Herbivory and seedling performance in a fragmented temperate forest of Chile

Forest fragmentation alters plant-animal interactions, including herbivory. Relying manipulative experiments, we test if the reduction in insect herbivory associated with forest fragmentation translates into increased seedling growth and survival of three tree common species (Aristotelia chilensis, Cryptocarya alba and Persea lingue) in forest fragments and continuous forests in coastal Maulino forest, central Chile. Furthermore, we test if after protecting seedlings from herbivorous insects, plant performance is increased regardless of forest fragmentation. Nursery grown seedlings were transplanted into four forest fragments and a continuous forest during 2002. Insects, important herbivores in this forest, were excluded from half the seedlings by repeated applications of insecticides. Compared to continuous forests, in forest fragments, herbivory was reduced in all three species, seedling growth was greater in A. chilensis and C. alba but not in P. lingue, and survivorship was unaffected by herbivory or fragmentation in all three species. Protecting seedlings from insects reduced herbivory in the continuous forest to similar levels attained in the forest fragments. No change in herbivory results from by protecting seedlings in forest fragments. These results confirm that insects are important herbivores in the Maulino forest and also support the hypothesis that fragmentation can have strong indirect effects on plant communities as mediated through trophic interactions.     

Exudation of carboxylates in Australian Proteaceae: chemical composition

Roots of a wide range of plant species exude carboxylates, such as citrate, into the rhizosphere. In the present study, seedlings of a range of Australian Banksia, Hakea and Dryandra species (Proteaceae) were assayed for their exudation of carboxylates. All of these species (Hakea prostrata, Hakea undulata, Hakea petiolaris, Hakea baxteri, Banksia grandis, Banksia prionotes, Banksia occidentalis and Dryandra sessilis) form cluster roots when grown in nutrient solution with a low phosphate concentration. Exudation of carboxylates was studied for cluster roots and non‐cluster roots separately, and for the entire root system. Cluster roots of these Proteaceae exuded malate, malonate, lactate, acetate, maleate, citrate, fumarate, cis‐ and trans‐aconitate. The relative contributions of each of these carboxylates differed between species. Malate, malonate, lactate, citrate and trans‐aconitate, however, were invariably present in large proportions of total carboxylate exudation. Non‐cluster roots of H. prostrata exuded a spectrum of carboxylates (mainly malonate, lactate and citrate), which differed somewhat from the exudation pattern of cluster roots (mainly malate, malonate, lactate and citrate). The rate of exudation for cluster roots of the seven species was approximately 1·6 nmol g^?1 FM s^?1, which is considerably higher than that reported for a variety of crop and native species that do or do not form cluster roots. Contrary to what occurs in the cluster roots of Lupinus albus, which release carboxylates accompanied by protons so that the rhizosphere is acidified, the present Proteaceae exude the carboxylates as anions without concomitant proton release. The role of carboxylates in the mobilization of phosphate and other nutrients from soil is discussed.     

Herbivory and plant competition reduce mountain beech seedling growth and establishment in New Zealand

The effect of herbivory and resource availability on the competitive ability of different plant species has been an area of intense debate amongst plant ecologists for at least two decades, but the interactive effects of herbivory and plant competition between woody and herbaceous plants are rarely studied and theory is poorly developed. This study used experimental manipulations on transplanted and naturally occurring mountain beech (Nothofagus solandri var. cliffortioides) seedlings to show the effects of deer browsing and competition from deer-induced, herbaceous turf communities on mountain beech regeneration in New Zealand. Differences in the species composition of these turfs had little effect on mountain beech seedling establishment, but turf removal increased seedling growth and survivorship, showing that competition with other plants had direct effects on mountain beech regeneration. Deer browsing reduced the establishment and growth of seedlings, but the size of this effect did not vary with light and nutrient availability. There was no immediate compositional response of turf communities following the removal of deer browsing. The addition of nutrients appeared to reduce the intensity of belowground competition (stem growth increased relative to root growth) and increase seedling mortality, but there was no effect of changing levels of light. These results showed simple and direct negative effects of deer browsing on mountain beech regeneration. Indirect negative effects on regeneration were caused by deer-induced turf communities. We found little evidence for interactive effects between herbivory, plant competition and the availability of light or nutrients on seedling regeneration, which suggests that these factors acted independently. Nomenclature: Beever et al. (1992); Parsons et al. (1995); Edgar and Connor (2000); and Brownsey and Smith-Dodsworth (2000). Raukaua simplex is described by Mitchell et al. (1997). Coprosma “taylorae” is referred to by Eagle (1986) and Halocarpus biformis, Phyllocladus alpinus, Podocarpus hallii and Podocarpus nivalis by Wilson and Galloway (1993).     

