Abundance of estuarine crab larvae is associated with tidal hydrologic variables

Abundances of brachyuran megalopae and juveniles were measured throughout consecutive tidal cycles during six 2 to 3 d sampling periods in summer 1992, and associated with rates of change of tidal hydrologic variables in the Newport River Estuary. Current speeds and rates of pressure change fitted sinusoidal (tidal) models well; however, rates of salinity and temperature change did not. Analysis of plankton samples taken during spring and neap tides showed peak abundances during nighttime rising tides for all taxonomic groups: Callinectes sapidus, Uca spp., Xanthidae, and Pinnixa spp. megalopae, and Pinnotheres spp. juveniles. Megalopal and juvenile abundances from time-intensive sampling were related to rates of changes in the hydrologic variables using stepwise logistic regression. No hydrologic variable accounted well for the presence of Uca spp. megalopae. Megalopal presence was best predicted by current speed for Pinnixa spp. megalopae, and rates of changes in pressure for xanthid megalopae and Pinnotheres spp. juveniles, and salinity for C. sapidus megalopae. These variables might act as cues causing megalopae to ascend into the water column at a particular point in the flooding tide, and subsequently descend to or near the bottom prior to ebb flow. In this way, larvae which develop on the continental shelf or lower estuary undergo transport up the estuary by behaviorally altering their swimming activity and depth concurrent with tidal changes.     

Rhythmicity of larval release in three species of intertidal brachyuran crabs (Crustacea: Brachyura) from Inhaca Island (Mozambique)

 A study of rhythmicity of larval release in three species of intertidal brachyuran crabs, based on laboratory and field experiments, was undertaken at Inhaca Island, southern Mozambique, using Leptodius exaratus and Macrophthalmus grandidieri from December 1994 to January 1995, and Arcotheres palaensis from April to July 1995. L. exaratus and M. grandidieri showed a semi-lunar cycle in larval release. The release of larvae for L. exaratus, a species having conspicuous larvae, occurred in the first half of the night, after the post-crepuscular high tide, which suggests maximisation of protection of larvae from visual predation. The larval release activity matched the late spring and early neap tides. Results from the field were similar to those from the laboratory. M. grandidieri, having inconspicuous larvae, did not show a pattern related to the light–dark cycle and hatched during spring tides (around full and new moons) to maximise larval dispersion. A. palaensis, living inside the host mussel which inhabits the lowest section of the intertidal zone, did not show a relation with moon phase, tidal or light–dark cycles. Received: 16 February 1999 / Accepted: 8 December 1999     

Endogenous control of timing of metamorphosis in megalopae of the shore crab Carcinus maenas

Batches of hundreds of freshly collected megalopae of the shore crab Carcinus maenas (L.) showed persistent circatidal rhythms of moulting to the juvenile crab stage when maintained in constant laboratory conditions. Peaks of moulting occurred around expected times of high tides, with few megalopae moulting at other times. In larvae collected offshore, the highest tidally-timed peak of metamorphosis occurred during the second to fifth expected times of high tide, and metamorphosis of 50% of each batch took about 22 h or longer. In contrast, in larvae collected at the water's edge, 70% metamorphosed during the first expected episode of high tide, within 6 to 8 h after collection. However, although inshore megalopae moulted before offshore ones, the tidal timing of moulting remained unaltered whether megalopae were collected at neaps or springs, from the water's edge or farther offshore, in the presence or absence of natural substratum, and under various light–dark and salinity regimes. Metamorphosis of C. maenas megalopae around the times of high tides may enhance settlement into the upper intertidal zone. Early juveniles of the crab apparently prefer that zone as they are most abundant there and, unlike adults, do not undertake up-and-down-shore migration with tides. The present finding demonstrates, for the first time, endogenous physiological timing of circatidal periodicity in the metamorphic moult of crab megalopae, suggesting that endogenous factors, as well as exogenous ones should be taken into account in considering the process of settlement by crab megalopae. Received: 21 February 1996 / Accepted: 27 November 1996     

Variability in abundances of fishes associated with seagrass habitats in relation to diets of predatory fishes

