Evaluating restoration trajectories using DNA metabarcoding of ground-dwelling and airborne invertebrates and associated plant communities.

Invertebrates are important for restoration processes as they are key drivers of many landscape-scale ecosystem functions; including pollination, nutrient cycling and soil formation. However, invertebrates are often overlooked in restoration monitoring because they are highly diverse, poorly described, and time-consuming to survey, and require increasingly scarce taxonomic expertise to enable identification. DNA metabarcoding is a relatively new tool for rapid survey that is able to address some of these concerns, and provide information about the taxa with which invertebrates are interacting via food webs and habitat. Here, we evaluate how invertebrate communities may be used to determine ecosystem trajectories during restoration. We collected ground-dwelling and airborne invertebrates across chronosequences of mine-site restoration in three ecologically disparate locations in Western Australia and identified invertebrate and plant communities using DNA metabarcoding. Ground-dwelling invertebrates showed the clearest restoration signals, with communities becoming more similar to reference communities over time. These patterns were weaker in airborne invertebrates, which have higher dispersal abilities and therefore less local fidelity to environmental conditions. Although we detected directional changes in community composition indicative of invertebrate recovery, patterns observed were inconsistent between study locations. The inclusion of plant assays allowed identification of plant species, as well as potential food sources and habitat. We demonstrate that DNA metabarcoding of invertebrate communities can be used to evaluate restoration trajectories. Testing and incorporating new monitoring techniques such as DNA metabarcoding is critical to improving restoration outcomes.

Dominance-diversity relationships in ant communities differ with invasion.

The relationship between levels of dominance and species richness is highly contentious, especially in ant communities. The dominance-impoverishment rule states that high levels of dominance only occur in species-poor communities, but there appear to be many cases of high levels of dominance in highly diverse communities. The extent to which dominant species limit local richness through competitive exclusion remains unclear, but such exclusion appears more apparent for non-native rather than native dominant species. Here we perform the first global analysis of the relationship between behavioral dominance and species richness. We used data from 1,293 local assemblages of ground-dwelling ants distributed across five continents to document the generality of the dominance-impoverishment rule, and to identify the biotic and abiotic conditions under which it does and does not apply. We found that the behavioral dominance-diversity relationship varies greatly, and depends on whether dominant species are native or non-native, whether dominance is considered as occurrence or relative abundance, and on variation in mean annual temperature. There were declines in diversity with increasing dominance in invaded communities, but diversity increased with increasing dominance in native communities. These patterns occur along the global temperature gradient. However, positive and negative relationships are strongest in the hottest sites. We also found that climate regulates the degree of behavioral dominance, but differently from how it shapes species richness. Our findings imply that, despite strong competitive interactions among ants, competitive exclusion is not a major driver of local richness in native ant communities. Although the dominance-impoverishment rule applies to invaded communities, we propose an alternative dominance-diversification rule for native communities.

A global database of ant species abundances.

What forces structure ecological assemblages? A key limitation to general insights about assemblage structure is the availability of data that are collected at a small spatial grain (local assemblages) and a large spatial extent (global coverage). Here, we present published and unpublished data from 51 ,388 ant abundance and occurrence records of more than 2,693 species and 7,953 morphospecies from local assemblages collected at 4,212 locations around the world. Ants were selected because they are diverse and abundant globally, comprise a large fraction of animal biomass in most terrestrial communities, and are key contributors to a range of ecosystem functions. Data were collected between 1949 and 2014, and include, for each geo-referenced sampling site, both the identity of the ants collected and details of sampling design, habitat type, and degree of disturbance. The aim of compiling this data set was to provide comprehensive species abundance data in order to test relationships between assemblage structure and environmental and biogeographic factors. Data were collected using a variety of standardized methods, such as pitfall and Winkler traps, and will be valuable for studies investigating large-scale forces structuring local assemblages. Understanding such relationships is particularly critical under current rates of global change. We encourage authors holding additional data on systematically collected ant assemblages, especially those in dry and cold, and remote areas, to contact us and contribute their data to this growing data set.

