The ecology of two sandy beaches in south west India. II. Notes on Emerita holthuisi

The mole crab Emerita holthuisi Sankolli occurred on two beaches, at Shertallai and at Cochin, in southwest India. Observations on the size-frequency of samples for Shertallai suggest that there were two main periods of recruitment, one in the pre-monsoon,months of February and March, and one during the monsoon, although there may have been some recruitment throught the year. At Cochin, E. holthuisi was present in the pre-monsoon period, but disappeared from the beach during the monsoon.     

A preliminary account of two sandy beaches in South West India

Observations were made on 2 beaches during the premonsoon period (January to March) when conditions were relatively stable. The beach below the berm at Cochin is 14 m wide with a slope of 1 in 8.5, and at Shertallai 40 m wide with a slope of 1 in 16. The well-sorted sand has a mean particle size of 275 at Cochin and 175 at Shertallai. The amount of chlorophyll attached to sand grains is very small. Chlorophyll in the water varies between 0.9 and 13.0 g/l. Carbon in the sand amounts to 210 to 630 g/g, and in the water to 355 to 3660 g/l. Throughout observations, the water temperature remained about 31°C, but on the dry sand surface it reached 50°C. Fauna zonation on the 2 beaches is similar to that of other tropical beaches; crabs of the genus Ocypode and the isopod Eurydice sp. occur at high water mark, the polychaetes of the genera Glycera, Onuphis, Scoloplos and Lumbrinereis in the mid-intertidal region and below; the tidal migrants Bullia melanoides (Gastropoda), Donax incarnatus and D. spiculum (Bivalvia) and Emerita holthuisi (Crustacea) inhabit the wash zone of the surf. The average biomass is 0.73 g carbon/m² at Shertallai and 0.85 g C/m² at Cochin, where fewer species are present. This compares with a value of 0.63 g C/m² for a sandy beach on the west coast of Scotland.     

Indications of sodium,chloride, and water exchange across the gills of the striped dogfish Poroderma africanum

The ecology of a large sandy beach at Calangute, Goa, Central West India over a 22-month period is described from the point of view of changes in the distribution and abundance of the macrofauna, and of the physical environment of the beach; the dominant effect of the annual monsoon is clearly shown. The results from Calangute are compared with a nearby beach (Colva) and from earlier studies in Southern India. The biology of the dominant wedge-clam Donax incarnatus Gmelin is described, in particular its more rapid breeding-cycle and smaller size at Calangute, compared with other populations studied. The annual production of D. incarnatus has been calculated as 14.7 kcal/m²/year, and has been shown to be relatively low in comparison with other bivalve populations, due to high mortality and the extensive environmental disruption caused by the annual monsoon. It is suggested that the conditions at Calangute are most suitable for settlement, and that the small size of D. incarnatus here is related to the short life span of 6 to 7 months of all individuals.     

Observations on the biology of Donax trunculus and D. vittatus at Ile d'Oléron (French Atlantic coast)

Observations have been made on the biology of two species of the bivalve genus Donax, on two beaches, Vertbois and St. Trojan, on the Ile d'Oléron, French Atlantic coast. The two species displayed different vertical distribution: D. trunculus Linné in the intertidal; and D. vittatus (da Costa) between low-water spring tide (LWST) and 5 to 6m depth, with only the fringe of the population extending to the shore. D. trunculus also showed differential distribution by size or age, the youngest individuals being highest on the shore, and the oldest near LWST. The two species exhibited a similar pattern of seasonal recruitment and growth, but in both there were differences in growth characteristics between the two beaches, individuals at St. Trojan growing to a larger size than those at Vertbois. The mean pattern of mortality in the two areas was similar, and productivity per 100 recruits surviving to the first winter following settlement was greater at St. Trojan than at Vertbois. For D. trunculus the mean population density at Vertbois was higher and the production per unit area on the two beaches was similar. Production to biomass (B) and elimination to biomass (B) ratios were similar for the two beaches for both species, and while production was greater for D. trunculus at both beaches, the B and B ratios were higher for D. vittatus. Comparisons with other published data reveal that the rate of growth of D. trunculus is similar throughout its range from Brittany to the Mediterranean Sea, while that of D. vittatus shows more variation. In both species, there is a trend of increasing P/B and E/B ratios from north to south throughout the geographical range.     