The spatial, diel and tidal variability in the abundance of piscivorous fishes and their teleost prey, and the dietary composition of predatory fishes were investigated in beds of Heterozostera tasmanica within Port Phillip Bay, Australia, from September 1997 to February 1998. Predatory and prey fish assemblages were sampled from beds of H. tasmanica at three locations during each combination of diel (day and night) and tidal (high and low) cycles. Pelagic and benthic crustaceans represented >60% by abundance of the diets of all predatory fishes. Seven species, 54% of all predatory fishes, were piscivorous. These piscivores consumed individuals from seven families, 36.8% of the fish families being associated with seagrass. Western Australian salmon, Arripis truttacea (Arripidae) (n = 174) and yank flathead, Platycephalus speculator (Platycephalidae) (n = 46) were the most abundant piscivores. A. truttacea consumed larval/post-larval atherinids, gobiids and sillaginids. P. speculator consumed late-juvenile/adult atherinids, clinids and gobiids. While the abundances of piscivores varied between locations (P < 0.001) and diel periods (P = 0.028), the relative differences in piscivore abundance between sites and diel periods were not consistent between tides. The abundances of A. truttacea varied in a complex way amongst sites, diel period and tidal cycle, as shown by a three-way interaction between these factors (P = 0.026). Only during diurnal periods at St. Leonards was the abundance of A. truttacea significantly higher during high than low tides (P < 0.001). During the other diel periods at each site, the abundance of A. truttacea did not vary. P. speculator was significantly more abundant nocturnally (P = 0.017). The abundance of small (prey) fishes varied significantly amongst sites (P < 0.001). During the day, the abundance of small fishes did not vary between high and low tides (P = 0.185), but their nocturnal abundance was greater during low tide (P < 0.001). Atherinids (n = 1732) and sillaginids (n = 1623) were the most abundant families of small fishes. Atherinids were significantly more abundant nocturnally (P = 0.005) and during low tides (P = 0.029), and varied significantly amongst sites (P < 0.001). Sillaginids varied significantly only amongst sites (P < 0.001). Seagrass beds provide a foraging habitat for a diverse assemblage of predatory fishes, many of which are piscivorous. Anti-predator behaviour and amongst-location variability in abundances of piscivorous fishes may explain some of the diel and tidal, and broad-scale spatial patterns in small-fish abundances. Received: 23 July 1999 / Accepted: 18 January 2000     

Flood tide transport of blue crab, Callinectes sapidus, postlarvae: behavioral responses to salinity and turbulence

Blue crab (Callinectes sapidus) postlarvae (megalopae) use flood tide transport to move upstream in estuaries during nocturnal flood tides. The megalopae have a endogenous diel rhythm in activity that is inconsistent with this tidally timed behavior. Thus, it is hypothesized that this behavior is regulated by behavioral changes in response to exogenous cues associated with tidal currents. In a laboratory flow tank, blue crab megalopae were exposed to simultaneous changes in salinity and turbulence to simulate tides in an estuary. On simulated flood tides, megalopae ascended upon exposure to a salinity increase, remained swimming during times of high turbulence, and descended at times of low turbulence. Turbulence stimulated swimming for several hours, approximating the duration of tidal currents in estuaries. Swimming was inhibited by decreasing salinity on simulated ebb tides. These results support a model for regulation of flood tide transport by blue crab megalopae as follows: (1) blue crab megalopae are stimulated to swim into the water column by increasing salinity associated with flood tide; (2) megalopae remain swimming during flood tide in response to high levels of turbulence; (3) megalopae descend at the end of flood tide, when current speed and turbulence decline to low levels; and (4) megalopae are inhibited from swimming on ebb tides by the associated salinity decrease. This is the first model for regulation of flood tide transport in a species lacking a tidal rhythm in activity.     

