Carabid beetle condition,reproduction and density in winter oilseed rape affected by field and landscape parameters

The carabid beetles Amara similata and Poecilus cupreus are abundant in Central European winter oilseed rape (OSR) fields and potential antagonists of OSR pests. Therefore, they were investigated in 29 OSR fields relative to the influence of field and landscape parameters on their nutritional condition, reproductive potential and activity density. Nutritional condition was measured by a condition factor (CF). Fecundity of female beetles was expressed by the number of ripe oocytes in the ovaries. Activity density comprised the number of individuals caught with pitfall traps during their peak of reproductive period. Soil productivity and pest abundance (as a proxy of local prey availability) were considered as field parameters. Landscape parameters comprised the distance to the next fallow and the percentage of crop area around each study field. In eight of ten cases, pest abundance proved to be the most important factor explaining carabid characteristics, indicating that P. cupreus and A. similata consume insect pests in OSR fields. Pest abundance influenced the CF of both species and sexes positively. Oocyte numbers of A. similata were negatively related to the distance to the next fallow. Oocyte numbers of P. cupreus tended to increase with increasing pest abundance. Activity density of female A. similata was negatively influenced by soil productivity. Activity density of male A. similata and of female and male P. cupreus was negatively influenced by pest abundance. When analysing exclusively the influence of severe OSR pests (Meligethes aeneus, Ceutorhynchus napi and Dasineura brassicae), abundance of larval M. aeneus and C. napi was most important in explaining the CF and activity density of male A. similata, and male and female P. cupreus. M. aeneus and C. napi could be an essential prey for the carabids studied which may reduce the pests, thereby contributing to the protection of OSR from pest herbivory.     

Migration to and dispersal from oilseed rape by the pollen beetle,Meligethes aeneus,in relation to wind direction

1 Host‐plant‐odour‐induced upwind anemotaxis is accepted as the main mechanism by which herbivorous insects find their host plant within an heterogenous environment, but field data supporting this hypothesis are scarce. 2 The flight directions of the pollen beetle Meligethes aeneus to and from a plot of winter oilseed rape and the direction of the wind were recorded concurrently. The beetles were sampled by eight double‐sided window traps encircling the plot. Distal sides of the traps, relative to the plot, sampled the beetles as they flew towards the plot, whereas the proximal sides of the traps sampled them as they flew away from the plot. Paired data on daily catch of beetles in distal or proximal traps and the volume of air impacting each trap were compared. 3 Correlations between daily catch of M. aeneus into distal traps and trap wind volume were negative, indicating that flights by both overwintered‐ and new‐generation insects towards the plot were upwind. 4 Correlations between daily catch of M. aeneus into proximal traps and trap wind volume varied with generation. Catch of overwintered‐generation M. aeneus was negatively correlated with trap wind volume, indicating that flight was upwind. Proximal trap catch of the new‐generation M. aeneus was weakly/modestly positively correlated with trap wind volume, indicating that flights were downwind or crosswind. 5 Understanding the effect of wind direction on flight of M. aeneus holds potential for improving the forecasting of their arrival and spatial distribution on the crop for integrated pest management.     

Wind‐modulated landscape effects on colonization of Brussels sprouts by insect pests and their syrphid antagonists

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The Distribution of Tardigrades Upwind and Downwindof a Missouri Coal-Burning Power Plant

Significant differences occurred in the density of tardigrades, rotifers, and nematodes and the diversity of tardigrades between collecting sites located upwind and downwind from a coal-burning power plant in Missouri. The oak tree species and lichen genera also varied in the two areas. Tardigrade and rotifer densities were greater in upwind sites, whereas nematode density was higher in downwind samples. One tardigrade species (Ramazzottius sp.) was found only at the upwind sites, and one species (Echiniscus sp.) was only in the downwind samples. In contrast, three species (Macrobiotus sp., Minibiotus sp., and Milnesium tardigradum) were found both upwind and downwind but in different densities in the two areas. The study presents baseline data for long-term monitoring of the effects of environmental factors on nematode and rotifer densities as well as tardigrade density and diversity.     

