Egg-brooding, body size and predation risk in planktonic marine copepods

We investigated the interacting effects of copepod body size and the presence or absence of egg masses on the risk of predation by a visual predator. We conducted selection experiments involving three-spined sticklebacks (Gasterosteus aculeatus) and copepods ranging in body mass from 0.5 to 740 μg C: Oithona similis, Corycaeus anglicus, Pseudocalanus newmani, P. moultoni, Pseudodiaptomus marinus, and Paraeuchaeta elongata. We found that sticklebacks selected ovigerous females of the two smallest-bodied species of copepods (Oithona similis and Corycaeus anglicus). In contrast, the fish showed no significant selection for ovigerous females of the remaining, larger-bodied species. Unexpectedly, egg mass position (i.e., in a ventral, dorsal or lateral location on the urosome) appeared to influence predation risk more than did body size, resulting in higher predation risk for the cyclopoid and poecilostomatoid species than for the calanoid species we tested. Although the sticklebacks showed no statistically significant preference for ovigerous females of any of the four calanoid species, for each species the overall proportion of ovigerous females ingested was slightly greater than 0.50. Thus, whether body size influences the susceptibility of egg-brooding calanoid copepods to predation remains an open question. Received: 24 August 1998 / Accepted: 6 July 1999     

Effects of hydroperiod and predation on a Mississippi River floodplain invertebrate community

 The objective of this study was to determine if pond permanence and vertebrate predation (by fish and waterfowl) affect invertebrate community structure in the mudflat habitat of floodplain ponds. Invertebrate communities were studied for 1 year in four Mississippi River floodplain ponds with different hydroperiods. Pond 1 experienced five dry periods, pond 2 experienced four, pond 3 dried once, and standing water remained in pond 4 for the entire year. Vertebrate predator exclusion treatments (all access, no access, small-fish access and cage controls) were placed in all ponds. As pond duration increased, predatory invertebrate richness and abundance increased while overall invertebrate richness and abundance decreased. With the exception of the cladoceran Diaphanosoma, all commonly encountered taxa were strongly affected by pond permanence in terms of abundance, biomass and, generally, individual biomass. Taxa were nearly early divided between those that were more abundant in less permanent ponds and those that were more abundant in longer-duration ponds. Invertebrate taxa richness, abundance, and total biomass were lower in the all-access treatment than in the treatments that restricted predator access, and these effects were stronger in the more permanent ponds. In general, there were no significant differences in responses to the treatments with small-fish access and no access. These results support models that predict relatively weak effects of predation in frequently disturbed habitats. Received: 30 May 1995 / Accepted: 21 June 1996     

Factors affecting body size and fat content in a digger wasp

Body size is one of the most important life history traits. In mass-provisioning solitary Hymenoptera, the maximum attainable adult size is not under the control of the larva but is limited by the amount of resources provided by the mother. I investigated the effect of the amount of different maternal resources and potentially interfering abiotic (temperature) and biotic (fungus infestation) factors on offspring body size and fat reserves in a solitary digger wasp, the European beewolf, Philanthus triangulum F. (Hymenoptera, Sphecidae). Females provide different resources for their progeny that might influence progeny size (egg, brood cell, and paralysed honey-bees as food). The number of bees provisioned explained the largest proportion of variation in cocoon length. With an increasing number of bees (one to four), progeny gained less weight per bee. Relative fat content increased with size. With a given number of bees, males were smaller than females. The duration of the feeding period was independent of the number of bees in a brood cell but decreased with increasing ambient temperatures (20, 25, 30°C). Cocoon size was influenced by temperature but the effect was not uniform. Cocoons from brood cells containing two and three bees were larger at 25°C than at 20°C; those at 30°C did not differ from those at either lower temperature. However, in brood cells containing one bee, cocoon length was independent of temperature. Sublethal levels of fungus infestation may have a small negative effect on cocoon size. Different temperatures during hibernation (8 vs 13°C) did not affect the size or fat content of emerging adults. These results on a mass-provisioning hunting wasp are compared with the well-studied herbivorous insects. Received: 12 April 1999 / Accepted: 3 December 1999     

Estimation of fine root mortality and growth from simple measurements: a method based on system dynamics