The ichthyofaunal composition of the Elbe Estuary: an analysis in space and time

A stow net (90 m² opening) was used, at regular intervals between July 1989 and June 1993, to collect fish over ca. 4 h on both the flood and ebb tides at five sites distributed at regularly spaced distances throughout the Elbe Estuary. Overall, 53.4% of the species were marine, and some of these were abundant, e.g.Clupea harengus, Pleuronectes flesus, Pomatoschistus minutus, Merlangius merlangus and Sprattus sprattus. However, in terms of number of individuals, such species contributed only 9.1%, compared with 90.0% by anadromous species, which was predominantly due to exceptionally high catches of Osmerus eperlanus. The ichthyofaunal compositions on the flood and ebb tides were similar, even downstream where salinities change markedly during each tidal cycle. Although this is probably due in part to groups of fish being swept first upstream on the flood tide and then back downstream on the ebb tide, several species clearly tended to occupy particular regions of the estuary. Each year the species composition changed sequentially from the most downstream site (max. salinity=ca. 31.4‰) to the most upstream site (max. salinity=ca. 1.5‰), mainly as a result of a sequential decline in the number and abundance of marine species and a progressive increase in the contributions of both anadromous and freshwater species. However, the marine species diagnostic of differences in the compositions among regions varied between years. Thus, for example, the diagnostic species for the downstream sites were C. harengus and M. merlangus in 1990/1991 and S. sprattus in 1992/1993, reflecting differences in the recruitment strengths of these marine species in the two years. The ichthyofaunal composition exhibited pronounced annual cyclical changes in each region and thus occurred irrespective of whether or not the salinity in a region underwent pronounced changes during the year. This cyclicity was attributable to intra-annual variations in the times of recruitment of certain marine species, particularly in the downstream region, and to seasonal migrations of diadromous species into each region, as well as to the movements of freshwater species into the upstream region in winter and early spring. Received: 18 November 1999 / Accepted: 25 October 2000     

Rhythmic patterns of abundance in small sublittoral crustaceans: variety in the synchrony with day/night and tidal cycles

Temporal fluctuations of abundance (or emergence) in small benthic and planktonic crustaceans were studied in shallow subtidal waters (1.5 to 3.5 m in tide height). The abundances were more or less rhythmic, and showed wide diversity ranging from very clear nocturnal patterns to patterns in sychrony with the tidal cycle alone. These abundance patterns were classified into categories relating to the degree of synchrony with day/night and tidal cycles. Nocturnal patterns were especially strong in benthic crustaceans, which would be inactive during the daytime, being attached to algae and stones or disappearing into rock crevices, and actively swim in the water at night. Mysis larvae also showed a clear nocturnal pattern. Their lifestyle might be similar to that of many benthic animals. Other planktonic crustaceans drifting in the water showed weak nocturnal patterns. In some planktonic crustaceans (e.g., Calanoida), the ratio of abundance in the surface and bottom samples was reversed between day and night. Their pattern might be a manifestation of weak diel vertical movement between day and night. Furthermore, most patterns of zooplankton and benthos were modified in synchrony with tides to various degrees. Small crustaceans may respond to changes of hydrologic variables fluctuating with the tides, which may exogenously produce a weak tidal component in their emergence patterns. Received: 12 January 1998 / Accepted: 29 August 1998     

Circatidal rhythm of an intertidal crab, Hemigrapsus sanguineus : synchrony with unequal tide height and involvement of a light-response mechanism

The circatidal rhythm of intertidal animals may reflect the inequality of the tides. In addition, a light-sensitive mechanism may be involved in their internal timing systems. To test these hypotheses, the larval release activity of the intertidal crab Hemigrapsus sanguineus was monitored under different light conditions in the laboratory. Under a 24-h light–dark (LD) cycle with the phase similar to the field, the activity coincided with the times of high tide in the field and showed a tidal rhythm. This rhythm free-ran in constant, dim-light conditions, suggesting that the timing is controlled by an endogenous clock. When the population was exposed to a 24-h LD cycle with the phase changed from that in the field, the tidal rhythm was phase-shifted; while the light cycle advanced in phase from the field caused a phase-advance of the rhythm, that delayed in phase induced a phase-delay of the rhythm. Thus, a light-response mechanism is definitely involved in circatidal timing systems. But the population rhythm showed a large variability among individuals, associated with the phase-shift, and the magnitude of the phase-shift did not accurately correspond to that of the light cycle. These results suggest that the light-response system can control the phase of the rhythm less stronger than that in estuarine crabs. Most releases occurred at higher high tides, but the release of some females obviously occurred at lower high tides. The larval release pattern thus could not be accounted for by a simple synchrony with higher high tides. Hatching of H. sanguineus occurred after a “hatching program” of 49.5 to 52.5 h. This program is initiated by some factor (hatching-program inducing stumuli: HPIS) transmitted from the female to the embryos. We speculated that this factor is effectively transmitted to the embryos when the habitat is exposed to air, i.e., at lower low-tide periods, and that once each embryo is stimulated, hatching occurs synchronously 2 d later during high tide. The release of HPIS is probably controlled by the circatidal clock of the female, and the 24-h LD cycle may participate in shifting this timing to the opposite low tide. Received: 14 January 1997 / Accepted: 18 February 1997     

