Early life history of the congrid eels gnathophis habenatus and G. incognitos in New Zealand waters

Larvae (leptocephali) of Gnathophis habenatus (Richardson, 1848) and G. incognitas Castle, 1963 occur off Castlepoint throughout most of the year (not sampled December‐February), In general they are smallest in late summer (March) and ilargest in mid‐spring (October‐November), with metamorphosis to the juvenile in early summer of the year of spawning. The two species therefore have a larval life of approximately 10 months. The early life of these two species in Australian waters, and of G. capensis (Kaup) off southern Africa, agrees well with these observations. Eel eggs collected in the East Cape region of New Zealand and tentatively identified as those of Gnathophis confirm the spawning times (March—April) suggested by the sizes of leptocephali.     

Changes in life cycle of the Ponto‐Caspian amphipod Pontogammarus robustoides at in the northernmost part of its range

The aim of this study was to investigate the dynamics of abundance, population structure and reproductive variables of the invasive Ponto‐Caspian amphipod Pontogammarus robustoides in its northernmost population, located in the Baltic Sea. Pontogammarus robustoides population variables were studied every month from May to October over 7 years (2002–2008) at two locations in the eastern part of the Gulf of Finland, one site in the north and one in the south. Observations of macrozoobenthos were also conducted once per year in July at 12 sites in this gulf during 2002–2014. This was to monitor the distribution area of P. robustoides and its abundance/biomass in different parts of the Gulf. The results showed that P. robustoides is a widely spread species in the Gulf, preferring to colonize stone‐sandy, vegetated and littoral habitats, from the shore to a depth of 5 m, with water salinity of 0.2–3 ppt. This species co‐exists with other amphipods, such as the Baikalian Gmelinoides fasciatus, the North Atlantic Gammarus tigrinus and the native Gammarus spp. The abundance of P. robustoides varied between the two study sites and among years (2002–2008), reaching a maximum level of 1000 ind.·m^?2 for adults and 3000 ind.·m^?2 for juveniles. The temperature‐dependent duration of egg development (embryogenesis) in the amphipod and the annual number of generations in P. robustoides populations varied among years. Three generations were recorded in most years and two generations in the coldest years. Clutch size (or fecundity) was dependent on the body length of female and averaged between 33 and 43 eggs per clutch for females ranging in body length from 11.2 to 12.2 mm. A maximum of 75 eggs were observed in a female with a body length of 16 mm. This paper concludes that life‐cycle variables change when P. robustoides spreads from the southern (Ponto‐Caspian region) to northern (Baltic Sea) range areas. In the studied northernmost location, the fecundity and number of generations of this amphipod were 1.5–2‐fold lower compared with the more southern locations. In addition, these variables and P. robustoides abundance were different between the coldest and warmest years at study sites that testify about their distinct dependence on local environmental conditions; further changes in life cycle of this species may be expected together with current climate warming.     

Is seagrass an important nursery habitat for the Caribbean spiny lobster,panulirus argus,in Florida?

Caribbean spiny lobster (Panulirus argus) settle preferentially in macroalgal‐covered hard‐bottom habitat, but seagrass is more prevalent in Florida (United States) and the Caribbean, so even low settlement of lobsters within seagrass could contribute substantially to recruitment if post‐settlement survival and growth were high. We tested the role of seagrass and hard‐bottom habitats for P. argus recruitment in three ways. We first explored possible density‐dependent regulation of early benthic juvenile lobster survival within cages deployed in seagrass and hard‐bottom habitats. Second, we compared settlement and survival of P. argus in both habitats, by comparing the recovery of microwire‐tagged early benthic juveniles from patches of seagrass and hard‐bottom. Finally, we assessed the relative abundance of juvenile lobsters in each habitat by deploying artificial structures in seagrass sites and compared these data with data from similar deployments of artificial structures in hard‐bottom habitat in other years. More early benthic juvenile lobsters were recovered from cages placed in hard‐bottom than in seagrass, but mortality of the early benthic life stage was high in both habitats. In regional surveys, the mean number of lobsters recovered from artificial shelters deployed within seagrass was lower than in any year that we sampled hard‐bottom, indicating that fewer lobsters reside naturally in seagrass, particularly large juveniles >40 mm carapace length. The greater abundance (and likely survival) of juvenile P. argus that we observed in hard‐bottom habitat as opposed to seagrass, combined with previous studies demonstrating that postlarval P. argus are attracted to, settle in, and metamorphose more quickly in red macroalgae, confirm that macroalgae‐dominated hard‐bottom habitat appears to be the preferred and more optimal nursery for Caribbean spiny lobster.     