Variation in plant functional traits across and within four species of Western Australian Banksia (Proteaceae) along a natural climate gradient

Plant traits are fundamental components of the ecological strategies of plants, relating to how plants acquire and use resources. Their study provides insight into the dynamics of species geographical ranges in changing environments. Here, we assessed the variation in trait values at contrasting points along an environmental gradient to provide insight into the flexibility of species response to environmental heterogeneity. Firstly, we identified how commonly measured functional traits of four congeneric species (Banksia baxteri, B. coccinea, B. media and B. quercifolia) varied along a longitudinal gradient in the South Western Australian Floristic Region. This regional gradient provides significant variation in moisture, temperature and soil nutrients: soil nitrogen content decreases with declining rainfall and increasing temperature. We hypothesized that (i) the regional pattern in trait–environment associations across the species would match those observed on a global scale and (ii) that the direction and slopes of the within‐species relationships would be similar to those across species for each of the measured traits. Along the regional gradient we observed strong shifts in trait values, and cross‐species relationships followed the expected trend: specific leaf area was significantly lower, and leaf N_area and seed dry mass significantly higher, at the drier end of the rainfall gradient. However, traits within species were generally not well correlated with habitat factors: we found weak patterns among populations, either due to the small ecological gradient or perhaps because fine‐scale structuring among populations (at a micro‐evolutionary scale) was low due to high gene flow within species. Understanding how species traits shift as a result of climatic influences, both at the regional (across species) and local (within species) scale, provides insight into plant adaptation to the environment. Such studies have important applications for conservation biology and population management in the face of global change.     

Ecology and biogeography in 3D: The case of the Australian Proteaceae

The key biophysical pressures shaping the ecology and evolution of species can be broadly aggregated into three dimensions: environmental conditions, disturbance regimes and biotic interactions. The relative importance of each dimension varies over time and space, and in most cases multiple dimensions need to be addressed to adequately understand the habitat and functional traits of species at broad spatial and phylogenetic scales. However, it is currently common to consider only one or two selective pressures even when studying large clades. We illustrate the importance of the all‐inclusive multidimensional approach with reference to the large and iconic plant family, Proteaceae: we review life‐history traits related to these three dimensions for the 46 genera occurring in Australia and show that this family can be considered the product of a long history of harsh environments, recurrent fires and strong faunal interactions. Because most Proteaceae species occur in fire‐prone ecosystems and possess fire‐adaptive traits that are both ancient and essential for their survival, disturbance by fire is likely to explain much of this family's ecology, evolution and distribution. Approaches that only examine prevailing environmental variables may fail to identify the mechanisms that drive a taxon's biogeography; they need to consider the likely mechanisms of adaptation and accept or reject plausible alternative hypotheses as the evidence allows. As multidisciplinary teams that consider all aspects of a taxon's ecology are assembled, and databases and numerical tools become increasingly available, studies on the ecology, biogeography and diversity of organisms at broader spatial and phylogenetic scales will arrive at more realistic conclusions.     

For everything a season: Smoke-induced seed germination and seedling recruitment in a Western Australian Banksia woodland

Abstract The influence of factors associated with fire on seed germination of Australian native species is generally well documented, but examples involving the use of smoke as a fire analogue for ecological research remain limited. The role of season of treatment in the efficacy of smoke as a promotive germination agent was investigated over two growing seasons using natural soil stored seedbanks in Banksia woodland near Perth, Western Australia. Smoke was applied to unburnt sites in the autumn, winter and spring of 1994. Germinant emergence and seedling survival of 37 species representing 18 families was monitored in both unburnt sites and in adjacent, recently burnt sites until the second spring after treatment (October 1995). Recruitment from seed was found to be profoundly affected by the season in which dormancy breaking treatment had been applied. The promotive effect extended beyond the initial year of application. For the majority of the species investigated, application of smoke to unburnt sites in autumn promoted a significantly greater germination response than treatment in winter or spring. In only three cases (introduced annuals, the Fabaceae and Hibbenia amplexicaulis) did autumn smoke treatment not yield better germination than in summer-burnt counterparts. However, in almost half of the cases examined, proportions of seedlings surviving past their first summer after emergence in burnt areas were consistently greater than those in smoked or untreated sites. Most notably, no seedlings emerging during the spring of the first year of study survived into the following summer. Implications of the results with respect to future seed bank research and management of native vegetation are discussed.     

