Oviposition,larval abundance,in situ larval growth and recruitment of the herbivorous gastropodLacuna vincta in kelp canopies in Barkley Sound,Vancouver Island (British Columbia)

We investigated recruitment of the herbivorous gastropodLacuna vincta (Montagu, 1803) in the canopies ofMacrocystis integrifolia andNereocystis luetkeana beds in Barkley Sound, Vancouver Island (British Colombia), from 1987 to 1989. Four factors influencing intensity and patterns of recruitment were studied: (1) seasonality of oviposition, (2) larval abundance, (3) growth of larvae in the field and (4) larval settlement. Egg masses were abundant on low intertidal algae but were scarce in kelp canopies. Although egg masses could be found almost year-round, a distinct and intense period of oviposition occurred during winter and spring. Intracapsular development lasted 2.5 to 3.5 wk before planktotrophic veligers emerged. The duration of the planktonic period, 7 to 9 wk, was determined through an in situ study of cohorts ofLacuna spp. larvae present in the plankton between January and June 1988. The general timing of the onset of the spring peak recruitment period was predicted from these cohorts. Primary periods of recruitment ofL. vincta in the canopy occurred in April–May (average density up to 383.9 juveniles m^–2 blades), with a second period of lower intensity in the late summer—fall period. We observed similar trends between abundance of advanced larvae (> 500µm) in the plankton and recruitment rates in kelp canopies. Although adults were occasionally observed in the canopy, newly metamorphosed juveniles consistently dominated the habitat. The persistance of small juveniles (0.7 to 1.5 mm), rapid declines in density shortly after recruitment, and SCUBA observations of drifting individuals suggest that juveniles migrate to the under-canopy or low intertidal area after a brief period of growth on kelp blades.     

UV-radiation versus grazing pressure: long-term floating of kelp rafts (<Emphasis Type="Italic">Macrocystis pyrifera</Emphasis>) is facilitated by efficient photoacclimation but undermined by grazing losses

Large quantities of floating macroalgae are traveling in coastal waters of the SE Pacific and in other temperate climate zones. While afloat, these algae are potentially exposed to full solar radiation, including UVA and UVB, which can have profound effects on their physiological and growth performance. Latitudinal variations in UV-radiation (UVR) are hypothesized to affect floating algae differently with higher impacts at low latitudes than at high latitudes. In addition, UVR together with grazing might accelerate the demise of floating kelps. This hypothesis was tested with outdoor laboratory experiments in which sporophytes of the giant kelp Macrocystis pyrifera (L.) C. Agardh were exposed to a combination of different UVR regimes (PAR only, PAR + UV) and grazing at three sites along the Chilean coast (20°S, 30°S, and 40°S). A latitudinal trend in irradiance was detected with increasing values from 40°S to 20°S. Surprisingly, floating M. pyrifera responded with a high acclimation potential within this latitudinal UVR gradient. At 20°S, floating kelps were slightly sensitive to UVR, which was reflected in reduced blade growth. At 30°S, physiological responses were hardly affected by the prevailing irradiance but sporophyte growth and thus persistence mainly depended on the presence or absence of amphipod grazers. At high latitudes, grazing had only minor impacts on algal biomass and blade growth, and kelps thrived well under all tested environmental conditions. Overall, our results reveal that floating M. pyrifera was only slightly affected by UVR and that sporophytes can efficiently acclimate over a latitudinal UVR gradient that spans from 20°S to 40°S. Given this high acclimation potential, we suggest that these (and possibly other) positively buoyant algae are important dispersal agents over a wide range of temperate latitude conditions.     