This study is a further development of a dynamic compartment-flow analysis, intended as an analytical tool for the empirical estimation of fine root growth, mortality and decomposition in forest soil. General properties of the dynamic system are utilised to interpret relatively simple measurements of standing biomass, necromass, and decomposition, in order to derive estimates of the process rates. The method is based on the finding that the ratio of fine root necromass to biomass is related to the specific rates of decomposition, mortality, and net growth. If the decomposition rate is measured and the net growth trend is determined from live root measurements, mortality and gross growth can be estimated using these relationships, provided certain regularity requirements are met. These requirements are explicated, such that the estimates can be easily assessed for reliability. To illustrate the use of the method, it was applied to the estimation of specific mortality rates in seven Scots pine stands of different ages and site types. A reanalysis of a previous sequential coring study yielded consistent results. The advantage of this method is that, unlike the standard analysis of sequential cores, it accounts for the possibility of simultaneous growth, mortality and decomposition. It is therefore applicable to situations with no apparent fluctuations or trend in the biomass and necromass levels. No minimum sampling interval is required; hence the method also allows for more extensive or prolonged studies. Received: 1 June 1999 / Accepted: 20 January 2000     

Interception and retention of macroscopic bryophyte fragments by branch substrates in a tropical cloud forest: an experimental and demographic approach

Although the significance of canopy plant communities to ecosystem function is well documented, the process by which such communities become established in trees remains poorly known. Colonization of tree surfaces by canopy-dwelling plants often begins with the establishment of bryophytes, so the conditions that affect the dispersal of bryophytes in the forest canopy merit study. We assessed success rates of one mechanism of bryophyte propagation, the aerial dispersal of macroscopic fragments, using an experimental approach. We quantified interception and retention of marked fragments released from a 36 cm×36 cm grid 50 cm above branches of saplings and mature trees of the species Ocotea tonduzii in a montane cloud forest in Costa Rica. Only 1% of bryophyte fragments dropped over sapling crowns in this manner were retained for the 6-month duration of the study, while branches in the forest canopy with intact epiphyte loads and branches that had been stripped of their epiphytes retained 24% and 5%, respectively. Our results suggest that larger-diameter branches and the presence of other epiphytes can both improve the retention of bryophyte fragments on canopy branches. Further work will be needed to address the relative roles of other dispersal mechanisms (spores, gemmae, microscopic bryophyte fragments) and the dynamics of growth and establishment of macroscopic bryophyte fragments following their interception. Received: 1 March 1999 / Accepted: 24 August 1999     

On the role of body size in a tri-trophic metapopulation model

 A particular tri-trophic (resource, prey, predator) metapopulation model with dispersal of preys and predators is considered in this paper. The analysis is carried out numerically, by finding the bifurcations of the equilibria and of the limit cycles with respect to prey and predator body sizes. Two routes to chaos are identified. One is characterized by an intriguing cascade of flip and tangent bifurcations of limit cycles, while the other corresponds to the crisis of a strange attractor. The results are summarized by partitioning the space of body sizes in eight subregions, each one of which is associated to a different asymptotic behavior of the system. Emphasis is put on the possibility of having different modes of coexistence (stationary, cyclic, and chaotic) and/or extinction of the predator population. Received 1 August 1995; received in revised form 8 January     

Use of an ultrasound cell retention system for the size fractionation of somatic embryos of woody species

 The potential of ultrasonic standing waves for trapping suspended particles was utilized successfully for differential size fractionation of plant somatic embryos. In a flow-through resonator equipped with a 200-kHz piezoceramic transducer, embryos of different sizes corresponding to different developmental stages could be retained by varying the electric power input and flow speed. The system was initially established for carrot (Daucus carota) somatic embryos and subsequently adapted for the larger-sized embryos of the woody species cork oak (Quercus suber), grapevine (Vitis Berlandieri × rupestris) and cherry (Prunus incisa × serrula). Separation performance was confirmed by analysing the different fractions for the expression of homeobox genes which are differentially expressed during embryogenesis. No inhibitory effects of embryos on short- and long-term development could be observed. Received: 6 December 1999 / Revised: 8 March 2000 / Accepted: 13 March 2000     

Interactive effects of habitat selection, food supply and predation on recruitment of an estuarine fish

