Density dependent and density independent relationships during a twenty-seven year study of the population dynamics of the benthic macroinvertebrate community of a chemically unstable lake

Data are given on net precipitation, water chemistry, vegetation cover and the population dynamics of 20 taxa of benthic macroinvertebrates from the total of 40 recorded ln the lake. The relationships between data sets were investigated by bivariate analysis. The relationships between abiotic factors are best described by linear equations as are some relationships between Sigara concinna, Sigara dorsalis, Potamopyrgus jenkinsi and environmental factors. The relationships of numbers of Gammarus tigrinus with temperature, Sigara stagnalis with conductivity, Corixidae with time and Sigara lateralis, Theromyzon tessulatum and Piscicola geometra with vegetation cover are best described by exponential equations. The relationships between the population dynamics of pairs of certain taxa viz. Gammarus tigrinus and G. duebeni, G. tigrinus and Asellus aquaticus, G. tigrinus and Corixidae, Sigara dorsalis and S. concinna, are best described by power equations. The relationship of vegetation cover with time and the population dynamics of many taxa are best described by logarithmic logistic equations.Both density dependent and density independent relationships appear to be responsible for changes in numbers and taxa within the total community. Density independent relationships are most important at times of environmental change while density dependent relationships, though operating continuously, are most important when conditions are stable.     

Recent drastic changes in the gammarid fauna (Crustacea, Amphipoda) of the Vistula River deltaic system in Poland caused by alien invaders

During the last decade of 20th century, the nonindigenous gammarid species Gammarus tigrinus, Dikerogammarus haemobaphes, Pontogammarus robustoides and Obesogammarus crassus invaded the lower Vistula River and its deltaic, partly brackish regions. G. tigrinus, an oligohaline North‐American species, was introduced to western Europe in the 1950s; the remaining three species are oligohaline/freshwater Ponto‐Caspian species. All these species are now invading central and western Europe using the network of man‐made canals connecting different European river systems. In the Vistula River, the native European freshwater gammarid species Gammarus pulex and G. varsoviensis were replaced in the 1920s by the Ponto‐Caspian Chaetogammarus ischnus (syn. Echinogammarus ischnus), which in turn has been outnumbered by the more recent invasions of D. haemobaphes and P. robustoides. In brackish waters, the native Atlantic‐boreal species Gammarus zaddachi and Gammarus duebeni are replaced or at least outnumbered by G. tigrinus, P. robustoides and O. crassus. Possible invasion routes are discussed.     

Invasions of alien gammarid species and retreat of natives in the Vistula Lagoon (Baltic Sea, Poland)

During the last decades of the twentieth century, the alien gammarid species Gammarus tigrinus, Dikerogammarus haemobaphes, Pontogammarus robustoides and Obesogammarus crassus invaded the lower Vistula River and its deltaic, partly brackish regions. In brackish waters of the Vistula Lagoon the native Atlantic-boreal species Gammarus zaddachi and Gammarus duebeni have been replaced or at least outnumbered by the aliens. As compared to our earlier studies, through the years 1998–2004 we could observe nearly total decline of the native gammarid populations along the coasts of the Lagoon, and overdomination of the North-American G. tigrinus in most places. Possible reasons for the observed phenomena are e.g. increasing pollution and eutrophication of the Lagoon accompanied by competition between the native and the alien species.     

Intraguild predation and species exclusions in amphipods: the interaction of behaviour, physiology and environment

