Echoes of ancient introgression punctuate stable genomic lineages in the evolution of figs.

Studies investigating the evolution of flowering plants have long focused on isolating mechanisms such as pollinator specificity. Some recent studies have proposed a role for introgressive hybridization between species, recognizing that isolating processes such as pollinator specialization may not be complete barriers to hybridization. Occasional hybridization may therefore lead to distinct yet reproductively connected lineages. We investigate the balance between introgression and reproductive isolation in a diverse clade using a densely sampled phylogenomic study of fig trees (Ficus, Moraceae). Codiversification with specialized pollinating wasps (Agaonidae) is recognized as a major engine of fig diversity, leading to about 850 species. Nevertheless, some studies have focused on the importance of hybridization in Ficus, highlighting the consequences of pollinator sharing. Here, we employ dense taxon sampling (520 species) throughout Moraceae and 1,751 loci to investigate phylogenetic relationships and the prevalence of introgression among species throughout the history of Ficus. We present a well-resolved phylogenomic backbone for Ficus, providing a solid foundation for an updated classification. Our results paint a picture of phylogenetically stable evolution within lineages punctuated by occasional local introgression events likely mediated by local pollinator sharing, illustrated by clear cases of cytoplasmic introgression that have been nearly drowned out of the nuclear genome through subsequent lineage fidelity. The phylogenetic history of figs thus highlights that while hybridization is an important process in plant evolution, the mere ability of species to hybridize locally does not necessarily translate into ongoing introgression between distant lineages, particularly in the presence of obligate plant-pollinator relationships.

Effects of lianas on forest biogeochemistry during their lives and afterlives.

Climate change and other anthropogenic disturbances are increasing liana abundance and biomass in many tropical and subtropical forests. While the effects of living lianas on species diversity, ecosystem carbon, and nutrient dynamics are receiving increasing attention, the role of dead lianas in forest ecosystems has been little studied and is poorly understood. Trees and lianas coexist as the major woody components of forests worldwide, but they have very different ecological strategies, with lianas relying on trees for mechanical support. Consequently, trees and lianas have evolved highly divergent stem, leaf, and root traits. Here we show that this trait divergence is likely to persist after death, into the afterlives of these organs, leading to divergent effects on forest biogeochemistry. We introduce a conceptual framework combining horizontal, vertical, and time dimensions for the effects of liana proliferation and liana tissue decomposition on ecosystem carbon and nutrient cycling. We propose a series of empirical studies comparing traits between lianas and trees to answer questions concerning the influence of trait afterlives on the decomposability of liana and tree organs. Such studies will increase our understanding of the contribution of lianas to terrestrial biogeochemical cycling, and help predict the effects of their increasing abundance.

Exploring systematic biases, rooting methods and morphological evidence to unravel the evolutionary history of the genus Ficus (Moraceae).

Despite many attempts in the Sanger sequencing era, the phylogeny of fig trees remains unresolved, which limits our ability to analyze the evolution of key traits that may have contributed to their evolutionary and ecological success. We used restriction-site-associated DNA sequencing (c. 420 kb) and 102 morphological characters to elucidate the relationships between 70 species of Ficus. To increase phylogenetic information for higher-level relationships, we targeted conserved regions and assembled paired reads into long loci to enable the retrieval of homologous loci in outgroup genomes. We compared morphological and molecular results to highlight discrepancies and reveal possible inference bias. For the first time, we recovered a monophyletic subgenus Urostigma (stranglers) and a clade with all gynodioecious Ficus. However, we show, with a new approach based on iterative principal component analysis, that it is not (and will probably never be) possible to homogenize evolutionary rates and GC content for all taxa before phylogenetic inference. Four competing positions for the root of the molecular tree are possible. The placement of section Pharmacosycea as sister to other fig trees is not supported by morphological data and considered a result of a long-branch attraction artefact to the outgroups. Regarding morphological features and indirect evidence from the pollinator tree of life, the topology that divides Ficus into monoecious versus gynodioecious species appears most plausible. It seems most likely that the ancestor of fig trees was a freestanding tree and active pollination is inferred as the ancestral state, contrary to previous hypotheses. However, ambiguity remains on the ancestral breeding system. Despite morphological plasticity, we advocate restoring a central role to morphology in our understanding of the evolution of Ficus, as it can help detect systematic errors that appear more pronounced with larger molecular datasets.

