Population genetics of orchid bees in a fragmented tropical landscape

The prospects for persistence of bees living in fragmented landscapes is a topic of considerable interest due to bees’ importance as pollinators of agricultural crops and wild plants, coupled with the ubiquity of native habitat loss and evidence that bees may be declining worldwide. Population persistence in fragmented areas depends on dispersal potential and maintenance of gene flow among fragments of habitat. Here we used population genetic techniques to characterize, for two equally abundant orchid bee species that differ in their physiology and ecology, levels of genetic differentiation among fragments of tropical forest in southeastern Costa Rica in a ~200 km² landscape. We measured population differentiation with ϕ_PT (an analogue to the traditional summary statistic Fst), as well as two measures that may more accurately reflect the level of differentiation when highly variable loci are used: G’st and D_est. We also calculated pairwise genetic distances among individuals and conducted Mantel tests to test the correlation of genetic and geographic distance, for each species. We found strong differences in genetic structure between the species. Contrary to our expectations, each measure of genetic structure revealed that the larger-bodied species, Eulaema bombiformis, had higher levels of differentiation than the smaller species, Euglossa championi. Furthermore, for Eulaema bombiformis there was a significant positive correlation of genetic and geographic distance while for Euglossa championi there was no significant positive correlation. Our results demonstrate that bee species can have strikingly different levels of gene flow in fragmented habitats, and that body size may not always act as a useful proxy for dispersal, even in closely related taxa.     

Drought effects on seedling survival in a tropical moist forest

The amount and seasonality of rainfall varies strongly in the tropics, and plant species abundance, distribution and diversity are correlated with rainfall. Drought periods leading to plant stress occur not only in dry forests, but also in moist and even wet forests. We quantified experimentally the effect of drought on survival of first year seedlings of 28 co-occurring tropical woody plant species in the understory of a tropical moist forest. The seedlings were transplanted to plots and subjected to a drought and an irrigation treatment for 22 weeks during the dry season. Drought effects on mortality and wilting behavior varied greatly among species, so that relative survival in the dry treatment ranged from 0% to about 100% of that in the irrigated treatment. Drought stress was the main factor in mortality, causing about 90% (median) of the total mortality observed in the dry treatment. In almost half of the species, the difference in survival between treatments was not significant even after 22 weeks, implying that many of the species are well adapted to drought in this forest. Relative drought survival was significantly higher in species associated with dry habitats than in those associated with wet habitats, and in species with higher abundance on the dry side of the Isthmus of Panama, than in those more abundant on the wet side. These data show that differential species survival in response to drought, combined with variation in soil moisture availability, may be important for species distribution at the local and regional scale in many tropical forests.     

The effects of cattle grazing on plant-pollinator communities in a fragmented Mediterranean landscape

The main aims of this study were to assess grazing impacts on bee communities in fragmented mediterranean shrubland (phrygana) and woodland habitats that also experience frequent wildfires, and to explain the mechanisms by which these impacts occur. Fieldwork was carried out in 1999 and 2000 on Mount Carmel, in northern Israel, a known hot-spot for bee diversity. Habitats with a range of post-burn ages and varying intensities of cattle grazing were surveyed by transect recording, grazing levels, and the diversity and abundance of both flowers and bees were measured. The species richness of both bees and flowers were highest at moderate to high grazing intensities, and path-analysis indicated that the effects of both grazing and fire on bee diversity were mediated mainly through changes in flower diversity, herb flowers being more important than shrubs. The abundance of bees increased with intensified grazing pressure even at the highest levels surveyed. Surprisingly though, changes in bee abundance at high grazing levels were not caused directly by changes in flower cover. The variation in bee abundance may have been due to higher numbers of solitary bees from the family Halictidae in grazed sites, where compacted ground (nesting resource) and composites (forage resource) were abundant. The effects of grazing on plants were clearest in the intermediate-aged sites, where cattle inhibited the growth of some of the dominant shrubs, creating or maintaining more open patches where light-demanding herbs could grow, thus allowing a diverse flora to develop. Overall, bee communities benefit from a relatively high level of grazing in phrygana. Although bee and flower diversity may decrease under very heavy grazing, the present levels of grazing on Mount Carmel appear to have only beneficial effects on the bee community.     

