Interactions ofMonoporeia affinis (Lindström) (Amphipoda) with sedentary Chironomidae

Interannual population oscillations ofMonoporeia affinis and sedentary chironomids are negatively correlated in Lake Mälaren. Sedentary chironomid abundance regressed againstM. affinis density was highly significant at a lag response of one year (adj. R²=0.54, P=0.0001). The inverse correlation between Tanytarsini (Micropsectra sp. andTanytarsus sp.) andM. affinis densities supports a recruitment limitation hypothesis,viz. that the growth or survival of early instar chironomids is low at high amphipod densities. Microcosm studies showed density-dependent effects on the growth and survival of second instarCh. riparius larvae across an amphipod density gradient. No significant effects were found on growth or survival of third instars. These findings support the chironomid recruitment limitation hypothesis. Amphipod predation on early instar chironomid larvae is suggested as a population controlling mechanism.     

Intraspecific interactions between adults and juveniles of the subtidal limpet,Patelloida mufria

Summary Regular monitoring of populations of the subtidal limpet Patelloida mufria (Hedley), which were at naturally large densities (>1000/m²), showed that at the times when there were peaks in juvenile recruitment, the mortality of adults increased greatly. This produced a cyclical pattern in the density of adults, suggesting the juveniles play a partial regulatory role. Experimental manipulations were done to test the influence of overall density and the density of juveniles on the growth and survival of adults. These indicated that an increase in overall density reduced the rate of growth of individuals and increased the rate of adult mortality during the peaks in juvenile recruitment. Similarly, removing these juveniles as they recruited averted this decline in adult abundance. The proposed mechanism to explain these results is that the juveniles have a competitive advantage over adults as they are much smaller and therefore require less energy to survive. It is hypothesisied, therefore, that there will be a threshold of adult size below which the recruits will lose their competitive advantage. Furthermore, if their density became too great, the cover of crustose algae (which is the main food of this species) would be removed resulting in a rapid decline in the numbers of P. mufria. Such an occurrence could be the ultimate regulatory agent for this species.     

Complex interactions between native and invasive fish: the simultaneous effects of multiple negative interactions

We suggest that the ultimate outcome of interactions between native species and invasive species (extinction or coexistence) depends on the number of simultaneous negative interactions (competition and predation), which depends on relative body sizes of the species. Multiple simultaneous interactions may constrain the ability of native species to trade fitness components (i.e., reduced growth for reduced risk of predation) causing a spiral to extinction. We found evidence for five types of interactions between the adults and juveniles of introduced western mosquitofish (Gambusia affinis) and the juveniles of native least chub (Iotichthys phlegethontis). We added ten large (23–28 mm) and seven small (9–13 mm) young-of-the-year (YOY) least chub to replicate enclosures with zero, low, and high densities of mosquitofish in a desert spring ecosystem. Treatments with mosquitofish reduced the average survival of least chub by one-third. No small YOY least chub survived in enclosures with high mosquitofish densities. We also performed two laboratory experiments to determine mortality to predation, aggressiveness, and habitat selection of least chub in the presence of mosquitofish. Mean mortality of least chub due to predation by large mosquitofish was 69.7% over a 3-h trial. Least chub were less aggressive, selected protected habitats (Potamogeton spp.), and were more stationary in the presence of mosquitofish where the dominance hierarchy was large mosquitofish>>large least chubsmall mosquitofish>>small least chub. Least chub juveniles appear to be figuratively caught in a vice. Rapid growth to a size refuge could reduce the risk of predation, but the simultaneous effects of competition decreased least chub growth and prolonged the period when juveniles were vulnerable to mosquitofish predation.     