Size‐frequency distributions of Joania cordata and Argyrotheca cuneata (Brachiopoda: Megathyrididae) from the Central Tyrrhenian Sea

Size‐frequency distributions can support reliable inferences concerning population dynamics of brachiopods, but only a few data are available so far. In this study, length and width frequency distributions of dead specimens of the Recent brachiopods Joania cordata and Argyrotheca cuneata from the Marine Protected Area ‘Secche di Tor Paterno’, Central Tyrrhenian Sea, Italy (41°35′ N, 12°20′ E), are reported in order to add new data about size‐frequency distributions of brachiopods. The studied specimens came from death assemblages in the coralligenous substrate, in the Posidonia oceanica meadows, and in the sand channels. The observed patterns vary from left‐skewed (J. cordata) to right‐skewed (A. cuneata), indicating respectively a low and high mortality of smaller individuals. Significant differences between the coralligenous substrate and the P. oceanica meadow were observed for both species, revealing a variation among different habitats. All length and width distributions are clearly polymodal, but the biological meaning of the peaks is difficult to interpret, as the two species seem to have a 2‐year life span. A biometric analysis of shell sizes revealed that length and width are the most variable parameters during the growth of the animal.     

Individual Variation in the Covering Behaviour of the Shallow Water Sea Urchin Paracentrotus lividus

Abstract. A number of recent field‐based studies have investigated the possible functions of covering behaviour in a range of echinoid species. Some of these have found important relationships between covering behaviour and environmental factors, such as light (in particular, ultra‐violet light) and covering item availability. However, these studies have often shown considerable within‐species variation, thus making covering functionality both within and between species difficult to interpret. The present study experimentally investigates individual variability in covering behaviour characteristics of the sea urchin Paracentrotus lividus (Lamarck). Data were collected bimonthly over a 1‐year period at Lough Hyne, Ireland, by observing P. lividus in outdoor aquaria. Significant variations in covering behaviour characteristics were observed as a function of individual size and mobility but not as a function of sex. A significant relationship between water temperature, mobility and covering behaviour was also found and is hypothesized to have implications for grazing activity in P. lividus. In contrast to previous field studies of P. lividus at Lough Hyne, no summer ‘peak’ in the proportion of individuals covering was observed. However, there was a significant relationship between local solar radiation and ‘time to first cover’. Overall, these data highlight (i) the ecological significance of individual variation in P. lividus covering behaviour and (ii) the importance of investigating multiple factors as potential determinants of echinoid covering behaviour.     

Growth and population dynamics of the non‐indigenous species Branchiomma luctuosum Grube (Annelida,Sabellidae) in the Ionian Sea (Mediterranean Sea)