Insect pests in winter oilseed rape affected by field and landscape characteristics

Winter oilseed rape (OSR, Brassica napus) cropping is often associated with an intensive use of pesticides. The transformation of structurally rich landscapes into more monotonous landscapes may be partly responsible for this, because non-crop habitats believed to benefit natural enemies have been eliminated. We examined the influence of field (soil quality, nitrogen fertilization) and landscape characteristics (OSR area and isolation, non-crop area, landscape diversity, proportions of grassy fallows and woody areas) on three major European OSR pest groups: pollen beetles, stem weevils, and brassica pod midges. Twenty-nine landscape sectors ranging from structurally poor to complex were studied at eight spatial scales (radii 250–2000 m) centered in the studied OSR fields. Abundances of pollen beetles and stem weevils were significantly positively correlated with soil quality and negatively related to OSR area in the surroundings. Generally, abundances of all groups were positively related to woody areas, but not related to grassy fallow area. Pod midges and stem weevils tended to respond primarily to landscape variables at small (250–500 m) and medium (1000–1250 m) scales, while pollen beetles responded at medium to large (1000–2000 m) scales. The results are discussed in relation to differences in overwintering strategies and mobility of pest insects. Strategies at the field and landscape level, aiming to reduce pest pressure in OSR fields, are also discussed.     

Regional-scale impact of the weed biocontrol agent Mecinus janthinus on Dalmatian toadflax (Linaria dalmatica)

Dalmatian toadflax (Linaria dalmatica (L.) Mill.) is an important invasive plant on rangelands throughout western North America. In 1991, the stem-mining weevil, Mecinus janthinus Germar, was introduced into Canada from Europe as a classical biological control agent to reduce toadflax densities and improve rangelands, particularly in British Columbia. To determine if the program was a success at a regional level, this paper answers three key questions: (1) has M. janthinus spread throughout the study area, (2) is M. janthinus causing a decline in toadflax plant size or density at the regional scale, and (3) has the distribution of toadflax plants changed following M. janthinus introduction? These questions are answered by combining historical survey data and mensurative experimental data on plant and weevil densities across a 40,000 km² area in southern British Columbia. The results show that through a combination of intentional redistribution and natural dispersal weevils have spread throughout the study area. Stem densities at naturally colonized sites and historical release sites were equally low. Across weevil populations between 3 and 10 years old, weevil densities peaked in the eighth year, and there was a negative relationship between weevil density and stem length. Between 2000 and 2007, toadflax patches were found to both decrease in density and become more fragmented over time, with 15% of patches disappearing completely. These findings show that M. janthinus has had a significant negative impact on both the density and distribution of Dalmatian toadflax throughout a large part of its range in British Columbia.     

Plant Quantity Affects Development and Survival of a Gregarious Insect Herbivore and Its Endoparasitoid Wasp

Virtually all studies of plant-herbivore-natural enemy interactions focus on plant quality as the major constraint on development and survival. However, for many gregarious feeding insect herbivores that feed on small or ephemeral plants, the quantity of resources is much more limiting, yet this area has received virtually no attention. Here, in both lab and semi-field experiments using tents containing variably sized clusters of food plants, we studied the effects of periodic food deprivation in a tri-trophic system where quantitative constraints are profoundly important on insect performance. The large cabbage white Pieris brassicae, is a specialist herbivore of relatively small wild brassicaceous plants that grow in variable densities, with black mustard (Brassica nigra) being one of the most important. Larvae of P. brassicae are in turn attacked by a specialist endoparasitoid wasp, Cotesia glomerata. Increasing the length of food deprivation of newly molted final instar caterpillars significantly decreased herbivore and parasitoid survival and biomass, but shortened their development time. Moreover, the ability of caterpillars to recover when provided with food again was correlated with the length of the food deprivation period. In outdoor tents with natural vegetation, we created conditions similar to those faced by P. brassicae in nature by manipulating plant density. Low densities of B. nigra lead to potential starvation of P. brassicae broods and their parasitoids, replicating nutritional conditions of the lab experiments. The ability of both unparasitized and parasitized caterpillars to find corner plants was similar but decreased with central plant density. Survival of both the herbivore and parasitoid increased with plant density and was higher for unparasitized than for parasitized caterpillars. Our results, in comparison with previous studies, reveal that quantitative constraints are far more important that qualitative constraints on the performance of gregarious insect herbivores and their gregarious parasitoids in nature.     

