Talitrids on the upper shore and strand-line

27-10-2004
Researched byGeorgina Budd Refereed byThis information is not refereed.
EUNIS CodeA2.211 EUNIS NameTalitrids on the upper shore and strandline

Summary

UK and Ireland classification

EUNIS 2008A2.211Talitrids on the upper shore and strandline
EUNIS 2006A2.211Talitrids on the upper shore and strandline
JNCC 2004LS.LSa.St.TalTalitrids on the upper shore and strand-line
1997 BiotopeLS.LGS.S.TalTalitrid amphipods in decomposing seaweed on the strand-line

Description

A community of talitrid amphipods may occur on any shore where decomposing seaweed accumulates on the extreme upper shore strand-line. The community occurs on a wide variety of sediment shores composed of shingle and mixed substrata through to fine sands, but may also occur on mixed and rocky shores in some circumstances. The decaying seaweed provides cover and humidity for Talitrus saltator and other components of the community. The amphipods Orchestia spp. are also often present, as well as enchytraeid oligochaetes. Polychaetes, molluscs and other crustaceans may be brought in on the tide, but are not necessarily associated with the infaunal community. Further analysis of the data may determine that Orchestia spp. are associated with a denser strand and that there are differences in the community dependant upon the substratum-type. Talitrus saltator may occur further down the shore, almost invariably accompanied by burrowing amphipods such as Bathyporeia spp. (LGS.AEur). (Information taken from the Marine Biotope Classification for Britain and Ireland, Version 97.06: Connor et al., 1997a, b).

Recorded distribution in Britain and Ireland

A common and widespread biotope around Britain and Ireland that is probably under recorded at many locations owing to its ephemeral nature.

Depth range

Strandline

Additional information

Strand-lines are ephemeral habitats owing to the methods of their formation, but may be more permanent and extensive features particularly in sheltered embayments or estuaries. The strand-line is a fringe habitat, neither fully marine nor terrestrial, it is consequently colonized by invertebrates from both ecosystems. On exposed shores the strand-line is of particular importance because it acts as a precursor to sand dunes. Strand-lines enhance the stabilization of the foreshore by supplementing the organic and moisture content of the substratum so that pioneering plants such as sea sandwort, Honkenya peploides, sea rocket, Cakile maritima, and saltwort, Salsola kali may eventually establish (Shackley & Llewellyn, 1997). This 'open tall herb community' develops best on shores that receive large inputs of detached macroalgae and windblown sand (Ignaciuk & Lee, 1980). Such plants trap sand and favour the development of embryo dunes and subsequent fore dunes (Salisbury, 1952; Chapman, 1976; Davidson et al., 1991). In areas subjected to intensive recreational use and consequently where mechanical beach cleaning is practised, dune formation and stability could be adversely affected (Davidson et al., 1991).

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JNCC

Habitat review

Ecology

Ecological and functional relationships

  • Of marine invertebrates, the order Amphipoda (sand hoppers) is dominant in the biotope. Three genera are common amongst the strand-line, Talitrus, Talorchestia and Orchestia, feeding on the stranded seaweed. Such amphipods are responsible for most of the primary consumption of surface material. The feeding activity of the amphipods serves to fragment algal matter (Harrison, 1977). Fragmentation has been identified as being central to the control of decomposition rates and subsequently the productivity of food chains based on algal material (Robertson & Mann, 1980). Fragmentation of macroalgae increases the decomposition rate by reducing particle size, allowing a greater surface area for microbial action and the excretion of nitrogen rich materials enhances microbial growth (Robertson & Mann, 1980).
  • Large numbers of Coleoptera such as the black, smooth ground beetle, Broscus cephalotes, and the scarce Nebria complanata (restricted to the south west: Davidson et al., 1991; Fowles, 1994), frequent the biotope to feed on talitrids and insect larvae (Llewellyn & Shackley, 1996). Several species of darkling beetles (Tenebrionidae) and rove beetles (Staphylinidae), live in the sand several centimetres below the strand-line deposits.
  • Koop & Griffiths (1982) reported a distinct relationship between the distribution of strand-line macro and meio- fauna and their food source: >82% of the biomass of both groups was concentrated beneath the most recent strand-line on a beach on the Cape Peninsula, South Africa.
  • Mites of the genera Halolaelaps and Phaulodinychus (Gamasida), in addition to the Histiostoma (Acaridida) (Acarina: Chelicerata), occur in strand-line debris. All three mites disperse between strand-lines via talitrid amphipods. Talitrid amphipods offer two principle advantages over insect hosts. Firstly, talitrid amphipods, such as Talitrus saltator, migrate between strand-lines throughout the year allowing continual mite dispersal and secondly juvenile talidrids are sufficiently large to support several mites (Pugh et al., 1997).
  • Large numbers of birds feed along the strand-line at certain times of year, including waders, corvids and passerines, in addition to many seabirds (Pienkowski, 1982; Cramp & Simmons, 1983; Lack, 1986, Cramp, 1988; Davidson et al., 1991).
  • Terrestrial mammals such as foxes, voles, mice and rats also frequent the strand-line to feed (Shackley & Llewellyn, 1997).

Seasonal and longer term change

  • The biotope is ephemeral. The amount of macroalgae stranded on the shore is likely to vary seasonally, with deposits being particularly plentiful after winter storms that dislodge algae from the rocks.
  • Griffiths & Stenton-Dozey (1981) attempted to follow successional changes in the fauna of strand-line algal detritus and changes in its condition.
    • Bacteria colonized the stranded macroalgae within 24 hours (Koop & Lucas, 1983).
    • Amphipods and dipteran flies dominated the biomass during the early stages, followed by beetles later on as the algae dried.
    • Changes were apparent between algal debris that had been deposited singly and that deposited in banks. Both types of deposit lost half of their dry mass within the first seven days following stranding indicating a rapid rate of utilization by consumer organisms.
    • Single strands of algae lost moisture more rapidly than banked algae. For instance, the moisture content of single strands fell from 80% at 3 days to 22% after 6 days, whilst banks of algae retained a moisture content of 53% after 6 days.
  • Adults of the most common genus of dipterous flies, Fucellia and Coelopa, are always present, as they are opportunistic colonizers, able to move rapidly between strand-line deposits. However, they are exceptionally abundant in summer and autumn, coinciding with the presence of many larvae. This probably reflects a summer/autumn breeding peak (Stenton-Dozey & Griffiths, 1980).
  • In Britain during the day, Talitrus saltator occurs above the high-tide line, either buried within the sand at depths of between 10-30 cm or within high shore deposits of stranded algae (Keith Hiscock, pers. comm..), prior to emerging at night to forage intertidally on the strand-line (Williams, 1983b). During the winter, quiescent populations are found burrowed above the extreme high water spring level (EHWS), as deep as 50 -100 cm (Bregazzi & Naylor, 1972; Williams, J.A., 1976)

