocols for cluster analysis and multidimensional scaling. This included eliminating all colonial species from the analysis since these did not represent counts. Also, species which occurred in 10% or less of all samples were designated as rare and removed from the analysis. The remaining species counts which represented the bulk of the data were square root transformed to allow intermediate abundant species to contribute to the similarity matrix calculated using the Bray-Curtis Index. Cluster and non-parametric multidimensional scaling analyses were performed on the resulting Bray-Curtis coefficients. Significant differences (P<0.05) between species grouped according to cluster analysis were determined using the PRIMER SIMPROF routine. For all analyses in this study, α = 0.05 was the level for significance. When statistical significance is given, it is for presentation purposes only for reasons described earlier. RESULTS Harvesting had a significant impact on rockweed Ascophyllum nodosum biomass (t= 2.117, P= 0.046) (Figure 6A). Biomass of rockweed in the experimental plot was greater following harvesting. September biomass was significantly different from July in the harvested plot (Student Newman-Keuls Test, q= 4.032, P=0.007), while the control plot showed no significant change (Figure 6B). There was a significant (P<0.05) direct linear relationship between the number of thalli and weight