Responses of estuarine crab megalopae to pressure,salinity and light: Implications for flood-tide transport

The megalopal larval stage of many estuarine brachyuran crabs appears to return to adult habitats by undergoing rhythmic vertical migrations which result in saltatory up-estuary transport on flood tides. Larval ascent into the water column during rising tides may be cued by changing hydrologic variables. To test this hypothesis, we investigated the responses of field-caught megalopae of the blue crab Callinectes sapidus and the fiddler crab Uca spp. to constant rates of pressure and salinity change under laboratory conditions. For both genera, pressure changes resulted in increased movement (barokinesis) and upward migration in the test chamber, with C. sapidus megalopae having a lower response threshold (2.8×10^-2 mbar s^-1) than Uca spp. larvae (5×10^-2 mbar s^-1). Similarly, larvae ascended in response to increasing salinity, with C. sapidus larvae being more sensitive. Larvae were negatively phototactic and failed to respond to pressure increases at light levels above 1.0×10¹⁵ and 1.0×10¹³ photons m^-2 s^-1 for C. sapidus and Uca spp. megalopae, respectively. Such responses are thought to explain the low abundances of larvae in the water column during daytime flood tides. Nevertheless, threshold sensitivities to increasing pressure for both genera were above levels experienced during floodtide conditions in the field. Similarly, it is unlikely that increasing salinity is sufficient to induce ascent in Uca spp. postlarvae. However, rates of salinity increase during midflood tide typically reach levels necessary to induce an ascent in C. sapidus megalopae. These results are consistent with the hypothesis that fiddler crab megalopae utilize an endogenous activity rhythm for flood-tide transport, while blue crab megalopae rely upon external cues, especially salinity changes, to time their sojourns in the water column.     

Endogenous swimming rhythms in estuarine crab megalopae: implications for flood-tide transport

Up-estuary migration of crab larvae to adult habitats is thought to be accomplished by selective tidal transport in which late-stage larvae enter the water column on flood tides and remain on or near the bottom on ebb tides. This study measured endogenous rhythms in swimming by the last larval stage (megalopa) of blue crabs Callinectes sapidus and fiddler crabs Uca spp. Previous field studies found that megalopae of both species were only abundant in the estuarine water column on nocturnal rising tides. Megalopae were collected from the Newport River Estuary, North Carolina (34°41N; 76°40W) during August–September 1992 and swimming activity was recorded for 4.5 to 7 d under constant conditions with a video system. Rhythms exhibited by both genera in the laboratory were not identical to those recorded in the field. Uca spp. displayed a circatidal rhythm, with maximum swimming occurring near the time of high tide in the field. Rhythm amplitude increased when crushed oyster shells were present, which suggested that megalopae bury or cling to the substrate during quiescent periods. In contrast, C. sapidus had a circadian rhythm in which maximum swimming coincided with the day phase in the field. In most trials, the activity of blue crab megalopae was unrelated to the expected tidal cycle. It was concluded that a tidal rhythm in swimming was the behavioral basis of flood-tide transport for fiddler crab larvae. The endogenous rhythm in blue crabs does not participate in transport, which probably results from behavioral responses to environmental cues associated with flood tide.     