Habitat structure and complexity

On sandy shores, the strand-line is an ideal place for many species to live. The high organic content and water retaining capacity contrast with the relatively sterile and fast draining sand elsewhere. Seaweed in the strand-line is likely to be in various states of decay, older dryer material towards high water spring mark and fresh material towards low water, so that a plethora of microhabitats is available for colonization and utilization by the different life stages of different species, e.g. adult and juvenile wrack flies prefer different locations (see recruitment processes). In addition, differences in microclimate and decomposition rate exist between algae that is deposited in banks and algae deposited singly. The interstitial environment of the sand beneath the strand-line also differs to that of the surrounding area owing to the high concentrations of dissolved organic matter (DOM) (Koop & Griffiths, 1982).

Productivity

Several studies on South African beaches (Robertson & Mann, 1980; Koop & Griffiths, 1982; Koop et al., 1982; Griffiths & Stenton-Dozey, 1981) have examined aspects of the roles of macrofauna, meiofauna and bacteria in the productivity of the strand-line environment.
Production is predominantly secondary. Carbon fixed during primary production by macroalgae in other habitats enters the detrital pathway, the decomposition of which is a key process in the channelling of energy and cycling of nutrients. The feeding activity of amphipods in particular serves to fragment algal detritus (Harrison, 1977). Fragmentation has been identified as being central to the control of decomposition rates and subsequently the productivity of food chains based on algal material (Robertson & Mann, 1980), its role in stream and lacustrine ecosystems has been well documented (e.g. Cummins, 1974). Fragmentation of macroalgae increases the decomposition rate by reducing particle size, allowing a greater surface area for microbial action and the excretion of nitrogen rich materials enhances microbial growth (Robertson & Mann, 1980). Whilst macro- and meiofauna play a vital role in fragmentation of organic particles, bacteria are overwhelmingly important in the productivity of strand-line ecosystems (Koop et al., 1982). Annual turnover estimates (P/B) suggest that bacteria may account for about 87% of annual strand-line production, with meiofauna and macrofauna accounting for 10% and 3% respectively (Koop et al., 1982).

Recruitment processes

Many marine and otherwise terrestrial species utilize the strand-line deposits of wrack as a refuge in which to breed. Within the strand-line algal debris, it is possible for the vulnerable juvenile life stages to survive in a favourable microclimate and ready supply of nourishment. For instance:
  • The sand hopper, Talitrus saltator broods its eggs and has an annual univoltine reproductive cycle (one generation reaching maturity each year). As in all crustaceans, mating and the release of juveniles is synchronised with the moult cycle. Juveniles may be found from May through until September, but peak reproductive activity occurs in August. The breeding cycle in Talitrus saltator is shorter than in other intertidal amphipods and is controlled by daylength irrespective of air and sea temperature (Williams, 1978). Juveniles reach maturity before autumn, over-winter and breed the following summer.
  • Terrestrial dipterous flies, especially Fucellia maritima and Fucellia fucorum, utilize the strand-line macroalgae for the deposition of their eggs. The flies favour the drier wrack beds, and three larval instars (stages between moults) are passed there prior to pupation. Larvae are not adapted to living in wet wrack owing to a lack of hairs on their posterior spiracles and large spines on the ventral surface which are present in other wrack flies. Emergence of adults is sudden towards the end of March and adults remain in abundance until the end of September (Egglishaw, 1960). In contrast to the larvae, adults are most attracted to wet wrack. Other flies, such as Coelopa frigida, Coelopa pilipes and Thoracochaeta zosterae breed in the wrack beds and would not be found on the shore but for the accumulations of wrack in which to live and breed (Eltringham, 1971).
  • Some beetles and centipedes also complete their reproductive cycle in the wrack beds. The staphylinid beetle, Bleduis spectabilis burrows in the sand, its tunnels have a side chamber in which the larvae develop. Parents supply the larvae with food and ventilate the burrow. The centipede Hydoschendyla submarina has become wholly adapted to life in the littoral zone. The female lays eggs that are impermeable, and so are not affected osmotically if inundated by seawater (Eltringham, 1971).

Time for community to reach maturity

The biotope is ephemeral in nature, consequently in order to utilize the resources that the stranded debris provides, the community reaches maturity within a few weeks. Such rapid colonization is achievable owing to the fact that species of the community originate from both terrestrial (e.g. flies, centipedes and beetles) and marine (e.g. sand hoppers) environments so can migrate quickly from adjacent habitats.

Additional information

Amphipods, such as Talitrus saltator, are useful in strand-line population assessments and monitoring surveys (Shackley & Llewellyn, 1997). They are always present at and around the most recent high water strand-line deposit and possess a well defined endogenous and circadian locomotor activity pattern (Bregazzi, 1972; Bregazzi & Naylor, 1972; Williams, 1980) that controls their daily migration to recently deposited strand-line algae.

Preferences & Distribution

Recorded distribution in Britain and IrelandA common and widespread biotope around Britain and Ireland that is probably under recorded at many locations owing to its ephemeral nature.

Habitat preferences

Depth Range Strandline
Water clarity preferences
Limiting Nutrients
Salinity Full (30-40 psu)
Physiographic Open coast
Biological Zone Supralittoral
Substratum Gravel / shingle, Sand
Tidal
Wave Exposed, Moderately exposed, Sheltered, Very sheltered
Other preferences Deposits of organic debris, especially macroalgae

Additional Information

LGS.Tal is an ephemeral habitat occurring where the tide carries and deposits seaweed and other debris. Such material will decompose, be transported elsewhere and the quantity vary owing to factors such as tidal cycle and prevailing weather conditions.

Species composition

Species found especially in this biotope

  • Armadillidium album
  • Coelopa frigida
  • Coelopa pilipes
  • Enchytraeidae spp.
  • Fucellia fucorum
  • Fucellia maritima
  • Orchestia gammarellus
  • Talitrus saltator
  • Talorchestia deshayessii

Rare or scarce species associated with this biotope

  • Rumex rupestris

Additional information

No text entered.

Sensitivity reviewHow is sensitivity assessed?