Regeneration response and seedling bank dynamics on a Dendroctonus rufipennis‐killed Picea engelmannii landscape

Question: How does regeneration response to a host‐specific, high‐severity, infrequent Dendroctonus rufipennis outbreak differ from our conceptualization of high‐severity, infrequent/low‐severity, frequent disturbance regimes in Picea engelmannii‐Abies lasiocarpa communities? Location: Southern Utah, USA. Methods: One hundred and seven plots across a high‐elevation P. engelmannii forest were sampled to reconstruct pre‐outbreak overstory and seedling bank densities, and calculate their associated metrics of diversity. Decade of establishment by seedling bank trees indicated “chronic” and “pulse” regenerators. Results: The post‐outbreak overstory and seedling bank were dominated by A. lasiocarpa. Although Pinus flexilis, Pinus ponderosa, Picea pungens, and Psuedotsuga menziesii were present in the overstory, they were virtually absent in the seedling bank. Seedling bank recruitment of A. lasiocarpa and P. engelmannii has been occurring chronically for at least the last ～205 and ～152 years, respectively. A pulse response of seedling bank Populus tremuloides was apparent; however, results were complicated by intense ungulate browsing. Conclusions: Despite some similarities to the high‐severity, infrequent/low‐severity, frequent conceptualization of regeneration response to disturbance, the high‐severity D. rufipennis outbreak is best described by explicitly considering host specificity and severity. Although, the outbreak simultaneously promoted both a pulse of P. tremuloides and a release of chronically regenerated A. lasiocarpa, the P. tremuloides response was generally masked by ungulate browsing, and the regeneration response came overwhelmingly from the A. lasiocarpa seedling bank. In this landscape, once dominated by P. engelmannii, the chronically regenerating seedling bank, typically thought to take advantage of canopy gaps associated with low‐severity disturbances, is poised to dominate forest reorganization in response to the host‐specific outbreak.     

Pollen dispersal and genetic structure of the tropical tree Dipteryx panamensis in a fragmented Costa Rican landscape

In the face of widespread deforestation, the conservation of rainforest trees relies increasingly on their ability to maintain reproductive processes in fragmented landscapes. Here, we analysed nine microsatellite loci for 218 adults and 325 progeny of the tree Dipteryx panamensis in Costa Rica. Pollen dispersal distances, genetic diversity, genetic structure and spatial autocorrelation were determined for populations in four habitats: continuous forest, forest fragments, pastures adjacent to fragments and isolated pastures. We predicted longer but less frequent pollen movements among increasingly isolated trees. This pattern would lead to lower outcrossing rates for pasture trees, as well as lower genetic diversity and increased structure and spatial autocorrelation among their progeny. Results generally followed these expectations, with the shortest pollen dispersal among continuous forest trees (240 m), moderate distances for fragment (343 m) and adjacent pasture (317 m) populations, and distances of up to 2.3 km in isolated pastures (mean: 557 m). Variance around pollen dispersal estimates also increased with fragmentation, suggesting altered pollination conditions. Outcrossing rates were lower for pasture trees and we found greater spatial autocorrelation and genetic structure among their progeny, as well as a trend towards lower heterozygosity. Paternal reproductive dominance, the pollen contributions from individual fathers, did not vary among habitats, but we did document asymmetric pollen flow between pasture and adjacent fragment populations. We conclude that long-distance pollen dispersal helps maintain gene flow for D. panamensis in this fragmented landscape, but pasture and isolated pasture populations are still at risk of long-term genetic erosion.     

Frankia inoculation,soil biota,and host tissue amendment influence Casuarina nodulation capacity of a tropical soil

The effects of soil biota, Frankia inoculation and tissue amendment on nodulation capacity of a soil was investigated in a factorial study using bulked soil from beneath a Casuarina cunninghamiana tree and bioassays with C. cunninghamiana seedlings as capture plants. Nodulation capacities were determined from soils incubated in sterile jars at 21 °C for 1, 7, and 28 days, after receiving all combinations of the following treatments: ± steam pasteurization, ± inoculation with Frankia isolate CjI82001, and ± amendment with different concentrations of Casuarina cladode extracts. Soil respiration within sealed containers was determined periodically during the incubation period as a measure of overall microbial activity. Soil respiration, and thus overall microbial activity, was positively correlated with increasing concentrations of Casuarina cladode extracts. The nodulation capacity of soils inoculated with Frankia strain Cj82001 decreased over time, while those of unpasteurized soils without inoculation either increased or remained unaffected. The mean nodulation capacity of unpasteurized soil inoculated with Frankia CjI82001 was two to three times greater than the sum of values for unpasteurized and inoculated pasteurized soils. Our results suggest a positive synergism between soil biota as a whole and Frankia inoculum with respect to host infection.     

