Halophila ovalis in the Tropical Atlantic Ocean

A species of seagrass in the genus Halophila was found growing in a shallow lagoon on the west shore of Antigua in the Caribbean West Indies. Genetic analysis showed the plants were Halophila ovalis. In addition, the samples had no genetic deviation (using nrDNA sequences) from Halophila johnsonii, considered to be an endemic and endangered species in Florida, USA. Morphological analysis demonstrated the Antiguan Halophila to be well within the range of plant characteristics previously described in the literature for H. ovalis, except for leaf width and number of seeds per fruit, and again, not different from H. johnsonii and very closely related to H. ovalis from the Indo-Pacific. Ours is the first report of H. ovalis in the Tropical Atlantic bioregion.     

Photophysiological responses of Halophila johnsonii to experimental hyposaline and hyper-CDOM conditions

The endemic seagrass Halophila johnsonii grows intertidally to 3 m deep, in both marine and riverine influenced habitats of eastern Florida. Salinity and chromophoric dissolved organic matter (CDOM) levels widely fluctuate across this broad habitat range, changing tidally and with variable influx of freshwater from watershed runoff, river discharge and stochastic storm events. CDOM exponentially absorbs light in the UV to blue wavelengths, affecting optical water quality. H. johnsonii produces 15 flavonoid compounds that maximally absorb in the UV range. These flavonoids are thought to function as UV-protectants (UVP) in high-light and UV-intense environments. This mesocosm study examined the photosynthetic capacity, quantum efficiency and pigment content of H. johnsonii under experimental treatments of three salinities (10, 20 and 30) with and without CDOM. Main treatment effects and possible interactive effects at both short- (1 day to 1 week) and longer-term (1 month) time scales were examined. There were no significant CDOM or CDOM x salinity effects over any of the experimental treatment durations. There was 100% mortality of plants at salinity 10 after 10 days regardless of water color. UVP content of leaves was not affected by CDOM in this study, but there was significant variation in UVP in response to salinity. Our results do not support the primary role of UVP in this species as a sunscreen, but indicate that different salinity environments contribute to changes in the levels of these flavonoids. The UVP response to salinity stress response was not mitigated by a decrease in UV-radiation (increased CDOM) as H. johnsonii continued to put energy into the production of the carbon-rich flavonoids regardless of potential UV-stress. The experimental results indicate that prolonged hypo-salinity conditions are an important environmental factor to manage in the limited geographic range of H. johnsonii.     

Megagametogenesis in Halophila johnsonii, a threatened seagrass with no known seeds,and the seed-producing Halophila decipiens (Hydrocharitaceae)

Megagametogenesis, the development of a megaspore into an embryo sac, has been identified in the seagrass Halophila johnsonii, a threatened species with no known sexual reproduction or seeds. Megagametogenesis in H. johnsonii was compared with megagametophyte development in Halophila decipiens, a related species known to readily produce viable seeds. In both species, ovules were structurally similar, megaspore mother cells were seen in premeiotic ovules, and linear tetrads and megagametophytes with two to eight nuclei were present in postmeiotic ovules. However, H. decipiens postmeiotic ovules had a chalazal pouch that was absent in the postmeiotic ovules of H. johnsonii. Late-stage H. decipiens ovules also contained embryos, indicating that they had been fertilized, whereas all late-stage H. johnsonii ovules were degrading and showed no signs of fertilization. These observations suggest that meiosis does occur in H. johnsonii megasporocytes, leading to the formation of viable megagametophytes and egg cells that could be fertilized if pollination occurred. Thus, the lack of seed set is due to a lack of pollination rather than any loss of capacity to produce seeds in this species.     

