Genetic variation and biochemical systematics in the marine bryozoan Alcyonidium mytili

For many years the taxonomic position of the marine ctenostome bryozoan Alcyonidium mytili Dalyell (1848) has been in doubt, with some authorities regarding it as being conspecific with A. polyoum (Hassall). We have set out to resolve this problem using enzyme electrophoresis to examine the genetics of the population structure in the two species. The results show beyond reasonable doubt that sympatric populations of the two species are indeed from quite separate and non-interbreeding gene pools. An examination of various allopatric populations shows a new species of A. mytili to exist sympatrically with the other at Langstone Harbour, Hampshire, England. This species can be readily distinguished on morphological as well as genetic ground. Samples of A. mytili from Guernsey also show substantial genetic differences from other populations examined. It is tentatively concluded that this population also represents a separate gene pool and consequently should be recognised as a third species of A. mytili.     

Sexual selection and the evolution of bird song: A test of the Hamilton-Zuk hypothesis

Summary Hamilton and Zuk (1982) suggested that secondary sexual characters evolve because they allow females to assess a potential mate's ability to resist parasites. A prediction of this theory is that the degree of elaboration of secondary sexual characters should be positively correlated with parasite load across species. In support of their hypothesis, Hamilton and Zuk reported a correlation across North American passerine species between haematozoa prevalence and both brightness and song complexity and variety, scored on a subjective six point scale. Here we show that this relationship is confounded by phylogenetic associations. We use quantitative data on song duration, inter-song interval, song continuity, song rate, song versatility, and song and syllable repertoire size for 131 species of European and North American passerines to test the Hamilton-Zuk hypothesis. Across species, there are significant negative relationships between haematozoa prevalence and song continuity, contrary to the direction predicted by Hamilton and Zuk. In accordance with their prediction, there is a positive correlation with song versatility. However, these relationships come about through taxonomic associations: within taxa there are no consistent relationships between any of the song variables and haematozoa prevalence. None of the other song variables correlate with haematozoa prevalence. We conclude that there is no evidence of an association between song elaboration and parasites. Offprint requests to: A.F. Read     

Cross-shelf distribution of copepods and fish larvae across the central Great Barrier Reef

The distribution of total dry weight of zooplankton, copepod numbers and ichthyoplankton across the outer continental shelf in the central Great Barrier Reef was examined at bi-weekly intervals for three months over summer of 1983. Copepods were sampled (236 m net) within 10 m of the surface and within 10 m of the bottom. Mean densities in surface waters decreased markedly from the mid-shelf to outer shelf and the Coral Sea, but no cross-shelf gradient occurred in the bottom-water. Densities of copepods on the mid-shelf (surface and bottom waters) and in bottom-waters of the outer shelf were typically ca. 400 m^–3. Significantly lower densities (ca. 100 m^–3) occurred in surface waters of the outer shelf, except during outbursts of Acartia australis, when densities in these waters differed little from those elsewhere on the shelf. In oceanic waters, 10 km from the outer shelf station, copepod densities in surface waters were ca. 40 m^–3. Four of the five most abundant copepod taxa in surface waters, Paracalanus spp., Eucalanus crassus, Acrocalanus gracilis and Canthocalanus pauper, tended to be most abundant at the mid-shelf end of the transect. Acartia australis was sporadically very abundant in surface waters of the outer shelf, as was Paracalanus spp. in bottom-water of the outer shelf. An assemblage of Coral Sea species of copepod occurred in bottom-water of the outer shelf during two major intrusions, but not at other times. Densities of all common species varied considerably between cruises. Maximum densities of all common species except A. australis tended to be associated with diatom blooms linked to intrusions but a bloom did not necessarily mean all common species were abundant. Fish larvae included both reef and non-reef taxa, with reef taxa predominating on the outer shelf (approx 2:1 in density of individuals) and non-reef taxa dominating in nearshore samples (approx 2:1). Nine of the ten most abundant taxa analysed showed highly significant variation in numbers among stations and all but one of these also exhibited significant station x cruise interactions. Interactions generally reflected changes in the rank importance of adjacent stations from one cruise to the next or lack of any significant cross-shelf variation on some cruises where overall abundance of the taxa was low.     

