The effectiveness of post-contact defenses in a prey with no pre-contact detection

Most empirical and theoretical papers on prey–predator interactions are for animals with long-range detection, animals that can detect and react to predators long before these touch the prey. Heavy-bodied and chemically defended harvestmen (Arachnida, Opiliones) are an exception to this general pattern and rely on contact to detect arthropod predators. We examined the interactions between the Brazilian wandering spider Ctenus ornatus with harvestmen (Mischonyx cuspidatus) or control prey (Gryllus sp. and M. cuspidatus immature, both with soft integuments). Considering a prey–predator system in which fleeing from or reacting to a predator at a distance is not possible, we predicted both a high survival value of near-range defense mechanisms and that mortality would be higher in the absence of such defense mechanisms. We also expected the predator to behave differently when interacting with harvestmen or with a control prey without such defense mechanisms. Our results from laboratory experiments partially matched our predictions: First of all, histological sections showed that the integument of adult harvestmen is thicker than that of immature harvestmen and that of crickets. Adult harvestmen were less preyed upon than the control prey; the heavy armature increases the survival rate but the secretions from the scent glands do not. The predator did behave differently when attacking harvestmen compared to crickets. Despite the large size difference between predator and harvestmen, the protection provided by the armature allowed some of the harvestmen to survive encounters without pre-contact detection, thus greatly reducing the reliance on long-range detection to survive encounters with predators. Harvestmen call for theoretical and empirical work on prey–predator interactions that take into account the possibility that prey may not detect the predator before contact is established.     

A contribution to the functional morphology of the midgut gland in phalangiid harvestmen Gyas annulatus and Gyas titanus during their life cycle

The structure of the midgut gland and its changes in different seasons have been examined in the harvestmen Gyas annulatus and Gyas titanus (Arachnida: Opiliones: Phalangiidae). In both species, in the epithelium of the midgut gland two different types of cells are present: secretory and digestive ones. The secretory cells are characterized by plentiful rER and secretory granula. The digestive cells are characterized by an apical system of tubules. Both cells are connected by prominent specialized junctions. If a secretory cell is in contact with a digestive cell, rER cisterna are in close vicinity and parallel to these junctions. As found light- and electron microscopically and also histochemically, glycogen and lipids are stored in both cells. In both species, glycogen was seen to be used as energy compound during overwintering. At the end of their life, the digestive cells develop into excretory ones, containing metabolic wastes.     

Experimental demonstration of close-range olfaction and contact chemoreception in the Brazilian harvestman, Iporangaia pustulosa

We studied the ability to detect food by close-range olfaction and contact chemoreception in the harvestman Iporangaia pustulosa Mello-Leitão (Opiliones: Laniatores: Gonyleptidae). We first tested the reaction of individuals towards tasteless (pure agar), aversive (agar with salt), and food-intake stimulating substrates (agar with saccharose). Only the substrate containing saccharose was consumed. Contact (mainly with legs II) was necessary for detection of the agar and, before ingestion, the stimulus was always tapped with legs I. In the second experiment, we observed the behavior of individuals in an arena with a screened plastic box containing pieces of Tenebrio molitor L. (Coleoptera: Tenebrionidae) larvae. Individuals spent more time on the box containing food than on the control. In the third experiment, in an arena identical to that used in Experiment 2, we introduced a live but motionless T. molitor larva in the box. There was no difference between experimental and control treatments. We also observed the behavior of I. pustulosa in an arena containing live isopods. In first capture attempts, isopods were only detected upon contact, mainly with legs I. Our results suggest that (i) I. pustulosa is capable of detecting food only by its chemical properties; (ii) food with weak odor may not be detected by close-range olfaction; and (iii) legs I and II are important for food detection but, before ingestion, legs I are used to examine potential food items.     

