Article

Growth and reproduction of Megabalanus tintinnabulum (Crustacea: Cirripedia) in coastal waters of Pakistan, North Arabian Sea

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Abstract

The growth and reproductive/ brooding pattern of Megabalanus tintinnabulum were studied from January 2012 to December 2013 on two rocky shores, Buleji and Manora, Karachi, the northern Arabian Sea. Population parameters were estimated using the von Bertalanffy growth function (VBGF) and ELEFAN 1 with the aid of FiSAT software. The population of M. tintinnabulum showed variability in density, size and growth rates between two rocky shores. The average density of M. tintinnabulum was higher at Buleji (101.9 ± 15.02 individual m⁻²) than Manora (76.0 ± 8.05 individual m⁻²). Though the estimated asymptotic length (L∞) of M. tintinnabulum was 51.98 mm at both sites, the growth coefficient (K) value was higher at Manora (0.63 year −1) than Buleji (0.44 year −1). The population of M. tintinnabulum grew at a faster rate attaining the length of 31.8 mm in one year at Manora as compared to length of 23.7 mm at Buelji. Longevity in M. tintinnabulum was 4.3 years at Manora and 6.4 years at Buleji, while the natural mortality rate (M) was 1.131 year−1 and 0.894 year−1 at Manora and Buleji, respectively. Though mature ovaries and testes were found throughout the year but the brooding individuals were observed during the winter (November to February) period showing that lower temperatures (20-24 ºC) are preferred over higher temperatures (29-34 ºC). The brooding capacity was same at two rocky shores but the numbers of brooders were higher at Manora than Buleji. The reproductive capacity was 7,914 ± 303 of eggs per brood.

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... Among the more straightforward traits to describe, the population size structure is underreported for barnacles in general. For Megabalanus, there are only a few accounts of the size structure of populations on natural substrates (Ali and Ayub 2021), with no reports from the endemic regions range of M. coccopoma, given that the endemic range of M. tintinnabulum is not known. ...
... Data on the maximum size of species is also elusive. For instance, for M. tintinnabulum, there are only three documented measurements by Karande and Palekar (1966), Annadale (1911) and, more recently with Ali and Ayub (2021) Estimations of growth in historical records are often based on the size of organisms after known events. For instance, Darwin (1854) deduced the growth of M. tintinnabulum from Samuel Stutchbury's observation of specimens approximately 5 cm in basal diameter on a vessel at sea for only a year. ...
... Apolinario (2001Apolinario ( , 2003 investigated the recruitment and reproduction of M. coccopoma and M. tintinnabulum in southeastern Brazil. Ali and Ayub (2021) explored growth and reproduction of M. tintinnabulum in Pakistan. For Megabalanus azoricus (Pilsbry 1916) Pham et al. (2011) described the recruitment, growth, and Dionisio et al. (2007) its reproductive biology, respectively. ...
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A demographic model for an open population with space-limited recruitment has been previously proposed and tested using the barnacle Balanus glandula on a single shore in California. The model makes a number of qualitative predictions about population structure. At small scales, areas of high settlement should have on average less free space and fewer cohorts. At a larger unspecified scale (>34.6 cm(2)) cohort structure should be more variable in areas of high settlement. We attempted to test these qualitative predictions on a number of different spatial scales using the barnacle Semibalanus balanoides. A recruitment gradient was defined for S. balanoides around the coast of Anglesey, Wales, with the north having lower recruitment than the south regardless of year. Given this recruitment gradient, a nested ANOVA was used to examine the qualitative predictions of the demographic model at different scales. The qualitative predictions of the demographic model were not generally supported by an analysis of data from Anglesey. Free space and small-scale population variance did not vary between areas of different recruitment density. Barnacle size variation was greater on high recruitment shores at an intermediate scale (100 m) as predicted by the model. However, this difference was not statistically significant. Even at the scale of individual 5 x 5 cm photographs there was no evidence of a positive relationship between barnacle size variance and free space. Possible reasons for the mismatch of the demographic model predictions and empirical observations are discussed. There may be insufficient spatial sampling, insufficient recruitment difference, large-scale variation in mortality and growth rates, or the model assumptions may be incorrect. One important difference between the 2 levels of recruitment is that low recruitment sites had a constant patch size below 25 cm(2), whereas the high recruitment sites did not. If patch scale is variable, the predictions of the demographic model will be irrelevant as there is no single scale at which they can be observed. Use of the nested ANOVA technique with logarithmic spatial scales also allows hypotheses to be made about the important spatial scales for future demographic studies using manipulative experiments. For population variance, free space and opercular length the important scales were small (<1 m), meso- (100 m), and large scales (>10 km).