The ecology of hydromedusae from the Cochin area

A survey has been made of medusae found in plankton samples taken fortnightly over a period of a year at 2 fixed stations in the waters near Cochin, Kerala, India. Twenty two species were identified, 8 of them were recorded for the first time in this region. The 2 stations were found to be controlled by different hydrographical regimes; that at Aroor is entirely characterised by backwater conditions, with salinity afternating from fresh water to 32 S according to the monsoon cycle; the station at Fairway Buoy is also dominated by the monsoon cycle, with mixed, low-salinity (11 to 13) waters at the surface, and upwelled Arabian Sea water in the bottom layer during the Southwest monsoon Life cycles are discussed and the population dynamics of Eutima commensalis at Aroor are analysed. A mathematical model is proposed for more detailed population dynamics studies of plankton species in the Cochin Backwaters, taking E. commensalis as an example.     

The ecology of two sandy beaches in south west India. I. Seasonal changes in physical and chemical factors,and in the macrofauna

Changes during one year in the fauna of two beaches in south India are described in relation to the nutrient content of the water and sand, and the physical changes brought about by the south-west monsoon. At both beaches, the most stable conditions were reached in the pre-monsoon months when the greatest number of species occurred and when there was marked zonation. During the monsoon, erosion took place and only actively migrating species remained to form a permanent element of the beach fauna. Data on population density and biomass indicate markedly seasonal recruitment and rapid growth of individual species, most of which are suspension feeders.     

The life cycle of Eutima commensalis sp. nov. (Eutimidae,Hydromedusae)

Developmental stages of the medusa Eutima commensalis sp. nov. were obtained from plankton collections made at 2 fixed stations, one at Fareway Buoy and the other at Aroor in the Cochin backwater system. Six stages were arbitrarily assigned and described. The hydroid and the medusa of this form were found to occur in these waters throughout the year, except during the monsoon months from June to September. The hydroid is endocommensalic on wood-boring molluscs such as Nausitora hedleyi, Teredo furcifera and Martesia striata.     

Biology,population dynamics and secondary production of<Emphasis Type="Italic"> Tylos europaeus</Emphasis> (Isopoda,Tylidae) on the western coast of Portugal

The biology, population dynamics, and production of Tylos europaeus were studied in two sandy beaches of the western coast of Portugal. At both sites, reproduction occurred seasonally, from April to July, with only one new cohort produced per year. Regarding population dynamics, cohort-splitting events were detected in males at the beginning of the reproduction period (April/May), resulting in two groups with distinct growth rates (fast-growing vs slow-growing males). Different biological characteristics were consequently detected in these two groups, namely regarding body size, lifespan, and contribution to the reproductive effort. Lifespan was estimated as approximately 3 years, for females and fast-growing males, and 4 years for slow-growing males. Cohort-splitting among males appeared as a possible strategy to cope with the highly male-biased sex ratios observed, which could lead to a strong male-male competition for mating. T. europaeus appeared as an annual species, with a univoltine life-cycle (one generation per year), and iteroparous females reproducing twice during their lifespan. Average growth production (P) was estimated at 0.082 g.m^–2.yr^–1 AFDW (ash-free dry weight) and the average annual biomass () (standing stock) at 0.052 g.m^–2, resulting in a P/ ratio of 1.58. These results produced baseline information for the construction of a population-dynamics model and highlighted the potential of this species as an environmental quality-assessment bioindicator on sandy shores.Communicated by S.A. Poulet, Roscoff     

Coexistence and reproductive isolation of the sympatric echinoids<Emphasis Type="Italic"> Diadema savignyi</Emphasis> Michelin and<Emphasis Type="Italic"> Diadema setosum</Emphasis> (Leske) on Kenyan coral reefs