Evidence for predatory flatworms as organizers of zooplankton and mosquito community structure in rice fields

I present evidence from two field studies and one laboratory experiment that the predatory flatworm, tentatively indentified as Mesostoma nr. lingua, can be important in organizing invertebrate communities in rice fields. In the first field study, I assessed the importance of various predators including two fishes, in explaining among-plot variation in mosquito abundance during a five week interval. Mosquitofish (Gambusia affinis) abundance was not significantly associated with densities of either mosquito, Culex tarsalis or Anopheles freeborni, except for the final week. Associations between green sunfish (Lepomis cyanellus) juveniles and mosquitoes were demonstrated for only A. freeborni — a positive association. Densities of Mesostoma were negatively associated with C. tarsalis and A. freeborni densities. C. tarsalis, protected from flatworms in small-mesh cages, had a much greater survival rate than those that were exposed in large-mesh cages. C. tarsalis survival, when left unprotected in large-mesh cages, was negatively correlated with field densities of flatworms and with the number of flatworms found inside the cages. Survival was also positively correlated with field densities of mosquitoes.In the second field study, Mesostoma densities were negatively correlated with C. tarsalis, A. freeborni, Cladocera and Ostracoda in rice field enclosures early in the season. Cyclopoid copepod densities were not significantly associated with this predator.In a laboratory experiment, predation by Mesostoma on C. tarsalis larvae was reduced when presented with an alternative prey, Moina micrura (a cladoceran).These results suggest that flatworms, via predation, can have large effects on the structure of mosquito and zooplankton communities in rice fields.     

Multiple density dependence in two sub-populations of the amphipod Monoporeia affinis: a potential for alternative equilibria

Possible mechanisms for differences in population densities and dynamics were investigated in the amphipod Monoporeia affinis at two deep sites in the northern Bothnian Sea. The two sites were sampled yearly for 10 years. Average sizes, growth and mortality of the different age-classes were estimated from the cohort structure of the two populations. Laboratory experiments also investigated the ability of the common predatory isopod Saduria entomon to cause densitydependent (DD) mortality of the prey M. affinis. At site A, 43 m depth, the average density of M. affinis was twice as high as at site B, 81 m depth. The fluctuations in density were asynchronous between the two sites. Recruitment and subadult sizes of Monoporeia affinis were density dependent at both sites. The main functional difference between the two populations seemed to be the DD mortality of the 1 + cohort that occurred only at the low-density site B. A corresponding DD mortality was found in the predation experiments at densities of 1 + m. affinis corresponding to those found at site B. The potential importance of the predator was also indicated by a significant negative correlation between the biomass of S. entomon and the rate of change in M. affinis density in the field. The similarities in the abiotic factors between the two sites suggested that differences in carrying capacity should be small. The results could be explained by the predation regulation hypothesis for the low-density population at site B, while at site A M. affinis seemed to be regulated by intra-specific competition and limited by predation. It is suggested that in this simple predator-prey system there is potential for the existence of alternative equilibria.     

Distribution and abundance of the Louisiana red swamp crayfish Procambarus clarkii Girard at Lake Naivasha,Kenya between 1987 and 1999

The abundance and habitat distribution of Procambarus clarkii, an exotic herbivore at Lake Naivasha, Kenya, was studied between 1987 and 1999. Its growth and morphometrics were similar to other locations in the world where the species have been studied. P. clarkii was found primarily amongst the floating, exotic vegetation where it achieved densities of over 500 m^–2 by the end of the study. It is suggested that these densities became possible because the habitat offers protection against predation and an adequate food supply of bacterial-enriched detritus exists in the absence of green plant material. Both adults and juveniles forage on the lake bed at night.     

Distribution and life cycle of the North American Spionid polychaeteMarenzelleria viridis (Verrill, 1873) in the Ems estuary

In 1983 the first specimens of the North American spionid polychaeteMarenzelleria viridis were found along the European mainland shore in the Ems estuary. Since then, this polychaete has spread over several estuaries around the North Sea and the Baltic. In the inner part of the Ems estuary juveniles were predominantly present in muddy sediments high in the intertidal zone; in more sandy sediments at higher salinities juveniles and adults co-occured. Detailed information was obtained at a muddy and at a sandy station. Gametes were present in the coelomic fluid from November through March. In May new recruits were found in the sediment samples, reaching densities of over 10⁵ M⁻² at the muddy station, andc. 2000 m⁻² at the sandy station. During summer, densities decreased at the muddy station, coinciding with a density increase at the sandy station, suggesting migration of juveniles from a nursery to the adult habitat. Cage experiments showed that the decrease of juveniles at the muddy station could be attributed to migration and not to mortality due to predation. At the sandy habitat palps and anterior parts ofM. viridis made up 4–11% of the stomach content of juvenile plaice (Pleuronectes platessa). In juvenile flounder (Platichthys flesus) only in Aprilc. 10% of the stomach content consisted ofM. viridis. During 1983–1990 increasing densities ofM. viridis at the sandy habitat coincided with a reduced abundance ofNereis diversicolor, however, this inverse relationship was not found to be statistically significant. Density fluctuations ofM. viridis andCorophium volutator showed a significant positive relationship, the cause of which is not yet understood.     

