Assessment of space utilisation in a subtidal temperate bryozoan community

On subtidal hard substrata, Bryozoa are often very abundant, both in terms of numbers of colonies and numbers of species. To investigate how so many species are able to co-exist, often in close proximity to each other, subtidal material was dredged from an area of bryozoan-rich coarse shelly sea-bed off the south-west of the Isle of Man (British Isles) in autumn 1984 and 1985. A survey was performed on the distribution of colonies of eight common, encrusting cheilostome bryozoan species across the surfaces of four types of empty, disarticulated bivalve shells. Each species was found to be significantly commoner on certain substratum types such as rough surfaces or inner concave surfaces. Fenestrulina malusii (Audouin) showed a further, more specific, tendency to be signficantly more abundant on the grooves of the corrugated inner surfaces of left valves of Pecten maximus (L.). In general however there was much overlap in distribution and it is proposed that such large numbers of species are able to co-exist because of the evolution of different life-history strategies ensuring that each species uses the available temporally and spatially heterogenous substrata in subtly different ways; some species, for example, occupying substrata that are optimal to bryozoan growth and other species, by reaching maturity earlier, being able to inhabit more ephemeral suboptimal substrata.     

Inhibition and facilitation of bryozoan and ascidian settlement by natural multi-species biofilms: effects of film age and the roles of active and passive larval attachment

Sessile marine invertebrate larvae can recognize suitable settlement substrata by using various environmental cues, including organic/microbial biofilms. In laboratory choice assays, the effect of biofilms of varying ages on the settlement behaviour of two fouling organisms was assessed. The species included the arborescent cheilostome bryozoan Bugula flabellata (Thompson) and the solitary ascidian Ciona intestinalis (L.), both of which are characteristic of temperate sublittoral hard substratum assemblages in northwest Europe. Experiments were carried out using polystyrene petri dish substrata preconditioned with multispecies biofilms from natural laboratory-aquarium seawater for 1, 3, 6 or 12 d. Unfilmed (new, initially sterile) dishes were used as control substrata. Whereas the coronate larvae of B. flabellata generally were inhibited by biofilming, irrespective of film age, the settlement of tadpole larvae of C. intestinalis was facilitated on biofilmed substrata, and numbers of settled larvae generally increased with biofilm age: the highest mean numbers were counted on 12 d-old biofilms. In C. intestinalis, settlement and metamorphosis are processes which can be temporally separate and are possibly induced by different environmental cues. This study therefore distinguished between C. intestinalis larvae which were attached to the biofilm surface by the anterior, and those larvae entrapped by the biofilm but not settled in the conventional meaning of the term. As reported in previous studies, we did observe that such entrapped larvae could subsequently attach and develop successfully into sessile juveniles. Both the numbers of “attached” and “trapped” tadpoles increased with biofilm age. Assuming that “settlement” is essentially a process involving the active behavioural response of larvae to environmental cues, it seems that the facilitated attachment of C. intestinalis onto biofilmed substrata is due to the combined effect of active habitat selection and passive deposition/“entrapment” of larvae onto the “sticky” substratum. Received: 21 August 1996 / Accepted: 21 November 1996     

Assessing the strength of climate and land-use influences on montane epiphyte communities

Epiphytes, air plants that are structurally dependent on trees, are a keystone group in tropical forests; they support the food and habitat needs of animals and influence water and nutrient cycles. They reach peak diversity in humid montane forests. Climate predictions for Central American mountains include increased temperatures, altered precipitation seasonality, and increased cloud base heights, all of which may challenge epiphytes. Although remaining montane forests are highly fragmented, many tropical agricultural systems include trees that host epiphytes, allowing epiphyte communities to persist even in landscapes with lower forest connectivity. I used structural equations models to test the relative effects of climate, land use, tree characteristics, and biotic interactions on vascular epiphyte diversity with data from 31 shade coffee farms and 2 protected forests in northern Nicaragua. I also tested substrate preferences of common species with randomization tests. Tree size, tree diversity, and climate all affected epiphyte richness, but the effect of climate was almost entirely mediated by bryophyte cover. Bryophytes showed strong sensitivity to mean annual temperature and insolation. Many ferns and some orchids were positively associated with bryophyte mats, whereas bromeliads tended to establish among lichen or on bare bark. The tight relationships between bryophytes and climate and between bryophytes and vascular epiphytes indicated that relatively small climate changes could result in rapid, cascading losses of montane epiphyte communities. Currently, shade coffee farms can support high bryophyte cover and diverse vascular epiphyte assemblages when larger, older trees are present. Agroforests serve as valuable reservoirs for epiphyte biodiversity and may be important early-warning systems as the climate changes.     

