Eight years of ant‐exclusion from citrus canopies: effects on the arthropod assemblage and on fruit yield

• 1 Ants are important generalist predators in most terrestrial ecosystems. However, because many ant species are also hemipteran mutualists, their role in agriculture has generally been considered to be negative for plants.
• 2 In the present study, we report an experiment in ant‐exclusion from tree canopies in an organic citrus grove with two main objectives: (i) to examine whether the absence of ants increased the abundance of some beneficial arthropods and reduced the attack of some pests such as aphids and (ii) to examine whether ant‐exclusion increased the fruit yield of citrus trees.
• 3 The exclusion of ants from tree canopies had positive effects on the arthropod assemblage and on fruit yield. However, the 8‐year duration of the experiment can be divided into two periods with contrasting results. In the first period, the arthropod assemblage was only slightly affected, except for a greater density of aphids in ant‐excluded trees; in addition, fruit yield was higher in ant‐excluded trees than in the control ones. In the second period, ant‐exclusion increased the abundance of most arthropod groups, although the previous positive effect on fruit yield was no longer observed.
• 4 There are two main conclusions of the present study. First, from an applied perspective, ant‐exclusion from tree canopies is not a sound management alternative in citrus plantations in the Mediterranean. Second, the 8‐year duration of the experiment highlighted the importance of long‐term experiments in community ecology and biological control because the effects observed in the first 4 years of the experiment were very different from what occurred subsequently.

     

Effects of the concurrent exclusion of ants and earwigs on aphid abundance in an organic citrus grove

Based on the well-known mutualism between ants (Hymenoptera: Formicidae) and aphids (Homoptera: Aphididae), we conducted a five-year experiment of ant-exclusion from the canopies of citrus trees as a possible method of biological control of aphids. However, our results showed that the exclusion of ants from the canopies increased, instead of reducing, aphid abundance. To explain this unexpected result, we reasoned that the exclusion of ants from the canopies might also have excluded crawling insects that prey on aphids, such as the European earwig (Forficula auricularia L., Dermaptera: Forficulidae). Such a possibility is supported by the negative relationship between aphid density and the abundance of earwigs, consistent with a top-down control of aphids by earwigs. In contrast, the abundance of other aphid predators (Coleoptera: Coccinellidae, and Heteroptera) had no such negative effect on aphid density but a positive one, suggesting a bottom-up control, and showed no differences between control and ant-excluded trees. Thus, the most likely explanation for the increase in aphid abundance in the ant-excluded trees is the absence of earwigs from the canopies of the experimental trees, providing further evidence of the major role that earwigs play as control agents of aphids in cultivated trees.     

Abundance,interannual variation and potential pest predator role of two co‐occurring earwig species in citrus canopies

Earwigs are usually considered pest predators in orchards. Because of its worldwide distribution, most research on earwigs focuses on the European earwig Forficula auricularia Linnaeus (Insecta: Dermaptera: Forficulidae). However, very little is known of this species in Mediterranean citrus orchards. Earwigs and aphids were collected monthly during 5 years (2006–2010) from citrus canopies. Two species of earwigs were found: F. auricularia and Forficula pubescens Gené (=Guanchia pubescens), with the latter seldom cited in the literature. The goals of this study were (i) to document the abundance of these two earwig species in Mediterranean citrus canopies; (ii) to determine whether they are positively or negatively associated with each other, or randomly distributed; (iii) to measure the interannual variation of the abundance of both species during a 5‐year period and (iv) to evaluate the potential role of earwigs as pest predators in citrus canopies. As compared to colder regions, F. auricularia active period in citrus canopies in our study site lasted longer. Both species co‐occurred randomly in canopies. In 2006, both species showed approximately the same abundance, but in 2010, F. pubescens abundance in canopies was 28 times greater than that of F. auricularia. The potential role of earwigs as pest predators is higher in the Mediterranean than in other colder regions, because of the longer active period. F. auricularia is a sedentary generalist predator, already present in citrus canopies at the onset of most pest outbreaks, while F. pubescens arrived later to the canopies, but most likely was abundant enough to contribute in the control of citrus pests.     