 Seagrass meadows are often important habitats for newly recruited juvenile fishes. Although substantial effort has gone into documenting patterns of association of fishes with attributes of seagrass beds, experimental investigations of why fish use seagrass habitats are rare. We performed two short-term manipulative field experiments to test (1) the effects of food supply on growth and densities of fish, and (2) effects of predation on the density and size distribution of fish recruits, and how this varies among habitat types. Experiments were conducted in Galveston Bay, Texas, and we focused on the common estuarine fish, pinfish Lagodon rhomboides. In the first experiment, replicate artifical seagrass and sand plots were either supplemented with food or left as controls. Recruitment of pinfish was significantly greater to seagrass than sand habitats; however, we detected no effect of food supplementation on the abundance of recruits in either habitat. Pinfish recruits in artifical seagrass grew at a significantly faster rate than those in sand habitats, and fish supplemented with food exhibited a greater growth rate than controls in both sand and artifical grass habitats. In our second experiment, we provided artificial seagrass and sand habitats with and without predator access. Predator access was manipulated with cages, and two-sided cages served as controls. Recruitment was significantly greater to the cage versus cage-control treatment, and this effect did not vary between habitats. In addition, the standard length of pinfish recruits was significantly larger in the predator access than in the predator exclusion treatment, suggesting size-selective predation on smaller settlers or density-dependent growth. Our results indicate that the impact of predation on pinfish recruits is equivalent in both sand and vegetated habitats, and thus differential predation does not explain the higher recruitment of pinfish to vegetated than to nonvegetated habitats. Since predators may disproportionately affect smaller fish, and a limited food resource appears to be more effectively utilized by fish in vegetated than in unvegetated habitats, we hypothesize that pinfish recruits may select vegetated habitats because high growth rates allow them to achieve a size that is relatively safe from predation more quickly. Received: 10 October 1996 / Accepted: 5 April 1997     

Biodiversity-productivity relations: an experimental evaluation of mechanisms

To examine the mechanisms underlying productivity-diversity relationships, we manipulated nutrient levels in replicate small-scale artificial habitat units in the marine subtidal zone and followed the process of community assembly. In contrast to most enrichment studies, algal diversity increased in enriched habitats relative to controls along with biomass – a result that may be explained by the low nutrient status of the region. Both the total number of faunal species and the total number of individuals were also significantly greater in enriched habitats, but the relationship between algal resources and faunal diversity did not support the resource heterogeneity hypothesis. Received: 14 June 1999 / Accepted: 29 October 1999     

Initial steps in lymph node metastasis formation in an experimental system: possible involvement of recognition by macrophage C-type lectins

 We used histological observations and experiments with fluorescent cell tracers to investigate the roles of tissue macrophages in recognition through a galactose/N-acetylgalactosamine-specific C-type lectin (mMGL) in lymph node metastasis formation by mouse ovarian tumor OV2944-HM-1 (HM-1) cells. Lymph node metastasis from subcutaneous sites was shown to be initiated by the entry of tumor cells into the subcapsular sinus of lymph nodes where mMGL-positive cells were mainly located. To investigate whether mMGL-positive cells contributed to host resistance against lymph node metastasis, we repeatedly treated mice bearing transplanted tumors with an mMGL-blocking monoclonal antibody that was known to inhibit mMGL binding to its ligands. The number of HM-1 cells recovered from lymph nodes 2 weeks after subcutaneous injections was significantly greater when the mice were treated with the blocking anti-mMGL antibody. These results suggested that mMGL-positive macrophages contributed to the host's defense against lymph node metastasis. Received: 30 July 1999 / Accepted: 1 November 1999     

Effect of patch size and plant density of Paterson"s curse (Echium plantagineum) on the oviposition of a specialist weevil, Mogulones larvatus

This study examines the oviposition response of a specialist weevil (Mogulones larvatus) to patches of its host, the noxious weed Paterson"s curse/salvation Jane (Echium plantagineum). We simultaneously examined the effect of patch size and plant density (and their interaction), on the recorded oviposition patterns. Our results show that oviposition first occurred on the largest patches with the highest number of plants. However, there was no significant effect of patch size or number of plants per patch at the end of the experiment. At this time the level of attack per plant was negatively correlated with plant density. This negative effect of density on the level of oviposition was not mediated by a reduction in plant size at higher densities. The pattern observed may indicate a risk-spreading strategy by females. Received: 15 July 1999 / Accepted: 14 April 2000     

Choice of optimal oviposition sites by Hoplobatrachus occipitalis (Anura: Ranidae) in an unpredictable and patchy environment