Climate mediates the effects of disturbance on ant assemblage structure.

Many studies have focused on the impacts of climate change on biological assemblages, yet little is known about how climate interacts with other major anthropogenic influences on biodiversity, such as habitat disturbance. Using a unique global database of 1128 local ant assemblages, we examined whether climate mediates the effects of habitat disturbance on assemblage structure at a global scale. Species richness and evenness were associated positively with temperature, and negatively with disturbance. However, the interaction among temperature, precipitation and disturbance shaped species richness and evenness. The effect was manifested through a failure of species richness to increase substantially with temperature in transformed habitats at low precipitation. At low precipitation levels, evenness increased with temperature in undisturbed sites, peaked at medium temperatures in disturbed sites and remained low in transformed sites. In warmer climates with lower rainfall, the effects of increasing disturbance on species richness and evenness were akin to decreases in temperature of up to 9°C. Anthropogenic disturbance and ongoing climate change may interact in complicated ways to shape the structure of assemblages, with hot, arid environments likely to be at greatest risk.

The critical role of ants in the extensive dispersal of Acacia seeds revealed by genetic parentage assignment.

Ants are prominent seed dispersal agents in many ecosystems, and dispersal distances are small in comparison with vertebrate dispersal agents. However, the distance and distribution of ant-mediated dispersal in arid/semi-arid environments remains poorly explored. We used microsatellite markers and parentage assignment to quantify the distance and distribution of dispersed seeds of Acacia karina, retrieved from the middens of Iridomyrmex agilis and Melophorus turneri perthensis. From parentage assignment, we could not distinguish the maternal from each parent pair assigned to each seed, so we applied two approaches to estimate dispersal distances, one conservative (CONS), where the parent closest to the ant midden was considered to be maternal, and the second where both parents were deemed equally likely (EL) to be maternal, and used both distances. Parentage was assigned to 124 seeds from eight middens. Maximum seed dispersal distances detected were 417 m (CONS) and 423 m (EL), more than double the estimated global maximum. Mean seed dispersal distances of 40 m (±5.8 SE) (CONS) and 79 m (±6.4 SE) (EL) exceeded the published global average of 2.24 m (±7.19 SD) by at least one order of magnitude. For both approaches and both ant species, seed dispersal was predominantly (44-84% of all seeds) within 50 m from the maternal source, with fewer dispersal events at longer distances. Ants in this semi-arid environment have demonstrated a greater capacity to disperse seeds than estimated elsewhere, which highlights their important role in this system, and suggests significant novel ecological and evolutionary consequences for myrmecochorous species in arid/semi-arid Australia.

Correlations between physical and chemical defences in plants: tradeoffs, syndromes, or just many different ways to skin a herbivorous cat?

Most plant species have a range of traits that deter herbivores. However, understanding of how different defences are related to one another is surprisingly weak. Many authors argue that defence traits trade off against one another, while others argue that they form coordinated defence syndromes. We collected a dataset of unprecedented taxonomic and geographic scope (261 species spanning 80 families, from 75 sites across the globe) to investigate relationships among four chemical and six physical defences. Five of the 45 pairwise correlations between defence traits were significant and three of these were tradeoffs. The relationship between species' overall chemical and physical defence levels was marginally nonsignificant (P = 0.08), and remained nonsignificant after accounting for phylogeny, growth form and abundance. Neither categorical principal component analysis (PCA) nor hierarchical cluster analysis supported the idea that species displayed defence syndromes. Our results do not support arguments for tradeoffs or for coordinated defence syndromes. Rather, plants display a range of combinations of defence traits. We suggest this lack of consistent defence syndromes may be adaptive, resulting from selective pressure to deploy a different combination of defences to coexisting species.