Effects of ant competition and bird predation on the spider assemblage of a citrus grove

Characterizing intraguild interactions is key to improving understanding of food webs because they are major forces in the structuring of communities. Spiders are generalist predators with intermediate positions in the food web that establish intraguild interactions with ants and birds, which respectively compete with and prey on them. Research has also found interactions between birds and ants, potentially resulting in non-additive effects of both groups on arthropod assemblages, although studies of their combined impacts with tests for multiple-predator effects are scarce. We thus aimed to discern the relative effect of ants and birds on the spider assemblage of a citrus grove. We used a split-plot design to factorially exclude these groups over 2 years, preventing ants reaching the canopies by placing sticky bands around tree trunks, and birds by enclosing groups of trees in cages. We sampled spiders from the canopies (beating) and the ground (pitfalls) every 3 months, and we identified them to species. We found a strong influence of ants on the canopy spider assemblage, mainly through a negative effect on the families Araneidae and Theridiidae. Since spiders’ weights from ant-excluded and control trees were similar, these results suggest interference competition of ants on spiders rather than competitive exploitation. Bird exclusion did not affect the spider assemblage, contrasting with other studies reporting a marked predatory pressure of birds on spiders; nor were there any non-additive effects of ants and birds. Our findings show that spider assemblages are not uniformly affected by intraguild competitors.     

Potential predation of non-webbuilding spider assemblage on cotton pests Helicoverpa armigera and Spodoptera littoralis (Lepidoptera: Noctuidae)

The relative feeding rates and preferences of a hunting-spider assemblage inhabiting southern Spanish cotton fields for two major cotton pests, Helicoverpa armigera (Hübner) and Spodoptera littoralis (Boisdubal) (Lepidoptera: Noctuidae), were analyzed under laboratory conditions. First, a no-choice feeding test was used to determine relative feeding rates for hunting-spider families and species, offering a fixed number of 10 neonate larvae of H. armigera or S. littoralis and observing predation after 2 h, 4 h, 8 h, and 24 h. In a second test, Drosophila melanogaster, a very palatable alternative prey, was used to determine the degree of preference for cotton pest larvae. The mean number of first-instar lepidoptera larvae consumed by hunting spiders after 24 h was 8.57±0.25. As expected, spiders showed no preference for either of the two cotton pest species H. armigera and S. littoralis over the other. Results also showed that cursorial spiders of the families Miturigidae (represented here by Cheiracanthium pelasgicum) and Philodromidae consumed significantly higher percentages of larvae than crab spiders belonging to the Thomisidae family after 2 h and 24 h, respectively. In the prey choice test, Cheiracantium pelasgicum displayed a strong preference for cotton pest larvae while Thomisidae and Oxyopidae showed no significant preference. In addition, as the attack sequence progressed, Ch. pelasgicum showed a clear tendency towards the alternation of prey while Thomisidae, and more irregularly Oxyopidae, maintained their preference for D. melanogaster. These findings confirmed both the considerable potential value of some cursorial spiders (e.g. Ch. pelasgicum) in the biological control of lepidopteran cotton pests and the relatively low impact of other hunting spiders, e.g. Thomisidae, on pests of this kind.     

Ant exclusion in citrus over an 8-year period reveals a pervasive yet changing effect of ants on a Mediterranean spider assemblage

Ants and spiders are ubiquitous generalist predators that exert top-down control on herbivore populations. Research shows that intraguild interactions between ants and spiders can negatively affect spider populations, but there is a lack of long-term research documenting the strength of such interactions and the potentially different effects of ants on the diverse array of species in a spider assemblage. Similarly, the suitability of family-level surrogates for finding patterns revealed by species-level data (taxonomic sufficiency) has almost never been tested in spider assemblages. We present a long-term study in which we tested the impact of ants on the spider assemblage of a Mediterranean citrus grove by performing sequential 1-year experimental exclusions on tree canopies for 8 years. We found that ants had a widespread influence on the spider assemblage, although the effect was only evident in the last 5 years of the study. During those years, ants negatively affected many spiders, and effects were especially strong for sedentary spiders. Analyses at the family level also detected assemblage differences between treatments, but they concealed the different responses to ant exclusion shown by some related spider species. Our findings show that the effects of experimental manipulations in ecology can vary greatly over time and highlight the need for long-term studies to document species interactions.     

