Experimental assessment of factors influencing microhabitat selection by the two tidepool fishes Oligocottus maculosus and O. snyderi

Oligocottus maculosus Girard and O. snyderi Greeley are two similar tidepool fishes with differing, but overlapping, intertidal distribution patterns. The factors influencing microhabitat selection and distribution patterns were experimentally assessed to elucidate the mechanisms by which the two species partition the resources of the intertidal environment. O. snyderi displays a much more rigid substrate-cover requirement. This, combined with its stenothermal nature serve to strongly restrict the intertidal distribution of this species. O. maculosus has a more generalized substrate-cover requirement and is also known to be eurythermal and euryhaline. Thus, O. maculosus is a ubiquitous intertidal species. Other behavioral attributes of O. maculosus, such as its tide-related locomotor activity and preference for shallow water, facilitate its particular intertidal distribution pattern. Similar depth preferences or tide-related activity were not observed in O. snyderi. Interspecific competition and related agonistic behavior were absent from both species and are probably not important to their microhabitat selection and intertidal distribution patterns.Based on a portion of a dissertation submitted in partial fulfillment of the requirements for the Ph.D. degree at the University of British Columbia, Vancouver, B.C., Canada     

Temporal and spatial settlement patterns of sympatric hermit crabs and the influence of shell resource availability

Larvae of marine organisms often need specific resources or environments at settlement, and their success at settlement might be strongly influenced by the abundance and distribution of such specific resources. The larvae of hermit crabs need small shells to settle, so it is thought that the distribution and abundance of small shells influence the settlement pattern of hermit crabs. To investigate the influence of small shell distribution on the settlement of pagurid hermit crab larvae, we conducted a field experiment at an intertidal rocky shore in Hakodate Bay, Japan. From the line-transect sampling in the field, we found that Pagurus middendorffii settled extensively in the offshore side of the intertidal zone while P. nigrofascia settled in the uppermost area of the intertidal zone. Small shells were most abundant in a narrow shallow trough, slightly offshore from the uppermost area of the intertidal zone. For both species, settler abundance was high where adults were abundant, but settler abundance did not appear to be related to shells abundance. An experiment to clarify settlement patterns showed that larval recruits tended to be similar to those in the line-transect sampling of settlers. Thus shells may not be a primary factor affecting settlement patterns at relatively large scale within the intertidal flat. However, when we analyzed the relationship of settlers and shells separately within each transect, the distribution of settlers was well explained by shell resource availability. Therefore on a smaller scale, shell availability may influence the number of settlers. Settlement periods of P. middendorffii and P. nigrofascia fully overlapped, so their larvae probably were affected by similar transport factors, such as current and tidal movement. Nevertheless they showed different spatial patterns of settlement.Communicated by T. Ikeda, Hakodate     

Assessment of intertidal growth and capacity adaptations in suspension-feeding bivalves

Curves relating instantaneous growth rate to aerial exposure were determined for six species of bivalves in laboratory and shore experiments. A dimensionless index equatable with the relative intertidal growth performance of a species was calculated by integration of the growth curve after converting both growth rate and aerial exposure to decimal fractions. Intertidal growth performance of the bivalves tested corresponded well with their natural levels of occurrence on the shore, and improved in the following order: Modiolus modiolus < Argopecten irradians < Ostrea edulis < Mytilus edulis < Crassostrea virginica. Geukensia demissa, for which an index value could not be determined, grew faster intertidally than subtidally. The relative contributions made to intertidal growth performance by energy-conserving and energy-supplementing capacity adaptations were assessed by resolving the growth curves into energy-input and energy-loss components. The rate of energy loss due to intertidal exposure was lower in the high-shore species, and also less affected by harsher subaerial conditions, than in the low-shore species. Moreover, M. edulis and C. virginica showed abilities to supplement energy input such that growth per unit immersion time was better at certain intertidal levels than subtidally. Energy conservation and supplementation in these forms made roughly equal contributions to their improved intertidal growth relative to species occurring lower on the shore.     