The population dynamics and gametogenesis of the non‐indigenous polychaete species Branchiomma luctuosum Grube, 1869 (Annelida, Sabellidae) has been investigated at three sites in the Taranto Seas (Ionian Sea, Mediterranean Sea). The species, probably introduced from the Red Sea, has been reported in the Mediterranean Sea since 1983. The species is hermaphrodite, and the reproductive season is between June and October when the largest mean size of oocytes was recorded together with the presence of mature spermatozoa. Small oocytes are present in specimens reaching about 20–25 mm in length. Therefore oogenesis seems to begin early during the first year of life, but the first reproduction can occur when the worms attain a larger size and are at least 6 months of age. Although most of the individuals reproduce seasonally within a discrete period, some individuals can reproduce in different periods during the year, the oogenesis of individuals not being synchronous. A life span of at least 2 years is highlighted, with a faster growth rate during the first months (about 20 mm per month) decreasing to about 10 mm from the 3rd to the 8th months and slowing down again after the worm reaches 100 mm in size. Some differences in growth performance are discussed, enhanced by comparing the sites located at greater depth (5–7 m) and those located at 0.5 m depth, together with the possible interactions of this alien species with the autochthonous sabellid Sabella spallanzanii.     

Propagation and growth of Ponnotheres pholadis living in mantle cavity of Hiatula acuta

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Population Dynamics and Secondary Production of Emerita brasiliensis (Crustacea: Hippidae) at Prainha Beach, Brazil

Abstract. The secondary production and population dynamics of the mole crab Emerita brasiliensis Schmitt, 1935 (Decapoda: Hippidae) were studied by taking monthly samples from June 1993 to May 1995 at each of three intertidal transects at Prainha beach, Brazil. The lifespan was ca. 8 months for males and females, but females showed higher growth, mortality, secondary production, and turnover rate. The higher production in spring versus autumn and winter was related to intense recruitment during that period. The population production was estimated at between 39.86 and 46.88 g (AFDW) · m^?2 · a^?1 for the first year (June 93–May 94) and between 150.95 and 156.07 g (AFDW) · m^?2 · a^?1 for the second year (June 94–May 95); the mean annual biomass was 4.91 and 23.09 g (AFDW) · m^?2, respectively. High P/B rates, between ca. 6 and 9 · a^?1, reflected the fast growth, high mortality, and low lifespan of the population, characterized by a high percentage of recently recruited individuals.     

The invertebrate fauna of a forest stream and its association with fine particulate matter

The invertebrate fauna colonising wooden blocks placed in a beech forest stream was studied over a period of 1 year. At least 31 species were taken during the course of the study, with 3 species of Chironomidae predominating numerically. Most species were associated principally with fine particulate matter (FPM) deposited on the upper surfaces of blocks, although a stonefly, Spaniocerca zelandica, was most abundant on the barer surfaces. Of the Chironomidae, Paucispinig‐era approximata and Polypedilum canum had clearly defined life history patterns with emergence from February to April and October to January, respectively. Psectrocladius sp. had a poorly defined life history. Gut contents of 16 insect species showed that 4 were predominantly predatory, and the remainder were detritivores. The latter could be categorised as shredders (large particle detritivores) or collector‐browsers which mainly ingested particles in the dominant (<50 μm) size range. Oxygen consumption by fine stream sediments was similar at different times of year which suggests that the food quality of detritus may tie relatively constant at all times.     

Temporal Dynamics and Biomass Partitioning in Three Adriatic Seagrass Species: Posidonia oceanica,Cymodocea nodosa,Zostera marina