Genetic variation in populations of the threatened seagrass Halophila beccarii (Hydrocharitaceae)

Halophila beccarii (Hydrocharitaceae), a small monoecious seagrass, has been listed as a threatened species. In this study, a total of 106 samples were collected from four Chinese populations located at the northern limit of its distribution range. Using six polymorphic microsatellites, we found low genetic variation in this species, in which the mean number of alleles per locus was 2.8, and 16 multi-locus genotypes were revealed. In the four populations, the mean number of alleles per locus ranged from 1.2 to 2, one to eight genotypes were found, and clonal diversity ranged from 0 to 0.23; observed and expected heterozygosity ranged from 0.17 to 0.40 and from 0.09 to 0.24, respectively. Strong genetic differentiation was found among the populations, and the standardized fixation index (F′_ST) was 0.787. Species traits (i.e., clonal growth) and bottleneck effects due to drastic population fluctuation may contribute to the observed low genetic variation.     

Occurrence of Halophila baillonii meadows in Belize,Central America

Halophila baillonii Ascherson was found in Belize, Central America in 2003 and 2005. The observation extends the known range of this seagrass species to the western Caribbean. H. baillonii was previously recorded only in the eastern Caribbean and at one Pacific site in Panama. Both fruits and flowers of H. baillonii were observed at two locations in Belize in 2005. H. baillonii in Belize is an important food for manatee, forms a productive seagrass-based ecosystem, and is adversely affected by shoreline development and watershed run-off.     

Chemical analysis of flavonoid constituents of the seagrass Halophila stipulacea: First finding of malonylated derivatives in marine phanerogams

The flavonoid fraction from the butanol extract of a Mediterranean sample of the seagrass Halophila stipulacea was chemically analyzed. A new malonylated flavone glucoside, genkwanin-4′-O-(6“-malonyl-glucopyranoside) (3), was isolated together with known flavone glucosides 4-9, previously reported only from terrestrial sources. The structure of 3 was established by means of spectroscopic techniques, mainly NMR methods.     

Distributional range extension of the seagrass Halophila nipponica into coastal waters off the Korean peninsula

Eight temperate seagrass species (five in the genus Zostera, two in the genus Phyllospadix, and Ruppia maritima) have been previously reported in coastal waters off the Korean peninsula, which lies between 33°N and 43°N. Recently, a species of Halophila, a genus which occurs predominantly in tropical and subtropical areas, has been observed on the southern coast of Korea for the first time. The species was identified as Halophila nipponica. H. nipponica is distributed in warm temperate regions of Japan influenced by the warm Tsushima Current and was previously unknown outside the Japanese archipelago. Thus, we are able to report a range extension into Korea. The Korean Halophila meadow that we observed covered an area of about 2.1 ha, with average shoot density of about 1300 m⁻². We measured morphological features of vegetative and reproductive organs between June and September 2007. Morphological and reproductive features of the Halophila species in Korea were similar to those of the species in Japan. Increased water temperature in the coastal waters of Korea may at least partially account for the persistence of this new population.     

The photosynthetic performance of the tropical seagrass Halophila ovalis in the upper intertidal

Some ecophysiological adaptation strategies of the tropical seagrass Halophila ovalis were investigated with respect to this plant's ability to grow in the upper intertidal in either monospecific pools (but not together with other intertidal species) or emergent and exposed to high temperatures and irradiances during several hours every day. It was found that Halophila ovalis could raise the pH in simulated pools to 8.6, while the two other major (biomass wise) intertidal seagrasses raised the pH to 8.8 (Cymodocea rotundata) and 9.2 (Thalassia hemprichii). In situ, midday pH values of 8.5, 8.7 and 9.0 were recorded in pools inhabiting the three species, respectively. It was further found that photosynthetic electron transport rates (ETR) measured continuously in situ for one leaf during a diurnal cycle followed the irradiance up to a daily maximal value of 2200 μmol photons m⁻² s⁻¹, but dropped by ca. 50% when the leaf became air exposed just before noon. This drop in ETR upon emergence was verified by “point measurements”, while ETRs of leaves that grew in small pools formed as the tide receded followed the irradiance more closely. Similarly, F_v/F_m measured after 15 min of dark adaptation decreased by ca. 50% in emergent leaves during midday, but maintained higher daily values in the submerged leaves.     

