Molecular and morphometric variation in European populations of the articulate brachiopod Terebratulina retusa

Molecular and morphometric variation within and between population samples of the articulate brachiopod Terebratulina spp., collected in 1985–1987 from a Norwegian fjord, sea-lochs and coastal sites in western Scotland, the southern English Channel (Brittany) and the western Mediterranean, were measured by the analysis of variation in the lengths of mitochondrial DNA (mtDNA) fragments produced by digestion with nine restriction endonucleases and by multivariate statistical analysis of six selected morphometric parameters. Nucleotide difference within each population sample was high. Nucleotide difference between population samples from the Scottish sites, both those that are tidally contiguous and those that appear to be geographically isolated, were not significantly different from zero. Nucleotide differences between the population samples from Norway, Brittany, Scotland and the western Mediterranean were also very low. Morphometric analysis confirmed the absence of substantial differentiation.     

The structure of subtidal algal and invertebrate assemblages at the Chatham Islands,New Zealand

We examined the distribution and abundance of organisms on subtidal rocky reefs at nine sites around the Chatham Islands, a remote group 780 km east of southern New Zealand. We sampled five depth strata ranging from 1 to<16 m to identify spatial patterns in the abundance of algae and invertebrates and to assess their variation within and among sites. This information is used to discuss hypotheses concerning community structure at this remote locality. Several patterns were apparent. The immediate subtidal was occupied by the southern bull kelp Durvillaea spp. A suite of 11 fucalean species were dominant to a depth of 10 m with an average abundance of 28 m^-2, while one species, Carpophyllum flexuosum, occurred mostly in deeper water. Only two laminarian species of algae were present at the islands. The indigenous Lessonia tholiformis was abundant at 2.5 to 15 m and was not found in deeper water, while the giant kelp Macrocystis pyrifera was abundant at two sites in 12 to 18 m. The commercially valuable abalone Haliotis iris was extremely abundant in shallow water, with an overall mean of 6 m^-2 at 5 m. The sea urchin Evechinus chloroticus was common, but reached high densities only in small (<25 m²) patches. The characteristic urchin-dominated zones reported in kelp beds world-wide were not seen. There was considerable site-to-site variation in the occurrence and abundance of individual species. Some differences between sites were associated with shelter from swell (e.g. M. pyrifera was found only in sheltered sites) and physical habitat (e.g. juvenile H. iris were found only beneath boulders inshore), but much of the variation could not be explained by physical or depth-related factors alone. We hypothesize that the differences in these kelp bed assemblages compared to mainland New Zealand are partially due to the high degree of endemism at the Chatham Islands. Local variation cannot be explained by herbivory, and is most likely the result of the various life-history characteristics of the major habitat-forming species, the large brown algae.     

Digestion in a subtidal population of Mercenaria mercenaria (Bivalvia)

The digestive processes of Mercenaria mercenaria (Linn.) were studied in a natural subtidal population. Hourly sampling of 4 to 5 quahogs was conducted over one 25 h period in August, 1978, at Woods Hole, Ma., USA. Crystalline style length did not vary significantly with time, tide, particulate C, N or suspended material in the water. Changes in secretory activity of the style sac epithelium were not evident from histological sections. Based on cellular morphological differences, digestive tubules were individually classified into one of 4 categories, indicative of the state of intracellular digestion within the digestive cells. All 4 tubule types were present in individuals from each sample hour, signifying that intracellular digestion occurs continually within the digestive gland. A 3 h time lag is evident between times of peak C, N or suspended materials and apparent peak levels of absorptive (Type II) tubules.Contribution No. 76 Marine Science Institute, Northeastern University; Nahant, MA 01908, USA     

Abundance of the New Zealand subantarctic southern right whale population estimated from photo-identification and genotype mark-recapture

The abundance of New Zealand subantarctic southern right whales (Eubalaena australis) was estimated for the first time using mark-recapture methods based on photo-identification and microsatellite genotyping (13 loci). Individual identification photographs of 383 whales and microsatellite genotypes of 235 whales were collected during annual austral winter field surveys from 1995 to 1998. Given the 4-year survey period and lack of geographic and demographic closure, we estimated super-population abundance using the POPAN Jolly-Seber model implemented in the software programme MARK. Models with constant survivorship but time-varying capture probability and probability of entry into the population were the most suitable due to the survey design. This provided estimates of abundance in 1998 of 908 non-calf whales (95% C.L. = 755, 1,123) for the photo-identification and 910 non-calf whales (95% C.L. = 641, 1,354) for the microsatellite genotype data sets. The current estimate of 900 whales may represent less than 5% of the pre-whaling abundance in New Zealand waters.     