Settlement times of blue crab (Callinectes sapidus) megalopae during flood-tide transport

Settlement by blue crab (Callinectes sapidus Rathbun) megalopae on artificial settlement substrates was monitored relative to tidal currents throughout ten nights from July to September 1997 in which the phase relationship between tides and the light dark cycle differed. Most megalopae were in intermolt, and the total number settling to collectors sampled at hourly intervals was greater than totals on collectors immersed all night. Maximum settlement occurred at slack water before ebb tide (SBE), with a smaller peak at slack water before flood tide (SBF). These results support the hypothesis that during flood-tide transport (FTT) blue crab megalopae remain swimming during flood tide at night in response to water turbulence and settle in response to the decline in turbulence occurring near SBE. Settlement peaks near SBF can be explained by a behavioral response of megalopae to increasing salinity at the beginning of flood tide, which results in an ascent response lasting only a few minutes. Depth maintenance in the water column is not maintained at SBF because of low water turbulence. Since light inhibits swimming and upward movement into the water column, settlement, and, presumably, transport were reduced when SBE occurred near the times of sunrise and sunset. Collectively, these results suggest that the phase relationship between the tide and light: dark cycles affects FTT, the timing of settlement, and behaviors associated with habitat selection. Published online: 9 August 2002     

Habitat selectivity of megalopae and juvenile mud crabs (<Emphasis Type="Italic">Scylla serrata</Emphasis>): implications for recruitment mechanism

Megalopae of several crab species exhibit active habitat selection when settling. These megalopae usually select structurally complex habitats which can provide refuge and food. The portunid mud crab, Scylla serrata, is commonly found within the muddy estuaries of the Indo-West Pacific after attaining a carapace width >40 mm. Despite substantial efforts, the recruitment mechanism of juvenile mud crabs to estuaries is not understood because their megalopae and early stage crablets (carapace width <30 mm) are rarely found. We used laboratory experiments to determine whether megalopae and early stage crablets are selective among three estuarine habitats which commonly occur in Queensland, Australia. These animals were placed in arenas where they had a choice of habitats: seagrass, mud or sand, and arenas where they had no choice. Contrary to the associations exhibited by other portunid crab megalopae, S. serrata megalopae were not selective among these estuarine habitats, suggesting that they tend not to encounter these habitats, or, gain no advantage by selecting one over the others. The crablets, however, strongly selected seagrass, suggesting that residing within seagrass is beneficial to the crablets and likely increases survival. This supports the model that for S. serrata, crablets and not megalopae tend to colonise estuaries, since a selective behaviour has evolved within crablets but not megalopae.     

Endogenous swimming rhythms of blue crab, Callinectes sapidus , megalopae: effects of offshore and estuarine cues

Larvae of the blue crab Callinectes sapidus Rathbun develop on the continental shelf. The postlarval stage (megalopa) occurs near the surface and is transported shoreward by wind-driven surface currents. It then uses selective tidal stream transport for migration up an estuary. Endogenous swimming rhythms were measured under constant dark conditions in the laboratory in megalopae collected from the Newport River Estuary (North Carolina), the Delaware Bay, and offshore from the Newport River Estuary. Megalopae from all areas had a similar circadian activity rhythm, in which they swam during the time of the day phase in the field and were inactive at night. This rhythm predicts the presence of a reverse, diel, vertical-migration pattern offshore which would contribute to the location of megalopae near the surface during the day. The rhythm lacks obvious ecological significance in estuaries because it does not contribute to selective tidal stream transport and would increase vulnerability to visual predators during the day. Attempts to entrain a circatidal rhythm in swimming by cyclic and step changes in salinity were unsuccessful, as the circadian rhythm persisted. The rhythm also continued in the presence of the eelgrass Zostera marina, which is a site of settlement and metamorphosis in the field. Thus, megalopae enter estuaries with a solar day rhythm in activity. This rhythm, however, is not expressed, because light inhibits swimming during the day upon exposure to estuarine water. Since this light inhibition is removed in offshore waters, the rhythm would be expressed if, after entering an estuary, megalopae were transported back to offshore areas. Received: 19 December 1995 / Accepted: 2 August 1996     

Mass spawning by the sea urchin Evechinus chloroticus (Echinodermata: Echinoidea) in a New Zealand fiord