Effects of petroleum products on bull kelp (Nereocystis luetkeana)

Although research has been conducted on the effects of oil on the giant kelp Macrocystis pyrifera, no similar studies have been completed on bull kelp, Nereocystis leutkeana, the dominant kelp in Washington State, British Columbia, and Alaska. The effects of three petroleum products [diesel fuel, intermediate fuel oil (IFO), and crude oil] were tested before and after weathering on N. luetkeana. Whole plants were exposed to petroleum product for 4 or 24 h and then transferred to the field; observations on the condition of the plants were made daily for 7 d. In addition, controlled bioassays were performed to measure the effects of petroleum exposure on net photosynthetic rate (NP) and respiration rate (R), using light-and dark-bottle techniques. These experiments verified the susceptibility of N. luetkeana tissue to the damaging effects of direct exposure to several oil types. The 4 h exposures to weathered diesel and unweathered IFO, and 24 h exposures to unweathered and weathered diesel and IFO resulted in moderate to severe damage to kelp tissue (i.e., clearly delineated bleached line accompanied by tissue necrosis). Weathered diesel was more toxic than unweathered diesel. The most severe damage to bull kelp was concentrated at the meristematic zone (junction of stipe and bulb) where new tissue growth occurs. Petroleum type significantly affected stipe and blade NP, R, and NP:R ratios. Diesel treatments had a greater negative effect on NP than did the IFO treatments. Based on these experiments, the relative ranking of the damaging effects of petroleum treatment on bull kelp are weathered diesel>unweathered IFO>unweathered diesel>weathered IFO>unweathered crude>weathered crude.     

Latitudinal symbiont zonation in <Emphasis Type="Italic">Stylophora pistillata</Emphasis> from southeast Africa

Studies on latitudinal gradients in Symbiodinium diversity on scleractinian corals are largely restricted to warm-water low latitude locations, and it appears that there is a shift in symbiont distributions with increasing latitude. The Symbiodinium assemblages of high latitude coral communities have largely been undocumented despite occupying an important transitional zone between tropical and temperate regions. Using a combination of the internal transcribed spacer region 2 (ITS2) and denaturing gradient gel electrophoresis (DGGE), we assessed the cladal and subcladal variability of Symbiodinium in the widely distributed species Stylophora pistillata along a latitudinal transect in southeast African waters which extended into high latitude locations. All colonies examined belonged to clade C. Six unique ITS2-DGGE banding profiles (designated C_spa to C_spf) were observed, which showed a latitudinal distribution from north to south, most likely a result of a gradient in water temperature and irradiance driven by riverine input in the southern regions. Sequence analysis revealed that all sequences except one did not match previously identified clade C sub-types, probably due to the lack of regional information in the Western Indian Ocean when compared to the Caribbean and Pacific. This study further supports the applicability of ITS2-DGGE in studies on Symbiodinium diversity, and highlights that potentially ecologically informative biogeographic patterns may be overlooked when only cladal designations are employed.     

Combined sclerochronologic and oxygen isotope analysis of gastropod shells (Gibbula cineraria, North Sea): life-history traits and utility as a high-resolution environmental archive for kelp forests

The grey top-shell, Gibbula cineraria is a common member of temperate to cold water kelp forest communities, but its longevity and the age structure of its populations remains unresolved. Combined measurements of shell growth rates (sclerochronology) and oxygen isotope composition allow analysis of rate and timing of shell growth. Eight specimens were analyzed from the southern North Sea (near Helgoland, German Bight). Three age groups were identified but external measurements (width, height, ornamentation patterns and number of whorls) and shell weight are not adequate for ontogenetic age discrimination. Stable oxygen isotope data is consistent with shell growth during the interval from April to December in isotopic equilibrium with seawater, and growth increments exhibit strong tidal controls with fortnightly bundles well preserved. Reliable environmental proxy data (water temperature) can be extracted from the shell aragonite using conventional stable oxygen isotope analyses, with a temporal resolution of days attainable during intervals of maximum growth, but annual extremes are not always recorded in the shell. While demonstrating the utility of G. cineraria as a environmental and potential paleoenvironmental proxy for kelp forest habitats, its longevity has been significantly overestimated.     