1. Data from field surveys, laboratory experiments and computer simulations of community dynamics revealed that a novel interaction among intraguild predation, physiological adaptation and environment may explain the complex distributions of two putatively competing aquatic amphipods. 2. Gammarus pulex and G. tigrinus both thrive in fresh and oligohaline waters in western Europe. However, the native European G. pulex excludes the invading North American G. tigrinus from freshwaters of relatively low conductivity, whereas the reverse occurs at higher conductivities. Additionally, there is much spatio-temporal fluctuation in the patterns of coexistence of these species. 3. Laboratory experiments in The Netherlands and Ireland revealed that mutual predation of moulting individuals occurred frequently between these species. However, predation frequencies were differentially in favour of G. pulex under the ionic conditions to which this species is physiologically adapted (freshwater). On the other hand, predation was not differential under the ionic conditions to which G. tigrinus is physiologically adapted (oligohaline water). 4. A mathematical model, which extends the logistic equation to include mutual intraguild predation, simulated interactions over a range of values of relevant population parameters. This indicated that G. pulex would be excluded when balanced instantaneous rates of mutual predation were combined with the known greater reproductive output of G. tigrinus. However, this reproductive advantage is overcome by any relatively small bias in the instantaneous rate of predation favouring G. pulex, leading to the exclusion of G. tigrinus. This occurs even when the reproductive advantage to G. tigrinus is relatively large. Moreover, the model generated ‘switches’ in species dominance that are determined by the relative values of reproductive rate and mutual predation. The time taken to ‘switch’ may explain the transient periods of apparent coexistence of these species observed in the field. 5. The complex community dynamics of such species may thus be understood in terms of variation in the intensity of species interactions mediated by behavioural, physiological and environmental factors.     

The role of ion regulation in the control of the distribution of Gammarus tigrinus (Sexton) in salt-polluted rivers

Fluctuating salinities at different sites on the German salt-polluted rivers Werra and Weser were compared with extracellular ion levels of specimens of Gammarus tigrinus (Sexton; Amphipoda, Crustacea), collected at the same sites. G. tigrinus regulated haemolymph concentrations of inorganic anions (Cl⁻, SO²⁻ ₄, PO³⁻ ₄) and cations (Na⁺, K⁺, Mg²⁺, Ca²⁺) during fluctuations of salt pollution in the upper Weser. This capacity to regulate varying levels of salt pollution in the upper Weser, correlated well with the distribution of the brackish amphipods in this river ecosystem. G. tigrinus tolerated periods of Na⁺ and Cl⁻ stress (>380 mmol l⁻¹) without compensating these maxima by regulating extracellular Na⁺ and Cl⁻. However, during such bursts of Na⁺ and Cl⁻ stress in Werra and Weser, the ability to regulate extracellular [K⁺] at river water K⁺ stress of ≥6.0 mmol l⁻¹ may explain why this brackish species has been more successful in these rivers than its competitors like Gammarus pulex. The present investigation demonstrates that the water salinity affects the [NO⁻ ₃] in the haemolymph of G. tigrinus. With increasing hypo-osmotic stress the animals accumulate increasing amounts of NO⁻ ₃. A simultaneous increase in stream water [NO⁻ ₃] causes an additional accumulation of NO⁻ ₃ in the haemolymph. The high extent of accumulation indicates that active ion transport systems may be involved. The accumulation of NO⁻ ₃ in the haemolymph has low physiological consequences to G. tigrinus, but when hypo-osmotically stressed under anoxic conditions, nitrite formed by the reduction of nitrate may have an adverse affect on the metabolism of G. tigrinus. Accepted: 4 October 1999     

Is <Emphasis Type="Italic">Gammarus tigrinus</Emphasis> (Crustacea,Amphipoda) becoming cosmopolitan through shipping? Predicting its potential invasive range using ecological niche modeling

While the intensity of global shipping has increased dramatically over the last decades, species exchange between continents has likewise intensified. Ballast water of ships is recognized playing a major role in this process. Many of the larger sea ports have become bridgeheads for invasions. Ecological niche modeling is used to investigate the potential invasive range and high invasive risk ports of the North American amphipod Gammarus tigrinus. Sixty-two occurrences of G. tigrinus in its native range (North America) and 34 environmental data sets were compiled. Data on dispersal distances were used via ecological niche modeling to analyze the invasive potential of G. tigrinus. The invasive risk of large ports was analyzed according to modeling result, as well as their salinity in the main oceanic routes of the world. G. tigrinus had a rapid range extension on the British Isles and in the rest of Western Europe. Now it is invading the countries surrounding the Baltic Sea. Worldwide it has a vast potential invasive range. It has a high invasive risk for many large ports along the main oceanic routes, among which the ports of Shanghai, Buenos Aires and Montevideo have the highest invasive risk. G. tigrinus may become cosmopolitan through shipping, and this possibility is increasing. Particular emphasis should be placed on preventing human-mediated dispersal. Ports may be the first places G. tigrinus invades. This study can identify high invasive risk ports, especially those at risk of introduced North America species. More importantly, the water of large ports should be monitored regularly for exotic aquatic organisms that may survive temporarily or permanently.     