Physiological responses of Mytilus edulis and Cardium edule to aerial exposure

Physiological responses of two bivalves (Mytilus edulis L. and Cardium edule L.) to intertidal conditions were studied. Specimens were collected from S. W. England in autumn/winter, 1980, and acclimatized to either intertidal or subtidal regimes before measurement of rates of heat dissipation and oxygen uptake during 5 h of air exposure, and rates of , particle clearance, ammonia excretion, and food-absorption efficiency during 7 h of reimmersion. Subtidal individuals were either intermittently or continuously fed in order to distinguish the effects of periodic food supply from the effects of air exposure. Specimens of M. edulis had low aerial rates of (14 to 20% of aquatic rate), and was greater than the energy equivalent of , indicating that they were largely anaerobic. In contrast, C. edule air-gaped and had higher aerial rates of and (50 to 75% of aquatic rate). There were behavioural and metabolic differences in the responses of intertidally and subtidally acclimatized mussels and cockles to air exposure. Intertidal individuals of both species were more quiescent, had lower aerial rates of and , and showed a conditioned response at the expected time of reimmersion. The reduction in aerial rate of was an energy-saving mechanism and the payment of the oxygen-debt within 2 h of reimmersion represented a significant cost. The heat increment associated with feeding and digestion was estimated as 15 to 17% of the oxygen uptake by M. edulis during all stages of recovery. M. edulis adapted to the intertidal regime by reducing its time-averaged aerial and aquatic rates of ammonia excretion. In contrast, C. edule maintained a high aerial and aquatic rate of ammonia excretion. The clearance rates of M. edulis recovered rapidly (0.5 to 1.5 h) following reimmersion, whereas those of C. edule recovered more slowly, particularly for the subtidal individuals following acute exposure (>4 h). There was no evidence of increased clearance rate or absorption efficiency by intertidal individuals to compensate for the loss of feeding time. Intertidally acclimatized individuals of M. edulis and C. edule had more energy available for growth (scope for growth) integrated over a 12 h period and higher growth efficiencies than subtidal individuals subjected to acute air exposure and intermittent feeding regimes. This was the result of reduced aerial and aquatic rates of energy expenditure, a relatively small cost in terms of the payment of oxygen-debt and a rapid recovery of clearance rate following reimmersion.     

Spatial and temporal patterns of recuruitment of juvenile coral reef fishes to coral habitats within “One Tree Lagoon”, great barrier reef

Recruitment of juvenile fishes to coral grids, each comprising 4 colonies of 3 species of coral (Acropora formosa, Seriatopora hystrix and Pocillopora damicornis), was examined at 4 widely separated sites within the lagoon of One Tree Reef over two successive summers and intervening months (November 1976–January 1978). Recruitment was highly seasonal, with most recruitment occurring during summer. For many species, numbers settling differed greatly from year to year with total numbers (over all sites) differing as much as an order of magnitude between summers. Many fish species demonstrated marked preference among the three coral species as settling sites. The distribution of each of the 20 commonest species across the 4 lagoon sites differed significantly from a random pattern. Differences of an order of magnitude were common in the numbers of a given species recruited to different sites. Each site was preferred by at least one species. In each of 5 cases examined, the pattern of settlement of the species across the 4 sites changed significantly from one summer to the next. The distribution of recruits of a number of species corresponded to the distribution of adults, but for other species there was no correspondence. It is concluded that, at the spatial scale examined, patterns of recruitment of some taxa are consistently more variable than those of other taxa. The implications of spatial variability of recruitment for the distribution of adult fish is discussed and the importance of being able to discriminate such natural variability from other kinds of change is stressed.     

Distribution of the cladoceran Podon polyphemoides in the Chesapeake Bay

The distribution of the cladoceran Podon polyphemoides (Leuckart) in the Chesapeake Bay (USA) estuarine system was determined by a quantitative pump sampling method, and the patterns of abundance were correlated with temperature and salinity distributions. The species was seasonally recurrent, with distinct population maxima in the central portion of the bay. Population densities in excess of 60,000 podonids/m³ have been recorded. The podonids first appeared in the spring in the shallow tributaries, when water temperatures near the bottom reached 6°C. The vernal populations disappeared when summer temperatures exceeded 27°C, but reappeared in the fall as the water cooled. The species was euryhaline and eurythermal in its distribution, but the greatest concentrations were attained within relatively narrow zones of temperatures between 11^o and 26°C, and salinities between 8 and 18. The production of males, sexual females and sexual eggs occurred both in the spring and the fall between the thermal limits of 11^o and 17°C.     

