Community stability: effects of limpet removal and reintroduction in a rocky intertidal community

Summary The stability of a high rocky intertidal community was assessed in a controlled field experiment in which the most common consumers, limpets, were temporarily removed. Compared to the unmanipulated plots, the exclusion plots developed greater algal abundance and altered species composition of both algae and barnacles. The community was not perturbed beyond its capacity to recover, since the community structure of the limpet-removal plots converged on the structure of the unmanipulated plots following limpet reintroduction. Different components of the community recovered at different rates, depending on whether or not the species had a size-related escape from the limpets. Algae had no size-related escape from limpets. The difference in algal abundance between manipulated and unmanipulated plots lasted less than six months after limpet reintroduction. Barnacles, however, had a size-related escape from limpets and therefore recovered more slowly. The difference in barnacle species composition between the perturbed and unperturbed plots lasted for 17 months after limpet reintroduction. The length of the limpet removal period (16 or 28 months) did not appear to affect the rate of community recovery.     

Temporal variation and succession in an algal-dominated high intertidal assemblage

We determined whether temporal variation and succession were similar among sites with similar species composition by sampling unmanipulated and cleared plots in a high intertidal assemblage dominated by Endocladia muricata and Mastocarpus papillatus. Sampling was done for 6 years at six sites spanning over 4° of latitude in California. Ten 1×2-m permanent plots were chosen in the central portion of the assemblage at each site. Four of these served as unmanipulated controls, three were cleared (scraped and burned) in the spring of 1985, and three were cleared in the fall of 1985. The cover of sessile and density of motile species were determined by subsampling within the plots from 1985 until 1991. Recovery of the clearings was determined by their similarity to the controls. The algae E. muricata, M. papillatus, and Fucus gardneri, and the barnacle Balanus glandula, were the most abundant sessile organisms in the control plots, although the latter never exceeded 12% cover at any site. The grazing gastropods Littorina scutulata/plena, various limpets, and Tegula funebralis were the most common mobile organisms. The species composition of the common species remained constant in the control plots over the study period and there were few large changes in relative abundance. Significant seasonal variation was detected in 11 species but variation was commonly site-specific.Ephemeral algae were abundant during early succession at only two of the six sites, and barnacle cover was low (<15% cover) at four sites and moderate (15-50% cover) at the remaining two throughout succession. Recovery rate varied considerably among sites and between times of clearing (1-10%/month). Correlations between ephemeral algae and grazer abundance, and between these variables and recovery rate were not significant. The effects of grazers on recovery rate were only evident at one site where they appeared to reduce an initially high cover of ephemeral algae and delay the establishment of perennials. Some of the largest differences in recovery rate were between clearing times, associated with differences in the phenology of the dominant perennial algae. In spite of these differences, most plots recovered by the end of the study period.These results indicate that the assemblages in the control plots at each site were relatively stable and, while successional pathways and processes varied, the assemblage at most sites still recovered. Current models, based largely on biological interactions, that attempt to explain within assemblage structure and succession were not broadly applicable.     

Predation by gulls on crabs in rocky intertidal and shallow subtidal zones of the Gulf of Maine

Most organisms in intertidal areas are marine in origin; many have distributions that extend into the subtidal zone. Terrestrial predators such as mammals and birds may exploit these animals during low tide and can have considerable effects on intertidal food webs. Several studies have shown that avian predators are capable of reducing densities of sessile and slow-moving intertidal invertebrates but very few studies have considered avian predation on mobile invertebrate predators such as crabs. In this study, we investigated predation by Great Black-backed Gulls (Larus marinus Linnaeus) on three species of crabs (Cancer borealis Stimpson, Cancer irroratus Say, and Carcinus maenas Linnaeus). The study was at Appledore Island, ME (a gull breeding island) and 8 other sites throughout the Gulf of Maine, including breeding islands and mainland sites. On Appledore Island, intertidal and subtidal zones provided over one-third of prey remains found at gull nests, and crabs were a substantial proportion (∼ 30% to 40%) of the total remains. Similarly, collections of prey remains from intertidal areas indicated that crabs were by far the most common marine prey. C. borealis was eaten far more often and C. irroratus and C. maenas less often than expected at each site. Comparing numbers of carapaces to densities of crabs in low intertidal and shallow subtidal zones at each site, we estimated that gulls remove between 15% and 64% of C. borealis during diurnal low tides. The proportion of C. borealis eaten by gulls was independent of proximity to a gull colony. Approximately 97% of the outer coast of Maine is within 20 km of a breeding island. Thus, a lot of gull predation on crabs may occur throughout the Gulf of Maine during summer months. Crabs are important predators of other invertebrates; if predation by gulls reduces the number of crabs in intertidal and shallow subtidal areas, gulls may have important indirect effects on intertidal food webs.     