Glacial-driven vicariance in the amphipod Gammarus duebeni

We have examined the genetic diversity using mitochondrial COI and ND2 sequence data from 306 specimens of the amphi-Atlantic-distributed amphipod Gammarus duebeni. Marine populations from the Atlantic Ocean, the Baltic and North Sea, as well as freshwater populations from Ireland, Cornwall and Brittany were analysed.G. duebeni is a complex of five allopatric lineages. Freshwater populations result from multiple invasions of marine ancestors, represented by distinct lineages. We interpret the recent distribution of lineages as the outcome of a series of spatio-temporal vicariant events caused by Pleistocene glaciations and sea level changes. The freshwater lineages are therefore regarded as ‘glacial relicts’. Furthermore, inter-specific competition with, for example, Gammarus pulex (which is absent in Ireland and western Brittany) may be another important determinant in the distribution of freshwater G. duebeni. In Ireland and Brittany, three freshwater refugia are suggested. The significantly limited gene flow detected among marine populations is more likely due to inter-specific competition than to salinity. The G. duebeni-complex represents a model system for the study of allopatric speciation accompanied by major habitat shifts. The pattern of spatio-temporal origins of the freshwater entities we describe here provides an excellent system for investigating evolutionary adaptations to the freshwater environment. Our data did not confirm the presently used subspecies classification but are only preliminary in the absence of nuclear genetic analyses.     

Rheotaktisches Verhalten einigerGammarus-Arten bei verschiedenem Sauerstoffgehalt des Wassers

Zusammenfassung 1. Zur Untersuchung kleinerer Wassertiere wurde eine Durchflußapparatur entwickelt, die eine stufenlose Regelung einer gerichteten Strömung sowie die Einstellung einer konstanten Gaskonzentration erlaubt.2. Das rheotaktische Verhalten vonGammarus pulex, G. roeseli, G. fossarum, G. tigrinus undG. salinus wurde bei einer Temperatur von 15° C unter verschiedenen Sauerstoffkonzentrationen getestet.3. Die höchste rheotaktische Aktivität der untersuchten Amphipoden wurde bei geringen Strömungsgeschwindigkeiten (5 cm/sec) konstatiert.4. Diese Aktivität erfuhr mit abnehmendem O₂-Gehalt eine Steigerung bis zu einem Aufwanderungsmaximum, das bei einer kritischen O₂-Konzentration lag. Noch geringerer Sauerstoffgehalt bewirkte Abwanderung in Strömungsrichtung, rasche Aktivitätsabnahme und führte meist zum Tod der Versuchstiere.5. Die so definierte kritische O₂-Konzentration lag fürGammarus pulex bei 2,7 mg O₂/l fürG. roeseli bei 3,1 mg O₂/l, fürG. fossarum bei 5,3 mg O₂/l und für die BrackwasserformG. tigrinus bei 3,5 mg O₂/l. Die euryhaline ArtG. salinus hatte keine ausgeprägte Rheotaxis; lediglich die Grenze vom Auf- zum Abwandern konnte definiert werden (2,5 mg O₂/l).

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Rheotactic behaviour of someGammarus species in different oxygen concentrations of the water

An experimental vessel has been developed which makes it possible to observe the behaviour of gammarids and other aquatic invertebrates in various water current speeds and oxygen concentrations. Studies have been conducted with the following limnic and brackish-water amphipods:Gammarus pulex (L.),G. roeseli Gervais,G. fossarum Koch,G. tigrinus Sexton andG. salinus Spooner. In adequate oxygen concentrations, the gammarids tested show a moderate positive rheotaxis. Lethal and sublethal oxygen concentrations, however, lead to negative rheotaxis. The maximum degree of upstream movement is attained in the region of the critical oxygen concentration. Under the experimental conditions (15° C, water current 5 cm/sec), these maxima vary according to species. The maximum occurred at 2.7 mg O₂/l forG. pulex, 3.1 mg O₂/l forG. roeseli, 3.5 mg O₂/l forG. tigrinus, and 5.3 mg O₂/l forG. fossarum. No such maximum was found forG. salinus. Oxygen deficit causesG. salinus to move downstream at 2.5 mg O₂/l.

