Predation and intertidal zonation of barnacles at Leigh,New Zealand

Experiments using cages at four shore levels were carried out to determine the part played by the feeding of the gastropods Neothais scalaris and Lepsiella scobina on the zonation of three species of intertidal barnacles: Chamaesipho brunnea, C. columna, and Epopella plicata. The lower limit of the two larger barnacle species was determined by predation, but C. columna was affected only in the absence of the other two barnacles.     

Culture and morphology of some New Zealand barnacles (Crustacea: Cirripedia)

The larval stages of the acorn barnacles Chamaesipho columna, Chamaesipho brunnea, Elminius plicatus, Elminius modestus, Balanus trigonus, and Tetraclita purpurascens are described from larvae reared in the laboratory. A key and a table are included for separating larval stages and species.

Skeletonema costatum appears to have a wide application as a food in the rearing of larvae of the Balanidae.

The two Chamaesipho species show typical chthamalid characters of hispid antennal setae and unilobed labrum. The similarity between the larvae of Elminius plicatus and Tetraclita purpurascens supports the classification of E. plicatus as a tetraclitid.     

Recent trends in numbers of four species of penguins at the Prince Edward Islands

Four species of penguin breed regularly at South Africa's Prince Edward Islands: king penguin Aptenodytes patagonicus, gentoo penguin Pygoscelis papua, macaroni penguin Eudyptes chrysolophus and southern rockhopper penguin E. chrysocome. In December 2008, it was estimated that some 65 000 pairs of king penguins were incubating eggs at Marion Island, the larger of the two islands in the group, and 2 000 pairs at Prince Edward Island. At Marion Island from 1987 to 2008, there was no long-term trend in numbers of king penguin chicks that survived to the end of the winter period, but there was considerable fluctuation in chick production in the 1990s. It was roughly estimated that on average 88% of king penguin chicks survived the winter period (from April to September/October). Numbers of gentoo penguins at Marion Island decreased from more than 1 300 pairs in the mid-1990s to fewer than 800 pairs in 2003, and then increased to almost 1 100 pairs in 2008 as breeding success improved. Between 1994/1995 and 2008/2009, numbers of macaroni and southern rockhopper penguins at Marion Island decreased by about 30% and 70% respectively. In 2008/2009, some 290 000 pairs of macaroni penguins bred at this island, mostly in two large colonies where there was a progressive decrease in the density of nests. At both these colonies, decreases in numbers breeding followed outbreaks of disease. Inadequate breeding success has influenced the decreases of macaroni and rockhopper penguins. In 2008/2009, some 42 000 pairs of southern rockhopper penguins bred at Marion Island and 12 000 pairs of macaroni penguins and 38 000 pairs of southern rockhopper penguins at Prince Edward Island.     

Genetic connectivity among populations of two congeneric direct-developing whelks varies across spatial scales

We examined the phylogeographic structure of two congeneric, direct-developing, intertidal whelks endemic to New Zealand, Cominella virgata and C. maculosa. A total of 855 specimens were collected from 23 rocky shore locations and we used them to determine the level of DNA sequence variation at the mitochondrial DNA (mtDNA) cytochrome c oxidase subunit I gene. Cominella virgata and C. maculosa have relatively high levels of haplotype diversity but low levels of nucleotide diversity. Sample sites formed regional groups that were composed of genetically differentiated populations, a finding consistent with their limited dispersal abilities. Cominella virgata showed higher levels of regional differentiation than C. maculosa, and there were also species-specific differences in genetic patterns within some regions. Both species exhibited genetic homogenisation among proximate sites within groups, suggesting some dispersal does occur between areas that are not connected by contiguous rocky shore habitat.     