The sympatric echinoids Diadema savignyi and D. setosum coexist in shallow reef lagoons throughout East Africa. The reproductive strategies of these echinoids were studied to investigate reproductive isolation as a possible mechanism for maintaining the coexistence of these closely related species. The annual reproductive cycle and lunar periodicity were determined by gonad index measurements, histological examination of gametogenesis, and induction of spawning with injections of KCl. The peak reproductive period of D. savignyi coincided with the north-east monsoon period (when light and temperatures are high) as gonad indices were high (>8%) beginning in February and peaked at 9.7% in May. Gonad indices subsequently rapidly decreased (by 26%) in June at the beginning of the cooler south-east monsoon period. However, the presence of sperm and ova in most months of the year indicates continuous gametogenesis with reduced reproductive effort during the cooler months. The annual cycle for D. setosum showed less of a seasonal trend as gonad indices remained above 7% throughout much of the year but tended to be highest when temperatures were lower. This is the first confirmation of continuous reproduction in these two species at the equator. The reproductive patterns of both species remained consistent over 2 years of sampling. Both species exhibited a synchronized lunar spawning periodicity during the 3 months sampled, with D. setosum spawning on lunar days 8–10 and D. savignyi spawning after the full moon (lunar days 17–18). Whereas spawning in D. savignyi was very tightly synchronized, 20% of D. setosum individuals still spawned after the peak spawning period. The coexistence of these closely related species appears to be maintained by temporal reproductive isolation during the lunar spawning period reinforced by seasonal differences in reproductive effort.Communicated by O. Kinne, Oldendorf/Luhe     

Population dynamics and production of the planktonic copepods in a eutrophic inlet of the Inland Sea of Japan. IV. Pseudodiaptomus marinus, the egg-carrying calanoid

Population dynamics and production of the egg-carrying calanoid copepod Pseudodiaptomus marinus were studied for a year in Fukuyama Harbor, a eutrophic inlet of the Inland Sea of Japan. This species was perennial, with a large numerical peak in June and small peaks in September/October and November/December. During the study period, at least 11 generations could be detected. For each generation, the stage-specific survival from egg to Copepodite Stage (C) V was determined; it was very high during early life stages (egg to NIII), and gradually decreased beyond. On average, 94% of eggs recruited into NIII, which is strongly contrasted with very high (>ca. 90%) mortality during the corresponding stages for free-spawning copepods, i.e. Acartia omorii, Centropages abdominalis and Paracalans sp. This demonstrates that the egg-carrying strategy has a great advantage to reduce mortality in egg stage. The biomass of this species showed marked seasonal variations largely in parallel with numerical abundance. The instantaneous somatic growth rate increased linearly with temperature. The population production rate was estimated as the sum of somatic growth of larval stages and egg production of adult females; the annual integration was 51.0 mg C m⁻³ yr⁻¹ or 0.38 g C m⁻² yr⁻¹. Received: 11 November 1996 / Accepted: 7 December 1996     

Growth kinetics and nutrient requirements of two tropical marine phytoplankters

Growth rates of nutrient-depleted cells of Biddulphia sinensis Greville and Ceratium furca (Ehrenberg) Claparède et Lachmann as a function of phosphate and nitrate, follow the hyperbolic expression of the Michaelis-Menten equation. variations in the half-saturation constants (K _s) for both species in each nutrient gave an index of their capacity to utilize these two nutrients when available either singly or in combination. The maximum growth rates (_max) of the two species did not follow the trends shown by K _s. Seasonal abundance of the two species in the Cochin Backwater (an Indian tropical estuary) showed that, at low concentrations of nutrients, C. furca becomes predominant; high nutrient concentrations present in the estuary during the monsoon months tend to increase the abundance of B. sinensis. The differences in the K _s values of the two organisms seem to reflect their observed seasonal abundance in the estuary.     