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There are approximately a dozen species of commercially interesting barnacles worldwide, some of which have been cultured on a semi-industrial scale. These species are listed and information is provided with regard to geographical distribution, landings and prices. Traditionally, ‘goose’ barnacles (four species) are considered to be the most important for consumption. World production already stands at 500 tonnes year−1, but this species has not been cultured to date. Some ‘acorn’ barnacles are also consumed (seven species), with harvest levels per species that do not exceed 200 tonnes year−1 and selling prices that can reach US$17/kg. ‘Acorn’ barnacle culture on a world scale is still developing. Nevertheless, production has occurred on a semi-industrial scale; specifically, spat have been collected from the wild and grown in suspended systems. Farming trials have focused on two species of acorn barnacles: Austromegabalanus psittacus (Molina 1782) ‘picoroco’ in Chile and Megabalanus azoricus (Pilsbry 1916) ‘craca’ in Portugal. The large-scale production of these crustaceans will depend on the optimization of spat collection from the wild and/or the parallel development of mass production technologies for larvae (hatcheries). In addition, further development will be achieved by opening up new markets for commercialization.
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Tetraclita squamosa and Tetraclita japonica are common, intertidal barnacles. In Hong Kong, the population dynamics of the two species exhibited spatial and temporal variation on two semi-exposed shores. T. squamosa produced egg masses from May–June and annual settlement and recruitment occurred from June–July. In contrast, settlement and recruitment of T. japonica was sparse from March–May, but intensified in July–October. Mature gonads and egg masses were, however, only present from September–November, suggesting the larvae from the two settlement pulses originated from other locations as well as Hong Kong populations. Settlement intensity and post-recruitment mortality of the two species varied between sites, possibly due to spatial variation in free rock space, physical transport of larvae and abundance of food in the water column.
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Balanus amphitrite, an acorn barnacle, is distinctly euryhaline, eurythermal and a dominant fouling organism found in warm and temperate waters throughout the world. In this study, the influence of temperature and food concentration on the reproductive biology of this species collected from a tropical habitat was evaluated. Adult barnacles were maintained at 20, 25 and 30°C temperatures at different concentrations of food (50, 100, 150 and 200 Artemia ind−1day−1). In this previously believed obligatory cross-fertilizing hermaphrodite, self-fertilization was observed. The rise in temperature from 20 to 30°C resulted in a longer interbreeding interval (6–7days, 200 Artemia ind−1day−1; 11–13days, 50 Artemia ind−1day−1). Computed carbon gained through feeding during the interbreeding interval indicated an inverse relationship to the temperature. At 20°C, although a greater amount of carbon was gained through feeding, the numbers of larvae produced were fivefold less when compared to those raised at 30°C. At 20°C, 2.3μg C was required to produce a single larva, whereas at 30°C it was 0.4μg C. A rise in rearing temperature also influenced the molting rate positively. Observations on temporal variation in the gonad development of this species in a tropical coastal environment influenced by the monsoons indicated gonad development to be positively related to chlorophyll a concentration.
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The present work addresses the effect of environmental factors (icing, water temperature, food availability) on the ecology of the patellid limpet Nacella (Patinigera) concinna, in a bay on the Antarctic Peninsula. Sampling was conducted at three depths (intertidal, 5m, 10m) from February 1987 to January 1988. Temperature was recorded and concentrations of Chlorophyll a were measured on the bottom, in the water and in the ice-water layer. The limpets were measured, weighed and a condition coefficient for somatic and gonadal mass was calculated. Their ages were estimated through size frequency distribution analysis and a seasonalized von Bertalanffy growth model was applied. The intertidal subpopulation migrated to deeper levels at the beginning of the icing season and recolonized the intertidal zone after ice retreat. The growth rate was very low (von Bertalanffy K 0.08). Growth rates showed important seasonal variations, with maxima during December and January. Nacella (P.) concinna spawns once a year and spawning coincided with raising water temperature (from -1.33C to-0.84C), and probably was also related to increasing spring food availability. Body mass increased during periods of high standing stock of microphytobenthos, revealing that ice-algae and phytoplankton were of minor importance as food sources for limpets in Esperanza Bay.
