Nurse plants vs. nurse objects: effects of woody plants and rocky cavities on the recruitment of the Pilosocereus leucocephalus columnar cactus

BACKGROUND AND AIMS: Most studies on cactus recruitment have focused on the role of woody plants as seedling facilitators. Although the spatial association of cacti with objects had been described, the mechanisms underlying this association remain unknown. The aims of this study were to identify which mechanisms facilitate the establishment of a columnar cactus under the shade and protection of objects and to compare these mechanisms with those involved in plant-plant facilitation. METHODS: Three split-split-plot field experiments were conducted to compare the effects of two microhabitats (inside rocky cavities and beneath plant canopies) on seed removal, germination, seedling survivorship and dry weight. Flat, open spaces were used as the control. For each microhabitat, the effect of seed or seedling protection and substrate limitation were explored; aboveground microclimate and some soil properties were also characterized. KEY RESULTS: The permanence of superficial seeds was greater inside rocky cavities than beneath woody plant canopies or on flat, open areas. Germination was similar in cavities and beneath plant canopies, but significantly higher than on flat, open areas. Seedling survivorship was greater beneath plant canopies than inside cavities or on flat, open spaces. CONCLUSIONS: The mechanisms of plant facilitation are different from those of object facilitation. There are seed-seedling conflicts involved in the recruitment of P. leucocephalus: nurse plants favour mainly seedling survivorship by providing a suitable microenvironment, while nurse objects mainly favour seed permanence, by protecting them from predators.     

Regeneration of Cyperaceae, with particular reference to seed ecology and seed banks

Cyperaceae (sedges) are an important component of many ecosystems. To understand better their regeneration, we examined seed ecology, including dispersal, seed characteristics, and germination behavior that relate to seed bank development and persistence. We also evaluated sedge seed banks from 104 studies, representing a wide array of habitats. Sedge seed bank development and persistence were associated with germination and dormancy traits, namely: dormancy level, seasonal dormancy patterns, and requirement for light, alternating temperatures, and aerobic conditions. Interplay of traits appears to have resulted in low-risk germination strategies adapted to exploit infrequent occurrence of gaps and facilitate formation of persistent seed banks. A variety of dispersal modes and morphological adaptations occurred, but many species had no apparent specialized structures. The main dispersal vectors were water and then animals. About 216 species, in 21 genera, were recorded in the seed bank survey. High densities (>50,000 m⁻²) occurred occasionally in wet habitats, but generally values were low (<500 m⁻² in 70% of entries). Species richness was also generally low (mean 4.8 species study⁻¹), but ranged from 10 to 33 species in certain wetlands. Our studies showed varied reproductive strategies within habitats, persistence, and ability of many species to colonize disturbed habitats. Overall, seed banks tended to be persistent (>1 year). Maximum longevity ranged between 10 and 295 years, but for certain species viability was lost in <3 years. Seeds of many sedges occurred in deeper soil layers to depths >1 m. Seed production, low in rhizomatous species, ranged between 0 and 345,000 seeds m⁻² year⁻¹. Amphicarpy or pseudo-viviparous plantlets occurred in limited numbers of species. The relation between seed production, seed rain, and seed bank is largely obscure and awaits further investigation. For successful restoration and species conservation projects, seed banks (or a source of seeds) are necessary, combined with suitable germination and establishment conditions. Future seed bank studies are considered.     

Vertical structure of seed banks and the impact of depth of burial on recruitment in two temporary marshes

The structure of the seed bank (including Chara oospores), in relation to depth within the sediment and disturbance, was studied in two Rhône delta temporary marshes for two years. The seeds of all species were concentrated in the top 2 cm of sediment with very low numbers beeing found below 4 cm. When an exclosure eliminated disturbances of the sediment by animals, the vertical repartition of seeds at site 2 was more pronounced than outside the exclosure.In experiment 1, the emergence capacity of seeds from different depths and buried under layers of sterile equivalent to those in the field was measured. Depending of the species, 22 to 98% of the seeds germinated from unburied seeds in the top 2 cm. Only 1% of the oospores of Chara (from site 2) at 2 to 4 cm depth in the sediment emerged.In experiment 2, surface seed bank samples were placed under 0, 2 or 4 cm sterile sediment depth. The samples contained numerous recent seeds and the emergence percentage reached 41% (for Ruppia maritima). Only the seeds of Zannichellia spp failed to germinate from a depth of 2 cm or more. The emergence percentage from 2 cm depth or more was always lower than at the surface. These experiments showed that both burial and ageing of seeds decrease germination capacity.The majority of the active seeds located at the surface germinate when the marsh is flooded. Seeds located between 2 and 4 cm can be brought back to the surface by disturbances and play the role of a reserve involved in maintenance of populations that go without seed production for one or some years.     