Explanation

Many species, both terrestrial and marine frequent the strand-line habitat as it is transitional between the two environments. Talitrus saltator is considered to be an important characterizing species. It is always present at and around the most recent high water strand-line deposit, as it possess a well defined endogenous and circadian locomotor activity pattern (Bregazzi, 1972; Bregazzi & Naylor, 1972; Williams, 1980), that controls its daily migration to recently deposited strand-line algae. Other species within the biotope are opportunistic colonizers, e.g. wrack flies, with highly mobile adult stages that are capable of migrating considerable distances to reach strand-line material, but which would not be present in the intertidal zone but for the presence of strand-line debris.

Species indicative of sensitivity

Community ImportanceSpecies nameCommon Name
Important characterizingTalitrus saltatorA sand hopper

Physical Pressures

 IntoleranceRecoverabilitySensitivitySpecies RichnessEvidence/Confidence
High Very high Low Major decline High
Substratum loss, in this instance, the deposited macroalgae and other organic debris, would cause a loss of habitat for the strand-line community. Intolerance has been assessed to be high. Species utilizing the stranded material are likely to be removed along with the material and the habitat would be destroyed.
The benchmark against which intolerance is assessed assumes a single event, so following deposition of fresh macroalgae, recovery of the community would be expected to be very rapid in terms of the species present (e.g. many species would migrate to the strand-line from the terrestrial habitats and sand hoppers would buried in the substratum awaiting the arrival of a new strand-line) but may not attain their former abundance for several moths as a considerable proportion of characterizing species would be lost. At the benchmark level, a recoverability of very high has been made.
However, repeated removal of the substratum within a short space of time, e.g. as a result of mechanical raking for the purposes of beach cleaning, would be expected to impact upon the recovery of the strand-line community. A proportion of the population (e.g. sand hoppers, beetles, mites, flies etc.) would be removed or disturbed each time, including important juvenile stages, so that recovery would have to occur from a diminishing population and may take a considerable period of time from the point that the activity ceased. Some species in particular would be at risk. Amphipods, such as Talitrus saltator have an annual univoltine reproductive cycle (only one generation reaches maturity each year) (Williams, 1978). Newly hatched juveniles are unable to bury themselves in the sand to avoid desiccation and remain in amongst the freshly deposited strand-line debris, which maintains an 85-90% relative humidity over low tide (Williamson, 1951). The continuous removal of strand-line algae, even over the summer months will in the long term, effectively destroy the population (Llewellyn & Shackley, 1996). A much longer recovery period would be expected and it is questionable whether the community would recover at all following an impact of extended duration. For instance, the ability of amphipods to colonize over wider areas (e.g. > 200m) may be restricted by their endogenous pattern of activity that generally restricts movement over a relatively short distance in the intertidal zone (see Bregazzi & Naylor, 1972; Lincoln, 1979; Scapini et al., 1992 ).
Low Immediate Not sensitive Minor decline Low
Many of the species inhabiting the biotope are highly mobile adult forms, e.g. wrack flies, that would avoid being physically covered by additional sediment. A uniform layer of 5 cm of sediment would bury the strand-line material and the species active within it. The habitat would be temporarily lost to those species, mainly terrestrial, that were able to move away. Adult sand hoppers, such as Talitrus saltator, are likely to be capable of burrowing through additional sediment, as the species are capable of burrowing to depths between 10-30 cm (Williams, 1983b). Newly hatched juveniles are unable to bury themselves in the sand to avoid desiccation and remain in amongst the freshly deposited strand-line debris, which maintains an 85-90% relative humidity over low tide (Williamson, 1951). Although juveniles may not be able to bury through the additional sediment to regain the surface and fresh deposits of macroalgal debris, it is likely that the seaweed debris would itself maintain a sufficiently open structure under the sediment for vulnerable juvenile stages to survive. Intolerance has been assessed to be low, but would be expected to be higher if the smothering material was viscous. Recoverability, in terms of the species present and abundance, has been assessed to be immediate (within a few days) as characterizing species would either remain in situ or are sufficiently mobile to rapidly return, e.g. flies.
Not relevant Not relevant Not relevant Not relevant Not relevant
The community is unlikely to be affected by an increase in the concentration of suspended sediment in the water column, as the habitat is created by the deposition of macroalgae and other organic debris on the ebb tide. An intolerance assessment was not considered relevant.
Not sensitive* No change Low
The community is unlikely to be affected by a decrease in the concentration of suspended sediment in the water column, as the habitat is created by the deposition of macroalgae and other organic debris on the ebb tide. An intolerance assessment was not considered relevant.
Not relevant Not relevant Not relevant Not relevant Not relevant
Moisture conservation is a major stress factor to crustaceans on intertidal beaches (Hurley, 1959, 1968) and high-shore and terrestrial sandy beach amphipods such as Talitrus saltator are highly dependant on behavioural mechanisms to locate and maintain humid microhabitats during the diurnal, quiescent phase of their activity cycle (e.g. Williamson, 1951). An intolerance of high would have been recorded but for the ability of the species to avoid exposure to desiccating conditions. For instance, Williams (1983b) found a significant statistical correlation between the distribution of Talitrus saltator and sand moisture content. Talitrus saltator burrows down into the beach until sand with at least 2 % moisture content is encountered. The burrows of Talorchestia deshayesii are usually associated with the immediate area of the high water mark and freshly deposited algae, which would ameliorate moisture conservation problems, whilst Orchestia spp. combat desiccation by remaining burrowed within strand-line debris and below stones during its diurnal quiescent period in a manner similar to juvenile infaunal talitrids (Williams, 1983b). Unprotected individuals of Talitrus saltator above the substratum survived only approximately 0.5 -1 hours in air, a longer period than Talorchestia deshayesii and Orchestia gammarellus (Williamson, 1951). An intolerance assessment of not relevant has been made because the strand-line populations are protected from desiccation by their activity cycle and behaviour, otherwise desiccation would cause considerable mortalities.
Tolerant Not relevant Not relevant No change Moderate
The strand-line habitat is created as the tide, which carries macroalgal debris towards the shore, ebbs and deposits the material on the shore. The biotope has been assessed to be not sensitive to emergence, because it is created when the tide is out.
Not sensitive* Not relevant
A decrease in emergence is a factor considered not to be of relevance to this biotope as the habitat is created by the deposition of macroalgal debris on the shore as the tide ebbs.
Not relevant Not relevant Not relevant Not relevant Not relevant
The community is unlikely to be affected by an increase in water flow rate, as the habitat is created by the deposition of macroalgae and other organic debris on the ebb tide. An intolerance assessment was not considered relevant.
Low Very high Moderate Minor decline Moderate