Effects of livestock breed and grazing pressure on ground‐dwelling arthropods in Cantabrian heathlands

Abstract 1. The effects of different goat breeds and grazing pressures on epigeal arachnids (Lycosidae and Opiliones) and ground beetles (Carabidae) were assessed in a heath–gorse shrubland located in the north of the Iberian Peninsula. Three treatments (low grazing pressure with Cashmere breed, high grazing pressure with Cashmere breed, and high grazing pressure with local Celtiberic breed) with three replicates of each treatment were allocated randomly to nine plots. 2. Arthropod fauna abundance, diversity, and species richness were estimated using pitfall trapping over 3 years (2003, 2004, and 2005). Vegetation cover, composition, and height were assessed along transects and above‐ground plant biomass was sampled in quadrats. 3. Lower grazing pressure led to lower herbaceous cover and higher heather cover than high grazing pressure. Overall, it did not enhance arthropod diversity or abundance, but the abundance and species richness of carabids, and abundance of lycosids like Pardosa pullata, were higher under high stocking rates, whereas Opiliones were favoured by the low stocking rates. Although it has been assumed that traditional breeds enhance biodiversity, the present study found no significant differences for both arthropod fauna and vegetation between the goat breeds. 4. Arthropod–vegetation relationships were examined by constrained ordination. Fauna assemblages followed a gradient from closed canopy shrubland areas, in the lowest grazing pressure, to grassier open canopy areas resulting from higher grazing pressure, revealing the importance of grazing management as a driver of arthropod community structure of the heathland.     

Ground predator abundance affects prey removal in highbush blueberry (<Emphasis Type="BoldItalic">Vaccinium corymbosum</Emphasis>) fields and can be altered by aisle ground covers

Habitat management to conserve natural enemies has increased biological control of insect pests in various cropping systems [Annu. Rev. Entomol. 45: 175–201, 2000]. We wanted to determine if insect predation in highbush blueberry, Vaccinium corymbosum L. (Ericales: Ericaceae), is influenced by manipulation of edaphic arthropod community and whether management of ground cover in aisles between blueberry rows enhances this community. The first question was studied in blueberry plots bounded by trenches permitting selective movement into plots (ingress) or out of plots (egress), as well as unbounded control plots. We observed a significant effect of boundary type on the arthropod communities relative abundance as measured with pitfall traps, with relative abundance highest in ingress plots, intermediate in control plots and lowest in egress plots. Effects of ground arthropod abundance on predation rates were assessed with onion fly, Delia antiqua (Meigen) (Diptera: Anthomyiidae), pupae as sentinel prey. Pupa recovery was greatest in egress boundary plots, intermediate in control plots and lowest in ingress boundary plots. Regression analyses indicate pupal recovery rate decreased as a function of carabid abundance as well as the abundance of non-insect ground predators. To determine if ground cover management influenced natural enemy abundance, aisles were clean cultivated or planted with three ground covers (clover, ryegrass, or buckwheat). Increasing ground cover had a significant effect on the relative abundance of Harpalus pensylvanicus De Geer (Coleoptera: Carabidae). In addition to conserving natural enemies for control of blueberry insect pests, we discuss additional benefits of ground covers that may increase their utility for blueberry production.     

The three-dimensional macronutrient niche of an invasive generalist predator

1. Harvestmen are predators, although they suspected of being partly omnivorous by including fruits in their diet. 2. In the present study, the right-angled mixture triangle (RMT) was used to analyse the macronutritional niche of the harvestman Opilio canestrinii, an invasive species in northern Europe. The study design followed a double-test procedure in which the animals were subjected to two self-selection tests: the first one immediately after being caught in the field and the second after 1 week of ad libitum feeding during which the animals became satiated and nutritionally balanced. A comparison of results from the two tests indicates whether the animals were food limited in the field, and whether they were limited by a particular macronutrient. 3. Females were found to be food limited in the field, whereas males were not. Opilio canestrinii had a target intake of 28% lipid : 52% protein : 20% carbohydrate [i.e. with a considerable proportion of carbohydrate (sugar) in the diet]. Both sexes were non-protein limited in the field, with sugar being more limiting than lipid. 4. The results indicate considerable inclusion of plant-derived sugar in the natural diet. This conclusion was supported by a separate experiment showing enhanced performance (survival, gain in mass) in animals whose only energy supply was fresh fruit. 5. It is concluded that the harvestman is best characterized as an omnivorous predator.     

Regional distribution patterns can predict the local habitat specialization of arachnids in heterogeneous landscapes of the Atlantic Forest

Aim

This study formally evaluates the ability of three models to use geographical data on species distribution to predict the habitat use patterns of species in heterogeneous landscapes.

Location

Species and habitats in the Brazilian Atlantic Rain Forest were investigated.