     

Exotic and indigenous freshwater gammarid species: physiological tolerance to water temperature in relation to ionic content of the water

Bioinvasions by closely related species often lead to niche competition between exotic and indigenous species. The outcome of this competition is partly determined by differences in physiological tolerance of the competing species to the environmental conditions of the colonised habitat. Physiological tolerance of the invading gammarid species Gammarus tigrinus, Echinogammarus ischnus and Dikerogammarus villosus and the indigenous gammarid species Gammarus pulex, Gammarus roeseli and Gammarus fossarum from Dutch waters was studied in the laboratory by comparing their pleopod beats at rest at different water temperatures, which reflect the gammarid's oxygen consumption. Pleopod beat frequencies increased from a minimum ventilatory activity of 0 beats per minute at 1 °C to maximum activity of up to 300 beats per minute at temperatures between 25 °C and 35 °C. At the state of maximum activity, a further increase in temperature was followed by a strong decrease in pleopod beat frequency, indicating acute stress, and subsequently mortality. Frequency response patterns of invading and indigenous gammarids were found to be highly similar, indicating a wide tolerance to temperature for all species. The tolerance of D. villosus, however, was reduced in brook water, indicating a lower competitive ability in relatively ion-poor water. G. tigrinus survived at higher temperatures in the more ion-rich, polluted waters than the indigenous gammarids, indicating a wider physiological tolerance and thus a higher competitive ability in these waters.     

Formation of phenol-protein complexes and their use by two stream invertebrates

Ethanol and water extracts of maple leaves and pine needles were analyzed for proteins, amino acids, sugars and phenolics. Leachates were mixed with a dissolved fungal cellulase. Within 24 h, insoluble particles formed, consisting of phenolics, proteins and amino acids. When exposed to an alkaline pH, or to a 0.04% solution of the surfactant lysolecithin, these particles released amino acids and proteins. Surface tensions of the gut fluids of Gammarus tigrinus and Tipula caloptera were considerably lower than that of distilled water, suggesting the presence of surfactants. Gut fluid of T. caloptera contained enzymes capable of digesting the proteins of particles formed with maple water extracts. The other particles did not appear susceptible to these enzymes. There was no evidence that G. tigrinus was able to digest the proteins of any of the particles examined.     

Invader–invader interactions in relation to environmental heterogeneity leads to zonation of two invasive amphipods, Dikerogammarus villosus (Sowinsky) and Gammarus tigrinus Sexton: amphipod pilot species project (AMPIS) report 6

As biological invasions continue, interactions occur not only between invaders and natives, but increasingly new invaders come into contact with previous invaders. Whilst this can lead to species replacements, co-existence may occur, but we lack knowledge of processes driving such patterns. Since environmental heterogeneity can determine species richness and co-existence, the present study examines habitat use and its mediation of the predatory interaction between invasive aquatic amphipods, the Ponto-Caspian Dikerogammarus villosus and the N. American Gammarus tigrinus. In the Dutch Lake IJsselmeer, we found broad segregation of D. villosus and G. tigrinus by habitat type, the former predominating in the boulder zone and the latter in the soft sediment. However, the two species co-exist in the boulder zone, both on the short and longer terms. We used an experimental simulation of habitat heterogeneity and show that both species utilize crevices, different sized holes in a plastic grid, non-randomly. These amphipods appear to optimise the use of holes with respect to their ‘C-shape’ body size. When placed together, D. villosus adults preyed on G. tigrinus adults and juveniles, while G. tigrinus adults preyed on D. villosus juveniles. Juveniles were also predators and both species were cannibalistic. However, the impact on G. tigrinus of the superior intraguild predator, D. villosus, was significantly reduced where experimental grids were present as compared to absent. This mitigation of intraguild predation between the two species in complex habitats may explain the co-existence of these two invasive species.     