Microsite and elevation zone effects on seed pilferage,germination, and seedling survival during early whitebark pine recruitment

Tree recruitment is a spatially structured process that may undergo change over time because of variation in postdispersal processes. We examined seed pilferage, seed germination, and seedling survival in whitebark pine to determine whether 1) microsite type alters the initial spatial pattern of seed caches, 2) higher abiotic stress (i.e. higher elevations) exacerbates spatial distribution changes, and 3) these postdispersal processes are spatially clustered. At two study areas, we created a seed distribution pattern by burying seed caches in microsite types frequently used by whitebark pine's avian seed disperser (Clark's nutcracker) in upper subalpine forest and at treeline, the latter characterized by high abiotic environmental stress. We monitored caches for two years for pilferage, germination, and seedling survival. Odds of pilferage (both study areas), germination (northern study area), and survival (southern study area) were higher at treeline relative to subalpine forest. At the southern study area, we found higher odds of 1) pilferage near rocks and trees relative to no object in subalpine forest, 2) germination near rocks relative to trees within both elevation zones, and 3) seedling survival near rocks and trees relative to no object at treeline. No microsite effects were detected at the northern study area. Findings indicated that the microsite distribution of seed caches changes with seed/seedling stage. Higher odds of seedling survival near rocks and trees were observed at treeline, suggesting abiotic stress may limit safe site availability, thereby shifting the spatial distribution toward protective microsites. Higher odds of pilferage at treeline, however, suggest rodents may limit treeline recruitment. Further, odds of pilferage were higher near rocks and trees relative to no object in subalpine forest but did not differ among microsites at treeline, suggesting pilferage can modulate the spatial structure of regeneration, a finding supported by limited clustering of postdispersal processes.     

Effect of seed predation on seed bank size and seedling recruitment of bush lupine (Lupinus arboreus)

Whether seed consumers affect plant establishment is an important unresolved question in plant population biology. Seed consumption is ubiquitous; at issue is whether seedling recruitment is limited by safe-sites or seeds. If most seeds inhabit sites unsuitable for germination, post-dispersal seed consumption primarily removes seeds that would otherwise never contribute to the population and granivory has minimal impacts on plant abundance. Alternatively, if most seeds ultimately germinate before they lose viability, there is greater potential for seed consumption to affect plant recruitment. Of the many studies on seed consumption, few ask how seed loss affects seedling recruitment for species with long-lived seed banks. We examined post-dispersal seed predation and seedling emergence in bush lupine (Lupinus arboreus), a woody leguminous shrub of coastal grasslands and dunes in California. We followed the fate of seeds in paired experimental seed plots that were either protected or exposed to rodent granivores in grassland and dune habitats. Significantly more seeds were removed by rodents in dunes than grasslands. In dunes, where rodent granivory was greatest (65% and 86% of seeds removed from plots by rodents in two successive years), there is a sparse seed bank (6.6 seeds m⁻²), and granivory significantly reduced seedling emergence (in the same two years, 18% and 19.4% fewer seedlings emerged from exposed versus protected plots), suggesting seed rather than safe-site limited seedling recruitment. In contrast, rodents removed an average of 6% and 56% of seeds from grassland plots during the same two years, and the grassland seed bank is 43-fold that of the dunes (288 seeds m⁻²). Even high seed consumption in the second year of the study only marginally influenced recruitment because seeds that escaped predation remained dormant. Burial of seeds in both habitats significantly reduced the percentage of seeds removed by rodents. Results suggest that granivores exert strong but habitat-dependent effects on lupine seed survival and seedling emergence. Received: 24 October 1996 / Accepted: 4 February 1997     