Pelagic-benthic coupling in a subtidal system of the North-Western Mediterranean

Suspended particulate matter, zooplankton, and macrobenthos dynamics were investigated in a shallow area of the Ligurian Sea (north-west Mediterranean) characterized by wide temporal variability over an annual cycle. As indicated by multivariate analyses, the seasonal dynamics can be summarized as follows: (1) a late winter-early spring phytoplankton bloom followed by high zooplankton and macrobenthos densities during the spring months; (2) low-quality particulate suspended matter in summer, and an increase in the importance of zooplankton taxa with a wide range of feeding strategies, a decrease in macrofaunal abundance, and an increase in deposit-feeders and predators; and (3) a second phytoplankton bloom in autumn, followed by an increase in copepod density and a low macrofaunal abundance. In conclusion, pelagic and benthic communities in the coastal area of the Ligurian Sea mainly seem to be controlled bottom-up. Our results suggest that the quality of the particulate organic matter may play an important role in determining the temporal changes of both plankton and benthic assemblages, while the direct influence of other environmental features (such as sediment grain size) is relevant only for some macrobenthic taxa (e.g. crustaceans).     

Functional morphology of the gonads of the articulate brachiopodTerebratulina retusa

The reproductive anatomy, and ultrastructural features of the gonads of the articulate brachiopodTerebratulina retusa (Linnaeus), are documented based on collections made between October 1985 and October 1986 from the Firth of Lorn, west coast of Scotland. This species is dioecious, and maturity is achieved in both sexes at shell lengths greater than ~5.5 mm. There is no obvious external sexual dimorphism except for slight differences in the coloration of the gonads; testes are white/cream, ovaries are yellow/orange. The gonads occur as four palmate lobes, a pair in each valve. Gonads are formed within a mantle sinus (vascula genitalia), which is an anterior extension of the coelom, that opens posteriorly into the visceral cavity and to the exterior via a pair of metanephridia. The latter serve as gonoducts during spawning. Gametes are borne on genital lamellae formed from a reticulate lattice of connective tissue. The lamellae are an extension of the ileoparietal band and are fused along one margin to the inner mantle membrane. Developing oocytes are closely affixed to the genital lamellae and originate from a pool of proliferating germ cells at its base. Vitellogenic oocytes that are at an advanced stage are released from the genital lamellae, but are retained within thevascula genitalia. Liberated oocytes continue to accumulate yolk and eventually occlude thevascula genitalia, before being spawned. Coelomocytes were identified within the gonads. In spent females these cells appear to be phagocytic and involved in the resorption of necrotic material, while in the male they may serve as nutritive cells.     

Contrast in the importance of arrow squid as prey of male New Zealand sea lions and New Zealand fur seals at The Snares,subantarctic New Zealand

New Zealand sea lions (Phocarctos hookeri) are threatened by incidental bycatch in the trawl fishery for southern arrow squid (Nototodarus sloanii). An overlap between the fishery and foraging sea lions has previously been interpreted as one piece of evidence supporting resource competition for squid. However, there is currently no consensus about the importance of squid in the diet of New Zealand sea lions. Therefore, we investigated this importance independently of spatial and temporal differences in squid availability through a simultaneous study with sympatric New Zealand fur seals (Arctocephalus forsteri), a species known to target arrow squid. Diet sampling at The Snares (48°01′S 166°32′E), subantarctic New Zealand, in February 2012 coincided with peak annual catch in the nearby squid fishery. Diets were deduced by analyses of diagnostic prey remains from scats (faeces) and casts (regurgitations). The contribution of each prey species to the diet was quantified using the per cent index of relative importance (% IRI) that combined frequency of occurrence, mass and number of prey items. Arrow squid was a minor component in sea lion scats (2 % IRI), and none was found in their casts. In contrast, arrow squid was the major component in fur seal scats and casts (93 and 99 % IRI, respectively). This study found that New Zealand sea lions ate minimal squid at a time when squid was clearly available as evidenced by the diet of New Zealand fur seals; hence, there was no indication of resource competition between sea lions and the squid fishery.     

