Host location and selection by the symbiotic sargassum crab <Emphasis Type="Italic">Portunus sayi</Emphasis>: the role of chemical,visual and tactile cues

The Sargassum community consists of a unique and diverse assemblage of symbiotic fauna critical to pelagic food chains. Associated symbionts presumably have adaptations to assist in finding Sargassum. In situ scattered Sargassum patches accumulate as they are pushed toward the shoreline (via wind, waves, currents or tides) and are frequently less than 1 m apart and in depths of 10 cm or less as the patches approach the shoreline Crabs, and other symbiotic fauna, must relocate to another patch that is seaward in direction or likely perish as their current patch will likely become beached. This study investigated sensory cues used for host location and selection by the Sargassum crab, Portunus sayi. Chemical detection trials were conducted with a two-chamber choice apparatus with Sargassum spp. and Thalassia testudinum as habitat source odors. Visual detection trials (devoid of chemical cues) and habitat selection trials were conducted in which crabs were given a choice between hosts. Results showed that P. sayi responded to chemicals from Sargassum spp. Crabs visually located host habitats but did not visually distinguish between different hosts. In host selection trials, crabs selected Sargassum spp. over artificial Sargassum and T. testudinum. These results suggest that crabs isolated from Sargassum likely use chemoreception; within visual proximity of a potential patch, crabs likely use both chemical and visual information.     

Energy transformations by the sargassum fish,Histrio histrio (L.)

Ingestion, egestion, growth, and respiration measurements were made in the laboratory on three size-classes of the Sargassum fish, Histrio histrio(L.). Comparison of three size-classes showed changes from high growth and low respiration and assimilation in small Histrio to reduced growth and increased respiration and assimilation in larger fish. A high assimilation and growth efficiency suggest that Histrio is adapted to a food-limited environment (Sargasso Sea). Energy requirements of the smallest size-class (10–29 mm) were compared to the energy provided by their dominant food source, the Sargassum shrimp Latreutes fucorum (Fabricius) and Leander tenuicornis (Say). The annual energy requirement per individual was 1.4 times greater than the food energy available. This discrepancy is attributed to the difficulty in extrapolating laboratory studies to the field and the difficulty in obtaining adequate population size measurements of the Sargassum community fauna.     

Effects of marine algal proximate composition on methane yields

The suitability of differentGracilaria spp. and twoSargassum species for bioconversion to methane was determined through bioassays of methane yield.Gracilaria species and strains were excellent feedstocks for high methane yields, ranging from 0.28 to 0.40 m³ kg^–1 volatile solids added. These yields ranged from 58 to 95% of theoretical stoichiometric yields. Methane yields were highly correlated with acid soluble carbohydrate components of theGracilaria spp. BothSargassum fluitans andS. pteropleuron were poor feedstocks, with methane yields ranging from 0.12 to 0.19 m³ kg^–1 volatile solids added, 27 to 46% of theoretical stoichiometric yields, respectively. The various tissue types of theseSargassum species were also poor feedstocks for anaerobic digestion to methane. While there is no clear explanation for the low methane yields, the twoSargassum spp. appear to contain a high proportion of an insoluble, non-extractable component which may not be available as a substrate for bioconversion to methane.     

Testing systematic concepts of Sargassum (Fucales, Phaeophyceae) using portions of the rbcLS operon

The taxonomic and phylogenetic concepts within the Sargassum C. Agardh (Sargassaceae) species complex were evaluated through molecular phylogenetic analyses using portions of the chloroplast encoded rbcLS Operon. According to more conservative sequences (rbcL), Turbinaria (Turner) J. Agardh is a close and well‐supported sister lineage to the Sargassum species complex and an appropriate external outgroup for analyses of subgenera and subsections within Sargassum. Both rbcL and more rapidly evolving rbcLS spacer sequences indicated that the East Asiatic genus Myagropsis (Mertens et Turner) Fensholt, along with Sargassum sinicola Setchell et Gardner, represent the closest lineage to Sargassum and form appropriate internal outgroups. The rbcLS spacer region supported three of four subgeneric designations by J. Agardh and sectional levels within the subgenus Sargassum. However, some aspects of Agardh's system were not supported: many of the subsectional ranks or the phyletic concepts; Phyllotrichia was not monophyletic as a subgenus, and its species were also not the most ancestral of Sargassum; and subgenus Sargassum was not the most derived subgenus within the genus. This modern phylogeny suggests a deep evolutionary history for subgenus Sargassum with rapid speciation in closely related subsections and series, and a sister relationship between subgenera Arthrophycus and Bactrophycus.     