Morphological and molecular characterizations of the Gregarina sp. (Apicomplexa: Protozoa) parasitizing on Phaedon brassicae (Coleoptera: Chrysomelidae)

A gregarine parasite (Eugregarinida: Gregarinidae) was observed in the population of daikon leaf beetle, Phaedon brassicae Baly, (Coleoptera: Chrysomelidae) in Korea. Gregarines are well known species-specific parasites of various Arthropoda. This Gregarina sp. also confirmed a species-specific parasite in P. brassicae. Based on 1.727 kb of 18S rDNA sequence (FJ481523), this gregarine species was grouped in eugregarine and a 5.258 kb of full length rDNA replicon was cloned (JF412715). We also observed interaction of trophozoite or gamonto of gregarine and epithelium of a host midgut using a light microscope and a scanning electron microscope. Although the developmental period of the infected host is delayed half a day in every larval stage, there was no significant difference in the developmental period of P. brassicae whether Gregarina sp. was infected or not. Gregarina sp. was a kind of facultative parasite from P. brassicae. This is the first report of a gregarine parasite in P. brassicae.     

Waterfowl grazing effects on submerged macrophytes in a shallow mediterranean lake

Waterfowl exclusion cages were set up in Sentiz Lake, an eutrophic shallow lake in León (NW of Spain) in order to determine the role of waterfowl herbivory on macrophyte biomass and species composition. Total macrophyte biomass was high during the study (250 g DW m⁻² in summer). The macrophyte community was mainly formed by Myriophyllum alterniflorum (95% cover), Ceratophyllum demersum (5%) and Potamogeton gramineus (<0.5%). High densities of co-occurring coots (Fulica atra; 24 ind/ha) and ducks (Anas penelope, A. strepera and A. platyrhynchos; 18 ind/ha) did not have a significant effect on macrophyte biomass in the lake. There were no statistical differences between total biomass inside and outside the exclosures, although plant biomass reached a higher value inside the cages than in the lake. Biomass species composition was significantly different inside and outside exclosures; C. demersum was more abundant in the cages than in the lake. P. gramineus, almost absent in the lake, became co-dominant with M. alterniflorum in some exclosures. The detailed study of M. alterniflorum flower buds in summer showed significant herbivory by coots. Flower bud abundance was lower in the lake (35% lower in June; 85% lower in July) than under waterfowl exclusion. The effect of waterfowl on macrophyte biomass in Mediterranean wetlands seems to be negligible as compared to effects identified in northern European lakes. Apart from an important role in dispersal, waterfowl in Mediterranean areas have a strong qualitative effect on the structure of plant communities by selecting most palatable species or their reproductive structures.     

Role of Proteolysis in Determining Potency of Bacillus thuringiensis Cry1Ac δ-Endotoxin

Bacillus thuringiensis protein δ-endotoxins are toxic to a variety of different insect species. Larvicidal potency depends on the completion of a number of steps in the mode of action of the toxin. Here, we investigated the role of proteolytic processing in determining the potency of the B. thuringiensis Cry1Ac δ-endotoxin towards Pieris brassicae (family: Pieridae) and Mamestra brassicae (family: Noctuidae). In bioassays, Cry1Ac was over 2,000 times more active against P. brassicae than against M. brassicae larvae. Using gut juice purified from both insects, we processed Cry1Ac to soluble forms that had the same N terminus and the same apparent molecular weight. However, extended proteolysis of Cry1Ac in vitro with proteases from both insects resulted in the formation of an insoluble aggregate. With proteases from P. brassicae, the Cry1Ac-susceptible insect, Cry1Ac was processed to an insoluble product with a molecular mass of ~56 kDa, whereas proteases from M. brassicae, the non-susceptible insect, generated products with molecular masses of ~58, ~40, and ~20 kDa. N-terminal sequencing of the insoluble products revealed that both insects cleaved Cry1Ac within domain I, but M. brassicae proteases also cleaved the toxin at Arg423 in domain II. A similar pattern of processing was observed in vivo. When Arg423 was replaced with Gln or Ser, the resulting mutant toxins resisted degradation by M. brassicae proteases. However, this mutation had little effect on toxicity to M. brassicae. Differential processing of membrane-bound Cry1Ac was also observed in qualitative binding experiments performed with brush border membrane vesicles from the two insects and in midguts isolated from toxin-treated insects.     

