Seagrass Meadows Modify Drag Forces on the Shell of the Fan Mussel <Emphasis Type="Italic">Pinna nobilis</Emphasis>

We assess the sheltering effect of Posidonia oceanica meadows on drag forces exerted on shells of the fan mussel Pinna nobilis. We examine a range of shell sizes under four unidirectional flow speeds (0.05–0.34 m s⁻¹) and two oscillating regimes. Three meadow densities are evaluated and a control without vegetation. We found that the attenuating effect of the meadow on drag forces experienced by bivalves is determined by the form of the hydrodynamic energy, e.g., as unidirectional flow or wave action. In tidal currents, the meadow protects most sizes of bivalves, with a higher efficiency for dense meadows, while in wave dominant zones the meadow reduces drag forces for bivalves with shell areas below a threshold of 0.019 m², whereas larger animals experience increased drag forces within the meadow independent of meadow density. Reduction of shoot density in seagrass meadows might therefore not affect the effectiveness of the canopy to reduce drag forces on associated species like the fan mussel in wave-dominated areas while increased storm frequency could result into losses of larger individuals during periods of high wave action.     

Fish assemblages in natural versus well-established recolonized seagrass meadows

Studies of fish assemblages between natural and newly recolonized (<4 yr) seagrass meadows have shown no significant differences in community composition between meadow types. However, comparison of natural and well-established (31 yr) recolonized seagrass meadows in the Indian River Lagoon, Florida, showed that, although patterns in fish assemblages are complex and not always consistent, differences were evident. Species richness was higher in natural meadows during spring and autumn while density and species richness were higher in recolonized meadows during summer. Juveniles of all but the five most abundant species were more common in one or the other meadow type. Additionally, species composition was distinctly different between recolonized and natural seagrass meadows, as indicated by UPGMA cluster analysis based on the Morisita-Horn similarity index, Spearman'sr _s (r _s>0.05 in all but one case), and a maximum of only 58.5% species in common. There were also significant differences in the length-frequency distribution for six of seven abundant species. Our results suggest that a well-established recolonized seagrass meadow has the potential to maintain species complements distinct from nearby natural meadows. Reasons for our differing results may include differences in seagrass morphology and collecting techniques between our study and the former studies. Additionally, species may have a longer time to establish specific habitat-use patterns in well-established compared to newly-formed recolonized meadows. Recolonized seagrass meadows appear to be as suitable a habitat as natural meadows for juvenile and small adult fishes.     

Laguna madre: Seagrass changes continue decades after salinity reduction

Vegetation maps of the lower Laguna Madre prepared from surveys conducted in 1965–1967, 1974–1976, and 1988 document a >330 km² decrease in cover byHalodule wrightii, an increase of almost 190 km² in other seagrass species, and an increase of 140 km² in bare bottom. Loss in seagrass cover is confined to deeper parts of the laguna; turbidity caused by maintenance dredging is the suspected cause. The species shifts are consistent with observed reductions in salinity maxima. Although the hydrological alterations and climatic shift responsible for moderating the salinity regime occurred between 1948 and 1965, the biological changes continue. Establishment of patches away from source meadows appears to be the process for displacing species that limits their rate of expansion into suitable habitat in this elongate embayment.     

Testing the predictive power of seagrass depth limit models

The power of equations predicting seagrass depth limit (Z_c) from light extinction (K _z) was tested on data on seagrass depth limits collected from the literature. The test data set comprised 424 reports of seagrass colonization depth and water transparency, including data for 10 seagrass species. This data set confirmed the strong negative relationship betweenZ _c andK _z. The regression equation in Duarte (1991) overestimated the realized seagrass colonization depths at colonization depths < 5 m, while there was no prediction bias above this threshold. These results indicated that seagrass colonizing turbid waters (K _z 0.27 m^-1) have higher apparent light requirements than those growing in clearer waters. The relationship between seagrass colonization depth and light attenuation shifts at a threshold of light attenuation of 0.27 m^-1, requiring separate equations to predictZ _c for seagrass growing in more turbid waters and clearer waters, and to set targets for seagrass restoration and conservation efforts.     