Changes in Fungal Communities across a Forest Disturbance Gradient.

Deforestation has a substantial impact on aboveground biodiversity, but the response of belowground soil fungi remains poorly understood. In a tropical montane rainforest in southwestern China, plots were established along a forest degradation gradient ranging from mature and regenerated forests to open land to examine the impacts of forest degradation and deforestation on ecosystem diversity and function. Here, we evaluated the changes in belowground fungal diversity and community composition using a metabarcoding approach. Soil saprotrophic fungal richness declined with increasing forest disturbance. For example, Penicillium spp. (phosphorus [P]-solubilizing fungi) dominated in mature forest but were less abundant in regenerating forests and showed the lowest abundance in open land sites. Conversely, the abundance of facultative pathogenic fungi increased along the disturbance gradient. The decline in soil saprophytic fungi may be a direct result of forest disturbance or it may be associated with increased availability of soil phosphorus indirectly through an increase in soil pH. The increase in abundance of facultative pathogenic fungi may be related to reduced competition with saprotrophic fungi, changes in microclimate, or increased spore rain. These results demonstrate a loss of dominant P-solubilizing saprotrophic fungi along the disturbance gradient, indicating a change from soil P limitation in mature tropical forests to soil C limitation in deforested sites. The increased prevalence of pathogenic fungi may inhibit plant succession following deforestation. Overall, this research demonstrates that soil fungi can be used as a sensitive indicator for soil health to evaluate the consequences of forest disturbance.IMPORTANCE The soil fungal functional group changes in response to forest disturbance and indicates a close interaction between the aboveground plant community and the belowground soil biological community. Soil saprotrophic fungi declined in relative abundance with increasing forest disturbance. At the same time, the relative abundance of facultative pathogenic fungi increased. The loss of saprotrophic fungal richness and abundance may have been a direct result of forest disturbance or an indirect result of changes in soil pH and soil P. Furthermore, the dominant P-solubilizing saprotrophic fungi were replaced by diverse facultative pathogenic fungi, which have weaker C decomposition ability. These changes potentially indicate a shift from soil phosphate limitation to carbon limitation following deforestation. This study suggests that changes in fungal functional group composition can be used as an indicator of the effects of forest disturbance on soil carbon and nutrients.

Agro-ecological options for fall armyworm (Spodoptera frugiperda JE Smith) management: Providing low-cost, smallholder friendly solutions to an invasive pest.

Fall armyworm (FAW), a voracious agricultural pest native to North and South America, was first detected on the African continent in 2016 and has subsequently spread throughout the continent and across Asia. It has been predicted that FAW could cause up to $US13 billion per annum in crop losses throughout sub-Saharan Africa, thereby threatening the livelihoods of millions of poor farmers. In their haste to respond to FAW governments may promote indiscriminate use of chemical pesticides which, aside from human health and environmental risks, could undermine smallholder pest management strategies that depend to a large degree on natural enemies. Agro-ecological approaches offer culturally appropriate low-cost pest control strategies that can be readily integrated into existing efforts to improve smallholder incomes and resilience through sustainable intensification. Such approaches should therefore be promoted as a core component of integrated pest management (IPM) programmes for FAW in combination with crop breeding for pest resistance, classical biological control and selective use of safe pesticides. Nonetheless, the suitability of agro-ecological measures for reducing FAW densities and impact need to be carefully assessed across varied environmental and socio-economic conditions before they can be proposed for wide-scale implementation. To support this process, we review evidence for the efficacy of potential agro-ecological measures for controlling FAW and other pests, consider the associated risks, and draw attention to critical knowledge gaps. The evidence indicates that several measures can be adopted immediately. These include (i) sustainable soil fertility management, especially measures that maintain or restore soil organic carbon; (ii) intercropping with appropriately selected companion plants; and (iii) diversifying the farm environment through management of (semi)natural habitats at multiple spatial scales. Nevertheless, we recommend embedding trials into upscaling programmes so that the costs and benefits of these interventions may be determined across the diverse biophysical and socio-economic contexts that are found in the invaded range.