Reproductive biomass allocation in three Sargassum species

Allocation of biomass to sexual reproductive (receptacle) tissue and vegetative (holdfast) tissue differed absolutely and relatively in three Sargassum species that form the bulk of the intertidal algal canopy in the northern Gulf of California. Sargassum herporhizum devoted a greater proportion of its thallus mass into its rhizoidal holdfast than did S. sinicola var. camouii or S. johnstonii, whose holdfasts are solid, more compact, and composed of a lower percentage of water. Conversely, more sexual receptacle tissue was produced by these two species with small holdfasts during the spring reproductive period. Sargassum sinicola var. camouii, which is the only species of the three that becomes fertile in the fall, produces a comparable amount of sexual tissue during this second period of reproduction. Removal of Sargassum from single-species patches showed that canopy regrowth by S. herporhizum with its encroaching rhizoidal holdfast was more rapid and complete than that of the other two species, which invest most of their reproductive efforts into sexual propagules that can disperse long distances. Sargassum herporhizum also displayed a more rapid and complete recovery of canopy cover in patches cleared of thalli and in control patches following the annual summer dieback. These two divergent modes of reproductive biomass allocation suggest that ability to encroach upon nearby open sites and ability to colonize distant discrete islands of suitable habitat represent two distinct reproductive strategies requiring different patterns of biomass allocation. Moreover, for energetic reasons, a species may not be able to excel at both modes of reproduction.     

Ontogenetic changes in habitat selection during settlement in a coral reef fish: ecological determinants and sensory mechanisms

The behavior of marine larvae during and after settlement can help shape the distribution and abundance of benthic juveniles and therefore the intensity of ecological interactions on reefs. Several laboratory choice-chamber experiments were conducted to explore sensory capabilities and behavioral responses to ecological stimuli to better understand habitat selection by “pre-metamorphic” (larval) and “post-metamorphic” (juvenile) stages of a coral reef fish (Thalassoma hardwicke). T. hardwicke larvae were attracted to benthic macroalgae (Turbinaria ornata and Sargassum mangarevasae), while slightly older post-metamorphosed juveniles chose to occupy live coral colonies (Pocillopora damicornis). Habitat choices of larvae were primarily based upon visual cues and were not influenced by the presence of older conspecifics. In contrast, juveniles selected live coral colonies and preferred those occupied by older conspecifics; choices made by juveniles were based upon both visual and olfactory cues from conspecifics. Overall, the laboratory experiments suggest that early life-history stages of T. hardwicke use a range of sensory modalities that vary through ontogeny, to effectively detect and possibly discriminate among different microhabitats for settlement and later occupation. Habitat selection, based upon cues provided by environmental features and/or by conspecifics, might have important consequences for subsequent competitive interactions.     

Rising CO2 concentrations affect settlement behaviour of larval damselfishes

Reef fish larvae actively select preferred benthic habitat, relying on olfactory, visual and acoustic cues to discriminate between microhabitats at settlement. Recent studies show exposure to elevated carbon dioxide (CO₂) impairs olfactory cue recognition in larval reef fishes. However, whether this alters the behaviour of settling fish or disrupts habitat selection is unknown. Here, the effect of elevated CO₂ on larval behaviour and habitat selection at settlement was tested in three species of damselfishes (family Pomacentridae) that differ in their pattern of habitat use: Pomacentrus amboinensis (a habitat generalist), Pomacentrus chrysurus (a rubble specialist) and Pomacentrus moluccensis (a live coral specialist). Settlement-stage larvae were exposed to current-day CO₂ levels or CO₂ concentrations that could occur by 2100 (700 and 850 ppm) based on IPCC emission scenarios. First, pair-wise choice tests were performed using a two-channel flume chamber to test olfactory discrimination between hard coral, soft coral and coral rubble habitats. The habitat selected by settling fish was then compared among treatments using a multi-choice settlement experiment conducted overnight. Finally, settlement timing between treatments was compared across two lunar cycles for one of the species, P. chrysurus. Exposure to elevated CO₂ disrupted the ability of larvae to discriminate between habitat odours in olfactory trials. However, this had no effect on the habitats selected at settlement when all sensory cues were available. The timing of settlement was dramatically altered by CO₂ exposure, with control fish exhibiting peak settlement around the new moon, whereas fish exposed to 850 ppm CO₂ displaying highest settlement rates around the full moon. These results suggest larvae can rely on other sensory information, such as visual cues, to compensate for impaired olfactory ability when selecting settlement habitat at small spatial scales. However, rising CO₂ could cause larvae to settle at unfavourable times, with potential consequences for larval survival and population replenishment.     