Recovery of benthic communities at Macquarie Island (sub-Antarctic) following a small oil spill

Intertidal communities at Macquarie Island (sub-Antarctic) were re-evaluated seven years after a small oil spill resulting from the grounding of the “Nella Dan”, and six years after the first assessment of biological impact. Sampling was conducted to evaluate community structure in three zones of the exposed rocky shore (upper red, kelp, and lower red) and in samples of Durvillaea antarctica holdfasts. There were no significant differences between the community structure in oiled and control locations in any of the three shore zones, but holdfast macrofaunal communities at oiled sites still showed evidence of impact. Holdfast community-structure in samples from heavily oiled sites showed moderate levels of recovery, with increased abundances of species which were considered sensitive to the oiling in the first post-impact studies and decreased abundances of opportunistic polychaete and oligochaete worms. In contrast, samples collected from the moderately oiled location at Secluded Bay showed little evidence of recovery. Holdfasts at this site were filled with sediment containing traces of diesel oil, and the macrofaunal community was dominated by opportunistic worms. This study clearly indicates that even small incidents of anthropogenic perturbation can have long-lasting consequences for marine communities at Macquarie Island. Received: 23 July 1997 / Accepted: 5 February 1998     

Population and reproductive biology of two sibling amphipod species from ascidians and sponges

 The population dynamics and reproductive biology of an ascidian- and a sponge-dwelling amphipod were examined. The two undescribed amphipod species, Leucothoe“ascidicola” and L.“spongicola”, are closely related to each other, and occur in ascidians and sponges, respectively, along the Florida Atlantic coast. L. “ascidicola” was abundant in solitary ascidians during fall 1997, disappeared during spring/early summer, and became abundant again in September 1998. During the time when L. “ascidicola” were absent from their hosts, a copepod became a frequent inhabitant of the ascidians but disappeared again when L.“ascidicola” returned to the ascidians in September 1998. The numbers of L.“spongicola” in sponges increased substantially during spring, when high reproductive activity was observed. Following this reproductive peak, both adult and juvenile amphipods apparently left the sponges, and during the summer amphipod numbers in the sponges were very low. Another small amphipod species, which often co-occurred with L.“spongicola”, showed less seasonal variation and was found in sponges throughout the whole study period. The percentage of ovigerous females per host unit was usually lower in the ascidian-dwelling than in the sponge-dwelling amphipods. In solitary ascidians, L.“ascidicola” amphipods usually occurred in groups of several adults, yet there never was more than one ovigerous female per ascidian. In contrast, several ovigerous L.“spongicola” females were found to cohabit in the same spongocoel. This suggests that intrasexual aggression may be stronger among reproductive amphipod females in the ascidians than in sponges. The size distributions of juvenile cohorts indicate that juvenile L.“ascidicola” remain for relatively long time periods in the parental ascidian, where they may reach sexual maturity. In contrast, in L.“spongicola”, only cohorts of very small juveniles could be identified, indicating that juveniles disperse shortly after emerging from the female's brood pouch. It is concluded that extended parental care is of very short duration or does not occur in the sponge-dwelling amphipod L. “spongicola”, possibly because fast-growing sponges with a highly branched spongocoel system do not allow long-lasting coexistence of parent-offspring groups. In contrast, the discrete character of the solitary ascidians may enhance the potential for exclusion of other species, resource monopolization by reproductive females, and furthermore for long-lasting extended parental care in the ascidian-dwelling amphipod. Groups of single parents together with cohorts of large juveniles are reported in the literature for amphipods and isopods from brachiopods, bivalves and ascidians, suggesting that these discrete biotic microhabitats may favor the evolution of extended parental care in peracarid crustaceans. Received: 30 July 1999 / Accepted: 8 May 2000     