Biomass rather than growth rate determines variation in net primary production by giant kelp

Net primary production (NPP) is influenced by disturbance-driven fluctuations in foliar standing crop (FSC) and resource-driven fluctuations in rates of recruitment and growth, yet most studies of NPP have focused primarily on factors influencing growth. We quantified NPP, FSC, recruitment, and growth rate for the giant kelp, Macrocystis pyrifera, at three kelp forests in southern California, U.S.A., over a 54-month period and determined the relative roles of FSC, recruitment, and growth rate in contributing to variation in annual NPP. Net primary production averaged between 0.42 and 2.38 kg dry mass x m(-2) x yr(-1) at the three sites. The initial FSC present at the beginning of the growth year and the recruitment of new plants during the year explained 63% and 21% of the interannual variation observed in NPP, respectively. The previous year's NPP and disturbance from waves collectively accounted for 80% of the interannual variation in initial FSC. No correlation was found between annual growth rate (i.e., the amount of new kelp mass produced per unit of existing kelp mass) and annual NPP (i.e., the amount of new kelp mass produced per unit area of ocean bottom), largely because annual growth rate was consistent compared to initial FSC and recruitment, which fluctuated greatly among years and sites. Although growth rate was a poor predictor of variation in annual NPP, it was principally responsible for the high mean values observed for NPP by Macrocystis. These high mean values reflected rapid growth (average of approximately 2% per day) of a relatively small standing crop (maximum annual mean = 444 g dry mass/m2) that replaced itself approximately seven times per year. Disturbance-driven variability in FSC may be generally important in explaining variation in NPP, yet it is rarely examined because cycles of disturbance and recovery occur over timescales of decades or more in many systems. Considerable insight into how variation in FSC drives variation in NPP may be gained by studying systems such as giant kelp forests that are characterized by frequent disturbance and rapid rates of growth and recruitment.     

Potential latitudinal variation in egg size and number of a geographically widespread reef fish,revealed by common-environment experiments

Variation in maternal reproductive traits was examined in field and reared populations of a geographically widespread reef fish, Pomacentrus coelestis (Pomacentridae), drawn from three different latitudes in Japan. Size-specific clutch size and clutch weight of wild fish increased with increasing latitude. Conversely, latitudinal variation in egg size of wild fish was obscure in same-season comparisons, probably because of the temperature effect on egg size. Common-environment experiments conducted at three temperatures showed that egg size decreased with increasing temperature in all populations. In the experiments, egg size, clutch size and clutch weight differed among populations at all temperatures, showing clear latitudinal clines. Females from low latitude spawned larger eggs at every experimental temperature. Size-specific clutch size and weight were greater in females from high latitude. Thus, the northern fish had a larger reproductive output per spawning and a larger number of smaller eggs in a spawning. Such interpopualtion variation in this fish is likely to be partially genetically based, although environmental effects on the variation cannot be entirely ruled out. This study provides evidence of potential latitudinal variation in the egg size and number in a coastal fish, by common-environment experiments. The close correspondence between latitudes and these maternal reproductive traits may be a consequence of local adaptation.     

Geographical patterns in reproductive biology of the Pan-American sandy beach isopod<Emphasis Type="Italic"> Excirolana braziliensis</Emphasis>

Eleven populations of the Pan-American sandy beach isopod Excirolana braziliensis, distributed from tropical (9°N) to temperate (39°S) sandy beaches in Atlantic and Pacific Oceans, were analyzed to determine latitudinal variations in: breeding and recruitment patterns, sex ratios, size of ovigerous females and of juveniles and length–fecundity relationships. E. braziliensis exhibited strong latitudinal patterns in all reproductive traits throughout its distribution range. Breeding and recruitment shifted from continuous to seasonal from tropical to temperate beaches, having a predominance of females at higher latitudes. In agreement with the latitudinal gradient hypothesis, ovigerous females inhabiting tropical sandy beaches (low latitudes) were smaller, became sexually mature at smaller sizes and had lower individual fecundity than on temperate beaches. Juveniles were also smallest at low latitudes. Between-ocean comparisons showed very similar reproductive characteristics for roughly the same latitude. These linked reproductive parameters suggest that the intensity of breeding effort is associated with the duration of the breeding season and geographically size-related characteristics of the species. Geographic variations in the breeding and recruitment seasons, as well as in individual fecundity, size structure of mature females and sex ratios, are proposed to have major consequences in explaining local variations in population demography. Our paper also reinforces the notion that sandy beach animals are highly plastic.Communicated by O. Kinne, Oldendorf/Luhe     