Relative importance of perch and facilitative effects on nucleation in tropical woodland in Malawi

Individual trees in open vegetation such as woodlands can act as “nuclei” for the colonization of forest tree species, which consequently lead to the formation of forest patches. This phenomenon is known as nucleation. The mechanism of nucleation is generally attributed to two factors: trees provide perches for frugivores that increase seed deposition (perch effect), and tree crowns ameliorate environmental conditions, which improves seedling establishment (facilitative effect). Few studies have attempted to distinguish the relative importance of these two factors. In this study, I separated these two effects in a woodland in northern Malawi. I chose Ficus natalensis as a potential nuclei tree because large individuals of this species are commonly located at the center of forest patches within open woodland at the study site. I monitored several environmental variables, seedling survival, seedling composition, and seed rain at three microsites: under F. natalensis, under Brachystegia floribunda (a dominant woodland species), and in open sites. Both tree species provided similar favorable conditions for the establishment of forest species compared to open sites. Thus, the survival of forest tree seedlings under F. natalensis and B. floribunda was similar, and substantially higher than seedling survival in open sites. However, communities of naturally occurring seedlings differed significantly between F. natalensis and B. floribunda. These results indicate that the facilitative effect alone cannot explain the nucleation pattern. I attribute this result to the perch effect of F. natalensis because the forest seedling species recorded under F. natalensis reportedly have small, brightly colored diaspores, which are indicative of dispersal by birds. Seed deposition of forest species under F. natalensis was significantly higher than that under B. floribunda or in open sites. My findings reinforce the idea that trees will lead to nucleation when they enhance seed deposition and have a positive effect on the post-dispersal stage.     

The influence of herbivory and weather on the vital rates of two closely related cactus species

Herbivory has long been recognized as a significant driver of plant population dynamics, yet its effects along environmental gradients are unclear. Understanding how weather modulates plant–insect interactions can be particularly important for predicting the consequences of exotic insect invasions, and an explicit consideration of weather may help explain why the impact can vary greatly across space and time. We surveyed two native prickly pear cactus species (genus Opuntia) in the Florida panhandle, USA, and their specialist insect herbivores (the invasive South American cactus moth, Cactoblastis cactorum, and three native insect species) for five years across six sites. We used generalized linear mixed models to assess the impact of herbivory and weather on plant relative growth rate (RGR) and sexual reproduction, and we used Fisher's exact test to estimate the impact of herbivory on survival. Weather variables (precipitation and temperature) were consistently significant predictors of vital rate variation for both cactus species, in contrast to the limited and varied impacts of insect herbivory. Weather only significantly influenced the impact of herbivory on Opuntia humifusa fruit production. The relationships of RGR and fruit production with precipitation suggest that precipitation serves as a cue in determining the trade‐off in the allocation of resources to growth or fruit production. The presence of the native bug explained vital rate variation for both cactus species, whereas the invasive moth explained variation only for O. stricta. Despite the inconsistent effect of herbivory across vital rates and cactus species, almost half of O. stricta plants declined in size, and the invasive insect negatively affected RGR and fruit production. Given that fruit production was strongly size‐dependent, this suggests that O. stricta populations at the locations surveyed are transitioning to a size distribution of predominantly smaller sizes and with reduced sexual reproduction potential.     

Negative and positive interactions among plants: effects of competitors and litter on seedling emergence and growth of forest and grassland species

Living plant neighbours, but also their dead aboveground remains (i.e. litter), may individually exert negative or positive effects on plant recruitment. Although living plants and litter co‐occur in most ecosystems, few studies have addressed their combined effects, and conclusions are ambivalent. Therefore, we examined the response in terms of seedling emergence and growth of herbaceous grassland and forest species to different litter types and amounts and the presence of competitors. We conducted a pot experiment testing the effects of litter type (grass, oak), litter amount (low, medium, high) and interspecific competition (presence or absence of four Festuca arundinacea individuals) on seedling emergence and biomass of four congeneric pairs of hemicryptophytes from two habitat types (woodland, grassland). Interactions between litter and competition were weak. Litter presence increased competitor biomass. It also had positive effects on seedling emergence at low litter amounts and negative effects at high litter amounts, while competition had no effect on seedling emergence. Seedling biomass was negatively affected by the presence of competitors, and this effect was stronger in combination with high amounts of litter. Litter affected seedling emergence while competition determined the biomass of the emerged individuals, both affecting early stages of seedling recruitment. High litter accumulation also reduced seedling biomass, but this effect seemed to be additive to competitor effects. This suggests that live and dead plant mass can affect species recruitment in natural systems, but the mechanisms by which they operate and their timing differ.     