Putting plant resistance traits on the map: a test of the idea that plants are better defended at lower latitudes.

• It has long been believed that plant species from the tropics have higher levels of traits associated with resistance to herbivores than do species from higher latitudes. A meta-analysis recently showed that the published literature does not support this theory. However, the idea has never been tested using data gathered with consistent methods from a wide range of latitudes. • We quantified the relationship between latitude and a broad range of chemical and physical traits across 301 species from 75 sites world-wide. • Six putative resistance traits, including tannins, the concentration of lipids (an indicator of oils, waxes and resins), and leaf toughness were greater in high-latitude species. Six traits, including cyanide production and the presence of spines, were unrelated to latitude. Only ash content (an indicator of inorganic substances such as calcium oxalates and phytoliths) and the properties of species with delayed greening were higher in the tropics. • Our results do not support the hypothesis that tropical plants have higher levels of resistance traits than do plants from higher latitudes. If anything, plants have higher resistance toward the poles. The greater resistance traits of high-latitude species might be explained by the greater cost of losing a given amount of leaf tissue in low-productivity environments.

Canopy and litter ant assemblages share similar climate-species density relationships.

Tropical forest canopies house most of the globe's diversity, yet little is known about global patterns and drivers of canopy diversity. Here, we present models of ant species density, using climate, abundance and habitat (i.e. canopy versus litter) as predictors. Ant species density is positively associated with temperature and precipitation, and negatively (or non-significantly) associated with two metrics of seasonality, precipitation seasonality and temperature range. Ant species density was significantly higher in canopy samples, but this difference disappeared once abundance was considered. Thus, apparent differences in species density between canopy and litter samples are probably owing to differences in abundance-diversity relationships, and not differences in climate-diversity relationships. Thus, it appears that canopy and litter ant assemblages share a common abundance-diversity relationship influenced by similar but not identical climatic drivers.

The ecology of a keystone seed disperser, the ant Rhytidoponera violacea.

Rhytidoponera violacea (Forel) (Hymenoptera: Formicidae) is a keystone seed disperser in Kwongan heathl and habitats of southwestern Australia. Like many myrmecochorous ants, little is known about the basic biology of this species. In this study various aspects of the biology of R. violacea were examined and the researchers evaluated how these characteristics may influence seed dispersal. R. violacea nesting habits (relatively shallow nests), foraging behavior (scramble competitor and lax food selection criteria), and other life history characteristics complement their role as a mutualist that interacts with the seeds of many plant species.

Climatic drivers of hemispheric asymmetry in global patterns of ant species richness.

Although many taxa show a latitudinal gradient in richness, the relationship between latitude and species richness is often asymmetrical between the northern and southern hemispheres. Here we examine the latitudinal pattern of species richness across 1003 local ant assemblages. We find latitudinal asymmetry, with southern hemisphere sites being more diverse than northern hemisphere sites. Most of this asymmetry could be explained statistically by differences in contemporary climate. Local ant species richness was positively associated with temperature, but negatively (although weakly) associated with temperature range and precipitation. After contemporary climate was accounted for, a modest difference in diversity between hemispheres persisted, suggesting that factors other than contemporary climate contributed to the hemispherical asymmetry. The most parsimonious explanation for this remaining asymmetry is that greater climate change since the Eocene in the northern than in the southern hemisphere has led to more extinctions in the northern hemisphere with consequent effects on local ant species richness.

Info-gap theory and robust design of surveillance for invasive species: the case study of Barrow Island.