Factors in the distribution of intertidal,estuarine polychaetes: a field experiment with Nereis (Hediste) diversicolor and Nephtys hombergi in the Tamar at Plymouth

A field experiment was conducted over 10 weeks from July to mid-September, 1984, on a mudflat in the lower reaches of the Tamar Estuary at Plymouth, to test the relative importance of environmental factors and species interactions in controlling the distribution of two polychaete species. Nereis (Hediste) diversicolor from an upper shore site were transplanted into a lower shore site nearby which was characterized by the presence of Nephtys hombergi. 65–71% of N. diversicolor which were transplanted in their own sediments, with no contact with N. hombergi, survived the experimental period. The appearance of some small stages of N. diversicolor in these treatments was attributed in part to recruitment from the overlying water. N. diversicolor transplanted directly into N. hombergi-sediment survived in inverse proportion to their numbers and only the lowest densities (equivalent to 480 m⁻²) survived as well as those in their own sediment. Given the choice to invade the other species' sediment, more N. hombergi than N. diversicolor finally made the change, and there was some evidence that the smallest N. diversicolor suffered predation in consequence. Otherwise there was little evidence for inter-species interaction. There appeared to be some disruption of oocyte maturation of the N. diversicolor which may have been due to changes in tidal rhythm and salinity at the experimental site.     

Trophic relationships on a fucoid shore in south-western Iceland as revealed by stable isotope analyses, laboratory experiments, field observations and gut analyses

Rocky shores in the North Atlantic are known for their zonation patterns of both algae and animals, which can be expected to greatly affect food availability to consumers at different height levels on the shore. We tested the hypothesis that consumers would feed on the most abundant suitable food source in their surroundings. In total 36 species/taxa of common primary producers and consumers were sampled for stable isotope analyses from a sheltered fucoid shore at Hvassahraun in south-western Iceland. A selection of these species was also collected seasonally and from different height levels. Feeding experiments, field observations and gut analyses were also conducted. Our results were in good overall agreement with pre-existing knowledge of trophic relationships in the rocky intertidal. Consumers often appeared to be assimilating carbon and nitrogen from the most common diet in their immediate surroundings. The predator Nucella lapillus was thus feeding on different prey at different height levels in accordance with different densities of prey species. When tested in the laboratory, individuals taken from low on the shore would ignore the gastropod Littorina obtusata, uncommon at that height level, even when starved, while individuals from mid-shore readily ate the gastropod. This indicated that some kind of learned behaviour was involved. There were, however, important exceptions, most noteworthy the relatively small contribution to herbivores, both slow moving (the gastropod L. obtusata) and fast moving (the isopod Idotea granulosa and the amphipod Gammarus obtusatus) of the dominant alga at this site, Ascophyllum nodosum. The recent colonizer Fucus serratus seemed to be favoured. Selective feeding was indicated both by isotope signatures as well as by results of feeding experiments. Seasonal migrations of both slow and fast moving species could partly explain patterns observed.     

Movement patterns of coastal bottlenose dolphins in the presence of a fast-flowing,prevailing current: shore-based observations at Cape Vidal,South Africa

The presence and movement of Indo-Pacific bottlenose dolphins Tursiops aduncus were investigated using shore-based observations made during a humpback whale Megaptera novaeangliae migration survey at Cape Vidal, South Africa, undertaken between June and October 1988–1991. Occurrence was analysed as counts of dolphin sightings per hour within a generalised estimating equation framework. Bottlenose dolphin sightings decreased throughout the daily survey period, and there was a small seasonal peak in sightings during August. Data on movement trajectory were collected using a theodolite for 60% of the groups of dolphins sighted. Most groups were observed travelling in a northerly direction, with few groups recorded moving south. A generalised linear model indicated that northward-travelling dolphins were located closer to shore, travelled at higher speeds, and occurred in bigger groups under low-tide conditions, than those seen moving in a southerly direction. The southward-flowing Agulhas Current is close to shore at Cape Vidal, where the continental shelf is particularly narrow. We suggest that bottlenose dolphins in this area have adopted a specific movement regime to cope with the consistent fast-flowing currents that dominate their environment at Cape Vidal.     

Sewage-derived bacteria monitored in a marine water column by means of captive mussels

Batches of mussels, Mytilus edulis, obtained from a sewage-free shore were introduced into the water column at a sewage polluted site and the changed content (Escherichia coli, faecal streptococci) was assessed after immersion periods of 2 to 40 h. Rapid bacterial uptake was demonstrated; changes were analysed with respect to immersion period, revealing that 2h was satisfactory. The numbers of bacteria varied with height in the water column; mussels from the sea bed contained higher numbers of E. coli than mussels immersed just below the surface except for mussels withdrawn at, and for about 3h after, low water (LW). These observations were shown to be related to the characteristics of the e effluent and tidal systems operative at the site.     