Genetic diversity in adult and seedling populations of <Emphasis Type="Italic">Primula vulgaris</Emphasis> in a fragmented agricultural landscape

Habitat fragmentation is known to generally reduce the size of plant populations and increase their isolation, leading to genetic erosion and increased between-population genetic differentiation. In Flanders (northern Belgium) Primula vulgaris is very rare and declining. Populations have incurred strong fragmentation for the last decades and are now restricted to a few highly fragmented areas in an intensively used agricultural landscape. Previous studies showed that small populations of this long-lived perennial herb still maintained high levels of genetic variation and low genetic differentiation. This pattern can either indicate recent gene flow or represent historical variation. Therefore, we used polymorphic microsatellite loci to investigate genetic variation and structure in adult (which may still reflect historical variation) and seedling (recent generation, thus affected by current processes) life stages. The recent generation (seedlings) showed a significant loss of observed heterozygosity (H _o) together with lower expected heterozygosity (H _e), a trend for higher inbreeding levels (F _IS) and higher differentiation (F _ST) between populations compared to the adult generation. This might result from (1) a reduction in effective population size, (2) higher inbreeding levels in the seedlings, (3) a higher survival of heterozygotes over time due to a higher fitness of heterozygotes (heterosis) and/or a lower fitness of homozygotes (inbreeding depression), (4) overlapping generations in the adult life stage, or (5) a lack of establishment of new (inbred) adults from seedlings due to degraded habitat conditions. Combining restoration of both habitat quality and gene flow between populations may be indispensable to ensure a sustainable conservation of fragmented populations.     

Ant-nest soil and seedling growth in a neotropical ant-dispersed herb

Summary A major hypothesis concerning the benefits of myrmecochory, seed dispersal by ants, to plants is that ant nests are nutrient-enriched microsites that are beneficial to seedling growth. We experimentally test this hypothesis for a neotropical myrmecochore, Calathea ovandensis, asking two questions: 1) is soil of nests of a seed-dispersing ant chemically or structurally distinct from surrounding soils, and 2) do seedlings grow better in soil collected from ant nests than in randomly collected soil? We found that although ant-nest soil was significantly enriched in nitrate-nitrogen, magnesium, iron, manganese, cadmium and percent organic matter compared to randomly collected soil, seedling growth was not significantly improved by ant-nest soil.     

Effect of soil biota on growth and allocation by Eucalyptus microcarpa

We examined growth of Eucalyptus microcarpa seedlings in soil collected from four sites in southeastern Australia, in which retired pasture land has been revegetated with mixed plantings of Eucalyptus and Acacia species. Revegetation of farm land in southeastern Australia is an area of major investment. The focus of the study was to examine the influence of soil biota on seedling growth and its possible interaction with soil enrichment from a legume (Acacia) and decomposition rates. We used a soil freezing treatment (−80°C for 3 days) to retard the soil biota, with the expectation that invertebrates in particular would be killed. Soil freezing did not cause a nutrient pulse, but did reduce the level of ammonium in soil. Nitrate levels increased with time in pots, regardless of the soil treatment. Decomposition rates measured using cellulose substrate were significantly reduced by the freeze treatment, but only for approximately 90 days. Eucalyptus microcarpa seedlings grown in freeze-treated soil were approximately 40% smaller (total biomass), had marginally lower LAR (leaf area ratio), and significantly lower LMA (leaf mass per area). Low LMA indicates that leaves are either thinner in cross-section or less dense. We hypothesise that both the poor growth of seedlings and production of less robust leaves are consequences of reduced availability of soil nutrients due to the diminished soil biota after freeze treatment. Litter under Acacia was richer in nitrogen than litter under Eucalyptus but there was no difference in nitrogen content of soil, and consequently no soil source effects on plant growth or decomposition. We suggest that variation in the soil biota has the potential to greatly enhance or hinder the success of revegetation on retired agricultural land, but enrichment of soil by decomposition of nitrogen rich litter in these sites requires longer than the 8–15 years since they were revegetated.     