Diurnal movements of chloroplasts in Halophila stipulacea and their effect on PAM fluorometric measurements of photosynthetic rates

Since diurnal chloroplast movements in Halophila stipulacea were described by Drew in 1979, this phenomenon has not been studied further for seagrasses. In addition to an apparent photoprotective role, such movements may affect the measurements of photosynthetic rates based on pulse amplitude modulated (PAM) fluorometry. This is because calculations of electron transport rates (ETR) are directly affected by the light absorption of the leaves (or the so-called absorption factor, AF), the latter of which changes with the movements of the chloroplasts. In this work, we therefore determined chloroplast clumping and dispersal, and measured AFs, chlorophyll contents and PAM fluorescence diurnally for H. stipulacea grown under two irradiance regimes. Diurnal chloroplast clumping occurred in high-light grown (HL) plants (∼450 μmol photons m⁻² s⁻¹ during midday), which was accompanied by a decrease in AF values (from 0.56 in the early morning to 0.34 at midday) but not in the chlorophyll content. Also, non-photochemical quenching (measured as NPQ) increased during the day in these plants. No such chloroplast movements and, thus, no diurnal changes in AF values (0.60 ± 0.04 throughout the day), and no changes in NPQ, were found in low-light grown (LL) plants (∼150 μmol photons m⁻² s⁻¹ during midday). As a consequence of the chloroplast clumping in HL plants, and its effect on AF values, maximal ETRs did not differ significantly between HL and LL plants. This finding thus shows the importance of taking into account changing AF values along the day when calculating ETRs of H. stipulacea, and other seagrasses potentially featuring diurnally changing AFs, under high-irradiance conditions.     

Wavelength-specific photosynthetic responses of Halophila johnsonii from marine-influenced versus river-influenced habitats

The seagrass Halophila johnsonii Eiseman grows from the upper intertidal to 3 m depths in habitats ranging from near-marine inlets to tidal riverine. These habitats have distinct optical characteristics, primarily due to variable concentrations of watershed-derived chromophoric dissolved organic matter (CDOM), which increases the attenuation of short-wavelength (blue and UV) light. H. johnsonii contains a suite of flavonoids that are thought to serve as UV-protective pigments (UVP). In this study, photosynthetic responses at specific wavelengths were compared between plants from a river-influenced high-CDOM habitat (Oleta River) and those from an adjacent marine, low-CDOM environment (Haulover Inlet) in Florida. Oxygen flux was used to measure dark respiration and photosynthesis under near-constant radiant energy at nine specific wavelengths from 340 to 730 nm. Riverine plants had higher gross photosynthetic rates and quantum efficiencies than inlet plants at the shortest wavelengths (350, 400 and 450 nm), while inlet plant photosynthetic rates and quantum efficiencies were higher at the two longest wavelengths measured (694 and 730 nm). Riverine plants also exhibited greater variation in photosynthetic responses across the spectrum and more variable pigment levels among replicates. Chlorophyll a and b concentrations were significantly greater in riverine plants suggesting that they were more shade-acclimated compared to the marine populations. Differences in wavelength-specific photosynthetic responses and chlorophyll levels indicate that the riverine plants were blue-shade acclimated. The higher and more variable UVP levels in the riverine population were not consistent with shade acclimation; however, these flavonoid pigments may protect chloroplasts from photodamage during short-term, high intensity irradiance conditions that occur over the course of tidal cycles at this highly fluctuating riverine site.     

Seed bank,biomass, and productivity of Halophila decipiens, a deep water seagrass on the west Florida continental shelf

One of the largest contiguous seagrass ecosystems in the world is located on the shallow continental shelf adjacent to the west coast of Florida, USA and is comprised of seasonally ephemeral Halophila decipiens meadows. Little is known about the demography of the west Florida shelf H. decipiens, which may produce 4.56 × 10⁸ g C day⁻¹ or more during the peak growing season. We documented seagrass distribution, biomass, and productivity, and density of sediment seed reserves, seedlings, flowers and fruits on the southeastern portion of the west Florida shelf by sampling along a transect at three stations in 10, 15, and 20 m water depth. Biomass, flower, fruit, seedling, and seed bank densities tended to be highest at stations in 10–15 m water depth and lowest at 20 m. Flowers and fruit were most prevalent during summer cruises (June and August 1999, July 2000). Seedling germination occurred during summer, fall (October 1999), and winter (January 2000) sampling events, with the highest seedling densities present during the winter. Seed bank density remained consistent through time. A Category I hurricane with sustained winds of 120 km h⁻¹ passed over the stations, but only limited impact on H. decipiens biomass was observed. The presence of a persistent seed bank provides for recovery after storm disturbance, annual reestablishment of populations, and continual maintenance of the 20,000 km² of deep water seagrass habitat present on the west Florida shelf.     