Use of RAPD in differentiation of selected species of <Emphasis Type="Italic">Sargassum</Emphasis> (Sargassaceae,Phaeophyta)

Sargassum C. Agardh is one of the most common but little understood genera of Phaeophyta in Malaysia. The difficulty in species delineation is due to morphological plasticity. A combination of morphology and random amplified polymorphic DNA (RAPD) studies of selected Sargassum species was carried out to have a better understanding of the taxonomy. Primer OPA13 was found to be good for discriminating between Sargassum species. Sargassum binderi was shown to be different from S. oligocystum (S_D>0.5 = 14.11%), indicating the importance of the vesicle and receptacle in species differentiation. S. baccularia was clearly separated out from S. polycystum and S. stolonifolium using primer OPA13. RAPD analysis showed that the presence of the stolon is an important character for separating S. baccularia (no stolon) from S. polycystum (stolon) and S. stolonifolium (stolon). Presented at the 6th Meeting of the Asian Pacific Society of Applied Phycology, Manila, Philippines.     

Phylogenetic relationships of Sargassum (Sargassaceae, Phaeophyceae) with reference to a taxonomic revision of the section Phyllocystae based on ITS-2 nrDNA sequences

Sequences of the end of the 5.8S gene and the internal transcribed spacer 2 (ITS‐2) of nuclear ribosomal DNA have been determined for 19 species of the brown algal genus Sargassum (Sargassaceae), representing three subgenera and eight sections (sections are in parentheses): Phyllotrichia, Bactrophycus (Teretia, Spongocarpus, Halochloa and Repentia) and Sargassum (Acanthocarpicae, Malacocarpicae, Zygocarpicae) to assess the taxonomic position of the section Phyllocystae traditionally included within the Bactrophycus. The sequence of Myagropsis myagroides (Mertens ex Turner) Fensholt (Sargassaceae) was used as an outgroup. Sequences of ITS‐2 were analyzed using neighbor‐joining, parsimony and maximum likelihood methods. The results showed the existence of three clades in Sargassum, corresponding to the three subgenera. The subgenus Phyllotrichia is positioned near the outgroup. Two robust clades were obtained, one corresponding to the subgenus Bactrophycus and the other to the subgenus Sargassum. Sargassum mcclurei Setchell and Sargassum quinhonense Nguyen, the two Phyllocystae investigated, are close to species belonging to the section Zygocarpicae in the subgenus Sargassum. A transfer of the section Phyllocystae to the subgenus Sargassum is therefore proposed on the basis of molecular data (ITS‐2) and morphological data (receptacles and basal leaf).     

A MORPHOLOGICAL AND MOLECULAR STUDY OF AUSTRAL SARGASSUM (FUCALES,PHAEOPHYCEAE) SUPPORTS THE RECOGNITION OF PHYLLOTRICHA AT GENUS LEVEL,WITH FURTHER ADDITIONS TO THE GENUS SARGASSOPSIS1

