The distribution and expansion of the invasive seagrass Halophila stipulacea in Dominica, West Indies, with a preliminary report from St. Lucia

The seagrass Halophila stipulacea Forsskål, native to the Red Sea, is an invasive species in the Mediterranean that was recently observed offshore Grenada, in the Caribbean. Here, we document the presence of this seagrass in Dominica and St. Lucia, demonstrating it has spread across part of the eastern Caribbean. H. stipulacea in Dominica was present in seven locations along the west coast covering more than 22.9 ha of the benthos, at depths from 2 to 18 m. Populations were concentrated in or adjacent to bays frequented by recreational or commercial boats, likely vectors for the introduction. Morphological features varied from bed to bed, with depth being the predominant driving factor. H. stipulacea had a rapid mean lateral bed expansion rate of 0.5 cm d⁻¹, with a maximum rate of >6 cm d⁻¹. H. stipulacea patches often occurred exclusive of the otherwise dominant seagrasses of the Caribbean. The potential for the expansion of H. stipulacea, combined with its tolerance for a wide spectrum of environmental conditions, positions it as a potential threat to local and regional biodiversity.     

The short-term influence of herbivory near patch reefs varies between seagrass species

The coexistence of multiple species within a trophic level can be regulated by consumer preferences and nutrient supply, but the influence of these factors on the co-occurrence of seagrass species is not well understood. We examined the biomass and density responses of two seagrass species in the Florida Keys Reef Tract to grazing pressure near patch reefs, and evaluated how nutrient enrichment impacted herbivory dynamics. We transplanted Halodule wrightii (shoalgrass) sprigs into caged and uncaged plots in a Thalassia testudinum (turtlegrass) bed near a patch reef. Nutrients (N and P) were added to half of the experimental plots. We recorded changes in seagrass shoot density, and after three months, we measured above- and belowground biomass and tissue nutrient content of both species. Herbivory immediately and strongly impacted H. wrightii. Within six days of transplantation, herbivory reduced the density of uncaged H. wrightii by over 80%, resulting in a decrease in above- and belowground biomass of nearly an order of magnitude. T. testudinum shoot density and belowground biomass were not affected by herbivory, but aboveground biomass and leaf surface area were higher within cages, suggesting that although herbivory influenced both seagrass species, T. testudinum was more resistant to herbivory pressure than H. wrightii. Nutrient addition did not alter herbivory rates or the biomass of either species over the short-term duration of this study. In both species, nutrient addition had little effect on the tissue nutrient content of seagrass leaves, and N:P was near the 30:1 threshold that suggested a balance between N and P. The different impacts of grazing on these two seagrass species suggest that herbivory may be an important regulator of the distribution of multiple seagrass species near herbivore refuges like patch reefs in the Caribbean.     

Monitoring of physically restored seagrass meadows reveals a slow rate of recovery for Thalassia testudinum

Physical damage by motor vessels is a widespread problem for seagrass meadows, with hull and propeller strikes accounting for thousands of acres of impaired habitat in Florida, United States, alone. Because the excavations can become topographically unstable, and because recolonization and succession of seagrasses can require decades to reach climax, Thalassia testudinum‐dominated communities, there has been increasing efforts to regrade and stabilize impacted sediments, and to speed succession. A prior project involving eight vessel groundings in two hydrodynamic settings (high and low energy) examined the relative efficacy of capping injuries with sand‐filled fabric tubes or limestone pea rock, followed by planting of fast‐growing seagrass species and nutrient amendment using bird‐roosting stake deployments. Monitoring after 4 years showed recruitment of fast‐growing, subordinate species: Syringodium filiforme or Halodule wrightii, particularly in low‐energy environments; however, T. testudinum had not yet returned to natural densities. The current study extended monitoring an additional 3 years. At 7 years posteffort, T. testudinum recolonization was still incomplete. Of the eight sites, only three had statistically recovered. In low‐energy areas, H. wrightii cover was greater than in reference meadows, and a strong inverse relationship between H. wrightii and T. testudinum was observed. One explanation is that residual nutrients from bird feces switched the competitive outcome. We demonstrate using seagrass tissue N:P that nutrients delivered via bird stakes remained in the sediments of low‐energy environments, and argue that prolonged fertilization resulted in competitive advantages for H. wrightii, depressing T. testudinum recruitment and delaying recovery of the targeted seagrass community.     