The survival and growth of Gammarus tigrinus Sexton (Crustacea: Amphipoda) in relation to salinity and temperature

Field collections from an inland saline lake indicated that a rapid increase in numbers of a population of Gammarus tigrinus was positively correlated with high summer temperatures and salinities. The laboratory experiments on growth are an attempt to determine whether high summer temperatures or optimum salinity were primarily responsible for the rapid increase in numbers. Furthermore, G. tigrinus was not found in either of two adjacent lakes; one non-saline, the other at a higher salinity. The laboratory experiments on survival are an attempt to determine whether the restricted distribution of G. tigrinus was related to salinity. The implications of the field and laboratory studies are discussed and compared with other data from Europe and North America.     

Intersexuality in the amphipod Gammarus duebeni results from incomplete feminisation by the vertically transmitted parasitic sex ratio distorter Nosema granulosis

We investigated the effects of parasitism and temperature on the production of intersexes in the amphipod Gammarus duebeni from a field population at Cumbrae, Scotland. There was significant temporal variation in intersex frequency which ranged from 0.5 to 5.2% in monthly field collections. Prevalence of Nosema granulosis, a feminising microsporidian parasite, also varied temporally and there was a significant correlation between parasite prevalence and intersex frequency in the field. Intersexes (16.3 ± 0.4 mg) were larger than true females (14.8 ± 0.1 mg) but produced fewer eggs thus demonstrating a cost of intersexuality. Intersexes were less likely to be paired than true females. In a breeding experiment, only females infected by N. granulosis produced intersex offspring. Temperature had no effect on intersex frequency. Intersexes therefore appear to be the result of incomplete feminisation by N. granulosis in this population. These results contrast with previous studies of G. duebeni from different populations which found that intersexes were the result of abnormal development under environmental sex determination. We suggest that intersexuality may be induced by both environmental and parasitic factors in populations of G. duebeni. This revised version was published online in August 2006 with corrections to the Cover Date.     

Interference competition between alien invasive gammaridean species

The relative abundances of gammaridean species in the river Rhine have profoundly changed since the invasion of Dikerogammarus villosus in 1994/1995. This study tested whether these changes in gammaridean dominance could have been determined by interspecific competition and unequal mortality, for example by intraguild predation (IGP). Single and two species tests have been carried out in aquariums provided with all substrata present in the main channel of the Rhine. Changes in substratum choice, increased swimming activity and increased mortality of a species were used as indicators of interspecific competition during interaction between gammaridean species. Interspecific competition and mortality between the most abundant invasive gammaridean species in the Rhine, viz. Gammarus tigrinus, Echinogammarus ischnus and Dikerogammarus villosus were tested. In single-species experiments, G. tigrinus and D. villosus showed similar preferences for a stony substratum, whereas E. ischnus mostly occupied the water column. The two-species aquarium experiments indicated direct interference competition for substratum and unequal mortality between G. tigrinus and D. villosus, with D. villosus being the stronger competitor. Competitive stress was influenced by population density, was size-dependent and varied between the different types of substratum due to substratum choice. G. tigrinus did not show any behaviour indicative of interference competition in the presence of E. ischnus, and neither did E. ischnus or D. villosus in the presence of any of the other gammarideans. Swimming in the water layer may already enable E. ischnus to minimise its encounters with the stone-dwelling D. villosus and G. tigrinus. To maximise the encounters between E. ischnus and D. villosus, a fish (Lepomis gibbosus) was added to occupy the water layer during the aquarium experiments. E. ischnus showed a higher mortality in the presence of both D. villosus and fish, probably due to increased stress, as shelter opportunities to escape the predators had been minimised. The study shows that interference competition between gammaridean species can explain the replacement of the North American invader G. tigrinus by D. villosus in the river Rhine. E. ischnus and D. villosus both Ponto-Caspian invaders did not show interference competition in our experiments and co-exist in the Rhine.     

The breeding behaviour of Gammarus duebeni

The breeding behaviour of a population of Gammarus duebeni is investigated. The breeding system includes a precopulatory guarding phase by a male of a female and the length of this phase varies markedly over the season. Large size is not of as great advantage in reproductive competition between males as in a stream population of G. pulex. In the spring of 1981 oligochete worms, Nais elinguis, were found in the brood pouches of some females; eggs were seldom present with the worms but rather amorphous debris. The occurrence of the worms and the debris in the females' pouches coincided with the pattern of abundance of the worms previously reported.     