Comparative species composition and relative abundance of decapod crustaceans in marine habitats of Panamá

The community structure of the decapod crustacean fauna of 7 tropical, shallowwater, marine habitats (sandy beaches, mangrove swamps and rocky intertidal habitats on both the Pacific and Caribbean coasts of Panama, and Pocillopora damicornis coral habitat of the Bay of Panama) were examined and analyzed for species composition and relative abundances. Collections from the 7 habitats yielded 4361 individuals, representing 236 species. The number of species per habitat was (Pacific, Caribbean): sandy beach (16, 7); mangrove (20, 17); P. damicornis (53); rocky intertidal (78, 67). There were more species represented by more individuals in the Pacific habitats. An index of faunal similarity was calculated for each pair (Pacific-Caribbean) of habitats. This index is the number of ecologically similar congeneric species which occurred in both habitats expressed as a percentage of the total number of species present in the pair of habitats. For the sandy beach communities there were three Pacific species which were similar to three Caribbean species, a similarity of 6/23 or 26%. The index of similarity for the mangrove communities is 54% and for the rocky intertidal communities it is 37%. The P. damicornis community has affinities with the Pacific rocky intertidal community (18%), with that of the Caribbean rocky intertidal (16%) and with that of Indo-West Pacific pocilloporid corals (20%). A few specialized species dominated each of the communities. The habitats and the number of species accounting for over half of the individuals present are (Pacific, Caribbean): sandy beach (1, 1); mangrove (4, 4); P. damicornis (5); and rocky intertidal (3, 6). Most of the species in each community were represented by one or a few individuals.     

Diel variation and seasonal consistency in the fish assemblage of the non-estuarine Sulaibikhat Bay,Kuwait

In the period between September 1986 and August 1988, using an otter trawl, a total of 50 species from 30 families were captured in Sulaibikhat Bay, Kuwait. The dominant species were Liza carinata (Valenciennes), Pomadasys stridens (Forrskal) and Leiognathus brevirostris (Valenciennes). Although the total number of fishes captured in 1986–1987 and 1987–1988 differed by an order of magnitude, the seasonal changes in numbers, biomass and number of species in the intertidal and subtidal zones were consistent. Unbalanced three-way analysis of variance showed that significantly larger numbers of fishes and number of species, but not biomass, were captured in the intertidal at night compared to the intertidal during the day. This difference may be due to both increased net efficiency and movement of fishes into the intertidal zone at night to avoid piscivorous fishes that move from deep water into shallower water. During the day few fish are present in the intertidal zone and this may be to avoid predation by piscivorous birds. Several species of fishes were shown to have different patterns of behaviour during the diel period.     

Variation and persistence of the middle rocky intertidal community of central Chile,with and without human harvesting

Humans are an important intertidal predator in central Chile. Following a five-year study we report on the effects of the exclusion of humans from the rocky intertidal at Las Cruces, central Chile. The middle intertidal of harvested and non-harvested areas diverged in species diversity and composition during the experiment. In harvested areas the middle intertidal rocky shore was dominated throughout the study by a monoculture of mussels, Perumytilus purpuratus. When humans were excluded, the middle intertidal community switched to one dominated by barnacles (predominantly Jehlius cirratus and Chthamalus scabrosus); this community has persisted for at least three years, despite the presence of forces e.g. mussel larvae, that have the potential to alter the community structure. Such changes were mediated by the muricid gastropod Concholepas concholepas, a key-stone predator. As a consequence of the above changes, the species diversity, H', (primary space occupiers) in the non-harvested area increased from H'=0 at the beginning of the study in 1983 (when the middle intertidal community was dominated by mussels), to values ca H'=2 toward the middle of the study in 1984 (which coincided with the maximum predatory impact of C. concholepas) and subsequently decreasing to ca H'=0.5 at the end of the study in 1987 (when the middle intertida community was dominated by barnacles).Please address all correspondence to J. C. Castilla, Department of Zoology, University of Washington, Seattle, Washington 98195, USA, until March 1990     