Risk Assessment Studies: Detailed Host Range Testing of Wild-Type Cabbage Moth, Mamestra brassicae (Lepidoptera: Noctuidae), Nuclear Polyhedrosis Virus

The host range of a multiply enveloped nuclear polyhedrosis virus (NPV) (Baculoviridae) isolated from the cabbage moth, Mamestra brassicae (Lepidoptera: Noctuidae), was determined by challenging a wide range of insect species with high (10⁶ polyhedral inclusion bodies) and low (10³ polyhedral inclusion bodies) doses of the virus. The identity of the progeny virus was confirmed by dot blotting. Analysis of 50% lethal dose was carried out on selected species, and the progeny virus was identified by using restriction enzyme analysis and Southern blotting. Other than the Lepidoptera, none of the species tested was susceptible to M. brassicae NPV. Within the Lepidoptera, M. brassicae NPV was infective to members of four families (Noctuidae, Geometridae, Yponomeutidae, and Nymphalidae). Of 66 lepidopterous species tested, M. brassicae NPV was cross-infective to 32 of them; however, 91% of the susceptible species were in the Noctuidae. The relevance of host range data in risk assessment studies is discussed.     

Attraction of the predator,Harmonia axyridis (Coleoptera: Coccinellidae), to its prey,Myzus persicae (Hemiptera: Aphididae), feeding on Chinese cabbage

The predatory multicolored Asian lady beetle, Harmonia axyridis, was attracted to volatiles released from Chinese cabbage infested by the green peach aphid, Myzus persicae, in T-tube olfactometer choice tests. However, lady beetle adults and larvae did not respond to clean air, Chinese cabbage alone or green peach aphid alone. Of different prey densities, H. axyridis adults were most attracted to Chinese cabbage infested by 60 M. persicae adults after 24 h. However, H. axyridis larvae were not attracted to Chinese cabbage infested by M. persicae. Mechanically damaged Chinese cabbage attracted neither lady beetle adults nor larvae. Predatory adults were attracted to 60 M. persicae adults after 24 and 48 h, and to 90 M. persicae adults after 12 h, suggesting that the predatory response depends on the prey density. Lady adult beetles did not prefer the volatiles induced by Diamondback moth, Plutella xylostella, indicating that specific host insect specificity attracts respective natural enemies. It can be explained that the volatile compounds emitted from the host plant as a result of herbivore attack preferred by the specific insect species.     

Interference between the macroalga Caulerpa prolifera and the seagrass Halodule wrightii

A field experiment was conducted to examine interactions between the seagrass Halodule wrightii and the macroalga Caulerpa prolifera in the Indian River Lagoon, FL, USA, and further if the outcome of the interactions between the two species was influenced by water depth. The experiment involved the manipulation of neighbor presence in plots established at the shallowest (50 cm) and deepest (80 cm) depths at which the two species co-existed at adequate densities to perform removal experiments. Shoot and frond densities were measured at the beginning (April), middle (July) and end (October) of the 6-month growing season, and above- and below-ground biomass values were determined at the end of the experiment (October). In the middle of the growing season H. wrightii had higher shoot densities and greater biomass in plots where C. prolifera had been removed at both water depths. This same pattern in shoot density and biomass for H. wrightii also occurred at the end of the growing season at the 80 cm depth. C. prolifera occurred at higher densities and greater biomass in the 80 cm depth range, but was generally unaffected by the presence of H. wrightii at either depth.     

Potential for oilseed rape resistance in pollen beetle control

Breeding for plant resistance to insect pests is a classic strategy in integrated management, but it has never been developed for use against European pests of oilseed rape (Brassica napus) (OSR), especially one of the most damaging ones, the pollen beetle (Meligethes aeneus). In this paper we look at the three strategies that could be employed to improve OSR resistance (based on transgenes, relatives of B. napus or OSR natural variation) and review our current knowledge as to how these strategies could be put into practice. We identify the drawbacks which are specific to the pollen beetle that could impede breeding programs for resistance, and propose an approach to circumvent them. Finally, we detail the steps of the interaction between OSR and the pollen beetle that could be targeted in order to improve plant resistance (host plant location, adult survival, adult feeding, egg production and oviposition, larval development) and discuss their efficiency and durability potential.     