Asexual propagation in the coral reef macroalga Halimeda (Chlorophyta, Bryopsidales): production, dispersal and attachment of small fragments

Siphonous, green macroalgae of the genus Halimeda are ubiquitous and ecologically important in tropical and subtropical marine environments. It has been hypothesized that the abundance of Halimeda on coral reefs is in part due to the ability of this genus to propagate asexually via vegetative fragmentation. However, vegetative fragmentation has only been documented for H. discoidea in a laboratory setting. To test the hypothesis that vegetative fragmentation contributes to field populations of Halimeda, we examined three aspects of fragmentation by H. tuna (Ellis and Solander) Lamouroux, H. opuntia (Linneaus) Lamouroux and H. goreaui Taylor on Conch Reef in the Florida Keys: (1) short-term (8 days) and long-term (14 weeks) fragment survival and rhizoid production in the laboratory and field (7 and 21 m), (2) size of the fragment pool and (3) influences of herbivory and water motion on production and dispersal of fragments. Although morphologically similar to H. discoidea, only a small percentage of H. tuna fragments survived. Fragments of H. opuntia and H. goreaui were more robust, and survival and rhizoid production were positively correlated with size in short-term trials. In 14-week field trials, one-third or fewer fragments of any species survived at 7 m, potentially because fragments were covered by large amounts of sediment. Survivors included some buried, seemingly dead individuals that turned green when exposed to light, highlighting the remarkable ability of this genus to survive disturbances. There was much less sediment accumulation at 21 m, where more fragments survived. Most (93%) eight-segment fragments of H. opuntia produced attachment rhizoids by the end of the 14-week trial. Overall, a range of 4.7-9.4 fragments of Halimeda m⁻² day⁻¹ were found on Conch Reef; most fragments were generated by H. goreaui. Fish bite marks were evident on 75-85% of the individuals of H. tuna and the number of bites per thallus ranged from 1 to 23. Herbivorous reef fish commonly fed on all three species of Halimeda. Some fish consumed the biomass, while others rejected most bites. For example, 83% of bites were rejected by the blue-striped grunt. Dispersal distances for rejected bites ranged from 0 to 31 m. Water motion was also responsible for fragment dispersal; experimentally produced fragments moved up to 48 cm day⁻¹. Results presented here suggest that asexual propagation of fragments of Halimeda is an important component of the life-history of this genus and vegetative fragmentation contributes to the abundance of this genus on coral reefs.     

Environmentally-controlled,density-dependent secondary dispersal in a local estuarine crab population

The mechanisms driving the pelagic secondary dispersal of aquatic organisms following initial settlement to benthic habitats are poorly characterized. We examined the physical environmental (wind, diel cycle, tidal phase) and biological (ontogenetic, density-dependent) factors that contribute to the secondary dispersal of a benthic marine invertebrate, the blue crab (Callinectes sapidus) in Pamlico Sound, NC, USA. Field studies conducted in relatively large (0.05 km²) seagrass beds determined that secondary dispersal is primarily undertaken by the earliest juvenile blue crab instar stages (J1 crabs). These crabs emigrated pelagically from seagrass settlement habitats using nighttime flood tides during average wind conditions (speed ~5 m s^–1). Moreover, the secondary dispersal of J1 crabs was density-dependent and regulated by intra-cohort (J1) crab density in seagrass. Our results suggest that dispersal occurs rapidly following settlement, and promotes blue crab metapopulation persistence by redistributing juveniles from high-density settlement habitats to areas characterized by low postlarval supply. Collectively, these data indicate that blue crab secondary dispersal is an active process under behavioral control and can alter initial distribution patterns established during settlement. This study highlights the necessity of considering secondary dispersal in ecological studies to improve our understanding of population dynamics of benthic organisms.     