Minimal predation upon meiofauna by endobenthic macrofauna in the Exe Estuary,south west England

Field manipulation experiments were performed in the Exe Estuary, south west England, in October 1988, to investigate the importance of the meiofaunamacroinfauna trophic link in benthic trophodynamics. Four hypothetically meiofauna-predacious endobenthic macrofauna species were selected for manipulation using the criteria of high abundance and different modes of feeding: Cerastoderma edule (filter-feeder), Nereis diversicolor (omnivorous scavenger), Ophelia bicornis (sand-ingester), Scrobicularia plana (deposit-feeder). Enclosures constructed from plastic tubing, 63 m nylon monofilament mesh and galvanized steel were deployed, containing adult members of these taxa at densities raised to approximately four times that of the surrounding sediment. The experiments ran for 12 tidal cycles. Differences in phyletic meiofaunal abundance between treatment and control enclosures at the termination of the experiment were assessed using both uni- and multivariate dataanalysis techniques. Only two univariate significant differences (p<0.05) existed for the N. diversicolor treatment and two for the S. plana treatment. Annelida, Turbellaria and copepod nauplii were the only meiofauna taxa affected. No univariate significant differences were recorded for either the C. edule or O. bicornis treatments. Multidimensional scaling ordination of the data revealed no consistent changes in community composition between treatments and controls. It is concluded that the experiments provide evidence of minimal predation by macroinfauna upon meiofauna.     

Vertical distribution and feeding patterns of midwater fish in the central equatorial Atlantic II. Sternoptychidae

The vertical distribution of seven sternoptychid species was examined from RMT 1+8 samples collected aboard R. V. Meteor in March-April 1979 and from Royal Research Ship R.R.S. Discovery in July 1974 in the central equatorial Atlantic. During daytime sternoptychids occupied depths between 200 and 1250 m, with Sternoptyx pseudobscura living deepest, centering between 800 and 900 m, and Argyropelecus sladeni most shallow, aggregating predominantly at 300 and 400 m. They are all considered limited or partial migrants, ascending only some 100 and 200 m towards the surface at night. Only A. sladeni was observed to enter the epipelagic zone (0 and 200 m).-Feeding patterns were investigated from stomach content analyses of Sternoptyx diaphana, S. pseudobscura, Argyropelecus sladeni and A. affinis. Additional stomach contents were analysed from samples of S. diaphana, A. hemigymnus and A. olfersi collected in June 1985 from F.R.V. Walther Herwig in the temperate NE Atlantic at 46°N, 17°W by means of the Engel Trawl. The food spectrum of the six species is generally described, and additional dietary evidence regarding calanoid copepod prey is provided for four of these taxa. All sternoptychid species investigated were planktivorous, feeding predominantly on copepods and ostracods, except for the largest size class, which preyed heavily on euphausiids and amphipods. The relationship of predator size towards prey type and prey size is analysed for both Sternoptyx species. Of these, S. pseudobscura in particular exhibits taxonomic selectivity towards polychaete prey. The diet of both species of Sternoptyx included a number of epipelagic or even neustonic calanoid copepod species which contributed more than 50% of the total copepod population by numbers. So far it is not known how the predators find access to prey organisms of the upper 200 m, as netfeeding is considered unlikely. Cyclopoid copepods of the genus Sapphirina were observed as dietary component particular of S. diaphana.     

The toxicity of aluminum to aquatic species in the US

In August 1988 the US Environmental Protection Agency (EPA) published an ambient water quality criteria document for the protection of aquatic organisms from the toxic effects of aluminum. The EPA water quality criteria were developed utilizing procedures described in theGuidelines for Deriving Numerical National Water Quality Criteria for the Protection of Aquatic Organisms and Their Uses, and after careful analysis of the latest toxicological information available to EPA on the adverse effects of aluminum on aquatic vertebrates, invertebrates and plants.The EPA criteria recommend that the four-day average concentration of aluminum not exceed 87 g L^–1 more than once every three years on the average when the ambient pH is between 6.5 and 9.0 to provide protection from chronic toxicity. The criteria also recommend that the one-hour average concentration of aluminum not exceed 750 g L^–1 more than once every three years on the average when the ambient pH is between 6.5 and 9.0 to provide protection from acute toxicity.Acute toxicity data for 20 species of freshwater aquatic organisms and chronic toxicity data for five species of freshwater aquatic organisms were utilized to develop the EPA water quality criteria. Striped bass and brook trout were observed to be the two most sensitive North American species to the toxic effects of aluminum. Aluminum toxicity was also observed to be increased at lower pH.     

