Seasonality of reproduction in the cushion star Odontaster validus at Signy Island, Antarctica

The reproductive and larval biology of the common Antarctic cushion-star Odontaster validus Koehler (Echinodermata: Asteroidea) were studied in a shallow-water population at Signy Island in the maritime Antarctic. The gonad index peaked in April/May, and injection with 1-methyladenine indicated that competence to spawn peaked in May/June. Gastrulae were first detected in May, and maximal water-column larval densities were measured in July. Reproduction was therefore highly seasonal, although not all individuals appeared to spawn each year. The pyloric caeca index increased in summer (typically November to March, but variable between years) and decreased during winter (June to October), suggesting that feeding activity was seasonal. All aspects of biology investigated showed significant interannual variability. Comparison with previous studies in the high Antarctic at McMurdo Sound indicated a similar if more pronounced seasonality at Signy, but the warmer seawater temperatures in the maritime Antarctic (particularly in late winter/early spring from September onwards) resulted in faster larval development at Signy. Received: 21 April 1997 / Accepted: 28 January 1998     

Stereological methods applied to reproductive cycle of Tapes rhomboides

A population of Tapes rhomboides (Pennant) in the Bay of St. Malo, France, was studied for one and a half years (July 1984–October 1985) to determine spawning frequency and fecundity under natural conditions, using the techniques of qualitative histological staging, condition index calculation and quantitative stereology. Spawning took place twice a year, in late May and in July/September. There was an extended winter resting period. Gonad development and oocyte production were positively correlated with female body size. The annual fecundity in a 40 mm clam was about 5x10⁵. Stereological techniques provided accurate information on oocyte dynamics within a gonad and the relationship with atretic processes. Each oocyte generation from the onset of the cycle to the winter resting stage was spawned eventually if conditions were suitable, or the oocytes were resorbed. Atresia of oocytes was seen throughout the reproductive period, but especially after the spring spawning and at the end of the summer.     

Histological investigation of the reproductive cycles of the limpets <Emphasis Type="Italic">Patella </Emphasis><Emphasis Type="Italic">vulgata</Emphasis> and <Emphasis Type="Italic">Patella ulyssiponensis</Emphasis>

This study devised a staging system for, and monitored, the gonad development of the limpet species Patella vulgata and Patella ulyssiponensis on the South West coast of Ireland using histological techniques. Maturation began in the males of both species in January and in the females it began in March. There was no statistical difference in gonad development between sexes and between species. Spawning in the male P. vulgata occurred from September to December 2003 and in September and October 2004. In female P. vulgata spawning occurred from October to December 2003, no spawning of females was observed in 2004. In male P. ulyssiponensis spawning occurred in November and December 2003 and from September 2004 to December 2004. Spawning was observed from November 2003 to January 2004 and in September 2004 in female P. ulyssiponensis. Sex ratios also varied between the species and between months sampled. Nevertheless more males were observed in both species.     

Strengthening recruitment of exploited scallops Pecten maximus with ocean warming

There is evidence that ocean warming has effects on the ecology, including recruitment dynamics, of marine organisms. In association with rising mean spring temperatures in the Irish Sea, a time-series of juvenile scallop Pecten maximus density around the Isle of Man showed a significant increasing trend since 1991. Favorable conditions (warmer water and correspondingly greater food availability) during gonad development can increase scallop gamete production. We examined the possibility that ocean warming has directly increased recruitment of exploited P. maximus around the Isle of Man by enhancing gonad development. From 1991–2007, there was a significant positive correlation between scallop recruitment and mean spring (the main period of gonad development) temperature in the year of larval settlement. Detrended (i.e., accounting for a time effect) recruitment data showed a marginally non-significant correlation to temperature. Gonadal somatic index of adult scallops and temperature were positively correlated. These relationships support the hypothesis that greater gamete production associated with ocean warming may be primarily responsible for observed increases in recruitment success and CPUE in a commercially important shellfish stock.     