Postsettlement patterns of habitat use by sciaenid fishes in subtropical seagrass meadows

Spatial and temporal patterns of distribution and abundance were examined for postsettlement sciaenids collected from seagrass meadows in the Aransas Estuary, Texas. Overall, 5443 sciaenid larvae and early juveniles were identified from biweekly epibenthic sled collections taken from August 1994 to August 1995. Eight species were present in seagrass meadows, with five accounting for over 99.9% of sciaenids collected: silver perch (Bairdiella chrysoura), spotted seatrout (Cynoscion nebulosus), spot (Leiostomus xanthurus), Atlantic croaker (Micropogonias undulatus), and red drum (Sciaenops ocellatus). Settlement to seagrass meadows was partitioned temporally with little overlap among the five species. Postsettlers from inshore spawners (B. chrysoura, C. nebulosus, S. ocellatus) inhabited seagrass meadows during the spring and summer, while individuals from offshore spawners (L. xanthurus, M. undulatus) were present in the late fall and winter. Densities ofB. chrysoura, C. nebulosus, S. ocellatus were highest for small individuals (4–8 mm SL) and these taxa remained in seagrass sites through the early juvenile stage. Conversely,L. xanthurus andM. undulatus maintained longer pelagic periods and generally entered seagrass meadows at larger sizes (10–14 mm SL). Moreover, these taxa were only temporary residents of selected seagrass meadows, apparently migrating to alternative habitats shortly after arrival. During peak settlement, mean and maximum densities among species ranged from 0.1 m^?2 to 0.8 m^?2 and 0.7 m^?2 to 23.8 m^?2, respectively. Density and mean size of possettlement sciaenids differed significantly between seagrass species (Halodule wrightii, Thalassia testudinum) and among sites within the estuary.     

The relationship of vegetation and soil differentiation during the formation of black-soil-type degraded meadows in the headwater of the Qinghai-Tibetan Plateau, China

In alpine meadow ecosystems, considerable spatial heterogeneity in forb-dominant vegetation exists as a result of severe grassland degeneration; however, there is limited quantitative information on the vegetative differences between degenerated and pristine grasslands. Therefore, a field study, which seeks to identify the edaphic factors driving the variation in plant composition and distribution, was conducted in a severely degraded alpine meadow located in the Qinghai-Tibetan Plateau, NW China. Five meadows, an original meadow and four degraded meadows, were used to determine the differentiation and relationships between the vegetation and soil of degraded alpine meadows. The dominated species of these degraded meadows are Ligularia virgaurea–Artemisia gmelinii (LA), Oxytropis ochrocephala–Leontopodium nanum (OL), Aconitum pendulum–Potentilla anserina (AP) and Stellera chamaejasme–Artemisia nanschanica (SA), respectively. The results indicate that vegetation cover, grass biomass, species number and diversity indices clearly decrease from the original to the degraded meadow. Soil water, clay and nutrient content are also reduced with grassland degradation in surface and subsoil layers. The joint study of floristic and edaphic variables confirms that the soil features, especially the bulk density, sand content, pH, salinity, N and K, mainly determine the establishment of vegetation in the severely degraded fields of this study. These results may be useful for alpine grassland ecosystem restoration and management.     

Effects of Seagrass Rhizospheres on Sediment Redox Conditions in SE Asian Coastal Ecosystems

We examined the rhizosphere structure of 14 seagrass meadows (seven mixed, three Enhalus acoroides, two Zostera japonica, one Thalassia hemprichii, and one Halophila ovalis) in the Philippines and Vietnam and tested their effect on sediment redox potential by comparing the redox potential in vegetated vs unvegetated sediments. The effect of seagrass photosynthesis on sediment redox potential was tested in an E. acoroides meadow during a short-term (2-day) clipping experiment. In all the meadows, the centroidal depth (i.e., depth comprising 50%) of seagrass belowground biomass was within the top 15 cm sediment layer. Redox potentials in vegetated sediments tended to be higher than those in adjacent unvegetated ones; sediment redox potential anomaly ranged from −61 to 133 mV across the meadows. The centroidal depths of positive redox potential anomaly and seagrass root biomass were significantly correlated across the meadows investigated (type II regression analysis, slope = 0.90, lower confidence limit [CL] = 0.42 upper CL = 1.82, R ² = 0.59, p < 0.01). Experimental removal of E. acoroides leaves resulted in a decrease in rhizosphere redox potential by 20 mV, further confirming the positive effect of seagrass roots and rhizomes on sediment redox potential and, thus, the general conditions for microbial processes in the coastal zone.     