Long-term changes in populations of subtidal bivalves (Abra alba and A. prismatica) from the Bay of Morlaix (Western English Channel)

Two populations of Abra alba (Wood) and one of A. prismatica (Montagu) (Mollusca: Bivalvia) were studied over a 10 yr period (1977–1987) in two muddy fine-sand subtidal communities of the Bay of Morlaix, France. The survey provided an example of long-term changes in the three Abra spp. populations, which displayed synchronized changes, with a regular annual cycle and increasing densities during 1979–1980 related to the higher concentration of organic matter resulting from the Amoco Cadiz oil spill in March 1978. A. alba rapidly adapts its demographic strategy to eutrophic conditions by increasing its reproductive potential, growth, and abundance. During times of eutrophication, A. alba has three spawning periods and three recruitments per year as opposed to two spawning periods and two recruitments per year during oligotrophic conditions. Growth of the juveniles of this species is insignificant until spring for individuals recruited in the autumn, whereas individuals which settle during spring or summer display immediate rapid growth. A. prismatica has a low capacity to adapt to eutrophic conditions. It has one annual period of sexual maturation at the end of the summer, with spawning in September–October and settlement beginning in mid-November. Growth of the juveniles after settlement is also insignificant until April. These results enable comparison of the demographic strategies of these two sympatric species.     

Zooplankton of the Bristol Channel and Severn Estuary. The distribution of four copepods in relation to salinity

The seasonal variations in distribution and abundance of the common zooplankton species in the Bristol Channel and Severn Estuary were related to the salinity regimes observed over the period November 1973 to February 1975. The dominant constituents in all regions were the calanoid copepods, which reached maximum densities in July: approximately 100 times their winter levels. Four zooplankton assemblages were recognised using an objective classification program which computed similarity coefficients and used group-average sorting. The assemblages existed along the salinity gradient observed from the Severn Estuary to the Celtic Sea. The assemblages were classified as true estuarine, estuarine and marine, euryhaline marine and stenohaline marine and were characterized by the copepods Eurytemora affinis (Poppe) (<30S), Acartia bifilosa var. inermis (rose) (27 to 33.5S), Centropages hamatus (Lilljeborg) (31 to 35S) and Calanus helgolandicus (Claus) (>33S), respectively.     

Proclivity and effectiveness in gall defence by soldiers in five species of gall-inducing thrips: benefits of morphological caste dimorphism in two species (<Emphasis Type="Italic">Kladothrips intermedius</Emphasis> and<Emphasis Type="Italic"> K. habrus</Emphasis>)

The essence of eusociality is a trade-off between producing ones own offspring and helping collateral kin via such activities as defence, foraging and brood rearing. This trade-off often involves morphological differences between helper and reproductive castes, but the advantages, correlates and phylogenetic context of morphological caste differentiation have seldom been analysed. Six species of Australian gall-inducing thrips on Acacia show morphological polymorphism. One morph, referred to as a soldier, has reduced wings and antennae but greatly enlarged fore-femora, which are thought to be adaptations for gall defence. The other, dispersing morph, has fully developed wings and relatively slight fore-femora. Here, we quantify the defensive behaviour of soldier morphs, and compare soldier and foundresses, using behavioural assays designed to measure proclivity to attack kleptoparasites (specialised invaders in the genus Koptothrips) and effectiveness in killing them. In all five species investigated, soldiers were able to kill Koptothrips. Moreover, the effectiveness of soldiers was relatively high in the more-derived species in the phylogeny of the clade, Kladothrips intermedius, K. habrus and K. waterhousei. Soldiers of K. intermedius and K. habrus also killed kleptoparasites more effectively than did foundresses, and K. habrus soldiers exhibited higher proclivity to attack than did foundresses. Data from naturally invaded galls demonstrate that soldiers in the field do kill Koptothrips, and vice versa. These results show that soldiers of Australian gall thrips are motivated and effective for gall defence.Communicated by D. Gwynne     

Occurrence and characteristics of unusual protistan symbionts from surgeonfishes (Acanthuridae) of the Great Barrier Reef,Australia

The occurrence of unusual symbiotic microorganisms was examined in the intestines of a range of fish from the Great Barrier Reef, Australia. The fish taxa examined included 26 species of the family Acanthuridae, as well as representatives of phylogenetically related and herbivorous taxa. The microorganisms, referred to as protists, were only found in herbivorous and detritivorous members of the Acanthuridae. Protists were not found in planktivorous acanthurids, nor in any members of the families Kyphosidae, Pomacentridae, Scaridae, Zanclidae, Siganidae and Bleniidae we examined. In addition, protists were absent from the herbivorous acanthurids A. xanthopterus and A. nigricans. A range of protist forms, characterized by differences in size (8 to 417 m), shape and mode of cell division (daughter-cell production and binary fission), was observed. The occurrence of these forms appeared to be correlated with host feedingecology. Large forms (>100 m) of the protists were only found in acanthurids which fed over hard-reef substrata. Smaller forms were found in sand-grazing and detritivorous species. One of the protist forms appears identical to protists previously reported from Red Sea acanthurids.     