Population biology of the annual grassTriplasis purpurea in relation to distance from shore on Staten Island, New York

Four populations of the native annual grassTriplasis purpurea were surveyed on coastal beaches along the south shore of Staten Island, NY, to determine the potential of this species to colonize shoreline habitats mostly devoid of other vegetation. If the species can establish and maintain dense populations, it may have conservation value for urban beaches disturbed by human activities. For two populations, survivorship, growth, and reproduction were monitored at different distances from shore to determine the ability of this species to maintain viable populations. At three sites,T. purpurea occurred in >75% of all quadrats and the highest density was 1195 plants/m² at 74 m from shore in one recently disturbed site. Density generally increased with increasing distances from shore at low tide (from ca. 40 – 90 m). Plants showed the greatest growth and reproduction at close distances to shore (30 – 40 m); part of this effect was due to density in one population, but when density effects were removed statistically, there still remained a decline in growth and reproduction with increasing distance from shore. Improved vigor nearest to shore may be due to continual sand deposition. Survivorship showed a Type I pattern, with low mortality throughout the growing season. By colonizing newly-deposited and continually shifting sands,T. purpurea can contribute to the earliest stages of ecological succession along disturbed beaches in eastern North America and may be valuable to the development and management of urban coastal plant communities.     

Population biology of the annual grass Triplasis purpurea in relation to distance from shore on Staten Island, New York

Four populations of the native annual grassTriplasis purpurea were surveyed on coastal beaches along the south shore of Staten Island, NY, to determine the potential of this species to colonize shoreline habitats mostly devoid of other vegetation. If the species can establish and maintain dense populations, it may have conservation value for urban beaches disturbed by human activities. For two populations, survivorship, growth, and reproduction were monitored at different distances from shore to determine the ability of this species to maintain viable populations. At three sites,T. purpurea occurred in >75% of all quadrats and the highest density was 1195 plants/m² at 74 m from shore in one recently disturbed site. Density generally increased with increasing distances from shore at low tide (from ca. 40–90 m). Plants showed the greatest growth and reproduction at close distances to shore (30–40 m); part of this effect was due to density in one population, but when density effects were removed statistically, there still remained a decline in growth and reproduction with increasing distance from shore. Improved vigor nearest to shore may be due to continual sand deposition. Survivorship showed a Type I pattern, with low mortality throughout the growing season. By colonizing newly-deposited and continually shifting sands,T. purpurea can contribute to the earliest stages of ecological succession along disturbed beaches in eastern North America and may be valuable to the development and management of urban coastal plant communities.     

Growth,mortality and recruitment of the yellow clam Mesodesma mactroides on Uruguayan beaches

We studied some aspects of the population dynamics of the yellow clam Mesodesma mactroides (Deshayes, 1854) which inhabits the eastern sandy beaches of Uruguay. Observations were made from March 1983 through March 1985 (harvesting season), and additional data from January 1988 to December 1989 (closed season) were also included in the analyses. Parameters of growth and mortality were estimated by methods that make use of age or length information. An age/length relationship was constructed by counting growth rings. Results were consistent with those obtained with modal class progression analysis and other length/frequency methods. The growth curve obtained from the age/length relationship did not account for seasonality in growth rate, and a seasonal growth model explained growth oscillations in a better way. The instantaneous coefficient of natural mortality (M) estimated for the closed season was 1.64, whereas an increase in fishing mortality (F) was observed during 1984. Age composition did not change during the 2 yr of study, and the first 6 mm class accounted for over 50% of the population. The main contribution of recruits was observed between late summer and early fall (February to April), and the recruitment pattern suggested one main settlement period per year. Growth and recruitment success appeared to be regulated by density-dependence processes. Some implications for management are considered in this context. Methodological aspects concerning the reliability of empirical equations and programs for estimating growth and mortality are discussed.     

The annual reproductive cycles of Uca annulipes,Portunus pelagicus and Metapenaeus affinis (Decapoda: Crustacea) from the South-west coast of India

Employing the gonad index method, the reproductive cycles of three decapod crustaceans, Uca annulipes (Latreille), Portunus pelagicus (Linnaeus) and Metapenaeus affinis (Milne-Edwards) have been studied. In these crustaceans breeding is not continuous all the year round, but extends over several months of the year with distinct peak periods of gonadal activity. The male and female reproductive cycles are not concurrent. The peak of the reproductive cycle of males occurs slightly earlier in the breeding season than that of females. These studies indicate the possibility of production of successive broods of eggs during the same breeding season. In these species, the low saline conditions of the monsoon period are unfavourable for breeding. The medium and high saline conditions during the post-monsoon and pre-monsoon months, respectively, with plenty of planktonic food for the larvae, seem to be the favourable periods for breeding activity.     