Plant diversity accurately predicts insect diversity in two tropical landscapes.

Plant diversity surely determines arthropod diversity, but only moderate correlations between arthropod and plant species richness had been observed until Basset et al. (Science, 338, 2012 and 1481) finally undertook an unprecedentedly comprehensive sampling of a tropical forest and demonstrated that plant species richness could indeed accurately predict arthropod species richness. We now require a high-throughput pipeline to operationalize this result so that we can (i) test competing explanations for tropical arthropod megadiversity, (ii) improve estimates of global eukaryotic species diversity, and (iii) use plant and arthropod communities as efficient proxies for each other, thus improving the efficiency of conservation planning and of detecting forest degradation and recovery. We therefore applied metabarcoding to Malaise-trap samples across two tropical landscapes in China. We demonstrate that plant species richness can accurately predict arthropod (mostly insect) species richness and that plant and insect community compositions are highly correlated, even in landscapes that are large, heterogeneous and anthropogenically modified. Finally, we review how metabarcoding makes feasible highly replicated tests of the major competing explanations for tropical megadiversity.

Impacts of hunting on tropical forests in Southeast Asia.

Although deforestation and forest degradation have long been considered the most significant threats to tropical biodiversity, across Southeast Asia (Northeast India, Indochina, Sundaland, Philippines) substantial areas of natural habitat have few wild animals (>1 kg), bar a few hunting-tolerant species. To document hunting impacts on vertebrate populations regionally, we conducted an extensive literature review, including papers in local journals and reports of governmental and nongovernmental agencies. Evidence from multiple sites indicated animal populations declined precipitously across the region since approximately 1980, and many species are now extirpated from substantial portions of their former ranges. Hunting is by far the greatest immediate threat to the survival of most of the region's endangered vertebrates. Causes of recent overhunting include improved access to forests and markets, improved hunting technology, and escalating demand for wild meat, wildlife-derived medicinal products, and wild animals as pets. Although hunters often take common species, such as pigs or rats, for their own consumption, they take rarer species opportunistically and sell surplus meat and commercially valuable products. There is also widespread targeted hunting of high-value species. Consequently, as currently practiced, hunting cannot be considered sustainable anywhere in the region, and in most places enforcement of protected-area and protected-species legislation is weak. The international community's focus on cross-border trade fails to address overexploitation of wildlife because hunting and the sale of wild meat is largely a local issue and most of the harvest is consumed in villages, rural towns, and nearby cities. In addition to improved enforcement, efforts to engage hunters and manage wildlife populations through sustainable hunting practices are urgently needed. Unless there is a step change in efforts to reduce wildlife exploitation to sustainable levels, the region will likely lose most of its iconic species, and many others besides, within the next few years.

Generic lures attract cerambycid beetles in a tropical montane rain forest in southern China.

