Swimming ability of eels (<Emphasis Type="Italic">Anguilla rostrata,Conger oceanicus</Emphasis>) at estuarine ingress: contrasting patterns of cross-shelf transport?

The transport of eel early life stages may be critical to their population dynamics. This transport from ocean spawning to freshwater, estuarine and coastal nursery areas is a combination of physical and biological processes (including swimming behavior). In New Jersey, USA, the American eel (Anguilla rostrata) enters estuaries as glass eels (48.7–68.1 mm TL) in contrast to the Conger eel (Conger oceanicus) that enters as larger (metamorphosing) leptocephali (68.3–117.8 mm TL). To begin to understand the mechanisms of cross-shelf transport for these species, we measured the potential swimming capability (critical swimming speed, U _crit) under ambient conditions throughout the ingress season. A. rostrata glass eels were collected over many months (January–June) at a range of temperatures (4–21°C), with relative condition declining over the course of the ingress period as temperatures warmed. C. oceanicus occurred later in the season (April–June) and at warmer temperatures (14–24.5°C). Mean U _crit values for A. rostrata (11.7–13.3 cm s⁻¹) and C. oceanicus (14.7–18.6 cm s⁻¹) were comparable, but variable, with portions of the variability explained by water temperature, relative condition, ontogenetic stage, and fish length. Travel times to Little Egg Inlet, New Jersey, estimated using 50% U _crit values, indicate it would take A. rostrata ~30 and ~60 days to swim from the shelf edge and Gulf Stream, respectively. Travel times for C. oceanicus were shorter, ~20 days from the shelf edge, and ~45 days from the Gulf Stream. Despite differences in life stage, our results indicate both species are competent swimmers, and suggest they are capable of swimming from the Gulf Stream and/or edge of the continental shelf to estuarine inlets.     

Ontogenic change of gill chloride cells in leptocephalus and glass eel stages of the Japanese eel, <Emphasis Type="Italic">Anguilla japonica</Emphasis>

The development of gill chloride cells was examined in premetamorphic larvae (leptocephali) and juveniles (glass eels) of the Japanese eel, Anguilla japonica. Branchial chloride cells were detected by immunocytochemistry using an antiserum specific for Na⁺,K⁺-ATPase. The specificity and availability of the antiserum for the detection of Japanese eel chloride cells were confirmed by Western blot analysis. The chloride cells first appeared on the developing gill filaments in a mid larval stage of leptocephalus (32.2 mm). Both immunoreactivity and the number of chloride cells gradually increased as the fish grew to a late stage of leptocephalus over 54 mm. In glass eels just after metamorphosis, gill lamellae developed from the gill filaments, and a rich population of chloride cells was observed in the gill filaments. In glass eels collected at a coastal area, chloride cells were extensively distributed in the gill filaments. The chloride cell size decreased progressively in glass eels transferred from seawater (SW) to freshwater (FW), whereas there was no difference in cell number. In contrast, some Na⁺,K⁺-ATPase immunoreaction distinct from typical chloride cells was observed in the gill lamellae throughout FW-transferred fish, but disappeared in control fish maintained in SW for 14 days. These findings indicate that the gill and gill chloride cells developed slowly during the extremely long larval stage, followed by rapid differentiation during a short period of metamorphosis. The excellent euryhalinity of glass eels may be due to the presence of the filament chloride cells and lamellar Na⁺,K⁺-ATPase-immunoreaction, presumably being responsible for SW and FW adaptation, respectively.     

三丁基锡和菲对鳗鲡的毒性效应（Toxic Effects of Tributyltin and Phenanthrene on Eels）

采用腹腔注射方式(每周1次,共5次)将生活于淡水或海水中的日本鳗鲡(Anguilla japonica)暴露于三丁基锡氯化物(TBTCl)和菲(Phe),以研究这两种污染物对鳗鲡的毒性效应以及该效应与鱼体所处环境盐度的关系.鳗鲡暴露于TBTCl或Phe1w后,其肝脏和脾脏的组织结构就已发生明显病变,且病变程度随着暴露时间的延长而加剧,其中海水鳗鲡比淡水鳗鲡表现更为严重;但肝体指数(HSI)和性腺组织结构均无明显变化.暴露于TBTCl4～5w后,海水鳗鲡肝脏的谷胱甘肽-S-转移酶(GST)活性比对照组显著升高,而淡水鳗鲡组无明显变化;暴露于TBTCl或Phe3w后,淡水鳗鲡的血浆皮质醇水平显著升高,但海水鳗鲡组无明显变化.TBTCl和Phe对淡水或海水鳗鲡的血浆雌二醇(E2)水平均无明显影响.以上结果表明:有机锡和菲对鳗鲡的肝脏、脾脏有毒性作用,对皮质醇分泌具有内分泌干扰作用,且该效应与暴露时间和鱼体所处环境盐度有关.     