Variation in bradyspory and seedling recruitment without fire among populations of Banksia serrata (Proteaceae)

Abstract Long-term canopy storage of seeds in fruits (bradyspory) varies among species and is thought to be related to the fire regime experienced by the plant community. This study sought to quantify intraspecific variation in bradyspory among nine populations of Banksia serrata (Proteaceae) on the south coast of New South Wales. Marked differences were found between populations, ranging from more than 30% of follicles open at two coastal sites, to less than 5% of follicles open at two inland heath sites. Low levels of bradyspory occurred particularly in coastal sites which also had Banksia integrifolia, a non-bradysporous species, present. In addition to spontaneous follicle opening without fire, fall of infructes-cences to the ground after breaking of branches was observed to be a major contributor to seed release. At three sites where fire had not occurred for many years, the amount of seedling establishment since the last fire corresponded well with levels of bradyspory; there were no inter-fire recruits at the site with the strongest bradyspory, while at the site with the weakest bradyspory there were numerous inter-fire plants, some with lignotubers large enough to be able to tolerate the next fire. It is argued that populations of B. serrata exhibit variable patterns of recruitment, depending on the site characteristics, ranging from complete dependence on fire for population maintenance to bet-hedging (spreading potential recruitment between fire events and inter-fire periods). Variations in populations and reproductive characteristics among sites make the application of demographic models to management decisions habitat-specific.     

Geographic range size, seedling ecophysiology and phenotypic plasticity in Australian Acacia species

Aim The degree to which eco‐physiological traits critical to seedling establishment are related to differences in geographic range size among species is not well understood. Here, we first tested the idea that seedling eco‐physiological attributes associated with establishment differ between narrowly distributed and geographically widespread plant species. Secondly, we tested the notion that species occupying wide geographic ranges have greater phenotypic plasticity in response to the environment than contrasted species with more restricted distributions. Location Eastern Australia. Methods We compared five pairs of geographically restricted and widespread Acacia species grown under glasshouse conditions for differences in seedling relative growth rate and associated allocational, morphological and physiological traits. We then examined whether widespread species displayed greater phenotypic plasticity in these traits than narrowly distributed species in response to changes in soil water availability. Results Neither relative growth rate nor any measure of biomass accumulation or allocation differed significantly between seedlings of narrowly distributed and widespread species. In addition, the plasticity of biomass allocation was not greater in widespread species. However, the leaflets of widespread species had higher photosynthetic capacity and greater plasticity of water use efficiency than the leaflets of narrowly distributed species. Main conclusions We demonstrated fundamental differences in the physiology and plasticity of leaflets of widespread and narrowly distributed species. The greater plasticity of these seedling leaflet traits may allow widespread Acacia species to utilize a wider range of environmental conditions in relation to soil moisture than restricted Acacia species. However, we did not find corresponding differences in mean or plasticity of seedling growth and allocational traits. In general, we suggest that relationships between rarity and species traits are both context and taxon specific.     

Pollination disruption by European honeybees in the Australian bird-pollinated shrubGrevillea barklyana (Proteaceae)

European honeybees (Apis mellifera) were less efficient pollinators ofGrevillea barklyana than nectar-feeding birds. Nectar-collecting honeybees did not contact reproductive parts of flowers. Pollen-collecting honeybees preferentially visited malestage flowers but rarely visited female-stage flowers. Fruit set on caged inflorescences that allowed access to honeybees but excluded birds was reduced by more than 50% compared to inflorescences that were visited by both types of visitors. Further, fruit set on caged inflorescences was less than on bagged inflorescences that excluded both birds and honeybees, indicating that pollen removal by bees decreased opportunities for delayed autonomous selfing in the absence of birds. Although fruit set was not pollen-limited at the study site, pollen removal by honeybees would decrease fruit set in small populations where birds are scarce. In addition, pollen removal by honeybees would reduce opportunities for outcrossing and reproductive success through male function. Although honeybees have been in Australia for insufficient time to have exerted selection on floral traits, evolutionary shifts in response to these animals are likely to occur in the future.     