The attraction of cerambycid beetles to 10 known cerambycid pheromones was tested in a tropical montane rain forest in southern China. From 28 May to 25 June 2010, 1,526 cerambycids representing 71 species were captured in pheromone-baited traps, with 14 species accounting for 92% of the specimens. Test compounds with a 3-hydroxyalkan-2-one or 2,3-alkanediol motif attracted significant numbers of both sexes for eight species in the subfamily Cerambycinae, including species of Demonax, Rhaphuma, and Xylotrechus. Rhaphuma horsfieldi (White) was the only species that was strongly attracted to more than one test compound, with significant attraction to both (2R*,3R*)-2,3-hexanediol and (2R*,3R*)-2,3-octanediol. Within the Lamiinae, males and females of five species were significantly attracted to 2-(undecyloxy) ethanol, including Acalolepta formosana (Breuning), Monochamus bimaculatus Gahan, Pharsalia subgemmata (Thomson), Pseudomacrochenus antennatus (Gahan), and Xenohammus bimaculatus Schwarzer. Only male Megopis costipennis White (Prioninae) were significantly attracted to (2R*,3S*)-2,3-octanediol, suggesting that this compound may be a sex pheromone component for this species. To date, 2,3-octanediols have only been reported as aggregation pheromone components for cerambycids in the subfamily Cerambycinae. Our results support the hypothesis that both closely related (congeners) and more distantly related cerambycid species (different tribes and even subfamilies) may share pheromone components. Our results also demonstrate that traps baited with even a limited number of different classes of pheromones may be useful tools for surveying cerambycid diversity, as well as for detecting and monitoring particular species, especially those that have the potential to be invasive pests in other parts of the world.

The effects of invertebrates on wood decomposition across the world.

Invertebrates and microorganisms are important but climate-dependent agents of wood decomposition globally. In this meta-analysis, we investigated what drives the invertebrate effect on wood decomposition worldwide. Globally, we found wood decomposition rates were on average approximately 40% higher when invertebrates were present compared to when they were excluded. This effect was most pronounced in the tropics, owing mainly to the activities of termites. The invertebrate effect was stronger for woody debris without bark as well as for that of larger diameter, possibly reflecting bark- and diameter-mediated differences in fungal colonisation or activity rates relative to those of invertebrates. Our meta-analysis shows similar overall invertebrate effect sizes on decomposition of woody debris derived from angiosperms and gymnosperms globally. Our results suggest the existence of critical interactions between microorganism colonisation and the invertebrate contribution to wood decomposition. To improve biogeochemical models, a better quantification of invertebrate contributions to wood decomposition is needed.

Consequences of defaunation for a tropical tree community.

Hunting affects a considerably greater area of the tropical forest biome than deforestation and logging combined. Often even large remote protected areas are depleted of a substantial proportion of their vertebrate fauna. However, understanding of the long-term ecological consequences of defaunation in tropical forests remains poor. Using tree census data from a large-scale plot monitored over a 15-year period since the approximate onset of intense hunting, we provide a comprehensive assessment of the immediate consequences of defaunation for a tropical tree community. Our data strongly suggest that over-hunting has engendered pervasive changes in tree population spatial structure and dynamics, leading to a consistent decline in local tree diversity over time. However, we do not find any support for suggestions that over-hunting reduces above-ground biomass or biomass accumulation rate in this forest. To maintain critical ecosystem processes in tropical forests increased efforts are required to protect and restore wildlife populations.

An extreme case of plant-insect codiversification: figs and fig-pollinating wasps.

It is thought that speciation in phytophagous insects is often due to colonization of novel host plants, because radiations of plant and insect lineages are typically asynchronous. Recent phylogenetic comparisons have supported this model of diversification for both insect herbivores and specialized pollinators. An exceptional case where contemporaneous plant-insect diversification might be expected is the obligate mutualism between fig trees (Ficus species, Moraceae) and their pollinating wasps (Agaonidae, Hymenoptera). The ubiquity and ecological significance of this mutualism in tropical and subtropical ecosystems has long intrigued biologists, but the systematic challenge posed by >750 interacting species pairs has hindered progress toward understanding its evolutionary history. In particular, taxon sampling and analytical tools have been insufficient for large-scale cophylogenetic analyses. Here, we sampled nearly 200 interacting pairs of fig and wasp species from across the globe. Two supermatrices were assembled: on an average, wasps had sequences from 77% of 6 genes (5.6 kb), figs had sequences from 60% of 5 genes (5.5 kb), and overall 850 new DNA sequences were generated for this study. We also developed a new analytical tool, Jane 2, for event-based phylogenetic reconciliation analysis of very large data sets. Separate Bayesian phylogenetic analyses for figs and fig wasps under relaxed molecular clock assumptions indicate Cretaceous diversification of crown groups and contemporaneous divergence for nearly half of all fig and pollinator lineages. Event-based cophylogenetic analyses further support the codiversification hypothesis. Biogeographic analyses indicate that the present-day distribution of fig and pollinator lineages is consistent with a Eurasian origin and subsequent dispersal, rather than with Gondwanan vicariance. Overall, our findings indicate that the fig-pollinator mutualism represents an extreme case among plant-insect interactions of coordinated dispersal and long-term codiversification. [Biogeography; coevolution; cospeciation; host switching; long-branch attraction; phylogeny.].