The community is unlikely to be affected by a decrease in water flow rate as the habitat is created by the deposition of macroalgae and other organic debris on the ebb tide. An intolerance assessment was not considered relevant.
Low Immediate Not sensitive No change Moderate
The important characterizing species of the strand-line, Talitrus saltator, occurs to the south of the British Isles, so is likely to be tolerant of a chronic temperature increase of 2°C. Bregazzi & Naylor (1972) observed that the timing of activity was temporarily advanced by increased temperature but otherwise the activity pattern possessed a large measure of temperature independence. Specimens brought in to laboratory conditions from a field temperature of 10.5°C were introduced to (within 3 hours) and maintained for 15 days at constant temperatures of 15, 20 and 25°C. For Talitrus saltator maintained at the highest temperatures the activity mid-point advanced by as much as three hours to occur before midnight. However, alterations in activity were compensated for within two to ten days. Acute temperature increases may therefore temporarily disrupt activity of the sand hopper and other similar species, but owing to insufficient evidence for adverse effects in the field intolerance has been assessed to be low. Immediate recovery has been recorded as the locomotor activity rhythm is synchronized within a few days.
Intermediate Immediate Moderate Decline Moderate
The important characterizing species of the strand-line, Talitrus saltator, remains inactive in high shore burrows for much of the winter in more northern latitudes. In the laboratory, exposure to low temperature (2 or 3°C) was accompanied by the onset of inactivity, a precipitous decrease in oxygen uptake and a marked increase in the concentrations of the major ions in the haemolymph (Spicer et al., 1994). In addition to causing a complete cessation of activity, chilling (2-3°C for 8 hours) also causes a delay in the successive activity peaks following return to normal temperatures. Maximum delay occurred if chilling began during the inactive period of the sand hopper and was of equal duration to that of the chill. At other times the delay was less than that of the chill (Bregazzi, 1972). Thus it is possible that exposure to decreased temperatures in the field would enforce a period of inactivity causing disruption to the species normal behaviour with potential consequences for the maintenance of a position with appropriate moisture, e.g. the substratum may be come too dry or the temporary burrow become inundated with water. The effects of an unusually cold winter are likely to be a simple physical one, whereby quiescent sand hoppers freeze within the substratum, causing cell and tissue damage and eventually rupture of cell and body walls. Other supralittoral members of the Talitridae with a similar habit to Talitrus saltator were reported to be adversely affected by the severe winter of 1962/63. In particular, sand hoppers of the genus Orchestia were found dead in considerable numbers (Crisp, 1964). However, intolerance has been assessed to be low at the benchmark level as the behaviour of the sand hopper is likely to be disrupted by mild chilling, whilst death as a result of freezing is probable only in severe winters. Recovery from mild chilling has been assessed to be immediate following an initial disruption to its activity.
Not relevant Not relevant Not relevant Not relevant Not relevant
The community is unlikely to be affected by the light attenuating effects of an increase in turbidity within the water column, as the habitat is created by the deposition of macroalgae and other organic debris on the ebb tide. An intolerance assessment was not considered relevant.
Not sensitive* Not relevant
The community is unlikely to be affected by a decrease in turbidity in the water column, as the habitat is created by the deposition of macroalgae and other organic debris on the ebb tide. An intolerance assessment was not considered relevant.
Tolerant* Not relevant Not sensitive* Rise Moderate
The species characteristic of the biotope are unlikely to be directly affected by an increase in wave exposure. However, increased wave exposure may be of indirect benefit to strand-line populations. Storm wave erosion is likely to be an important factor in determining the quantity and quality of strand-line debris deposited on the beach. Increased wave exposure onshore is likely to increase the quantity of debris washed-up and available for colonization by strand-line fauna. For example, Shackley & Llewellyn (1994) recorded a positive correlation between strand-line debris weight and total amphipod numbers at beaches in South Wales. The community would benefit from the additional habitat/food resources and an intolerance assessment of not sensitive* has been made.
Low Very high Moderate Minor decline Moderate
The species characteristic of the biotope are unlikely to be directly affected by a decrease in wave exposure. However, decreased wave exposure may affect the abundance of shore strand-line populations. Storms are likely to be an important factor in determining the quantity of strand-line debris deposited on the beach. Reduced wave exposure over the period of a year is likely to affect the quantity of debris washed-up on the shore. As a consequence of reduced habitat availability the abundance of strand-line invertebrate populations may be reduced, owing to increased competition for resources. As the viability of some species may be affected an intolerance assessment of low has been made. However, fluctuation in the quantity of stranded debris is likely to be a normal feature of the biotope, as it is characteristically ephemeral, and species populations are likely to be able to cope but may fluctuate accordingly. Recoverability has consequently been assessed to be very high.
Tolerant* Not relevant Not sensitive* No change Low
The invertebrate fauna of the biotope are unlikely to be sensitive to noise at the benchmark level. However, birds that frequent the biotope to feed may be disrupted and their feeding efficiency impaired, although this would benefit invertebrate species. An assessment of not sensitive* has been made.
Tolerant* Not relevant Not sensitive* No change Low
The invertebrate fauna of the biotope possess visual acuity and are able to detect changes in light for purposes of navigation and probably are able to detect prey items within their visual envelope. However, apart from flies which would be temporarily disturbed by the approach of machinery, other important characterizing species of the biotope are unlikely to be disturbed by visual presence. Some species of bird that frequent the biotope to feed are likely to be disturbed by the visual presence of machinery and people in the vicinity of the strand-line, and their feeding efficiency reduced, although this would benefit invertebrate species. An assessment of not sensitive* has been made.
Tolerant Not relevant Not sensitive Not relevant Moderate
This biotope is subject to physical disturbance due to the rising and falling of the tide, wave action, and the movement of marine debris, including strand line material. Human trampling, and in this specific case, mechanical beach cleaning/raking, are potential sources of additional abrasion and physical disturbance. Adults of the many terrestrial species that exploit the biotope are highly mobile, e.g. wrack flies, and are likely to avoid disturbance. During the day, species such as Talitrus saltator, usually remain burrowed in the sand or amongst the algal debris (to avoid desiccation), so their environmental position may offer considerable protection from physical disturbance caused by trampling. Therefore, an overall assessment of not sensitivte has been made. However, the biotope is likely to be highly intolerance of substratum loss caused by mechanical beach cleaning/raking (see 'substratum loss', above).
Tolerant Immediate Not sensitive No change Moderate
Species within the biotope are mobile and in the event of displacement form strand-line material, would be able to rapidly recolonize debris elsewhere in the vicinity. An assessment of not sensitive has been made.

