The deep-water species of Halimeda Lamouroux (Halimedaceae,Chlorophyta) from San Salvador Island,Bahamas: species composition,distribution and depth records

Caribbean species of Halimeda from the steep slopes of San Salvador Island, Bahamas, were collected along vertical transects between 25 and 255 m, on the northeast, northwest, west, southwest and south sides of the island, using the Harbor Branch Oceanographic Institution's submersible Johnson Sea Link I. The characteristics delineating species (segment and utricle size) were assessed for selected species, and the depth and western Atlantic distributions of the species reviewed. Halimeda copiosa was found to show a variation of surface utricle diameter with depth, with the deeper plants having 15% larger diameter utricles than the shallower plants. Of the seven species, one variety and one form of Halimeda studied from our submersible dive sites, H. copiosa, H. cryptica and H. gracilis grew to the greatest depths, from 120 to 150-m depths. H. lacrimosa var. globosa was found growing to 91 m, and H. tuna f. platydisca and H. discoidea to a depth of 73 m. Those of lesser depth, to 61 m, were H. goreauii, H. tuna f. tuna and H. lacrimosa var. lacrimosa. These represent substantial increases in depth distribution of the nine taxa found.     

Productivity and nutrient relationships in psammophytic versus epilithic forms of Bryopsidales (Chlorophyta): comparisons based on a short-term physiological assay

Members of the green algal order Bryopsidales (= Caulerpales) are important calcifying agents of tropical reefs and comprise two fundamentally different life-form groups: (1) epilithic species with limited attachment structures and (2) psammophytic forms that have extensive subterranean rhizoidal systems. Because the shallow-water habitats of the former have relatively low nitrogen (N) to phosphorus (P_i) ratios compared to the pore waters of the sedimentary carbonate-rich substrata in which the latter are anchored, we hypothesized that epilithic forms should tend to be relatively more limited by N, while psammophytic species should tend to show P_i limitation. In partial support of the hypothesis, light-saturated net photosynthesis (P_max) in the epilithic forms, Halimeda opuntia, H. lacrimosa and H. copiosa, tended to be enhanced by N, while P_i was inhibitory or had no effect. In contrast, the psammophytic forms, Udotea sp., U. conglutinata, H. monde, H. tuna and H. simulans, tended to be stimulated more by P_i, whereas N had little effect. The utility of a bioassay to assess macroalgal nutrient limitation, based on a physiological response (net P_max) to short-term nutrient pulses, is demonstrated.     

Chemical defense and chemical variation in some tropical Pacific species of Halimeda (Halimedaceae; Chlorophyta)

Over a dozen species of the genus Halimeda have been chemically investigated and found to produce the diterpenoid metabolites halimedatrial (1) and halimedatetraacetate (2) in varying concentrations. These meabolites have been proposed to play a role in chemical defense against herbivores based on their chemical structures and their demonstrated biological activities in laboratory and aquarium assays. We examined and compared the feeding deterrent effects of these two compounds tovard herbivorous fishes in field experiments on Guam reefs. Halimedatrial is a more effective feeding deterrent than halimedatetraacetate. It is the major secondary metabolite in young Halimeda macroloba and in the newly produced segments of growing plants. The organic extracts from young plants and new segments were significantly more deterrent than extracts from mature plant tissue. Some populations of Halimeda growing in reef-slope habitats, where herbivory is intense, also have high concentrations of halimedatrial. We compared extracts between reef slope and reef flat collections of Halimeda opuntia on Guam and Pohnpei (= Ponape), and H. discoidea and H. macroloba on Guam. We found that halimedtrial was the major metabolite in reef-slope collections of H. opuntia from Pohnpei and Pago Bay, Guam, and that halimedatetraacetate was the major metabolite a non-reef slope populations. In the cases examined, chemical defenses were greatest in (1) plant parts and (2) populations that were at greatest risk to herbivores.     

