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.     

Effects of large Saduria entomon (Isopoda) on spatial distribution of their small S. entomon and Monoporeia affinis (Amphipoda) prey

We performed laboratory experiments to investigate the effects of predator avoidance and numerical effects of predation on spatial distribution of small Saduria entomon (Isopoda) and Monoporeia affinis (Amphipoda), with large S. entomon as predators. The horizontal distribution and mortality of the prey species, separately and together, were studied in aquaria with a spatial horizontal refuge. We also estimated effects of refuge on mortality of small S. entomon and M. affinis by experiments without the refuge net. In addition, we investigated whether predation risk from large S. entomon influenced the swimming activity of M. affinis, to clarify the mechanisms behind the spatial distribution. Both small S. entomon and M. affinis avoided large S. entomon. The avoidance behaviour of M. fffinis contributed about 10 times more to the high proportion in the refuge than numerical effects of predation. Due to the low mortality of small S. entomon the avoidance behaviour of this species was even more important for the spatial distribution. The combined effect of avoidance behaviour and predation in both species was aggregation, producting a positive correlation between the species in density. M. affinis showed two types of avoidance behaviour. In the activity experiments they reduced activity by 36% and buried themselves in the sediment. In the refuge experiments we also observed avoidance behaviour with the emigration rate from the predator compartment being twice the immigration rate. The refuge did not lower predation mortality in M. affinis, probably due to the small scale of the experimental units in relation to the mobility of the species. Predation mortality in small S. entomon was higher in absence of a refuge and especially high in absence of M. affinis.     

Predation by the isopod Saduria entomon on the amphipods Monoporeia affinis and Pontoporeia femorata: experiments on prey vulnerability

Summary Predation by Saduria entomon on the depositfeeding amphipods Monoporeia (=Pontoporeia) affinis and Pontoporeia femorata was studied in laboratory experiments. Prey vulnerabilities were compared in singleprey treatments and in mixed-prey treatments, where the proportions of the two species were varied at one total density. In a pilot experiment, P. femorata was the more vulnerable prey, both in single-prey and mixed-prey treatments. In later experiments, the amphipod species were equally vulnerable in single-prey treatments, while in mixed-prey treatments Saduria preyed preferentially on M. affinis, even when this prey was rare, i.e. it did not switch. We suggest that the different result in the later experiments may have been due to a change in the search mode of the isopods.     

Paternal effects on juvenile growth and survival in spring peepers (Hyla crucifer)

Summary In 1986 and 1987 we crossed severalHyla crucifer females to males that obtained mates in nature and to large and small body sized males that did not do so. Offspring resulting from these crosses were raised as pairs of juveniles in 1986, or in isolation in 1987 to assess sire effects on juvenile growth and survival in a competitive and a non-competitive context. When raised as part of a pair of juveniles, individuals sired by natural mates had lower growth rates than their fullsibs raised alone. The reduction in growth was of a similar magnitude whether the cohabiting juvenile was sired by a large or small bachelor. When raised together, individuals sired by natural mates had higher growth rates (indicative of competitive superiority) than their halfsibs sired by large bachelor males, but competed as equals with the progeny of small bachelor males. Progeny of large bachelors competed with progeny of small bachelors as equals. When raised individually in 1987, juveniles sired by natural mates had lower growth rates than their halfsibs sired by bachelor males. Paternal size or mating status had marginally non-significant effects on juvenile survival in 1986 and no effect in 1987. The larger individual in a pair was more likely to survive in 1986. In neither year was juvenile growth rate correlated with prior growth rate as a larva. These experiments suggest that the mating system can affect the distribution of juvenile growth rates (and possible survival rates), and thereby have important fitness consequences long after pair formation. These effects are, however, inconsistent either among sets of individual males or among juvenile environments.     

The regeneration niche of the grey mangrove (Avicennia marina): effects of salinity,light and sediment factors on establishment,growth and survival in the field

Field observations of seedlings and saplings of Avicennia marina showed patterns that correlated with salinity, light and sediment. Models that account for these observations were subsequently tested in a series of field experiments. Establishment varied within an estuary under controlled conditions but was not related to salinity or sediment type. Seedling survival was uniform over 3 years regardless of position in estuary and sediment type. Seedling densities and survival under canopies or in canopy gaps were not significantly different. However, seedling growth and density of saplings were greater in canopy gaps. Experiments involving manipulations of canopies showed no differences in seedling survival under canopies or in light gaps, but addition of slow-release fertilizer enhanced growth and survival in canopy gaps and under canopies. Long-term comparison of areas denuded of a canopy and with sediment disturbance showed enhanced establishment and survival when compared with areas with canopy gaps but with undisturbed sediments. Overall there appears to be no restriction to establishment of propagules within mangrove stands other than the supply of propagules and tidal or wave action. In contrast, recruitment to the sapling stage appears to be restricted by light and sediment resources. We suggest that propagules need to establish in a regeneration niche for seedling recruitment to the sapling stage. This differs from the view that seedlings in the under-storey are analogous to a seed pool in the soil.     