Chemical Pressures

 IntoleranceRecoverabilitySensitivityRichnessEvidence/Confidence
High High Moderate Major decline Low
In general, crustaceans are widely reported to be intolerant to synthetic chemicals (Cole et al., 1999) and intolerance to some specific chemicals has been observed in amphipods. Amphipods have been reported to be intolerant to TBT and leachates from antifouling paints (Laughlin et al., 1982). Numbers of the sand hopper, Talitrus saltator, were found in a lethargic state at the base of dunes at Constantine Bay (Cornwall), after spraying with the BP1002 an oil dispersant detergent after the Torrey Canyon oil spill (Smith, 1968). Intolerance has been assessed to be high.
Recovery assumes the deterioration of the contaminant, but is likely to be variable. In the case of the important characterizing species, Talitrus saltator, if a remnant population of young adults survived to breed, recovery would begin within a year. However, it has an annual univoltine reproductive cycle (only one generation reaches maturity each year) (Williams, 1978). So if a population were to be killed, recovery would take much longer and be reliant on the recolonization of adults able to breed. However, the ability of amphipods to colonize over wider areas may be restricted by their endogenous pattern of activity that generally restricts movement over a relatively short distance in the intertidal zone (see Bregazzi & Naylor, 1972; Lincoln, 1979; Scapini et al., 1992 ). Recoverability has been assessed to be high.
Heavy metal contamination
No information Not relevant No information Not relevant Not relevant
Talitrus saltator has been used as a spatial and temporal heavy metal biomonitor (Rainbow et al., 1989, 1998; Fialkowski et al., 2000). Bioavailable sources of trace metals to talitrids are available in solution and in food, the latter consisting of decaying macrophytic material on the strand-line. Such material acts as an adsorption site for heavy metals locally, as sandy substrata does not adsorb contaminants as easily as other substrata. The species is an efficient bioaccumulator of heavy metals whose moult cycle does not interfere with its biomonitoring potential. Specimens of the sand hopper from the Isle of Cumbrae, a non metal polluted site in the Clyde, Scotland, had zinc concentrations between 145-181 µg /Zn/g and copper concentrations of 35.8 µg/Cu/g (Rainbow & Moore, 1990). In comparison, Talitrus saltator from a heavy metal polluted site in Dulas Bay, Anglesey, Wales (Foster et al., 1978; Boult et al., 1994) had a zinc concentration of 306 µ g/Zn/g and a copper concentration of 112 µg/Cu/g. In the Gulf of Gdansk, Poland, comparable concentrations for zinc were in the region of 200-400µ g/Zn/g with bottom sediment zinc concentrations of 0-20 µg/g and 40µ g/g in the most polluted areas (Fialkowski et al., 2000). It is likely that the most significant contamination pathway to the amphipod is that of pollutants adsorbed to vegetative matter that is consumed rather than that concentrated in the water column. However, insufficient information has been recorded as no evidence concerning the effects of heavy metal contamination on the community as a whole was found.
Hydrocarbon contamination
High High Moderate Major decline Moderate
intolerance to hydrocarbon contamination has been assessed to be high. Supralittoral sediment habitats immediately adjacent to the littoral zone can be susceptible to damage from oil pollution and any subsequent attempts to remove the oil by scraping off the sediment surface. Oil which reaches the shore following a pollution incident generally gets concentrated along the high tide mark. Oil deposits on the strand-line and amongst seaweed would probably incapacitate and kill, by smothering and toxic effects, a considerable proportion of invertebrates that are found in strand-line debris. For instance, following the Torrey Canyon oil tanker spill in 1967 quantities of Talitrus saltator were found dead at Sennen, Cornwall, as were other scavengers of the strand-line, e.g. Ligia and Orchestia. Signs of oil dispersant detergent damage were reported at Constantine Bay (Cornwall) where sand hoppers were found in a lethargic state at the base of dunes after spraying with detergent (Smith, 1968).
Shackley & Llewellyn (1997) monitored shores with dune systems at Pendine and Pembury within Carmarthen Bay, that received oil spilt by the Sea Empress tanker in February 1996. Strand-line material at the two beaches contained quantities of oiled material and small particles of oil (2-5 mm in diameter) became mixed in with the sediment. However, Pendine was amongst the initial areas to become contaminated and received more viscous oil than Pembury, where oil appeared later and in a more weathered form. Tar balls persisted within the sediment at Pendine a year after the spill, whilst very little oiled material was found at Pembury a year later. Whilst physical and biological factors are important in determining the amphipod populations on such shores and differ between localities, differences were found in the abundance of amphipods between the two shores that could not be accounted for by physical and biological processes alone. Shackley & Llewellyn (1997) suspected that the persistence of oil at the strand-line and in the sediment beneath was affecting the strand-line community. Oil amongst strand-line material and in the sediment may affect the viability of species and/or it may simply deter species from colonizing. Recovery of the community is likely to vary according to the extent of oil pollution. Oil may be responsible for the decimation of amphipod populations, unless a remnant population survives buried in the substratum or in refuges higher than the tide mark. Some species in particular would be at risk. Amphipods, such as Talitrus saltator have an annual univoltine reproductive cycle (only one generation reaches maturity each year) (Williams, 1978). Newly hatched juveniles are unable to bury themselves in the sand to avoid desiccation and remain in amongst the freshly deposited strand-line debris, which maintains an 85-90% relative humidity over low tide (Williamson, 1951), so oil pollution could effectively remove the breeding population and recovery consequently protracted. Terrestrial species including coleopteran insects and dipteran flies are likely to colonize the strand-line rapidly following the deterioration of oil. Recoverability has been assessed to be high, as it may take more than a year for amphipod populations to recover to former abundances.
Radionuclide contamination
No information Not relevant No information Not relevant Not relevant
Insufficient
information.
Changes in nutrient levels
Not relevant Not relevant Not relevant Not relevant Not relevant
The community is unlikely to be directly affected by an increase in the concentration of dissolved nutrients in the water column, as the food resource that the community utilizes is in the form of macroalgal debris. An assessment of not relevant has been made.
Not relevant Not relevant Not relevant Not relevant Not relevant
The strand-line occurs in the supralittoral biological zone, where the benchmark increases in salinity are unlikely to occur. An assessment of not relevant has been recorded.
Tolerant Not sensitive* No change Low
The community may experience periods of freshwater inundation owing to episodes of rain. The highly mobile adult forms of wrack flies are likely to go elsewhere, whilst sand hoppers such as Talitrus saltator and other strand-line fauna are likely to seek protection within the strand-line debris. Talitrus saltator demonstrated the ability to hyper-regulate at lower external salinity concentrations, maintaining a haemolymph concentration between 750-850 mOsm (Morritt, 1988). It is likely to tolerate a short term decrease in salinity, resulting from episodes of rain. An assessment of not sensitive has been made.
Not relevant Not relevant Not relevant Not relevant Not relevant
At the benchmark level, intolerance is assessed against changes in the amount of dissolved oxygen in the water column. The strand-line habitat is created as the tide ebbs and deposits organic debris on the shore. Species inhabiting the strand-line are either fully terrestrial, or are marine species that have assumed a terrestrial mode of life, and all can therefore respire in air. An assessment of not relevant has been made.