Methods

Based on empirical data on harvestmen and scorpions, we estimated the strength of species association with preferred habitat and classified them as habitat generalists or habitat specialists. We compared these empirical results with predictions made using data on species range size (model 1), species occurrence in biomes (model 2) and species occurrence in habitats within the biomes (model 3).

Results

We used 1,278 records of eight harvestman and two scorpion species that had specific determination and enough sampling numbers to allow safe identification of habitat specialization. We observed the following: (1) the extension of species occurrence did not influence the strength of species–habitat association (estimated by IndVal), which led us to reject model 1; (2) species habitat specialization derived from occurrences in biomes was 60% coincident with the classification derived from empirical data. This value is not different enough from the value expected by chance for these data, which also led us to reject model 2; and (3) species classification derived from secondary data about the habitats used had a significant coincidence of 80% with the empirical classification, which led us to accept model 3.

Main conclusions

For correct classification of species habitat specialization using secondary distributional data, we recommend that future studies consider using the most accurate information available on the habitats used by species. Especially for megadiverse and understudied groups, information about habitats used is not easy to obtain, so it is important for researchers and institutions to register and disseminate this information, which could support many other studies.

     

Experimental test of the mechanism underlying sexual segregation at communal roosts of harvestmen (Prionostemma spp.)

Closely related, ecologically similar species often roost in distinctly different habitats, and roosting patterns also vary within species in relation to sex, age and season. The causes of such variation are not well understood at either a proximate or ultimate level. We studied communal roosting in two congeneric species of Prionostemma harvestmen at a rainforest site in Nicaragua. Previous research showed that Prionostemma sp. 1 forms male‐biased communal roosts in tree‐root cavities, while Prionostemma sp. 2 forms communal roosts of variable but temporally stable sex ratios on spiny palms. Here, we investigate potential mechanisms underlying variation in roosting site choice between and within these syntopic species. First, we present the results of a field experiment designed to probe the mechanism underlying skewed roost sex ratios in Prionostemma sp. 2. Previous studies have suggested that these harvestmen use conspecific scent to locate communal roosts and that new roosts can be established via group translocation. Therefore, to test the hypothesis that skewed roost sex ratios in this species arise from sex differences in scent marks, we translocated single‐sex groups of ca. 30 individuals to each of 20 previously unoccupied spiny palms. Female release sites attracted new recruits of both sexes, while male release sites attracted almost exclusively males. We infer that Prionostemma sp. 2 females preferentially roost in sites scent‐marked by females and that this mechanism is sufficient to explain the skewed roost ratios. Further adding to knowledge of Prionostemma roosting behavior, we show that Prionostemma sp. 1 forms female‐biased communal roosts on spiny palms, that some roosts contain both species, and that the species composition is stable on a time scale of at least 2 weeks. To the best of our knowledge, this study is the first experimental test of mechanisms underlying sexual segregation at communal roosts in any taxon.     

Comparative performance of double‐digest RAD sequencing across divergent arachnid lineages

Next‐generation sequencing technologies now allow researchers of non‐model systems to perform genome‐based studies without the requirement of a (often unavailable) closely related genomic reference. We evaluated the role of restriction endonuclease (RE) selection in double‐digest restriction‐site‐associated DNA sequencing (ddRADseq) by generating reduced representation genome‐wide data using four different RE combinations. Our expectation was that RE selections targeting longer, more complex restriction sites would recover fewer loci than RE with shorter, less complex sites. We sequenced a diverse sample of non‐model arachnids, including five congeneric pairs of harvestmen (Opiliones) and four pairs of spiders (Araneae). Sample pairs consisted of either conspecifics or closely related congeneric taxa, and in total 26 sample pair analyses were tested. Sequence demultiplexing, read clustering and variant calling were performed in the pyRAD program. The 6‐base pair cutter EcoRI combined with methylated site‐specific 4‐base pair cutter MspI produced, on average, the greatest numbers of intra‐individual loci and shared loci per sample pair. As expected, the number of shared loci recovered for a sample pair covaried with the degree of genetic divergence, estimated with cytochrome oxidase I sequences, although this relationship was non‐linear. Our comparative results will prove useful in guiding protocol selection for ddRADseq experiments on many arachnid taxa where reference genomes, even from closely related species, are unavailable.