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Environmental heterogeneity can promote coexistence, and is therefore predicted to increase invasibility of communities, but decrease invasion impacts. We examined the role of local-scale environmental heterogeneity in promoting the coexistence of the invasive seagrass, Zostera japonica, and its native congener Zostera marina in a patch mosaic. In its introduced range, Z. japonica often co-occurs with the native Z. marina in a patch mosaic associated with intertidal microtopography (centimeter to decimeter relief over meter to decameter distances). Here, Z. marina inhabits depressions that retain water during low tides, and Z. japonica inhabits well-drained mounds. Transplant experiments revealed that Z. marina suppressed Z. japonica shoot densities, more so in pools than on mounds. Z. marina suppressed Z. japonica above-ground and below-ground biomass by 47 and 19 % respectively, on mounds, and by over 60 % in pools. Z. marina shoot densities and biomass were 40 and 95 % lower, respectively, on mounds, regardless of Z. japonica presence. Topographic context remained the most influential predictor of Z. marina responses, even when we transplanted Z. marina into higher densities of Z. japonica. These results indicate that the native Z. marina is physiologically restricted from mounds and competitively excludes the introduced Z. japonica from pools. We provide empirical evidence of local-scale heterogeneity promoting coexistence of an invasive and a native macrophyte, supporting the hypothesis that environmental heterogeneity increases invasibility and decreases invasion impacts. Furthermore, active control of the invader in mixed beds is unlikely to benefit the native.

In the Northeast Pacific the nonnative seagrass Zostera japonica frequently exists at the same sites as the native seagrass Zostera marina. Although at some sites their vertical distributions overlap, at most sites in the Pacific Northwest there is a distinctive unvegetated zone between them. The objective of this study was to better understand why a gap between the lower limit of Z. japonica and the upper limit of Z. marina exists. To address this issue we carried out transplant experiments, conducted in situ monitoring of existing Z. japonica patches, and collected sediment samples at South Beach on Shaw Island, Washington, during the spring and summer of 2006. Transplant and in situ monitoring data indicate that survival and performance of Z. japonica are reduced lower in the intertidal zone. In addition, Z. japonica patches tended to be smaller and more spaced out at lower tidal heights. Although we found no Z. japonica seeds within or outside extant Z. japonica patches, high transplant mortality indicates that Z. japonica dispersal limitation is an unlikely cause of the unvegetated gap zone. Our field observations further suggest that herbivory, bioturbation, and epiphytes are unlikely causes of the gap pattern at our study site. Instead, we hypothesize that light limitation prevents Z. japonica from occurring lower in the intertidal. A review of published vertical distribution data for both Zostera species indicates that the lower limit of Z. japonica is relatively invariant among sites. In contrast, the upper limit of Z. marina is highly variable, ranging by more than 4 m within some subregions in Washington State. Consequently we hypothesize that intersite variability in the vertical distribution of Z. marina is the primary driver of spatial variability in the presence of the unvegetated gap.

Mangrove stands and seagrass beds grow along the coasts of the United Arab Emirates (UAE). These marine plant species are concentrated in specific tidal zones along sheltered intertidal coastlines in association with estuaries and lagoons; mangroves fringe the coastline encroaching into the lower intertidal region, whereas seagrasses populate the adjacent deeper water. In most cases, the distribution of mangroves and seagrasses does not overlap. It is important to monitor the geographic extent and health of mangrove forests and seagrass beds, which serve as an important habitat for juvenile marine species. Remote-sensing data for the Khor Al Bazam area in the vicinity of Abu Dhabi, UAE, covering the years of 1994, 2000, and 2003, were used to determine the change in mangrove and seagrass cover. Since 1994, there has been an increase in mangrove cover, likely because of plantation activity, the closure of nearby shipyards, and an increase in public awareness regarding mangrove preservation. Althou...