Sargassum subgenus Phyllotricha currently includes seven species restricted to Australian and New Zealand coasts. A recent study of Cystoseira and other Sargassaceae genera based on mitochondrial 23S DNA and chloroplast‐encoded psbA sequences resulted in the most widely distributed species of subgenus Phyllotricha, Sargassum decurrens, being transferred to the reinstated monospecific Sargassopsis Trevisan. The fate of the residual six Phyllotricha species, however, was not considered. The present study examines these Phyllotricha species, alongside other Sargassum subgenera, Sargassopsis, Sirophysalis trinodis (formerly Cystoseira trinodis) and the New Zealand endemic Carpophyllum Greville, using morphological evidence and the molecular phylogenetic markers cox3, ITS‐2 and the rbcL–S spacer. Our results suggest both the genus Sargassum and Sargassum subgenus Phyllotricha are polyphyletic as currently circumscribed. Four S. subgen. Phyllotricha species, i.e. S. sonderi, S. decipiens, S. varians and S. verruculosum, form a monophyletic group sister to the genus Carpophyllum, and S. peronii is genetically identical to S. decurrens with regard to all three loci. We propose the resurrection of the genus Phyllotricha Areschoug, with type species Phyllotricha sonderi, and include the new combinations Phyllotricha decipiens, Phyllotricha varians and Phyllotricha verruculosum. Sargassum peronii, S. heteromorphum and S. kendrickii are transferred to Sargassopsis and Sargassum peronii is considered a synonym of Sargassopsis decurrens.     

Multiple cues from multiple habitats: Effect on metamorphosis of the Florida stone crab, Menippe mercenaria

The Florida stone crab, Menippe mercenaria, is an economically and ecologically important species that ranges from North Carolina throughout the Caribbean and the southeastern Gulf of Mexico. However, there is little known about its early life history stages as compared to other commercially important species in the region. The goal of this research was to examine effects of putative cues on metamorphosis from the megalopa stage to the first juvenile stage. Our study investigated the effect of water-soluble exudates from four substrata, as well as natural biofilms, and exudates from adult stone crabs. In addition, the influence of natural substrata was compared to that of artificial substrata. Adult exudate had no significant effect on metamorphosis, despite a wide range of tested concentrations. In contrast, there was a significant effect on mean time to metamorphosis in experimental groups exposed to multiple cues associated with the brown alga Sargassum fluitans, rubble from stone crab habitat, the eastern oyster Crassostrea virginica, and biofilms associated with the oyster. Furthermore, we provide evidence for metamorphic responses to water-soluble chemical cues, as well as biochemical and physical cues associated with different substrata. Overall results were coherent with the relevant body of previous work on metamorphosis of brachyuran crab larvae and indicate that both physical and chemical cues are important factors in facilitating the settlement and metamorphosis of M. mercenaria larvae in juvenile nursery habitat.     

Growth and size distribution of some large brown algae in Ohori,east coast of Korea

About fifty individuals of four dominant large brown algae were collected by random sampling at monthly intervals using SCUBA. Size frequencies of Sargassum confusum and S. horneri were concentrated mainly in the small size classes during the initial growth phase, but were more or less evenly distributed in most size classes during the fast growing phase. Undaria pinnatifida and Costaria costata showed the same tendency as Sargassum spp., that frequency distribution became even when the plants reached their maximum growth. The kelp species recorded their maximum lengths in March, whereas Sargassum spp. showed their peaks in early summer. The relationships between the length and weight in different growth phases appeared to be dependent on the phenology of each species.     

Recognition and use of ascidian hosts, and mate acquisition by the symbiotic pea crab Tunicotheres moseri (Rathbun, 1918): the role of chemical, visual and tactile cues

Many crustaceans live in intimate association with larger host animals, mostly for shelter and food. Host recognition and location behaviors by these crustaceans, and other symbionts, are vital in the life history of most symbioses. The pinnotherid crab Tunicotheres moseri (Rathbun, 1918) is a common symbiont with various solitary ascidians throughout its reported range of Jamaica, Venezuela, and West Florida. Sensory cues affecting host recognition, host use, and mate acquisition in the West Florida crab population were the focus of this study. T. moseri responded to host generated and conspecific (males responded to non-gravid females) waterborne chemical cues. Although host generated waterborne chemical cues were recognized by T. moseri, access to direct contact (which could involve chemical, visual and tactile cues) with host ascidians triggered much stronger responses. Results of conditioning trials suggest that T. moseri is a generalist, but prefers Styela plicata even after conditioning with the novel host species Molgula occidentalis and Phallusia nigra. These results show that T. moseri employs the use of multiple senses in establishing symbioses with host tunicates, and the plasticity of the responses allow potentially novel tunicates to serve as hosts. Finally, this study shows that male T. moseri, which typically only enter tunicates for breeding purposes, use waterborne cues to find tunicates housing potential female mates.     