Allozyme and colour differences between two sibling species of the heteronemertean Lineus torquatus from the Sea of Japan

Brown and cherry colour morphs of the heteronemertean Lineus torquatus Coe, 1901 from the Vostok Bay of the Sea of Japan were compared using allele frequencies at 18 isozyme loci. These morphs shared no common alleles at 9 of the 18 loci. The observed genetic differences are strong evidence that the brown and cherry colour morphs, which occur sympatrically, are different species. Nei's genetic distance between these species (D=1.135) is considerably higher than typical D values for congeneric animal species. As with much previous work on nemerteans, intraspecific allozymic variability in both species was high, with mean expected heterozygosity values 0.286±0.049 and 0.159±0.051 for L. torquatus (brown) and L. torquatus (cherry) species, respectively.     

Diversity of swimming behavior in hydromedusae as related to feeding and utilization of space

Feeding behaviors of the following 4 species of hydromedusae are described from field and laboratory observations: Probosidactyla flavicirrata, Stomotoca atra, Phialidium gregarium and Polyorchis penicillatus. Feeding efficiency of medusae has previously been considered equivalent to fishing with a given amount (combined tentacle length) of adhesive fishing line; however, detailed observation shows that behavior of medusae greatly modifies the fishing capacity of each species. It is hypothesized that in addition to (1) tentacle number and length, the following factors strongly influence feeding efficiency: (2) tentacle posture, (3) velocity of tentacles moving through water (4) swimming pattern of medusa, (5) streamlining effects of medusa bell on water flow, (6) diameter of prey, (7) swimming pattern and velocity of prey. Each species of hydromedusa utilizes the above factors in different combinations.Mailing address     

Sequential female choice and the previous male effect in sticklebacks

Summary Female choice, identified as a major force in sexual selection theory, has recently been demonstrated in a number of species. These tests concentrated on simultaneous choice situations although females have to compare males sequentially in most territorial species, which is the more demanding task. Here it is shown that female three-spined sticklebacks, Gasterosteus aculeatus L., rate sequentially presented males according to their brightness. With increasing costs of sampling the females become less choosy. Furthermore, a male's attractiveness has a significant effect on the female's rating of the next male; a given male is rated higher when preceded by a duller male than by a brighter one and vice versa. Female sticklebacks use a stochastic decision rule in sequential mate choice that is attuned to the attractiveness of the present and previously encountered male. This demonstration of a previous male effect not only indicates an efficient mechanism for finding the best of a number of males but also extends the applicability of sexual selection theory.Offprint requests to: T.C.M. Bakker     

Biology of penaeid prawns in the Suez Canal lakes

A study was made from January 1988 to March 1989 of the penaid prawns in the Great Bitter Lake and Lake Timsah located in the central part of the Suez Canal. Two species of Red Sea origin were investigated,Metapenaeus stebbingi andTrachypenaeus curvirostris; the former is by far the commoner. Both species displayed seasonal breeding over the period April to October, with peaks in the early and late parts of the season.M. stebbingi had two main cohorts each year. A spring-spawned cohort survived 16 mo, and attained modal sizes of 23 mm carapare length (CL) for females and 16 to 17 mm CL for males. The autumn-spawned cohort survived 13 mo, and reached modal sizes of 20 mm and 15 mm CL for males and females, respectively. The reasons for the success ofM. stebbingi in the Canal lakes, in contrast to other Lessepsian penaeids which have flourished in the Mediterranean Sea, have been considered. The main factors are that it is essentially a shallow-water species, with excellent tolerance of salinity variations and adverse environments. However, the Canal lake populations have smaller maximum sizes, and mature earlier, than those in open seas.     