The current distribution of the introduced seagrass Zostera japonica is restricted to the mid- to upper intertidal zone in the coastal Pacific Northwest region of North America. The climate in this region is cool and wet, becoming hotter and dryer with increasing distance southward. Since temperature is likely to be an important factor affecting distribution of this species, growth of two populations located near the northern and southern limits of its established range along the Pacific Coast of North America were measured in an experimental setting across a range of temperatures typical of those in the field during the growing season (10, 20, and 30 °C). The effects of temperature and population were both significant. Leaf elongation, growth, and areal productivity rates of the northern population were consistently lower than those of the southern population. Across the range of temperatures, mean leaf elongation rates ranged from 0.47 to 1.40 cm 2 shoot −1 d −1 ; mean growth rates ranged from 0.19 to 0.52 mg dry wt shoot −1 d −1 . Mean areal productivity ranged from 0.54 to 1.92 g dry wt m −2 d −1 . Maximum rates of leaf elongation, growth, and areal productivity for both populations were observed at 20 °C. However, leaf elongation, growth, and areal productivity of the northern population declined markedly at 30 °C, whereas no comparable declines were observed for the southern population. This suggests that Z. japonica populations near the southern limits of its established range may be better adapted to warmer temperatures than populations near the northern range limits and further range extensions southward along the California coast may be likely. These differences could be important in predicting the outcome of competitive interactions between native and introduced seagrass species, and in determining future patterns of distribution and zonation of Pacific Coast seagrasses.

Photosynthetic processes in Zostera japonica, an upper intertidal species, were found to be more severely affected by desiccation than Z. marina, a lower intertidal and subtidal species, at comparable levels of tissue water content. The data indicate that photosynthetic responses to desiccation at the level of the individual leaf are insufficient to explain observed patterns of intertidal seagrass zonation. Desiccation tolerance in seagrasses is more likely to involve a complex interaction of morphological traits and growth strategies at the level of the whole plant, such as downsizing (e.g. smaller, narrower leaves), reduced structural rigidity and increased rates of leaf abscission and leaf turnover.

The objectives of this study were to determine the distribution and abundance of Zostera marina (eelgrass) in relation to the distribution of the mat forming bacteria Beggiatoa spp., and the levels of sulfide and organic material (wood waste) in the sediment. Underwater videography and intertidal surveys were used to map the distribution and abundance of Z. marina beds and Beggiatoa in the nearshore area of Commencement Bay, WA (USA), a location that has a long history of sawmill activity. Zostera marina occurred from the intertidal to −6 m mean lower low water (MLLW) on sandy substrates in areas with low levels of sulfide ( 200 μm) occurred where there were significant amounts of organic material in the sediments, which was found to be wood waste that had been discarded from sawmills. Zostera marina was absent from the intertidal and occurred at lower densities in areas with high sulfide levels. In contrast, mats of Beggiatoa were only found in areas where the sulfide levels were >1000 μm and there were significant deposits of wood. Thus, the negative correlation between the distribution and abundance of Z. marina and Beggiatoa suggests that the presence of Beggiatoa mats could be used as a biological indicator of inhibiting levels of hydrogen sulfide in the marine environment.

We gathered sequence information from the nuclear 5.8S rDNA gene and associated internal transcribed spacers, ITS-1 and ITS-2 (5.8S rDNA/ITS), and the chloroplast maturase K ( matK ) gene, from Zostera samples collected from subtidal habitats in Monterey and Santa Barbara (Isla Vista) bays, California, to test the hypothesis that these plants are conspecific with Z. asiatica Miki of Asia. Sequences from approximately 520 base pairs of the nuclear 5.8S rDNA/ITS obtained from the subtidal Monterey and Isla Vista Zostera samples were identical to homologous sequences obtained from Z. marina collected from intertidal habitats in Japan, Alaska, Oregon and California. Similarly, sequences from the matK gene from the subtidal Zostera samples were identical to matK sequences obtained from Z. marina collected from intertidal habitats in Japan, Alaska, Oregon and California, but differed from Z. asiatica sequences accessioned into GenBank. This suggests the subtidal plants are conspecific with Z. marina , not Z. asiatica . However, we found that herbarium samples accessioned into the Kyoto University Herbarium, determined to be Z. asiatica , yielded 5.8S rDNA/ITS sequences consistent with either Z. japonica , in two cases, or Z. marina , in one case. Similar results were observed for the chloroplast matK gene; we found haplotypes that were inconsistent with published matK sequences from Z. asiatica collected from Japan. These results underscore the need for closer examination of the relationship between Z. marina along the Pacific Coast of North America, and Z. asiatica of Asia, for the retention and verification of specimens examined in scientific studies, and for assessment of the usefulness of morphological characters in the determination of taxonomic relationships within Zosteraceae.