Phytogeography, Biology and Conservation of Western Australian Epacridaceae

The Epacridaceae of Western Australia comprise 181 named speciesdistributed in 17 genera. One hundred and seventy seven species(98% of the total) and ten genera are endemic to the region.within Western Australia, all members are confined to the south,with a bimodal pattern of species diversity centred on the sandplainsand uplands around the Stirling Ranges and Mount Lesueur.Leucopogon, the largest genus, reflects this pattern, however, speciesdiversity is greatest on the south coastal sandplains. All species are woody shrubs, mostly found on neutral or acidicsandy soils. Plants of at least 120 species are killed by fire;Astrolomacontainsthe only significant number of resprouting species. Althoughsome species are in flower every month every year, autumn/winterflowering is a common feature of many members of the family.All species are biotically pollinated. Bird pollination is foundin 18 species in six genera. The remaining species are insectpollinated. Bees are the major pollinators inLeucopogon, ConostephiumandColeanthera.Moths and butterflies are important pollinators inAndersonia,Leucopogon,LysinemaandStyphelia. Flies are minor pollinators of some speciesofAndersonia, LeucopogonandMonotoca. All species studied areprotandrous and outbreeding, with onlyCosmeliabeing capableof self pollination. Fruits are fleshy inAstroloma, Brachyloma,Croninia, in certain species ofLeucopogonandStypheliaand areanimal dispersed. Other genera have dry, dehiscent fruit withgravity or wind dispersed seeds. Many species have relatively narrow distribution ranges, 90species having ranges of over 300km, 54 species have rangesof 100–300km and 37 have ranges of less than 100km (andthere are many undescribed species in this group). The conservationstatus of many species is still poorly known. Dieback diseaseis a major threat to the family, and could cause the extinctionof several species in the wild. Western Australia; Epacridaceae; phytogeography; biology; pollination; dispersal; conservation; disease     

Pyric Herbivory and the Nexus Between Forage,Fire and Native and Introduced Large Grazing Herbivores in Australian Tropical Savannas

Earth’s tropical savannas typically support high biomass of diverse grazing herbivores that depend on a highly fluctuating resource: high-quality forage. An annual wet–dry cycle, fire and herbivory combine to influence forage quality and availability throughout the year. In the savannas of northern Australia, a depauperate suite of large native (marsupial) herbivores (wallaroos [Osphranter spp.] and the agile wallaby [Notamacropus agilis]) compete for resources with non-native large herbivores introduced in the late nineteenth century, particularly bovines (feral and managed cattle [Bos spp.] and feral water buffalo [Bubalus bubalis]) that now dominate the landscape. Anecdotal reports of recent population declines of large macropods and negative impacts of bovines highlight the need to better understand the complex relationship between forage, fire and abundance of native and introduced large herbivores. The pyric herbivory conceptual model, which posits complex feedbacks between fire and herbivory and was developed outside Australia, predicts that native and introduced large herbivores will both respond positively to post-fire forage production in Australian savannas where they co-occur. We used grazing exclosures, forage biomass and nutrient analyses and motion-sensor camera-trapping to evaluate the overall robustness of the pyric herbivory model in the Australian context, specifically whether forage quantity and quality are impacted by herbivory, season and fire activity, and which forage attributes most influence large grazing herbivore abundance. Forage quantity, as measured by live, dead and total herbaceous biomass and proportion of biomass alive, was higher inside herbivore exclosures, even at relatively low densities of herbivores. Forage quality, as measured by fibre content, was not affected by herbivory, however, crude protein content of live herbaceous biomass was greater outside herbivore exclosures. Recent fire was an important predictor of all measures of forage quantity and quality. Recent fire occurrence decreased overall quantity (biomass) but increased quality (decreased fibre content and increased crude protein content); late dry season fires resulted in forage with the highest crude protein content. The predictions of the pyric herbivory conceptual model are consistent with observations of the feeding behaviour of introduced bovines and some large macropods in northern Australian savannas, lending support to the global generality of pyric herbivory in fire-prone grassy biomes.