Intraspecific behaviors and major cheliped sexual dimorphism in three congeneric callianassid shrimp

Decapod callianassid shrimps are usually solitary occupants of their burrows. They are known to show distinct sexual dimorphism of the major cheliped, which is used as a weapon for intraspecific fighting. Three species of Nihonotrypaea occur in an estuary in southern Japan; they consist of two tidal flat species (N. harmandi; N. japonica) and one boulder beach species (N. petalura), with maximum population densities of 1,400, 340, and 12 m^–2, respectively. The major cheliped size and total length of shrimp were recorded from each population. The degrees of major cheliped sexual dimorphism were ordered as N. harmandi >N. japonica >N. petalura. In the laboratory, intra- and intersexual behaviors at forced encounters between two shrimps were recorded, for the former behavior throughout the year and the latter in the non-breeding season. At their intersected burrows, the shrimps either fought or retreated or filled the burrow crack. Males interacted aggressively with each other, with the intensity being N. petalura >N. harmandi N. japonica. Females of the tidal flat species were non-aggressive, while those of N. petalura were as aggressive with each other as were males. Intersexually, males of all species and females of N. petalura were much less aggressive than intrasexually. In N. petalura only, burrow-sharing behavior between sexes occasionally occurred. The interspecific difference in these behaviors is in parallel with the degree of major cheliped sexual dimorphism. Different intensities of intrasexual competition for mates could have been imposed by the different population densities of these species.Communicated by T. Ikeda, Hakodate     

Spatial and temporal variation in the diet of a high trophic level predator,the Australian fur seal (<Emphasis Type="Italic">Arctocephalus pusillus doriferus</Emphasis>)

This study quantifies the manner in which Australian fur seals, Arctocephalus pusillus doriferus, use their prey in a spatial and temporal context. We analysed 977 scat and 66 regurgitate samples collected from Tasmanian breeding colonies and haul-outs between 1994 and 2000. Diagnostic prey remains identified in the scats represented 35 fish taxa and 8 cephalopod taxa. The main taxa identified in scats, where frequency of occurrence was 10%, were leatherjacket species (family Monocanthidae), redbait (Emmelichthys nitidus), barracouta (Thyrsites atun), jack mackerel (Trachurus declivis) and red cod (Pseudophysis bachus). Regurgitates were dominated by cephalopods, primarily Goulds squid (Nototodarus gouldi), Octopus maorum, O. berrima/pallidus and Sepia apama. Discriminant function analyses indicated that there were generally no significant differences in the composition of the diet between colonies within a year, suggesting that prey distribution is fairly uniform throughout Bass Strait at those time scales. The diet at breeding colonies, however, exhibited significant inter- and intra-annual variation, determined by the presence of several key taxa, such as barracouta and a species of scorpionfish (family Scorpaenidae). The diet composition also varied regionally, between Bass Strait and southern Tasmania in spring 1999 and autumn 2000, with redbait, barracouta and a species of scorpionfish identified as the main taxa contributing to this difference. Redbait occurred in the diet only in southern Tasmania, whereas barracouta and scorpionfish occurred only in Bass Strait.Electronic Supplementary Material Supplementary material is available in the online version of this article at http://dx.doi.org/10.1007/s00227-003-1219-0.Communicated by G.F. Humphrey, Sydney     

Distribution of algae on tropical rocky shores: spatial and temporal patterns of non-coralline encrusting algae in Hong Kong

Encrusting algae have been described as dominant space occupying species on rocky shores around the world. Despite their abundance, however, most studies classify species under generic names (e.g. Ralfsia sp.) or as a functional group (e.g. encrusting algae), thereby underestimating the number of species present and their ecological importance. Studies on six rocky shores of varying exposure, in Hong Kong, recorded eight common species of encrusting algae. The greatest abundance of encrusting algae was recorded on shores of intermediate exposure, where four distinct zonation bands could be identified; a cyanobacterial Kyrtuthrix-Zone in the upper midlittoral, a Bare-Zone below this, a Mixed-Zone in the lower midlittoral and a Coralline-Zone in the infralittoral fringe. Abundance declined on shores of greater and lower exposure to wave action, where bivalves and barnacles were competitively dominant. Certain species were found in greater abundance on exposed shores (e.g. Dermocarpa sp. and Hildenbrandia occidentalis), while others preferred more sheltered shores (e.g. Hildenbrandia prototypus and Kyrtuthrix maculans). With the exception of some cyanobacterial crusts, the abundance of encrusting algae was always greatest towards the low shore, an area of decreased physical stress and increased herbivore density. Zonation patterns showed seasonal variation associated with the monsoonal climate of Hong Kong. Most species increased in abundance during the cool season, while during the summer months the cover and vertical extent of encrusting algae decreased in relation to summer temperatures, although K. maculans increased in abundance during the summer. On Hong Kong shores, encrusting algae have a high species richness and exhibit within-functional group spatial and temporal variation which is mediated by herbivory and seasonal, physical stresses.