An experimental test of the effects of variation in recruitment intensity on intertidal community composition

The rocky intertidal has been a model system for experimentally testing hypotheses regarding the factors that structure natural communities. Many ecologists have proposed that changes in the intensity of recruitment of individuals into a community should influence community structure. Past work investigating this hypothesis has primarily been surveys of recruitment and community structure across large spatial scales. Surprisingly, no researchers to date have manipulated recruitment into a rocky intertidal community to assess the response of interactions of the whole community to variation in recruitment intensity. We manipulated the densities of Balanus glandula and Chthamalus spp. recruits across a four-fold difference at two sites spanning the Monterey Bay, California, USA, to experimentally test if differences in recruitment intensity influenced initial changes in community composition and if these changes persisted through time. The results of this work indicate that differences in recruitment influence community composition initially, but that these changes can be short-lived.     

Effects of small-scale disturbances of canopy and grazing on intertidal assemblages on the Swedish west coast

The effects of small-scale disturbances (80×30-cm plots) of canopy and grazers on intertidal assemblages were investigated in this 4-year experiment on sheltered rocky shores on the Swedish west coast. Canopy disturbances due to ice scouring were mimicked by removal of adult plants of the seaweed Ascophyllum nodosum (L.) Le Joli. Density of the main epilithic grazing gastropods, Littorina spp., was lowered by exclosure and handpicking. Based on earlier experiments in other areas, the general hypothesis was that canopy removal and grazer exclosure, alone or in combination, should increase the recruitment of A. nodosum or other fucoid juveniles, and change the structure of the understorey assemblage.There was an effect of canopy removal on the development of this assemblage, lasting for more than 31 months. Both increased and decreased abundances of species were found as short-term effects, but there was also a longer-term effect with increased abundance. Grazer exclosure was only effective in combination with canopy removal, causing a short-term increase in ephemeral green algae. Short-term effects of canopy removal were also the increase in recruitment of Semibalanus balanoides (Linnaeus) and the decrease of the red alga Hildenbrandia rubra (Sommerfelt) Meneghini. Fast recruitment and growth of fucoid species (Fucus serratus L. and F. vesiculosus L.) restored the canopy and conditions of the understorey within 18 months. Thus, the canopy removal changed the physical conditions for the understorey, making it possible for other species to coexist in this community. Surprisingly, no effect of canopy removal or grazer exclusion was found on the recruitment of juvenile A. nodosum, neither by canopy removal nor grazer exclosure. The lack of such effects might be due to the early mortality caused by other grazers (small, mobile crustaceans), or to the low density of periwinkles on these shores. However, despite the patchy and generally low recruitment of A. nodosum juveniles, observations suggested that the cover of A. nodosum in manipulated patches would return to initial levels, either by recruitment or regrowth of small holdfasts and from growth of edge plants.     

Interactive effects of food availability and aerial body temperature on the survival of two intertidal Mytilus species

Intertidal organisms must episodically contend with the rigors of both the terrestrial and the marine environments. While body temperatures during high tide are driven primarily by water temperature, aerial body temperatures are driven by multiple environmental factors such that temperature of an organism during low tide is usually quite different from air temperature. Thus, whereas decades of research have investigated the effects of water temperature on intertidal species, considerably less is known about the physiological impacts of temperature during aerial exposure at low tide, especially with regard to the interaction of aerial body temperature with other stressors. We examined the interactive effects of aerial body temperature and food supply on the survival of two intertidal blue mussels, Mytilus galloprovincialis and Mytilus trossulus. Survival was monitored for nine weeks using a simulated tidal cycle, with two levels of food and three levels of aerial body temperature (30, 25, and 20 °C). Decreased food supply significantly reduced the survival of mussels, but only under the 30 °C treatment. In the other two thermal regimes there were no significant effect of food on survival. When aerial body temperatures are high, food availability may have a greater effect on intertidal organisms. Decreases in ocean productivity have been linked to increased in ocean temperatures, thus intertidal organisms may become more susceptible to thermal stress as climates shift.     