Pseudoreplication in tropical forests and the resulting effects on biodiversity conservation.

Tropical forest ecosystems are threatened by habitat conversion and other anthropogenic actions. Timber production forests can augment the conservation value of primary forest reserves, but studies of logging effects often yield contradictory findings and thus inhibit efforts to develop clear conservation strategies. We hypothesized that much of this variability reflects a common methodological flaw, simple pseudoreplication, that confounds logging effects with preexisting spatial variation. We reviewed recent studies of the effects of logging on biodiversity in tropical forests (n = 77) and found that 68% were definitively pseudoreplicated while only 7% were definitively free of pseudoreplication. The remaining proportion could not be clearly categorized. In addition, we collected compositional data on 7 taxa in 24 primary forest research plots and systematically analyzed subsets of these plots to calculate the probability that a pseudoreplicated comparison would incorrectly identify a treatment effect. Rates of false inference (i.e., the spurious detection of a treatment effect) were >0.5 for 2 taxa, 0.3-0.5 for 2 taxa, and <0.3 for 3 taxa. Our findings demonstrate that tropical conservation strategies are being informed by a body of literature that is rife with unwarranted inferences. Addressing pseudoreplication is essential for accurately assessing biodiversity in logged forests, identifying the relative merits of specific management practices and landscape configurations, and effectively balancing conservation with timber production in tropical forests.

Understanding the impact of fall armyworm (Spodoptera frugiperda J. E. Smith) leaf damage on maize yields.

Fall armyworm (Spodoptera frugiperda J. E. Smith), a serious pest of maize and other cereals, recently invaded the Old World potentially threatening the food security and incomes of millions of smallholder farmers. Being able to assess the impacts of a pest on yields is fundamental to developing Integrated Pest Management (IPM) approaches. Hence, working with an early maturing, medium maturing and late maturing variety, we inoculated maize plants with 2nd instar S. frugiperda larvae at V5, V8, V12, VT and R1 growth stages to investigate the effects of FAW induced damage on yield. Different plants were inoculated 0-3 times and larvae were removed after 1 or 2 weeks to generate a wide range of damage profiles. We scored plants for leaf damage at 3, 5 and 7 weeks after emergence (WAE) using the 9 point Davis scale. While at harvest we assessed ear damage (1-9 scale), and recorded plant height and grain yield per plant. We used Structural Equation Models to assess the direct effects of leaf damage on yield and indirect effects via plant height. For the early and medium maturing varieties leaf damage at 3 and 5 WAE, respectively, had significant negative linear effects on grain yield. In the late maturing variety, leaf damage at 7 WAE had an indirect effect on yield through a significant negative linear effect on plant height. However, despite the controlled screenhouse conditions, in all three varieties leaf damage explained less than 3% of the variation in yield at the plant level. Overall, these results indicate that S. frugiperda induced leaf damage has a slight but detectable impact on yield at a specific plant developmental stage, and our models will contribute to the development of decision-support tools for IPM. However, given the low average yields obtained by smallholders in sub-Saharan Africa and the relatively low levels of FAW induced leaf damage recorded in most areas, IPM strategies should focus on interventions aimed at improving plant vigour (e.g. through integrated soil fertility management) and the role of natural enemies, as these are likely to result in greater yield gains at lower cost than a focus on FAW control.