Presence of an adipokinetic peptide in the corpus cardiacum of Dermaptera but not in the neurohaemal aorta, and chemical and functional identification of the peptide

Corpora cardiaca of the earwigs Labidura riparia and Forficula auricularia contain a substance that causes hyperlipaemia in migratory locusts and hypertrehalosaemia in the American cockroaches. A conspecific bioassay in L. riparia revealed that this factor is lipid-mobilizing. Isolation of the neuropeptide was achieved by single-step RP-HPLC. The primary structure of the earwig adipokinetic peptide was elucidated by automated Edman degradation in combination with matrix-assisted laser desorption/ionization mass spectrometry. It is a blocked octapeptide, pGlu-Val-Asn-Phe-Ser-Thr-Gly-Trp-NH₂, previously denoted as Grb-AKH and first identified in Gryllus bimaculatus ( Gäde & Rinehart 1987 ). The synthetic peptide co-chromatographed under various conditions with the native peptide and, in biological assays, resulted in lipid-mobilization in L. riparia when injected in low concentration.     

Impact of avian and arthropod predation on lepidopteran caterpillar densities and plant productivity in an ephemeral agroecosystem

Abstract.  1. Most studies evaluating the combined impact of spiders and other predators on herbivore densities in agroecosystems have focused primarily on their trophic connections with invertebrate predators (e.g. carabids, chrysopids); however linkages among spiders and vertebrate predators may also help structure the population dynamics of insect herbivores. A field experiment was conducted to examine the impact of avian and spider predation on lepidopteran caterpillar densities and plant productivity within a Brassica agroecosystem.
2. Arthropod abundance, leaf-chewing damage, and final plant productivity associated with broccoli, Brassica oleracea L. (var. italica ), were recorded for four treatments: (1) bird present but spiders removed; (2) both birds and spiders present; (3) birds excluded, spiders present; and (4) birds and spiders both excluded.
3. Densities of Artogeia rapae L. (Lepidoptera: Pieridae) and Trichoplusia ni Hübner (Lepidoptera: Noctuidae) large caterpillars and post feeding stages were reduced significantly by bird predation. The abundance of large caterpillars was also reduced on spider-inhabited plants during early plant growth; however the assemblage of birds and spiders did not suppress caterpillar densities more significantly than either predator alone.
4. Plants protected by birds, spiders, and birds plus spiders sustained less folivory attributable to leaf chewing caterpillars than check plants. Plant productivity was also greater for predator-protected plants than check plants.
5. Although spiders and parasitoids were responsible for some of the mortality inflicted upon lepidopteran caterpillars, it was concluded that in this study system, birds are the most important natural enemies of folivores.     

The effects of deer exclusion on the development of a Mediterranean plant community affected by a wildfire

Despite the resilience of Mediterranean ecosystems to fire, the ecological restoration of burned plant communities can be hindered by ungulate herbivores, particularly in areas with high population densities. This study compares the postfire development of a shrub community with and without deer, after a wildfire occurred in 2003 in a protected area in Central Portugal. We monitored 12 fenced and 12 unfenced plots 2, 3, 4, and 8 years after fire. Within each plot, we established a linear transect and measured the monospecific canopy projections (plant patches). Five plant community indicators (patch number, average patch length, average patch height, patch cover, and patch phytovolume per square meter) were obtained. The diameter and height of individuals of the most abundant shrub species (Cistus salvifolius, Erica scoparia, Myrtus communis, Pistacia lentiscus, Rubus ulmifolius, and Ulex jussiaei) were also measured. These measurements were used as response variables in generalized linear mixed models in order to assess the effects of time‐after‐fire and fencing, on the development of the plant community. Patch height and phytovolume had a significantly higher growth in fenced plots. At the species level, C. salvifolius, M. communis, R. ulmifolius, and U. jussiaei showed a higher growth across time both in height and in diameter, in the absence of herbivory. This work shows that deer exclusion needs to be considered when aiming at the postfire restoration of Mediterranean shrub communities.     

Ants and plant size shape the structure of the arthropod community of Hirtella myrmecophila, an Amazonian ant-plant

Abstract.  1. To investigate the influence of plant size and the presence of resident ants on the arthropod community associated with the ant-plant Hirtella myrmecophila , ants ( Allomerus octoarticulatus ) were experimentally removed from plants of various sizes, with a second group of similar-sized plants remaining as controls. The abundance and diversity of arthropods on these plants were determined at 45-day intervals for 7 months. Collected arthropods were assigned into one of three predetermined groups: herbivores, spiders (mostly web-building spiders), or 'tourists' (mostly non-resident ants, hemipterans, dipterans, or scarabeid beetles).
2. Ant removal had a positive influence on the abundance and diversity of herbivores, but no measurable effects on the abundance and diversity of 'tourists', while the effect on spider abundance was significant only when data from Dipoena sp. nov. were removed from the analysis. While A. octoarticulatus had a negative effect on most spiders, it favoured Dipoena sp. nov., which was found to be a specialist predator of A. octoarticulatus .
3. Plant size positively affected the abundance and diversity of 'tourists' in plants with and without ants, but for herbivores and spiders this effect was dependent on ant treatment. In ant-maintained plants the per-plant abundance of herbivores and spiders was independent of plant size, whereas in ant-removed plants it was not.
4. These results suggest that A. octoarticulatus affects all arthropods found on its host plant, except those presenting an occasional and temporary association with the plant, and that the magnitude of ant effects on the susceptible guilds increases as plant size increases.     