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.     

Why spacing behavior does not stabilize density in cyclic populations of microtine rodents

Summary There are several published hypotheses that consider spacing behavior to be a significant factor causing the multiannual density fluctuations characteristic of some microtine rodent populations. Recent modeling efforts have concluded, however, that spacing behavior should have a stabilizing rather than a destabilizing effect on population dynamics. Why doesn't spacing behavior stabilize these cyclic populations? We argue that while spacing behavior does have a stabilizing influence on population dynamics by limiting the number of breeding individuals, reproduction continues and population size is not limited in an asymptotic manner. Rather, microtine social organization produces demographic changes within a population that allow density cycles to occur under certain conditions. Using a simulation model, we demonstrate that in a strongly seasonal environment populations with low density dependence in reproduction will cycle whereas populations with high density dependence in reproduction will have relatively stable densities. Given such complicating factors as the annual species nature of microtine rodents, occasionally intense predation, and the tendency for territoriality to break down during the non-breeding season, individuals with low density dependence in reproduction will always be able to invade and eventually dominate populations with high density dependence in reproduction, regardless of the resulting destabilization of population dynamics.     

Density and intercohort priority effects on larval <Emphasis Type="Italic">Salamandra salamandra</Emphasis> in temporary pools

Priority effects, i.e., effects of an early cohort on the performance of a later cohort, are generally studied between, and not within, species. The paucity of intraspecific assessments does not reflect a lack of ecological importance, but the technical problem associated with differentiating between conspecific cohorts. Here, we examine priority and density-dependent effects on larval Salamandra salamandra infraimmaculata. Larvae deposited by their mother early in the season have increased risk of desiccation, as rains at the beginning of the season are less frequent and unpredictable. However, breeding later may incur a high cost through conspecific priority effects, including cannibalism and competition. In an outdoor artificial pool experiment, we established densities of 0, 1, 2, 4 or 6 newly born larvae per pool (∼30 l), and 40 days later, added a second cohort of three newly born larvae to each pool. We differentiated between cohorts using natural individual-specific markings. For the early cohort, increasing density decreased survival and size at metamorphosis, and increased time to metamorphosis. For the late cohort, survival was 100% in pools without early-cohort larvae, but ranged between 13 and 33% in the presence of early-cohort larvae. Time to metamorphosis was significantly longer in the presence of low vs high densities of early-cohort larvae. Results suggest that early-cohort larvae are mainly subjected to exploitative competition and cannibalism mediated by food limitation, and that late-cohort larvae are subjected to cannibalism and interference due to size asymmetry between cohorts. The strong priority effects suggest that Salamandra females could increase their fitness by adjusting the number of larvae they deposit in specific pools to avoid cannibalism and intraspecific competition.     

Diet overlaps and feeding relationships between the year classes in the yellow perch (Perca flavescens)

Synopsis A comparative study of the diets of five year classes of perch in Lake Opinicon showed Year class 0 to be mainly Cladocera feeders, the Year class 11 to take a diversified range of insect larvae, whereas from Year class V onwards Anisoptera nymphs, decapods, and fish dominated the diet. Sizes of the dominant food items changed strikingly with growth, prey weights of 0.1, 0.5–1.0, and 100–250 mg predominating in these three age groupings. There were marked month to month changes in types and proportions of different prey consumed.Overall and month to month feeding overlaps between consecutive year classes were considerable, most values ranging from 0.68–1.0 by the Levins formula. However, diet overlap dropped sharply with increasing age gap.Month to month dietary changes and year class differences were interpreted relative to benthic invertebrate abundance cycles. Months of increased diet overlap were characteristically those when favored prey types peaked, thereby simultaneously attracting several year classes to them. Hence increased diet. overlap is not necessarily an indicator of increased intraspecific competition.Habitat differences between the year classes of perch in Lake Opinicon were partial and concerned mainly the smaller, and larger, fish relative to those in the middle size range.     