A mass synchronous spawning of the sea urchin Evechinus chloroticus (Valenciennes) was observed in situ in Doubtful Sound, a large New Zealand fiord. Spawning occurred between 17:30 hrs and 18:30 hrs on 27 January 1994 and coincided with a full moon, spring tides and a period of decreasing sea temperatures. During spawning, the sea urchins formed a dense spawning aggregation of both sexes, with >90% of the urchins observed spawning at the time. Spawned gametes clouded the water column, and some were eaten by small labrid fish species. The spawning, which may have been as widespread as 40 km, marked a 42 to 50% decrease in gonad indices and resulted in a widespread, dense cohort of  E. chloroticus larvae within the fiord. Received: 25 September 1997 / Accepted: 6 March 1998     

Plasticity in the temporal organization of behaviour in the limpet <Emphasis Type="Italic">Cellana grata</Emphasis>

The behaviour of intertidal consumers is often tightly constrained to tidal movements, although activity patterns can vary within these constraints. Spatio-temporal variability in behaviour of a limpet, Cellana grata, was analysed over different tidal conditions (spring and neap tides) and during different times of the year (one summer and one winter) at sites in Hong Kong. Activity was generally dictated by tidal movements, being concentrated when animals were awash. Plasticity in behaviour was observed, with some limpets anticipating activity during the summer period and delaying activity during winter time. Limpets were active for a time equal, or slightly less, than time awash. As the time awash exceeded ~14–16 h, however, activity duration decreased. Within this general pattern, tidal variation as well as variation among times of the year was noted, with the lowest dependence on time awash being recorded during winter neap tides. Limpets showed a slight preference for being active during nighttime, which was particularly evident when animals were emersed during the summer period. Although the basic activity in C. grata is constrained to a specific temporal window, this limpet is able to modulate its foraging strategies and resting height, according to local, daily changes in environmental conditions.     

Spatial and temporal strategies in the surface activity of some sandy beach arthropods living along the French Atlantic Coast

During a field study on the eco-ethology of sandy beach arthropods conducted along the French Atlantic Coast it emerged thatTylos europaeus, Phaleria cadaverina andLabidura riparia were among the most abundant species that lived in the eulittoral. Tetradirectional pitfall traps, placed from the base of the dune to the tidal limits, were used. The traps intercepted surface-active arthropods, and the number of traps varied according to the levels of the tides. Data were recorded hourly during two periods corresponding to a spring tide and the following neap tide. At the saine time the most important environmental parameters were registered each hour. The results indicate that all three species show prevalently nocturnal surface activities. ForT. europaeus andPh. cadaverina an influence of the synodic and tidal phases was found on the zonation of the surface activity and on the extent of movement. The latter was due to the greater or smaller distances that the isopods and tenebrionids had to move from the diurnal burrowing sites to the foraging zones. ForL. riparia no variations in the total mean zonations were found between the two synodic phases, and only at neap tide were seaward, followed by landward movements found. Multiple regression analysis showed that the environmental parameters registered in this period were never limiting, as threshold values in this case probably were not reached.     

Foraging and homing behaviour in the high-shore, crevice-dwelling limpet Helcion pectunculus (Prosobranchia: Patellidae)

The foraging activity of the high-shore, crevice-dwelling limpet Helcion pectunculus (Gmelin, 1791) (Prosobranchia: Patellidae) from an exposed shore on the eastern Cape coast of South Africa was monitored. Activity was compared during spring and neap tides and between spring, summer, autumn and winter. Rhythms of activity in this limpet varied depending upon microhabitat; individuals inhabiting both east- and west-facing rock surfaces were active during nocturnal low tides, but limpets on west-facing rock surfaces were also active during daytime low tides, whilst in the shade. Individuals travelled further during foraging excursions in winter (xˉ=85.53 cm) than in either spring (xˉ=55.7 cm) or summer (xˉ=48.8 cm) and also during spring low tides (xˉ=89.8 cm) compared with neaps (xˉ=40.9 cm). H. pectunculus exhibited rigid homing to a fixed scar within a crevice, and feeding excursions were found to consist of three distinct phases: a relatively rapid outward phase, a much slower foraging phase and a rapid homeward phase. Whether or not these limpets graze throughout an excursion is not known. Foraging was always highly directional, with a mean vector which took limpets onto an area of the rock face with the highest microalgal biomass. The fact that H. pectunculus took advantage of optimal feeding areas and memorized their location to enable return visits suggests a learning component in herbivorous gastropod foraging behaviour. Received: 29 December 1997 / Accepted: 8 May 1998     