Influences of sedimentation and hydrodynamics on the spatial distribution of Alexandrium catenella/tamarense resting cysts in a shellfish farming lagoon impacted by toxic blooms

Since resting cysts are a potential seeding source for blooms, the presence of these cysts in sediments is a marker of an established population for a number of harmful algal species. The spatial patterns of cyst density in relation to sediment characteristics and hydrodynamics are still largely misunderstood. This study investigated the spatial distribution of resting cysts belonging to the Alexandrium tamarense species complex (Dinophyceae) in sediments of a Mediterranean coastal lagoon (Thau Lagoon, France). This lagoon, hosting shellfish farming, is regularly impacted by toxic Alexandrium catenella blooms. The average cyst density across the whole lagoon was rather low, <20 cysts g⁻¹ of dry sediment (DS). However, densities varied widely among sampled stations, with the highest density (∼440 cysts g⁻¹ DS) recorded in a shallow cove named Crique-de-l’Angle, which is the only area where dense blooms of A. catenella and A. tamarense have been recorded in the years preceding this survey. An analysis using spatial autoregressive models demonstrated that cyst densities were highly spatially autocorrelated (indicating that close stations tended to have more similar cyst densities) with accumulation sites. With respect to sediment characteristics (5 granulometric fractions <2 mm and biochemical components), the highest densities were found in silty sediments containing high proportions of water and organic matter. Nevertheless, the linkage between cyst density and sediment structure was not always verified; this reflected the influence of hydrodynamics on the sedimentation of cysts and sediment particles, and on the dispersal of cysts away from the bloom area by wind-induced currents, suggesting that hydrodynamics was responsible for the spatially autocorrelated distribution of cyst densities.     

Pollen beetles are consumed by ground‐ and foliage‐dwelling spiders in winter oilseed rape

Pollen beetles, Meligethes aeneus (Fabricius) (Coleoptera: Nitidulidae), are major pests in oilseed rape (OSR), Brassica napus L. (Brassicaceae). Among the predator species in the generalist predator complex present in OSR fields, wolf spiders (Araneae: Lycosidae) are found on the ground and cobweb spiders (Araneae: Theridiidae) build webs in the foliage. Here we study the incidence of predation of pollen beetles by these two spider groups using DNA‐based molecular analysis. Wolf spiders of the genus Pardosa and the cobweb spider, Theridion impressum L. Koch, were each collected in three winter OSR fields over a period of about 3 weeks. Pollen beetle densities as well as the occurrence of predators and alternative prey were monitored. In total, 13.8% of the collected Pardosa spp. tested positive for pollen beetle DNA in the PCR analyses, whereas 51.7%T. impressum were positive. The likelihood of detecting pollen beetle DNA in the gut contents of both spider groups was positively related to pollen beetle larval density. The implications of these results for conservation biological control and future studies of food webs in OSR are discussed.     

Habitat fragmentation determines diversity of annual plant communities at landscape and fine spatial scales

The aim of this study was to disentangle the effects of landscape configuration (i.e., fragment area, connectivity, and proximity to a busy highway) on the assembly of annual plant communities at different spatial scales. Our main hypothesis was that larger and more connected fragments would have higher species densities per plot and this may result in differences in turnover and nestedness patterns at the fine spatial scales where plants interact. Specifically, since Mediterranean annuals are known to form strong competitive hierarchies, we expected to find a nested pattern of beta diversity due to sequential species loss. The study area was a fragmented gypsum habitat in central Spain with a semiarid climate where two fragmentation drivers coexist: agricultural practices and a roadway. Larger fragments had higher species densities per plot (20 × 20 m). Nevertheless, we detected no effect on the species assembly at fine spatial scales (30 × 30 cm). However, when the fragment connectivity was high the species that appeared in poor quadrats (30 × 30 cm) comprised a subset of the species in rich quadrats. These results agree well with the establishment of strong competitive hierarchies among annual species. The distance to the highway influenced the identity of the species established in the community (i.e., species composition) at fine spatial scales, but we detected no effect on species turnover, nestedness, or species densities. The main conclusion of our study is that the effects of habitat fragmentation extend beyond the landscape scale and they determine the spatial assembly at fine spatial scales.     