Development of epiphytic communities on eelgrass (Zostera marina) along a nutrient gradient in a Danish estuary

The effect of nutrient enrichment on epiphyte development was examined by following the seasonal development of epiphyte biomass on eelgrass (Zostera marina L.) at four localities along a nutrient gradient in Roskilde Fjord, Denmark between March and December 1982. In the most nutrient-poor area, epiphyte biomass followed a distinct bimodal seasonal pattern with maxima in spring and early fall. Low nutrient availability and a high rate of eelgrass leaf renewal kept epiphyte biomass at a low level throughout the summer period. Unlike phytoplankton, the epiphytic community was not stimulated by nutrient enrichment during spring, however, from May through August, the biomass of both components increased exponentially with increasing concentrations of total N in the water. Along the nutrient gradient, phytoplankton biomass increased 5- to 10-fold, while epiphyte biomass increased 50- to 100-fold. Thus differences in nutrient conditions among study sites were more clearly reflected by epiphytes than phytoplankton.Contribution No. 419 from the Freshwater-Biological Laboratory, University of Copenhagen     

Butterfly effects in mimicry? Combining signal and taste can twist the relationship of Müllerian co-mimics

Müllerian co-mimics are aposematic species that resemble each other; sharing a warning signal is thought to be mutually beneficial for the co-mimics by reducing per capita predation risk. In Batesian mimicry, edible mimics avoid predation by resembling an aposematic model species. The protection of both the model and the mimic is weakened when the mimics are abundant compared to the models. The quasi-Batesian view suggests that defended (Müllerian) co-mimics, when unequal in their defences, could also show a Batesian-like trend of increasing mortality with increasing abundance of a less defended “mimic”. We manipulated frequencies of unequally distasteful artificial co-mimics that were prey for great tits. The co-mimics had different signals (imperfect mimicry) but were equally preferred by the birds when palatable. Unexpectedly, when unpalatable, one of the signals was easier for the birds to learn to avoid. Consequently, during predator learning, the signal design of the prey strongly affected mortality of the co-mimics; there was an interaction between the signal and frequency treatments, but increasing the frequency of a less defended “mimic” did not increase co-mimic mortalities as predicted. In contrast, in a memory test that followed, the effect of signal design disappeared; if the birds had experienced high frequency of “mimics” during learning, co-mimic mortalities did subsequently increase. Since the effect of co-mimic frequencies on mortalities changed depending on the signal design of the prey and predator experience, the results suggest that mimetic relationship may be an unpredictable interplay of several factors in addition to taste and abundance.     

Foliose algal assemblages and deforested barren areas: phlorotannin content,sea urchin grazing and holdfast community structure in the Aleutian dragon kelp,Eualaria fistulosa

In the Aleutian Archipelago, two distinct organizational states of kelp forest communities exist, foliose algal assemblages and deforested barren areas. The canopy-forming kelp Eualaria fistulosa can be found in both states, although it is much less abundant in the deforested state. In contrast, sea urchins also occur in both states, but they are considerably more abundant in the deforested state. This study determined whether the protective phlorotannin content in E. fistulosa sporophylls that originated in foliose algal assemblages would differ from those that originated in deforested areas. We also examined sea urchin grazing rates on these sporophylls to determine whether there were any differences in grazing rates related to the origin of the sporophylls. Next, we determined whether taxon richness and abundance of E. fistulosa holdfast communities would differ depending on the origin of the holdfast. Our results showed that sporophylls that were collected in barren areas generally had higher phlorotannin content than sporophylls found in foliose algal assemblages, although phlorotannin content varied across the study area. Grazing rates on sporophyll tissue and holdfast community structure did not differ between foliose algal assemblages and barren areas. These results show that phlorotannin content is greatest in areas under high grazing pressure and that phlorotannins may possibly protect E. fistulosa and the holdfast communities from being grazed in barren areas.     