Ecology of hemiepiphytism in fig species is based on evolutionary correlation of hydraulics and carbon economy.

Woody hemiepiphytic species (Hs) are important components of tropical rain forests, and they have been hypothesized to differ from non-hemiepiphytic tree species (NHs) in adaptations relating to water relations and carbon economy; but few studies have been conducted comparing ecophysiological traits between the two growth forms especially in an evolutionary context. Using common-garden plants of the genus Ficus, functional traits related to plant hydraulics and carbon economy were compared for seven NHs and seven Hs in their adult terrestrial "tree-like" growth phase. We used phylogenetically independent contrasts to test the hypothesis that differences in water availability selected for contrasting suites of traits in Hs and NHs, driving evolutionary correlations among functional traits including hydraulic conductivity and photosynthetic traits. Species of the two growth forms differed in functional traits; Hs had substantially lower xylem hydraulic conductivity and stomatal conductance, and higher instantaneous photosynthetic water use efficiency. Leaf morphological and structural traits also differed strikingly between the two growth forms. The Hs had significantly smaller leaves, higher leaf mass per area (LMA), and smaller xylem vessel lumen diameters. Across all the species, hydraulic conductivity was positively correlated with leaf gas exchange indicating high degrees of hydraulic-photosynthetic coordination. More importantly, these correlations were supported by correlations implemented on phylogenetic independent contrasts, suggesting that most trait correlations arose through repeated convergent evolution rather than as a result of chance events in the deep nodes of the lineage. Vatiation in xylem hydraulic conductivity was also centrally associated with a suite of other functional traits related to carbon economy and growth, such as LMA, water use efficiency, leaf nutrient concentration, and photosynthetic nutrient use efficiency, indicating important physiological constraints or trade-offs among functional traits. Shifts in this trait cluster apparently related to the adaptation to drought-prone canopy growth during the early life cycle of Hs and clearly affected ecophysiology of the later terrestrial stage of these species. Evolutionary flexibility in hydraulics and associated traits might be one basis for the hyper-diversification of Ficus species in tropical rain forests.

Rapid Assessment of Cerambycid Beetle Biodiversity in a Tropical Rainforest in Yunnan Province, China, Using a Multicomponent Pheromone Lure.

The Cerambycidae comprise a large and ecologically important family of wood-boring beetles. The purpose of this study was to examine the effectiveness of a generic lure as a potential monitoring tool. Working in a subtropical forest in southwest China, we set traps baited with generic lures at ground level (1 m) and canopy height (~18 m) across 22 randomly located forest plots (12 regenerating forest, 10 mature forest). Three stations were established per plot and each plot was trapped for 7 days in May-June 2013. In total, 4541 beetles of 71 species were caught, including 26 species with 10 or more individuals. We used Hierarchical Modeling of Species Communities (HMSC) to analyze the data and produced informative models for 18 species, showing that trap height, slope, elevation, and leaf-area index were important determinants of cerambycid distribution. Our results demonstrate the potential for using generic lures to detect and monitor cerambycid populations, both for regulatory purposes and for the study of cerambycid beetle ecology. Further research should focus on refining lure blends, and on repeated sampling to determine temporal and spatial dynamics of cerambycid communities.

Upscaling tropical restoration to deliver environmental benefits and socially equitable outcomes.

The UN Decade on Ecosystem Restoration offers immense potential to return hundreds of millions of hectares of degraded tropical landscapes to functioning ecosystems. Well-designed restoration can tackle multiple Sustainable Development Goals, driving synergistic benefits for biodiversity, ecosystem services, agricultural and timber production, and local livelihoods at large spatial scales. To deliver on this potential, restoration efforts must recognise and reduce trade-offs among objectives, and minimize competition with food production and conservation of native ecosystems. Restoration initiatives also need to confront core environmental challenges of climate change and inappropriate planting in savanna biomes, be robustly funded over the long term, and address issues of poor governance, inadequate land tenure, and socio-cultural disparities in benefits and costs. Tackling these issues using the landscape approach is vital to realising the potential for restoration to break the cycle of land degradation and poverty, and deliver on its core environmental and social promises.