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The golden mussel, Limnoperna fortunei, is an invasive freshwater bivalve. Since its introduction to Argentina, it had caused damage to the native fauna as well as economic damage to industries of the region. Here, we describe the growth of L. fortunei in a natural temperate environment in Argentina. Age was estimated according to the modal progression method. The constants in the von Bertalanffy growth model were adjusted by an iterative algorithm. Three annual cohorts had similar growth rates. The estimated t 0 for each cohort showed a temporary displacement in relation to the spawning period.
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1. A method of producing a pre-arranged pattern and density of barnacle settlement is described, which makes use of the tendency for barnacle cyprids to settle in small pits. The pits had a negligible influence on growth-rate after the first 2 weeks. This method was used to study the influence of water flow, population density and orientation on the rate of growth of B. balanoides in the Menai Straits, North Wales. 2. Barnacles parasitized by Hemioniscus balani grew more slowly than unparasitized individuals. 3. The passage of an uninterrupted current of water over the barnacles was the most important factor in promoting rapid growth. 4. Population density appreciably influenced growth in volume only when the individuals were closely packed, either touching or nearly touching. Even at densities as high as 4 individuals per cm2, much of the loss in cross-sectional area caused by space limitation was compensated for by increase in height. The effect of close packing in itself did not appear to be harmful. 5. In a unidirectional current, there was little evidence that the leading individuals in a group grew appreciably faster than the trailing ones. 6. Orientation with the rostrum pointing to the current source conferred an advantage resulting in slightly enhanced growth. 7. It is shown that a barnacle whose growth is isometric and determined only by the rate of intake of food from a current of constant nutrient content would grow at a constant linear rate. This is approximately true for the greater part of the growth curve of the species investigated. Early growth just after metamorphosis is slower, probably because the rate of water flow is reduced in the immediate vicinity of the solid substratum. Growth slows down after maturity in all species as a consequence of metabolic changes.
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1. The reproductive cycles and brood production of Chthamalus fissus, Balanus glandula and Tetraclita squamosa from central California are compared. C. fissus produces about 16 small broods from March through October. B. glandula produces three to six relatively large broods from December or January to May. T. squamosa incubates only about three intermediate-sized broods from June through September.2. Brooding in C. fissus is regulated by food availability, and yolk for no more than one brood is stored at a time. Feeding in the laboratory elicited high brooding frequencies during periods when brooding activity and food levels in the field were low, and the frequency of brooding was directly proportional to the size of the food dosage. Temperature and photoperiod did not affect brooding frequencies. B. glandula rapidly stores nutrients in the ovary for about three broods during summer. Cold temperatures induce early brooding in the laboratory during late fall and early winter, and the population in the war...
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The yellow clam, Mesodesma mactroides, is an intertidal bivalve typical from sandy beaches of the South American Atlantic coast. Growth parameters of southernmost populations of M. mactroides were studied and compared with other populations. Thin shell sections were examined to describe internal shell layers and to contrast with external shell transparency. Periodicity of deposition of external growth increments was studied recording the degree of transparency of the shell border. Growth patterns were determined using modal progression analysis from size frequency distributions, analysis of external shell increments, and size-at-age data derived from inner shell layers. Growth parameters were described using the von Bertalanffy growth model. Both internal and external patterns were coincident and exhibited a succession of one translucent and one opaque region. The transparent region was deposited during summer. Growth differences found between populations may be related to unequal size of first ring in both beaches. This feature may originate from asynchrony in spawning and recruitment. The monthly analysis of shell length size frequency distribution shows that growth of M. mactroides is seasonal. Estimations of asymptotic size of studied populations and others located at the southern (coldest) half of the geographical range of distribution suggest a negative relation with latitude.
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Marked seasonality of ship-observed chlorophyll concentrations in the upper layers of the central and northern parts of the Arabian Sea is established for three hydrographically defined offshore areas. A peak related to the southwest (summer) monsoon occurs, either with or without a northeast (winter) monsoon-related peak. Both appear to depend on the establishment of a deep mixed layer and the concomitant nutrient supply, but at any station the relation between pigment content and mixed-layer depth or nitrate concentration is usually obscure.