Effect of retentive markers on the dynamics of settlement: The case of arthropod silk

During a settlement decision, the presence of conspecifics is crucial to species subject to Allee effects, for which the number of founders affects the subsequent growth of the colony. Marking the area (physically or chemically) conveys information about the number of conspecifics present in a new patch. Here, we study how an individual affinity for the marker affects the dynamics of a foundation process. A generic population model is presented, in which marking and affinity for the marker are at stake. Our results show that population size thresholds can appear, below which settlement is not possible. This model is then used to study the dynamics of migration and aggregation in a set of interconnected populations. We show that affinity for the marker can induce asymmetries in the population distribution. Anelosimus eximius is a social spider subject to Allee effects, for which silk potentially acts as a marker. We test our predictions with field experiments involving two populations of A. eximius in a Y-shaped setup. The agreement between our experimental and theoretical results strongly supports the validity of the model. This allows us to use the model to estimate a realistic set of parameters of biological significance to this social spider.     

Strong positive effects of termites on savanna bird abundance and diversity are amplified by large herbivore exclusion

Vast areas of the African savanna landscapes are characterized by tree‐covered Macrotermes termite mounds embedded within a relatively open savanna matrix. In concert with termites, large herbivores are important determinants of savanna woody vegetation cover. The relative cover of woody species has considerable effects on savanna function. Despite the potentially important ecological relationships between termite mounds, woody plants, large herbivores, and birds, these associations have previously received surprisingly little attention. We experimentally studied the effects of termites and large herbivores on the avian community in Lake Mburo National Park, Uganda, where woody vegetation is essentially limited to termite mounds. Our experiment comprised of four treatments in nine replicates; unfenced termite mounds, fenced mounds (excluding large mammals), unfenced adjacent savanna, and fenced savanna. We recorded species identity, abundance, and behavior of all birds observed on these plots over a two‐month period, from late dry until wet season. Birds used termite mounds almost exclusively, with only 3.5% of observations occurring in the treeless intermound savanna matrix. Mean abundance and species richness of birds doubled on fenced (large herbivores excluded) compared to unfenced mounds. Feeding behavior increased when large mammals were excluded from mounds, both in absolute number of observed individuals, and relative to other behaviors. This study documents the fundamental positive impact of Macrotermes termites on bird abundance and diversity in an African savanna. Birds play crucial functional roles in savanna ecosystems, for example, by dispersing fruits or regulating herbivorous insect populations. Thus, the role of birds in savanna dynamics depends on the distribution and abundance of termite mounds.     

Ant assemblage structure on cocoa trees in smallholder farms in the Centre Region of Cameroon

We investigated the ant community structure in cocoa farms in the Centre Region of Cameroon. Ants were collected on the cocoa trees during the years 2006 and 2007 using chemical knock‐down. We tested the hypothesis of the existence of deterministic factor in the structuration of ant mosaic using C‐Score; we assessed the relationship between the numerical dominant and subdominant ant species using Spearman correlation test and discussed on the influence of vegetation structure and farm management on the ant community structure. A total of 53 ant species belonging to 20 genera and five subfamilies were identified from a set of 51,525 workers collected. C‐score analysis supported the hypothesis that ant community were structured by competition. Negative relationships were found between dominant ant species. Farming practices which were mainly pruning, chemical treatment and habitat structure appeared to influence the ecological status and distribution of dominant ant species.     