Photoinhibition of photosynthesis in a sun and a shade species of the red algal genus Porphyra

Gametophytes of two species of Porphyra collected around San Juan Island, Washington in 1986 and acclimated to low light conditions in culture showed different resistances to photoinhibition of photosynthesis. The intertidal species P. perforata J. Agardh exhibited photoinhibition at one-third the rate exhibited by the subtidal species P. nereocystis Anderson following treatments at 2000 mol photons m^-2 s^-1 under conditions of full hydration and optimal temperature. The greater resistance of P. perforata to photoinhibition could not be attributed to reduced photosynthetic pigment concentration, higher photosynthetic capacity, avoidance of light by chloroplast movement or to enhanced rates of photorespiration. Total carotenoid concentrations were similar in the two species. It is probable that the mechanisms of this resistance are operating at the level of the thylakoid membranes. Resistance to photoinhibition represents an adaptation of photosynthesis in P. perforata which may contribute to its persistance in the extreme environment of its intertidal habitat.     

A Marine Pollution Study of Northeast Coastal Water Off Taiwan Island

Abstract

The coastal water of northeast Taiwan island, called ‘Yin-Yang Hai’ for its distinct yellow colour compared with blue offshore water, was investigated from 1989 to 1990 by the authors. Biological study showed the dominant species of plankton to be Copepoda, Cladocera, planktonic eggs and Diatoma. Dominant species of benthos were young crabs, Amphipoda and Annelida, with Amphipoda usually occurring in heavily polluted areas. Heavy metal data showed that the concentration of copper was high. the copper and iron concentration in algae of the intertidal zone was also high. the concentrations of iron and copper in inshore water were also higher than in offshore water. By comparison of the pH and salinity distribution of this area, we conclude that this coastal water has been polluted by acid waste water from coastal industry. the suspended solids concentration in sea water is high. Flocculation occurring at the boundary of fresh and saline water might be a reason for the distinct yellow colour of the water of this area. Further study is required.     

Influence of seasonal food abundance and quality on the feeding habits of an opportunistic feeder,the intertidal crab <Emphasis Type="Italic">Pachygrapsus marmoratus</Emphasis>

The abundance of trophic sources on the intertidal zone is discontinuous and their supply can vary both in a predictable or unpredictable way. The Mediterranean semi-terrestrial crab Pachygrapsus marmoratus, is known, as adult, to entirely rely on the intertidal trophic sources, and, consequently, it faces the fluctuations of nutritional sources and quality. To clarify the relationships between the feeding habits of an Italian population of P. marmoratus and the temporal variation of its food sources, we carried out a 2-year sampling protocol. Data on seasonal variation in composition of intertidal food item assemblages, on the average content in N and C of the commonest algae, on seasonal changes in crabs feeding habits were collected and compared using a suite of multivariate and univariate techniques. Results showed that P. marmoratus takes advantage of the recruitment phase of the most common invertebrates, affecting and controlling the abundance even of those species whose adults are out of its reach. It can act both as a herbivore, and as a carnivore that shifts between bivalves, more abundant in spring/summer, and the periwinkle Melaraphe neritoides, during winter. In conclusion, we provide evidences on the food choice of this common rocky shore species, which is very plastic and capable of relying on many trophic sources, possibly influencing the abundance and/or the population structure of a number of intertidal populations.     

Sand stress as a non-determinant of habitat segregation of indigenous (Perna perna) and invasive (Mytilus galloprovincialis) mussels in South Africa