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The reproductive cycles of two intertidal barnacles, Chthamalus stellatus (Poli) and C. montagui Southward, were studied at Lough Hyne Marine Reserve, Cork, Ireland, over 28 months. In both species there were seasonal trends in development of the male and female reproductive organs with, for example, the ovaries regenerating while eggs were being brooded. Breeding occurred mainly during June–August but also at other times. The sizes of the ova and eggs of the two species were compared and the timing of the cycles relative to other areas considered.
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Adult Tesseropora rosea (Krauss) and Tetraclitella purpurascens (Wood) are mostly found in the eulittoral (barnacle) zone of rocky seashores in New South Wales. Below this zone most space is occupied by the tube-worm Galeolaria caespitosa (Lamarck) or by various species of macroalgae. Within the eulittoral zone, T. rosea are mostly on sunny areas of rock exposed to relatively strong wave-action. T. purpurascens are present mainly in crevices, caves, and under ledges where there is considerable shade.Cyprids of both species settled on sandstone plates and on experimentally cleared areas in the barnacle and Galeolaria zones. Neither species settled where the substratum was already covered by algae or Galeolaria. No spat of T. purpurascens were found in sunny areas of the barnacle zone. T. rosea, however, settled in cleared substrata in sunny and shaded areas. Neither species settled in the littoral fringe above the upper limit of distribution of adults. On boulders transferred to high levels of the shore during a storm, small T. purpurascens died within a few weeks.Barnacles of both species which had settled in experimentally cleared areas in the Galeolaria zone survived and grew. In these areas some T. purpurascens were killed by being smothered by tube-worms which settled after the barnacles. This probably happens to T. rosea, but was not demonstrated experimentally. In the Galeolaria zone, both species of barnacles were very quickly smothered and killed by macroalgae growing over them, except where these were experimentally removed.Within the barnacle zone, all newly-settled spat of T. purpurascens transferred to sunny sites died within two months, whilst many of those in shaded sites survived. In areas where wave-action was strong, spat of T. rosea survived and grew well in sunny areas, but survived better in the shade. Under a ledge, however, where wave-action was reduced, all the T. rosea in sunny sites, and most of those in shaded sites died within two months; many newly-settled T. purpurascens survived in the shade in this area.The grazing limpet Cellana tramoserica (Sowerby) dislodged and crushed some newly-settled T. rosea and reduced survival in some sunny areas. T. rosea settled preferentially on bare rock and were rarely found on the shells of adult barnacles. Thus, the density of spat was greater where adult barnacles were absent. In contrast, many newly-settled T. purpurascens were found on the shells of adults of their own species in shaded areas; they also settled on cleared rock. Because T. purpurascens tended to settle amongst and on adults, and in crevices and confined areas, they were not much affected by limpets. When newly-settled T. purpurascens were in high densities, they had lower survival than in areas with reduced densities, because of squashing and smothering by each other.The upper and lower limits of vertical distribution (zonation) of these two species of barnacles are determined primarily by the settlement of cyprids. Neither species settled at the highest levels on the shore. Whether this was due to the decreasing time of submersion during high tide towards the top of the shore, or a result of preferences for settlement site is unknown. Even if cyprids were to settle in the littoral fringe, the spat would die very quickly probably as a result of desiccation. Below the barnacle zone, the entire substratum is usually occupied by other sessile species, particularly macroalgae, on which the barnacles do not settle. In experimentally cleared areas below the barnacle zone, or in any naturally cleared areas both species settled, and could survive the physical conditions. Newly-settled spat were, however, overgrown and killed by algae and Galeolaria.Within the barnacle zone, T. purpurascens is restricted to shaded areas because of the inability of newly-settled spat to survive the physical stresses of high temperature and desiccation in sunny habitats. T. rosea appears to be excluded from shaded areas by a combination of the lack of suitable substrata on which to settle, and the effects of reduced water-flow in many crevices and under ledges. T. rosea survived better in areas with strong wave-action and can survive in shaded areas where water-flow is not reduced by the topography of the substratum.