The REVEALS model,a new tool to estimate past regional plant abundance from pollen data in large lakes: validation in southern Sweden

The REVEALS model was developed to reconstruct quantitatively regional vegetation abundance (in a 10⁴–10⁵ km² area) from pollen assemblages in large lakes (≥100–500 ha). This model corrects for biases in pollen percentages caused by inter‐taxonomic differences in pollen productivity and dispersal. This paper presents the first case study to validate REVEALS, using empirical data from southern Sweden. Percentage cover of modern regional vegetation in Skåne and Småland, two contrasting vegetation regions, was predicted with REVEALS for 26 key taxa, using pollen assemblages from surface sediments in 10 large lakes, and compared to the actual vegetation within 10⁴ km² compiled from satellite data, forestry inventories, crop statistics, aerial photographs, and vegetation inventories. REVEALS works well in predicting the percentage cover of large vegetation units such as total trees (wooded land), total herbs (open land), total conifers and total broad‐leaved trees, and it provides reasonable estimates for individual taxa, including Pinus, Picea, Betula, Corylus, Alnus, Tilia, Salix spp., Juniperus, Poaceae, Cyperaceae, Cerealia and Secale. The results show great potential for REVEALS applications, including (1) quantitative reconstructions of past regional land cover important for palaeoclimatology and nature conservation, and (2) local‐scale reconstruction of vegetation (<1 km² up to ～ 5 km² area) relevant for palaeoecology and archaeology. Copyright © 2007 John Wiley & Sons, Ltd.     

Changes in Seagrass-associated fish and crustacean communities on Florida Bay mud banks: The effects of recent ecosystem changes?

The fauna of seagrass-covered mud banks in Florida Bay, documented in the mid 1980s prior to recent seagrass die-off, phytoplankton blooms, and other ecosystem changes, was reexamined in the mid 1990s for faunal changes that might be associated with environmental perturbations. During both decades, decapod crustaceans and fishes were collected with 1-m² throw traps from seagrass beds at six sites that differ in the amount of freshwater and/or marine influence and in seagrass community metrics. The most common faunal changes were declines in seagrass-canopydwelling forms and increases in benthic forms. At three sites with relatively lush seagrass meadows, above-ground seagrass standing crop declined and abundance of the benthic predatory fishOpsanus beta increased. The degree of faunal change among these sites appeared to be related either to salinity variability or to the degree of exposure to the ecosystem changes that have taken place in Florida Bay. At two sites with poorly developed seagrass meadows, seagrass standing crop and canopy height did not change significantly between decades, but there was an increase in shoot density and total leaf area. The animal communities at these sites were characterized by significant increases in the abundance of benthic crustaceans. At the site on the edge of Rankin Lake, the basin where seagrass die-off was first observed in Florida Bay during 1987, seagrass standing crop, canopy height, shoot density, and leaf area declined significantly between decades, but species richness of both crustaceans and fishes increased. The abundance of canopy-dwelling crustaceans and fishes declined markedly at this site, whereas the abundance of benthic forms less dependent on seagrass cover generally increased. In retrospect, we believe the fauma at this site during the 1980s, characterized by high productivity but few species, was already showing signs of the stresses that led to the seagrass die-off that began in 1987.     

A comparison of early juvenile red drum densities among various habitat types in Galveston Bay, Texas

Seagrass meadows are often cited as important nursery areas for newly settled red drum even though many estuaries, such as Galveston Bay, Texas, support large numbers of red drum and have limited seagrass cover, suggesting the use of alternate nursery areas. We examined patterns of habitat use for newly settled red drum at six sampling areas in Galveston Bay; two areas had seagrass beds and four areas had no seagrass. We measured densities in different habitat types using epibenthic sleds and enclosure samplers. Peak recruitment of young red drum to the estuary occurred during September through December. Highest densities of new settlers were found in seagrass meadows (primarilyHalodule wrightii), but when seagrass was absent, the highest densities of red drum occurred along theSpartina alterniflora marsh edge interface. Densities were relatively low on nonvegetated bottom away from the marsh edge. We also examined density patterns in other habitat types at selected sampling areas and found no red drum within marsh vegetation away from the marsh edge interface (5 and 10 m into the marsh interior). Oyster reefCrassostrea virginica was sampled using lift nets, and we found no red drum using this habitat, although adjacent seagrass and marsh interface habitats were used. Even though red drum densities in marsh edge were low relative to seagrass, the large areal extent of marshes in the bay complex probably makes marsh edge the most important nursery habitat for red drum in Galveston Bay.     