Predation risk assessment by larval reef fishes during settlement-site selection

Predation rates of marine species are often highest during the transition from the pelagic to the benthic life stage. Consequently, the ability to assess predation risk when selecting a settlement site can be critical to survival. In this study, pairwise choice trials were used to determine whether larvae of three species of anemonefish (Amphiprion melanopus, A. percula and Premnas biaculeatus) are able to (1) assess the predation risk of potential anemone settlement sites through olfactory cues alone and (2) alter their settlement choices depending on the options available (host or non-host anemone). When predation risk was assessed with host and non-host anemone species independently, all species of anemonefish significantly chose the odor associated with the low-risk settlement option over the high-risk site. Most importantly, all species of anemonefish selected water with olfactory cues from their host anemone regardless of predation risk when paired against non-host anemone odor. These results demonstrate that larval reef fishes can use olfactory cues for complex risk assessment during settlement-site selection; however, locating the correct habitat is the most important factor when selecting a settlement site.     

Does restoration of a habitat‐forming seaweed restore associated faunal diversity?

Declines of habitat‐forming organisms in terrestrial and marine systems can lead to changes in community‐wide biodiversity. The dominant habitat‐forming macroalga Phyllospora comosa (Fucales) went locally extinct along the metropolitan coastline of Sydney in the 1980s. However, the consequences of that disappearance to the associated faunal diversity in these habitats, and whether Phyllospora is ecologically redundant with respect to the biodiversity it supports, are not known. Efforts are underway to restore Phyllospora, and the capacity to enhance local biodiversity is an important component of the rationale for restoration. We compared epifaunal diversity (abundances and composition) between Phyllospora and two other co‐occurring habitat‐forming algae, the kelp Ecklonia radiata and the fucoid Sargassum vestitum, and determined whether Phyllospora transplanted to Sydney developed different epifaunal communities than undisturbed thalli and controls. Where the 3 species naturally co‐occurred, Phyllospora supported different abundances of taxa than Ecklonia and Sargassum, as well as different composition at finer scales, which suggests that this species is not completely redundant and that its disappearance may have affected local biodiversity. Similarly, assemblages on transplanted Phyllospora differed from those on Ecklonia and Sargassum at restored sites, but did not always resemble assemblages from extant natural Phyllospora populations, even 18 months after transplantation. These experiments indicate that restoration of key habitat‐forming seaweeds not only recovers the algal species but also reduces risks of losing habitat diversity for epifauna and their consumers. However, restoration of all the original biodiversity associated with these seaweeds can be a difficult, complex, and long‐term process.     

Response of a native,herbivorous snail to the introduced seaweed <Emphasis Type="Italic">Sargassum muticum</Emphasis>

The role of native consumers in mediating biological invasions is poorly understood. In theory, there are reasons to expect both strong and weak effects of native consumers on non-native species. However, non-native ranges may include multiple regions or even continents, each with its own suite of consumers and invader–consumer interactions may play out differently in different places and times. In this Washington State (USA) study we found that the common herbivorous snail Lacuna vincta was 2–9 times more abundant on the non-native seaweed Sargassum muticum, compared to native kelps. Choice feeding trials with fresh tissue and artificial foods both suggest that S. muticum is a preferred food for Lacuna vincta. Lab experiments indicated that L. vincta did not experience diminished predation by two common predators on Sargassum muticum compared to native kelp hosts. Our results suggest that Sargassum experiences considerable herbivory by Lacuna vincta in our study region, a conclusion that is consistent with previous work and our own field observations. In our system, L. vincta and S. muticum have been coexisting in the same habitats for at least 50 years and available data suggest that it acquired a preference for S. muticum more than 30 years after the initial invasion. Comparison of our results to recent work on Sargassum–herbivore interactions in Europe suggests that the response of native consumer communities to S. muticum varies both within and among regions. Geographic and temporal variation in the response of native consumers are likely to be hallmarks of many large-scale invasions.     