How to be an invasive gammarid (Amphipoda: Gammaroidea)–comparison of life history traits

About six life history and two ecological traits of gammarid species occurring in Central European waters were compared in order to identify the characters of successful invader. The species were (1) natives: Gammarus fossarum, G. pulex, G. lacustris, G. varsoviensis, G. balcanicus, G. leopoliensis, G. roeselii, and (2) aliens: Gammarus tigrinus, Chaetogammarus ischnus (=Echinogammarus ischnus), Pontogammarus robustoides, Obesogammarus crassus, Dikerogammarus haemobaphes, D. villosus. Generally the alien species were characterised by a combination of large brood size, high partial fecundity, early maturation and by appearance of higher number of generations per year. Also, these species presented higher tolerance towards severe environmental conditions, i.e. elevated salinity and human degradation of the environment. The above features seem to facilitate the colonisation of new areas and competition with native species -- a phenomenon that has been currently observed in various parts of Europe.     

Die Veränderung populationsdynamischer Parameter bei Gammarus tigrinus Sexton (Crustacea: Amphipoda) als Ausdruck subletaler Effekte durch die Wechselwirkung von Temperatur und cadmiumkontaminiertem Futter

Changes in the Population Dynamics of Gammarus tigrinus SEXTON (Crustacea: Amphipoda) as Expression of Sublethal Effects by Reciprocal Interactions of Temperature and Cadmium Enriched Food The population dynamics of Gammarus tigrinus SEXTON were studied in a flow-through system under semi-natural conditions over a period of 9 months. Elevated temperatures and Cadmiumenriched food alter lifespan of generations, growth rate, number of individuals and time to reach maturity, even before elevated Cadmium concentrations can be detected in the animals. Reciprocal interactions of Cadmium intake via food and/or water can be modified by the experimental design, because it depends on the experimental animals, on the Cadmium concentration and on the temperature regime with constant or cyclic conditions.     

Interference competition among native and invader amphipods

Aquarium experiments were used to study indications of interference competition, such as substratum choice shifts, swimming activities and mortality of invasive and indigenous gammarids in each other's presence. The more recent invaders Gammarus tigrinus and Dikerogammarus villosus were more likely to prefer stone substratum, whereas the native Gammarus pulex and an earlier invader Gammarus roeseli were found more frequently in the water layer. Sand was the least likely substratum to be chosen by any of the species. G. pulex and G. roeseli did not alter their substratum preference in each other's presence. In the presence of D. villosus, G. pulex shifted towards smaller stones and increased its swimming activities, whereas D. villosus did not change its behaviour in the presence of G. pulex. These shifts may indicate interference competition, with D. villosus being the stronger competitor. The greatest shifts in substratum preference arose when one species had occupied a substratum before the other one was introduced, especially when D. villosus was already present before G. pulex was introduced, possibly indicating pre-emptive competition. Swimming activities of G. pulex increased in the presence of D. villosus, whereas D. villosus spent little time swimming. Mortality was comparable between the different experiments without any indication of predation. The effect of Intra Guild Predation (IGP) may not be reflected adequately by short-time experiments as moults occurred seldom during the experiments. Although no IGP was observed during our experiments, habitat shifts occurred, which may indicate that competitive interactions are apparent before IGP starts. Such shifts may serve to avoid intraguild competition.     

Strahlenbiologische Untersuchungen an Gammariden (Crustacea,Amphipoda)