Surveillance for invasive non-indigenous species (NIS) is an integral part of a quarantine system. Estimating the efficiency of a surveillance strategy relies on many uncertain parameters estimated by experts, such as the efficiency of its components in face of the specific NIS, the ability of the NIS to inhabit different environments, and so on. Due to the importance of detecting an invasive NIS within a critical period of time, it is crucial that these uncertainties be accounted for in the design of the surveillance system. We formulate a detection model that takes into account, in addition to structured sampling for incursive NIS, incidental detection by untrained workers. We use info-gap theory for satisficing (not minimizing) the probability of detection, while at the same time maximizing the robustness to uncertainty. We demonstrate the trade-off between robustness to uncertainty, and an increase in the required probability of detection. An empirical example based on the detection of Pheidole megacephala on Barrow Island demonstrates the use of info-gap analysis to select a surveillance strategy.

Agricultural matrices affect ground ant assemblage composition inside forest fragments.

The establishment of agricultural matrices generally involves deforestation, which leads to fragmentation of the remaining forest. This fragmentation can affect forest dynamics both positively and negatively. Since most animal species are affected, certain groups can be used to measure the impact of such fragmentation. This study aimed to measure the impacts of agricultural crops (matrices) on ant communities of adjacent lower montane Atlantic rainforest fragments. We sampled nine forest fragments at locations surrounded by different agricultural matrices, namely: coffee (3 replicates); sugarcane (3); and pasture (3). At each site we installed pitfall traps along a 500 m transect from the interior of the matrix to the interior of the fragment (20 pitfall traps ~25 m apart). Each transect was partitioned into four categories: interior of the matrix; edge of the matrix; edge of the fragment; and interior of the fragment. For each sample site, we measured ant species richness and ant community composition within each transect category. Ant richness and composition differed between fragments and matrices. Each sample location had a specific composition of ants, probably because of the influence of the nature and management of the agricultural matrices. Species composition in the coffee matrix had the highest similarity to its corresponding fragment. The variability in species composition within forest fragments surrounded by pasture was greatest when compared with forest fragments surrounded by sugarcane or, to a lesser extent, coffee. Functional guild composition differed between locations, but the most representative guild was 'generalist' both in the agricultural matrices and forest fragments. Our results are important for understanding how agricultural matrices act on ant communities, and also, how these isolated forest fragments could act as an island of biodiversity in an 'ocean of crops'.

[The bamboo Merostachys fischeriana (Bambusoideae: Bambuseae) as a canopy habitat for ants of Neotropical Montane Forest]. / O bambu Merostachys fischeriana (Bambusoideae: Bambuseae) como habitat para formigas de Floresta Tropical Montana.

Although Merostachys fischeriana is very abundant in the Brazilian Atlantic Rainforest, little attention has been paid to the biological interactions with other animals. The present study describes some of the interactions between ants and this bamboo species. The experiment was carried out in a fragment of a montane tropical forest in the Parque Estadual do Itacolomi, near Ouro Preto, MG, Brazil. Thirty culms of bamboo were randomly collected. The ants were obtained by direct collection from nodes and internodes. Morphometric variables of the bamboo were recorded for characterization of potential ant habitat. Merostachys fischeriana grows in rosettes as a thin bamboo (average = 1,0 cm; se = 0,27; n = 20) and is tall enough to reach the upper canopy of this low forest (average = 9,1 m; se = 2,72; n = 20). Fifteen ant species were sampled. Brachymyrmex heeri Forel was the most abundant in the nodes, while Camponotus crassus Mayr (Hymenoptera: Formicidae) was the most abundant in the internodes. The composition of the species that inhabit the internodes was different from the composition in the node (Q-test: Q = 3,76; P = 0,05). The level of occupation was defined by the number of holes (F = 10,33; P < 0,01), the number of internodes in the canopy (F = 6,84; P = 0,01) and the length of the culm (F = 7,52; P = 0,01). The plant's morphology allowed the occurrence of additional species of ants in the canopy and influenced the composition of the entire ant assemblage.

Ants sow the seeds of global diversification in flowering plants.