Demographic and genetic connectivity: the role and consequences of reproduction,dispersal and recruitment in seagrasses

Accurate estimation of connectivity among populations is fundamental for determining the drivers of population resilience, genetic diversity, adaptation and speciation. However the separation and quantification of contemporary versus historical connectivity remains a major challenge. This review focuses on marine angiosperms, seagrasses, that are fundamental to the health and productivity of temperate and tropical coastal marine environments globally. Our objective is to understand better the role of sexual reproduction and recruitment in influencing demographic and genetic connectivity among seagrass populations through an integrated multidisciplinary assessment of our present ecological, genetic, and demographic understanding, with hydrodynamic modelling of transport. We investigate (i) the demographic consequences of sexual reproduction, dispersal and recruitment in seagrasses, (ii) contemporary transport of seagrass pollen, fruits and seed, and vegetative fragments with a focus on hydrodynamic and particle transport models, and (iii) contemporary genetic connectivity among seagrass meadows as inferred through the application of genetic markers. New approaches are reviewed, followed by a summary outlining future directions for research: integrating seascape genetic approaches; incorporating hydrodynamic modelling for dispersal of pollen, seeds and vegetative fragments; integrating studies across broader geographic ranges; and incorporating non‐equilibrium modelling. These approaches will lead to a more integrated understanding of the role of contemporary dispersal and recruitment in the persistence and evolution of seagrasses.     

Resource translocation within seagrass clones: allometric scaling to plant size and productivity

The allometric scaling of resource demand and translocation within seagrass clones to plant size (i.e. shoot mass and rhizome diameter), shoot production and leaf turnover was examined in situ in eight seagrass species (Cymodocea nodosa, Cymodocea serrulata, Halophila stipulacea, Halodule uninervis, Posidonia oceanica, Thalassodendron ciliatum, Thalassia hemprichii and Zostera noltii), encompassing most of the size range present in seagrass flora. One fully developed shoot on each experimental rhizome was incubated for 2–3 h with a pulse of NaH¹³CO₃ (235 μmol) and ¹⁵NH₄Cl (40 μmol). The mobilisation of incorporated tracers across the clone was examined 4 days later. Carbon and nitrogen demand for shoot production across seagrass species scaled at half of the shoot mass, whereas seagrass leaves incorporated tracers (¹³C and ¹⁵N) at rates proportional to the shoot mass. The shoots of all seagrass species shared resources with neighbours, particularly with younger ones. The time scales of physiological integration and the absolute amount of resources shared by seagrass ramets scaled at 2.5 power of the rhizome diameter. Hence, the ramets of larger species were physiologically connected for longer time scales and share larger absolute amounts of resources with neighbours than those of smaller species. The different pattern of resource translocation exhibited by seagrasses helps explain the ecological role displayed by these species and the success of large seagrasses colonising nutrient-poor coastal areas, where they often dominate.     

Can floral consumption by fish shape traits of seagrass flowers?