An experimental study of population regulation in the salamander,Notophthalmus viridescens dorsalis (Urodela: Salamandridae)

Summary The roles of density-dependent larval survival and cannibalism of larvae as potential mechanisms of population regulation in the newt (Notophthalmus viridescens dorsalis) were evaluated in laboratory and field experiments. In laboratory containers, adults cannibalized larvae and large larvae cannibalized smaller larvae. In artificial ponds, larval survival did not depend on initial larval density. No cannibalism could be demonstrated in the complex environment, although the experiment was powerful enough to detect an ecologically relevant difference in survival. Adult growth was negatively correlated with the final biomass of larval newts, suggesting that the two life stages competed for resources. Larval growth rates were negatively correlated with final larval density, suggesting that larvae competed with each other. The proportion of larvae that became sexually mature at age 7 months (paedomorphs and adults that skipped the eft stage) varied inversely with larval density. Therefore, the potential regulatory mechanisms identified in this study are competition within and between life stages.     

Comment arising from a paper by Wittmer et al.: hypothesis testing for top-down and bottom-up effects in woodland caribou population dynamics

Conservation strategies for populations of woodland caribou Rangifer tarandus caribou frequently emphasize the importance of predator–prey relationships and the availability of lichen-rich late seral forests, yet the importance of summer diet and forage availability to woodland caribou survival is poorly understood. In a recent article, Wittmer et al. (Can J Zool 83:407–418, 2005b) concluded that woodland caribou in British Columbia were declining as a consequence of increased predation that was facilitated by habitat alteration. Their conclusion is consistent with the findings of other authors who have suggested that predation is the most important proximal factor limiting woodland caribou populations (Bergerud and Elliot in Can J Zool 64:1515–1529, 1986; Edmonds in Can J Zool 66:817–826, 1988; Rettie and Messier in Can J Zool 76:251–259, 1998; Hayes et al. in Wildl Monogr 152:1–35, 2003). Wittmer et al. (Can J Zool 83:407–418, 2005b) presented three alternative, contrasting hypotheses for caribou decline that differed in terms of predicted differences in instantaneous rates of increase, pregnancy rates, causes of mortality, and seasonal vulnerability to mortality (Table 1, p 258). These authors rejected the hypotheses that food or an interaction between food and predation was responsible for observed declines in caribou populations; however, the use of pregnancy rate, mortality season and cause of mortality to contrast the alternative hypotheses is problematic. We argue here that the data employed in their study were insufficient to properly evaluate a predation-sensitive foraging hypothesis for caribou decline. Empirical data on seasonal forage availability and quality and plane of nutrition of caribou would be required to test the competing hypotheses. We suggest that methodological limitations in studies of woodland caribou population dynamics prohibit proper evaluation of the mechanism of caribou population declines and fail to elucidate potential interactions between top-down and bottom-up effects on populations. An erratum to this article can be found at     

Population biology of the broad-horned flour beetle,Gnathocerus cornutus (F.) II. Crowding effects of larvae on their survival and development

Crowding effects of larvae on survival and development were examined for the broad-horned flour beetle, Gnathocerus cornutus (F.). The larvae matured about 3 weeks after hatching regardless of their densities, but pupation was severely hindered by crowding. There existed an upper limit for the number of the pupae produced and its mechanism was studied by a statistical analysis of the distribution patterns of pupal cells and the experiment in which glass tubes were artificially supplied in addition as pupation site. These studies show that G. cornutus larvae have a habit to construct cells for pupation and this habit leads to a contest competition for pupation site at high densities. The significance of the contest competition for population regulation was discussed comparing the results on Tribolium confusumJacqueline duVal .     

Patterns of Solidago altissima ramet growth and mortality: the role of below-ground ramet connections

Summary For the rhizomatous perennial, Solidago altissima, I identified clonal fragments in the field, mapped ramet spatial locations, and documented patterns of ramet recruitment, growth, and mortality. Parent ramet size influenced the size and number of daughter ramets produced, and small ramets had lower survivorship and fecundity than large ramets. Similarly, small rhizomes tended to develop into small ramets, and ramets that survived to produce daughter ramets had longer parent-daughter rhizome connections than ramets that did not survive. In addition, most ramets that died during the growing season were connected to (genetically identical) ramets that persisted. There were large size inequalities among rhizomes, ramets, and clonal fragments. Inequalities in the size of ramets increased during the early part of the growing season, then decreased at the end of the season; similar patterns were observed for the growth of clonal fragments. In both instances, the decrease in size inequality could be attributed to the mortality of small individuals (ramets or clonal fragments). I found little evidence that ramet size hierarchies were structured by intraspecific competition. For example, path analyses and randomization tests indicated that size variation among S. altissima ramets was influenced little by the size of their near neighbors (but was influenced by parent size and rhizome size). In addition, within-season variation for the relative size and growth rate of individual ramets led to poor correlations between early and final ramet size; this result suggests that there was no stable hierarchy of dominant and suppressed ramets. I discuss implications of my results for contrasting interpretations of clonal plant population dynamics.     