Direct and indirect effects of predation by Saduria entomon (Isopoda) on the size-structure of Monoporeia affinis (Amphipoda)

We performed a 6-month laboratory experiment to investigate the direct and indirect effects of predation by the benthic invertebrate predator Saduria entomon on the growth and survival of Monopreia affinis prey individuals in different age-cohorts at low and high prey densities. The experimental results were compared with changes of growth and abundance in corresponding age-cohorts of M. affinis at sites with different S. entomon and M. affinis densities in the deep sublittoral zone of the Bothnian Sea during the same year. In the experiment, the presence of S. entomon reduced growth rate of M. affinis in the 1-year and 2-year age-cohorts at low amphipod density. Increased refuge use by M. affinis, expressed as a decrease in swimming activity in the presence of S. entomon, is suggested to have reduced feeding rate and therefore growth of the amphipods. The recruitment of M. affinis offspring was reduced in presence of S. entomon. In the field, the growth rate of amphipods in the 1-year cohort increased with increasing S. entomon density at low amphipod density. We found no corresponding increase of M. affinis growth in the 2-year cohort. The positive effect on 1-year amphipod growth indicated that predation reduced intra-cohort competition of M. affinis and increased growth of surviving prey at high predator density. In both the experiment and the field data we found indications of size-selective predation on smaller M. affinis specimens. This was because of the changed ratio between number of individuals in the juvenile age-cohorts and lower recruitment of amphipod offspring connected to S. entomon density. The experimental results and field data suggest that predation by S. entomon may have both direct and indirect effects on the size-structure of M. affinis populations. Received: 22 June 1998 / Accepted: 15 March 1999     

Physiological consequences of changes in life form of the Mexican epiphyte Tillandsia deppeana (Bromeliaceae)

Summary The heterophyllous epiphyte Tillandsia deppeana exhibits an atmospheric habit as a juvenile and a tank form as an adult. Both juveniles and adults utilize C₃ photosynthesis. This is the first report of an atmospheric form of Tillandsia which does not exhibit CAM. Photosynthetic saturation occurred at approximately 10% of full sunlight in both forms, but the adults exhibited greater rates of photosynthesis at all levels of irradiance. The adults also had a higher and broader photosynthetic temperature optimum than did the juveniles. The adults transpired at greater rates than the juveniles; however, the water use efficiencies of both forms were similar and were high for C₃ plants. In both forms the photosynthetic rate decreased in response to a decrease in humidity. After 8 days without water the juveniles were able to fix CO₂ throughout the day. The adults, however, exhibited a net loss of CO₂ on the second day without water and thereafter. These results indicate that the water-conservative atmospheric juvenile of T. deppeana is well adapted to establishment in the epiphytic habitat.     

Ex situ evaluation of impacts of invasive mosquitofish on the imperiled Barrens topminnow

Central to the protection of native species is an understanding of impacts of actual or potential invasive species and also the mechanisms through which those impacts are mediated. The introduction and spread of western mosquitofish, Gambusia affinis, into spring systems of the Barrens Plateau region of middle Tennessee is a concern for native species such as the Barrens topminnow, Fundulus julisia. We investigated whether mosquitofish might act as predators on early life stages of topminnows as well as affect the physiological well being of adults through aggressive interactions. A short-term, 24-h laboratory study with mosquitofish and topminnows demonstrated the vulnerability of young topminnow life stages to large mosquitofish predation and aggression. Survival of topminnow young, <16 mm total length (TL), was 0% and was attributed to predation by mosquitofish. Survival of juveniles, 20–30 mm TL, was 25%; juveniles mostly succumbed (post 24-h) to injuries inflicted by large mosquitofish. Adult topminnow survival was 100% but adults faced injury risk, primarily during the initial stages of their interaction with large mosquitofish. A long-term, 60-day laboratory study with syntopic and allotopic populations of adult topminnows and mosquitofish failed to detect any negative impacts on topminnows due to coexistence. Survival, growth, and fecundity of adult topminnows syntopic with mosquitofish were not different from the allotopic population, although injury risk in the form of fin damage was greater syntopically. Thus, predation and aggression towards young topminnows may be the primary mechanisms by which western mosquitofish jeopardize the persistence of native Barrens topminnows in the wild. Our results reemphasize the danger to native aquatic biodiversity of unregulated introductions of Gambusia species.     