Conditional outcomes of facilitation by a habitat-modifying subtidal bivalve

Facilitation by habitat modifiers is common in ecological communities, but the potential for temporal and spatial variations in environmental conditions to modify the outcome of these interactions and influence the strength of feedbacks is poorly understood. Suspension-feeding bivalves are important habitat modifiers that can facilitate surrounding communities by providing refuge from predation and changing boundary flows and through the production of organically enriched biodeposits. However, numerous studies have highlighted the problem of finding generalizable patterns. We tested the strength and generality of the relationship between the large suspension-feeding bivalve Atrina zelandica and surrounding macrofauna and hypothesized that facilitation by Atrina is conditional and modulated by site-specific suspended sediment concentration (SSC), which influences the quantity and quality of biodeposit production. We found temporally consistent patterns of higher rates of biodeposition and increased abundance and species richness in close proximity to Atrina under low SSC conditions. Facilitation strength decreased with increasing SSC, suggesting that the facilitation effect of Atrina is reduced and reversed along this environmental stress gradient.     

A population study of the polychaete Capitella capitata at plymouth

A population of Capitella capitata (Fabricius) was sampled from January, 1973 to February, 1974. Size of the worms was measured and sexual maturity determined. There is an extensive breeding season, spawning occurring asynchronously throughout the year. Females containing ripe oocytes were almost always present, but most females at any one time contained immature oocytes. A high percentage of males contained active sperm. Oocytes are not released into the coelomic fluid until almost fully developed, and all are released at a single spawning. A large number of eggs are produced at each spawning and there is some evidence to suggest that larval development may be completely benthonic. Gametes first develop in 7 to 8 month old worms and take about 4 months to complete development. Analyses of the female population structure suggests that there are 3 breeding classes. The growth rate is estimated at 30 mm per year, and few worms are believed to survive longer than 2 years.     

New Approaches Toward a Better Understanding of the Decline of Takahe (Porphyrio mantelli) in New Zealand

Island populations of endemic birds are known to be particularly susceptible to extinction. This paper focuses on factors behind past and present declines of the highly endangered Takahe ( Porphyrio mantelli ), a flightless rail endemic to New Zealand. Subfossil evidence indicates that Takahe were once found throughout this island nation, being most abundant along forest margins and streams in lowland regions. Their numbers have declined dramatically since human colonization about 800–1000 years ago, probably due to a combination of habitat destruction and hunting by early Polynesian settlers. Today, the wild Takahe population consists of about 100 adult birds in an isolated alpine habitat and approximately 30 individuals recently released on several small offshore islands. Despite protection and intensive management, including removal of an introduced competitor (red deer, Cervus elaphus ), the alpine population has continued to decline. In contrast, the Takahe's nearest relative, the Pukeko ( Porphyrio porphyrio ) has expanded its range across New Zealand despite heavy hunting pressure since its colonization from Australia less than 1000 years ago. We suggest that, unlike Pukeko, Takahe lack appropriate behavioral responses to cope with mammalian predators such as stoats ( Mustela erminea ), which have been introduced relatively recently by European colonists. A study comparing predator defense behavior between these two closely related species is currently underway.     

A genetic analysis of the pea crabs (Decapoda: Pinnotheridae) of New Zealand

Seven populations of the pea crabPinnotheres atrinicola Page were sampled from around the North Island of New Zealand from February to October 1987, and individuals were scored at 23 presumptive enzymatic loci. For a brachyuran crab,P. atrinicola revealed high levels of polymorphism and heterozygosity. Of the loci scored, phosphoglucose isomerase (Gpi) and phosphoglucomutase (Pgm) were distinguished by high variability ( =0.602 and 0.526, respectively). A clinal variation in electromorph frequency was evident at several loci, and atGpi in particular. Statistical analyses revealed that, despite relatively small genetic distance separation, a high degree of structuring was present between the geographic populations. The degree of population subdivision observed in this study is atypical of brachyuran crabs. It is suggested that the genetic differentiation observed between pea crab populations is maintained by life-history attributes and current movements which restrict gene flow between populations and, to some extent, by random genetic drift.Publication No. 41 from the Evolutionary Genetics Laboratory, University of Auckland     

Mortality estimates from repeated visual censuses of a parrotfish (Sparisoma viride) population: demographic implications