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[The effect of temperature on breeding activity and moulting of the tropical barnacle, Chthamalus malayensis was studied under rearing conditions in the laboratory. Both breeding activity and moulting frequency were influenced by temperature. Temperature seems to be the main external factor regulating both breeding and moulting in C. malayensis in Hong Kong. The temperature above which breeding begins in C. malayensis agrees well with the natural breeding season. L'effet de la température sur l'activité de reproduction et la mue du cirripède tropical, Chthamalus malayensis a été étudié dans des conditions d'élevage au laboratoire. L'activité reproductive et la fréquence de mue étaient influencées toutes deux par la température. La température semble être le principal facteur externe régulant à la fois la ponte et la mue chez C. malayensis à Hong Kong. La température à laquelle la reproduction commence chez C. malayensis s'accorde bien avec la saison naturelle de reproduction., The effect of temperature on breeding activity and moulting of the tropical barnacle, Chthamalus malayensis was studied under rearing conditions in the laboratory. Both breeding activity and moulting frequency were influenced by temperature. Temperature seems to be the main external factor regulating both breeding and moulting in C. malayensis in Hong Kong. The temperature above which breeding begins in C. malayensis agrees well with the natural breeding season. L'effet de la température sur l'activité de reproduction et la mue du cirripède tropical, Chthamalus malayensis a été étudié dans des conditions d'élevage au laboratoire. L'activité reproductive et la fréquence de mue étaient influencées toutes deux par la température. La température semble être le principal facteur externe régulant à la fois la ponte et la mue chez C. malayensis à Hong Kong. La température à laquelle la reproduction commence chez C. malayensis s'accorde bien avec la saison naturelle de reproduction.]
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An account is given of the stomach contents of some common barnacles; these include both intertidal and sublittoral species. The contents reflect to some extent the environment, species taken from the same place at the same time having similar contents. With some species, a particle size analysis was possible on a pooled sample of stomach contents. It is evident that much debris is present, and that particulate matter with a diameter less than 2 μ can be taken. Detritus may be an important source of nutriment with sublittoral species. Small particles cannot be taken by the captorial activity of the cirri. It is considered that glandular secretions at the base of the cirri, where there are numerous setae, enmesh the food, which is then carried forward to the mouth parts to be rejected or accepted.
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A new computer-based method is presented for the "tracing" of growth curves through fish length-frequency samples sequentially arranged in time. The method allows for an objective extraction of growth parameters whenever representative samples are available from a fish population. The method is suited both to the analysis of growth in tropical stocks and in temperate stocks, where it oscillates seasonally. A full program listing with user's instructions is available from the authors.
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Long-term fluctuations (3-9 years) in the abundance of the barnacle Chthamalus dentatus were determined from archival photographs of permanent 0,25 m2 quadrats in the mid-littoral at 11 sites along the Transkei coast. These data were combined with a more detailed analysis of growth and recruitment over one year. Recruitment occurred in spring and autumn, but periodogram analyses revealed additional cycles with frequencies of 24-36 months. There was no geographic pattern in long-term differences in abundance between sites nor were these related to human disturbance. Recruitment intensity was significantly related to mean abundance of adults, which may indicate limited larval dispersal or gregarious settlement. Maximum aperture length increased by 3,0 mm in the first year but increased by only 1 mm·year−1 in the third year. Individual growth was variable, with small individuals often overtaking larger ones. There were few large individuals, those of 7,0 mm or more constituting only 3% of the population. An analysis of longevity indicated a mean lifespan of 1,5 + 0,5 years.
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The distribution of the warm-water barnacle, Balanus perforatus, was surveyed along the south coast of England and the north-east coast of France between 1993 and 2001, repeating work carried out between the 1940s and 1960s. The species has recovered from catastrophic mortality during the severe winter of 1962–1963 and was found over 120 km (UK) and 190 km (France) east of previous records on both sides of the Channel. The presence of the species in the eastern Channel refutes suggestions in the 1950s that larvae, and hence adults, would not be found east of the Isle of Wight because of reproductive sterility close to the limits of distribution. Brooding of specimens translocated to Bembridge, Isle of Wight, commenced in May, earlier than previously observed in British waters, and continued until September. The stage of embryo development at Bembridge in mid-August was comparable to that of the large population at Lyme Regis, Dorset 100 km further west. However the size of brood per standard body weight was greater at Lyme Regis. Factors influencing the rate of colonization and further geographic range extension of the species as a possible result of climate change, are discussed.