Recent studies found seed size variation within the seagrass Zostera marina, one of nine species in the genus Zostera. The objectives of this study were to determine if variation also exists in the seeds of two other species Zostera japonica and Zostera asiatica within this genus. Results indicate that: (1) length and weight varied between two populations (one indigenous population from Akkeshi-Ko, Japan, and one exotic population from Willapa Bay, Washington, USA) of the small-bodied intertidal seagrass species Z. japonica, and (2) seed-size classes were discernable. Preliminary investigations were also initiated with a Japanese population of Z. asiatica, a large-bodied subtidal seagrass species. Z. japonica seeds from the exotic population were significantly (P < 0.001) longer and heavier when compared to those from the indigenous population, a finding which may help explain both the process of the earlier introduction and the recent expansion of this exotic in the northeastern Pacific. Also, preliminary results indicate that Z. asiatica seeds are heavier than both those of Z. marina and Z. japonica, which suggests that larger seeds may be associated with large-bodied plants in this genus, an observation that should direct future seed ecology studies within the genus. These findings demonstrate that, similar to the study of terrestrial angiosperms, investigations designed to describe the comparative ecology of marine seed-bearing plants should include an evaluation of seed size.

Two species of seagrasses frequently co-occur with extensive thalassinid shrimp populations and aquaculture operations in the intertidal zone of estuaries along the west coast of North America. Although thalassinid shrimp are known to be strong bioturbators and affect both aquaculture operations and benthic intertidal community structure, few studies have investigated shrimp–seagrass interactions. Application of the pesticide carbaryl to control shrimp populations for oyster aquaculture in Willapa Bay, Washington provided us with an experimental tool to investigate one such interaction. We found that the seagrass Zostera japonica colonized areas where ghost shrimp ( Neotrypaea californiensis ) had been removed via carbaryl application. We applied carbaryl to small (900 m 2 ) experimental plots and compared seagrass colonization on these to that on control plots where shrimp remained abundant (100 m −2 ). The cumulative proportion of Z. japonica seeds and sprouts was slightly higher in the surface layer of treated plots (presumably due to the lack of shrimp bioturbation distributing them to depth), but seedling abundance was not significantly different between treated (no shrimp) and untreated control plots when they first emerged in early spring. As the season progressed however, and shrimp became more active, fewer seedlings survived in the untreated areas, and those that did survive grew much more slowly than those in the treated plots. We suspect that this was due to the effects of shrimp bioturbation and either light limitation (shoots that survived were much smaller) or direct burial and loss. Although it is an introduced plant, the natural distribution of Z. japonica is high in the intertidal zone and it is often separated from its congener Zostera marina by an expansive sandflat that is dominated by the ghost shrimp N. californiensis in west coast estuaries. The treatment of intertidal oysterbeds with carbaryl clearly reduces abundance of shrimp in this zone and we documented the same pattern of seagrass colonization on a commercial oyster bed and lack of seagrass in an adjacent unsprayed area. Density of native seagrass Z. marina shoots was also enhanced in plots treated with carbaryl, but only at lower tidal elevations or in intertidal pools where it could survive. We believe the removal of shrimp will continue to broaden the distribution of Z. japonica in Washington coastal estuaries where carbaryl use is permitted and add an interesting perspective to this controversial management issue.

In 1997 and 1998 seeds were collected from the seagrass, Zostera marina L. (Zosteraceae) from Long Island, New York, USA (var. typica), Shaw Island, Washington, USA (var. latifolia) and Izembek Lagoon, Alaska, USA (var. izembekensis) to determine if seed size varied between the varieties, if seed size varied within the same variety, and if the relationship between seed length and seed weight was linear. Measurements of seed length revealed statistically significant differences in this metric between varieties associated with smaller intertidal plants (var. typica and var. izembekensis) and larger plants from the subtidal region (var. latifolia). It was also found that seed length varied within the same variety and verified that the cross-sectional shape of these seeds was primarily elliptical. Using seed cross-sectional area and weight from a randomly selected batch of seeds from the three locations sampled in 1998, a linear model was created to determine seed weight and seeds from var. latifolia were found to be heavier than seeds from var. typica. The potential advantage of heavy seeds for subtidal seeds in search of safe sites is discussed, together with the merit of using the model to calculate the weight of other elliptical seeds within the Zosteraceae and other seagrass families. © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 142, 281–288.