Estuarine epifauna recruit despite periodic hypoxia stress

In some estuaries, the recruitment of epifaunal benthic invertebrates coincides with a significant environmental stress, low water-column dissolved oxygen, termed hypoxia (̀ mg O₂ l^-1). Recruitment of epifaunal species was measured in the lower York River, a subestuary of the Chesapeake Bay, USA, which experiences predictable, periodic hypoxia associated with neap/spring tidal cycling during summer. Recruitment substrata were exposed during 48-h deployments in two areas with differing levels of hypoxia, and epifauna were allowed to recruit during periods of low oxygen (neap tides) and high oxygen (spring tides) in 1996 and 1997. Recruitment was often high during neap tides, even when severe oxygen depletion (<0.5 mg O₂ l^-1, <0.71 ml O₂ l^-1) occurred during deployments; indeed, peak recruitment episodes of several dominant epifaunal taxa, and of total epifauna, coincided with hypoxic events during both summers. Increased recruitment during neap tides suggests that factors besides hypoxia influenced recruitment in the York River; these factors may have included changes in larval availability and lower current speeds. This study illustrates how the relationship between recruitment and large-scale stresses, such as hypoxia, may be difficult to predict, since large-scale stresses are often correlated with numerous other factors. Short-term hypoxia appears to have little effect on recruitment in the field for many epifaunal species in this ecosystem, which may explain, in part, why substrata exposed for longer durations (1 month) in this system showed few effects of hypoxia on community composition or diversity. High larval tolerance of hypoxic stress may allow communities to persist even though the summer hypoxia season coincides with the recruitment of many epifaunal species. Electronic Supplementary Material is available if you access this article at http://dx.doi.org/10.1007/s00227-002-0930-6. On that page (frame on the left side), a link takes you directly to the supplementary material.     

Can synchronous spawning be predicted from environmental parameters? A case study of the lugworm Arenicola marina

The annual epidemic spawning period of a Scottish population of Arenicola marina (L.) has been recorded over a period of 13 yr. This population spawns between mid-October and mid-November in a discrete spawning event over a period of 4 to 5 d. Endocrine manipulation experiments showed that spawning is induced in females only if sufficient titres of PMH (prostomial maturation hormone) are present in the prostomia. These levels are attained during the 2 to 3 wk prior to the natural spawning date. The East Sands, St. Andrews population always spawns during periods of spring tides regardless of tidal amplitude or whether they are full- or new-moon tides. Meteorological data, including sea-temperature data were collected for each year, and correlation of the environmental data with spawning time was attempted. Correlation of spawning times with weather patterns showed that mean daily air pressures were significantly higher during the spawning period than from September to November as a whole. Evidence also suggests that a reduction in sea temperature is required prior to spawning. A significant moderate negative correlation was found between May to July air temperatures and spawning date, suggesting that higher May to July temperatures may induce early spawning. Daily rainfall and wind speed were also lower during the spawning period, but not significantly so. These results indicate that air pressure (or changes therein) may act as a final spawning cue, and the advantages of this are discussed in relation to fertilization success. A model of the interplay between environmental parameters and the endocrine mechanisms controlling the induction of spawning is proposed. Higher than average summer temperatures may advance gametogenesis to bring the population into a state of maturity (full-size oocytes, well-developed sperm morulae), and may also advance spawning time. Once the population has completed gametogenesis, a drop in sea temperature is then required to trigger an increase in endocrine titres within the prostomium, without which spawning cannot be induced by prostomial injection. The population spawns on spring tides; however a lack of clement weather coinciding with the spring tide will result in population-wide spawning being aborted, as in 1996. Clement weather (high pressure, low rainfall and wind speed) in conjunction with spring tides permits spawning to proceed to completion. Received: 21 June 1999 / Accepted: 25 January 2000