Cosmetophilus vonones, N. G., N. Sp., (Apicomplexa: Actinocephalidae) in the Harvestman Vonones sayi (Arachnida: Cosmetidae)

The gregarine Cosmetophilus vonones , n. g., n. sp. (Sporozoasida: Actinocephalidae: Acanthosporinae) from the intestine and intestinal caeca of the harvestman Vonones sayi (Simon) (Opiliones: Cosmetidae) is described. The new internal parasite is the first recorded from a harvestman in the New World and the second from the opilion suborder Laniatores. In addition to the records from the type locality in western Texas, additional collections are reported from the same host in central and eastern Texas and Tennessee.     

Hedgerows support rich communities of harvestmen (Opiliones) in upland agricultural landscape

Hedgerows play important roles in agricultural landscapes and they increase biodiversity by providing habitat refugia for species sensitive to agricultural disturbance. We have studied the characteristics of the communities of harvestmen (Opiliones) inhabiting hedgerows. Harvestmen are terrestrial arachnids associated with soil surface and subsurface and sensitive to land-use and disturbance. We were specifically interested in quantifying how hedgerow characteristics (e.g., elevation, length, width, connectivity, and plant diversity) affect harvestman diversity and community composition. We expected harvestman diversity and community composition to be positively related to both hedgerow size (area, length, width) and biological attributes of hedgerows (e.g., connectivity, plant community diversity). We surveyed hedgerow characteristics and harvestman communities of 20 hedgerows in an upland agricultural landscape in the Western Carpathians. Hedgerow characteristics were measured in the field or derived from GIS layers and we used correlation and ordination methods to relate them to harvestman community metrics. We found surprisingly high taxonomic richness of harvestmen within the studied hedgerows (15 species). Importantly, the Shannon index of harvestman communities was positively related to hedgerow length and hedgerow tree layer species richness and diversity. Harvestman community composition varied with hedgerow area and width. Despite their small total area, hedgerows represented an important habitat for diverse harvestman communities and hedgerow attributes such as size and tree diversity significantly affected the composition and the Shannon index of harvestman communities. Thus, greater lengths and widths of hedgerows can provide habitats for a higher Shannon index of harvestman communities, within surrounding agricultural landscapes.     

Harvestman (Opiliones) fauna associated with Maine lowbush blueberry fields in the major production areas of Washington and Hancock counties

Over a period of 19 yr, the harvestman (Opiliones) community associated with the lowbush blueberry agro-ecosystem in Maine was studied. Eight species representing five genera, four subfamilies, and two families of harvestmen belonging to the suborder Eupnoi were collected. The harvestman community was dominated by two introduced, synanthropic species: Phalangium opilio in all but 1 yr (that year dominated by Rilaena triangularis). Rilaena was recorded for the first time from eastern North America. Relative abundance of harvestman adults increases throughout the season and the temporal pattern of trap capture does not refute speculated life cycles of the harvestmen being univoltine with overwintering eggs. Some blueberry management practices were found to affect trap capture. We did find that on average (with opposite results 1 yr) trap captures are greater in pruned fields than in fruit-bearing fields. Organic fields were found to have higher relative abundance of harvestmen than conventionally managed fields. Conventionally managed fields with reduced-risk insecticides showed no difference in harvestmen relative abundance compared with those conventionally managed fields using the older more persistent organophosphate insecticides. Insecticide trials with common insecticides used in blueberry insect pest management showed that the organophosphate insecticide, phosmet, and the pyrethroid insecticide, esfenvalerate, were detrimental to P. opilio adults when exposed to leaf residues, whereas the reduced-risk insecticide, spinosad, showed no negative effects compared with nonsprayed foliage.     