Reproductive biology of the commercial sea cucumber Holothuria fuscogilva in the Solomon Islands

 Reproduction of Holothuria fuscogilva (Selenka, 1867) in the Solomon Islands was investigated over a 4 yr period (1994 to 1998) by macroscopic and microscopic examination of the gonad tubules, the gonad index (GI) method, histological examination of gametogenesis, and spawning-induction trials. The gonad consisted of numerous tubules that dominated the coelom of gravid specimens. New tubules appeared in March, and grew in size and extent of branching until they reached their maximum size and maturity in August. Spawning occurred from August to October, with the majority of gametes released during October, although it was only partial in many individuals. After spawning, the tubules appeared wrinkled and resorbed into the gonad basis. A five-stage gonad maturity scale based on the macroscopic appearance of the gonad tubules corresponded with discrete stages of gametogenesis identified by histology. Gametogenesis was initiated in mid-March, with oogenesis and spermatogenesis occurring in parallel, followed by the growing stage (May to July) which was marked by active gamete development. Successful induction of spawning during the breeding period corroborated the GI and histological data. The uniform growth of gonad tubules indicated that H. fuscogilva in the Solomon Islands does not conform to the progressive tubule recruitment model described for other holothurians. An important application of this study is that the appearance of gonad tubules, removed by biopsy, can be used to determine the gonad condition of wild adults or captive broodstock. Received: 28 June 1999 / Accepted: 8 March 2000     

Sexual reproduction of the reef coral Montastreacavernosa (Scleractinia: Faviidae) in the Santa Marta area, Caribbean coast of Colombia

Sexual reproduction of the reef-building coral Montastreacavernosa was studied in the Santa Marta area, Caribbean coast of Colombia, from October 1990 to October 1991. The area is subjected to a seasonal upwelling–outwelling regime. From microscopical analysis of tissue sections sampled each lunar month around the full moon, it was confirmed that this is a gonochoric, broadcasting species, with a single gametogenic cycle per year, and a 1:1 sex ratio. Oogenesis began a few weeks after spawning, and extended for 11 mo. The onset of spermatogenesis occurred just before the full moon of June, later than at other localities where sea-water temperatures during the first half of the year are not as low. The amount of reproductive tissue strongly increased in both sexes after July, in association with an increase in sea-water temperature and a decrease in photoperiod. The gonad index of fully mature female colonies was about four times lower than at other localities, perhaps due to the stressful seasonal regime. The male gonad index was lower than that of the female, indicating sex-related differences in the rates of biomass allocation to reproduction. As in other Caribbean localities, spawning occurred after the full moons of August and September. However, there was some evidence of a third spawning episode after the full moon of October, possibly associated with a delay in the occurrence of maximum sea-water temperatures in near-equatorial localities compared to higher latitudes. The probability of cross-fertilization in this species with a gonochoric breeding system and a broadcasting mode of reproduction is favored by its balanced sex ratio, its usually high within-reef zone abundance, and by localized and repeated spawning episodes, synchronized by lunar phase. Received: 19 April 1996 / Accepted: 24 September 1996     

Reproductive cycles of tropical sea cucumbers (Echinodermata: Holothuroidea) in southern Taiwan

Reproductive cycles of Afrocucumis africana (Dendrochirotida), Actinopyga echinites, Holothuria leucospilota, H. cinerascens, H. difficilis (Aspidochirotida), Synaptamaculata, Opheodesoma grisea, Patinapta taiwaniensis and Polycheira rufescens (Apodida), representing three orders of intertidal holothurians in southern Taiwan, were determined by gonad index and histological examination from March 1990 through September 1991. All nine species of holothurians have annual spawning periods lasting 2 to 4 mo in spring or summer. The early spawning of H. cinerascens and P. rufescens (suspension feeders) suggests that their feeding mode is related to food resources and may influence the reproductive period. In the direct-developing dendrochirotid (suspension feeder), gonad development occurs in cool months, and gametes are released at the beginning of warm months. In indirect-developing aspidochirotid and apodid (deposit feeder), gamete release occurs in late spring or summer and appears to be correlated with summer phytoplankton growth. The holothurians in the present study display spawning periodicity and duration similar to those of temperate species. Four species of holothurians lengthened spawning season with decreased latitude.     

Seasonal variations in the nucleic acid content and RNA:DNA ratio of the gonad of the scallopPecten maximus

The RNA and DNA contents of the male and female portions of the gonad of the hermaphroditic marine bivalve molluscPecten maximus L. have been monitored throughout an annual reproductive cycle, from October 1985–September 1986, for a population from the Baie de Seine (Normandy, France). The DNA content of the male gonad was consistently 5 to 10 times greater than that of the female gonad, whilst the RNA content was lower. This underlines the basic premise that spermatogenesis involves the production of large numbers of small gametes, whereas oogenesis involves the production of few, large, synthetically active gametes. The RNA:DNA ratio is a good indicator of sexual maturation in the male gonad. Two peaks of RNA:DNA ratio were observed for the female gonad, possibly corresponding to gonad restoration and vitellogenesis.     

