MARINE ECOLOGY PROGRESS SERIES Vol. 106: 147-156, 1994 Published March 17 Mar. Ecol. Prog. Ser. Patch dynamics of eelgrass Zostera marina Birgit Olesen, Kaj Sand-Jensen* Department of Plant Ecology, University of Aarhus, Nordlandsvej 68, DK-8240 Risskov, Denmark ABSTRACT: Eelgrass has declined extensively during the last decades following eutrophication of coastal regions of western Europe and the USA. Recent efforts have been taken to reduce nutrient loading, with the hope of restoring the former widespread populations. We studied fine-scale patch dynamics of eelgrass Zostera marina L, in permanent plots outside the continuous vegetation in a pro- tected embayrnent in Limfjorden, Denmark, to evaluate means of recruitment and rates of expansion and mortality of patches. The size distribution was dominated by small patches which were formed by seedlings at high rates during spring (0.16 to 0.76 m-'). Mortality was high and only low proportions (0 to 24 %) of the studied cohorts remained as individual patches 1.5 and 2.5 yr later. Patch mortality was restricted to small patches containing <32 shoots with a mean age of < 5 yr. The sharply declining mortality with increasing patch age and size is presumably due to improved anchoring, mutual physi- cal protection and physiological integration among the shoots. The lateral expansion of established patches by centrifugally growing horizontal rhizomes averaged 16 cm yr-l and was independent of patch size With this lateral growth the possible annual areal expansion will be faster in systems with many small patches (260% in 0.1 m' patches) and slower in systems with few large patches (19% in 10 m2patches). Successful recovery of the eelgrass vegetation in large systems is, therefore, dependent on seed production and dispersal and subsequent seedling establishment and patch growth. KEY WORDS: Zostera marina. Population . Colonization . Patch dynamics INTRODUCTION Eelgrass Zostera marina (L.) has undergone similar changes in areal cover throughout most of its distnbu- The areal cover of seagrasses undergoes extensive tion range in Europe and the USA, including a wide- natural fluctuations due to variable growth conditions spread recent decline (den Hartog & Polderman 1975, and catastrophic declines during storm events (Birch & Rasmussen 1977, Orth & Moore 1983a, Phillips 1984, Birch 1984, den Hartog 1987, Williams 1988, Larkum den Hartog 1987, Giesen et al. 1990). Encouraging & den Hartog 1989) and epidemic diseases (Giesen et examples of eelgrass recovery remain few (Wyer et al. al. 1990). More recently, cultural eutrophication and 1977, Verhagen & Nienhuis 1983, Harrison 1987). For intensified coastal development have severely reduced example, eelgrass covered about 7000 km2of the inner seagrass abundance and alarmed marine ecologists coastal Danish waters early in this century (Petersen and responsible politicians (Kemp et al. 1983, Cam- 1914).The 'eelgrass wasting disease' in the early 1930s bridge & McComb 1984, Giesen et al. 1990, Larkum & destroyed most eelgrass meadows, and in 1942 the West 1990). Although seagrass decline occurs world- areal cover of eelgrass was only 540 km2 (Rasmussen wide, studies of the dynamics of seagrass cover and the 1977). After a phase of re-establishment, eutrophica- associated colonization, expansion-recession, and mor- tion has subsequently led to further reduction of eel- tality processes are few (Birch & Birch 1984, Brouns grass cover since the 1970s (Orth & Moore 1983a, den 1987, Williams 1988, Duarte & Sand-Jensen 1990a, b, Hartog 1987). The eelgrass populations present today Williams 1990). are mainly small and confined to shallow protected areas and should have great difficulties in expanding 'Present address: Freshwater Biological Laboratory, Univer- to the larger, more exposed and deeper regions sity of Copenhagen, Helsingersgade 51, DK-3400 Hdler~d. because of insufficient seed input, greater physical dis- Denmark turbance and poorer light conditions than during the O Inter-Research 1994 148 Mar. Ecol. Prog. Ser. 106: 147-156, 1994 former periods of widespread eelgrass coverage. How- during the study period. A continuous eelgrass meadow ever, a planned reduction in nutrient loading to Danish covers the central deeper part of the embayment, occu- waters by 80% for P and 50% for N before 1995 aims pying 52% of the total area, and is bordered at its at reducing phytoplankton development and restoring upper limit by a fringe (50 to 100 m wide) of isolated the former eelgrass populations. The likelihood and eelgrass patches at mean depths between 0.4 and possible time scales of eelgrass recovery remain as yet 0.7 m. This investigation was performed within this unknown. shallow zone of eelgrass vegetation. Eelgrass meadows are maintained principally by Eelgrass patch dynamics. Size distribution of eel- vegetative production of lateral shoots (Olesen & Sand- grass patches (minimum registered area 0.3 m2)within Jensen 1994) and areal expansion