The gorgonian coral Acabaria biserialis: life history of a successful colonizer of artificial substrata

The azooxanthellate coral Acabaria biserialis Kükenthal, 1908 (Octocorallia: Alcyonacea) is highly abundant on the vertical underwater structures of the oil jetties at Eilat (Red Sea), but it is rare on the adjacent natural reefs. To understand its success on such artificial substrata we examined various aspects of its life history and population dynamics. A. biserialis is a gonochoric brooder. The percentage of colonies on the artificial substrata bearing gonads ranged from none (September 1994 and 1995) to 100% (January 1994, 1995 and April 1994). Mature oocytes and sperm sacs reached rather small maximum diameters of 240 and 160 μm, respectively, probably dictated by the small gastrovascular cavities. A continuous release of planulae was observed in the laboratory from March to July 1995. Planulation occurred during various lunar phases, and both by day and night. Recruitment of A. biserialis on PVC plates attached to the jetties coincided with the breeding period inferred from the laboratory findings. Recruits reached a maximum height of 3 cm within 3 months, thus exhibiting a remarkably rapid growth rate compared with other gorgonians. This high colony growth rate may compensate for the lack of clonal propagation in the studied A. biserialis population. The complex substrata of the jetties provide the conditions required for successful colonization, which include upside-down orientation, an adequate light regime and exposure to flow. The findings of our study may provide a useful contribution to the design of artificial reefs aimed at attracting rapid colonization by A. biserialis colonies. Received: 30 December 1998 / Accepted: 19 August 1999     

Epiphyte Biodiversity in the Coffee Agricultural Matrix: Canopy Stratification and Distance from Forest Fragments

Abstract: Quality of the agricultural matrix profoundly affects biodiversity and dispersal in agricultural areas. Vegetatively complex coffee agroecosystems maintain species richness at larger distances from the forest. Epiphytes colonize canopy trees and provide resources for birds and insects and thus effects of agricultural production on epiphytes may affect other species. We compared diversity, composition, and vertical stratification of epiphytes in a forest fragment and in two coffee farms differing in management intensity in southern Mexico. We also examined spatial distribution of epiphytes with respect to the forest fragment to examine quality of the two agricultural matrix types for epiphyte conservation. We sampled vascular epiphytes in a forest fragment, a shade polyculture farm, and a shade monoculture farm at 100 m, 200 m, and 400 m from the forest. Epiphyte and orchid richness was greater in the forest than in the monoculture but richness was similar in the forest and polyculture farm. Epiphyte species composition differed with habitat type, but not with distance from the forest. In the forest, epiphytes were distributed throughout tree canopies, but in the farms, epiphytes were primarily found on trunks and larger branches. Epiphyte richness and species similarity to forest species declined with distance from the forest fragment in the monoculture, but richness and similarity to forest species did not decline with distance from forest in the polyculture. This suggests polyculture coffee has greater conservation value. In contrast, monoculture coffee is likely a sink habitat for epiphytes dispersing from forests into coffee. Coffee farms differ from forests in terms of the habitat they provide and species composition, thus protecting forest fragments is essential for epiphyte conservation. Nonetheless, in agricultural landscapes, vegetatively complex coffee farms may contribute to conservation of epiphytes more than other agricultural land uses.     

How do nutrient conditions and species identity influence the impact of mesograzers in eelgrass-epiphyte systems?

Coastal eutrophication is thought to cause excessive growth of epiphytes in eelgrass beds, threatening the health and survival of these ecologically and economically valuable ecosystems worldwide. Mesograzers, small crustacean and gastropod grazers, have the potential to prevent seagrass loss by grazing preferentially and efficiently on epiphytes. We tested the impact of three mesograzers on epiphyte biomass and eelgrass productivity under threefold enriched nutrient concentrations in experimental indoor mesocosm systems under summer conditions. We compared the results with earlier identical experiments that were performed under ambient nutrient supply. The isopod Idotea baltica, the periwinkle Littorina littorea, and the small gastropod Rissoa membranacea significantly reduced epiphyte load under high nutrient supply with Rissoa being the most efficient grazer, but only high densities of Littorina and Rissoa had a significant positive effect on eelgrass productivity. Although all mesograzers increased epiphyte ingestion with higher nutrient load, most likely as a functional response to the quantitatively and qualitatively better food supply, the promotion of eelgrass growth by Idotea and Rissoa was diminished compared to the study performed under ambient nutrient supply. Littorina maintained the level of its positive impact on eelgrass productivity regardless of nutrient concentrations.     