Periodical sand inundation influences diversity and distribution of intertidal species throughout the world. This study investigates the effect of sand stress on survival and on habitat segregation of the two dominant mussel species living in South Africa, the invasive Mytilus galloprovincialis and the indigenous Perna perna. P. perna occupies a lower intertidal zone which, monthly surveys over 1.5 years showed, is covered by sand for longer periods than the higher M. galloprovincialis zone. Despite this, when buried under sand, P. perna mortality rates were significantly higher than those of M. galloprovincialis in both laboratory and in field experiments. Under anoxic condition, P. perna mortality rates were still significantly higher than those for M. galloprovincialis, but both species died later than when exposed to sand burial, underlining the importance of the physical action of sand on mussel internal organs. When buried, both species accumulate sediments within the shell valves while still alive, but the quantities are much greater for P. perna. This suggests that P. perna gills are more severely damaged by sand abrasion and could explain its higher mortality rates. M. galloprovincialis has longer labial palps than P. perna, indicating a higher particle sorting ability and consequently explaining its lower mortality rates when exposed to sand in suspension. Habitat segregation is often explained by physiological tolerances, but in this case, such explanations fail. Although sand stress strongly affects the survival of the two species, it does not explain their vertical zonation. Contrary to our expectations, the species that is less well adapted to cope with sand stress maintains dominance in a habitat where such stress is high. GI Zardi, KR Nicastro contributed equally to the work     

Measurement of chlorophyll fluorescence reveals mechanisms for habitat niche separation of the intertidal seagrasses <Emphasis Type="Italic">Thalassia hemprichii</Emphasis> and <Emphasis Type="Italic">Halodule uninervis</Emphasis>

In Taiwan, Thalassia hemprichii dominates the upper intertidal zone, whereas Halodule uninervis occupies the lower intertidal zone. We tested the hypothesis that T. hemprichii is better adapted to high irradiance and more resistant to air exposure than H. uninervis. The photosynthetic efficiency, damage, and extent of recovery were determined by measuring chlorophyll fluorescence using pulse amplitude modulated fluorometry. Both species growing in tidal pools, in response to high irradiance alone, revealed a small depression in maximal quantum yield of photosystem II (Fv/Fm) at noon. The second experiment compared the effect of air exposure alone and the combined effect of air exposure with high irradiance by interposing a shading screen on both species, growing in the intertidal zone over a diurnal cycle. Values of Fv/Fm of both the shaded and irradiated T. hemprichii remained high at low tide. However, H. uninervis exhibited a marked depression following air exposure and a synergistic depression under the combined effect. The experimental manipulations of exposure time demonstrated that the tolerance of T. hemprichii to the combined effect was longer and the recovery from air exposure following re-submersion was better than those of H. uninervis. Both species were more susceptible to the combined effect in the dry season than in the wet season. Our results suggest that air exposure is more important than high irradiance in constraining the distribution of H. uninervis in the upper intertidal zone. This was confirmed by transplantation experiments in which a rapid decline of H. uninervis was observed after transplantation into the upper intertidal zone. In the lower intertidal zone, measurements of the response of the photosynthetic electron transport rate to irradiance demonstrated that the transplanted T. hemprichii exhibited a sun-type response and H. uninervis a shade-type response.     

Comparisons of genetic population structures in four intertidal brachyuran species of contrasting habitat characteristics

Genetic population structures along the Japanese coast, analyzed by sequence data from the mitochondrial DNA COI region, were determined for four intertidal brachyuran species in the superfamily Thoracotremata (Ocypode ceratophthalma, Gaetice depressus, Chiromantes dehaani and Deiratonotus japonicus), which were characterized by different habitat requirements. O. ceratophthalma (seashore; supratidal sand) and C. dehaani (estuarine; supratidal marsh) showed no significant genetic differentiation among Japanese populations. The Japanese populations of O. ceratophthalma, however, were found to genetically differentiated from the Philippine population. G. depressus (seashore; intertidal cobbles) exhibited significant genetic differentiation between the Amami-Ohshima population and other local populations. D. japonicus (estuarine; intertidal cobbles) showed significant genetic differentiation among many local populations separated by about 30–1,200 km. The different patterns of genetic population structure recorded for the four species, thus, do not simply correspond to habitat type. An erratum to this article can be found at     

Comparisons of genetic population structures in four intertidal brachyuran species of contrasting habitat characteristics