Non-random dispersal by ants: Long-term field data versus model predictions of population spread of a forest herb

Myrmecochory, i.e. dispersal of seeds by ants towards and around their nests, plays an important role in temperate forests. Yet hardly any study has examined plant population spread over several years and the underlying joint contribution of a hierarchy of dispersal modes and plant demography. We used a seed-sowing approach with three replicates to examine colonization patterns of Melampyrum pratense, an annual myrmecochorous herb, in a mixed Scots pine forest in north-eastern Germany. Using a spatially explicit individual-based (SEIB) model population patterns over 4 years were explained by short-distance transport of seeds by small ant species with high nest densities, resulting in random spread. However, plant distributions in the field after another 4 years were clearly deviating from model predictions. Mean annual spread rate increased from 0.9 to 5.1 m per year, with a clear inhomogeneous component. Obviously, after a lag-phase of several years, non-random seed dispersal by large red wood ants (Formica rufa) was determining the species’ spread, thus resulting in stratified dispersal due to interactions with different-sized ant species. Hypotheses on stratified dispersal, on dispersal lag, and on non-random dispersal were verified using an extended SEIB model, by comparison of model outputs with field patterns (individual numbers, population areas, and maximum distances). Dispersal towards red wood ant nests together with seed loss during transport and redistribution around nests were essential features of the model extension. The observed lag-phase in the initiation of non-random, medium-distance transport was probably due to a change of ant behaviour towards a new food source of increasing importance, being a meaningful example for a lag-phase in local plant species invasion. The results demonstrate that field studies should check model predictions wherever possible. Future research will show whether or not the M. pratense–ant system is representative for migration patterns of similar animal dispersal systems after having crossed range edges by long-distance dispersal events.     

Spider assemblages in winter oilseed rape affected by landscape and site factors

Spiders are an abundant and diverse group of generalist predators in arable fields. Knowledge on what landscape and site factors affect this group can be valuable for efforts to reduce biodiversity loss in agricultural landscapes and can have implications for natural pest control. We investigated the impact of landscape and site factors on epigeic spiders in 29 winter oilseed rape fields (Brassica napus, OSR) embedded in differently structured landscapes in an agricultural region east of Vienna (Austria). Landscape factors included proportions of non‐crop areas, woody areas and fallows, lengths of road‐side strips and hedges, and landscape diversity at different spatial scales (r=250–2000 m). Site factors included OSR stand density, soil index, soil cultivation intensity, nitrogen fertilisation level, OSR vegetation cover in late autumn, and insecticide applications. Data were analysed using regression, ordination, and variation partitioning. Different characteristics of spider assemblages responded to different landscape factors at different spatial scales. Observed species richness showed the strongest positive reponse to proportions of woody areas at rather small scale (radius 500 m), but the relation remained significant up to the 1250 m radius. Standardised species richness was positively related to non‐crop area at the smallest scale (radius 250 m). Activity density was positively related to length of road‐side strips with maximum effects at large scale (radius 1750 m) and non‐crop area (radius 750 m). Site characteristics (stand density, insecticide applications, and late autumn ground cover) and landscape factors (woody areas and fallows at radius 500 m) were similarly important for explaining species composition. We interpret the scale‐dependency of relations as the result of differences in dispersal power of the studied spider species. These results demonstrate the important, scale dependent influence of natural and semi‐natural habitats on spider assemblages in arable fields.     

Effects of intraspecific and community density on the lifetime fecundity of long-lived shrubs

Intra- and interspecific density dependence has profound consequences for plant population and community dynamics. In long-lived plants, however, lifetime patterns and mechanisms of density dependence are difficult to study. Here, we examine effects of intraspecific and community density on the lifetime fecundity of two long-lived shrub species from South African Fynbos: Protea repens (animal-pollinated, hermaphroditic) and Leucadendron rubrum (wind-pollinated, dioecious). Both species are serotinous, retaining seeds in cones until fire kills the mother plant. We measured lifetime fecundity as the product of cone number, proportion of cones that are not damaged by predation and seed set (fertile seeds per intact cone). Intraspecific and community densities were quantified by counting individuals of target species and all Proteaceae in small- and large-scale neighbourhoods (10 m and 50 m radius) around each focal individual. Additionally, we determined the age and size of focal individuals. We found that lifetime fecundity of the wind-pollinated L. rubrum is density independent. In contrast, the lifetime fecundity of the animal-pollinated P. repens increases with large-scale intraspecific density and shows a hump-shaped relationship to large-scale community density. Community density has a hump-shaped effect on seed set (probably through partial absence of generalized pollinators at low and competition for pollinators at high densities) and negatively affects cone number per individual. For both species, plant age decreases seed set while increasing lifetime fecundity. The qualitative differences in the density dependence of lifetime fecundity may arise from differences between animal and wind pollination. In particular, interactions with generalized animal pollinators may cause community-level Allee effects with profound consequences for the future dynamics of long-lived plant populations and communities.