The cost of living slowly: metabolism,Q10 and repeatability in a South American harvestman

Abstract Resources are usually considered to be limited in caves and underground habitats. Therefore, lower metabolic rates of ectotherms in these environments should be advantageous. The standard metabolic rate (SMR) of the common harvestman Pachylus paessleri Roewer, 1913 (Arachnida, Opiliones) is determined at two ambient temperatures. Repeatability of SMR is estimated by two methods: (i) product–moment correlation on residuals of body mass and (ii) variance components. Estimations of the thermal sensitivity of metabolic rate (individual Q₁₀). SMR of P. paessleri at 20 °C show a mean of 19.01 ± 6.04 and 14.99 ± 3.64 μL CO₂ h^?1 for males and females, respectively. Thermal sensitivity of SMR is 2.11 ± 0.23 with a substantial coefficient of variation of 26.7%. In males, residuals of CO₂ release are significantly repeatable (r_p = 0.61) between measurements realized 5 months apart, which reflects the consistency of this trait over time. As typical soil inhabitants, the harvestman P. paessleri present a lower metabolic rate in comparison with arthropods of similar body mass (e.g. arachnids). Although the coefficient of thermal sensitivity is within of the range reported for arthropods inhabiting Mediterranean zones, it is lower than expected for an organism with such a low SMR. This appears to be the first report of repeatability of metabolic rate in a harvestman species.     

Epigean communities of harvestmen (Opiliones) in Pannonian Basin floodplain forests: an interaction with environmental parameters

During the years 2015–2016, we studied the fauna of epigeon in eight similar habitats in Slovakia and Serbia. The study material of Opiliones was obtained by a pitfall trap method. We focused on the comparison of the incidence of harvestmen in floodplain forests and ecotones alongside the Danube River in Slovakia and along the Tisa and Begej River in Serbia. We examined their dependence on anthropogenic impact (frequency of visits), cover of vegetation layers (herbs, shrubs and trees), plant diversity of vegetation layers, area of forest stands, distance to the forest edge, circumference of forest stands, age of forest stands, thickness of litter layer and physico-chemical properties of soil and litter (pH, conductivity, P, N, C, H). Total epigeic activity of harvestman showed significant positive relationship with proportion of N in litter. Community equitability was positively related with species richness of plant communities in shrub layer and negatively related with H content of soil. We did not find any significant correlations between diversity of harvestman communities and measured environmental characteristics.     

Influence of agricultural management form on the species structure of harvestman (Opiliones) communities

The paper is aimed at evaluation of the influence of two different agricultural management forms on harvestman (Opiliones) communities structure in relation to the optimization of land use on the model territory of Agricultural Cooperative (AC) Očová (Central Slovakia). The research was carried out using pitfall trapping through growing seasons from 2005 to 2007. Harvestmen were captured at four pairs of sites (8 sites in total). For each pair of sites the cultivation of the same crop with application of two different forms of agricultural management (conventional form and sustainable form with basic sustainable agro-environmental scheme) was characteristic. In total, 667 individuals of 8 species of the Phalangiidae family were captured during the research. The research results confirm the influence of management form on the structure of harvestman communities only partially. The obtained data indicate that the management form plays a significant role, especially in such agrocoenoses, in which the same crop is cultivated for several years (e.g., sites with alfalfa or permanent lawn), i.e., on the sites without ploughing or use of other similar agrotechnical measures. In these cases, the sites with the sustainable agro-environmental scheme showed higher diversity and equability of harvestman communities than the sites with the conventional form of agrotechnical management. Harvestman communities structure on sites with crop rotation and repeated ploughing was more influenced by additional factors than by the agricultural management form.     

Microstructural organization of the central nervous system in the harvestman Leiobunum japonicum (Arachnida: Opiliones)

Although the order Opiliones constitutes the third‐largest group of arachnids, this creature is still mysterious and has a rich unexplored field compared to what is known about insects and crustaceans. The order Opiliones is traditionally regarded as a close relative of mites, mainly because of morphological similarities in external body structure; however microstructural organization of the ganglionic neurons and nerves in the harvestman Leiobunum japonicum is quite similar to the central nervous system (CNS) in all extant arachnids. The CNS consists of a large neural cluster with paired appendicular nerves. The esophagus passes through the neural cluster and divides it into the upper supraesophageal ganglion (SpG) and the lower subesophageal ganglion (SbG). The dorsal part of the SpG has a quite condensed cell body compared with other parts of the CNS and has two main components, the protocerebrum and the cheliceral ganglion. The protocerebrum receives the optic nerves and has four main groups of neuropiles from the optic lobes, the superior central body, the lateral neuropils (corpora pedunculata) and the inferior neuropil. However, a pair of pedipalpal and four pairs of appendage nerves including several pairs of abdominal nerves arise from the nerve masses of the SbG.     