Influence of habitat,litter type,and soil invertebrates on leaf-litter decomposition in a fragmented Amazonian landscape

Amazonian forest fragments and second-growth forests often differ substantially from undisturbed forests in their microclimate, plant-species composition, and soil fauna. To determine if these changes could affect litter decomposition, we quantified the mass loss of two contrasting leaf-litter mixtures, in the presence or absence of soil macroinvertebrates, and in three forest habitats. Leaf-litter decomposition rates in second-growth forests (>10 years old) and in fragment edges (<100 m from the edge) did not differ from that in the forest interior (>250 m from the edges of primary forests). In all three habitats, experimental exclusion of soil invertebrates resulted in slower decomposition rates. Faunal-exclosure effects were stronger for litter of the primary forest, composed mostly of leaves of old-growth trees, than for litter of second-growth forests, which was dominated by leaves of successional species. The latter had a significantly lower initial concentration of N, higher C:N and lignin:N ratios, and decomposed at a slower rate than did litter from forest interiors. Our results indicate that land-cover changes in Amazonia affect decomposition mainly through changes in plant species composition, which in turn affect litter quality. Similar effects may occur on fragment edges, particularly on very disturbed edges, where successional trees become dominant. The drier microclimatic conditions in fragment edges and second-growth forests (>10 years old) did not appear to inhibit decomposition. Finally, although soil invertebrates play a key role in leaf-litter decomposition, we found no evidence that differences in the abundance, species richness, or species composition of invertebrates between disturbed and undisturbed forests significantly altered decomposition rates.     

The effects of mycorrhizal roots on litter decomposition, soil biota, and nutrients in a spodosolic soil

We studied the effects of mycorrhizal pitch pine (Pinus rigida) roots on litter decomposition, microbial biomass, nematode abundance and inorganic nutrients in the E horizon material of a spodosolic soil, using field microcosms created in a regenerating pitch pine stand in the New Jersey Pinelands. Pine roots stimulated litter decomposition by 18.7% by the end of the 29 month study. Both mass loss and N and P release from the litter were always higher in the presence of roots than in their absence. Nutrient concentrations in decomposing litter were similar, however, in the presence and absence of roots, which suggests that the roots present in the with-root treatment did not withdraw nutrients directly from the litter. The soil was slightly drier in the presence of roots, but there was no discernible effect on soil microbial biomass. The effects of roots on soil extractable inorganic nutrients were inconsistent. Roots, however, were consistently associated with higher numbers of soil nematodes. These results suggest that, in soils with low total C and N contents, roots stimulate greater activity of the soil biota, which contribute, in turn, to faster litter decomposition and nutrient release.Contribution No. 95-22 from the Institute of Marine and Coastal Sciences.Contribution No. 95-22 from the Institute of Marine and Coastal Sciences.     

Osmotic effects on radial growth rate and specific growth rate of three soil fungi

The radial growth rate on osmotically adjusted agar medium and the relative specific growth rate in osmotically adjusted liquid medium were determined for Rhizoctonia solani, Pythium ultimum, and Verticillium dahliae. On basal medium, an isolate of P. ultimum and R. solani had similar radial growth rates of 0.52 and 0.47 mm/h, respectively, whereas V. dahliae grew at a rate of 0.08 mm/h. Radial growth rate was reduced 50% at osmotic potentials of -16, -27, and -32 bars for P. ultimum, R. solani, and V. dahliae, respectively. No growth occurred at -32 bars for P. ultimum, -56.2 bars for R. solani, and -100 bars for V. dahlia. Specific growth rates in liquid culture were 0.011 h-1 for P. ultimum, 0.008 h-1 for V. dahliae, and 0.026 h-1 for R. solani. Ratios of radial growth rate (Kr) to specific growth rate (alphas) were computed for each fungus growing at different osmotic potentials. There was not a constant relationship between Kr on agar and alphas in liquid medium, e.g., Kr/alphas ratios varied from 8-41% from a mean ratio for a particular species. The results indicated that radial growth rate on osmotic agar was not useful as a measure of relative specific growth rate of a fungus in osmotically adjusted liquid medium.     