Growth of the kelp Ecklonia radiata near the northern limit of its range in Western Australia

The length, growth and survivorship of mature sporophytes of Ecklonia radiata (C.Ag.) J. Agardh were measured on two patch reefs within the lagoon of a high-latitude, coralreef atoll (Easter Group, Houtman Abrolhos; 28° to 29° S; 113°35 to 114°03E) for 1 yr (1982–1983). The sites differed in their proximity to the perimeter reef, but had similar regimes of temperature, light, nutrient concentration and water movement. Kelp length, growth rate and survivorship differed significantly between sites, although plant density was similar (less than l m^-2). At the site near the lagoon perimeter, the central lamina averaged 381 mm in length, grew at an annual mean rate of 1.28 mm d^-1, and 56% of tagged individuals had died after one year. The corresponding values for the site near the centre of the lagoon were: 257 mm, 0.75 mm d^-1 and 92%. Three other sites on the coastal limestone reefs near Perth (400 km to the south) were studied for various intervals over a 5 yr period (April 1979 to August 1984). The kelp from the low-latitude sites were smaller, grew more slowly, and suffered similar or greater mortality than their southern counterparts. Latitudinal comparisons were confounded by differences in kelp density between sites, but there is no evidence for density-dependent effects on the measured parameters across the range of natural densities observed. Growth rates at all sites were negatively correlated with ambient sea temperatures when these were above 20°C. There was no evidence of adaptation to the higher sea temperatures experienced at the Abrolhos, and temperature cannot be dismissed as a factor controlling the growth of the species near the northern limit of its distribution on the Western Australian coast. Other factors however must be involved in determining the latitudinal position of that limit.     

Inversion of plant dominance-diversity relationships along a latitudinal stress gradient

Species interactions affect plant diversity through the net effects of competition and facilitation, with the latter more prevalent in physically stressful environments when plant cover ameliorates abiotic stress. One explanation for species loss in invader-dominated systems is a shift in the competition-facilitation balance, with competition intensifying in areas formerly structured by facilitation. We test this possibility with a 10-site prairie meta-experiment along a 500-km latitudinal stress gradient, quantifying the relationships among abiotic stress, exotic dominance, and native plant recruitment over five years. The latitudinal gradient is inversely correlated with abiotic stress, with lower latitudes more moisture- and nutrient-limited. We observed strong negative effects by invasive dominant grasses on plant establishment, but only in northern sites with lower-stress environments. At these locations, disturbance was critical for recruitment by reducing the suppressive dominant (invasive) canopy. In more stressful environments to the south, the impacts of the dominant invaders on plant establishment became facilitative, and diversity was more limited by seed availability. Disturbance prevented recruitment because seedling survival depended on a protective plant canopy, presumably because the canopy reduced temperature or moisture stress. Seed limitation was similarly prevalent in all sites. Our work confirms the importance of facilitation as an organizing process for plants in higher-stress environments, even with transformations of species composition and dominance. It also demonstrates that the mechanisms regulating diversity, including invader impacts, can vary within the same plant community depending on environmental context. Because limits on native plant recruitment are environmentally contingent, management strategies that seek to increase diversity, including invader eradication, must account for site-level variations in the balance between biotic and abiotic constraints.     