Measuring depth of settlement in the red rock lobster,Jasus edwardsii

Settlement of the puerulus stage of Jasus edwardsii (Decapoda: Palinuridae) near the shore takes place over a wide depth range. It occurs intertidally, and, as determined by crevice collectors, also takes place down to depths of at least 50 m. At sites along the east coast of the North Island of New Zealand, depth of greatest settlement found on collectors varied with locality and time, but was within the upper 11.5 m. Surface‐buoyed collectors caught fewer animals than did collectors at greater depths. Settlement levels generally increased with depth to about 10–12 m, then decreased with increasing depth and distance from shore. The ability of/, edwardsii to settle over a wide depth range may improve its chances of recruitment. A device (closing crevice collector) for measuring puerulus settlement of J. edwardsii at depth is described.     

Interactions between Zostera capensis,Callianassa kraussi and Upogebia africana: deductions from field surveys in Langebaan Lagoon,South Africa

Callianassid sandprawns and eelgrasses have been postulated to have mutually negative effects on each other. The relationship between Callianassa kraussi and Zostera capensis was investigated in Langebaan Lagoon on the west coast of South Africa by correlative field surveys. Seven facts point to mutually antagonistic interactions between the species: (1) cover of Z. capensis was negatively correlated with density of C. kraussi; (2) Z. capensis was largely confined to the high shore, whereas C. kraussi occurred in unvegetated sandbanks below the eelgrass beds; (3) at sites where Z. capensis was absent, C. kraussi extended farther up the shore than in its presence; (4) C. kraussi reached a larger maximum size in sandbanks than in beds of Z. capensis; (5) immediately below eelgrass beds there was a 'frontier zone' where above-ground Z. capensis was absent but below-ground root-rhizomes appeared to exclude C. kraussi; (6) sediment penetrability was greater in sandbanks than in eelgrass beds; (7) rate of burrowing by C. kraussi was faster in unvegetated sandbanks than in eelgrass beds. Conversely, the density and sizes of the mudprawn Upogebia africana were greater in beds of Z. capensis than in sandflats dominated by C. kraussi. Although this evidence is correlative, it suggests that Z. capensis excludes C. kraussi because it stabilises the sediment and inhibits burrowing, and that C. kraussi prevents establishment of eelgrass beds by destabilising the sediment and suspending material that smothers the eelgrass. In the process, two distinct habitats are created. U. africana may benefit directly from eelgrass because it requires stable sediment for its semi-permanent U-tubes; but also indirectly because Z. capensis excludes C. kraussi, which may disrupt these feeding burrows. At the head of the lagoon where sediments are very fine and compact, U. africana occurs both in and below the eelgrass bed, and coexists with at least low densities of C. kraussi, although Z. capensis and C. kraussi are still negatively correlated.     

Factors influencing the distribution of coastal lichens Hydropunctaria maura and Wahlenbergiella mucosa

Responses to abiotic and biotic stresses that potentially drive the vertical zonation of the intertidal lichens Hydropunctaria maura, an upper littoral lichen, and Wahlenbergiella mucosa, a lower littoral lichen, were investigated in field and laboratory experiments. When transplanted, both lichens exhibited an inability to survive outside their normal vertical distribution range. W. mucosa appeared to be unable to tolerate prolonged periods of desiccation following translocation from lower to upper littoral regions, whereas H. maura was unable to survive in lower littoral zones possibly owing to increased grazing pressure. The effect of desiccation in both lichens was compared using pulse amplitude modulated chlorophyll fluorescence and infra‐red gas analysis; results indicated a more hydration‐dependent nature of W. mucosa. Photosynthetic (algal) pigments and phenolic compounds were determined in both lichen thalli, and a range of additional coastal lichens occupying a natural gradient from upper to lower shore levels. Pigment composition and concentration in both lichen thalli were similar whereas levels of phenolic compounds were up to three times higher in W. mucosa than H. maura. Pigment and phenolic concentration and composition exhibited some seasonality across 13 different lichens originating from different shore levels. Phenolic concentration increased towards the lower shore, suggesting a potential anti‐herbivory function. This marks the first study of pigments and phenolics in coastal lichen communities, and prompts further investigations on the particular physiological features of marine and maritime lichens that enable them to thrive in this extreme environment.     