The effects of anthropogenic nutrient enrichment on turtle grass (Thalassia testudinum) in Sarasota Bay,Florida

Four meadows of turtle grass (Thalassia testudinum Banks ex Konig) in Sarasota Bay, Florida were sampled on a bimonthly basis from June 1992 to July 1993 to determine spatial and temporal variation in short shoot density, biomass, productivity, and epiphyte loads. Concurrent with the seagrass sampling, quarterly water-quality monitoring was undertaken at ≥3 sites in the vicinity of each studied seagrass meadow. Three months after termination of the seagrass sampling effort, a biweekly water-quality monitoring program was instituted at two of the seagrass sampling sites. In addition, a nitrogen loading model was calibrated for the various watersheds influencing the seagrass meadows. Substantial spatial and temporal differences in turtle grass parameters but smaller spatial variation in water quality parameters are indicated by data from both the concurrent quarterly monitoring program and the biweekly monitoring program instituted after termination of the seagrass study. Turtle grass biomass and productivity were negatively correlated with watershed nitrogen loads, while water quality parameters did not clearly reflect differences in watershed nutrient inputs. We suggest that traditional water-quality monitoring programs can fail to detect the onset or continuance of nutrient-induced declines in seagrass health. Consequently, seagrass meadows should be monitored directly as a part of any effort to determine status and/or trends in the health of estuarine environments. *** DIRECT SUPPORT *** A01BY074 00029

     

Direct evidence of imbalanced seagrass (Posidonia oceanica) shoot population dynamics in the Spanish Mediterranean

Direct census of shoots tagged in permanent plots was used to assess the present (2000–2002)Posidonia oceanica population dynamics in 25 meadows along the Spanish Mediterranean Coast. Shoot density ranged from 154±8 to 1,551±454 shoots m⁻², absolute shoot mortality from 5±0 to 249±53 shoots m⁻² yr⁻¹, and absolute shoot recruitment from <5 ±1 to 62±42 shoots m⁻²yr⁻¹. Specific shoot mortality and recruitment rates, which are mathematically and statistically (p>0.05) independent of shoot density, varied from 0.015±0.006 to 0.282±0.138 yr⁻¹ and 0.018±0.005 to 0.302±0.093 yr⁻¹, respectively. Absolute shoot mortality rate was scaled to shoot density (Pearson correlation, r=0.78, p<0.0001), and variability in specific shoot recruitment rate was partially due to differences in the percentage of growing apexes, which produce most of the recruits within the population (Pearson correlation, r=0.50, p<0.001), demonstrating the existence of structural constraints on shoot demography. Shoot half-life was estimated to range from 2.5 to 60.4 yr and meadow turnover times between 6.7 yr and more than a century, provided current estimates of shoot mortality, recruitment rates, and density remain uniform. There were differences in shoot mortality and recruitment at the regional scale, with the meadows developing along the coast of the Spanish mainland experiencing the highest shoot mortality (Tukey test, p<0.05) and tending to exhibit the highest shoot recruitment. The low shoot recruitment did not balance shoot mortality in most (60%) of the meadows, showing a prevalence of declining populations among the 25 meadows studied (Wilcoxon ranked sign test, p<0.0005). This study demonstrates the power of direct census of seagrass shoots in permanent plots to evaluate the present status of seagrass meadows, to detect on-going population decline, and to provide some insight onto the possible factors involved. The incorporation of direct census of seagrass meadows to monitoring programs will help provide the early-warning signals necessary to support management decisions to conserve seagrass meadows.     

Epiphyte-grazer relationships in seagrass meadows: Consequences for seagrass growth and production

Studies of seagrass meadows have shown that the production of algal epiphytes attached to seagrass blades approaches 20% of the seagrass production and that epiphytes are more important as food for associated fauna than are the more refractory seagrass blades. Since epiphytes may compete with seagrasses for light and water column nutrients, excessive epiphytic fouling could have serious consequences for seagrass growth. We summarize much of the literature on epiphytegrazer relationships in seagrass meadows within the context of seagrass growth and production. We also provide insights from mathematical modeling simulations of these relationships for a Chesapeake BayZostera marina meadow. Finally we focus on future research needs for more completely understanding the influences that epiphyte grazers have on seagrass production.     