Changes in sea urchin populations after the destruction of kelp beds

In St. Margaret's Bay, Nova Scotia, Canada, there are large areas in which sea urchins (Strongylocentrotus droebachiensis) have eliminated beds of kelp (Laminaria spp.). Sites were identified where destruction of kelp beds had taken place 1, 2, 3, 3.5 and 4 years ago. With increase of time since kelp disappearance, the sea urchins showed decreased growth rate, reduced gonad size, but an increase in numbers resulting from high recruitment rates in the first two years after kelp bed destruction. These sea urchin populations, by their browsing, effectively prevent the regeneration of kelp. There is, as yet, no evidence of the sea urchin populations being starved out to allow the kelp to return.     

Colonisation of kelp imitations by epiphyte and holdfast fauna; a study of mobility patterns

The kelp Laminaria hyperborea provides a habitat for a diverse invertebrate community. Invertebrate abundance is particularly high in the kelp holdfasts and associated with epiphytes on the stipes. We investigated colonisation patterns of this fauna on artificial substrata (holdfast and epiphyte mimics) exposed for short time periods in the kelp forest in two seasons, summer and late autumn. Within 7 days, 99 taxa common on natural kelp substratum colonised the artificial substrata in large numbers, demonstrating high mobility and indicating that there may be a rapid exchange of individuals and species among algae. Gastropods and amphipods were the most abundant colonisers. In summer, a specific “epiphyte fauna” could be distinguished from a “holdfast fauna”, and species associated with epiphytes were significantly more abundant on the artificial substrata. Colonisation was more rapid in summer than in late autumn, which may be related to animal population density. The high dispersal rate can facilitate the interaction with other communities and with higher trophic levels in the kelp-forest food web. Published online: 13 August 2002     

Photosynthetic characteristics of giant kelp (Macrocystis pyrifera) determined in situ

Rates of net photosynthesis and nocturnal respiration by individual blades of the giant kelp Macrocystis pyrifera (L.) C. Agardh in southern California, were determined in situ by measuring oxygen production in polyethylene bags during spring/summer of 1983. Mature blades from different depths in the water column exhibited different photosynthetic characteristics. Blades from the surface canopy (0 to 1 m depth) exhibited higher photosynthetic capacity under saturating irradiance and higher photosynthetic efficiency at low irradiances than blades from 3 to 5 or 7 to 9 m depths. Saturating irradiance was lower for canopy blades than for deeper blades. Canopy blades showed no short-term photoinhibition, but photosynthetic rates of deeper blades were significantly reduced during 1 to 2 h incubations at high irradiances. Results of 1 to 2 wk acclimation experiments indicated that differences between photosynthetic characteristics of blades from different depths were primarily attributable to acclimation light conditions. Vertical displacement of blades within the kelp canopy occurred on a time-scale of 1 min to 1 h. Blades continually moved between the unshaded surface layer and deeper, shaded layers. Vertical movement did not maximize photosynthesis by individual blades; only a small proportion of blades making up a dense surface canopy maintained light-saturated photosynthetic rates during midday incubations. The relatively high photosynthetic rates exhibited by canopy blades over the entire range of light conditions probably resulted from acclimation to intermittent high and low irradiances, a consequence of vertical displacement. Vertical displacement also reduced the afternoon depression in photosynthesis of individual canopy blades. The overall effect of vertical displacement was optimization of total net photosynthesis by the kelp canopy and, therefore, optimization of whole-plant production.     