Genetic population structures along the Japanese coast, analyzed by sequence data from the mitochondrial DNA COI region, were determined for four intertidal brachyuran species in the superfamily Thoracotremata (Ocypode ceratophthalma, Gaetice depressus, Chiromantes dehaani and Deiratonotus japonicus), which were characterized by different habitat requirements. O. ceratophthalma (seashore; supratidal sand) and C. dehaani (estuarine; supratidal marsh) showed no significant genetic differentiation among Japanese populations. The Japanese populations of O. ceratophthalma, however, were found to genetically differentiated from the Philippine population. G. depressus (seashore; intertidal cobbles) exhibited significant genetic differentiation between the Amami-Ohshima population and other local populations. D. japonicus (estuarine; intertidal cobbles) showed significant genetic differentiation among many local populations separated by about 30–1,200 km. The different patterns of genetic population structure recorded for the four species, thus, do not simply correspond to habitat type.     

The effects of sand movement on intertidal seaweeds and selected invertebrates at Bound Rock,New Hampshire,USA

A seasonal study of sand movement and the benthic intertidal organisms at Bound Rock, New Hampshire, USA was conducted between November, 1973 and February, 1975. The site is subjected to irregular sand fluctuations, as well as diurnal, neapspring and major summer sand intrusions. The abundance and distribution of intertidal species was interrelated with the historical sand fluctuations at the area. For example, the lower limits of Mytilus edulis, Balanus balanoides and Porphyra umbilicalis approximated the highest summer sand elevations. Highly abraded rock surfaces in the lower intertidal zone were dominated by opportunistic annuals (e.g. Enteromorpha spp.) and perennial psammophytic or sand-loving seaweeds (e.g. Ahnfeltia plicata and Sphacelaria radicans). Overall, the intertidal seaweed populations at Bound Rock showed a lower number of perennials and fewer species than adjacent rocky shores. The low species diversity of seaweeds at the study site is attributable to unstable environmental conditions and a limited number of habitats. Even so, several psammophytic microhabitats are evident at Bound Rock depending upon the extent of sand burial and abrasion. The morphological and reproductive adaptations of several psammophytic species are discussed.Published with the approval of the Director of the New Hampshire Agricultural Experiment Station as Scientific Contribution No. 812.Scientific Contribution No. 53 of the Jackson Estuarine Laboratory.     

Correlations between Human Collecting and Intertidal Mollusc Populations on Rocky Shores

We tested whether the population structures of intertidal molluscs varied with the activity of human collectors in northern Port Phillip Bay, Victora, Australia. We examined eight sites, two of which had been protected for around 70 years by the area being used as a rifle range with the other six being visited frequently by human. The shores were examined in two different years, and in 1991 we counted humans and recorded their activities. At heavily visited shores, almost half of the people seen were simply walking, while 25% were actively collecting, despite regulations prohibiting the collection of intertidal gastritis. The remaining people were fishing or skydiving. At the protected shores, we observed only one collector and a handful of walkers over more than 20 observation periods, including weekdays, normal weekends, and holiday weekends. In 1989 and 1991, we compared size distributions of seven intertidal species between sites in the two categories. Four mollify species are taken for food and bait; the remaining three served as “controls”. The size distributions and abundance of noncarbonated species (alicized napalm, Leaderless viola, and emization ebonies) did not differ between visited and protected sites in either year, although most species did vary between individual sites. In contrast, three of the four collected species, alicize trampoliners, Austrocochlea constricta, and Nerita atramentosa, were significantly larger at the protected sites, and N. atramentosa was markedly less abundant at heavily visited sites. The fourth collected species, Turbo undulatus, has a distribution extending into the subtidal zone, and it is possible that intertidal populations ate replenished from deeper water. The changes in mean size of molluscs are well within the range reported from South America and South Africa under subsistence fishing, and they demonstrate that at least close to major urban centers, human recreational activities can have substantial effects in developed countries.     