Repeated visual censuses of different categories (juveniles, females, territorial and group males) of the stoplight parrotfish (Sparisoma viride, a protogynous hermaphrodite) over a 3-year period indicated a relatively stable size and structure of the adult population. This allowed estimates of size-specific mortality, sex change, and territory acquisition probabilities from previously reported growth rates. Comparison of the predicted number of survivors, sex changers, and territory take overs with field observations indicates that our estimates are quite reliable. However, rather large differences in mortality are obtained for the largest three size classes (>25 cm), which may be due to reduced accuracy of length estimates of large fish. A pooled mortality for these classes is therefore suggested as a more realistic estimate. The life-history implications of our findings are investigated by comparing the predicted survival and future reproductive success of fish that change sex at different sizes. Ten percent of the adults reach an age of ca. 17 years, once a length of 20 cm is attained, as predicted from the pooled estimates. Calculations based on these estimates indicate that the predicted reproductive output of a 15-year-old fish (2500 matings) is the same for early (i.e. at 20 cm) and late (>30 cm) sex changers and for an average fish subject to the estimated daily sex change and territory acquisition probabilities. These findings suggest that S. viride individuals are able to flexibly adjust the timing of sex change in an adaptive way to unpredictable local conditions. However, independent mortality estimates are needed to corroborate our present findings. Received: 2 January 1997 / Accepted: 27 January 1997     

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.     

Oxygen- and carbon-isotope analyses of the articulate brachiopodLaqueus californianus: a recorder of environmental changes in the subeuphotic zone

This study is the first systematic attempt to use oxygen and carbon isotopes in modern brachiopods to investigate seasonality and growth rates. A comparison of oxygen-isotope analyses of shells of living specimens ofLaqueus californianus dredged from 80 and 130 m off Santa Catalina Island, California, to available hydrographie data indicates that this articulate brachiopod secretes its shell in or close to oxygen isotopic equilibrium with ambient seawater. Periodic oxygen-isotope depletions appear to result from El Nino events. Unexpectedly low ¹³C values associated with the 1983 El Nino may be explained by increased bacterial activity or organic loading into the Southern California Bight associated with these warm-water pulses. Growth rates determined from annual cycles in ¹⁸O records are variable, but generally average between 2 and 3 mm yr^–1 for mature individuals. Because of the longevity ofL. californianus,¹⁸O profiles provide high-resolution seasonal temperature records spanning one to two decades. Our data suggest that oxygen isotopes in brachiopod shells can be utilized as monitors of environmental change in the subeuphotic zone.     

Patterns of shell repair in articulate brachiopods indicate size constitutes a refuge from predation

The cost of overcoming prey defenses relative to the value of internal tissues is a key criterion in predator/prey interactions. Optimal foraging theory predicts: (1) specific sizes of prey will result in the best returns to predators, and (2) there will often be a size at which the cost/benefit balance is low enough to effectively exclude predation. Data presented here on styles of repaired shell damage and size at which injury had been sustained was collected from samples of terebratulide brachiopods from the Antarctic Peninisula (Liothyrella uva), Falkland Islands (Magellania venosa and Terebratella dorsata) and Chile (M. venosa). The predominant form of damage on shells was indicative of predators attacking the valve margins. The modal size for repaired damage was more than 10 mm smaller than the modal size for the overall size distribution in each species and there were no repaired attacks in the largest size classes of any species. These data suggest that size forms a refuge from predation, as would be predicted by optimal foraging theory. The optimal sizes that predators appeared to attack vary between species, as do the sizes that provided a refuge from predation. High levels of multiple repairs (19% of the M. venosa population from the Falkland Islands sampled had 2 or more repairs) suggest that the mortality following attack is low, suggesting that many predators abandon their attacks.     

Reproductive periodicity in the euryalinid snake star Astrobrachion constrictum in a New Zealand fiord

The euryalinid snake star Astrobrachion constrictum (Farquhar), a normally deep-water species, exists within SCUBA diving range in New Zealand flords, and thus presents a unique opportunity for study. A population of A. constrictum in Doubtful Sound was investigated from February 1990 to July 1994 to determine the reproductive cycle. Unlike most ophiuroids, the gonads of A. constrictum are positioned along the aboral coelom in the proximal portion of the arms. The proximal 2 to 3 cm of the arms of individuals were thus removed on an approximately monthly basis for histological examination of the gonads. Image analysis was used to determine oocyte size-frequency distributions. A gonad index was calculated for each individual based on the ratio of cross-sectional area of the gonads to cross-sectional area of the arm. Maturity indices were calculated based on stage of development of the gonads of each individual. A. constrictum has a period of enhanced breeding activity in April–June, after which both the gonad and maturity indices decline. The increase in gametogenic activity during these months was correlated with rising sea temperature within the fiord. Although the females showed an annual pattern of gonad development, oogenesis was continuous throughout the year, with oocytes at different stages of development present in the gonads. Males also showed continuous spermatogenesis and, although the same annual pattern of gonad development was evident as seen in the females, it was less marked. The presence of females with large oocytes and ripe males throughout the study indicates that breeding in the Doubtful Sound population is prolonged, and possibly continuous, as found in many deep-sea ophiuroids.     