Competitive interactions between two successful molluscan invaders of freshwaters: an experimental study

The New Zealand mud snail Potamopyrgus antipodarum (Hydrobiidae) and the pulmonate Physella acuta (Physidae) have invaded freshwaters in many parts of the world and become established. They co-exist in many streams, lakes and ponds in New Zealand, often at high densities. In the present study the effects of intraspecific- and interspecific interactions between the two species on growth and reproductive output were examined in laboratory mesocosms. In 30-day experiments, growth of Potamopyrgus antipodarum was lower in high density treatments than controls providing evidence for competition at higher densities of both snail species. No competitive effect was obtained for Physella acuta when controls were compared with high-density treatments, but growth was reduced at high densities of conspecifics. Numbers of juveniles released by Potamopyrgus antipodarum in 40 day trials declined at high snail densities and were lowest at high densities of conspecifics. Egg production by Physella acuta was also reduced at high snail densities. However, when the two species were kept together at equal densities (total snail density twice that of controls), egg production by Physella acuta was significantly higher than in all other treatments, suggesting facilitation by the congenor. Lastly, in a 10-day experiment, Physella acuta grew faster in water conditioned by Potamopyrgus antipodarum than in Physa-conditioned water, whereas Potamopyrgus antipodarum showed no growth response to Physella-conditioned water. Overall, our results indicate that growth and reproductive output of both snail species are influenced more by the density of conspecifics than the presence and density of the other species. The successful co-existence of the two species in New Zealand freshwaters therefore may be a reflection, at least in part, of few competitive interactions between them.     

Intraspecific competition of Glyptotendipes paripes (Diptera: Chironomidae) larvae under laboratory conditions

Glyptotendipes paripes larvae were reared in wells of tissue culture plates, in groups of 2, 4, 8, 16, and 32 (representing densities of about 1,300, 2,600, 5,200, 10,400, and 20,800 larvae per m², respectively). Larval groups were supplied with one of two concentrations (low or high) of food and larvae were individually observed to evaluate the effects of density on mortality, growth, development, behavior, and adult body size. Increased larval densities resulted in higher mortality, as well as slower larval growth and development. The distribution of developmental time became flatter at higher density, with a wider range of values, or even became bimodal. This was a consequence of the most rapidly developing individuals at higher densities emerging as adults sooner than the fastest developing individuals at lower densities, although overall mean developmental time was longer at higher densities. At higher densities, growth and development of smaller larvae were slowed, based on the relative difference in body length between competitors. When larger competitors emerged as adults or died, the growth of smaller larvae may have accelerated, resulting in increased variability of developmental times. The effect of larval density on adult body size was complex, with the largest body size found at the lowest density and a second peak of adult size at high-middle densities, with smaller adult body sizes found at low-middle, and high densities. Similarly, as with developmental time, the range of body size increased with increasing density. Examined food concentrations had no effect on larval mortality, but significantly affected developmental time, growth rate, and adult body size. At higher densities, larvae spent more time gathering food and were engaged in aggressive or antagonistic behaviors.     