An experimental study of habitat selection by birds in a coffee plantation

Unique components of tropical habitats, such as abundant vascular epiphytes, influence the distribution of species and can contribute to the high diversity of many animal groups in the tropics. However, the role of such features in habitat selection and demography of individual species has not been established. Understanding the mechanisms of habitat selection requires both experimental manipulation of habitat structure and detailed estimation of the behavioral and demographic response of animals, e.g., changes in movement patterns and survival probabilities. Such studies have not been conducted in natural tropical forest, perhaps because of high habitat heterogeneity, high species diversity, and low abundances of potential target species. Agroforestry systems support a less diverse flora, with greater spatial homogeneity which, in turn, harbors lower overall species diversity with greater numerical dominance of common species, than natural forests. Furthermore, agroforestry systems are already extensively managed and lend themselves easily to larger scale habitat manipulations than protected natural forest. Thus, agroforestry systems provide a good model environment for beginning to understand processes underlying habitat selection in tropical forest animals. Here, we use multistate, capture-recapture models to investigate how the experimental removal of epiphytes affected monthly movement and survival probabilities of two resident bird species (Common Bush-Tanager [Chlorospingus ophthalmicus] and Golden-crowned Warbler [Basileuterus culicivorus]) in a Mexican shade coffee plantation. We established two paired plots of epiphyte removal and control. We found that Bush-Tanagers were at least five times more likely to emigrate from plots where epiphytes were removed compared to control plots. Habitat-specific movement patterns were not detected in the warbler. However, unlike the Golden-crowned Warbler, Common Bush-Tanagers depend upon epiphytes for nest sites and (seasonally) for foraging. These dispersal patterns imply that active habitat selection based on the presence or absence of epiphytes occurs in C. ophthalmicus on our study area. Survival rates did not vary with habitat in either species. Interestingly, in both species, survival was higher in the nonbreeding season, when birds were in mixed-species flocks. Movement by Common Bush-Tanagers into areas with epiphytes occurred mostly during the breeding season, when mortality-driven opportunity was greatest.     

Conservation of Vascular Epiphyte Diversity in Mexican Coffee Plantations

Abstract:  Coffee plantations have replaced many lower-montane forests in the Neotropics, and ongoing intensification is converting traditional polycultures with a variety of shade trees to plantations with no or monospecific shade trees. To evaluate the impact of coffee cultivation on epiphyte diversity, I surveyed vascular epiphytes on shade trees in nine different coffee plantations and compared records with those in four natural forests in central Veracruz, Mexico. Eighty-nine species occurred in coffee plantations, and 104 species occurred in natural forests. The number of epiphytic species in traditional polycultures with old shade trees, mostly of the genus Inga , was similar to that in forests, but plantations with small trees and sparse shade hosted fewer epiphytes than those with large trees. Epiphyte communities were, however, more homogeneous in coffee plantations than in forests, possibly because of a drier microclimate and the lack of large and long-lived trees. These results demonstrate the value of traditional polycultures for epiphyte diversity, show the importance of conserving large shade trees, and suggest that these coffee systems may not be suitable for all epiphytes.     