Genetic structure of the subtidal red alga Delisea pulchra

We used random amplified polymorphic DNA (RAPDs) to examine small-scale spatial genetic structure in the red alga Delisea pulchra (Greville) Montagne at two locations near Sydney, Australia. We examined genetic structure among plants at four spatial scales ranging from 2 km apart down to <50 cm apart between locations, among sites within locations, among quadrats within sites, and among plants within quadrats. Haploid stages of D. pulchra were absent from the populations studied, suggesting that they are maintained through asexual reproduction of diploid plants. Consistent with this, we found that 19 RAPD phenotypes scored in this study had multiple individuals, indicating the presence of clones in these populations. However, there were no RAPD phenotypes common to two locations separated by only 2 km. Analysis of molecular variance revealed that strong genetic differences occurred between plants from these two locations, with 46.3% of the total genetic variation occurring at this scale, most probably reflecting limited gene flow. Within each location, <25% of the genetic variation was attributable to differences among sites or quadrats, indicating gene flow at those smaller scales. Most of the variation within each location occurred at the smallest spatial scale, among plants within 0.25 m² quadrats. Nonetheless, some pairwise genetic distances (φST) between sites or quadrats within locations were large, indicating some genetic divergence on smaller scales. Genetic distance was independent of spatial distance within both locations, suggesting that fine-scale differences within locations were most probably caused by variation in fine-scale patterns of water movement or fine-scale natural selection. We assessed the impact of one potential selective agent, grazing sea urchins, on the fine-scale genetic structure of D. pulchra. There was no evidence that grazing by sea urchins affected the genetic structure of D. pulchra. In combination with demographic data, our results indicated that local populations of D. pulchra within locations were relatively open and that fine-scale genetic structure was probably constrained by gene flow. At the larger scale however, strong genetic differentiation indicated little gene flow between locations and restricted dispersal of spores. Received: 22 April 1999 / Accepted: 29 November 1999     

Seasonal growth of the giant kelp Macrocystis pyrifera in New Zealand

The seasonal growth pattern of Macrocystis pyrifera (L.) C. Agardh in New Zealand was determined from measurements of blade-elongation rates between March 1986 and December 1987. Growth rates remained relatively constant throughout most of the year, but declined significantly during the summer months. Seawater nitrate levels had a marked seasonal cycle, with concentrations of 0.3 μM detected in summer. The timing of the decline in internal nitrogen concentrations varied for different-aged blades, but occurred ∼1 mo after the decrease in seawater nitrate concentrations. Sufficiently high irradiance levels and seawater nutrient concentrations support active growth for most of the year, but growth is nutrient-limited during summer. No carbon accumulation during summer was observed. Between March and August 1988, growth estimates were also derived from node-initiation rates and stipe-elongation rates to permit comparisons with previous studies from the northern and southern hemispheres. Direct comparisons of the three growth parameters determined for non-terminated canopy and sub-surface fronds were used to assess possible differential resource allocation in the two frond classes. Received: 16 May 1997 / Accepted: 28 May 1997     

Condition cycle of the green-lipped mussel Perna canaliculus in New Zealand

The condition of the green-lipped mussel Perna canaliculus (Gmelin) from 7 sites around the northern half of New Zealand was determined monthly between December 1973 and February 1975. Seven separate estimates of condition were calculated for each sample; 4 condition indices, 2 percentage glycogen analyses and 1 estimate of percentage solids. All showed a similar annual cycle, with minimum values in winter (June/ August) rising to peak levels in spring (October/ December). The 7 sites were divided into 3 groups, (i) a central New Zealand group, characterised by high levels of condition index (CI) throughout most of the year >10 CI_weight for 8 to 11 months, average to high percentage glycogen (annualmeans 25 to 35% dry weight), and high percentage solids (annual means > 20% dry weight); (ii) an intertidal group, having low levels of CI (<10 CI_weight for 10 to 11 months), low percentage glycogen(mean <20%), and low percentage solids (mean <18%); and (iii) a northern group, showing intermediate values for each of the estimates of condition. Difficulties in defining the causes of changes in the condition cycle are discussed. There is an inverse relationship between CI and mussel size, resulting from changes in body proportions. An inverse relationship is also suggested between CI and water temperature. A precise index, $$CI_{weight} = \frac{{100 \times dry meat weight}}{{whole{\text{ weight - shell weight}}}}$$ is recommended for biological studies on mussels and a rapid simple index, $$CI_{commercial} = \frac{{100 \times wet meat weight}}{{whole{\text{ (live) weight}}}}$$ for use in the field and in mussel farming practice.