Role of plant phenology in mediating interactions between two biological control agents for spotted knapweed

The role of spotted knapweed phenology on the attack rate of two seed-head insects Urophora affinis and Larinus minutus was assessed in a series of field studies at four study sites in south-eastern British Columbia, Canada. Slow or later developing knapweed plants had more seed heads that contained only single or multiple U. affinis whereas early or faster developing plants had more seed heads containing L. minutus alone or in combination with U. affinis. L. minutus did not distinguish between seed heads with or without U. affinis larvae when laying eggs. However, seed heads with multiple U. affinis present, produced fewer L. minutus adults than expected. The probability of single or multiple U. affinis galls being present increased with seed-head diameter but was not affected by seed-head height. Attack by L. minutus increased with seed-head diameters >5 mm and was lower at plant heights above 50 cm. These results demonstrate two mechanisms that enable U. affinis to successfully coexist with L. minutus: differences between the species in their response to the developmental phenology of knapweed heads, and increased survivorship of U. affinis in heads with multiple U. affinis galls through niche interference competition. These mechanisms provide a possible explanation for the persistence of U. affinis populations on spotted knapweed, in spite of high levels of within seed-head mortality that have been observed with increasing L. minutus populations.     

Trophic interactions involving mysid shrimps (Mysidacea) in the near-bottom habitat in the Baltic Sea

In a six month mesocosm tank experiment, hypotheses were tested concerning the role of benthopelagic mysid shrimps (Mysidacea) in the near-bottom food web of the Bothnian Sea, in the northern Baltic Sea. The first hypothesis tested was that the mysids interact, through predation, with benthic deposit-feeding Monoporeia affinis amphipods. A second hypothesis tested was that the sediment type is important for the overwintering success of the mysids. Changes in abundance and mass were recorded for M. affinis and mysids when separate and when coexisting, in two sediment types differing in organic content (food level); soft muddy clay (rich) and fine sand (poor). Despite the fact that newborn M. affinis offspring, a plausible target for predation by mysids, were present in substantial numbers in the tanks, no consistent evidence for any interaction between these taxa was found. The biomass of mysids was slightly higher in the muddy clay than in the sand tanks, and the mechanism behind this substrate effect is discussed. A third hypothesis, that the mysids interact with near-bottom zooplankton, was investigated. The tanks were continually supplied with in situ near-bottom sea-water containing a semi-natural assemblage of near-bottom plankton. As a result of mysid predation, tanks with mysids had lower abundance and biomass of cyclopoid copepods than tanks without mysids. Thus, the major interaction found was predation on near-bottom zooplankton by mysids and it is suggested that this interaction could potentially be an important food link, especially during periods with low food availability in the pelagic system.     

Interaction of the Microbivorous Nematode Teratorhabditis dentifera and the Nematode-Pathogenic Fungus Hirsutella rhossiliensis

The fungus Hirsutella rhossiliensis is an obligate pathogen with a broad host range among nematodes. Microbivorous nematodes are abundant around plant roots and may serve as hosts for the fungus. Our objective was to determine the influence of the bacterial-feeding nematode Teratorhabditis dentifera on the abundance of H. rhossiliensis. Experiments were conducted in a growth chamber with pots containing pasteurized soil, the fungus, and potato plants. The abundance of infectious conidia was compared in pots with and without T. dentifera after 50 or 70 days. The nematode reached high densities (10-40/cm³ soil) but had no effect on the abundance of conidia. Many individuals were dauer juveniles, a stage that acquired conidia but did not become infected. To test whether this life stage could deplete the pool of conidia in soil, different proportions of dauer juveniles with (resistant) and without (susceptible) a sheath were added to H. rhossiliensis-infested soil. The number of conidia in the soil decreased with an increasing proportion of resistant nematodes. Different stages of T. dentifera appear to have opposing effects on H. rhossiliensis; while adults and regular juveniles acquire conidia, become infected, and produce new infectious conidia, dauer juveniles can deplete the supply of conidia.     

Utilisation of a tropical bay as a nursery area by sharks of the families Carcharhinidae and Sphyrnidae

Synopsis At least eight species of sharks of the families Carcharhinidae and Sphyrnidae use Cleveland Bay in northern Australia as a communal nursery area.Carcharhinus dussumieri, C. fitzroyensis, C. limbatus andC. tilstoni use the bay as a seasonal primary nursery, with juveniles occurring in it for only a few months each year immediately after birth. Alternatively,Carcharhinus sorrah, Rhizoprionodon acutus andR. taylori use the bay as a year-round primary and secondary nursery, with juveniles remaining in it up to the size at maturity. AdultR. taylori also persist in the bay, a behavioural pattern possibly explained by their small maximum size. While present immediately after birth the type of utilisation pattern displayed bySphyrna lewini could not be clarified in this study. Although diets of these species in the bay are similar, there is probably little direct competition for food due to the highly productive habitats in the bay supporting an abundance of food resources. The highest numbers of juveniles occur when prey species are the most abundant, and when temporal separation of some seasonally-occurring species of sharks in effect.     