BACKGROUND: The extraordinary diversification of angiosperm plants in the Cretaceous and Tertiary periods has produced an estimated 250,000-300,000 living angiosperm species and has fundamentally altered terrestrial ecosystems. Interactions with animals as pollinators or seed dispersers have long been suspected as drivers of angiosperm diversification, yet empirical examples remain sparse or inconclusive. Seed dispersal by ants (myrmecochory) may drive diversification as it can reduce extinction by providing selective advantages to plants and can increase speciation by enhancing geographical isolation by extremely limited dispersal distances. METHODOLOGY/PRINCIPAL FINDINGS: Using the most comprehensive sister-group comparison to date, we tested the hypothesis that myrmecochory leads to higher diversification rates in angiosperm plants. As predicted, diversification rates were substantially higher in ant-dispersed plants than in their non-myrmecochorous relatives. Data from 101 angiosperm lineages in 241 genera from all continents except Antarctica revealed that ant-dispersed lineages contained on average more than twice as many species as did their non-myrmecochorous sister groups. Contrasts in species diversity between sister groups demonstrated that diversification rates did not depend on seed dispersal mode in the sister group and were higher in myrmecochorous lineages in most biogeographic regions. CONCLUSIONS/SIGNIFICANCE: Myrmecochory, which has evolved independently at least 100 times in angiosperms and is estimated to be present in at least 77 families and 11 000 species, is a key evolutionary innovation and a globally important driver of plant diversity. Myrmecochory provides the best example to date for a consistent effect of any mutualism on large-scale diversification.

A keystone ant species promotes seed dispersal in a "diffuse" mutualism.

In order to understand the dynamics of co-evolution it is important to consider spatial variation in interaction dynamics. We examined the relative importance of ant activity, diversity and species identity in an ant seed dispersal mutualism at local, regional and continental scales. We also studied the determinants of seed dispersal rates and dispersal distances at eight sites in the Eneabba sandplain (29.63 S, 115.22 E), western Australia to understand local variation in seed dispersal rate and distance. To test the generality of the conclusions derived from the eight local sites, we established 16 sites along a 1650-km transect in western Australia, covering 11 degrees of latitude and a six-fold increase in rainfall, at which we sampled the ant assemblage, estimated ant species richness and ant activity and observed the removal rate of myrmecochorous seeds. We also assessed the importance of ant species identity at a continental scale via a review of studies carried out throughout Australia which examined ant seed dispersal. Among the eight sandplain shrubland sites, ant species identity, in particular the presence of one genus, Rhytidoponera, was associated with the most dispersal and above average dispersal distances. At the landscape scale, Rhytidoponera presence was the most important determinant of seed removal rate, while seed removal rate was negatively correlated with ant species richness and latitude. Most ant seed removal studies carried out throughout Australia reinforce our observations that Rhytidoponera species were particularly important seed dispersers. It is suggested that superficially diffuse mutualisms may depend greatly on the identity of particular partners. Even at large biogeographic scales, temporal and spatial variation in what are considered to be diffuse mutualisms may often be linked to variation in the abundance of particular partners, and be only weakly - or negatively - associated with the diversity of partners.

O bambu Merostachys fischeriana (Bambusoideae: Bambuseae) como habitat para formigas de Floresta Tropical Montana / The bamboo Merostachys fischeriana (Bambusoideae: Bambuseae) as a canopy habitat for ants of Neotropical Montane Forest