Abstract. The temporal dynamics of three seagrasses, Posidonia oceanica, Cymodocea nodosa and Zostera marina, was studied in different areas of the Adriatic Sea by analysing phenological parameters and biomass trends in different compartments of seagrass systems. For this purpose, samplings were conducted in 1997 once per season at each station, Otranto (southern Adriatic Sea) and Grado (northern Adriatic Sea). Structural parameters and biomass of plant compartments differed among seagrasses both in absolute values and in seasonal variability. P. oceanica was the largest plant, showing the highest number of leaves per shoot, highest leaf surface, Leaf Area Index and shoot weight. Z. marina was intermediate in size and had the longest leaves, whereas C. nodosa was the smallest seagrass. P. oceanica accounted for the highest total biomass (mean ± SE: 1895.9 ± 180.2 g DW · m^–2; CV = coefficient of variation: 19.0 %), considerably more than C. nodosa (mean ± SE: 410.4 ± 88.4 g DW·m^–2; CV: 43.1 %) and Z. marina (mean ± SE: 312.1 ± 75.1 g DW · m^–2; CV: 48.1 %), although the two latter species displayed a higher seasonal variability. Similarly, other features, such as shoot density, leaf surface, LAI, shoot weight and relative contributions of above‐ and below‐ground compartments, were less variable across seasons in P. oceanica than in the two other seagrasses, while leaf length showed the highest seasonal fluctuation in P. oceanica. As for biomass partitioning, C. nodosa showed a higher proportion of the below‐ground relative to above‐ground biomass (up to 90 %), with a distinct seasonality, whereas in P. oceanica the proportion of below‐ground biomass (around 80 %) was fairly constant during the year. We infer that in P. oceanica the seasonal forcing is probably buffered by the availability of internal resources stored permanently during the year in the below‐ground. In C. nodosa and Z. marina, instead, growth processes seem to be amplified by a greater influence of environmental factors.     

Spawning and fecundity of two flounder species in the Hauraki gulf,New Zealand

Spawning grounds and spawning times of the sand flounder, Rhombosolea plebeia (Richardson), and the yellow‐belly flounder, Rhombosolea leporina Gunther, in the Hauraki Gulf are described. The occurrence of female fish at different stages of ovarian development at different stations during the year, the distribution of eggs in the plankton, and changes in ovary weight during the year were used as indicators of spawning grounds and spawning times.

The sand flounder was found to spawn in the waters to the east of Waiheke and Ponui Islands, at the northern end of the Firth of Thames, from June to November.

Yellow‐belly flounders spawned during September, October, and November, slightly to the south of the sand flounder spawning grounds in a belt extending from Tapu, on the eastern side of the Firth of Thames, north‐westwards towards Ponui Island.

Fecundity of both species was approximately proportional to the weight of the fish, or to the length cubed, and less than proportional to the ovary weight. The mean fecundity of sand flounders in the Hauraki Gulf varied from approximately 100,000 eggs in a fish of 18 cm to 500,000 in a 30 cm fish. That of yellow‐belly flounders varied from approximately 250,000 eggs in a fish of 30 cm to 1.25 million in a 45 cm fish.     

Colony growth in the harvested octocoral Pseudopterogorgia acerosa in a Caribbean coral reef

Pseudopterogorgia acerosa is the tallest gorgonian octocoral found in the Caribbean. In the Cartagena area, Colombia, P. acerosa is commonly harvested as raw material for handicrafts, rendering important the study of its growth and population dynamics. Growth measurements of branches from multiple colonies at two different reefs during 1 year revealed that the main growth was due to rapid elongation of the main (‘mother’) branch and production of daughter branches from the same (or just a few) mother branch(es). Some small colonies doubled their original size in a year. The branches’ absolute growth rate was weakly correlated with size, with no significant relationships between branch size and percent growth. While the pinnate branching pattern in P. acerosa is similar to its congeners Pseudopterogorgia elisabethae and Pseudopterogorgia bipinnata, it produces taller colonies due to the faster growth rate in the primordial mother branch. This growth mode seems advantageous as it augments exposure to light and higher fluxes of suspended food in the benthic boundary layer. Post‐harvesting, clipped colonies exhibited clear signs of regeneration, even in cases where only the holdfast remained. Pseudopterogorgia acerosa seems to have similar regeneration capabilities respect to other Pseudopterogorgia species, where regenerating colonies may grow faster than intact ones.     