Morphology and Meiotic/Mitotic Behavior of B Cromosomes in a Japanese Harvestman, Metagagrella tenuipes (Arachnida: Opiliones): No Evidence for B Accumulation Mechanisms

Earlier, it has been demonstrated that wild populations of a Japanese harvestman Metagagrella tenuipes (Arachnida: Opiliones) are polymorphic for B chromosomes. In this paper, we present results of a study of the morphology and mitotic and meiotic behavior of the Bs. The B chromosomes varied considerably in size and proportion of eu- and heterochromatin. The single nucleolus organizing region, found in males, was located on a chromosome of the A complement. Some intercell variation in number of Bs may be explained by accidental chromosome losses during chromosome preparation. We also found no intertissue variation in number of Bs. There were also no differences in mean number of B chromosomes per individual among males and females, adult and subadult harvestmen. Segregation of Bs in mitotic and meiotic divisions was nonrandom; B chromosomes tended to segregate equally between daughter cells. The results obtained provide no support for the hypothesis of existence of B accumulation mechanism in this species.     

Two new species of harvestmen (Opiliones,Eupnoi, Neopilionidae) from Waitomo,New Zealand

Two new species of harvestman (Opiliones: Neopilionidae: Enantiobuninae) are described from the Waitomo region of the North Island, New Zealand, Forsteropsalis bona sp. n. and F. photophaga sp. n. Both have been collected within caves in the region, where predation on glow-worms Arachnocampa luminosa has been previously recorded for one or both species (misidentified as ‘Megalopsalis tumida’). However, both are regarded as troglophiles rather than strict troglobites due to the presence of specimens outside the cave systems, and the absence of troglobitic adaptations. Megalopsalis tumida (Forster, 1944) is identified as a junior synonym of Forsteropsalis fabulosa (Phillipps & Grimmett, 1932).     

A relict in New Caledonia: phylogenetic relationships of the family Troglosironidae (Opiliones: Cyphophthalmi)

The species richness and endemism of New Caledonia are traditionally held to result from the main island's Gondwanan origin and progressive diversification subsequent to extended isolation. Recent studies have challenged this hypothesis, promoting a scenario of recent origins and diversifications of New Caledonian arthropod groups. In the present study, the phylogeny of the endemic harvestman family Troglosironidae (Opiliones: Cyphophthalmi) is investigated using DNA sequence data from two nuclear ribosomal genes (18S rRNA and 28S rRNA) and two mitochondrial genes (the protein-coding cytochrome c oxidase subunit I and the ribosomal 16S rRNA). Phylogenetic analyses support the monophyly of Troglosironidae and a scenario of an ancient (> 200 Ma) origin of the family, with subsequent diversification of extant lineages in the Eocene. These results corroborate the relictual nature of taxa among New Caledonia's biota while being consistent with diversification in accordance with geological events in the Eocene.     

Sperm morphology of the neotropical harvestman Iporangaia pustulosa (Arachnida: Opiliones): comparative morphology and functional aspects

We describe herein the sperm morphology of the harvestman Iporangaia pustulosa. Adult males were dissected, the reproductive tract was schematized and the seminal vesicle was processed by light, transmission and scanning electron microscopy. The male reproductive tract is composed of a tubular testis, two deferent ducts, a seminal vesicle, a propulsive organ and a penis, similar to that observed in other Opiliones. The spermatozoa from the seminal vesicle are oval, aflagellate and immotile, presenting a nucleus surrounding an invagination of the cytoplasm, as well as a complex acrosome and projections on the cell surface. In the testis, spermatozoa are devoid of projections. In the seminal vesicle, they gradually acquire the projections with tufts adhering to it. Consequently, spermatozoa in various distinct stages of projection development can be found in the seminal vesicle. We believe that these projections (1) could help transport sperm along the male and perhaps female reproductive tracts; (2) are used to anchor the spermatozoa inside the female spermatheca in order to avoid mechanical displacement by the genitalia of other males and (3) may play a role in oocyte recognition. We propose that the evolution of aflagellarity in Opiliones is related to the unique morphology of the female reproductive tract. Since eggs are fertilized on the tip of the ovipositor just prior to being laid, there is no advantage favoring sperm mobility. Additionally, female sperm receptacles are small and males that produced small spermatozoa would have a higher chance of fertilizing more eggs.     