Seagrasses are marine flowering plants with hydrophilous pollination. This abiotic pollination by water assumes absence of flower-animal interaction, but animals can interfere in this process through consumption of reproductive structures. We studied predation on male flowers by fish for three dioecious seagrass species (Thalassia testudinum, Syringodium filiforme and Halodule wrightii) in the Mexican Caribbean. Seagrass flowers have a highly reduced or absent corolla and florivores directly consumed the anthers with pollen. The foliar structures (tepals, bracts or sheaths) protecting the male flower buds were removed by hand in situ. The floral buds were followed by videos or taking pictures at regular intervals and most (56–100 %, depending on seagrass species and experimental setting) artificially denuded male flower buds were consumed within 24 h by juvenile fish of various species. Histochemical analysis showed that the pollen and embedding mucilage were rich in polysaccharides and proteins, thus potentially nutritious. The seagrasses had copious production of pollen (between 0.2 and 1.2 × 10⁶ pollen per flower, depending on the species). But T. testudinum and S. filiforme were often pollen limited, and the probability of fruit set was reduced ~50 % when the females were at the distance of 1 and 5–6 m from the males flowers, respectively. Under natural conditions, depredation on pre-anthesis male flowers in the three species was low because flower bud emergence (few hours) and pollen release (1–4 h) were ephemeral processes. In addition, the release of pollen of T. testudinum occurred at dusk when herbivorous fish became inactive. These life-cycle characteristics aid to avoid excessive pollen consumption by fish, however, whether they are anti-predator strategies or mere adaptations for submarine pollination remains to be established.     

Starch grain morphology of the seagrasses Halodule wrightii, Ruppia maritima, Syringodium filiforme, and Thalassia testudinum

Starch grains are a ubiquitous component of plants that have been used in tandem with phytoliths, pollen, and macrofossils to reconstruct past floral diversity. This tool has yet to be fully explored for aquatic plants, specifically seagrasses, which lack phytoliths and are rarely preserved as macrofossils or pollen. If starch grains in seagrasses are morphologically distinct, this method has the potential to improve seagrass identification in the fossil record in such cases where its starch is preserved (e.g. scratches and occlusal surfaces of tooth enamel from seagrass consumers). The goals of this study were twofold: (1) to determine if starch is present in seagrass material and (2) to assess how starch grain morphology differs between different seagrasses.This study focused on four abundant and ecologically distinct seagrasses from the Caribbean: Halodule wrightii, Ruppia maritima, Syringodium filiforme, and Thalassia testudinum. Starch grains were observed in all species except S. filiforme. Grains from H. wrightii are typically observed in side-on orientation, are sub-round to angular, and are fairly small (3-19 μm, end-on). Grains of R. maritima are small spherical grains (4-8 μm) that have a centric hilum and a straight extinction cross with a median angle between the arms of 90°. Grains from T. testudinum are large (9-31 μm, end-on), conical in side-on and round/sub-round in end-on orientation, have a slightly eccentric hilum with an obvious particle, and prominent lamellae.Visual assessment and comparative statistics demonstrate that the morphology of starch grains from T. testudinum, R. maritima, and H. wrightii are significantly different. With more extensive research, there is potential for the positive identification of starch grains from an unknown seagrass. The ability to identify seagrass from starch grains could facilitate the identification of seagrasses in the fossil record and supply information on seagrass evolution and distribution, climate effects on seagrass distribution, and the diets of seagrass consumers.     

State of Thalassia testudinum Banks ex König meadows in the Veracruz Reef System,Veracruz, México

The vegetative development of the seagrass Thalassia testudinum was assessed in three reefs (Hornos, isla Sacrificios, isla de Enmedio) of the Veracruz Reef System (Veracruz, México) which are located at increasing distances from the mainland coast. Leaf nutrient content of T. testudinum suggests that the availability of nitrogen and phosphorus increases from Hornos reef to isla Sacrificios and isla de Enmedio, that is, as the distance to the coast increases. The total biomass of this species tended to increase similarly as a result of the large increase of the biomass of rhizomes and roots but not of leaves. In contrast to previous knowledge of the response of seagrasses along gradients of nutrient availability, the rhizome and roots to shoot biomass ratio increased by a factor of two as nutrient availability increased. The density of T. testudinum shoots and their mass, the LAI and leaf productivity, and the average number of leaves produced by a T. testudinum shoot in 1 year were lowest in the reef closest to shore (Hornos). Our results show that the vegetative development of T. testudinum in Hornos reef is restrained when compared with that in isla Sacrificios and isla de Enmedio and suggest that the differences of vegetative development of this species in the Veracruz Reef System might be driven by factors other than nutrient availability.     