Severe storm disturbances and reversal of community structure in a southern California kelp forest

Regular observations made over a period of 5 yr in four permanent transects provided data on plant, sea urchin, and fish densities which indicate that two unusually severe winter storms in 1980 (Storm I) and 1983 (Storm II) had different effects on a southern California kelp-forest community. Storm I removed all canopies of the giant kelp Macrocystis pyrifera, but spared most understory kelps, mainly Pterygophora californica. Hence, the previously large accumulation of detached drift kelp, mostly M. pyrifera, disappeared. Denied their preferred diet of drift kelp, the sea urchins Strongylocentrotus franciscanus and S. purpuratus then emerged from shelters to find alternative food. Without effective predators, they consumed most living plants, including the surviving understory kelps. This weakened the important detritus-based food chain, as indicated indirectly by declining abundances of algal turf and fish (Embiotocidae) that eat small animals living in turf. In 1983, Storm II reversed the process by eliminating exposed urchins, while clearing rock surfaces for widespread kelp settlement and growth. By summer 1984, the kelp grew to maturity to form extensive canopies despite elevated water temperatures during summer and fall of 1983. Thus, severe storms may have vastly different effects on community structure, depending on the state of the community before the disturbance.     

Seasonal growth in Laminaria longicruris: Relations with dissolved inorganic nutrients and internal reserves of nitrogen

Observations have been made on seasonal fluctuations in dissolved inorganic nutrients, internal reserves of nitrogen and growth rates in Laminaria longicruris. The onset of winter growth in shallow-water stations (6 and 9 m) correlated well with improved dissolved nitrate conditions in the sea. During the winter, reserves of NO ₃ ^- were accumulated by the plants and reached maximum values of 150 moles per g fresh weight in March. This represents a concentration factor of approximately 28,000 over the ambient levels, or an internal nitrogen reserve of 2.1% of the dry weight of the tissue. Depletion of this nitrogen pool followed the disappearance of the external NO ₃ ^- with a lag period of up to 2 months. Rapid kelp growth was measured during this period. Reserves of organic nitrogen also reached maximum values in March and declined slowly throughout the summer into autumn. It is suggested that the combined inorganic and organic nitrogen reserves sustain the rapid growth rates into July and at reduced rate through the late summer. Fertilization of an experimental perimental kelp bed with NaNO₃ increased the internal plant reserves of NO ₃ ^- and produced a much improved summer growth rate. The enriched plants developed very small reserves of carbohydrate during the rapid summer growth phase.NRCC No. 15549.     

Predator stimuli and calling behavior of Carolina chickadees (Poecile carolinensis), tufted titmice (Baeolophus bicolor), and white-breasted nuthatches (Sitta carolinensis)

Evidence from different chickadee species (Poecile genus) indicates that birds can modify the note composition of their “chick-a-dee” calls in the presence of predator stimuli. Here, we tested the effects of predator models and the distance of those models on calls of three species foraging together at feeding stations: Carolina chickadees (Poecile carolinensis) and tufted titmice (Baeolophus bicolor), both members of the Paridae family, and white-breasted nuthatches (Sitta carolinensis), a member of the Sittidae family. Model and distance affected seed-taking rates in all three species. “Chick-a-dee” calling rates were higher in the predator context for both chickadees and titmice, but we detected no predator context effects on “quank” call rates for nuthatches. Predator and distance contexts affected acoustic parameters of notes of the “chick-a-dee” calls of chickadees and titmice; no such effects were detected for nuthatch “quank” calls. These results suggest species differences in encoding of information in the primary social calls of these three species that commonly occur in multi-species flocks. Chickadees and titmice are “nuclear” species and nuthatches are “satellite” species, and these different roles might be related to the differences in vocal signaling that we detected.     

Intra- and interannual variability in the larval fish assemblage off Gran Canaria (Canary Islands) over 2005–2007