Effects of prohydrojasmon‐treated corn plants on attractiveness to parasitoids and the performance of their hosts

We investigated the effect of prohydrojasmon [propyl (1RS,2RS)‐(3‐oxo‐ 2‐pentylcyclopentyl) acetate] (PDJ) treatment of intact corn plants, on their attractiveness to the specialist endoparasitoid, Cotesia kariyai Watanabe (Hymenoptera: Braconidae), and on the performance of the common armyworm, Mythimna separata (Walker) (Lepidoptera: Noctuidae) under laboratory conditions. Attractiveness of C. kariyai to PDJ‐treated plants was studied in a wind tunnel, whereas performance of M. separata larvae was tested in plastic cages. The attractiveness of the treated plants increased with concentrations of PDJ increasing to 2 mm , which was equivalent to the attractiveness of host‐infested plants. PDJ‐treated corn plants emitted 16 volatile compounds (α‐pinene, β‐myrcene, (Z)‐3‐hexenyl acetate, limonene, (E)‐β‐ocimene, linalool, (E)‐4,8‐dimethyl‐1,3,7‐nonatriene, (+)‐cyclosativene, ylangene, (E)‐β‐farnesene, (E, E)‐4,8,12‐trimethyl‐1,3,7,11‐tridecatetraene, α‐bergamotene, γ‐cadinene, δ‐cadinene, α‐muulolene and nerolidol), most of which were observed in the headspace of host‐infested corn plants with some quantitative and qualitative differences. We also tested the effects of PDJ treatment on the performance of M. separata larvae. The survival rates of the larval and pupal stages were significantly lower at 2 mm level of PDJ. A significant decrease in weight at 6th stadium larvae was observed only at 2 mm level of PDJ. In contrast, PDJ treatment at all PDJ concentration levels caused significant reduction in weight of pupal stage as compared to control. These data suggested that PDJ, originally developed as a plant growth regulator, especially to induce coloring of fruits, has the potential to induce direct and indirect defenses in corn plants against common armyworm, M. separata.     

Tree growth variation in the tropical forest: understanding effects of temperature,rainfall and CO2

Tropical forest responses to climatic variability have important consequences for global carbon cycling, but are poorly understood. As empirical, correlative studies cannot disentangle the interactive effects of climatic variables on tree growth, we used a tree growth model (IBTREE) to unravel the climate effects on different physiological pathways and in turn on stem growth variation. We parameterized the model for canopy trees of Toona ciliata (Meliaceae) from a Thai monsoon forest and compared predicted and measured variation from a tree‐ring study over a 30‐year period. We used historical climatic variation of minimum and maximum day temperature, precipitation and carbon dioxide (CO₂) in different combinations to estimate the contribution of each climate factor in explaining the inter‐annual variation in stem growth. Running the model with only variation in maximum temperature and rainfall yielded stem growth patterns that explained almost 70% of the observed inter‐annual variation in stem growth. Our results show that maximum temperature had a strong negative effect on the stem growth by increasing respiration, reducing stomatal conductance and thus mitigating a higher transpiration demand, and – to a lesser extent – by directly reducing photosynthesis. Although stem growth was rather weakly sensitive to rain, stem growth variation responded strongly and positively to rainfall variation owing to the strong inter‐annual fluctuations in rainfall. Minimum temperature and atmospheric CO₂ concentration did not significantly contribute to explaining the inter‐annual variation in stem growth. Our innovative approach – combining a simulation model with historical data on tree‐ring growth and climate – allowed disentangling the effects of strongly correlated climate variables on growth through different physiological pathways. Similar studies on different species and in different forest types are needed to further improve our understanding of the sensitivity of tropical tree growth to climatic variability and change.     

Interactive effects of nitrate concentrations and carbon dioxide on the stoichiometry,biomass allocation and growth rate of submerged aquatic plants

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