is probably limited the studied embayment was measured in a 1000 m2 by the slow growth rate of horizontal rhizomes (Sand- area to include large size-classes. Jensen 1975).Colonization of new areas should depend Small eelgrass patches down to single shoots and on the dispersal and subsequent germination of seeds their dynamics were studied by mapping the vegeta- and, therefore, on the distance to reproductive stands tion at intervals of 3 to 12 mo in 2 permanent plots and the maintenance of adequate conditions for (120 m2)established in May 1990 (Plot I) and May 1991 seedling establishment and growth. Patch mortality is (Plot 11). Within each plot a grid was established by likely to decline with increasing patch size (e.g.Duarte placing permanent sticks at 2 m intervals, onto which & Sand-Jensen 1990a, Sand-Jensen & Madsen 1992) an aluminium frame (1 m') divided into 0.2 X 0.2 m because of mutual protection among neighbour plants. squares could be fitted during observations. On each Lateral spread and shoot growth may also depend on sampling date the position and dimensions of all patch size, such that shoot mortality may be high and patches were determined and each patch was identi- areal expansion slow until a certain critical minimum fied according to the previous recordings. The origin of patch size is reached (Duarte & Sand-Jensen 1990a, b). new patches appearing between sampling dates, We provide here a first step to evaluate eelgrass whether developed from seedlings or fragments of patch dynamics by determining (1) the formation and former larger patches, was recorded. Newly estab- size-dependent mortality of eelgrass patches, (2) the lished seedlings were identified from their morphology lateral spread of differently sized patches and (3) shoot (short, narrow-leaved shoots with 1 to 3 leaves). This growth rate as a function of patch size and position identification was confirmed outside the permanent within the patch. We restricted the investigation to a plot on harvested seedlings showing remains of the single site in a protected Danish embayment during seedling coat and a scorpioid rhizome base (Setchel a 2.5 yr period and concentrated on the short-term 1929). The number of shoots in each patch was dynamics of predominantly small patches character- counted, or in larger patches (> 0.1 m2)estimated from ized by high turnover. We acknowledge that more sites the area multiplied by the measured annual mean of different exposure and longer time periods, par- shoot density of 840 shoots m-2 (Olesen & Sand-Jensen ticularly for large patches, are needed to establish a 1994). Our patch dynamic analysis was based on the more comprehensive knowledge of eelgrass popula- distribution in logarithmic size classes that became tion dynamics. increasingly wider at higher numbers to make the procedure described above sufficiently accurate. Patch mortality was calculated as the relative number of METHODS patches lost within each size-class. Patch growth. Lateral expansion of patches was Study area. Eelgrass patch dynamics were investi- determined by measuring the elongation of eelgrass gated from May 1990 to September 1992 in a small rhizomes from the patch margin into bare sediments (1.6 km2) semi-enclosed embayment of Limfjorden, a for 38 patches ranging widely in size (diameter from shallow eutrophic brackish-water area (1500 km2) in 0.2 to 8.1 m). Permanent marks were placed at the northern Jutland, Denmark. The embayment studied is edge of each patch at 4 positions (N, S, E and W of 1.7 km long and 1.4 km wide at the entrance. The patch centre) in April 1990 and lateral growth relative effective fetch, estimated according to Smith (1979), to the marks was measured in July and October 1990 was short (0.51 km) towards the prevailing westerly and May and September 1991. winds, indicating relatively high protection. The em- Eelgrass shoot growth. The influence of patch size bayment is 2 m deep at the entrance, and slopes to a and distance to the patch edge on shoot growth was depth of about 1 m in the central part. Mean tidal studied in August by measuring leaf elongation rate range is 0.15 m, but wider fluctuations in water level normalized to leaf length of shoots at the centre of are caused by wind. Sallnity averages 26%0and water 5 patches of different size (diameters between 1.2 and temperature ranged from just below zero to 25OC 20 m) and along transects from the edge (20 cm inside Olesen & Sand-Jensen: F'atch dynam~csof eelgrass 149 the patch) to the centre of these 5 patches. We and most (85%) of them contained 1 or 2 shoots. New expressed shoot growth as relative leaf elongation patches were formed from seedlings which were the rate, analogous to the relative growth rate widely used main contnbutors to the high number of small patches as the standard for terrestrial plants, also to compen- in May 1990 and May 1991 as well as to the overall sate for the influence of different shoot sizes on increase in patch number (Figs.