Amphipods on seaweeds: partners or pests?

Summary Herbivorous marine amphipods have been implicated as important grazers on filamentous and ephemeral algae, and thus as beneficial to macrophytes in reducing overgrowth by epiphytic competitors. In North Carolina, USA, amphipods comprise 97% of all macroscopic animals inhabiting the abundant brown seaweed Sargassum filipendula, and peak in abundance between late winter and early summer. I used outdoor tank experiments to test the species-specific impact of common phytal amphipods on the growth of Sargassum and its epiphytes. The results show that seaweed-associated amphipods are a trophically diverse group that could either increase or decrease host fitness depending on their feeding preferences. The amphipods Ampithoe marcuzii, Caprella penantis, and Jassa falcata each significantly reduced growth of epiphytes on Sargassum plants relative to amphipod-free controls, while Ericthonius brasiliensis had no significant effect on Sargassum or its epiphytes. However, amphipod grazing was not necessarily beneficial to Sargassum. A. marcuzii consumed Sargassum in one outdoor tank experiment, reducing its mass by 11%, while Sargassum plants without amphipods grew by 81%. Epiphytes (mostly diatoms and the filamentous brown alga Ectocarpus siliculosus) and detritus remained abundant on these plants suggesting that A. marcuzii preferred the host to its epiphytes. Similarly, when given simultaneous access to Sargassum and to several common foliose and filamentous epiphytes in the lab, A. marcuzii ate Sargassum almost exclusively. The other three amphipods ate no macroalgae. In contrast to A. marcuzii, C. penantis consistently reduced epiphytes with no negative effect on Sargassum. Thus the species composition of the amphipod fauna can determine whether these animals increase or decrease seaweed fitness.     

Application of polymerase chain reaction (PCR) using random amplified polymorphic DNA (RAPD) primers in the molecular identification of selected Sargassum species (Phaeophyta,Fucales)

The random amplified polymorphic DNA-polymerase chain reaction (RAPD-PCR) was used for the molecular characterisation and identification of Sargassum spp. A total of 17 samples of Sargassum (Sargassaceae, Fucales) was obtained from various localities around Peninsular Malaysia and Singapore. On the basis of morphological characteristics, the samples were tentatively grouped into five species: Sargassum baccularia, S. glaucescens, S. oligocystum, S. polycystum and S. siliquosum. By RAPD-PCR, five of 31 random primers tested generated reproducible amplification products, and polymorphic loci were detected by four of them (OPA02, OPA03, OPA04, OPA13). The RAPD-PCR profiles did not correlate with the morphological grouping into five species and extensive variation was detected between different isolates of the same species. A 450 base pair fragment generated using OPA13 was detected in 12 of 17 samples of Sargassum. This fragment was also present in profiles from Turbinaria (Sargassaceae). This study suggests that RAPD-PCR is useful in discriminating individual samples of the genus Sargassum and in developing fingerprints for them.     

Sea-land transitions in isopods: pattern of symbiont distribution in two species of intertidal isopods <Emphasis Type="Italic">Ligia pallasii</Emphasis> and <Emphasis Type="Italic">Ligia occidentalis</Emphasis> in the Eastern Pacific

Studies of microbial associations of intertidal isopods in the primitive genus Ligia (Oniscidea, Isopoda) can help our understanding of the formation of symbioses during sea-land transitions, as terrestrial Oniscidean isopods have previously been found to house symbionts in their hepatopancreas. Ligia pallasii and Ligia occidentalis co-occur in the high intertidal zone along the Eastern Pacific with a large zone of range overlap and both species showing patchy distributions. In 16S rRNA clone libraries mycoplasma-like bacteria (Firmicutes), related to symbionts described from terrestrial isopods, were the most common bacteria present in both host species. There was greater overall microbial diversity in Ligia pallasii compared with L. occidentalis. Populations of both Ligia species along an extensive area of the eastern Pacific coastline were screened for the presence of mycoplasma-like symbionts with symbiont-specific primers. Symbionts were present in all host populations from both species but not in all individuals. Phylogenetically, symbionts of intertidal isopods cluster together. Host habitat, in addition to host phylogeny appears to influence the phylogenetic relation of symbionts.