Visual pigment,dark adaptation and rhodopsin renewal in the eye of Pontoporeia affinis (Crustacea,Amphipoda)

The benthic amphipod Pontoporeia affinis lives in the Baltic sea and in northern European lakes in an environment where very little light is available for vision. The eyes, consisting of 40–50 ommatidia, are correspondingly modified. Microspectrophotometric recordings on isolated eyes show the presence of at least two kinds of screening pigments in the ommatidia with maxima at 540–580 nm and 460–500 nm. Difference spectra obtained from the rhabdoms after exposure to red and blue light, respectively, give evidence of a single rhodopsin with its maximum at 548 nm and a 500-nm metarhodopsin. In ERG recordings sensitivity in the dark-adapted state, after saturating exposures to blue and to red light, stabilizes at levels determined by the rhodopsin concentration. No change is observed during 10–14 h after the beginning of dark adaptation. However, using animals pre-exposed with a strong red light and then kept in darkness, it is found that after a delay of 20–40 h sensitivity of the dark-adapted eye begins to increase and finally, after 5–6 days reaches a level corresponding to 100% rhodopsin. Thus, a slow renewal of rhodopsin appears to occur in darkness, where a photoisomerization of metarhodopsin is excluded.Abbreviations ERG electroretinogram - IR infrared - MSP microspectrophotometry     

Parasitoids of the goldenrod gall moth: effects of scale on spatial density dependence

Summary A parasitoid assemblage consisting of four hymenopteran species caused larval and pupal mortality of the host Gnorimoschema gallaeosolidaginis. In the absence of discrete host patches, the relationship between densities of hosts and parasitized hosts was examined on scales of 0.25, 1, 4, 16 and 25 m² within a 400 m² plot and between ten 16 m² plots within a 1·3 km² area. Within the 400 m² plot, positive density dependence was observed on scales of 1, 4 and 16 m². Rates of parasitism were found to be density independent across the ten 16 m² plots. The nature of parasitoid and host spatial patterns were examined at the 400 m² plot by using Goodall's (1974) paired-quadrat variance function, and by fitting the observed distribution of galls and parasitized galls to the Poisson and negative binomial on three scales. Postive density dependence at the 400 m² plot occurred in the context of a host density gradient and may indicate aggregation of search time by parasitoids. However, significant responses on scales of 4 and 16 m² may only reflect the sum of parasitoid responses on lower scales because of this gradient. Predominance of significant responses on a 1 m² scale may be related to the high degree of variation in host density, and limits to active searching range of parasitoids.     

Density, climate and varying return points: an analysis of long-term population fluctuations in the threatened European tree frog

Experimental research has identified many putative agents of amphibian decline, yet the population-level consequences of these agents remain unknown, owing to lack of information on compensatory density dependence in natural populations. Here, we investigate the relative importance of intrinsic (density-dependent) and extrinsic (climatic) factors impacting the dynamics of a tree frog (Hyla arborea) population over 22 years. A combination of log-linear density dependence and rainfall (with a 2-year time lag corresponding to development time) explain 75% of the variance in the rate of increase. Such fluctuations around a variable return point might be responsible for the seemingly erratic demography and disequilibrium dynamics of many amphibian populations.     

Competitive effects of the invasive shrub,Lonicera maackii (Rupr.) Herder (Caprifoliaceae), on the growth and survival of native tree seedlings