Spatial scale changes the relationship between beta diversity, species richness and latitude.

The relationship between ß-diversity and latitude still remains to be a core question in ecology because of the lack of consensus between studies. One hypothesis for the lack of consensus between studies is that spatial scale changes the relationship between latitude and ß-diversity. Here, we test this hypothesis using tree data from 15 large-scale forest plots (greater than or equal to 15 ha, diameter at breast height ≥ 1 cm) across a latitudinal gradient (3-30o) in the Asia-Pacific region. We found that the observed ß-diversity decreased with increasing latitude when sampling local tree communities at small spatial scale (grain size ≤0.1 ha), but the observed ß-diversity did not change with latitude when sampling at large spatial scales (greater than or equal to 0.25 ha). Differences in latitudinal ß-diversity gradients across spatial scales were caused by pooled species richness (γ-diversity), which influenced observed ß-diversity values at small spatial scales, but not at large spatial scales. Therefore, spatial scale changes the relationship between ß-diversity, γ-diversity and latitude, and improving sample representativeness avoids the γ-dependence of ß-diversity.

Selection of Native Tree Species for Subtropical Forest Restoration in Southwest China.

The use of native species in forest restoration has been increasingly recognized as an effective means of restoring ecosystem functions and biodiversity to degraded areas across the world. However, successful selection of species adapted to local conditions requires specific knowledge which is often lacking, especially in developing countries. In order to scale up forest restoration, experimental data on the responses of native species to propagation and restoration treatments across a range of local conditions are required. In this study, the restoration potential of 34 native tree species was evaluated based on nursery research and field planting experiments at a highly degraded site in a subtropical area of southwest China. We examined species performance in terms of germination rates as well as survival rates and growth over 2 years after planting. Of the 34 species examined, 25 had a germination percentage greater than 50%. Survivorship ranged from 0 to 97% across species and was greater than 50% for 20 species. Mean monthly growth increments varied between species. Pioneer species performed well, and 14 mid- and late-successional species performed reasonably well to very well in this study. However, the remaining 16 mid- and late-successional species performed poorly. These results indicate that carefully selected mid- and late-successional species can be effectively incorporated into mixed species plantings. This data can be used to inform restoration planning, helping to identify suitable species and so enhance the biodiversity and resilience of restored forests.

Does spatial variation in predation pressure modulate selection for aposematism?

It is widely believed that aposematic signals should be conspicuous, but in nature, they vary from highly conspicuous to near cryptic. Current theory, including the honest signal or trade-off hypotheses of the toxicity-conspicuousness relationship, cannot explain why adequately toxic species vary substantially in their conspicuousness. Through a study of similarly toxic Danainae (Nymphalidae) butterflies and their mimics that vary remarkably in their conspicuousness, we show that the benefits of conspicuousness vary along a gradient of predation pressure. Highly conspicuous butterflies experienced lower avian attack rates when background predation pressure was low, but attack rates increased rapidly as background predation pressure increased. Conversely, the least conspicuous butterflies experienced higher attack rates at low predation pressures, but at high predation pressures, they appeared to benefit from crypsis. Attack rates of intermediately conspicuous butterflies remained moderate and constant along the predation pressure gradient. Mimics had a similar pattern but higher attack rates than their models and mimics tended to imitate the signal of less attacked model species along the predation pressure gradient. Predation pressure modulated signal fitness provides a possible mechanism for the maintenance of variation in conspicuousness of aposematic signals, as well as the initial survival of conspicuous signals in cryptic populations in the process of aposematic signal evolution, and an alternative explanation for the evolutionary gain and loss of mimicry.