Evolution and phylogeography of Halimeda section Halimeda (Bryopsidales, Chlorophyta)

Nuclear ribosomal and plastid DNA sequences of specimens belonging to section Halimeda of the pantropical green seaweed genus Halimeda show that the group under scrutiny contains many more genetically delineable species than those recognized by classical taxonomy. Discordances between phylograms inferred from nuclear and plastid DNA sequences suggest that reticulate evolution has been involved in speciation within the clade. Nonetheless, our data do not allow ruling out certain alternative explanations for the discordances. Several pseudo-cryptic species are restricted to the margins of the generic distribution range. In a clade of H. cuneata sibling species from widely separated subtropical localities in the Indian Ocean, the South African sibling branches off first, leaving the Arabian and West Australian species as closest relatives. We hypothesize that geographic isolation of the siblings may have taken place following Pleistocene or Pliocene periods of climatic cooling during which subtropical species occupied larger distribution ranges. A more basal separation of Atlantic, Indo-Pacific, and Mediterranean species indicates vicariance. The alternative events that could have caused this vicariance are discussed.     

Diterpenoid metabolites from pacific marine algae of the order Caulerpales (Chlorophyta)

Chemical investigations of two new species of Udotea, reinvestigation of Tydemania expeditionis and Chlorodesmis fastigiata from Guam and Saipan, Marianas Islands, as well as studies of Caulerpa brownii from Southern Australia, have led to the isolation of four new related diterpenoids and several previously described metabolites. These metabolites possess antimicrobial activities and inhibit cell division in the fertilized sea urchin egg bioassay.     

A new species of Late Pleistocene rail (Aves: Rallidae) from Abaco, the Bahamas

We describe a new species of rail from the Sawmill Sink blue hole on Abaco Island in the northern Bahamas. Known from abundant, beautifully preserved Late Pleistocene fossils, Rallus cyanocavi sp. nov. was a medium-sized, flightless species that probably was endemic to the Little Bahama Bank, which is a carbonate platform surrounded by deeper water. We are uncertain whether R. cyanocavi survived into the Holocene, when higher sea levels transformed the Little Bahama Bank from a single large, Late Pleistocene island (ca. 12000 km²) to the scattering of smaller islands seen today, the largest of which is Abaco (1681 km²). Fossils of additional extinct, flightless species of Rallus probably await discovery on some of the 21 other carbonate banks that span the Bahamian Archipelago.     

Molecular systematics,historical ecology,and phylogeography of Halimeda (Bryopsidales)

Halimeda (Bryopsidales), a genus of calcified, segmented green seaweeds, abounds in reefs and lagoons throughout the tropics. To investigate phylogenetic, phylogeographic, and historic ecological relationships of the genus, the nuclear rDNA including the SSU and both ITS regions were sequenced. A maximum likelihood tree revealed the following: (1) there were anatomical and morphological synapomorphies for five well-supported lineages; (2) the last common ancestor of one lineage invaded sandy substrata; those of two other lineages established in wave-affected habitats, whereas the cenancestor of the remaining two lineages occupied sheltered rocky slopes. Yet, several species adapted to new habitats subsequently, resulting in several cases of convergence; (3) all lineages separated into Atlantic and Indo-Pacific daughters, likely resulting from the rise of the Panamanian Isthmus. Each daughter pair gave rise to additional convergent species in similar habitats in different oceans; (4) Halimeda opuntia, the only monophyletic pantropical species detected so far, dispersed from the Indo-Pacific into the Atlantic well after the closure event; (5) minor SSU-sequence differences across species and phylogeographic patterns of vicariance indicated a relatively recent diversification of the extant diversity. Cretaceous and Early Tertiary fossil look-alikes of modern species must then have resulted from iterative convergence.     