Associational resistance mediates predator--prey interactions in a marine subtidal system

There is a growing awareness of the role that indirect interactions play in influencing food webs and ecosystem structure. In this study, the hypothesis that crustose algal epibionts provide gastropods associational resistance from predation was investigated through field surveys and laboratory feeding assays. In rocky low intertidal/shallow subtidal systems in the northeast Pacific, several species of crustose algae (the red alga Peyssonnelia meridionalis and crustose corallines) can colonize the shells of living Tegula brunnea snails. The growth patterns of these epibiontic crustose algae allow them to cover their host’s surface completely, which may, in turn, protect their hosts from predation. A multi‐site field survey of T. brunnea revealed that >60% of snails were at least 75% covered with one or more species of crustose algae, with 35% fully covered, indicating that this is common in the field. Laboratory feeding assays revealed that sea stars, a primary predator of T. brunnea, distinguished among snails with different shell coverings; Pisaster consumed nearly three times as many bare (i.e. no crustose algae) snails as those covered with Peyssonnelia, while Pycnopodia consumed four times as many bare snails as those covered with crustose corallines. These results suggest that epibiont crustose algae can benefit their hosts via associational resistance; this finding may have implications for the role of associational resistance in trophic interactions.     

Abundance,Tidal Movement,Population Structure and Burrowing Rate of Emerita analoga (Anomura,Hippidae) at a Dissipative and a Reflective Sandy Beach in South Central Chile

Abstract. To evaluate the effects of beach morphodynamics upon the abundance, tidal movement, population structure and burrowing rate of the crab Emerita analoga (Stimpson) (Anomura, Hippidae) we sampled two beaches in south central Chile (ca. 42° S), Mar Brava and Ahui with dissipative and reflective characteristics, res­pectively. The swash zone at the dissipative beach was 5 – 6 times wider than that of the reflective beach. A at the dissipative beach, upwash speeds were higher and the number of effluent line crossings were lower by more than an order of magnitude. To examine the tidal movement of E. analoga, we collected crabs from 5 to 6 tidal levels of each beach every 2 h across 12 h of the tidal cycle. The intertidal distribution of crabs differed between beaches; i. e., at the dissipative beach they were primarily located at the swash zone, while at the reflective beach they were mostly located at the low tide level and shallow subtidal. The change in position of crabs was pronounced across the tidal cycle at the dissipative beach (Mar Brava), with most of the animals remaining in the active swash zone. Body size data were used to construct size frequency distributions for each population. Crabs from the dissipative beach reached larger sizes than those at the reflective beach. Sediments were coarser at the latter versus the former beach. Crabs burrowed at similar rates in the sand from both beaches, a result which supports the idea that E. analoga is a “sediment generalist” capable of burrowing successfully in a wide range of sediment types. This characteristic is likely a key to the broad success of this species on the full range of beach morphodynamic types along the coasts of South and North America.     

Spatial and Temporal Variations in the Zooplankton Community of Phosphorescent Bay, Puerto Rico

Nocturnal variations of zooplankton abundance and hydrographic conditions were examined at three locations (centre, north shore and south shore) in Phosphorescent Bay, Puerto Rico, from May 1992 through April 1993. Seven taxa accounted for approximately 96% of the annual mean zooplankton abundance:Oithonaspp. (43·5%),Acartia tonsa(31·5%), copepod nauplii (8·8%),Paracalanusspp.(6·7%), gastropod veligers (2·5%), larvaceans (1·7%) andPseudo-diaptomus cokeri(1·6%). Copepods dominated numerically throughout the year and comprised 94·3% of total zooplankton. Higher abundance of zooplankton (mean±1 SD=252 259±176 797 individuals m⁻³) was associated with cool water temperatures (24·9–27·4 °C) and dry conditions (0·3–2·9 cm precipitation/month) which prevailed between December and March relative to the period between April and November (warm/wet season) (mean±1 SD=59 773±26 861 individuals m⁻³), when temperature and precipitation were higher (27·3–30·3 °C, 3·1–20·6 cm month⁻¹). Fluctuations of zooplankton populations, particularly copepods, followed progressive increments in chlorophyllaconcentrations. This abundance pattern was consistent at the three sampling stations. Zooplankton abundance was higher on the north shore of the bay. The taxonomic composition of zooplankton was similar at the sampling stations studied.     