Analysis of occurrence patterns and biological factors of cetaceans based on long‐term and fine‐scale data from platforms of opportunity: Madeira Island as a case study

Management and conservation issues are addressed through the identification of areas of particular importance, which requires the acquisition of baseline information on species distribution and dynamics. These types of data are particularly difficult to obtain at high resolution for large marine vertebrates like cetaceans, given that dedicated surveys are complex and logistically expensive. This study uses daily presence–absence sighting data of cetaceans collected year‐round from whale‐watching boats to support the theory that fine‐scale data obtained from platforms of opportunity can provide valuable information on species occurrence and group dynamics. Data from 7,551 (daily) sightings comprising 22 species were collected from 3,527 surveyed days over 11 years (mean of 321 days per year, SD = 17) in the pelagic environment of Madeira Island. Cetaceans were observed on 92% of the surveyed days, and a mean of 15.4 (SD = 1.5), 8.2 (SD = 2.0) and 2.1 (SD = 1.2) species were recorded per year, month, and day, respectively. There were significant differences in the number of species per month (p < .001), with the highest diversity recorded in June. At least nine species, comprising 96% of all sightings, were found to use the Madeiran waters on a regular basis, such as the Atlantic spotted dolphin (Stenella frontalis), the short‐beaked common dolphin (Delphinus delphis), the bottlenose dolphin (Tursiops truncatus), and others featured in the Red List of the International Union for Conservation of Nature as Endangered, Vulnerable, and Data Deficient. In addition, 10 species were found to use the Madeiran waters for travelling, feeding, resting, socializing and calving, which suggests that the southern and southeastern waters of Madeira Island constitute an area of interest for cetaceans. This study characterizes the cetaceans’ community structure (occurrence, aggregation sizes, behaviours, proportion of calves, and inter‐specific relationships) of a poorly studied region, providing important information for managers. Finally, the advantages and limitations of using fine‐scale data from a type of platform of opportunity that is increasing along coastlines globally are discussed.     

Early development of Forsterygion varium,Gilloblennius decemdigitatus,and G. tripennis (Pisces: Tripterygiidae)

In the Wellington area (central New Zealand) Forsterygion varium (Forster, in Bloch & Schneider 1801) spawns from early June to late November, Gilloblennius decemdigitatus (Clarke, 1879) from early June to early October, and G. tripennis (Forster, in Bloch & Schneider 1801) from July to October. They deposit their eggs on firm substrates below the low tide level. Egg clusters are attended by an adult fish until hatching, which in the laboratory at 11.5–13.5°C, occurs after 18 days for F. varium and 20 days for G. decemdigitatus. Development to hatching for G. tripennis takes about 21 days at 13–15°c. The egg development of all three species is illustrated. The yolk‐sac larvae (prolarvae) of F. varium, G. decemdigitatus, and G. tripennis at hatching average 5.85, 5.03, and 5.72 mm standard length respectively. Yolk‐sac larvae of F. varium have a single row of 9–17 small, stellate melanophores along the ventral midline of the tail. The yolk‐sac larvae of G. tripennis have up to nine melanophores along the ventral midline of the tail, a medium‐sized melanophore above the optic lobes, and a further two above the medulla. The yolk‐sac larvae of G. decemdigitatus have three large mid‐dorsal melanophores interspersed with five smaller, paler pigment cells, and three large mid‐ventral melanophores interspersed with three of the smaller cells.     

High gene flow promotes the genetic homogeneity of the fish goby Pomatoschistus marmoratus (Risso, 1810) from Mar Menor coastal lagoon and adjacent marine waters (Spain)

The extreme environmental variability of coastal lagoons suggests that physical and ecological factors could contribute to the genetic divergence among populations occurring in lagoon and open‐coast environments. In this study we analysed the genetic variability of lagoon and marine samples of the sand goby, Pomatoschistus marmoratus (Risso, 1810) (Pisces: Gobiidae), on the SW Spain coast. A fragment of mitochondrial DNA control region (570 bp) was sequenced for 196 individuals collected in five localities: Lo Pagan, Los Urrutias and Playa Honda (Mar Menor coastal lagoon), and Veneziola and Mazarrón (Mediterranean Sea). The total haplotype diversity was h = 0.9424 ± 0.0229, and the total nucleotide diversity was π = 0.0108 ± 0.0058. Among‐sample genetic differentiation was not significant and small‐scale patterns in the distribution of haplotypes were not apparent. Gene flow and dispersal‐related life history traits may account for low genetic structure at a small spatial scale. The high genetic diversity found in P. marmoratus increases its potential to adapt to changing conditions of the Mar Menor coastal lagoon.     