Harvestman (Arachnida: Opiliones) species distribution along three Neotropical elevational gradients: an alternative rescue effect to explain Rapoport's rule?

Aim Relationships between elevation and litter‐dweller harvestman (Arachnida: Opiliones) species richness along three elevational gradients in the Brazilian Atlantic Forest were evaluated. Specifically, three candidate explanatory factors for the observed patterns were tested: (1) the mid‐domain effect, (2) the Rapoport effect, and (3) the influence of environmental variables on species density and specimen abundance. Location Cuscuzeiro, Corcovado and Capricórnio mountains, in Ubatuba (23°26′ S, 45°04′ W), a coastal municipality in São Paulo state, south‐eastern Brazil. Methods We recorded harvestman species and abundance through active sampling using 8 × 8‐m plots in both summer and winter. At each plot we measured the temperature, humidity and mean litter depth. Harvestman species richness per elevational band was the sum of all species recorded in each band, plus the species supposed to occur due to the interpolation of the upper and lower elevational records. Differences between observed and expected species richness per elevational band, based on the mid‐domain effect, were examined through a Monte Carlo simulation. The Rapoport effect was evaluated using both the midpoint method and a new procedure proposed here, the ‘specimen method’. We applied multiple regression analysis to evaluate the contribution of each environmental variable (elevation, temperature, humidity and litter depth) on species density and specimen abundance per plot. Results Harvestman abundance and species richness decreased at higher elevations in the three mountains. The decrease in species richness was not monotonic and showed a plateau of high species richness at lower elevations. The number of harvestman species per elevational band does not fit that predicted by the mid‐domain effect based solely on geometric constraints assuming hard boundaries. Species with their midpoints at higher elevations tended to cover broader elevational range sizes. Both the midpoint method and the specimen method detected evidence of the Rapoport effect in the data. At fine spatial scales, temperature and humidity had positive effects on species density and specimen abundance, while mean litter depth had no clear effect. These relationships, however, were not constant between seasons. Main conclusions Our results suggest that harvestman species density declines at higher elevations due to restrictions imposed by temperature and humidity. We found a pattern in species range distribution as predicted by the elevational Rapoport effect. However, the usual rescue effect proposed to explain the Rapoport effect does not apply in our study. Since the majority of harvestman species covering broader elevational ranges do not exhibit reduced abundance at low elevations, an alternative rescue effect is proposed here. According to this alternative rescue effect, the decrease in species richness at higher elevations occurs due to differential upper limits of species with source populations below mid‐elevations. The seasonal differences in the relationships between environmental variables and species richness/specimen abundance per plot is an indication that species occurrence on elevational gradients is seasonally dependent. Thus relationships and hypotheses based on data recorded over short time periods, or in a single season, should be viewed cautiously.     

No regulatory role for adult predation in cyclic population dynamics of the autumnal moth,Epirrita autumnata

1. Multiannual population cycles of geometrid moths are thought to be driven by trophic‐level interactions involving a delayed density‐dependent component. Predation on adult moths has been a little‐studied mechanism of this phenomenon. 2. Using 29 daytime and 61 night‐time predation trials in the field, we exposed living autumnal moth (Epirrita autumnata Borkhausen, Lepidoptera: Geometridae) females to their natural predators during each autumn throughout the 10‐year population cycle. 3. In our northern study location (70°N), insectivorous passerines had already migrated, and harvestman Mitopus morio Fabricius (Opiliones: Phalangiidae) was found to be the main predator of the adult moths. The predation mortality occurred mainly at night and was positively correlated with the minimum temperature measured during the predation trial. 4. Despite high annual variability in the degree of adult predation, both direct and delayed density‐dependent effects were weak and indicate that predation on adult moths in the autumn does not have any regulatory role in cyclic population dynamics of the autumnal moth in northern Fennoscandia.     