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The barnacle Balanus glandula is predominantly an open coast species in the Northeast Pacific. However, B. glandula densely inhabits estuaries where environmental conditions such as salinity and temperature drastically differ from the open coast. The increased variability of environmental conditions within an estuary can potentially affect spatial patterns of reproduction in B. glandula. I examined gonad maturity, reproductive periodicity and fecundity, and then calculated reproductive output in B. glandula populations distributed along an estuarine gradient. Results indicated that reproductive output of this intertidal barnacle decreased four times over a spatial scale of kilometers, as a transition occurred from a marine to a freshwater habitat along an estuarine gradient. Additionally, a higher proportion of the population had well-developed gonads in the oceanic end of the estuary compared to the riverine end. These results indicate how reproductive pattern can significantly vary over a spatial scale of kilometers, resulting in site-specific contributions of offspring to the larval pool.
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This study presents a historical review, a meta-analysis, and recommendations for users about weight–length relationships, condition factors and relative weight equations. The historical review traces the developments of the respective concepts. The meta-analysis explores 3929 weight–length relationships of the type W = aLb for 1773 species of fishes. It shows that 82% of the variance in a plot of log a over b can be explained by allometric versus isometric growth patterns and by different body shapes of the respective species. Across species median b = 3.03 is significantly larger than 3.0, thus indicating a tendency towards slightly positive-allometric growth (increase in relative body thickness or plumpness) in most fishes. The expected range of 2.5 < b < 3.5 is confirmed. Mean estimates of b outside this range are often based on only one or two weight–length relationships per species. However, true cases of strong allometric growth do exist and three examples are given. Within species, a plot of log a vs b can be used to detect outliers in weight–length relationships. An equation to calculate mean condition factors from weight–length relationships is given as Kmean = 100aLb−3. Relative weight Wrm = 100W/(amLbm) can be used for comparing the condition of individuals across populations, where am is the geometric mean of a and bm is the mean of b across all available weight–length relationships for a given species. Twelve recommendations for proper use and presentation of weight–length relationships, condition factors and relative weight are given.
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Aim Our aim in this paper is to present the first broad‐scale quantification of species abundance for rocky intertidal communities along the Pacific coast of North America. Here we examine the community‐level marine biogeographical patterns in the context of formerly described biogeographical regions, and we evaluate the combined effects of geographical distance and environmental conditions on patterns of species similarity across this region. Location Pacific coast of North America. Methods Data on the percentage cover of benthic marine organisms were collected at 67 rocky intertidal sites from south‐eastern Alaska, USA, to central Baja California Sur, Mexico. Cluster analysis and non‐metric multidimensional scaling were used to evaluate the spatial patterns of species similarity among sites relative to those of previously defined biogeographical regions. Matrices of similarity in species composition among all sites were computed and analysed with respect to geographical distance and long‐term mean sea surface temperature (SST) as a measure of environmental conditions. Results We found a high degree of spatial structure in the similarity of intertidal communities along the coast. Cluster analysis identified 13 major community structure ‘groups’. Although breaks between clusters of sites generally occurred at major biogeographical boundaries, some of the larger biogeographical regions contained several clusters of sites that did not group according to spatial position or identifiable coastal features. Additionally, there were several outliers – sites that grouped alone or with sites outside their region – for which localized features may play an important role in driving community structure. Patterns of species similarity at the large scale were highly correlated with geographical distance among sites and with SST. Importantly, we found community similarity to be highly correlated with long‐term mean SST while controlling for the effects of geographical distance. Main conclusions These findings reveal a high degree of spatial structure in the similarity of rocky intertidal communities of the north‐east Pacific, and are generally consistent with those of previously described biogeographical regions, with some notable differences. Breaks in similarity among clusters are generally coincident with known biogeographical and oceanographic discontinuities. The strong correlations between species similarity and both geographical position and SST suggest that both geography and oceanographic conditions have a large influence on patterns of intertidal community structure along the Pacific coast of North America.
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The effect of geographical differences in breeding cycles on the recruitment variation of the limpets — Patella vulgata L., P. aspera Rding and (to a limited extent) P. depressa Pennant — has been studied over much of their range in Britain and NW. Europe. In spite of considerable annual and local variation in recruitment success, broader patterns can be distinguished, which can be linked to spawning times and factors affecting the survival of newly-settled spat. The breeding cycles of P. vulgata and P. aspera differ across their ranges in that, in both species, spawning begins, and gametogenesis ends, earlier in the north and east than in the south and west. The cause of these differences can be correlated with geographical and annual differences in sea temperature over the potential breeding periods, and can be related to the regional incidence of conditions found experimentally to be necessary for successful settlement and survival of spat during a critical stage of their growth. The significance of this temperature window in determining the littoral and geographical distribution of the species is discussed.