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     

Ecological responses to reductions in freshwater supply and quality in South Africa’s estuaries: lessons for management and conservation

Fresh water, a fundamental element of all estuarine ecosystems, is South Africa’s most limited natural resource. Recent projections indicate that by the year 2020 the country will be utilizing all its exploitable freshwater sources. Steeply increasing demands by a rapidly growing population on this limited commodity have already resulted in a severe reduction of water supplies to natural users such as estuaries — this trend is predicted to increase in the future. Concurrent with excessive water abstraction, poor land husbandry (e.g. soil erosion) in many catchment basins and pollution (e.g. salinization) in return flows have led to a serious deterioration in water quality. In contrast, a review of estuarine responses to varying flow regimes stresses the strong dependence of local systems on riverine fresh water inputs of adequate quantity and quality. Freshwater dependence is i.a. expressed in: flooding events that scour accumulated sediments, riverine nutrient input to drive estuarine phyto- and zooplankton production, axial salinity gradients that increase habitat and species diversity, and maintenance of open tidal inlets that prevent salinity and temperature extremes and facilitate larval exchange, fish migrations and tidal flushing of salt marshes. Thus, estuarine conservation will have to encompass management of rivers and watersheds and play an increasingly political role in decision processes concerning water allocations among ‘human’ and ‘natural’ users.     

Changes in water and ion transport in isolated intestines of the eel during salt adaptation and migration

When Japanese cultured eels, Anguilla japonica, were transferred from freshwater to sea-water, their body weight decreased by 15% on the second day and then recovered gradually to the original weight within a week. Serum sodium content increased by 58% on the second day, and thereafter decreased to a constant level. Isolated intestines from these eels were bathed on both sides, mucosal and serosal, by identical Ringer solution. The water and sodium transort was very active from mucosa to serosa in such isolated intestines of sea-water-adapted eels. This active movement was particularly dominant during the week after transferring the eel to seawater, and a peak was reached on the fifth day. The molar ratio of water to sodium in the fluid moving across the intestinal wall was higher in sea-water-adapted eels than in freshwater eels. The isolated intestines taken from silver eels during their catadromous migration, but still in the river, showed a larger amount of water transport and a higher molar ratio of water to sodium than intestines from yellow sedentary eels. These adaptive properties are a prerequisite for the marine life of the eel.     

Winter mortality, growth, and behavior of young-of-the-year of four coastal fishes in New Jersey (USA) waters

Winter mortality has been hypothesized to select for large body size in young-of-the-year (YOY) fishes, yet substantiation of winter mortality and its cause(s) are available for few estuarine or marine species. We examined seasonal length distributions of wild populations of four common marine species, black sea bass (Centropristis striata), tautog (Tautoga onitis), cunner (Tautogolabrus adspersus), and smallmouth flounder (Etropus microstomus), and mortality (i.e., frequency of death), growth, and behavior of their YOY in the laboratory at ambient winter temperatures (mean 7°C, range 2-13°C) during a 135-day period (December 1992 through mid-April 1993) to establish potential causes of their mortality in the field. Young-of-the-year black sea bass experienced 100% mortality when water temperatures decreased to 2-3°C in February, emphasizing the importance of winter emigration from estuaries in this southern species. The low mortality of two labrid species, YOY tautog (14%) and YOY cunner (3%), was consistent with their northern distribution and year-round occurrence in estuarine and nearshore coastal waters. Laboratory mortality of YOY smallmouth flounder (33%) was higher for small (<35 mm total length) fish, suggesting that this small species may experience high winter mortality in estuaries and nearshore coastal waters. Seasonal differences in fish length result potentially from several mechanisms (e.g., mortality and/or migration) that are difficult to assess, but our laboratory experiments suggest that seasonal temperature changes cause size-specific mortality of YOY smallmouth flounder and offshore migration of YOY black sea bass.     