Complex structural effects of two hemispheric climatic oscillators on the regional spatio-temporal expansion of a threatened bird

Links between climatic conditions in the eastern equatorial Pacific and extratropical ecological processes remain unexplored. The analysis of a 20‐year time series of spatial and numeric dynamics of a threatened Mediterranean bird suggests, however, that such couplings can be remarkably complex. By providing a new ecological time‐series modelling approach, we were able to dissect the joint effects of the El Niño/Southern Oscillation (ENSO), the North Atlantic Oscillation (NAO), regional weather, population density and stochastic variability on the expansion dynamics of the White‐headed duck (Oxyura leococephala) in Spain. Our results suggest that the spatial and numeric dynamics of ducks between peak brood emergence and wintering were simultaneously affected by different climatic phenomena during different phases of their global cycles, involving time lags in the numeric dynamics. Strikingly, our results point to both the NAO and the ENSO as potentially major factors simultaneously forcing ecological processes in the Northern Hemisphere, and suggest a new pathway for non‐additive effects of climate in ecology.     

Seed dispersal of a fleshy-fruited invasive shrub is affected by changes in the frugivorous bird assemblage along an elevational gradient

Seed dispersal by birds constitutes an essential mechanism for ornithochorous exotic plants to successfully invade a new system. New biotic associations with native birds might facilitate the upward spread of exotic plants from the foothills into the high mountains. However, environmental changes associated with elevation are known to drive changes in bird assemblages, and it is not clear how elevation changes impact the seed dispersal service of ornithochorous invaders. We evaluated changes in frugivorous bird assemblages of one of the exotic shrubs (Cotoneaster franchetii, Rosaceae) with the broadest elevation range among woody invaders in the Córdoba Mountains (Argentina). We quantified frugivory interactions (including absolute and proportional fruit consumption by seed dispersers, pulp consumers, and seed predators) using 4-h observations of focal C. franchetii shrubs distributed across low-elevation, mid-elevation, and high-elevation sites (700, 1100, and 1800 m a.s.l., respectively; 15 individuals per elevational band and one site per elevation). Seed disperser richness was highest at the low- and mid-elevation sites (three species vs. one at the high-elevation site), but proportional and absolute fruit consumption of C. franchetii was highest at the high-elevation site (39.1%, 88 seeds at high-elevation and 7.7%, 20 seeds at low-elevation). The Chiguanco Thrush (Turdus chiguanco, Turdidae) was the only seed disperser species found at the highest elevation site. Fruit consumption by seed dispersers was positively related to their abundance and elevation. In a high mountain system, a single abundant generalist seed disperser, rather than a high richness of seed disperser species, can uphold an effective dispersal service for an invasive ornithochorous shrub. This pattern may facilitate the spread of such plants across higher elevational ranges, thereby promoting the invasion of other exotic ornithochorous plants into upper elevations.     

有机化梨园节肢动物群落时间动态的聚类分析

对北京顺义地区有机化梨园中2个不同栽培区节肢动物总群落、害虫及天敌亚群落分别进行时间动态的系统聚类。结果表明,在同一园区内,稀植区和密植区均以木虱为优势种类,相对丰盛度分别达0.3801、0.3300。总群落的24次调查结果稀植区可聚为5类,D=1.87,密植区聚为3类,D=1.80;害虫亚群落调查结果,当两区D=1.80时,稀植区可聚为3类,密植区可聚为4类;天敌亚群落调查结果稀植区聚为4类,D=2.12,密植区聚为5类,D=1.75。在早春的密植区和入秋的稀植区木虱发生量大,应加强监测,及时防治。     

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.     

Assessing the role of invasive weeds in the impact of successional habitats on the bird assemblage in overgrowing agriculture

Agricultural habitats are assumed to be biodiversity refuges. However, some studies treat agricultural land management as a cause of the biodiversity decline, to which habitat loss and heterogeneity may contribute. Between the crops, the successional habitats appear – ruderal plant communities and bush areas. Their influence on farmland biodiversity is unknown. This research assessed the impact of spatial relationships between agricultural areas, semi-natural meadows and successional habitats on the bird species richness, Shannon diversity index, and Faith’s phylogenetic diversity index. An additional habitat variable was the presence of weeds, i.e., invasive Caucasian hogweeds Heracleum sp., treated as crops in the past. The birds and habitats research was on 74 sites set in pairs (invaded vs control) in south-eastern Poland. Results showed that birds assembling in agricultural and semi-natural areas were more diverse and contained protected farmland species, while birds appearing in overgrown habitats (i.e., successional and invaded) were clumped with their habitat requirements. In the presence of plant invaders, ruderal habitats negatively affected the bird phylogenetic diversity index. In invaded sites, bush areas had no positive effects on the Shannon diversity index and species richness of birds, in contrast with control sites. The presented research suggests the need to re-evaluate the importance of successional non-crop habitats considered positive in agricultural landscapes if those habitats develop in areas with plant invasion.