Deterioration of Sediment Quality in Seagrass Meadows (<Emphasis Type="Italic">Posidonia oceanica</Emphasis>) Invaded by Macroalgae (<Emphasis Type="Italic">Caulerpa</Emphasis> sp.)

Species of the macroalgae Caulerpa sp. are increasingly being observed in meadows of the endemic Mediterranean seagrass Posidonia oceanica, and in particular Caulerpa taxifolia, has been considered as an invasive species leading to seagrass decline. Studies have so far failed to reveal the underlying mechanisms of the success of the macroalgae, and here, we examine how biogeochemical changes of the environment associated to indigenous (Caulerpa prolifera) and non-indigenous (Caulerpa racemosa and C. taxifolia) species affect the habitat of P. oceanica. Two of the species (C. prolifera and C. racemosa) affect the sediment biogeochemical conditions by increasing organic matter pools, microbial activity, and sulfide pools of the sediments, and limited effects were found for C. taxifolia. Biomass of the macroalgae contributed to the extent of impacts, and high sulfide invasion into the seagrasses and regression of the meadow were pronounced at the location with the highest Caulerpa biomass. This suggests that Caulerpa invasion contributes to seagrass decline probably because Caulerpa thrives better than the seagrasses in the modified environment.     

Oceanographic controls on shallow‐water temperate carbonate sedimentation: Spencer Gulf,South Australia

Spencer Gulf is a large (ca 22 000 km²), shallow (<60 m water depth) embayment with active heterozoan carbonate sedimentation. Gulf waters are metahaline (salinities 39 to 47‰) and warm‐temperate (ca 12 to ?28°C) with inverse estuarine circulation. The integrated approach of facies analysis paired with high‐resolution, monthly oceanographic data sets is used to pinpoint controls on sedimentation patterns with more confidence than heretofore possible for temperate systems. Biofragments – mainly bivalves, benthic foraminifera, bryozoans, coralline algae and echinoids – accumulate in five benthic environments: luxuriant seagrass meadows, patchy seagrass sand flats, rhodolith pavements, open gravel/sand plains and muddy seafloors. The biotic diversity of Spencer Gulf is remarkably high, considering the elevated seawater salinities. Echinoids and coralline algae (traditionally considered stenohaline organisms) are ubiquitous. Euphotic zone depth is interpreted as the primary control on environmental distribution, whereas seawater salinity, temperature, hydrodynamics and nutrient availability are viewed as secondary controls. Luxuriant seagrass meadows with carbonate muddy sands dominate brightly lit seafloors where waters have relatively low nutrient concentrations (ca 0 to 1 mg Chl‐a m^?3). Low‐diversity bivalve‐dominated deposits occur in meadows with highest seawater salinities and temperatures (43 to 47‰, up to 28°C). Patchy seagrass sand flats cover less‐illuminated seafloors. Open gravel/sand plains contain coarse bivalve–bryozoan sediments, interpreted as subphotic deposits, in waters with near normal marine salinities and moderate trophic resources (0·5 to 1·6 mg Chl‐a m^?3) to support diverse suspension feeders. Rhodolith pavements (coralline algal gravels) form where seagrass growth is arrested, either because of decreased water clarity due to elevated nutrients and associated phytoplankton growth (0·6 to 2 mg Chl‐a m^?3), or bottom waters that are too energetic for seagrasses (currents up to 2 m sec^?1). Muddy seafloors occur in low‐energy areas below the euphotic zone. The relationships between oceanographic influences and depositional patterns outlined in Spencer Gulf are valuable for environmental interpretations of other recent and ancient (particularly Neogene) high‐salinity and temperate carbonate systems worldwide.     