Relative contribution of migratory type on the reproduction of migrating silver eels, <Emphasis Type="Italic">Anguilla japonica</Emphasis>, collected off Shikoku Island,Japan

In order to understand the reproductive contribution among migratory types in the Japanese eel, Anguilla japonica, otolith strontium (Sr) and calcium (Ca) concentrations by X-ray electron microprobe analysis were examined for 37 silver eels collected in Kii Channel off Shikoku Island during the spawning migration season. The wide range of otolith Sr:Ca ratios indicated that the habitat use of A. japonica was not obligatory but facultative among fresh, brackish and marine waters during their growth phases after recruitment to the coastal areas as glass eels. Three migratory types, which were categorized as river eels, estuarine eels and sea eels were found. The estuarine eels were dominant (59%), followed by sea eels (22%) and river eels (19%). The low proportion of river eels from the spawning migration season suggested that the estuarine and sea eels inhabiting the nearby coastal areas might make a larger reproductive contribution to the next generation in this area.     

Factors influencing community structure and distribution of different life-cycle categories of fishes in shallow waters of a large Australian estuary

Fish were collected at regular intervals over 5 yr (February 1977 to December 1981) from ten shallow-water sites located throughout the lower, middle and upper regions of the large Swan Estuary in temperate southwestern Australia. Analysis of the catch data showed that the total number of species and total density of fishes were both influenced to a greater extent by site and season within the estuary than by year. The number of species and density of fishes within the whole system were greatest during the summer and autumn, when salinities and temperatures were at a maximum, and declined with distance from the estuary mouth. This reflects the trends shown by marine species, which comprise many species that occur only occasionally in the estuary (marine stragglers) and others which enter estuaries regularly and in considerable numbers (marine estuarine-opportunists). The density of marine estuarine-opportunists was also correlated with temperature, reflecting the tendency for the species of this category to congregate in the shallows during the summer and autumn. The similar seasonal aggregations of the single anadromous species and representatives of species that complete their whole life cycle in the estuary were frequently related to spawning. The density of the estuarine category was correlated neither with salinity nor distance from the estuary mouth. The number of estuarine species was also not correlated with distance from the estuary mouth. The density of freshwater species was inversely correlated with salinity and positively correlated with distance from the estuary mouth. The composition of the fish fauna changed progressively through the estuary, with that of the lower estuary being the most discrete. The composition also changed seasonally, particularly in the upper estuary where, during the winter and spring, the volume of freshwater discharge increased greatly and as a consequence the salinity declined markedly. The species diagnostic of the lower estuary were generally marine estuarine-opportunists, whereas those of the upper estuary typically belonged to either the estuarine or anadromous categories. The marine estuarine-opportunistMugil cephalus was, however, also one of the diagnostic species in the upper estuary during the winter and spring.Please address all correspondence and requests for reprints to I. C. Potter at Murdoch University     

Ontogeny of osmoregulation in the European sea bass Dicentrarchus labrax L.

 Under natural conditions, sea bass eggs hatch in the open sea and young-of-the-year sea bass are found close to estuaries and enter brackish and even freshwater lagoons in the Mediterranean and the eastern Atlantic. The ontogeny of osmoregulation in the European sea bass, Dicentrarchus labrax (Linnaeus, 1758), was studied in different developmental stages, from hatching to large juveniles, exposed to a range of salinities at 17 °C. The experiments were carried out from March to August 1998 in southern France. The type of hyper-hypo-osmotic regulation did not change during development. All stages hyper-regulated at low salinity (under 10–11‰) and hypo-regulated at higher salinities. The acquisition of the full ability to hypo-regulate occurred in four steps. Osmoregulatory capacity was size- and age-dependent and reached its maximum for fish 17–26 mm long, 63–86 days after hatching. The iso-osmotic salinity was 10.2–11.6‰. Our results suggest that early development of osmoregulatory ability, and thus of salinity tolerance in sea bass, may provide an advantageous flexibility for the timing of migration towards low-salinity habitats. Received: 22 May 2000 / Accepted: 18 December 2000     