Root mass distribution patterns under standardised conditions in species of Chionochloa and Festuca from New Zealand

The vertical biomass allocation patterns of roots grown under standardised conditions were determined for species representing the major New Zealand indigenous grass genera Chionochloa and Festuca. Ten ramets, each of 2–3 tillers from garden collections of each species were grown in irrigated vertical sand columns in a glasshouse, and harvested after 168 days. Chionochloa teretifolia, Chionochloa macra, and Chionochloa crassiusucula, characteristic of alpine environments failed to produce new roots and died. However, most of the Chionochloa taxa (Chionochloa beddiei, Chionochloa pallens, Chionochloa rigida ssp. rigida, Chionochloa rubra ssp. cuprea, Chionochloa vireta), developed extensive new roots that reached the base of the one metre sand column. Roots of Chionochloa cheesemanii and Chionochloa conspicua reached 80–90 cm depth. Two Festuca taxa (Festuca actae, Festuca luciarum) had roots to 1 m depth, and roots of Festuca coxii, Festuca matthewsii ssp. latifundii, Festuca matthewsii ssp. matthewsii, Festuca multinodis, and Festuca novae-zelandiae grew to 70–90 cm depth. The edaphic specialists (Festuca deflexa, Chionochloa spiralis, Chionochloa defracta) were all shallow rooting.Species of Festuca maintained at least 40% of the root mass in the upper 10 cm of the column and most of the Chionochloa taxa had less than 40% of root mass in the upper zone. Genotype level variation in root mass less than 10 cm deep was greater in Chionochloa than in Festuca, and least in the edaphic specialist grasses.     

Sediment on rocky intertidal reefs: Effects on early post-settlement stages of habitat-forming seaweeds

Modification of the coastal environment by human activities often leads to an increase in sedimentation of nearshore waters, with potential impacts on benthic marine assemblages. Here we assess the relationships between the levels of sedimentation, wave exposure and benthic organisms on rocky intertidal platforms around the Kaikoura Peninsula in southern New Zealand. We designed and tested five sediment traps to provide a tool for measuring the relative abundance of sediment across sites. Using field- and laboratory-based experiments, we tested hypotheses concerning whether different levels of sedimentation affected algal germling survival and algal zygote attachment, and whether the interactions of grazers and sediments affected germling survival. Levels of sediment and exposure were inversely related across seven sites. The fucoid alga Hormosira banksii characterized the more sedimented wave-sheltered and intermediately wave-exposed sites, with up to 80% cover in the lower mid-tidal zone, while the bull kelp Durvillaea antarctica characterized the three most wave-exposed sites. Grazing molluscs were found across all sites but species abundances varied by sediment and exposure levels. We did two 11-day trials testing the effects of different levels of sediment and different species of molluscan grazers on the survival of 1-week-old Hormosira germlings. Generally, there was no significant treatment effect of grazers, but mortality varied considerably among sites. In particular, one site had very high levels of sediment, which resulted in 100% mortality of germlings across all grazer treatments. Removing sediment at 1-week, 2-week and 4-week intervals made no difference to the survival of Hormosira germlings. In laboratory-based experiments, a light dusting of sediment reduced the percentage of zygotes of Hormosira by 34% and Durvillaea by 71% that attached to primary substratum, and a complete cover of sediment prevented attachment altogether. Overall, the effects of sediments and its interaction with molluscan grazing were highly variable but often large, particularly on the attachment of zygotes to primary substratum.     