Effect of autoclave sterilization of a tropical andept on seed germination and seedling growth

Summary Steam sterilization of a Typic Dystrandept in Costa Rica resulted in a six-fold increase in extractable Mn, to levels often considered toxic. Seeds of eight species, comprised of six successional taxa and two cultivars (soybean,Glycine max and raddish,Raphanus sativus) were planted in the sterilized soil and in unsterilized soil after delays of 1, 8, 15, and 28 days. Germination and mortality were not different in the two soils, indicating that steamsterilized soil can safely be used in seed traps. Six species (including both cultivars) grew better in unsterilized soil, but two of the native taxa (Phytolacca rivinoides andBocconia frutescens) grew significantly faster in sterilized soil.     

Nonlinearity of effects of invasive ecosystem engineers on abiotic soil properties and soil biota

Invasions of non‐indigenous species into natural communities are currently rated as one of the most important threats to biodiversity. Particularly exotic ecosystem engineers such as earthworms potentially have profound impacts on community assembly and functioning. We investigated the impact of invasion by the lumbricid earthworms into an aspen forest of the Canadian Rocky Mountains on soil organic matter, microorganisms and microarthropod communities. Building on the results of previous studies in this forest, we expected positive effects of Lumbricus terrestris middens and negative effects of Octolasion tyrtaeum on soil biota (increase and decrease in soil nutrient concentrations, microbial parameters and soil microarthropod density and diversity, respectively). Further, we expected that earthworm effects change with time. Combined results of previous and the present study suggest a wavelike colonization pattern for Dendrobaena octaedra and O. tyrtaeum and that indeed the impact of earthworms on soil biota changed with time, likely due to changes in earthworm density. Unexpectedly, L. terrestris middens neither affected soil abiotic nor soil biotic properties. By contrast and in contrast to our hypothesis, carbon and nitrogen concentration and C‐to‐N ratio in deeper soil layers increased in presence of O. tyrtaeum, thereby likely enhancing nutrient availability for soil microorganisms and microarthropods. Even though the density of this endogeic species was rather low, presence of O. tyrtaeum resulted in increased densities of a number of microarthropod taxa and increased microarthropod diversity. The results suggest that at low density, invasive ecosystem engineers, such as O. tyrtaeum, cause disturbances of intermediate strength thereby beneficially affecting soil microorganisms and most microarthropods. This contrasts earlier effects during the wavelike invasion of O. tyrtaeum into the aspen forest when densities of O. tyrtaeum were high resulting in generally detrimental effects on soil biota. The results emphasize the nonlinearity of earthworm effects on abiotic and biotic soil properties and call for further long‐term investigations.     

Trait-based community assembly of understory palms along a soil nutrient gradient in a lower montane tropical forest

Two opposing niche processes have been shown to shape the relationship between ecological traits and species distribution patterns: habitat filtering and competitive exclusion. Habitat filtering is expected to select for similar traits among coexisting species that share similar habitat conditions, whereas competitive exclusion is expected to limit the ecological similarity of coexisting species leading to trait differentiation. Here, we explore how functional traits vary among 19 understory palm species that differ in their distribution across a gradient of soil resource availability in lower montane forest in western Panama. We found evidence that habitat filtering influences species distribution patterns and shifts community-wide and intraspecific trait values. Differences in trait values among sites were more strongly related to soil nutrient availability than to variation in light or rainfall. Soil nutrient availability explained a significant amount of variation in site mean trait values for 4 of 15 functional traits. Site mean values of leaf nitrogen and phosphorus increased 37 and 64%, respectively, leaf carbon:nitrogen decreased 38%, and specific leaf area increased 29% with increasing soil nutrient availability. For Geonoma cuneata, the only species occurring at all sites, leaf phosphorus increased 34% and nitrogen:phosphorus decreased 42% with increasing soil nutrients. In addition to among-site variation, most morphological and leaf nutrient traits differed among coexisting species within sites, suggesting these traits may be important for niche differentiation. Hence, a combination of habitat filtering due to turnover in species composition and intraspecific variation along a soil nutrient gradient and site-specific niche differentiation among co-occurring species influences understory palm community structure in this lower montane forest.