Biological Pressures

 IntoleranceRecoverabilitySensitivityRichnessEvidence/Confidence
No information Not relevant No information Not relevant Not relevant
Wolbachia are endosymbiotic bacteria known to infect a wide range of arthropods and nematodes (Cordeaux et al., 2001). Wolbachia endosymbionts can alter their hosts reproduction. Cordeaux et al. (2001) found Wolbachia strains in the intertidal amphipods Talorchestia deshayessiiand Orchestia gammarellus living in the same habitats as semi-aquatic isopods whose populations typically have an infection level >35% (e.g. Sphaeroma rugicauda, Sphaeroma hookeri and the sea slater Ligia oceanica). Whilst reproductive impairment has been reported in isopod species, there is insufficient information to assess the likely impact of Wolbachia bacteria on amphipod populations inhabiting the strand-line. Insufficient
information has been recorded.
Tolerant Not relevant Not relevant Not relevant Low
No species alien to the British Isles are known to impact upon the community.
Not relevant Not relevant Not relevant Not relevant Not relevant
It is extremely unlikely that Talitrus saltator would be targeted for extraction and we have no evidence for the indirect effects of extraction of other species on this biotope.
Not relevant Not relevant Not relevant Not relevant Not relevant

Additional information

No text entered.

Importance review

Policy/Legislation

UK Biodiversity Action Plan Priority

Exploitation

In many popular tourist resorts, sandy beaches are cleaned by mechanical means, which involves the removal of litter, flotsam and natural strand-line debris. It is a criterion that in order for a beach to receive an EnCams Seaside Award or Blue Flag status that 'no algal or other vegetation may accumulate or decay on the beach, except in areas designated for a specific use, and as long as this does not constitute a nuisance' (EnCams, 2002; FEE, 2002). Consequently in order to gain such awards and maintain the amenity function of the beach, many local councils beach clean during the year but especially over the period between Easter and October. Such cleaning operations coincide with the reproductive period of strand-line fauna. Whilst litter presents a hazard and is unsightly, removal of natural strand-line detritus will impact upon the degradation and cycling of nutrients and potentially the stability of sand dune ecosystems.
Historically, stranded seaweed was collected as a garden conditioner but is probably no longer as widely practised.

Additional information

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Bibliography

  1. Ayari, A., Jelassi, R., Ghemari, C. & Nasri-Ammar, K., 2015. Locomotor activity patterns of two sympatric species Orchestia montagui and Orchestia gammarellus (Crustacea, Amphipoda). Biological Rhythm Research, 46 (6), 863-871.

  2. Backlund, H.O., 1945. Wrack fauna of Sweden and Finland: ecology and chorology. Opuscula Entomologica (Supplementum) 5, 1-236.
  3. Boult, S., Collins, D.N., White, K.N. & Curtis, C.D., 1994. Metal transport in a stream polluted by acid mine drainage - the Afon Goch, Anglesey, U.K. Environmental Pollution, 84, 279-284.
  4. Bregazzi, P.K. & Naylor, E., 1972. The locomotor activity rhythm of Talitrus saltator (Montagu) (Crustacea, Amphipoda). Journal of Experimental Biology, 57, 375-391.
  5. Bregazzi, P.K., 1972. The effects of low temperature upon the locomotor activity rhythm of Talitrus saltator (Montagu) (Crustacea: Amphipoda). Journal of Experimental Biology, 57, 393-399.
  6. Chapman, V.J., 1976. Coastal Vegetation. Permagon Press.
  7. Cole, S., Codling, I.D., Parr, W. & Zabel, T., 1999. Guidelines for managing water quality impacts within UK European Marine sites. Natura 2000 report prepared for the UK Marine SACs Project. 441 pp., Swindon: Water Research Council on behalf of EN, SNH, CCW, JNCC, SAMS and EHS. [UK Marine SACs Project.], http://www.ukmarinesac.org.uk/
  8. Connor, D.W., Brazier, D.P., Hill, T.O., & Northen, K.O., 1997b. Marine biotope classification for Britain and Ireland. Vol. 1. Littoral biotopes. Joint Nature Conservation Committee, Peterborough, JNCC Report no. 229, Version 97.06., Joint Nature Conservation Committee, Peterborough, JNCC Report No. 230, Version 97.06.
  9. Cordaux, R., Michel-Salzat, A. & Bouchon, D., 2001. Wolbachia infection in crustaceans: novel hosts and potential routes of transmission. Journal of Evolutionary Biology, 14, 237-243.
  10. Cramp, S. & Simmons, 1983. Handbook of the birds of Europe, the Middle East and North Africa. The birds of the Paleartic. Vol. III. Waders to Gulls. Oxford University Press.
  11. Cramp, S., 1988. Handbook of the birds of Europe, the Middle East and North Africa. The birds of the western Paleartic. Vol. V. Tryrant flycatchers to thrushes. Oxford University Press.
  12. Crisp, D.J. (ed.), 1964. The effects of the severe winter of 1962-63 on marine life in Britain. Journal of Animal Ecology, 33, 165-210.
  13. Cummins, K.W., 1974. Structure and function of stream ecosystems. Bioscience, 24, 631-641.
  14. Davidson, N.C., Laffoley, D., Doody, J.P., Way, L.S., Key, R., Drake, C.M., Pienkowski, M.W., Mitchell, M.R. & Duff, K.L., 1991. Nature Conservation and Estuaries in Great Britain. Peterborough: Nature Conservancy Council.
  15. Davies, C.E. & Moss, D., 1998. European Union Nature Information System (EUNIS) Habitat Classification. Report to European Topic Centre on Nature Conservation from the Institute of Terrestrial Ecology, Monks Wood, Cambridgeshire. [Final draft with further revisions to marine habitats.], Brussels: European Environment Agency.
  16. Egglishaw, H.J., 1960. The life history of Fucellia maritima (Haliday) (Diptera, Muscidae). Entomologist, 93, 225-231.
  17. Eltringham, S.K., 1971. Life in mud and sand. London: The English Universities Press Ltd.
  18. EnCams, 2002. Environmental Campaigns Seaside Awards. www.seasideawards.org.uk, 2002-08-06
  19. Fanini, L., Marchetti, G.M., Baczewska, A., Sztybor, K. & Scapini, F., 2012. Behavioural adaptation to different salinities in the sandhopper Talitrus saltator (Crustacea: Amphipoda): Mediterranean vs Baltic populations. Marine and Freshwater Research, 63 (3), 275-281.