The present study provides the longest and most intensive plankton and larval fish seasonal variability analysis in the Canary Islands and forms a basis for understanding life cycle scheduling and interactions among species, as well as the potential variability in transport processes of early life stages. Larval fish assemblages were studied weekly at Gran Canaria Island, Canary Islands, from January 2005 to June 2007, which represented two contrasting hydrological and biological periods. The former year was characterized by lower temperature and salinity that increased through 2006 and 2007. In contrast, chlorophyll concentration and mesozooplankton biomass decreased through the same period, especially when only the late winter bloom period was evaluated. However, ichthyoplankton abundance did not exhibit any clear pattern, as larger values were observed during 2006. The larval fish community of this oceanic island, located near the NW African upwelling, was composed of both neritic and oceanic taxa. Two families accounted for almost half of the collected larvae: Clupeidae (21.9%) and Myctophidae (20.5%). Although total larval concentration did not exhibit any seasonal peak linked to changes in zooplankton, the ichthyoplankton composition gradually changed during the year due to the high diversity and extended spawning periods of the fish species represented in samples. “Winter” and “summer” larval assemblages were identified, corresponding to the mixing and stratification periods of the water column, respectively. These assemblages were characterized by changes in the contribution of the most abundant annual taxa (Sardinella aurita, Cyclothone braueri, Ceratoscopelus spp. and Gobids) and by the presence of larvae of winter (Pagellus bogaraveo, Pomacentridae sp1) or summer spawners (Pomacentridae sp2, Trachinus draco, Arnoglossus thori, Tetraodontidae sp1). Upwelling filaments shed from the NW African coast reached the sampling area three times during this study, but changes in the local larval community were only detected in August 2005.     

Life cycles ofCalanoides acutus,Calanus simillimus andRhincalanus gigas (Copepoda: Calanoida) within the Scotia Sea

The life cycles and distribution of three dominant copepods,Calanoides acutus, Calanus simillimus andRhincalanus gigas were studied from the Discovery collections in the Scotia Sea earlier this century.C. simillimus is a sub-Antarctic species which mates in the top 250 m mainly in spring. The rapid development of the summer generation may allow a second mating period and a smaller second generation to appear in late summer.C. simillimus remains in the surface layers for a longer period thanCalanoides acutus orR. gigas, and its depth distribution is bimodal throughout the winter.R. gigas is most abundant in sub-Antarctic waters to the north of the Polar Front. It mates within the top 750 m later in spring, and development seems less synchronised than that of the other two species, with egg laying and the growth season being more protracted. Stages CIII and CIV are reached by the first autumn and further development resumes very early the following spring. It is not clear whether the majority then spawn or whether a further year may be needed to complete the life cycle. The predominantly Antarctic species,C. acutus mates below 750 m in middle to late winter and the summer generation develops rapidly to either CIV or CV. Its lifespan seems typically 1 yr, but some of the CVs which fail to moult and spawn in winter survive into their second summer, and their subsequent fate is uncertain. The cold-water speciesCalanus propinquus is comparatively rare in the Scotia Sea and aspects of its distribution and life cycle are briefly described for comparison. Regional variations in the timing of these events were apparent forC. simillimus and possiblyCalanoides acutus, but were not seen inR. gigas. Their geographic and vertical separation, together with their asynchronous life cycles support the concept of habitat-partitioning of these dominant herbivores.     

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.     

Habitat use by juvenile Atlantic cod (Gadus morhua) in the presence of an actively foraging and non-foraging predator

Experiments were conducted in the autumn and winter of 1992/93 to examine habitat use by juvenile (age 0+) Atlantic cod, Gadus morhua L., before, during and following exposure to a passive or actively foraging predator (age 3+ cod). Experiments presented groups of juvenile cod (n=5 fish/group) with one of two combinations of three substrates; (1) gravel, sand, and a patch of artificial kelp (kelp), or (2) cobble, sand, and kelp. Cobble is known to provide juvenile cod with a refuge from predation. Kelp was used to test the hypothesis that juvenile cod associate with fleshy macroalgae in nature because of the safety it provides from predators. There was little difference in habitat use by juvenile cod before, during or following exposure to a passive predator. Under these conditions, juvenile cod appeared to prefer finer grained mineral substrates and avoided the kelp. The extent of the juvenile response to a passive predator was to avoid the predator's location in the experimental tank. In contrast, juvenile cod showed a significant shift in habitat use when exposed to an actively foraging predator, hiding in cobble or, when cobble was not available, in kelp. Use of both these habitats resulted in a significant reduction in predation risk to the juvenile cod. Our results suggest that: (1) an association with kelp provides safety from predation to juvenile cod, and (2) juvenile cod are capable for assessing the risk a predator represents and adjust their response accordingly.