Seasonal changes in zooplankton abundance in the lower Rhine during 1987–1991

Zooplankton in the River Rhine was surveyed for five years at the Dutch sampling stations, Lobith (German/Dutch border) and Maassluis (at the point of discharge of the river into the North Sea). The zooplankton abundance showed an apparent seasonal pattern at both stations, characterized by low densities during the winter period, and higher densities during the summer period, with a spring peak. Zooplankton was dominated by rotifers at both stations, although during the winter periods the contribution of copepods was considerable. The rotifers were dominated byBrachionus angularis, B. calyciflorus, Keratella cochlearis andK. quadrata; the copepods by cyclopoid nauplii; the cladocerans by small-sized species mainly belonging toBosmina. At Maassluis the relative contribution of copepods was higher than at Lobith. Furthermore, the zooplankton at Maassluis included the speciesEurytemora affinis, characteristic for estuarine conditions. In spring, the rotifer density and water temperature and rotifer density and chlorophylla concentration were positively correlated. Furthermore, both rotifer density and chlorophylla were inversely correlated with discharge. The possible role of environmental factors (water temperature, chlorophyll content, discharge and biotic factors) controlling the river zooplankton dynamics is indicated.     

The response of three chydorid species to temperature, pH and food

The responses of three chydorid species, Chydorus sphaericus(O.F. Müller), Alona affinis (Leydig) and Alonopsis elongata (Sars) to temperature, pH and food type were examined. Egg development time of all species decreased with increasing temperature, although the degree of change was different for each species. C. sphaericus had the fastest development time at all temperatures, and A. elongata the slowest. pH also affected the egg development time of each species differently. A. elongata failed to reproduce at low and medium pH, the egg development time of C. sphaericus was fastest at high pH while that of A. affinis was fastest at low pH. Food type was found to have significant effects on the population growth of individual species. C. sphaericus populations grew equally well in all three food types provided, and grew more than the other two species when fed on an algae culture, and filtered pond water. A. affinisand A. elongata populations grew best when fed on a detritus food source. The responses of each species to the different variables tested are discussed in relation to field observations of their distribution and abundance.     

The influence of riparian vegetation on the functional organization of four Hong Kong stream communities

Pontoporeia affinis Lindström predominated profundal macroinvertebrate communities in eutrophic Lake Mälaren. Amphipod densities, at both stations studied, averaged > 50% of overall total macroinvertebrate densities. The life history of P. affinis was two years, with two cohorts distinguished, both exhibiting similar patterns in annual growth. During the sixteen years of this study, amphipod growth seemed to exhibit long-term cyclic fluctuations, with periods of 9–10 yrs. at Görväln and 5–6 yrs. at Grönsöfjärden. Amphipod growth was significantly (negatively) correlated with conspecific density and biomass. The mechanism(s) governing Pontoporeia population oscillations is (are) not clear. It is postulated that changes in food availability, as the result of overgrazing, may adversely affect Pontoporeia recruitment and subsequently be observed in the form of long-term population oscillations as noted in Lake Mälaren.     

Microsporidiosis in a Daphnia pulex population

The impact of parasitism by Thelohania sp., an intracellular microsporidian, on Daphnia pulex inhabiting a vernal pond was studied for three years. Three issues were considered: the distribution of parasites among hosts, the effect of parasites on individuals, and the impact of parasitism on population growth. Each year, Thelohania infected members of at least half of the generations of Daphnia produced in the pond. When the frequency of infection was low, parasites were found only in large adults. As infection frequency increased, parasitism spread downward through host size classes. However, parasites were rarely found in juveniles. Parasitism reduced clutch size drastically, increased mortality to a variable extent, had little impact on egg size or on per instar growth and none on molt frequency. Interaction with other stresses, such as food limitation, exacerbated some of these effects. Parasitism reduced instantaneous birth rate much more than it elevated instantaneous death rate. Population growth was reduced significantly but it is unlikely that Thelohania alone regulates the growth of this Daphnia population.