Activity and shelter use of an intertidal snail: effects of sex, reproductive condition and tidal cycle

Although activity patterns are an important component of the ecology of animals as they search for food, mates and appropriate habitats, substantial variation can be observed even within a single population. The source of this variation is the combination of intrinsic and extrinsic factors acting at an individual level. We documented the activity patterns of the ovoviviparous intertidal gastropod Littorina saxatilis (Olivi) over a series of tidal cycles to assess how differences in movements of individuals with different reproductive attributes (non-gravid females, gravid females and males) might explain intra-population variability in activity. Four behaviors were measured for four consecutive tidal cycles for each of 3 months: the magnitude, frequency, and orientation of movements, and the use of shelters (i.e., crevices). In addition, we correlated movements with variations in several environmental factors (temperature, wind, humidity) during low tide to investigate the possible risk sensitivity of the different reproductive groups. All groups were more active and moved greater distances during high tide. Females, especially gravid females, appeared to be more risk-sensitive as they moved shorter distances and used shelters more often. Gravid females were also the only group with a negative relationship between air temperature and movement during low tide. We interpret these behaviors as a means to increase the probability of survival of the brooded offspring carried by the female. Males were the only group to consistently have directionality in their movements and showed evidence of an “up-down” behavior for remaining in a restricted zone. Reproductive attributes appear to be able to explain part of the intra-population variability in foraging activity in natural populations and should be taken into account in the design and interpretation of behavioral and ecological studies.     

Analysis of competitive interactions between the limpets Patella depressa Pennant and Patella vulgata L. on the northern coast of Portugal

Inter- and intraspecific competitive interactions among the species Patella depressa Pennant and P. vulgata L. were analysed on the northern coast of Portugal, where both species co-occur in similar proportions. Increased (×2, ×4), normal and decreased (×1/2 intraspecific only) densities of limpets were used to test the effects of competition on the growth and mortality of the limpets, and competitive interactions between the different species. Fenced plots of 25×25 cm enclosing marked limpets at various densities were set up at a mid tidal level on the shore. Twelve treatments with three replicates of different combinations of densities and species were established. Mortality was recorded every 15 days and length was measured monthly during the 5 months of the experiment. At the end of the experiment, limpets were collected for biometry, sex determination and gonad staging. Both species of grazing molluscs showed increased mortality and reduced size and weight in increased density treatments. Limpets in decreased density treatments showed lower mortality and higher size. There were no significant differences between the effect of P. vulgata on P. depressa and the effect of P. depressa on P. vulgata on mortality and length. There were, however, decreases in weight due to intraspecific effects for both species. For P. vulgata, interspecific effects were much less than intraspecific effects and for P. depressa, the interspecific effects depended on density. The analysis of the gonad stage at the end of the experiment showed that the lower stages of development were most prevalent at increased densities and the occurrence of neuter individuals only in increased densities single species plots corroborated the evidence of a stronger intraspecific interaction. The availability of food was indirectly assessed by determination of chlorophyll concentration with spectrophotometric analysis of rock chips. Microalgal food assessment in this experiment, however, did not show any clear trends. The results were compared with previous studies of competition in grazing molluscs and discussed in relation to the limits of distribution of P. vulgata on the Portuguese coast.     

Phylogeny of the Bangia flora of New Zealand suggests a southern origin for Porphyra and Bangia (Bangiales, Rhodophyta)

Analysis of nuclear small subunit ribosomal RNA gene (18S rDNA) sequence data from 123 samples of the red algal genus Bangia from mainland New Zealand has revealed diversity exceeding that reported for the genus from any other region in the world. Our study resolves two New Zealand Bangia taxa basal to the order Bangiales, and five clades of Bangia, four of which include New Zealand members. The basal taxa are separated from each other by 139 bp and differ from all other Bangia taxa in the New Zealand region by 103-163 bp over approximately 1750 bp 18S rDNA sequence data. Our results reveal a Bangia flora of previously unsuspected richness, and show that the simple morphology of these organisms obscures significant levels of genetic diversity. The presence of high diversity and retention of basal taxa in New Zealand Bangia raises the prospect that the southern hemisphere, and particularly eastern Gondwana, is not only a centre of diversity, but a centre of origin for the modern Bangiales.     