Antipredator defenses along a latitudinal gradient in Rana temporaria

Antipredator defenses are expected to decrease toward higher latitudes because predation rates are predicted to decrease with latitude. However, latitudinal variation in predator avoidance and defense mechanisms has seldom been studied. We studied tadpole antipredator defenses in seven Rana temporaria populations collected along a 1500-km latitudinal gradient across Sweden, along which previous studies have found increasing tadpole growth and development rates. In a laboratory common garden experiment, we measured behavioral and morphological defenses by raising tadpoles in the presence and absence of a predator (Aeshna dragonfly larva) in two temperature treatments. We also estimated tadpole survival in the presence of free-ranging predators and compared predator densities between R. temporaria breeding ponds situated at low and high latitudes. Activity and foraging were generally positively correlated with latitude in the common garden experiment. While all populations responded to predator presence by decreasing activity and foraging, high-latitude populations maintained higher activity levels in the presence of the predator. All populations exhibited defensive morphology in body and tail shape. However, whereas tail depth tended to increase with latitude in the presence of predator, it did not change with latitude in the absence of the predator. Predator presence generally increased larval period and decreased growth rate. In the southern populations, predator presence tended to have a negative effect on metamorphic size, whereas in the northern populations predators had little or a positive effect on size. Latitude of origin had a strong effect on survival in the presence of a free-ranging predator, with high-latitude tadpoles experiencing higher mortality than those from the low latitudes. In the wild, predator densities were significantly lower in high-latitude than in mid-latitude breeding ponds. Although the higher activity level in the northern populations seems to confer a significant survival disadvantage under predation risk, it is probably needed to maintain the high growth and development rates. However, the occurrence of R. temporaria at high latitudes may be facilitated by the lower predator densities in the north.     

Effect of solar ultra-violet radiation on the kelp Ecklonia radiata

During spring and summer, 1982–1986, experiments were carried out near Marmion Reef, Western Australia. In summer, nearly 30% of the surface solar ultraviolet radiation (280 to 400 nm) penetrates offshore waters to 5 m depth. Experimental removal of the mature Ecklonia radiata kelp canopy in summer results in tissue damage, photopigment destruction, reduced growth, and low survivorship of subcanopy kelp sporophytes. These effects do not occur with canopy removal in winter. Laboratory experiments revealed that the UV component of radiation, rather than intense photosynthetically active radiation, was responsible for the inhibition of growth and photodamage. UV radiation probably affects survival of the settlement stages of E. radiata sporophytes, thus excluding them from otherwise suitable substrata in shallow waters. UV radiation is implicated in the reduction of canopy productivity in summer.     

Growth and utilization of internal nitrogen reserves by the giant kelp Macrocystis pyrifera in a low-nitrogen environment

An adult giant kelp plant (Macrocystis pyrifera), moved from an inshore kelp forest to an offshore, low-nitrogen environment near Santa Catalina Island, California (USA), maintained growth for 2 wk on internal nitrogen reserves. Frond elongation rates decreased significantly during the third week, and plant growth rate (wet wt) dropped from an initial inshore rate of 3.6 to 0.9% d^-1. During this 3 wk period, nitrogen contents and free amino acid concentrations decreased, while mannitol and dry contents increased in frond tissues. After depletion of internal nitrogen reserves, the nitrogen content of lamina and stipe tissues averaged 1.1 and 0.7% dry wt, respectively. The experimental plant was exposed to higher ambient nitrogen concentrations during the last 2 wk. Rates of frond elongation and plant growth increased, but nitrogen content and amino acids in frond tissues remained low. Of the total nitrogen contained in frond tissue of the plant before transplantation, 58% was used to support growth in the absence of significant external nitrogen supply. Amino acids constituted a small proportion of these internal nitrogen reserves. Net movement of nitrogen occurred within large fronds, but not between different frond size classes. The nitrogen content of holdfast tissue remained relatively constant at 2.4% dry wt and accounted for 18 to 29% of the total nitrogen. Holdfast nitrogen was not used to support growth of nitrogen-depleted fronds. In comparison to Laminaria longicruris, which is adapted to long seasonal periods of low nitrogen availability, M. pyrifera has small nitrogen-storage capacity. However, internal reserves of M. pyrifera appear adequate to make nitrogen starvation uncommon in southern California kelp forests.     