Variations in crab claw morphology and diet across contrasting inter‐tidal habitats

Crabs are important predators of inter‐tidal ecosystems, controlling the abundance and distribution of their prey populations. Often the same crab species occupies several habitats and, although their effects on prey have been quantified across habitats, crabs’ dietary and morphological responses to differing environmental influences have been overlooked. Here, we used the crabs Eriphia verrucosa and Pachygrapsus marmoratus as model species to examine differences in claw morphometry – size and wear – and diet between rocky shore and heterogeneous sand flat habitats. We predicted that, intra‐specifically, crabs from rocky shores would consume more hard‐shelled prey owing to their high availability and consequently, would display chelipeds with the following claw characteristics: a higher degree of claw damage, stronger musculature (higher propel height) and increased mechanical advantage (defined as the ratio of input lever length to output lever length) than crabs in the heterogeneous sand flat habitats. Sampling was performed in heterogeneous sand flat habitats and rocky shores of the Central Portuguese coast. For each crab species, carapace width, diet composition and several claw morphometric measures were recorded, revealing significant intra‐specific differences (using multivariate analysis) between shore types. We found that E. verrucosa and P. marmoratus consumed more hard prey on rocky shore than on sand flat habitats, which resulted in rocky shore crabs having more accentuated dentition wear and larger musculature than their sand flat habitat counterparts. We suggest that the strong response of crab claw morphometry to environmentally induced diet variations is an important mechanism in the successful adaptation of crab species to inhabit differing habitats. A major implication is that the impact of the same species on prey may vary largely with habitat type as a result of predation efficiency varying with claw condition.     

The interactive roles of predation and tidal elevation in structuring populations of the edible cockle,Cerastoderma edule

The size and age structure of the Cerastoderma edule population at Traeth Melynog, North Wales, varies dramatically with tidal level. In areas low on the shore up to 96% of cockle spat fail to survive their first summer, but mortality rate subsequently declines and remains at a low level. By contrast cockles high on the shore suffer moderate mortality during their first year (47%), but increasing rates thereafter. High-shore populations consequently consist mainly of smaller (younger) individuals and low-shore ones of a transient spatfall, plus a few larger and older individuals. The potential role of shore crabs, Carcinus maenas, and oystercatchers, Haematopus ostralegus in the determination of these patterns is assessed. Shore crabs move up into the intertidal to feed with each flood tide from about April to December. They selectively consume cockles < 15 mm in length, taking an estimated 236 × 10³ cockles, or 2432 g dry flesh year⁻¹ per linear meter of shoreline, mostly from lower shore levels. Oystercatchers are present only during winter and preferentially select large cockles of at least 20 mm length. They are estimated to remove 9 × 10³ cockles, or 1204 g dry flesh year⁻¹ per linear meter of shoreline, most of this from mid- and high-shore levels. These results indicate that shore crabs are far more important predators than previously suspected, taking 25 × the numbers and 2 × the biomass cosumed by oystercatchers. Predation also appears to be the key factor controlling the structure of the C. edule population. Crabs consume almost all the cockles settling low on the shore during their first summer, but avoid older individuals, which subsequently survive and grow well under low levels of oystercatcher predation. On the high shore, crabs are unimportant and the cockles survive well as they slowly grow into the size range attractive to oystercatchers. Thereafter they suffer increasingly severe winter mortality and are soon eliminated.     

Aspects of the ecology of a littoral chiton,Sypharochiton pellisekpentis (Mollusca: Polyplacophora)

On several Auckland shores, a littoral chiton, Sypharochiton pelliserpentis (Quoy and Gaimard, 1835), was widely distributed and common. At Castor Bay it was the commonest chiton, and its density equalled or exceeded that of the commonest limpet (Cellana spp.) over most of the inter‐tidal range. Spot measurements of population density were made at other sites including exposed and sheltered shores. The smallest animals were restricted to the lower shore in pools or on areas of rock which were slow to drain. Exclusive of these small animals, the population structure was similar in pools and water‐filled crevices situated either high or low on the shore. This was also true for populations of animals from drained surfaces, but, at both shore levels, the population structure in the two micro‐habitats (pools and drained surfaces) was distinct.