Mercury bioaccumulation in estuarine food webs

We tested for unintended mercury contamination problems associated with estuarine floodplain restoration projects of the Louisiana coastal zone, USA. Barataria Bay and Breton Sound are two neighboring deltaic estuaries that were isolated by levees from the Mississippi River about 100 years ago. These estuaries recently have been reconnected to the nutrient-rich Mississippi River, starting major river diversion (input) flows in 1991 for Breton Sound and in 2004 for Barataria Bay. We collected > 2100 fish over five years from 20 stations in these estuaries to test two hypotheses about Hg bioaccumulation: (H1) Background Hg bioaccumulation in fish would be highest in low-salinity upper reaches of estuaries, and (H2) recent river inputs to these upper estuarine areas would increase Hg bioaccumulation in fish food webs. For H1, we surveyed fish Hg concentrations at several stations along a salinity gradient in Barataria Bay in 2003-2004, a time when this estuary lacked strong river inputs. Results showed that average Hg concentrations in fish communities were lowest (150 ng/g dry mass) in higher salinity areas and -2.4x higher (350 ng/g) in low-salinity oligohaline and freshwater upper reaches of the estuary. For H2, we tested for enhanced Hg bioaccumulation following diversion onset in both estuaries. Fish communities from Breton Sound that had long-term (> 10 years) diversion inputs had -1.7x higher average Hg contents of 610 ng/g Hg vs. 350 ng/g background values. Shorter-term diversion inputs over 2-3 years in upper Barataria Bay did not result in strong Hg enrichments or stable C isotope increases seen in Breton Sound, even though N and S stable-isotope values indicated strong river inputs in both estuaries. It may be that epiphyte communities on abundant submerged aquatic vegetation (SAV) are important hotspots for Hg cycling in these estuaries, and observed lesser development of these epiphyte communities in upper Barataria Bay during the first years of diversion inputs may account for the lessened Hg bioaccumulation in fish. A management consideration from this study is that river restoration projects may unintentionally fertilize SAV and epiphyte-based food webs, leading to higher Hg bioaccumulation in river-impacted floodplains and their food webs.     

Respiratory responses of the estuarine mysid Neomysis integer (Peracarida: Mysidacea) in relation to a variable environment

The euryhaline mysid Neomysis integer (Peracarida: Mysidacea) is a common member of the hyperbenthos of the upper reaches of European estuaries. In the East Looe River Estuary (Cornwall, England), this species experiences extensive tidal and seasonal changes in temperature (3 to 15 °C) and salinity (1 to 34‰). In this investigation, the effects of temperature (5, 10 and 15 °C) and salinity (1, 10, 20 and 30‰) on the oxygen consumption of male and female N. integer are reported, and are related to field measurements to identify the adaptive responses of the respiratory physiology to such a variable environment. The general responses were similar for each sex; however, at any given temperature/salinity combination, male N. integer consumed more oxygen than females. The general trends were increased oxygen consumption with increasing temperature (Q₁₀ values ranged from ∼1.7 to 2.5) and decreased oxygen consumption with increasing salinity. Temperature and salinity interacted at high water antagonistically to minimise changes in mysid oxygen-consumption. When related to tidal fluctuations in temperature and salinity experienced by N. integer inhabiting the East Looe River Estuary, the results reveal how the respiratory physiology of this species is adapted to its variable environment. Received: 16 June 1998 / Accepted: 15 December 1998     

Effects of sudden changes in salinity on endogenous rhythms of the spotted sea bass Lateolabrax sp.

The endogenous rhythm of oxygen consumption in juvenile spotted sea bass (Lateolabrax sp.) was measured to test the effects of sudden changes in salinity on the metabolic activity. Mean oxygen consumption rates of this euryhaline fish decreased by 13.5 to 16.0% and 25.3 to 36.4% when they were transferred from 31.5 to 15‰ seawater and to fresh water (0‰), respectively. The maximum rate of oxygen consumption was observed between 18:00 and 19:00 hrs local time, 1 to 2 h before sunset, even though they were kept in constant darkness. The peaks of oxygen consumption occurred in 23.2- and 23.3-h intervals, which correspond with a circadian rhythm, as revealed by maximum entropy spectral analysis. A markedly weakened rhythm in oxygen consumption occurred from 8 to 10 d after onset of the experiments. This study indicates that spotted sea bass can withstand sudden drops in salinity from 31.5‰ to fresh water, and yet maintain a regular though somewhat dampened endogenous rhythm of oxygen consumption. Received: 16 June 1997 / Accepted: 3 February 1998     

Sensory ecology in a changing world: salinity alters conspecific recognition in an amphidromous fish, Pseudomugil signifer