Spawning periodicities coinciding with semidiurnal tidal rhythms in the chiton Acanthopleura japonica

Spawning of the chiton Acanthopleura japonica was examined in 1983 and 1984, by means of consecutive sampling of newly spawned eggs from Tanabe Bay, Japan. Abundant collection of the eggs was recorded just before the high tide peaks on the days around spring tide, predominantly in the morning, for 7 to 9d, and subordinately in the afternnon for 3 to 4d. Water temperature has no effect on spawning time.     

Seasonal abundance,distribution and growth of postlarval and juvenile grass shrimp (Palaemonetes pugio) in a Georgia,USA, salt marsh

Postlarval and juvenile grass shrimp (Palaemonetes pugio Holthuis) ≦15 mm total length (TL) were abundant at low tide in shallow aquatic microhabitats (i.e. puddles and films of residual tidal water) in the intertidal zone of a salt marsh on Sapelo Island, Georgia, USA from 1982 to 1984. The highest concentrations of young P. pugio occurred at 190 to 200 cm above mean low water, ∼10 to 20 cm below mean high water. The intertidal distributions of young grass shrimp expanded and contracted with changes in tidal amplitude. Postlarval grass shrimp (6 to 8 mm TL) continuously recruited into the intertidal marsh population from May until October, but densities varied in a regular pattern with peaks in abundance occurring at ∼2-week intervals, corresponding to spring tide periods in the lunartidal cycle. Although present nearly year-round in the intertidal marsh, juveniles (9 to 15 mm TL) were most abundant from August to October. Apparent growth rates of individuals up to 15 mm TL averaged 0.268±0.026 (mean±95% C.I.) mm d^-1 from May to October and 0.070±0.032 mm d^-1 in November and December. Unlike larger aquatic organisms, which can forage in the emergent marsh only when it is flooded by the tide, juvenile grass shrimp have constant access to intertidal resources. Although potentially important predators in this system, the role of young P. pugio in the trophic organization of salt marsh benthic invertebrate assemblages has yet to be examined. Contribution No. 576 of the University of Georgia Marine Institute     

The gametogenic cycle of<Emphasis Type="Italic"> Octopus vulgaris</Emphasis> (Mollusca: Cephalopoda) as observed on the Atlantic coast of Andalusia (south of Spain)

The present study reports the gametogenic cycle of the common octopus, Octopus vulgaris Cuvier, 1797 in the Atlantic coast of Andalusia between May 2001 and May 2002. A total of 673 specimens of total weight ranging from 30 to 3,732 g were analysed. The gonad development was defined by means of histological methods and gonadosomatic indexes. Four gametogenic stages were identified in males (I immature, II maturing, III mature, IV spawning) and in females (I immature, II maturing, III pre-spawning, IV spawning). The sex ratio obtained was not significantly different from 1:1 (P>0.01). Mature males were found during the whole year, September being the month when the highest rate of sexually active individuals occurred (48%). In females, the gametogenic cycle commenced in December and ended in November, and spawning occurred between April and October, with two maximum peaks in August and September (48% in each). Size at sexual maturity was 850 g of total body weight for males and 1,250 g of total body weight for females. Males mature earlier than females.Communicated by S.A. Poulet, Roscoff     

The Arctic sea butterfly<Emphasis Type="Italic"> Limacina helicina</Emphasis>: lipids and life strategy