Single measurements of salinity in the very small volumes of water in which chitons were living gave a range of 13.3–45.8‰. In laboratory experiments at normal temperatures this species could survive these fluctuations for the maximum time possible between tides. Desiccation experiments, in which weight loss was expressed as a percentage of total water content, showed weight loss to depend on size. No significant differences in weight loss were found between populations from the two shore levels and the two micro‐habitats considered, though there were slight survival differences. Desiccation is probably an important factor influencing population structure, but loss of up to 75% of water content could be tolerated before 50% mortality occurred. Osmotic and desiccation stress evoked a characteristic behaviour pattern.     

Thermal constraints on the respiration and excretion rates of krill,Euphausia hanseni and Nematoscelis megalops,in the northern Benguela upwelling system off Namibia

Rates of respiration and ammonia excretion of Euphausia hanseni and Nematoscelis megalops were determined experimentally at four temperatures representative of conditions encountered by these euphausiid species in the northern Benguela upwelling environment. The respiration rate increased from 7.7 µmol O₂ h^?1 g_ww ^?1 at 5 °C to 18.1 µmol O₂ h^?1 g_ww ^?1 at 20 °C in E. hanseni and from 7.0 µmol O₂ h^?1 g_ww ^?1 (5 °C) to 23.4 µmol O₂ h^?1 g_ww ^?1 (20 °C) in N. megalops. The impact of temperature on oxygen uptake of the two species differed significantly. Nematoscelis megalops showed thermal adaptations to temperatures between 5 °C and 10 °C (Q₁₀ = 1.9) and metabolic constraint was evident at higher temperatures (Q₁₀ = 2.6). In contrast, E. hanseni showed adaptations to temperatures of 10–20 °C (Q₁₀ = 1.5) and experienced metabolic depression below 10 °C (Q₁₀ = 2.6). Proteins were predominantly metabolised by E. hanseni in contrast to lipids by N. megalops. Carbon demand of N. megalops between 5 and 15 °C was lower than in E. hanseni versus equal food requirements at 20 °C. It is concluded that the two species display different physiological adaptations, based on their respective temperature adaptations, which are mirrored in their differential vertical positioning in the water column.     

Enhancement of polycyclic aromatic hydrocarbons in estuarine invertebrates by surface runoff at a decommissioned military fuel depot

Accumulation of polycyclic aromatic hydrocarbons (PAHs) was determined in blue mussels (Mytilus spp.) and shore crabs (Hemigrapsus sp.) at a recently closed military fuel depot in central San Francisco Bay, California. In April 1996, during a period of above average precipitation, specimens were collected at the depot, near the depot, and at sites 10 and 20 km south of the depot. Four weeks after the rains ended, blue mussels were again collected at the depot, and at two additional sites in the central Bay region. In April, total PAHs in mussels from the depot were significantly higher only than that in mussels collected 20 km from the depot; however, seven specific, substituted PAHs were higher at the depot than at all other sites. In June, only two of the 38 PAHs common in mussels in April were detected at the depot; these concentrations were comparable to ambient concentrations in mussels at the Bay. It seemed that bioavailability of PAHs at the depot was enhanced by rainfall, probably due to the mobilization of PAHs via groundwater into the Bay. Concentrations in mussels from chronically contaminated sites were about five times higher than mussels collected from the depot. Low PAH concentrations were detected in shore crabs near the depot, and the highest levels were not associated with the depot. Observed PAH concentrations are discussed in relation to upper trophic organisms.     

Distribution and ecology of the crabs Cyclograpsus lavauxi and C. insularum in Northern New Zealand

The differences between the crabs Cyclograpsus lavauxi (Milne Edwards, 1853) and C. insularum (Campbell and Griffin, 1966) are outlined. Cyclograpsus lavauxi has long been known to be very common in New Zealand, but C insularum was considered very rare and had been collected only six times since 1929.

In 1970, 60 spot surveys in the northern part of the North Island of New Zealand revealed 14 new populations of Cyclograpsus insularum.

Both species are found under boulders high on the beach. Cyclograpsus lavauxi occurs at maximum density higher on the shore than the maximum numbers of C. insularum, but the zones overlap widely. Cyclograpsus insularum occurs on substrates of smaller sand and silt content than those on which C. lavauxi is found.