Nutrient dynamics in 3 morphological different tropical seagrasses and their sediments

Seasonality of nutrient dynamics in three morphologically different seagrass species and their sediments was examined for 1 year between November 2006 and November 2007 at four sites in the Andaman Sea, Thailand. The smaller species, Cymodocea serrulata and Halophila ovalis, showed major seasonal variation in shoot density, above- and belowground biomass, much more than expected from seasonal changes in water temperature and light conditions. All parameters showed minimum values in the dry season due to desiccation during neap tides. In contrast Enhalus acoroides showed less seasonal variation. Only limited seasonality was found in tissue N content of all species, whereas tissue P content responded to the low P concentration in the water column during the wet season. There were no differences in sediment conditions among species, and nutrient pools were generally low. Furthermore there were no significant spatial differences in seagrass and sediment nutrient dynamics, despite varying anthropogenic activity at the study sites, reflecting the oligotrophic conditions in this region.     

New record for Halophila decipiens Ostenfeld in Kenya based on morphological and molecular evidence

The seagrass Halophila decipiens Ostenfeld was recorded for the first time in Kenya in 2003. It was growing in subtidal mixed meadows with H. ovalis in protected bays at a depth of 3 m. DNA sequence analysis and morphological characteristics confirmed the identification. It is possible that this species is more widely distributed in the east African region. There was very little sequence differentiation between the Kenyan H. decipiens and the other samples in the Indo-West Pacific, and between this region and the Atlantic Ocean, further validating from an evolutionary perspective recent dispersal of this species.     

Localization and antioxidant capacity of flavonoids from intertidal and subtidal Halophila johnsonii and Halophila decipiens

In this study, flavonoid localization, content and total antioxidant capacity in leaves of subtidal Halophila decipiens were compared to intertidal and subtidal Halophila johnsonii. H. johnsonii leaves had significantly higher flavonoid content (3.5 and 3.8 nmol quercetin equivalent mm⁻² leaf for intertidal and subtidal H. johnsonii, respectively) and antioxidant capacity (101.7 and 224.2 nmol Trolox equivalent mm⁻² leaf for intertidal and subtidal H. johnsonii, respectively) than H. decipiens leaves (1.4 nmol quercetin equivalent mm⁻² leaf and 21.0 nmol Trolox equivalent mm⁻² leaf). Flavonoid content did not significantly differ between intertidal and subtidal H. johnsonii, however, antioxidant capacity was significantly higher in subtidal plants. Confocal laser scanning microscopy of fresh leaf cross sections indicated that both species contained flavonoids in the cuticle, but only H. johnsonii contained intracellular flavonoids. Intracellular flavonoids are better situated to perform antioxidant functions in planta. These results suggest that flavonoid compounds in H. johnsonii are capable of sunscreen and antioxidant functions while an antioxidant role for flavonoids within H. decipiens is not supported.     

Posidonia oceanica in the Marmara Sea

The seagrass Posidonia oceanica is a stenohaline species endemic to the Mediterranean Sea, where it normally lives at a salinity of between 36.5 and 39.5 ppt. Surveys carried out at the North-eastern distribution limits revealed large beds in the Dardanelles Strait and isolated beds in the Marmara Sea, where the salinity ranges between 21.5 and 28 ppt. Microsatellite analysis of these low-salinity tolerant P. oceanica beds, show different signs of genetic isolation: excess of heterozygosity and a presence of fixed alleles. These particularities are rarely found in the whole distributional range of the species. Moreover, all the populations considered in the analysis have a very low genetic diversity in comparison with most of the meadows sampled throughout the Mediterranean Sea. Taking into consideration the genetic data, rhizome expansion rate and the actual extent of the isolated beds in the Marmara Sea and knowing the reproductive rate and dissemination characteristics of P. oceanica, we hypothesize that the isolated population of the Marmara Sea has been established since the Middle Holocene, before the catastrophic intrusion of brackish water into the Marmara Sea and the strong and persistent flow coming from the Black Sea.     