The sea butterfly Limacina helicina was collected from May to September 2001 in Kongsfjorden, Spitsbergen, to investigate population structure and body and lipid composition with regard to life cycle and reproductive strategy. Veligers and juveniles were only found in late autumn and spring, whereas females occurred from July to September. The size of the females increased until mid-August and decreased in September. Dry and lipid mass were closely related to size; dry mass increased exponentially and lipids linearly with size. The lipid content was highest in veligers (31.5% of dry mass) and juveniles (23.6%) but low in females (<10%). Phospholipids were the dominating lipid class followed by triacylglycerols. Females, veligers, and egg ribbons, all from September, were richest in phospholipids. Juveniles contained the highest amounts of triacylglycerols, whereas females had low levels in July and the beginning of August. In mid-August, levels of triacylglycerols were variable and higher. This suggests that females were in their main spawning period and the high variability in triacylglycerols points to different stages within the spawning cycle. Enhanced amounts of free fatty acids in females from July may be related to gonad development. The 16:1(n-7) fatty acid was more dominant in spring whereas 18:4(n-3) increased in summer and autumn, which reflects a change in diet from diatom-dominated food items in spring to dinoflagellates in summer/autumn. Small amounts of long-chain monounsaturated fatty acids suggest ingestion of copepods, and the fatty acid composition of veligers feeding on particulate matter. L. helicina has a one-year life cycle with peak spawning in August and over-winters as veligers that may grow to juveniles during the winter period. They metamorphose into juveniles during spring, develop to males in early summer, and mature into females in July and August.Communicated by M. Kühl, Helsingør     

Analysis of the relationship between gonads and hepatopancreas in males and females of the crab Sesarma intermedia,with reference to resource use and reproduction

At the beginning of the reproductive period in May, there was an increment in both the gonad and heaptopancreas. During the peak phase in June, with high ovary production, there was a decrease in the hepatosomatic index (HSI), particularly for the Ariake females. The minimum gonadosomatic index (GSI) of 2% in August and September for the Ariake and in September and October for the Hakata populations was inversely related to the increase of HSI. A second rise in GSI was observed in November at the 1% level as against the June peak of 5% for the Ariake and 3.5% for the Hakata females, and it remained at this level until December, while the HSI decreased to the minimum. The December minima for both GSI and HSI were about the same as the initial February values. The spermathesomatic index (SSI) had a slight increase to 0.8% in August, but remained constant at 0.5% throughout the year. Likewise, GSI for males was constant at the 0.5% level, with minor fluctuations, and did not show any clear relationship with the HSI as for female GSI. Females had higher hepatosomatic indices but the soma weighed less than males throughout the year. Again in May, medium-sized females had the highest GSI followed by large and then small-sized individuals. As the HSI steadily decreased, GSI increased with advancing egg development stages for ovigerous females. The quantitative stomach analyses showed that females had higher stomach quantity indices (SQI) in May than in September, when the breeding period had ended. Contrary results were observed for males. Also, female SQI in May was higher than that for males.     

Nocturnal feeding migrations of Nerita plicata,N. undata and N. textilis (Prosobranchia: Neritacea) on the rocky shores at Mkomani,Mombasa, Kenya

The vertical zonation of the three common rocky shore neritids at Mkomani, Mombasa, Kenya, Nerita plicata Linnaeus, N. undata Linnaeus, and N. textilis Dillwyn, as a function of feeding migrations and of size, was studied from 28 February to 24 March 1983. These snails perform feeding migrations at night starting at around mid-ebb tide and return to their resting positions with the flood tide. They remain in their resting positions throughout the day until the next nocturnal ebb tide. The direction of migration is sizerelated, with the larger snails of each species moving in the opposite vertical direction to the smaller ones, so that the populations as a whole exhibit no statistically significant net vertical displacement. The larger individuals of two of the species, N. plicata and N. undata, invariably move downwards to their feeding levels, while the smaller individuals move upwards; the larger individuals of N. textilis display a different pattern of migration, moving downwards on and around spring-tide days and upwards on and around neap tide days, while the smaller individuals move in the opposite directions. N. textilis rest above their feeding level around spring tides, and below that level around neap tides. It is demonstrated how these nocturnal migratory feeding rhythms are integrated into the spring-neap and seasonal cycles of the snails' daytime resting positions. The adaptive significance of these migrations is also discussed.     

Geographic,population, and seasonal patterns in the reproductive parameters of the sea urchin <Emphasis Type="Italic">Paracentrotus lividus</Emphasis>

Generalized additive models were used to synthesize the data from 52 publications on the gonad growth and spawning seasonality of Paracentrotus lividus and identify spatial and temporal patterns in the reproductive processes. According to our results, Atlantic populations spawn in spring and develop large gonads (6.4 ± 0.2% of fresh body weight (mean ± SE)), which increase in size toward higher latitudes. While in the Mediterranean, the gonads are smaller (3.1 ± 0.1%), possibly as the result of successive spawnings which impede nutrients accumulating in the gonads. Differences were also observed between habitats: gonad production was higher in seagrass meadows in the Mediterranean and in subtidal rocky substrates in the Atlantic. Food availability might cause these variations as well as the negative correlation observed between depth and the gonad index for rocky substrates. Sex and body size do not seem to influence greatly gonad growth.     