Population dynamics and production of the planktonic copepods in a eutrophic inlet of the Inland Sea of Japan. I. Centropages abdominalis

Population dynamics and production of the calanoid copepod Centropages abdominalis were studied from November 1986 to November 1987 in Fukuyama Harbor, in the central part of the Inland Sea of Japan. This species was present in the plankton during a cold-water period from November to June (temperature range: 8.9 to 21.1 °C), with a peak abundance (23 600 ind m^–3) in February. During this period, six generations could be detected, and each generation time agreed well with that predicted from food-satiated laboratory experiments, indicating that the natural population was not food-limited. The population suffered extremely high mortality during the period from egg to naupliar stage (N) II: only 0.02 to 4% of the eggs survived to NII. However, the mortality in stages older than NII was almost negligible. The growth rate of C. abdominalis increased exponentially with increasing temperature. Its biomass and production rate showed marked seasonal variations largely in parallel with numerical abundance. The estimated production between 7 November 1986 and 29 May 1987 was 355 mg C m^–3 or 2.66 g C m^–2, 95% of which occurred during February and March. The daily production rate to biomass ratio increased exponentially with temperature from 0.18 at 8.9°C to 0.37 at 19°C.     

Direct estimation of natural mortality rates for littoral marine fishes using populational data from a marine reserve

 Visual censuses conducted in a marine reserve (Medas Islands) were used to estimate the natural mortality rates (M) for five common fish species in the NW Mediterranean Sea (Coris julis, Diplodus annularis, D. sargus, Serranus cabrilla and Symphodus roissali). Visual censuses of these same five species were also performed at three sites in unprotected areas of the coast where both commercial and sport fishing activity was normal. Censuses were conducted over a 3 year period. Estimates of M in the 3 years displayed scant seasonal or interannual variation, which may mean that the populations were in equilibrium during that period. The results of this study showed that the relationships between M and the growth parameters and maximum life span were unclear, and considerable caution is therefore recommended when using indirect methods of estimating M based on those parameters. For certain species the values of M were equal to or greater than the estimated total mortality in the exploited areas. The virtual absence of piscivorous predators in the unprotected area as a consequence of the high level of fishing in that area contrasts with the high abundance of such predators in the marine reserve. Since predation is the main contributor to M, estimated mortality in the unprotected areas is attributable nearly entirely to fishing. It is suggested that M may vary according to alterations taking place in conditions in the ecosystem inhabited by a species and thus that use of a value of M for a pristine population cannot be extrapolated to exploited areas. Received: 18 January 2000 / Accepted: 14 July 2000     

Ecology of sandy beaches in Oman

The benthic macrofauna and physical features of 10 sandy beaches along the coast of Oman were surveyed quantitatively. This is a mesotidal regime mostly subject to low to moderate wave energy but more exposed in the south. Five northern beaches are tide-dominated, with low wave energy, and their profiles consist of a berm, a steep, swash-dominated upper shore and a broad tide-dominated terrace from mid-shore downwards. They are composed of moderately sorted fine to medium sands. Southern beaches experience greater wave energy, particularly during the summer southwest monsoon, and exhibit smoother, concave profiles with fine, fairly well sorted carbonate sand. 58 species and species groups were recorded, with crustanceans, polychaetes and molluscs dominant. In general species richness was high, at least 19–25 species per beach, but dry biomass moderate to low at 26–90g/m shoreline, with one high value of 450g/m. Total abundance was moderate at 3–73×10³ organisms/m of beach. Some zonation was evident with ocypodid crabs andTylos in the supralittoral, cirolanid isopods on the upper shore and a variety of species on the lower shore. The coast of Oman appears to constitute a single zoogeographic region, but with some regional differentiation between north and south due to varying physical conditions. Thus, OmanÕs beaches are characterized by tide-dominated morphodynamics and exceptionally high species richness.     