Synergistic effects of high temperature and sulfide on tropical seagrass

To examine the synergism of high temperature and sulfide on two dominant tropical seagrass species, a large-scale mesocosm experiment was conducted in which sulfide accumulation rates (SAR) were increased by adding labile carbon (glucose) to intact seagrass sediment cores across a range of temperatures. During the initial 10 d of the 38 d experiment, porewater SAR in cores increased 2- to 3-fold from 44 and 136 μmol L^− 1 d^− 1 at 28-29 °C to 80 and 308 μmol L^− 1 d^− 1 at 34-35 °C in Halodule wrightii and Thalassia testudinum cores, respectively. Labile C additions to the sediment resulted in SAR of 443 and 601 μmol L^− 1 d^− 1 at 28-29 °C and 758 to 1,557 μmol L^− 1 d^− 1 at 34-35 °C in H. wrightii and T. testudinum cores, respectively. Both T. testudinum and H. wrightii were highly thermal tolerant, demonstrating their tropical affinities and potential to adapt to high temperatures. While plants survived the 38 d temperature treatments, there was a clear thermal threshold above 33 °C where T. testudinum growth declined and leaf quantum efficiencies (Fv/Fm) fell below 0.7. At this threshold temperature, H. wrightii maintained shoot densities and leaf quantum efficiencies. Although H. wrightii showed a greater tolerance to high temperature, T. testudinum had a greater capacity to sustain biomass and short shoots under thermal stress with labile C enrichment, regardless of the fact that sulfide levels in the T. testudinum cores were 2 times higher than in the H. wrightii cores. Tropical seagrass tolerance to elevated temperatures, predicted in the future with global warming, should be considered in the context of the sediment-plant complex which incorporates the synergism of plant physiological responses and shifts in sulfur biogeochemistry leading to increased plant exposure to sulfides, a known toxin.     

A method for the culture of tropical seagrasses

Three tropical seagrass species were planted into 1.5 m² culture tanks and grown under the same conditions for 2 years. New shoot production and vegetation growth of both Syringodium filiforme Kütz. and Halodule wrightii Aschers. resulted in complete cover in monoculture tanks within the first year. The vegetative spread of Thalassia testudinum Banks ex König was slower than that of the other species. The culture of seagrasses in open mesocosm systems was most successful when continuous current circulation was maintained, water column nutrients were kept low, and extreme high temperatures (> 36°C) were avoided. Seagrass colonized and grew equally well in Indian River mud substratum and in quartz sand.     

Go with the flow: Fragment retention patterns shape the vegetative dispersal of aquatic plants in lowland streams

1. The dispersal of aquatic plant propagules is highly facilitated in streams due to flow. As many aquatic plants predominantly spread through vegetative propagules, the specific retention and thus drift distance of dispersed plant fragments largely contribute to the rapid spread along the course of a stream.
2. We determined fragment retention for four aquatic plant species (Elodea canadensis, Myriophyllum spicatum, Ceratophyllum demersum, Salvinia natans; representing four different common morpho-structural groups) in sections of small to medium-sized German streams with different levels of stream sinuosity.
3. The number of fragments showed a logistic decline over drift distance. In two small streams, 90% of drifting fragments were retained at distances (D₉₀) of only 5–9 m and 19–70 m, while higher D₉₀ values of 116–903 m and 153–2,367 m were determined for sections of a medium-sized stream. The likelihood of retention thereby decreased significantly with increasing stream size and was reduced in straightened stream sections.
4. Differences in retention were more strongly related to fragment buoyancy rather than fragment size and morphology. Increasing buoyancy significantly lowered the likelihood of fragment retention over drift distance by a factor of 3–8, whereas contrasting effects were documented for size and morphology of fragments.
5. The relevance of different obstacles was highly stream section-specific and depended on obstacle abundance, distribution, and the degree of submergence/emergence.
6. Our findings elucidate the dynamic retention patterns of plant fragments and highlight the strong interplay between extrinsic (stream) and intrinsic (fragment) properties. We conclude that straightened lowland streams of intermediate size promote the rapid dispersal of invasive aquatic plants and are particularly prone to invaders producing large amounts of small and highly buoyant plant fragments. Information on the species-specific fragment colonisation dynamics in the field is further required to improve our understanding of the vegetative dispersal capacity of invasive aquatic plants in stream ecosystems.