平潭岛东北部近岸海域大型污损生物群落结构特征

于2017年6月至2018年5月在平潭岛东北部近岸海域开展污损生物生态研究,探讨了该海域大型污损生物的群落结构特点及演替趋势。周年模拟挂板试验结果显示,该海域大型污损生物隶属13门92种,以广分布暖水种为主,其中主要优势种为营固着生活的网纹藤壶(Amphibalanus reticulatus)、大室别藻苔虫(Biflustra grandicella)、三角藤壶(Balanus trigonus)和中胚花筒螅(Ectopleura crocea),以及自由生活的廉形叶钩虾(Jassa falcata)和齿掌细身钩虾(Maeropsis serratipalma)等。虽然该海域全年均有生物附着,但附着强度的季节性差异显著,附着盛期为6月至8月(生物量介于7 326.0～12 970.0 g/m2之间,以湿质量计),12月至次年2月(生物量介于39.5～1 580.5 g/m2之间)为附着淡季,而且污损生物摄食类型以悬浮物食者为主。温度和盐度是影响污损生物附着和分布的主要环境因素,水体透明度、水动力条件、地表径流和养殖等人类活动对污损生物的分布也有重要影响。     

Life‐history studies on New Zealand Brachyura

Abbreviated or direct development is described in Pilumnus novaezealandiae Filhol, 1886 and P. lumpinus Bennett, 1964 from New Zealand. Embryonic development is separated into Nauplius, Metanauplius and embryonic zoeal stages. The ontogenetic position of the pre‐zoeal cuticle suggests that it represents the Metanauplius stage, and the absence of plumose pre‐zoeal aesthaetes in P. novaezealandiae implies relationship with the more recently evolved brachyuran families. The significance of the pre‐zoeal cuticle in brachyuran phylogeny is discussed. In P. noveazealandiae four embryonic zoeal phases are recognised as approximately equivalent to the four Zoea larval stages occurring in the majority of xanthid Brachyura. It was not possible to distinguish similar phases in embryonic zoeal development of P. lumpinus because of advanced anachronism which is normally associated with abbreviated development. The appearance of relatively advanced structures very early in ontogeny is attributed to the absence of the need for successive developmental stages to be adapted for free life. In the embryonic Zoea, carapace spines considered to assist with flotation and orientation, and natatory setae, are together reduced or absent in abbreviated development where such structures would be of no use within the egg.

In P. novaezealandiae and P. lumpinus increase in egg size during incubation is probably due to absorption of water by the embryo during development, and eclosion is initiated by rupture of the chorion through internal pressure thus created. Larvae of P. noveazealandiae hatch at a Megalopa larval stage and are retained beneath the pleon of the female crab. Those of P. lumpinus hatch as advanced, much‐modified, and non‐natatory Zoeae which are not retained by the parent. Simultaneous moulting of the pre‐zoeal and zoeal skins occurs in both species.

The Megalopa larva and first five juvenile crab stages of P. novaezealandiae are described and compared with equivalent stages of P. vestitus from Australia recorded by Hale (1931). The Megalopa larva of P. lumpinus is described. P. lumpinus is considered as the least evolved of the three species of Pilumnus possessing abbreviated development, while P. vestitus suggests a second stage, and the third and most advanced stage is shown by P. novaezealandiae Ecology of these three species is correlated with abbreviated development. Retention of potentially free‐swimming Megalopa larvae by the parent explains gregariousness among juvenile crabs and adults of both P. novaezealandiae and P. vestitus

It is probable that abbreviated development has evolved among marine and freshwater Brachyura as a response to habitat requirements of the adult crabs. Partial or complete abbreviation of the planktonic phase and retention of larvae would assist with retaining the stock within a restricted locality or ecological niche. Abbreviated development among Brachyura therefore has little phylogenetic significance.     