Morphology and sedimentology of Halimeda bioherms from the eastern Java Sea (Indonesia)

Halimeda bioherms, occurring primarily along the western and southern margins of Kalukalukuang Bank in the eastern Java Sea, display a wide variety of thicknesses and shapes. In general, high-frequency forms of the northern bank are replaced by thicker and lower frequency forms along the deeper southern margin. Sidescan sonar data suggest aperiodic reworking of shallow bioherm crests of the northern bank into features suggestive of bedforms. These features are not associated with deeper bioherms of the southern bank. Cores from the bioherms indicate that they consist mostly of disarticulated Halimeda plates set in a lime mud matrix composed largely of Halimeda fragments and foraminifera tests. Carbon-14 dating shows that, with exception of some deep southern bank examples, bioherms are actively accreting. Results of mineralogy and elemental chemistry on piston core PC-12 suggest cyclic variations in Mg-calcite (cement in Halimeda utricles), which may be related to periodic excursions of cold Pacific throughflow water onto the bank. Composition of the Mg-calcite (8.6 mole-%) suggests a temperature of formation of about 22 °C, which is 7 °C below average surface water temperatures. The carbon and oxygen isotope compositions of both aragonite and Mg-calcite phases are remarkably homogeneous, but were inconclusive with regard to the cold-water intrusion hypothesis. However, a lack of reef-building corals below a depth of 15 m, abundance of Halimeda bioherms on the western margin of K-Bank, where upwelling is predicted, extensive boring of sedimentary particles by endolithic boring algae, and high nutrient values of water at the thermocline (50–70 m deep) all support the incursion of cold, nutrient-rich water onto the bank. Upwelling and nutrient overloading are suggested as explanations for remarkable algal growth at the expense of reef-building corals.     

Ultrastructure of the spermatozoid in Halimeda tuna (chlorophyceae)

The mature Halimeda tuna spermatozoid was studied under the electron microscope. It is pear-shaped and biflagellate. The nucleus is an uncondensed structure except for a few opaque (chromatin) aggregations and shows a rounded profile. The endoplasmic reticulum is a rather well-developed system of rough cisternae engaged in synthesis and storage of proteins. Free ribosomes are numerous. A large mitochondrial apparatus shaped like a horseshoe lies in the anterior gamete region. Only one single plastid is found, and it exhibits a deeply indented outline, a partially structured matrix, osmiophilic globules, and three to four starch grains. The axoneme pattern is 9 + 2. B tubules show septate lumina. A peculiar structure provided with a three-layered shell covering materials of varying electrondensity lies on the upper surface of both basal bodies. The flagellar root system exhibits a cruciate pattern and sets having an inconstant number of microtubules – ie, three, four, or five.     

A comparison of line transect versus linear percentage sampling for evaluating stony coral (Scleractinia andMilleporina) community similarity and area coverage on reefs of the central Bahamas

Three methods of evaluating stony coral communities were used on selected reefs in the Exuma Cays Land and Sea Park (24°22N, 77°30W) in the central Bahamas. Shallow reefs (< 4 meters depth) were selected from aerial surveys based on size, location, and physical setting, and grouped into three community types: (1) channel patch reefs, (2) soft-coral-sponge patch reefs and (3) fringing reefs. Three survey techniques used to evaluate the stony coral communities were a) species presence and absence lists, b) linear percentage and c) line transects using 1 mx1 m grids. Data collected from these survey methods was used to calculate coral colony density, species area coverage, and species diversity based on colony number and based on linear (cm) coral cover. The linear percentage sampling was considered too convervative in determining distribution patterns of a reef community; this technique takes into account the massive reef framework species such asM. annularis. The line transect technique can account for both colony number and area coverage, thus is a better method for characterizing reef communities. Sample size considerations are discussed for future applications of survey techniques for ground-truthing digital images of small, shallow reef communities.     