  20. Fanini, L., Marchetti, G.M., Scapini, F. & Defeo, O., 2009. Effects of beach nourishment and groynes building on population and community descriptors of mobile arthropodofauna. Ecological Indicators, 9 (1), 167-178.

  21. Fanini, L., Zampicinini, G. & Pafilis, E., 2014. Beach parties: a case study on recreational human use of the beach and its effects on mobile arthropod fauna. Ethology Ecology & Evolution, 26 (1), 69-79.

  22. FEE, 2002. Foundation for Environmental Education, Blue Flag Awards. www.blueflag.org, 2002-08-06
  23. Fialkowski, W., Rainbow, P.S., Fialkowska, E. & Smith, B.D., 2000. Biomonitoring of trace metals along the Baltic coast of Poland using the sand hopper Talitrus saltator (Montagu) (Crustacea, Amphipoda). Ophelia, 52, 183-192.
  24. Foster, P., Hunt, D.T.E. & Morris, A.W., 1978. Metals in an acid mine stream and estuary. Science of the Total Environment, 9, 75-86.
  25. Fowles, A., 1994. Invertebrates of Wales: a review of important sites and species. Invertebrates of Wales: a review of important sites and species., JNCC Peterborough.
  26. Griffiths, C.L. & Stenton-Dozey, J., 1981. The fauna and rate of degradation of stranded kelp. Estuarine, Coastal and Shelf Science, 12, 645-653.
  27. Harrison, P.G., 1977. Decomposition of macrophyte detritus in seawater: effects of grazing by amphipods. Oikos, 28, 165-170.
  28. Healy, B., 1979. Records of Enchytraeidae (Oligochaeta) Ireland. Journal of Life Sciences, Royal Dublin Society, 1, 39-70.
  29. Hurley, D.E., 1959. Notes on the ecology and environmental adaptations of the terrestrial Amphipoda. Pacific Science, 13, 107-129.
  30. Hurley, D.E., 1968. Transition from water to land in amphipod crustaceans. American Zoologist, 8, 327-353.
  31. Ignaciuk, R. & Lee, J.A., 1980. The germination of four annual strand-line species. New Phytologist, 84, 581-591.
  32. Jelassi, R., Bohli-Abderrazak, D., Ayari, A. & Nasri-Ammar, K., 2015. Endogenous activity rhythm in Talitrus saltator, Britorchestia brito (Crustacea, Amphipoda) and Tylos europaeus (Crustacea, Isopoda) from Barkoukech Beach (Tabarka, Tunisia). Biological Rhythm Research, 46 (6), 873-886.

  33. JNCC, 2015. The Marine Habitat Classification for Britain and Ireland Version 15.03. JNCC: JNCC. 2015(20/05/2015). jncc.defra.gov.uk/MarineHabitatClassification
  34. JNCC (Joint Nature Conservation Committee), 1999. Marine Environment Resource Mapping And Information Database (MERMAID): Marine Nature Conservation Review Survey Database. [on-line] http://www.jncc.gov.uk/mermaid,
  35. Junoy, J., Castellanos, C., Vieitez, J.M. & Riera, R., 2013. Seven years of macroinfauna monitoring at Ladeira beach (Corrubedo Bay, NW Spain) after the Prestige oil spill. Oceanologia, 55 (2), 393-407.

  36. Junoy, J., Castellanos, C., Vieitez, J.M., de la Huz, M.R. & Lastra, M., 2005. The macroinfauna of the Galician sandy beaches (NW Spain) affected by the Prestige oil-spill. Marine Pollution Bulletin, 50 (5), 526-536.