     

Maize seedling morphology and defence hormone profiles,but not herbivory tolerance,were mediated by domestication and modern breeding

We addressed whether Zea seedling morphology relevant to performance, defence hormone profiles and tolerance of a phloem‐feeding, specialist herbivore were affected by two processes, plant domestication and modern breeding. Domestication effects were inferred through comparisons between Balsas teosintes (Zea mays parviglumis) and landrace maizes (Z. mays mays), and modern breeding effects through comparisons between landrace maizes and inbred maize lines. Specifically, we compared seedling forms (a composite measure of leaf length, average stem diameter, shoot wet weight, shoot dry weight, total root length, root wet weight, and root dry weight), shapes (forms scaled by seedling dry weight, a proxy for seedling size), and defence hormone profiles among Balsas teosinte and landrace and inbred line maizes, exposed or unexposed to feeding by Dalbulus maidis. Our results suggested that domestication as well as modern breeding strongly mediated both seedling form and shape. Form was more similar between landrace and inbred maize than between Balsas teosinte and landrace maize, suggesting that domestication affected seedling form more than modern breeding. In contrast, shape was more similar between Balsas teosinte and landrace maize than between landrace and inbred maizes, suggesting that modern breeding affected seedling shape more than domestication. Additionally, seedling shoot : root ratios appeared to have been mediated by domestication, but not by modern breeding. In broad terms, individual seedling structures relevant to seedling ecology in wild or managed environments, such as leaf and root lengths, and shoot and root masses, were enlarged with domestication and reduced with modern breeding. Herbivory did not affect seedling shape, but had a weak effect on form so that seedlings were slightly larger in the absence versus presence of D. maidis. Also, both domestication and modern breeding seem to have mediated seedling hormone profiles, with breeding more strongly mediating profiles than domestication. Jasmonic acid isoleucine (JA‐Ile) and salicylic acid (SA) were induced by herbivory in both teosinte and maize. The hormone profiles assays collectively suggested that domestication and modern breeding altered constitutive levels of SA, abscisic acid and JA‐related (JA‐Ile and oxo‐phytodienoic acid) hormone levels in seedlings, particularly by increasing the levels of SA and decreasing those of JA‐related hormones. Altogether, our results suggested that maize domestication and modern breeding significantly altered seedling form, shape, ecologically relevant morphological traits (e.g. leaf and root lengths, and shoot and root masses) and hormonal defences, but not tolerance of D. maidis herbivory.     

Climate change, plant migration, and range collapse in a global biodiversity hotspot: the Banksia (Proteaceae) of Western Australia

Climate change has already altered global patterns of biodiversity by modifying the geographic distributions of species. Forecasts based on bioclimatic envelop modeling of distributions of species suggests greater impacts can be expected in the future, but such projections are contingent on assumptions regarding future climate and migration rates of species. Here, we present a first assessment of the potential impact of climate change on a global biodiversity hotspot in southwestern Western Australia. Across three representative scenarios of future climate change, we simulated migration of 100 Banksia (Proteaceae) species at a rate of 5 km decade^?1 and compared projected impacts with those under the commonly applied, but acknowledged as inadequate, assumptions of ‘full‐’ and ‘no‐migration.’ Across all climate × migration scenarios, 66% of species were projected to decline, whereas only 6% were projected to expand or remain stable. Between 5% and 25% of species were projected to suffer range losses of 100% by 2080, depending mainly on climate scenario. Species losses were driven primarily by changes in current precipitation regimes, with the greatest losses of species projected to occur in a transition zone between wet coastal areas and interior arid regions and which is projected to become more arid in the future. Because the ranges of most species tended to collapse in all climate scenarios, we found that climate change impacts to flora of southwestern Western Australia may be large, even under optimistic assumptions regarding migration abilities. Taken together, our results suggest that the future of biodiversity in southwestern Western Australia may lie largely in the degree to which this hotspot experiences increased drought and in the ability of species to tolerate such decreases in precipitation. More broadly, our study is among a growing number of theoretical studies suggesting the impacts of future climate change on global biodiversity may be considerable.