Invasive plants are often associated with reduced cover of native plants, but rarely has competition between invasives and natives been assessed experimentally. The shrub Lonicera maackii, native to northeastern Asia, has invaded forests and old fields in numerous parts of eastern North America, and is associated with reduced tree seedling density in Ohio forests. A field experiment was conducted to test the effects of established L. maackii on the survival and growth of transplanted native tree species. The experiment examined above-ground competition (by removing L. maackii shoots) and below-ground competition (by trenching around transplanted seedlings). The effects of above-ground competition with L. maackii were generally more important than below-ground competition, though both were detected. Shoot treatment was the key determinant for the survival of all species except P. serotina, whereas trenching only enhanced survival for A. saccharum caged and P. serotina, and only in the shoot removal treatment. For the surviving seedlings, L. maackii shoot removal increased growth of A. saccharum seedlings protected with cages, but actually reduced the growth of unprotected Q. rubra and A. saccharum seedlings, indicating that L. maackii shoots confer some protection from deer browsing. Significant interactions between root and shoot treatment on Q. rubra growth parameters, specifically greatest growth in the shoot present & trenched treatment, is attributed to protection from deer browsing combined with release from below-ground competition. Despite this protective function of L. maackii shoots, the overall effect of this invasive shrub is increased mortality of native tree seedlings, suggesting it impacts the natural regeneration of secondary forests.     

Interactions within and between species at different densities of the bark beetleIps typographus and its predatorThanasimus formicarius

Interactions between the bark beetleIps typographus and one of its predators,Thanasimus formicarius, were investigated in caged spruce logs containing both species in eight different density combinations. Productively (offspring per female) ofIps was adversely affected by highIps gallery density as well as highThanasimus density.Thanasimus productivity was enhanced by highIps gallery density but negatively affected by highThanasimus density.Ips productivity andThanasimus developmental rate differed between tree individuals, probably owing to tree-related differences in phloem thickness. Relative predator-causedIps mortality was ca 20% higher at high gallery density (ca 300 egg galeries per m²) than at low gallery density (ca 100 egg galleries per m²), indicating that mortality was density dependent. This difference was due to the fact thatThanasimus larval density was positively related toIps gallery density. Mortality increased by ca 0.4% with each additionalThanasimus larva per m², independently of gallery density. Relative population levels ofIps andThanasimus were monitored with pheromone traps in two regions differing in theirIps typographus outbreak history. Absolute catch as well as theThanasimus/Ips catch ratio were ten times greater in the outbreak region than in the non-outbreak region. Coupled with the results in the caging experiment, this indicates thatT. formicarius responds numerically to changes inI. typographus numbers per unit bark area as well as toIps population changes at the regional level. The findings suggest that predation under bark may be a significant factor in supressingI. typographus outbreaks.     

Interactions between the branched-chain amino acids in the growth of Spirodela polyrhiza

The joint action of L-valine and L-isoleucine, L-leucine and L-isoleucine, and L-valine and L-leucine on the growth of Spirodela polyrhiza was established. The effect of one branched-chain amino acid on growth inhibition by another one was compared with the non-specific antagonisms which glycine and L-alanine exert on growth inhibition by singly supplied branched-chain amino acids. In this way specific and non-specific interactions could be distinguished. It appeared that: (1) L-isoleucine was a specific antagonist of L-valine; (2) L-leucine was a specific antagonist of L-isoleucine; (3) L-valine and L-leucine were synergistic growth inhibitors. Further, it was found that: (4) growth inhibition by L-leucine was specifically antagonized by simultaneously supplied L-valine and L-isoleucine; (5) an excess of L-isoleucine strongly inhibited the conversion of exogenous valine into leucine; (6) accumulation of valine was typical of isoleucine-induced growth inhibition. The results are consistent with the view that growth inhibition by L-valine and L-leucine is due to the blocking of acetohydroxy acid synthetase, the first common enzyme in the valine-isoleucine biosynthetic pathway. Growth inhibition by L-isoleucine, however, seems to result from inhibition of leucine synthesis at a step after 2-oxoisovaleric acid. Some aspects of the regulation of branched-chain amino acid biosynthesis in higher plants are discussed.     

The prevalence of non-indigenous species in southern California embayments and their effects on benthic macroinvertebrate communities

The prevalence of non-indigenous species (NIS) in southern California embayments was assessed from 123 Van Veen grab samples collected in nine bays and harbors during the summer of 1998. NIS occurred in all but two samples. They accounted for only 4.3% of the 633 taxa but contributed 27.5% of the abundance. There was no significant difference in the proportion of NIS abundance among ports harboring large vessels, small boat marinas, and areas where boats were not moored. Three species accounted for 92% of the NIS abundance: a spionid polychaete worm Pseudopolydora paucibranchiata, a mytilid bivalve Musculista senhousia, and a semelid bivalve Theora ubrica. The NIS did not appear to have a negative impact at the overall community level since NIS abundance was positively correlated with the abundance and richness of other species. This may be due to biogenic structures built by P. paucibranchiata and M. senhousia that enhance the abundances of other macrofauna.