     

Biochemical studies on Brachionus plicatilis: hydrolytic enzymes,integument proteins and composition of trophi

In Terminos Lagoon, México, more than 80 fish species use the mangrove and seagrass habitats. We studied nekton dynamics in an inlet seagrass system and a more sheltered seagrass/mangrove system located behind a barrier island. Seasonal community biomass ranges from 0.6 to 5.2 g wet wt m^-2. For the two habitats together, there are 28 dominant species. Eleven species were common to both areas: Sphoeroides testudineus (Linnaeus, 1758), Archosargus rhomboidalis Linnaeus, 1758, A. probatocephalus (Walbaum, 1792), Arius felis (Linnaeus, 1766), A. melanopus (Gunther, 1864), Eucinostomus gula (Cuvier & Valenciennes, 1830), Bairdiella chrysoura (Lacépède, 1803), Orthopristis chrysoptera (Linnaeus, 1766), Chilomycterus schoepfi (Walbaum, 1792), Opsanus beta (Goode & Bean, 1879) and Lutjanus griseus (Linnaeus, 1758). Ten species used exclusively the inlet seagrass system: Urolophus jamaicensis (Cuvier, 1817), Haemulon aurolineatum (Cuvier, 1829), H. bonariense (Cuvier, 1830), H. plumieri (Lacépède, 1802), Anisotremus virginicus (Linnaeus, 1758), Odontoscion dentex (Cuvier, 1830), Corvula sanctae-luciae (Jordan, 1890), Nicholsina usta (Valenciennes, 1839), Stephanolepis hispidus (Linnaeus, 1766) and Diodon hystrix Linnaeus, 1758. Seven species were dominant only in the seagrass/mangrove system: Anchoa mitchilli (Cuvier & Valenciennes, 1848), Scorpaena plumieri Bloch, 1789, Cynoscion nebulosus (Cuvier & Valenciennes, 1830), Diapterus rhombeus (Cuvier & Valenciennes, 1830), Bairdiella ronchus (Cuvier & Valenciennes, 1830), Cichlasoma urophtalmus (Günther, 1862) and Acanthostracion quadricornis (Linnaeus, 1758). Comparative analysis showed that periodic variation in biomass and diversity of fish assemblages in seagrass and seagrass/mangrove habitats were synchronized with sizes and densities of population, season of the year (dry, wet, ‘nortes’), circulation pattern, and patterns of primary production (phytoplankton, Thalassia testudinum Konig, 1805; and Rhizophora mangle Linnaeus). This analysis allowed the definition of 3 life-cycle patterns with a clear nektonic ‘seasonal programming’ following the timing of primary production in these critical habitats: (1) marine species which spawn in or near the inlet with eggs and larvae transported into and distributed throughout the lagoon by the predominant currents; (2) estuarine-marine species which spawn in different habitats of the lagoon and use the seagrass/mangrove system as a nursery area, and (3) species which complete their life history in the inlet seagrass and/or seagrass/mangrove systems. EPOMEX Program Contribution No. 0026.     