Role of larger herbvores in seagrass communities

The nutritional ecology of macroherbivores in seagrass meadows and the roles of grazing by urchins, fishes and green turtles in tropical systems and waterfowl in temperate systems are discussed in this review. Only a few species of animals graze on living seagrasses, and apparently only a small portion of the energy and nutrients in seagrasses is usually channeled through these herbivores. The general paucity of direct seagrass grazers may be a function of several factors in the composition of seagrasses, including availability of nitrogen compounds, presence of relatively high amounts of structural cell walls, and presence of toxic or inhibitory substances. The macroherbivores, however, can have a profound effect on the seagrass plants, on other grazers and fauna associated with the meadow, and on chemical and decompositional processes occurring within the meadow. Grazing can alter the nutrient content and digestibility of the plant, as well as its productivity. Removal of leaf material can influence interrelations among permanent and transient faunal residents. Grazing also interrupts the detritus cycle. Possible consequences of this disruption, either through acceleration or through decreased source input, and the enhancement of intersystem coupling by increased export and offsite fecal production, are discussed. The extent and magnitude of these effects and their ecological significance in the overall functioning of seagrass meadows only can be speculated, and probably are not uniform or of similar importance in both tropical and temperate seagrass systems. However, areas grazed by large herbivores provide natural experiments in which to test hypotheses on many functional relations in seagrass meadows.     

Capybara (<Emphasis Type="Italic">Hydrochaeris hydrochaeris</Emphasis> rodentia: Hydrochaeridae): A mammalian seagrass herbivore

This note reports an unknown trophic interaction between a mammalian herbivore, the capybara (Hydrochaeris hydrochaeris), and the seagrassRuppia maritima (wigeongrass) and compares the feeding behavior of capybaras to other seagrass grazers. Observations were made in Spring 2002 in the Barra Grande, a small, shallow, moderately stratified, bar-built estuary at Ilha Grande, Rio de Janeiro State, southeast Brazil. The activities of the capybaras were investigated and grazing impacts were quantified in situ. The capybaras were observed feeding onR. maritima during the day and aquatic feeding alternated with periods of feeding on land.R. maritima was the only submerged aquatic vegetation to be consumed by the capybaras. The feeding activity of the capybaras on wige ongrass consisted of alternately diving down to theR. maritima then surfacing; the capybaras spent a significantly greater amount of time under water. In the area where the capybaras foraged 18.1% of the seagrass meadow showed recent grazing scars. Vegetation of recently and not recently grazed areas were compared. Grazing scars, which were slightly elongated, were not completely devoid ofR. maritima but presented reduced standing crop: canopy height, shoot density, and shoot, rhizome, and root biomass were reduced in grazed areas. The grazing patterns observed in capybaras resembled those previously reported in the sirenia, mammals that include two seagrass-eating species.     

A new method to describe seagrass habitat sampled during fisheries-independent monitoring

We explain a new method of quantifying seagrass cover and describing seagrass species composition during fisheries-independent monitoring. This new method is similar to a point-intercept method developed to estimate arboreal crown cover, but it uses an aquascope designed for shallow water. The method does not require a diver. Seagrass cover (cover ratio) distinguished different percentage cover categories in 0.25-m² seagrass plots. Estimates of species composition determined by using the new method were most similar to those obtained by using estimates of aboveground biomass. Within each 141-m² area sampled with a 21.3-m fish seine, we accurately estimated seagrass cover ratio and species composition with six observations that typically required less than 6 total minutes. Within such areas, 42 trials were conducted to evaluate the precision with which different observers estimated seagrass cover ratio and species composition. In 98% of the trials, observers attained statistically similar estimates of cover ratio, and in 100% of the trials in areas with multiple seagrass species, observers attained statistically similar estimates of species composition. We conclude that the new method provided efficient and reasonably accurate means to quantify seagrass cover and species composition.     

Modeling nonlinear seagrass clonal growth: Assessing the efficiency of space occupation across the seagrass flora

The clonal growth of 9 seagrass species was modeled using a simulation model based on observed clonal growth rules (i.e., spacer length, rhizome elongation rates, branching rates, branching angle) and shoot mortality rates for seagrass species. The results of the model confirmed the occurrence of complex, nonlinear growth of seagrass clones derived from internal dynamics of space occupation. The modeled clones progressed from a diffuse-limited aggregation (DLA), dendritic growth, identified with a guerrilla strategy of space occupation, to a compact (Eden) growth, comparable to the phalanx strategy of space occupation, once internal recolonization of gaps, left by dead shoots within the clone, begins. The time at which seagrass clones shifted from diffuse limited to compact growth was predictable from the branching angle and frequency of the species and varied from 1 yr to several decades among species. As a consequence the growth behavior and the apparent growth strategy of the species changes with the development of the clones. The results of the model demonstrate that the emergent complexity of seagrass clonal growth is contained within the simple set of growth rules that can be used to represent clonal growth.