Morphology and ultrastructure of epilithic versus cryptic,microbial growth in lower Cambrian phosphorites from the Montagne Noire,France

The lower Cambrian grainy phosphorites of the northern Montagne Noire occur interbedded with grey to black, laminated to massive shales and limestones deposited along the edge of a continental shelf, associated with slope‐related facies and unstable substrates. The concentration of phosphate took place by repeated alternations of low sedimentation rates and condensation (hardgrounds), in situ early‐diagenetic precipitation of fluorapatite, winnowing and polyphase reworking of previously phosphatized skeletons and hardground‐derived clasts. The succession of repeated cycles of sedimentation, phosphate concentration, and reworking led to multi‐event phosphate deposits rich in allochthonous particles. Phosphogenesis was primarily mediated by microbial activity, which is evidenced by the abundance of phosphatized putative microbial remains. These occur as smooth and segmented filaments, sheaths, and ovoid‐shaped coccoids. These simple morphologies commonly form composite frameworks as a result of their aggregation and entanglement, leading to the record of biofilms, microbial mats, and complex networks. These infested the calcitic skeletonized microfossils that littered the substrate. Microbial activity evidences epilithic (anisotropic coatings on skeletons), euendolithic (perforating skeletal walls), and cryptoendolithic (lining inter‐ and intraparticulate pores) strategies, the latter dominated by bundles of filaments and globular clusters that grew along the cavities of helcionellids and hyoliths. According to their epilithic versus cryptic strategies, microbial populations that penetrated and dwelled inside hard skeletal substrates show different network and colonial morphologies. These early Cambrian shell concentrations were the loci of a stepwise colonization made by saprophytic to mutualistic, cyanobacterial–fungal consortia. Their euendolithic and cryptoendolithic ecological niches provided microbial refugia to manage the grazing impact mainly led by metazoans.     

Loss of psbS expression reduces vegetative growth, reproductive output, and light-limited, but not light-saturated, photosynthesis in Arabidopsis thaliana (Brassicaceae) grown in temperate light environments

Plants protect themselves against the deleterious effects of high light intensities by inducing a mechanism ubiquitous among plants known as energy dissipation, which safely converts excess light to heat before it can lead to the formation of free radicals. Mutants possessing a deletion of the psbS gene, such as the npq4 mutant, cannot perform energy dissipation and thus offer an opportunity to assess the importance of this process to plant function. In a temperate light environment, greenhouse-grown npq4 mutants of Arabidopsis thaliana had smaller rosette diameters and leaf numbers. The reduction in size observed in npq4 plants was associated with fewer floral stalks, fewer fruits, lower whole-plant and individual seed masses, and lower germination rates. In the field, npq4 mutants developed fewer fruits. After a controlled exposure to high light stress, both PSII efficiency and CO(2) assimilation were more significantly compromised in npq4 mutants at low light intensities, but not at high light intensities. Thus, the protective nature of energy dissipation manifests in light environments that include periods of high light, which predispose plants to PSII photoinactivation, and periods of low light, when PSII photoinactivation decreases the rate of photosynthesis.     

Effects of temperature, salinity, irradiance and diurnal periodicity on growth and photosynthesis in the diatom Nitzschia americana: light-limited growth

The effects of variations in temperature (10, 15, 20, 25, 30C)and salinity (8, 15, 20, 26, 32 p.p.t.) on cell size and ratesof photosynthesis and population growth were evaluated in axenic,light-limited (30 µE m^–2 s^–1) cultures ofan estuarine clone of the diatom ^{Nitzschia americana}. Experimentalconditions were chosen to reflect the range of natural conditionswhich occur in the clone's native environment, the Cape FearRiver Estuary, Nqrth Carolina. Rates of light-limited grossphotosynthesis; or photosynthetic efficiency (PSE), were determinedfrom short-term (1 h) ¹⁴C incubations. Diurnal variation inPSE was analyzed using ¹⁴C samples taken during times of estimatedmaximum and minimum rates of diurnal photosynthesis. The salinity-dependenttemperature response of PSE is characterized by a gradual increasein rates up to a temperature optimum at –25C, beyondwhich rates rapidly decline to zero at an upper lethal limit(30–40C). A similar pattern was observed in populationgrowth rates as a function of salinity and temperature. Independentof temperature, optimum salinity for growth was 26 p.p.t. Amaximum growth rate of 2.4 div d^–1 was measured at 25Cand 26 p.p.t. The effect of non-optimum salinity is a reductionin growth rates relative to a predicted temperature-dependentmaximum. Salinity-dependent patterns of variation in cell volume,in general, mirrored the response of population growth suchthat cultures with relatively high growth rates were dominatedby small cells. Significant diurnal variation was observed inPSE; maximum diurnal rates were generally 1.5–3.5 timesgreater than minimum diurnal rates.     