Zusammenfassung 1. Der Einfluß einmaliger Röntgenbestrahlungen auf die Überlebenszeiten der euryhalinen AmphipodenGammarus duebeni Lilljeborg,Gammarus salinus Spooner undGammarus zaddachi Sexton wurde bei einer konstanten Temperatur von 15⁰ C und einem Salzgehalt von 10 untersucht.2. Bestrahlungen mit Dosen von 625 und 1250 R führen beiG. duebeni zum vorzeitigen Tod von nur einigen Versuchstieren, Bestrahlungen mit 2500 R und höheren Dosen zum vorzeitigen Tod aller Versuchstiere.3. BeiG. duebeni sind die strahlenempfindlicher als die , die Jungtiere strahlenempfindlicher als adulte Individuen.4.G. salinus undG. zaddachi sind strahlenempfindlicher alsG. duebeni.5. Die LD₅₀-Kurven lassen drei Bereiche erkennen. Bei mittleren Dosen ändert sich die Zeitspanne, in der 50% der Versuchstiere sterben, nur wenig oder gar nicht mit der Dosis. Bei hohen und niedrigen Dosen ist die Überlebenszeit dosisabhängig.6. Die Strahlenempfindlichkeit vonG. duebeni war im ersten Quartal des Jahres 1967, in dem die Temperaturen über den langfristigen Monatsmitteln lagen, geringer als bei Flohkrebsen, die zu einem späteren Zeitpunkt gefangen wurden oder aus Laboratoriumszuchten stammten.7. Die Häutungsvorgänge, deren Beeinflussung nur beiG. duebeni untersucht wurde, werden durch Bestrahlungen mit 1250 R oder geringeren Dosen nicht beeinflußt.8. Nach einer Bestrahlung mit 20 000 R sind 2 Tage nach der Bestrahlung nur wenige oder gar keine Häutungen möglich. Nach Bestrahlungen mit 10 000 und 5000 R erfolgen die Häutungen verspätet.9. Als Folge von Bestrahlungen mit letalen Dosen nimmt die Zahl der Häutungen um den 30. Tag nach der Bestrahlung ab.10. Die Häutungsvorgänge waren bei Tieren, die im ersten Quartal 1967 gefangen und bestrahlt wurden, weniger beeinflußbar als bei Tieren, die aus Laboratoriumszuchten stammten oder zu einem späteren Zeitpunkt gefangen wurden.11. Nach Bestrahlungen mit letalen Dosen treten zwei kritische Phasen auf. Ein Teil der mit mittleren und der größte Teil der mit hohen Dosen bestrahlten Tiere stirbt bereits in der ersten kritischen Phase. Die zweite kritische Phase wird von allen mit niedrigen Dosen bestrahlten Tieren erreicht. Tiere, die sich in der ersten kritischen Phase häuten, haben eine geringe Lebenserwartung.12. Die Überlebenszeit des einzelnen Individuums hängt außer von der Bestrahlungsdosis von dem Zeitpunkt ab, zu dem es sich innerhalb der Versuchszeit häutet.

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Radiobiological investigations on gammarids (Crustacea, Amphipoda)

Effects of single exposures of X-radiation on survival were studied inGammarus duebeni, G. salinus andG. zaddachi under constant conditions of temperature (15⁰ C) and salinity (10 ). Effects on moulting were investigated inG. duebeni. The amphipods were irradiated with doses between 625 and 20,000 R.G. salinus andG. zaddachi are equally radioresistant (LD_50/30 : 1,700 R) but less resistant thanG. duebeni (LD_50/30(males) : 3,900 R; LD_{50/30(females)} : 3,500 R). At high doses, half of the test individuals die within a few days; at medium doses survival time is dose-independent; at lower doses survival time again increases with decreasing doses up to dose-ranges (below 1,250 R forG. duebeni, 1,000 R forG. salinus andG. zaddachi), at which only a few individuals die before their natural death. Subadult gammarids are less resistant;G. duebeni of 4 to 7 mm body length have a LD_50/30 of 2,200 R. Death distribution after medium doses indicates that at least two mechanisms are involved in acute mortality. A first mortality maximum occurs shortly after irradiation; subsequently, mortality decreases for a few days, and is followed by another maximum. At lower doses no individual dies during the first critical period. Shortening of survival time at higher doses results from survival of fewer individuals to the second critical period. Few or no moults occur inG. duebeni irradiated with 20,000 R following the second day after irradiation and after the 30th day in those individuals irradiated with 2,500 to 10,000 R; moulting is delayed after exposure to 5,000 to 10,000 R. Individuals ofG. duebeni taken from the field during the mild winter 1966/67 were more resistant to radiation, and moulting was less affected, than in laboratory-reared amphipods, or in those collected in the field during other seasons. Besides on irradiation dose, survival time of an individual depends on the time of its moulting in the course of an experiment.