Effects of Selective Logging on Bat Communities in the Southeastern Amazon

Abstract:  Although extensive areas of tropical forest are selectively logged each year, the responses of bat communities to this form of disturbance have rarely been examined. Our objectives were to (1) compare bat abundance, species composition, and feeding guild structure between unlogged and low-intensity selectively logged (1–4 logged stems/ha) sampling grids in the southeastern Amazon and (2) examine correlations between logging-induced changes in bat communities and forest structure. We captured bats in understory and canopy mist nets set in five 1-ha study grids in both logged and unlogged forest. We captured 996 individuals, representing 5 families, 32 genera, and 49 species. Abundances of nectarivorous and frugivorous taxa (Glossophaginae, Lonchophyllinae, Stenodermatinae, and Carolliinae) were higher at logged sites, where canopy openness and understory foliage density were greatest. In contrast, insectivorous and omnivorous species (Emballonuridae, Mormoopidae, Phyllostominae, and Vespertilionidae) were more abundant in unlogged sites, where canopy foliage density and variability in the understory stratum were greatest. Multivariate analyses indicated that understory bat species composition differed strongly between logged and unlogged sites but provided little evidence of logging effects for the canopy fauna. Different responses among feeding guilds and taxonomic groups appeared to be related to foraging and echolocation strategies and to changes in canopy cover and understory foliage densities. Our results suggest that even low-intensity logging modifies habitat structure, leading to changes in bat species composition.     

Seasonal variation in movement,aggregation and destructive grazing of the green sea urchin (<Emphasis Type="Italic">Strongylocentrotus droebachiensis</Emphasis>) in relation to wave action and sea temperature

Hydrodynamic forces are an important determinant of subtidal community structure, particularly when they limit the distribution and foraging ability of mobile consumers. We examined the effect of wave action on the rate of movement and destructive grazing of a kelp bed by the green sea urchin (Strongylocentrotus droebachiensis) under field conditions. We measured density and rate of advance at fixed intervals along ∼100 m of a grazing front over 1 year, and quantified individual movement rates in the barrens 5–10 m behind the urchin front using a time-lapse videography. Seasonal variation in the mean rate of advance of the front (range: 0–4 m month⁻¹) was explained by changes in urchin density at the front (120–360 individuals m⁻²), which in turn varied inversely with significant wave height (0.5–2 m). Water temperature (0.8–17.6°C) had no effect on the rate of advance or on urchin density (aggregation) at the front, except when temperature exceeded 17°C. Movement of individual urchins also was affected by wave action: we observed a significant decrease in speed and displacement of urchins with increasing significant wave height. Wave action had no effect on the proportion of urchins moving or the degree of linearity of their movements. We propose that the decrease in urchin density at the front associated with increased wave action, results from de-aggregation, which reduces the risk of dislodgement, combined with a reduction in urchin movement in barrens, which supplies new urchins to the front.     

The role of grazing by the lysianassid amphipod<Emphasis Type="Italic"> Orchomenella aahu</Emphasis> in dieback of the kelp<Emphasis Type="Italic"> Ecklonia radiata</Emphasis> in north-eastern New Zealand

The lysianassid amphipod Orchomenella aahu was associated with small-scale mass mortality of Ecklonia radiata, the dominant laminarian alga in north-eastern New Zealand. O. aahu burrowed into and hollowed out stipes with severe bleaching, accelerating mortality by 12–14 months. Meristoderm tissue of bleached plants contained approximately one-third of the phlorotannin content (~4% dry mass) present in unbleached, apparently healthier, sporophytes (~12% dry mass). Unbleached stages consistently lacked amphipod damage. O. aahu also colonised plants when non-lethal storm damage afforded entry into the cortical layers of the primary lamina, and ultimately the stipe. O. aahu typically consumed both medullary and cortical tissue of the stipes, forming networked brood-chambers within them, with increasing populations often reducing the plant to the holdfast. There was synchronised mortality across sites and between depths over spatial scales of metres. Following loss of the mature canopy, recruitment of E. radiata was generally within 2 months at most study sites where mortality had occurred.Communicated by G.F. Humphrey, Sydney