 The rock pools on the river bank of the Comoé National Park (West Africa) provide a very diverse and unpredictable environment for anuran larval development. Because rock pools differ considerably in biotic and abiotic parameters, it should be adaptive for reproducing anurans to choose the most suitable oviposition sites. During the beginning of each rainy season (March to May), from 1991 to 1995, we investigated the allocation of breeding sites by Hoplobatrachus occipitalis, counted the number of eggs, and measured several biotic and abiotic factors that might influence the choice of a spawning site. The probability of predation by conspecific cannibalistic tadpoles and the water-holding capacity (WHC) of pools were the best predictor of number of eggs laid. We experimentally investigated the influence of these two parameters on egg-laying and showed that adults potentially can assess the presence, density, and size of tadpoles in pools by chemical cues. Likewise, manipulation of the water-holding capacity caused a rapid change in egg-laying behavior. To assess the risk of desiccation, frogs have to visit familiar pools repeatedly to monitor the decrease in volume over time and thus gain information about the pools’ water-holding capacity. Received: 22 April 1996 / Accepted: 17 July 1996     

Influence of leaf size on tree architecture: first branch height and crown dimensions in tropical rain forest trees

 First branch height is an important attribute of sapling architecture, as it defines the height at which prolonged lateral growth is possible. First branch height, measured on saplings of 70 species in tropical rain forests of Australia, Costa Rica, Panama, and Sabah, Malaysia, was highly correlated with leaf blade and petiole length. The observed relationship, first branch height ∝ blade length × (petiole length)^0.5, implies that the ratio of first branch height to blade length increases somewhat with increasing leaf size, among species with a given ratio of petiole to blade length. Orthotropic species, with more or less radially symmetric arrangements of leaves on ascending axes, had a mean first branch height of 7x that observed for plagiotropic species, with planar leaf arrangements. The greater first branch height of orthotropic species was associated with their larger leaves and longer petioles. Plagiotropic species had wider crowns than orthotropic species in the sapling stage, as assessed at the Costa Rican site. Thus, leaf dimensions influence the dynamics of crown construction (or visa versa), as well as affecting leaf energy balance and gas exchange. Received: 5 September 1997 / Accepted: 3 March 1998     

Sequential bulked typing: a rapid approach for detecting QTLs

A strategy of DNA pooling aimed at identifying markers linked to quantitative trait loci (QTLs), ‘Sequential Bulked Typing’ (SBT), is presented. The method proposed consists in pooling DNA from consecutive pairs of individuals ranked phenotypically, i.e., pools are formed with individuals ranked (1st, 2nd), (3rd, 4th),…, (N-1st, Nth). The N/2 pools are subsequently amplified using the polymerase chain reaction (PCR). If the whole population is typed the number of PCRs per marker is halved with respect to individual typing (IT). But if this strategy is combined with selective genotyping of extreme individuals savings can be further increased. Two extreme cases are considered: in the first one (SBT⁰), it is assumed that only presence or absence of a given allele can be ascertained in a pool; in the second one (SBT¹), it is further assumed that differences between allele band intensities can be distinguished. The theory to estimate by maximum likelihood the QTL effect and its position with respect to flanking markers is presented. The behaviour of IT and SBT was studied using stochastic computer simulation in backcross and F₂ populations. Three percentages of subpattern distinction (0, 50 and 100%) two population sizes (n=1200 and 600) and two QTL effects (a=0.1 and 0.25 standard deviations) were considered. SBT¹ had the same power as individual genotyping at half the genotyping costs in all situations studied. Accuracy of QTL location is not increased with a dense number of markers, as opposed to individual typing. As a result DNA pooling is not useful for accurate location of the QTL but rather to pick up genome regions containing QTLs of at least moderate effect. The theory developed provides the general theoretical framework to deal with any DNA pooling strategy aimed at detecting QTLs. Received: 15 September 1997 / Accepted: 6 October 1997     

Spring ephemeral herbs and nitrogen cycling in a northern hardwood forest: an experimental test of the vernal dam hypothesis