Group fission and fusion processes are driven by state dependence, risk and the availability of information from others. Yet the availability of information changes under different environmental conditions, thus aiding or inhibiting group formation and maintenance. Chemical cues provide information on the location of individuals and can act as a mechanism for individuals to group together, although they can be greatly affected by environmental conditions. Using a flow channel, we studied how one shoaling fish species, the Pacific blue-eye (Pseudomugil signifer), responds to conspecific chemical cues (CCCs) in different environmental conditions (salinities). This species lives in estuarine environments, ranging in salinity from fresh to fully marine. P. signifer responded to CCCs in freshwater but not in saltwater. Furthermore, P. signifer did not respond to saltwater with CCCs added from freshwater. It took significantly longer for fish in saltwater, than in freshwater, to locate and join a shoal when only CCCs from the shoal were present. Finally, fish formed more cohesive shoals in freshwater than in brackish or saltwater. These results suggest that these fish do not rely on chemical cues in saltwater to locate conspecific shoals. Furthermore, the reduced amounts of these cues in saltwater may inhibit the maintenance of tight shoal structures. We suggest that fish utilise different sensory modalities in fresh or saltwater in order to locate one another, or the social structure of these groups is fundamentally different between these two water types. The importance of this study in relation to understanding how animals utilise and change different sensory modalities in varying environmental conditions is discussed.     

Changes in intestinal absorption and renal excretion of water during adaptation to sea-water in the Japanese eel

The rate of sea-water ingestion in Japanese cultured eels, Anguilla japonica, adapted to sea-water for one week, was 89±9.5 ml/kg/day; approximately 75% of the ingested water was absorbed by the intestine. Similar rates of ingestion and absorption of water were also found with sea-water-adapted rainbow trout, Salmo gairdnerii, and two species of marine teleosts, Stephanolepis cirrhifer and Goniistius zonatus. When cultured eels were transferred from freshwater to sea-water the amount of ingested and absorbed water inoreased gradually to a maximum on the 5^th day and then decreased to a constant level. The urine flow and glomerular filtration rate were found to be high in freshwater cultured eels, and both reduced markedly within 6 h after transferring the eel to sea-water. The reabsorption of water by the renal tubules was very small in both the freshwater and sea-water-adapted (6 h) eels. On the other hand, in the eels adapted to sea-water for 10 days the rate of urine flow was also very slight but the glomerular filtration rate recovered to that of the freshwater eels. Thus, water reabsorption by the tubules increased to a large degree. The results seem to indicate the existence of two distinct phases in the renal function during sea-water adaptation of cultured eels.     

Photosynthesis in Codium fragile (Chlorophyta) from a Nova Scotia estuary: responses to desiccation and hyposalinity

Codium fragile ssp. tomentosoides from Caribou Harbour, an estuarine site in the southern Gulf of St. Lawrence, was extremely tolerant to stresses from desiccation and reduced salinity. Photosynthetic responses of both rhizomatous and erect growth forms were measured using pulse amplitude modulation (PAM) fluorometry of chlorophyll a fluorescence to determine effective quantum yield (Φ_PSII) and relative electron transport rate (rETR). After 5 h of desiccation, thalli lost 20% of their mass, but still showed high levels of Φ_PSII. Thalli survived for at least 6 h in freshwater, and showed virtually complete recovery of photosynthetic capacity within a few hours of return to full seawater. Immersion in 8 psu showed virtually complete recovery until the 24 h treatment period. Combining desiccation and salinity stresses produced a synergistic effect, but plants still showed strong recovery even after 86% dehydration and reimmersion in 16 psu. These results suggest that the photosynthetic physiology of Codium fragile is highly adapted to growth in estuarine conditions.     

Temperature and salinity: two climate change stressors affecting early development of the New Zealand sea urchin Evechinus chloroticus

Temperature and salinity are important environmental factors affecting the normal functioning of marine animals, particularly animals such as sea urchins living in shallow waters and tide pools. Here, we evaluated the effect of different combinations of temperature and salinity on early embryos of the endemic New Zealand sea urchin Evechinus chloroticus. Animals were collected at Matheson’s Bay (36º18′17′′S; 174º47′51′′E) in north-eastern New Zealand in February 2013. Embryos were exposed to five salinities (29, 31, 34, 35 and 37 ppt) and two temperatures (18 and 21 °C) during the first 24 h of development. Low salinity (29 ppt) affected all parameters (fertilization, development rate, gastrulation and normal development), with ca. 50 % of embryos surviving at 29 ppt, whereas seawater temperature only affected development rate and gastrulation. An increase in temperature from 18 to 21 °C minimized the negative effect of low salinity (≤31 ppt) on development rate and gastrulation of E. chloroticus. Overall, the results of this study suggest that early embryos of E. chloroticus have developmental plasticity to withstand reductions in salinity up to 29 ppt; however, it is still unknown whether the surviving embryos will be able to complete larval development at low salinities, particularly whether the embryos and larvae are carried into extreme environments such as estuaries where salinity is even lower. Multistressor studies are very important for climate change research as multiple environmental factors will act together in the wild, having major consequences for development and recruitment of marine invertebrates.     