Although Merostachys fischeriana is very abundant in the Brazilian Atlantic Rainforest, little attention has been paid to the biological interactions with other animals. The present study describes some of the interactions between ants and this bamboo species. The experiment was carried out in a fragment of a montane tropical forest in the Parque Estadual do Itacolomi, near Ouro Preto, MG, Brazil. Thirty culms of bamboo were randomly collected. The ants were obtained by direct collection from nodes and internodes. Morphometric variables of the bamboo were recorded for characterization of potential ant habitat. Merostachys fischeriana grows in rosettes as a thin bamboo (average = 1,0 cm; se = 0,27; n = 20) and is tall enough to reach the upper canopy of this low forest (average = 9,1 m; se = 2,72; n = 20). Fifteen ant species were sampled. Brachymyrmex heeri Forel was the most abundant in the nodes, while Camponotus crassus Mayr (Hymenoptera: Formicidae) was the most abundant in the internodes. The composition of the species that inhabit the internodes was different from the composition in the node (Q-test: Q = 3,76; P = 0,05). The level of occupation was defined by the number of holes (F = 10,33; P < 0,01), the number of internodes in the canopy (F = 6,84; P = 0,01) and the length of the culm (F = 7,52; P = 0,01). The plant's morphology allowed the occurrence of additional species of ants in the canopy and influenced the composition of the entire ant assemblage.

Feeding and oviposition preferences of Ctenarytaina spatulata Taylor (Hemiptera, Psyllidae) for Eucalyptus spp. and other Myrtaceae in Brazil / Preferência de alimentação e oviposição de Ctenarytaina spatulata Taylor (Hemiptera, Psyllidae) por Eucalyptus spp. e outras Myrtaceae

Feeding and oviposition preferences of Ctenarytaina spatulata Taylor (Hemiptera, Psyllidae) for Eucalyptus spp. and other Myrtaceae in Brazil. The Australian psyllid, Ctenarytaina spatulata Taylor (Hemiptera, Psyllidae), was first detected in Brazil in 1994, where it was found on drought-affected shoots of Eucalyptus grandis in a plantation located in the northern part of Paraná State. The oviposition and feeding preferences of this psyllid were examined on 19 Eucalyptus species, one Eucalyptus hybrid (Cambiju), three Corymbia species and four native Myrtaceae species (Hexaclames edulis, Marlieria edulis, Plinia trunciflora, and Psydium sp.) under greenhouse conditions. The largest populations of C. spatulata were found on E. robusta and E. pellita, while sizeable infestations were also found on E. urophylla, E. grandis, and the Cambiju hybrid. The plants with the greatest symptoms of damage were E. grandis and E. resinifera. Eucalyptus cinerea, E. benthamii, E. pilularis, and E. dunnii were not infested and E. cloeziana was minimally infested. Among the Corymbia species, the number of eggs of C. spatulata was very low on C. citriodora and C. torelliana. No eggs and nymphs of C. spatulata were found on native Brazilian Myrtaceae. The number of eggs on plants was highly correlated with the subsequent levels of nymphs, suggesting that egg counts can be used as a viable monitoring tool to assist with the integrated management of this pest.

Ctenarytaina spatulata Taylor, 1997 (Hemiptera, Psyllidae), originária da Austrália, foi detectada no Brasil em 1994. Sua ocorrência foi inicialmente observada em E. grandis no Norte do Paraná associada à seca dos ponteiros. Com o objetivo de verificar a preferência deste psilídeo para postura e alimentação foram analisadas, em casa de vegetação, um híbrido e dezenove espécies do gênero Eucalyptus, três do gênero Corymbia e cinco mirtáceas nativas (Hexaclames edulis, Marlieria edulis, Plinia trunciflora and Psydium sp.). As maiores populações de C. spatulata ocorreram em E. robusta e E. pellita. No entanto, foram as espécies E. grandis e E. resinifera que apresentaram o maior número de plantas com sintomas de danos. As espécies E. cinerea, E. cloeziana, E. dunnii, E. benthamii, E. nitens, E. viminalis, E. pilularis e E. camaldulensis não apresentaram infestação por C. spatulata. Nas espécies de Corymbia foi observado um número muito reduzido de ovos de C. spatulata em C. citriodora e C. torelliana. Em nenhuma das espécies de Myrtaceae nativas foram observados ovos ou ninfas de C. spatulata. O número de ovos na planta foi altamente correlacionado com o número de ninfas, sugerindo que a contagem de ovos possa ser usada como uma ferramenta viável no monitoramento desta praga.