Tisular and population level responses to habitat harshness in sandy beaches: the reproductive strategy of <Emphasis Type="Italic">Donax hanleyanus</Emphasis>

Microscopic analysis and field sampling procedures were used to compare demographic and reproductive strategies of the intertidal wedge clam Donax hanleyanus (Bivalvia: Donacidae) in two exposed sandy beaches with contrasting morphodynamics (reflective vs. dissipative) during 13 consecutive months. Histological analysis showed that: (1) the reproductive cycle of D. hanleyanus was more extended in the dissipative beach, and this was true for all the three pre-active (beginning of gonadal activity), active (maturation) and spawning stages; and (2) males and females showed significantly smaller sizes at sexual maturity at the reflective beach. Even though successive increments in proportion and mature at size were observed, the sigmoid function was significantly steeper at the reflective Arachania for both sexes, suggesting an abrupt transition to maturity. Field sampling revealed a more extended recruitment period at the dissipative beach, where recruits were also significantly more abundant than at the reflective beach. These results give support for the habitat harshness hypothesis, which predicts that in intertidal species capable of sustaining populations across a wide spectrum of physical conditions, such as D. hanleyanus, abundance, recruitment, size at maturity and extent of reproductive and recruitment seasons increase from reflective to dissipative beaches. However, a recent hypothesis suggests that reflective beaches acting as sink populations were not sustained, because mature and spawning individuals of both sexes were found in the reflective beach throughout the study period. Thus, we suggest that post-settlement processes are critical in modulating population patterns for this bivalve.     

Growth and mortality rates of the fan mussel <Emphasis Type="Italic">Pinna nobilis</Emphasis> in Lake Vouliagmeni (Korinthiakos Gulf,Greece): a generalized additive modelling approach

The temporal patterns and the effect of shell size and depth on growth and mortality rates of the endangered fan mussel Pinna nobilis were investigated in the marine Lake Vouliagmeni (Korinthiakos Gulf, Greece). A total of 160 individuals were tagged and monitored monthly for a period of 17 months. At each visit, the size of the tagged individuals (shell width, w) was measured in situ and recorded. Any mortality event was also recorded and attributed to natural causes or (illegal) fishing. Growth and mortality rates were modeled with generalized additive models, which are non-parametric flexible models that free the researcher from the limiting concept of a strict parametric shape. The use of GAMs allowed the exploration of shapes of growth and mortality response curves in relation to predictor variables and allowed the fitting of statistical models that better agree with ecological theory. Growth rates had a seasonal pattern, with an extended period of very slow growth between late autumn and early spring, i.e., during the cold season, another short period of slow growth during August (when water temperatures reached their maximum values exceeding 29°C), and a peak in growth rates during late spring–early summer, probably related to an optimum combination of temperature and food availability. Growth rates varied with shell size, with a peak at w ∼4.5 cm, followed by a sharp decline to an approximately constant level, with sizes ranging from 9 to 15 cm, and a further decline with larger sizes down to almost zero for w > 20 cm. Growth rates did not vary substantially with depth. Although P. nobilis is a protected species in the EU and its fishing is strictly prohibited, fishing mortality was very high in Lake Vouliagmeni (much greater than natural mortality), especially during the hot season when the lake was crowded by summer visitors. The fan mussels were poached exclusively by free-diving and due to the high turbidity of the lake’s water, fishing mortality was higher in shallow areas (and mostly for large individuals) and was practically zero at depths >9 m. Due to fishing mortality, a size segregation of P. nobilis was observed in the lake: large individuals were restricted to deeper areas, while young and small individuals were more abundant in shallow areas where there was preferential recruitment. Natural mortality was strikingly size dependent and P. nobilis suffered high natural mortality during the first year of life; the probability of death by natural causes quickly diminished as the fan mussels grew in size. No depth-related differences in natural mortality were found.