In the late 1970s R.N. Muller and F.H. Bormann posited their ”vernal dam” hypothesis, stating that spring-ephemeral herbs in deciduous forests serve as a temporary sink for N when overstory trees are dormant, and then release this N later, in the summer, when the trees are active. This hypothesis has gained wide acceptance, yet two of its critical assumptions have never been experimentally tested: (1) that N taken up by spring ephemerals would otherwise be lost from the ecosystem, and (2) that N from senesced ephemeral tissues contributes to increased rates of summertime N mineralization. To test these assumptions, I quantified patterns of N cycling and loss from a set of paired plots, half of which served as controls and from half of which all spring-ephemeral plants were removed. There were no significant differences in NO₃ ^– leaching between plots with and without spring ephemeral vegetation. These results are consistent with the relatively low rates of N uptake by the dominant spring ephemeral, Allium tricoccum, and its apparent preference for NH₄ ⁺, which is far less mobile in soil than NO₃ ^–. In addition, based on sequential sampling, I found that soil microorganisms took up 8 times as much N during the spring than did spring-ephemeral herbs (microbial uptake=3.19 vs. plant uptake=0.41 g N m^–2), suggesting that microbial immobilization of N is the dominant sink for N during this season. Removal of spring ephemeral vegetation also had no effect on summertime rates of net N mineralization. Furthermore, the addition of spring ephemeral litter to soil+forest floor microcosms did not significantly increase rates of N mineralization in a laboratory incubation. Instead, this experiment demonstrated the overwhelming influence of forest floor litter in controlling the release of mineral N from these soils. Overall, neither assumption of the vernal dam hypothesis holds true in this ecosystem, where patterns of N cycling and loss appear to be dominated by microbial decomposition of forest floor material and soil organic matter. Received: 24 August 1999 / Accepted: 23 March 2000     

The effects of plant dispersion and prey density on parasitism rates in a naturally patchy habitat

Despite extensive research on parasitoid-prey interactions and especially the effects of heterogeneity in parasitism on stability, sources of heterogeneity other than prey density have been little investigated. This research examines parasitism rates by three parasitoid species in relationship to prey density and habitat spatial pattern. The herbivore Itame andersoni (Geometridae) inhabits a subdivided habitat created by patches of its host plant, Dryas drummondii, in the Wrangell Mountains of Alaska. Dryas colonizes glacial moraines and spreads clonally to form distinct patches. Habitat subdivision occurs both on the patch scale and on the larger spatial scale of sites due to patchy successional patterns. Itame is attacked by three parasitoids: an ichneumonid wasp (Campoletis sp.), a braconid wasp (Aleiodes n. sp.), and the tachinid fly (Phyrxe pecosensis). I performed a large survey study at five distinct sites and censused Itame density and parasitism rates in 206 plant patches for 1–3 years. Parasitism rates varied with both plant patch size and isolation and also between sites, and the highest rates of overall parasitism were in the smallest patches. However, the effects of both small- and large-scale heterogeneity on parasitism differed for the three parasitoid species. There was weak evidence that Itame density was positively correlated with parasitism for the braconid and tachinid at the patch scale, but density effects differed for different patch sizes, patch isolations, and sites. At the site scale, there was no evidence of positive, but some indication of negative density-dependent parasitism. These patterns do not appear to be driven by negative interactions between the three parasitoid species, but reflect, rather, individual differences in habitat use and response to prey density. Finally, there was no evidence that parasitism strongly impacted the population dynamics of Itame. These results demonstrate the importance of considering habitat pattern when examining spatial heterogeneity of parasitism and the impacts of parasitoids. Received: 3 June 1999 / Accepted: 4 October 1999     

Movement coordination and signalling in ravens (Corvus corax): an experimental field study

Vagrant non-breeding ravens frequently attract conspecifics to rich ephemeral food sources. There, grouping may allow them to overcome the defence of territorial breeders. Here, we focus on ravens making use of regular food supplies in a game park, where they divert food from the provision of park animals. We investigated if ravens foraging in the Cumberland game park (Grünau, Austria) are attentive towards one another when they experience some unpredictability in food provisioning. We confronted a group of 30–50 ravens with two different treatments. Ten minutes ahead of the feeding of either wolves or wild boars we showed buckets containing pieces of meat to the ravens flying overhead. In the reliable cue treatment (RCT), the meat was placed next to one of the two enclosures, whereas in the unreliable cue treatment (UCT), the buckets were placed simultaneously in front of both enclosures though only in one of the enclosures were the animals fed 10 min later. Thus, during RCT but not during UCT, ravens could predict where food would become available. Only during UCT, ravens moved in large groups between the two feeding sites. Many ravens moving at the same time in the same direction may indicate some co-ordination in space and time, which is most likely achieved by social attraction among individuals. Furthermore, the number of ravens approaching and leaving, respectively, a feeding site cross-correlated with a temporary increase in the rate of a food-associated call, the yell. This suggests that in addition to watching each other, calling may have contributed to group formation. Possible benefits of group formation during food inspection are discussed. Received: 9 February 2000 / Received in revised form: 30 May 2000 / Accepted: 1 June 2000