Respiration as a function of oxygen concentration in intertidal barnacles

Respiration as a function of oxygen concentration was studied in two species of intertidal barnacles: Balanus amphitrite amphitrite ¹ (Darwin) and B. tintinnabulum tintinnabulum (L.). A critical oxygen tension was observed in both species below which the respiratory regulation broke down. In B. amphitrite amphitrite the critical oxygen tension was 2.5 ml O₂/l, and in B. tintinnabulum tintinnabulum 3.5 ml O₂/l. Species differences and habitat relations were observed in both species in their respiratory adaptation to oxygen concentration. B. amphitrite amphitrite which inhabits oxygen-deficient areas was able to regulate to much lower concentrations than B. tintinnabulum tintinnabulum which inhabits oxygen-rich open intertidal regions.     

A diatom assemblage living below the surface of intertidal sand flats

The assemblages of species living on and in intertidal sand flats at Barnstable Harbor, Massachusetts (USA) are briefly described. An endopsammic assemblage of diatoms, mainly Amphora species, living below the surface is described for the first time in such a marine situation. Some aspects of its ecology and relationship to the other diatom assemblages living on the sediments are discussed.     

Desiccation enhanced nitrogen uptake rates in intertidal seaweeds

Desiccation increased nitrate and ammonium uptake rates upon resubmergence in late summer populations of the intertidal macroalgae Gigartina papillata (C.Ag.) J.Ag., Enteromorpha intestinalis (L.) Grev., Fucus distichus L., and Pelvetiopsis limitata (Setch) Gardn. The ratio of nitrogen uptake rates in desiccated plants to rates in hydrated plants (controls) was correlated with the position of the species in the intertidal zone. Gracilaria pacifica Abbott., the species occurring at the lowest shore level, showed no enhancement of nitrogen uptake following desiccation. The high intertidal species such as P. limitata and F. distichus showed a two-fold enhancement of nitrate and ammonium uptake following more extensive desiccation (>30%) and continued uptake even following severe desiccation (50 to 60%). After the plants had been desiccated, the increase in nitrate uptake rates upon submergence lasted much longer than a similar enhancement of ammonium uptake. The duration of the enhanced nitrate uptake was similar to the time required for total rehydration but the uptake rates were not related to the state of rehydration. The potential contribution that this enhanced nitrogen uptake following desiccation could make to total nitrogen procurement for growth is discussed. The experiments were carried out in 1979 or 1980 and repeated in 1981.This paper is dedicated to Dr. R. F. Scagel on the occasion of his retirement for his outstanding contribution to phycology     

Tolerance of high temperatures by some intertidal barnacles

The median lethal times of survival of the barnacles Elminius modestus, Balanus crenatus and B. balanoides, when continuously submerged at high temperatures, were determined for individuals collected in the summer and winter. In E. modestus and B. crenatus there was no seasonal change in high temperature tolerance. In B. balanoides, however, the adults were more susceptible in the winter than in the summer to temperatures a little below the upper lethal temperature. Using less comprehensive data for other British species of barnacles, it is concluded that, in general, the order of tolerance to high temperatures corresponds to the order of temperatures within the geographical and the intertidal distributions of the species. From the time-temperature-survival curves, intertidal barnacles are living closer to environmental temperatures than would be supposed on the basis of the measurement of the upper lethal temperature, which has been commonly measured for many species, but is of less ecological significance.     

Ecological effects of intertidal uplifting from nuclear testing

Uplifting of a portion of an intertidal rock bench by the Milrow underground nuclear test at Amchitka Island, Alaska affected algal abundance, species composition, and zonation. Parts of the populations of 8 species died off, primarily in the Hedophyllum sessile zone, and mostly within the first 6 months after the disturbance. The center of abundance of these species shifted seaward. Fucus distichus, a species of the upper intertidal, appeared in the area and recolonized a portion of the H. sessile zone. The observations indicate that intertidal algal communities at Amchitka Island are very sensitive to positive elevation changes.Contribution No. 376 of the College of Fisheries, University of Washinton, USA. This publication is based on work performed under U.S. Atomic Energy Commission Contract AT(26-1)-171 for Battelle Memorial Institute, Columbus Laboratories.