Response of predatory snails to a novel prey following the geographic range expansion of an intertidal barnacle

During a geographic range expansion, the success of a species colonizing a new region may be influenced by how resident predators respond to this novel prey. The volcano barnacle Tetraclita rubescens has recently expanded its range > 300 km northward along the coast of California, USA. In the historic portion of its range, Tetraclita is preyed upon by a southern dogwhelk, Nucella emarginata. In newly colonized northern habitats, Tetraclita has joined an assemblage of mid-intertidal barnacles preyed upon by two species of northern dogwhelks (N. ostrina and N. canaliculata). Using a combination of field enclosures and laboratory experiments, we tested the impacts of northern whelks on volcano barnacles relative to other barnacle species. In the field, juvenile Tetraclita were rarely consumed. In laboratory trials with larger barnacles, Tetraclita were nearly immune to drilling by northern whelks, perhaps due to superior morphological defenses. In all experiments, prior encounters with Tetraclita did not appear to increase the frequency of predation on this barnacle; northern whelk species rarely preyed on Tetraclita even when collected from a site where this barnacle was abundant. In contrast, the southern whelk N. emarginata readily preyed on Tetraclita in laboratory trials, perhaps reflecting a longer shared evolutionary history between these species. Overall, our results suggest that the success of a population established beyond a species' historical range boundary may be influenced by the functional traits of that species in relation to the composition and evolutionary history of the surrounding community.     

First observations of predation by New Zealand Greenshell mussels (Perna canaliculus) on zooplankton

Observations made overseas of predation by blue mussels and zebra mussels on mesozooplankton (>200 μm) have raised concern within New Zealand that the Greenshell mussel, Perna canaliculus, which is cultured in large tonnages throughout hundreds of marine farms within the New Zealand coastal zone, could exert ecologically detrimental effects by preying on zooplankton. We conducted experiments at Clova Bay, Pelorus Sound in May 2002 to determine the rates that P. canaliculus ingests prey, up to and including the mesozooplankton size range. Single mussels from farms were incubated with seawater enriched with zooplankton (>60 μm) in gently circulated 15-l pails. Depletion of chlorophyll-a (chl-a), ciliate microzooplankton, and nauplii, copepodites, and adults of copepods was determined over 5 h, relative to controls with no mussels. Two experiments were made over consecutive days. Gut contents of these experimental mussels, and of mussels examined soon after collection from a farm, were described.Gut contents of experimental and of freshly collected mussels (standard shell length ∼90 mm) had numerous copepod parts, whole copepods and larval bivalves present. Experimental mussels cleared chl-a and ciliates from 59- to 137-l individual⁻¹ day⁻¹, respectively, averaged across the two experiments. Faster ciliate than chl-a clearance was probably caused by the high proportion (56%) of phytoplankton below the retention size for P. canaliculus (ca. 5 μm) and by faster ciliate grazing in controls than treatments. The average clearance rates of adult, copepodite, and naupliar copepod stages by mussels were 20, 31, and 49 l individual⁻¹ day⁻¹, respectively. The clearance rates of each copepod stage were not significantly different between the two experiments. Clearance of nauplii was significantly greater than of adults and copepodites, while adult and copepodite clearance rates were nearly significantly different. The mean lengths of the adult, copepodite, and naupliar copepods were 430, 265, and 165 μm, respectively. The decreasing clearance rates with increasing size and development of prey (from ciliates, through naupliar, copepodite to adult copepods), suggested that prey escape ability, related to body size and/or morphology, affected capture rates. Mussel faecal samples indicated complete digestion of the gut contents. Pseudofaecal samples showed very low rejection rates of mesozooplankton by mussels. The results are considered in context of current biophysical modelling studies of impacts of large mussel farms in New Zealand. Designs of future experiments to improve accuracy of estimates of mesozooplankton clearance rates by P. canaliculus are considered.     