Nutrient enrichment promotes survival and dispersal of drifting fragments in an invasive tropical macroalga

The effect of nutrient availability on growth, survival, and photosynthetic performance of drifting fragments of the invasive red alga Hypnea musciformis was studied in Maui (Hawaii), where this species smothers native reef communities and forms localized blooms. H. musciformis does not sexually reproduce in Hawaii and drifting fragments represent the only pathway by which H. musciformis can disperse and invade new areas. Growth rates decreased with age and approached zero when fragments aged 32 days. Increased nutrient availability did not result in increased relative growth rates during this period. In contrast to growth, photosynthetic performance remained unaffected through time and showed no clear relationship with nutrient availability. Increased nutrient availability increased fragment survival and fragments survived for >2 months in the high nutrient treatment (3.0 μmolPO₄ + 30.0 μmolNH₄). This indicates that increased nutrient availability increases the dispersal potential of H. musciformis. Low growth rates of drifting Hypnea fragments increased recruitment success since attachment success of this epiphytic species decreased with increasing fragment size. H. musciformis thus uses resources for survival and maintenance rather than growth, resulting in long competency periods and optimal recruitment, which likely contribute to its success as an invader of Hawaiian reef communities.     

Ecosystem Structure and Function Still Altered Two Decades After Short-Term Fertilization of a Seagrass Meadow

An oligotrophic phosphorus (P) limited seagrass ecosystem in Florida Bay was experimentally fertilized in a unique way. Perches were installed to encourage seabirds to roost and deliver an external source of nutrients via defecation. Two treatments were examined: (1) a chronic 23-year fertilization and (2) an earlier 28-month fertilization that was discontinued when the chronic treatment was initiated. Because of the low mobility of P in carbonate sediments, we hypothesized long-term changes to ecosystem structure and function in both treatments. Structural changes in the chronic treatment included a shift in the dominant seagrass species from Thalassia testudinum to Halodule wrightii, large increases in epiphytic biomass and sediment chlorophyll-a, and a decline in species richness. Functional changes included increased benthic metabolism and quantum efficiency. Initial changes in the 28-month fertilization were similar, but after 23 years of nutrient depuration T. testudinum has reestablished itself as the dominant species. However, P remains elevated in the sediment and H. wrightii has maintained a presence. Functionally the discontinued treatment remains altered. Biomass exceeds that in the chronic treatment and indices of productivity, elevated relative to control, are not different from the chronic fertilization. Cessation of nutrient loading has resulted in a superficial return to the pre-disturbance character of the community, but due to the nature of P cycles functional changes persist.     

In situ nitrogen fixation associated with seagrasses in the Gulf of Elat (Red Sea)

In situ nitrogen fixation associated with the seagrass Halophila stipulacea, at the northern Gulf of Elat (Red Sea), is eight to ten times higher than that of nearby plant-free areas. A daily cycle of nitrogen fixation is evident, with rates during the day being seven times greater than during the night. Removal of seagrass leaves only from a patch within a seagrass bed gradually decreases nitrogen fixation activity, reaching the rates of plant-free areas after ten hours. A method devised for the in situ measurement of nitrogen fixation rates using belljars is described in detail. Nitrogen fixation rates in situ are higher than in the laboratory and lack the lag period typical to laboratory measurements. In laboratory experiments using intact plant samples, glucose enhances nitrogen fixation rates both in light and dark. Photosystem II inhibitor (3-3,4-dichloro-phenyl-1,1-dimethylurea) doubles nitrogen fixation rates in light. Both field and laboratory results indicate that light is essential for nitrogen fixation activity in the H. stipulacea bed possibly through its effect on cyanobacterial population that occupy the aerobic niches of the phyllosphere and on photosynthetic Rhodospirillacean bacteria that inhabit the anaerobic ones. Nitrogen fixation rates evident in H. stipulacea beds in situ account for a considerable portion of the biomass production by the seagrasses. The dependence of high nitrogenase activity by the diazotrophs on the presence of the seagrasses indicates the great importance of the seagrass community to the nitrogen cycle in its highly oligotrophic surroundings of the Gulf of Elat.     