  37. Koop, K. & Griffiths, C.L., 1982. The relative significance of bacteria, meio- and macrofauna on an exposed sandy beach. Marine Biology, 66, 295-300.
  38. Koop, K. & Lucas, M.I., 1983. Carbon flow and nutrient regeneration from the decomposition of macrophyte debris in a sandy beach microcosm. In Proceedings of the First international Symposium on Sandy Beaches, Port Elizabeth, South Africa, 17-21 January 1983. Sandy beaches as ecosystems, (ed. A. McLachlan & T. Erasmus), pp. 249-262. The Hague, Dr W. Junk.
  39. Koop, K., Newell, R.C. & Lucas, M.I., 1982. Microbial regeneration of nutrients from the decomposition of macrophyte debris on the shore. Marine Ecology Progress Series, 9, 91-96.
  40. Lack, P. (Ed.), 1986. The atlas of wintering birds in Britain and Ireland. T. & A.D. Poyser.
  41. Laughlin, R., Linden, O. & Guard, H., 1982. Toxicity of tributyltins and leachates from antifouling paints on marine amphipods. , Karlskrona: Institutet for Vattenoch Luftvardsforskning.
  42. Lincoln, R.J., 1979. British Marine Amphipoda: Gammaridea. London: British Museum (Natural History).
  43. Llewellyn, P.J. & Shackley, S.E., 1996. The effects of mechanical beach-cleaning on invertebrate populations. British Wildlife, 7, 147-155.
  44. McLachlan, A., 1983. Sandy beach ecology - a review. In Sandy beaches as ecosystems (ed. A. McLachlan & T. Erasmus), pp.321-381. The Hague: Dr W. Junk Publishers.
  45. Morritt, D., 1988. Osmoregulation in littoral and terrestrial talitroidean amphipods (Crustacea) from Britain. Journal of Experimental Marine Biology and Ecology, 123, 77-94.
  46. Picton, B.E. & Costello, M.J., 1998. BioMar biotope viewer: a guide to marine habitats, fauna and flora of Britain and Ireland. [CD-ROM] Environmental Sciences Unit, Trinity College, Dublin., http://www.itsligo.ie/biomar/
  47. Pienkowski, M.W., 1982. Diet and energy intake of grey and ringed plovers, Pluvialis squatarola and Charadrius hiaticula, in the non-breeding season. Journal of Zoology, 197, 511-549.
  48. Pugh, P.J.A., Llewllyn, P.J., Robinson, K. & Shackley, S.E., 1997. The association of phoretic mites (Acarina: Chelicerata) with sand-hoppers (Amphipoda: Crustacea) on the South Wales coast. Journal of Zoology, 243, 305-318.
  49. Rainbow, P.S. & Moore, P.G., 1990. Seasonal variation in copper and zinc concentrations in three talitrid amphipods (Crustacea). Hydrobiologia, 196, 65-72.
  50. Rainbow, P.S., Fialkowski, W. & Smith, B.D., 1998. The sand hopper Talitrus saltator as a trace metal biomonitor in the Gulf of Gdansk, Poland. Marine Pollution Bulletin, 36, 193-200.
  51. Rainbow, P.S., Moore, P.G. & Watson, D., 1989. Talitrid amphipods (Crustacea) as biomonitors for copper and zinc. Estuarine, Coastal and Shelf Science, 28, 567-582.
  52. Reyes-Martínez, M.J., Ruíz-Delgado, M.C., Sánchez-Moyano, J.E. & García-García, F.J., 2015. Response of intertidal sandy-beach macrofauna to human trampling: An urban vs. natural beach system approach. Marine Environmental Research, 103, 36-45.

  53. Robertson, A.I. & Mann, K.H., 1980. The role of amphipods in the initial fragmentation of eelgrass detritus in Nova Scotia, Canada. Marine Biology, 59, 63-69.
  54. Rodil, I.F., Lucena-Moya, P., Olabarria, C. & Arenas, F., 2015. Alteration of Macroalgal Subsidies by Climate-Associated tressors Affects Behavior of Wrack-Reliant Beach Consumers. Ecosystems, 18 (3), 428-440.

  55. Rossi, F., Olabarria, C., Incera, M. & Garrido, J., 2010. The trophic significance of the invasive seaweed Sargassum muticum in sandy beaches. Journal of Sea Research, 63 (1), 52-61.

  56. Salisbury, E., 1952. Downs and dunes. G. Bell & Sons.
  57. Scapini, F., Chelazzi, L., Colombini, I. & Fallaci, M., 1992. Surface activity, zonations and migrations of Talitrus saltator on a Mediterranean beach. Marine Biology, 112, 573-581.
  58. Shackley, S.E. & Llewellyn, P.J., 1994. Factors affecting the invertebrate population abundance of the strand-line, the littoral fringe and supralittoral zones, of soft sediment beaches. , Unpublished report, Countryside Council for Wales.
  59. Shackley, S.E. & Llewellyn, P.J., 1997. Sediment shore impact assessment/monitoring: strandline fauna. CCW Sea Empress Contract Science Report, 240., CCW Sea Empress Contract Science Report, 240.
  60. Smith, J.E. (ed.), 1968. 'Torrey Canyon'. Pollution and marine life. Cambridge: Cambridge University Press.
  61. Spicer, J.I., Morritt, D. & Taylor, A.C., 1994. Effect of low temperature on oxygen uptake and haemolymph ions in the sand hopper Talitrus saltator (Crustacea: Amphipoda). Journal of the Marine Biological Association of the United Kingdom, 74, 313-321.
  62. Stenton-Dozey, J. & Griffiths, C.L., 1980. Growth, consumption and respiration by larvae of the kelp-fly Fucellia capensis. South African Journal of Zoology, 15, 280-283.
  63. Ugolini, A., Pasquali, V., Baroni, D. & Ungherese, G., 2012. Behavioural responses of the supralittoral amphipod Talitrus saltator (Montagu) to trace metals contamination. Ecotoxicology, 21 (1), 139-147.

  64. Ugolini, A., Ungherese, G., Ciofini, M., Lapucci, A. & Camaiti, M., 2013. Microplastic debris in sandhoppers. Estuarine Coastal and Shelf Science, 129, 19-22.

  65. Ugolini, A., Ungherese, G., Ciofini, M., Lapucci, A. & Camaiti, M., 2013. Microplastic debris in sandhoppers. Estuarine Coastal and Shelf Science, 129, 19-22.

  66. Ugolini, A., Ungherese, G., Somigli, S., Galanti, G., Baroni, D., Borghini, F., Cipriani, N., Nebbiai, M., Passaponti, M. & Focardi, S., 2008. The amphipod Talitrus saltator as a bioindicator of human trampling on sandy beaches. Marine Environmental Research, 65 (4), 349-357.

  67. Williams, J.A., 1976. The effect of light on the locomotor activity and general biology of Talitrus saltator (Montagu). , Ph.D. thesis, Liverpool University, 258 pp.
  68. Williams, J.A., 1978. The annual pattern of reproduction of Talitrus saltator (Crustacea: Amphipoda: Talitidae). Journal of Zoology, 184, 231-244.
  69. Williams, J.A., 1980. The light response rhythm and seasonal entrainment of the endogenous circadian locomotor of Talitrus saltator (Crustacea: Amphipoda). Journal of the Marine Biological Association of the United Kingdom, 60, 773-785.
  70. Williams, J.A., 1983b. Environmental regulation of the burrow depth distribution of the sand-beach amphipod Talitrus saltator. Estuarine, Coastal and Shelf Science, 16, 291-298.
  71. Williamson, D.I., 1951. Studies in the biology of Talitridae (Crustacea, Amphipoda): effects of atmospheric humidity. Journal of the Marine Biological Association of the United Kingdom, 30, 73-90.

Citation

This review can be cited as:

Budd, G.C. 2004. Talitrids on the upper shore and strand-line. In Tyler-Walters H. and Hiscock K. (eds) Marine Life Information Network: Biology and Sensitivity Key Information Reviews, [on-line]. Plymouth: Marine Biological Association of the United Kingdom. Available from: http://www.marlin.ac.uk/habitat/detail/176

Last Updated: 27/10/2004