Extinction and recolonization of coastal megafauna following human arrival in New Zealand

Extinctions can dramatically reshape biological communities. As a case in point, ancient mass extinction events apparently facilitated dramatic new evolutionary radiations of surviving lineages. However, scientists have yet to fully understand the consequences of more recent biological upheaval, such as the megafaunal extinctions that occurred globally over the past 50 kyr. New Zealand was the world''s last large landmass to be colonized by humans, and its exceptional archaeological record documents a vast number of vertebrate extinctions in the immediate aftermath of Polynesian arrival approximately AD 1280. This recently colonized archipelago thus presents an outstanding opportunity to test for rapid biological responses to extinction. Here, we use ancient DNA (aDNA) analysis to show that extinction of an endemic sea lion lineage (Phocarctos spp.) apparently facilitated a subsequent northward range expansion of a previously subantarctic-limited lineage. This finding parallels a similar extinction–replacement event in penguins (Megadyptes spp.). In both cases, an endemic mainland clade was completely eliminated soon after human arrival, and then replaced by a genetically divergent clade from the remote subantarctic region, all within the space of a few centuries. These data suggest that ecological and demographic processes can play a role in constraining lineage distributions, even for highly dispersive species, and highlight the potential for dynamic biological responses to extinction.     

Spatial variation in the recruitment of the intertidal barnacles Chthamalus montagui Southward and Chthamalus stellatus (Poli) (Crustacea: Cirripedia) over an European scale

Spatial variation in the recruitment of the intertidal barnacles Chthamalus montagui and Chthamalus stellatus was examined over an European scale. The study was carried out using standardised protocols at a series of locations. The five locations chosen (SW Ireland, NW Spain, SW Portugal and NW and NE Italy) span a large part of the range of these species in Europe. The spatial scales were location (hundreds of kilometres) and shore (thousands of metres).Estimates of total cumulative recruitment (cyprids and metamorphs) summed over the year (April 1997 to March 1998) showed substantial variation between locations which was dependent on the species. Recruitment was highest in SW Portugal for C. montagui and in SW Ireland and NW Spain for C. stellatus. Overall recruitment of C. montagui was higher than that of C. stellatus at all locations except SW Ireland, where recruitment of the two species was not significantly different. There were significant differences among shores in each location.The recruitment period of both species varied with location, with recruitment beginning earlier further south. In general, recruitment of C. montagui and C. stellatus was recorded in 8 months in NW Spain and NE Italy, while only in 7 months in SW Ireland. Recruitment of C. montagui occurred in 10 months in SW Portugal, but no recruits of C. stellatus were found. In all locations there was at least one distinct peak of recruitment. In SW Ireland both species showed only one peak of recruitment, a month after initiation. At the more southerly Atlantic locations, as well as in the Mediterranean, two unequal peaks of recruitment were generally seen. During recruit census, the number of cyprids, in comparison to metamorphs, found at any location was very low. In SW Ireland and NW Spain cyprids of both species were found, while in SW Portugal and in the Mediterranean, only cyprids of C. montagui were found.     

Chemosystematic investigations of irregular diterpenes in Anisotome and related New Zealand Apiaceae

A chemosystematic HPLC-UV and HPLC-MS investigation of New Zealand members of the Apiaceae was performed. Diterpenes were identified and quantified in methanolic extracts from subaerial parts of 28 taxa and 54 samples of Aciphylla, Anisotome, Apium, Gingidia, Lignocarpa, Oreomyrrhis, and Scandia. Six diterpenes (1-2, 4-7) and four polyacetylenes (8-11) were identified. The known compounds were the diterpenes anisotomenoic acid 1, anisotomene-1-ol 2, 16-acetoxyanisotomenoic acid 4 and anisotomene-1,12-diol 5; and the polyacetylenes falcarinol 8, falcarindiol 9, (+)-9(Z),17-octadecadiene-12,14-diyne-1,11,16-triol 10, and (+)-9(Z),17-octadecadiene-12,14-diyne-1,11,16-triol 1-acetate 11. New irregular diterpenes 13,14-dihydroanisotom-12E-ene-1,14-diol 6 and 14-methoxy-13,14-dihydroanisotom-12E-ene-1-ol 7 were isolated from A. haastii. Isomers of the new semi-synthetic diterpene 16-hydroxyanisotomenoic acid 3 were detected in extracts of Anisitone flexuosa. Structure elucidation was performed by HR mass spectrometry and 1D and 2D NMR spectroscopy. In crude extracts, compounds were identified by their HPLC retention times and their on-line HPLC-UV and MS spectra. Anisotomene diterpenes occurred in eight out of 16 species of the genus Anisotome, but were not detected in any of the other genera. In contrast, polyacetylenes were present in all the genera investigated.     