Habitat selection of two gobies (Microgobius gulosus, Gobiosoma robustum): influence of structural complexity, competitive interactions, and presence of a predator

Herein I compare the relative importance of preference for structurally complex habitat against avoidance of competitors and predators in two benthic fishes common in the Gulf of Mexico. The code goby Gobiosoma robustum Ginsburg and clown goby Microgobius gulosus (Girard) are common, ecologically similar fishes found throughout the Gulf of Mexico and in the southeastern Atlantic Ocean. In Florida Bay, these fishes exhibit habitat partitioning: G. robustum is most abundant in seagrass-dominated areas while M. gulosus is most abundant in sparsely vegetated habitats. In a small-scale field survey, I documented the microhabitat use of these species where their distributions overlap. In a series of laboratory experiments, I presented each species with structured (artificial seagrass) versus nonstructured (bare sand) habitats and measured their frequency of choosing either habitat type. I then examined the use of structured versus nonstructured habitats when the two species were placed together in a mixed group. Finally, I placed a predator (Opsanus beta) in the experimental aquaria to determine how its presence influenced habitat selection. In the field, G. robustum was more abundant in seagrass and M. gulosus was more abundant in bare mud. In the laboratory, both species selected grass over sand in allopatry. However, in sympatry, M. gulosus occupied sand more often when paired with G. robustum than when alone. G. robustum appears to directly influence the habitat choice of M. gulosus: It seems that M. gulosus is pushed out of the structured habitat that is the preferred habitat of G. robustum. Thus, competition appears to modify the habitat selection of these species when they occur in sympatry. Additionally, the presence of the toadfish was a sufficient stimulus to provoke both M. gulosus and G. robustum to increase their selection for sand (compared to single-species treatments). Distribution patterns of M. gulosus and G. robustum likely result from a synthesis of various biotic and abiotic filters, including physiological tolerances to environmental factors, dispersal ability of larvae, and availability of food. Selection for structural complexity, competition, and presence of predators may further define the resulting pattern of distribution observed in the field.     

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.     

Distribution of exudate flavonoids in the genus Plectranthus

Thirty-four species of the genus Plectranthus (including species of the former genera Coleus and Solenostemon, fam. Lamiaceae) were surveyed for exudate flavonoids to see whether the distribution of these compounds would support a recent classification of the genus based on molecular and morphological characters. In this classification two major groups had been identified, the Coleus and Plectranthus clades. Only about 40% of the species, predominantly from the Plectranthus clade, were found to produce exudate flavonoids, which were mainly flavones. Flavanones were restricted to five species of the Plectranthus clade, whereas flavonols were only found in two species of the Coleus clade, Plectranthus montanus Benth. (synonyms Plectranthus marrubioides Hochst. ex Benth. and Plectranthus cylindraceus Hochst. ex Benth.) and Plectranthus pseudomarrubioides R.H.Willemse. Four of these flavonols were isolated from P. montanus and identified by NMR spectroscopy as the 3,7-dimethyl ether and 3,7,4′-trimethyl ether of quercetin and the 3,6,7-trimethyl ether and 3,6,7,4′-tetramethyl ether of quercetagetin. The remaining flavonols and flavones were identified by HPLC–UV and LC–MS of crude extracts on the basis of their UV and mass spectra, retention times and comparison with standards. Most flavonols were 3-methyl ethers and many of the flavones and flavonols were oxygenated at the 6-position. The most common flavones, occurring in both clades, were cirsimaritin and salvigenin, which are methoxylated at the 6- and 7-positions. 6-Hydroxylated flavones such as scutellarein and ladanein were restricted to species of the Plectranthus clade.