Effects of elevated temperature on seasonal in situ leaf productivity of Thalassia testudinum Banks ex König and Syringodium filiforme Kützing

Net leaf productivities of Thalassia testudinum Banks ex König and Syringodium filiforme Kützing, measured in situ over a 1-year period in a sub-tropical estuary that receives thermal addition from a local power plant, were significantly temperature dependent. Primary production in both species followed seasonal temperature variation, with increase in leaf dry weight as a proportion of total leaf biomass ranging from 0.12% to 2.54% day^?1 for T. testudinum and 0.33% to 3.80% day^?1 for S. filiforme. From September to May, S. filiforme exhibited significantly higher productivity at the thermally-impacted (experimental) station than at the control station. However, from June to August, S. filiforme productivity at the experimental station was significantly lower than at the control station. T. testudinum productivity showed similar trends, but interstation differences were not statistically different. Maximal growth occurred between 23 and 29°C for S. filiforme and 23 and 31°C for T. testudinum. Thus, the thermal addition generally enhanced the seagrass productivity of both species in the autumn, winter, and spring, but reduced that of S. filiforme in the summer, as the optimal temperature range of this species was exceeded. The fact that T. testudinum exhibited a less marked response to the thermal addition and had a greater optimal temperature range than S. filiforme supports its status as the more eurythermal of the 2 species.     

DIFFERENTIATION IN RESPONSE TO CHILLING TEMPERATURES AMONG POPULATIONS OF THREE MARINE SPERMATOPHYTES,THALASSIA TESTUDINUM,SYRINGODIUM FILIFORME AND HALODULE WRIGHTII

Upon exposure to chilling conditions, the seagrass populations of Thalassia testudinum Banks ex König, Syringodium filiforme Kütz., and Halodule wrightii Aschers. showed various amounts of leaf and plant damage that correlated with their origin in the Gulf of Mexico-Caribbean. Populations of more tropical origin in the southern Gulf and Caribbean showed the most chill damage and those of the northern Gulf showed the least injury from the exposure to low temperatures. Of the three seagrasses, Halodule showed greatest chill tolerance, Syringodium showed the least tolerance and Thalassia was intermediate. The uptake of ¹⁴C by leaves following exposure to chilling temperatures showed quantitative differences that correlated with the amount of leaf damage in the various populations. No significant changes in the fatty acids in total lipid extracts were noted in the Texas seagrasses after chilling and, therefore, their resistance to low temperature damage did not relate to changes in saturation of fatty acids. Although the growing conditions slightly altered the severity of the chilling effects, the differentiation of response to chilling among the seagrass populations is based on inherited properties.     

Fish larvae settling in seagrass: Effects of leaf density and an epiphytic alga

Settlement of fish to seagrass with low leaf densities was investigated by a field experiment in Botany Bay, NSW. Artificial seagrass units (ASU) were used to simulate isolated seagrass habitats of 0, 25, 50, 100, 200 and 400 leaves m^-2. A total of 369 fish of 21 species settled in the experimental units during a 6 week period. The labrid, Achoerodus viridis, made up almost 70% of these individuals. The abundance of recently settled A. viridis increased greatly between 0 and 25 leaves m ^-2, At more than 25 leaves m^-2, A. viridis settled in approximately equal numbers. Patterns of settlement for all other species combined were similar to those for A. viridis. During the experiment, an epiphytic alga. Giffordia mitchellae, colonized and grew on the ASU. The alga increased the structure associated with the experimental units. Treatment leaf densities could thus be regarded only as relative measures of shelter. At low leaf densities, fish settlement increased with increasing algal cover; at higher leaf densities, the opposite occurred. The rapid increase in abundance of recently settled fish over a relatively narrow range of leaf densities above zero implies that small increases in shelter can have large effects on settlement of larval fish.