Effects of seston variability on the clearance rate and absorption efficiency of the mussels Aulacomya maoriana, Mytilus galloprovincialis and Perna canaliculus from New Zealand

The clearance rates (CR in l g⁻¹ h⁻¹) and net absorption efficiencies (AE) of three co-occurring mytilid species were estimated to determine the effects of variation in ambient seston quantity (total particulate matter, TPM, mg l⁻¹) and quality (particulate organic matter, POM, mg l⁻¹; percent organic matter, PCOM=POM/TPM) on these physiological functions. CR_S estimates were significantly different among the species (Mytilus galloprovincialis>Perna canaliculus=Aulacomya maoriana), but were not correlated with differences in species-dependent mean body size (P. canaliculus>M. galloprovincialis>A. maoriana). AE estimates were independent of mean body weight, and did not differ among the species. For all three species, CR responded in a simple linear manner to seston organic content, either in terms of POM (for A. maoriana and M. galloprovincialis), or PCOM (for P. canaliculus). The AE response, although also of a simple linear type, was species-dependent and involved interactions of two or three seston components: TPM, POM and PCOM for A. maoriana; POM and PCOM for M. galloprovincialis; and TPM and PCOM for P. canaliculus. For all three species, seston variation explained 15-20% of the variation in CR and 52-59% of the variation in AE. Comparisons among the species indicated that two very different responses to variation in seston quantity and quality exist. First, significant differences in CR result from species-dependent differences in the magnitude of the response to seston variation. Second, species-dependent responses to variation in seston variation resulted in the similarity of the AE responses. Thus, the three species appear to have evolved different strategies for dealing with seston variation, but with the end result that their AE responses do not differ. Finally, no evidence was found of a negative association between CR and AE for any of the three species, suggesting that under the seston conditions experienced in this work, these species do not have the physiological compensatory capacity to reduce CR in order to increase AE. The importance of a comparative (i.e., multi-species) approach utilising ambient seston is emphasised by the findings of this research if we are to understand better the feeding and digestive physiologies of suspension-feeding organisms such as mussels.     

Bone growth marks reveal protracted growth in New Zealand kiwi (Aves,Apterygidae)

The presence of bone growth marks reflecting annual rhythms in the cortical bone of non-avian tetrapods is now established as a general phenomenon. In contrast, ornithurines (the theropod group including modern birds and their closest relatives) usually grow rapidly in less than a year, such that no annual rhythms are expressed in bone cortices, except scarce growth marks restricted to the outer cortical layer. So far, cyclical growth in modern birds has been restricted to the Eocene Diatryma, the extant parrot Amazona amazonica and the extinct New Zealand (NZ) moa (Dinornithidae). Here we show the presence of lines of arrested growth in the long bones of the living NZ kiwi (Apteryx spp., Apterygidae). Kiwis take 5–6 years to reach full adult body size, which indicates a delayed maturity and a slow reproductive cycle. Protracted growth probably evolved convergently in moa and kiwi sometime since the Middle Miocene, owing to the severe climatic cooling in the southwest Pacific and the absence of mammalian predators.     

Growth and reproduction ofPorphyra columbina Mont. (Bangiales,Rhodophyceae) from southern New Zealand

Changes in biomass and chemical composition, and the reproductive phenology ofPorphyra columbina Mont. were monitored at three sites in southern New Zealand over two growing seasons. Both temporal and spatial variations were found. Seasonal changes in biomass and chemical components were correlated with seawater nitrate concentrations and temperature. The summer decline in biomass was a result of the onset of unsuitable environmental conditions and the release of reproductive tissue. Under more suitable conditions, the decline in biomass was delayed. There was an inverse relationship between vegetative growth and reproduction. Reproductive plants first appeared in August at a time of increasing temperature, irradiance and daylength. Only larger plants which were mainly found in subsites low on the shore became reproductive. Plants sampled from high subsites had a shorter growth season, were generally smaller, had lower nitrogen and pigment content and were non-reproductive.Presented at the XIIIth International Seaweed Symposium, University of British Columbia, Vancouver, Canada, August 1989.