Influence of season,ontogeny and tide on the diet of the temperate marine herbivorous fish Odax pullus (Odacidae)

The diet of the temperate marine herbivorous fish Odax pullus (Pisces: Odacidae) was examined using gut-content analysis followed by principal-component analysis and analysis of variance. Fish were collected near Leigh, on the northeast coast of New Zealand, from February 1983 to September 1984. The data were categorised by size of fish, season, and state of tide. A major ontogenetic shift in diet was observed. Juveniles fed on animal material (crustaceans and gastropods), epiphytic rhodophytes, and some phaeophyte material. Adults fed almost exclusively on two phaeophyte taxa, the laminarian Ecklonia radiata and the fucoid Carpophyllum spp. Dietary selection was evident at the level of particular plant components; the diet of larger fish in spring samples was dominated by fucoid reproductive receptacles. There was no direct evidence of a tidal influence on diet composition of O. pullus, although the relative proportions of E. radiata and Carpophyllum spp. ingested by adult fish appeared to vary with tidal state. The volume of gut contents varied seasonally for all size classes, although the timing of peak annual food intake varied among size classes. Tidal state did not influence the volume of the gut contents. The ontogenetic and seasonal trends evident in the diet of O. pullus appeared to be related to a number of factors including relative gut length, gonad development, ontogenetic changes in feeding anatomy and seasonal changes in algal composition.     

Seasonal variations in biochemical constituents during the reproductive cycle of the female dog cockle Glycymeris glycymeris

In an attempt to describe the biochemical events associated with the main stages of the annual and reproductive cycles of the female dog cockle Glycymeris glycymeris L., we studied seasonal variations in the various stages of oocyte development of the ovaries, and the glycogen, total protein and total lipid content of five body tissues – adductor muscle, foot, tunic coat, visceral mass and mantle. From November 1991 to November 1994, microscopic examination of the ovaries and measurement of the tissue concentrations of glycogen, total proteins and total lipids in these five body tissues were made monthly on ten female dog cockles originating from the sea area around Douarnenez (south Brittany, France). Morphological studies revealed that in the population investigated the annual cycle is characterised by three major periods: a first period of vitellogenesis extending from February/March to April/May and preceding a spawning in spring; a second period of vitellogenesis extending from May/June to September/October and leading to either no spawning, a single autumnal spawning event, or to two spawning events in summer and autumn; and a third period extending from October/November to February/March and characterised by a high level of oocyte lysis. In the muscular body tissues of the dog cockle, i.e. the adductor muscles, the foot and the tunic coat (the muscular envelope containing the visceral mass), the concentrations of glycogen, total proteins and total lipids underwent very similar variations during the annual cycle. During each stage of vitellogenesis, a typical glycogen–protein–lipid sequence was observed in the muscular tissues that was characterised firstly by a peak of glycogen concentration 2 to 3 mo before spawning, followed by a peak in total proteins 1 mo before spawning, and finally by a peak in lipid content just before spawning. A similar glycogen–protein–lipid sequence was also recorded in the first half of the winter period. However, these events were followed by general atresia affecting all oocytes in the gonad. Maximum energetic value of biochemical constituents in females coincided with peaks in lipid content in the visceral mass and mantle. These biochemical events occurred principally immediately before and at the end of oocyte lysis (December/January). A drop in the total energetic value, affecting mainly the visceral mass and the mantle, was recorded each year during the period January to March, coinciding with the period of shell growth in this species. Our data clearly indicate that in female G. glycymeris all muscular tissues contribute to the storage of glycogen and proteins, and suggest that glycogen may be the source of energy triggering vitellogenesis. Biochemical and microscopic observations revealed that oocyte development takes place during the first half of winter, but that these oocytes undergo atresia in December/January. The metabolites produced from oocyte lysis could contribute to somatic growth, which occurs in late winter. Received: 3 March 1997 / Accepted: 23 July 1997     