Influence of habitat on abundance and size structure of a large temperate-reef fish, Achoerodus viridis (Pisces: Labridae)

The relationship between densities of Achoerodus viridis (Pisces: Labridae) and reef habitats at various localities within New South Wales (NSW), Australia was examined. Types of habitats were quantified at inner, mid and outer estuarine locations in each of two estuaries (Botany Bay and Port Jackson) to determine whether purported patterns of movement from estuaries could be related to differences in habitat. Although the same types of habitat were generally found at all locations, differences in the proportion of habitat types were found between shallow and deep reefs and among inner, mid and outer estuarine locations for both estuaries. Shallow habitats were usually dominated by Ecklonia radiata, turf and/or fringe habitat in Botany Bay, whereas deep sites were generally dominated by urchin-grazed barrens habitat and, sometimes, sponge- and ascidian-dominated deep reef. Shallow sites in Port Jackson were dominated by a mixture of habitats, as were deep reefs at mid-estuarine locations. Other deep reefs in Port Jackson were dominated by E. radiata (inner estuarine) or barrens (outer estuarine) habitat. Thus, patterns of habitat cover were not consistent between estuaries and numbers of fish could not be related to proportional representation of habitat on reefs along estuarine gradients. Univariate and multivariate analyses showed that there was little evidence that any size class of fish was correlated with the proportional representation of a particular habitat or group of habitats. Counts of fish that focused on barrens and E. radiata forest habitats over a period of 10 yr showed that similar numbers and all sizes of fish were found in the two types of habitat. Greater numbers of small fish were, however, found in the E. radiata forest habitat than in the barrens habitat. Estimates of abundance along the coast of NSW (100s to 1000 km) in a range of habitats (e.g. ascidian-dominated reef, kelp forest, urchin-grazed barrens) showed that there was no indication that a particular habitat consistently had greater numbers of A. viridis than other habitats. Therefore, A. viridis of a range of sizes appears to be flexible in its use of habitats on reefs. Received: 24 December 1997 / Accepted: 23 June 1998     

Vegetation succession of low estuarine marshes is affected by distance to navigation channel and changes in water level

Climate change and engineering activities have modified the hydrology and morphology of estuaries. However, the potential effects of these modifications on vegetation succession in estuarine marshes are still poorly understood. Therefore, we studied temporal changes in tidal habitats of the Elbe estuary over a period of 30 years. We compared vegetation maps from 1980 to 2010 and calculated the change in area of habitats with respect to three salinity and three elevational zones. To analyze the direction of the temporal change, we differentiated between progressive and regressive succession. By using regression tree models (conditional inference trees), we identified the most influential factors determining progressive or regressive succession of low marshes. The total area of the estuarine tidal marshes at the Elbe increased by 2 % from 1980 to 2010, but changes were unequal among the salinity zones. In the salt and brackish zones, the area covered by high marshes increased substantially but decreased in the tidal freshwater zone, while that covered by low marshes decreased in all the salinity zones. Additionally, we determined high persistence of tidal flats and high marshes, whereas only 19 to 28 % of the low marshes found in 1980 remained in 2010. In salt and brackish marshes, more than two-thirds of the area that had been identified as low marshes in 1980 had progressively developed into high marshes. In contrast, 44 % of the area of low marshes in tidal freshwater marshes showed regressive succession back into tidal flats. The distance to the navigation channel was the main factor determining successional direction in salt and brackish marshes. Here, greater proximity to the channel was correlated with higher risk of regressive succession. In tidal freshwater marshes, we identified both the distance to the navigation channel and the situation on the river shore (i.e. inner bank, outer bank or straight bank) as the main factors for marsh succession. Here, considerable engineering activities in the channel had simultaneously decreased the mean low water level and increased the mean high water level between 1980 and 2010, which led to an increase in tidal amplitude. It is quite likely that these changes negatively modified marsh distribution, increased regressive succession and, thus, lowered the quality of tidal freshwater marshes.