Reproductive and growth strategies of Idotea balthica basteri (Pallas, 1772) population in the Bizerte lagoon (Tunisia,Southern Mediterranean)

We analysed several life history traits of the marine isopod Idotea balthica basteri (Pallas, 1772) from the Bizerte lagoon, Southern Mediterranean Sea. Growth was continuous throughout the life of the animal with a high growth rate in the first life phase, and the growth curve was described according to von Bertalanffy's model. The lowest growth rate (0.23 mm) was recorded in winter (December, January and February) and the maximum rate (2.31 mm) between April and June. The total number of hatched eggs or embryos was positively correlated with the body length of ovigerous females. This population of I. balthica basteri was iteroparous, showing distinct strategies of reproduction. Large ovigerous females with high fecundity were collected during the whole sampling period, while breeding in smaller females with low fecundity was restricted to the period from late spring to early autumn, Manca size increased significantly with increasing female body size and there was also a significant trade‐off between manca size and the number of eggs per brood. Reproductive allocation, ranging between 17.1 ± 1.2% in winter and 23.2 ± 1.8% in summer, was positively correlated with female weight. Accordingly, parental investment in producing a juvenile varied between 1.02% per manca in winter to 3.38% in spring. Evaluated traits show that late summer and autumn cohorts have a K‐strategy, whereas cohorts born in winter and spring, and which exhibit a shorter life time, exhibit faster development, earlier reproduction and a smaller parental investment tending towards an r‐selected strategy.     

Feeding ecology,population structure and distribution of Pomatoschistus microps (Krøyer, 1838) and Pomatoschistus minutus (Pallas, 1770) in a temperate estuary,Portugal

The feeding ecology, growth and spatial–temporal abundance and distribution of Pomatoschistus microps (Krøyer, 1838) and Pomatoschistus minutus (Pallas, 1770) were studied between June 2003 and June 2004, based on beam trawl surveys and macrobenthic samples conducted in the Mondego estuary, Portugal. Polychaetes, molluscs and amphipods were the most important items in the diet of P. microps, while for P. minutus the dominant preys were polychaetes, mysids and decapods. Pomatoschistus microps' recruitment lasted for six months and was composed of three new cohorts per year. Pomatoschistus minutus had an uncommon long reproductive season, from April to November, and population segregation was found, corresponding to the two reproductive peaks. Populations of both species were composed mainly of 0-group individuals with sand gobies presenting a more extended life span. In the Mondego estuary, inter- and intra-specific spatial segregation occurred between the two species and between the 0+ and 1+ age groups of P. microps.     

Phytoplankton models and life history strategies

Phytoplankton models generally do not consider the initial phases of seed stocks and deal only with the vegetative growth phase, ignoring life history strategies. Some quantitatively important diatoms, such as species of Chaetoceros and Skeletonema, have special strategies with respect to the timing of the planktonic phase that cannot be explained purely on the basis of environmental clues. In Norwegian waters and elsewhere, the first Chaetoceros bloom of the growth season usually starts in mid March, initiated by C. socialis. Other Chaetoceros species appear in the water column later. Species of dinoflagcllates like Alexandrium also bloom at certain times of the year. In many cases, phytoplankton inocula originate from resuspension of bottom-dwelling spores or cysts rather than from residual planktonic vegetative cells, and it is probable that, in some species, inoculation events are controlled by endogenous biological clocks. The sequential appearance of different Chaetoceros species may be related to day-length-regulated germination of spores. Most Chaetoceros species have few generations, but they appear at opportunistic times in the plankton. In contrast, Skelelonema costatum and Scrippsiella trochoidea appear at any time of the year. Some modelling results can be improved by including the dynamics of phytoplankton seed stocks in the sediments.