Seasonal moulting patterns and the generation cycle of<Emphasis Type="Italic"> Calanus finmarchicus</Emphasis> in the NE Norwegian Sea,as inferred from gnathobase structures,and the size of gonads and oil sacs

The seasonal moulting pattern of Calanus finmarchicus, related to body size, oil content and depth distribution, was investigated on the Norwegian midshelf from February to October 1997. Both gonad and gnathobase development were well correlated with time, and proved to be good indices for tracing the moulting cycle. The growth and development of gonads preceded the formation of new teeth. Both gonad development and tooth formation patterns in overwintering stage V copepodites (CV) indicated that the most intensive moulting occurred in spring. In the upper 200 m layer, the proportion of copepods in premoult phases ranged from 10% at the beginning of February, 60–95% from the end of February through April, to 4–7% in mid-May. No significant latitudinally determined variations in the moulting pattern were observed. The upward migration of deep-dwelling overwintering CVs was not synchronous and was determined by the readiness to moult. In April, no CVs were found at depths >300 m, that may indicate the termination of the overwintering moulting from G_o. By the end of May, the large-sized, lipid-rich CVs of G₁ started descending to depth. The second period of moulting (i.e. most likely the formation of G₂) was only recorded to have taken place in the surface (0–200 m) component of the CV stock from July onwards. This group was estimated to make up 10–20% of CVs inhabiting the upper 200 m layer and to have originated from the smallest sized fraction of CVs. There was a consistent trend towards size-differential moulting from CVs, in both spring and summer. The large-sized CVs moulted into adults earlier in spring compared with the smaller ones. By mid-June, the large and lipid-rich CVs of G₁ disappeared from the uppermost layers, sinking to depths. In summer (July onwards), the moulting of small CVs is probably the main contributor to G₂ copepodites that completed the cycle through to CIV/CV by the end of October.Communicated by O. Kinne, Oldendorf/Luhe     

Geographic variation in development rate between populations of the teleost Fundulus heteroclitus

The teleost Fundulus heteroclitus (Linnaeus) employs a semilunar spawning strategy, whereby eggs deposited on a spring tide optimally hatch on the following spring tide. This spawning strategy constrains the development period to ≃2 wk, regardless of the mean water temperature, which varies throughout the species' range (Nova Scotia, Canada, to Florida, USA). We hypothesized that F. heteroclitus embryos would exhibit development-rate compensation among populations to ensure appropriate hatching during the spring tide. Development rates of embryos from Massachusetts and Florida were examined. Northern embryos had faster temperature-specific development rates than southern embryos, while data from the literature showed that embryos from Delaware have an intermediate development rate. Results from reciprocal hybrid crosses indicated that there may be a genetic basis for these differences. In addition, northern embryos were more cold-tolerant and southern embryos more heat-tolerant. Field studies showed that embryos in their local environments develop at similar rates despite large temperature differences. Embryos seldom, if ever, experience lethal temperatures in their native habitats, but would do so if exposed to the temperature regimes at the extreme ends of the species' range. Thus, F. heteroclitus populations along the Atlantic coast have specifically adapted their development rates and thermal tolerances to the local thermal regime. Received: 2 September 1996 / Accepted: 17 September 1996     

The reproductive cycle of the sea urchin Arbacia lixula in northwest Mediterranean: potential influence of temperature and photoperiod

We studied the reproductive cycle of the sea urchin Arbacia lixula in a subtidal population from northeast Spain over four years using a gonadosomatic index (GSI) and gonad histology. Our results show that the GSI of A. lixula follows a seasonal cycle which peaks in May–July and attains its lowest values in October–November every year. The time course of the GSI matched closely the photoperiod cycle. We also found a remarkable inter-annual variability in the maximum value of GSI, which correlated with mean water temperature during the gonad growth period (winter and spring). Gonad histology was also in agreement with a single gametogenic cycle per year in this species. We explored the application of circular statistics to present and analyse gonadal development data, which allowed us to adequately handle the high intra-individual variability detected, with several developmental stages commonly found within the same gonad. The picture that emerged is one of a gametogenic timing driven by photoperiod, while the amount of reproductive output is determined by temperature. This is coherent with the tropical origin of the species and lends support to recent warnings about an increase in the abundance of this species in the Mediterranean as a result of global warming, with associated increased impact potential in sublittoral communities.