A new model for the calcification of the green macro-alga Halimeda opuntia (Lamouroux)

Halimeda opuntia is a cosmopolitan marine calcifying green alga in shallow tropical marine environments. Besides Halimeda’s contribution to a diverse habitat, the alga is an important sediment producer. Fallen calcareous segments of Halimeda spp. are a major component of carbonate sediments in many tropical settings and play an important role in reef framework development and carbonate platform buildup. Consequently the calcification of H. opuntia accounts for large portions of the carbonate budget in tropical shallow marine ecosystems. Earlier studies investigating the calcification processes of Halimeda spp. have tended to focus on the microstructure or the physiology of the alga, thus overlooking the interaction of physiological and abiotic processes behind the formation of the skeleton. By analyzing microstructural skeletal features of Halimeda segments with the aid of scanning electron microscopy and relating their occurrence to known physiological processes, we have been able to identify the initiation of calcification within an organic matrix and demonstrate that biologically induced cementation is an important process in calcification. For the first time, we propose a model for the calcification of Halimeda spp. that considers both the alga’s physiology and the carbon chemistry of the seawater with respect to the development of different skeletal features. The presence of an organic matrix and earlier detected external carbonic anhydrase activity suggest that Halimeda spp. exhibit biotic precipitation of calcium carbonate, as many other species of marine organisms do. On the other hand, it is the formation of micro-anhedral carbonate through the alga’s metabolism that leads to a cementation of living segments. Precisely, this process allows H. opuntia to contribute substantial amounts of carbonate sediments to tropical shallow seas.     

Complex evolutionary transitions and the significance of c(3)-c(4) intermediate forms of photosynthesis in Molluginaceae

C(4) photosynthesis is a series of biochemical and structural modifications to C(3) photosynthesis that has evolved numerous times in flowering plants, despite requiring modification of up to hundreds of genes. To study the origin of C(4) photosynthesis, we reconstructed and dated the phylogeny of Molluginaceae, and identified C(4) taxa in the family. Two C(4) species, and three clades with traits intermediate between C(3) and C(4) plants were observed in Molluginaceae. C(3)-C(4) intermediacy evolved at least twice, and in at least one lineage was maintained for several million years. Analyses of the genes for phosphoenolpyruvate carboxylase, a key C(4) enzyme, indicate two independent origins of fully developed C(4) photosynthesis in the past 10 million years, both within what was previously classified as a single species, Mollugo cerviana. The propensity of Molluginaceae to evolve C(3)-C(4) and C(4) photosynthesis is likely due to several traits that acted as developmental enablers. Enlarged bundle sheath cells predisposed some lineages for the evolution of C(3)-C(4) intermediacy and the C(4) biochemistry emerged via co-option of photorespiratory recycling in C(3)-C(4) intermediates. These evolutionarily stable transitional stages likely increased the evolvability of C(4) photosynthesis under selection environments brought on by climate and atmospheric change in recent geological time.     

A preliminary study of keratinophilic fungi from Abaco Island,the Bahamas

The soil of two Bahamian communities on Abaco Island was examined for keratinophilic fungi. Isolated fungal species in soil identifies possible causal agents of dermatophytoses. Inhabitants of the area may have contaminated the soil, or species present may represent a potential source of new clinical cases. Fungal pathogens have been reported as free living soil saprophytes.     

A preliminary study of marine fungi from Abaco Island,the Bahamas

Marine soil, mangrove wood, and free floating algal collections were made in the shallow littoral zone of Abaco Island, The Bahamas. Material was subsequently examined in the laboratory for fungal populations inhabiting seawater shorelines of the island. The study follows previous mycological investigations of Abaco including the keratinophilic fungi (10) and the fresh water fungi (11).