Ecology of Arachnida alien to Europe

The ecology of selected species of spiders (Araneae), harvestmen (Opiliones), and the neglected tropical order Schizomida, alien to Europe, is discussed. Their geographic origins and pathways of introduction, by transportation with goods, are similar to other predatory terrestrial arthropods. Occurrence in buildings (synanthropy) is a prerequisite for range expansion in many arachnid species. Thelytokous parthenogenesis facilitated spread of a few spider, harvestman and schizomid species outside their native range. Both prey generalists and specialists (woodlice-eating Dysdera crocata and ant-eating Zodarion rubidum) are among species successfully expanding their range. Alien arachnids include individual species belonging to many different families. We stress the absence of true invasiveness of alien arachnids in Europe. Some unusual adverse effects of alien arachnids were recorded, such as pollution of buildings and clogged car fuel system ventilation tubes. Frequency of painful biting and threat to native biota are low.     

Communal Roost Site Selection in a Neotropical Harvestman: Habitat Limitation vs. Tradition

Many species have been reported to form roosting (resting, sleeping) aggregations at ‘traditional’ sites, but the alternative hypothesis that specific sites are used repeatedly because of habitat limitation is rarely tested. We studied the roosting behavior of a species of harvestman (Opiliones, Prionostemma sp.) at a lowland rainforest site in Nicaragua. Both sexes roosted by day in spiny palm trees, dispersed at dusk to forage, and rejoined aggregations just before dawn. The distribution of harvestmen among spiny palms was significantly clumped, and harvestman density did not correlate with spiny palm density. Aggregations formed repeatedly in a small subset of the available spiny palms and the same sites were used in two different years (2001, 2003). Nevertheless, the membership of aggregations was fluid; individual harvestmen were found at multiple roosts and moved up to 0.2 km per night. Translocated animals often returned to the roost where they had been released or nearby roosts but were never found at previously unused sites. The high consistency of site use but low site fidelity of individuals suggests that roost sites differed conspicuously (to the harvestmen) from sites that were not used. We found no univariate or multivariate differences between used and unused sites, however, in the characteristics of the trees or microclimate. These results conflict with the habitat limitation hypothesis but are consistent with the traditional site use hypothesis. The tradition may be mediated by a site‐labeling chemical, a mechanism that does not require individual site fidelity. We discuss these results in relation to the proposed functions of roosting aggregations.     

Polyphyly of Caddoidea,reinstatement of the family Acropsopilionidae in Dyspnoi,and a revised classification system of Palpatores (Arachnida,Opiliones)

Among the least studied harvestmen are the members of the family Caddidae sensu Shear, 1975 , a group of Opiliones with massive eyes and the putative sister group of the remaining Eupnoi. Caddids were originally described as two families, Caddidae and Acropsopilionidae, but these are currently treated as subfamilies of Caddidae. These minute arachnids are rarely collected and present some interesting biogeographical patterns, including a disjunct distribution between East Asia and eastern North America, and some of the few cases of trans‐Pacific genera in southern hemisphere Opiliones. We therefore obtained samples from most of the landmasses inhabited by Caddidae and undertook a phylogenetic study using nuclear and mitochondrial genes for as many samples as possible. Our results, based on a broad taxonomic sampling, surprisingly showed polyphyly of Caddidae, with the genus Caddo forming the sister group of the remaining Eupnoi, whereas the southern hemisphere genera, many of which were originally placed in Acropsopilionidae, within Dyspnoi, formed the sister clade of the remaining Dyspnoi. In addition, the more recently described genus Hesperopilio, from Western Australia and Chile, was unrelated to either Caddidae or Acropsopilionidae, despite having the supposedly diagnostic large ocularium, and instead appeared deeply nested within the Eupnoi superfamily Phalangioidea. Our results are robust to analytical treatment and to homology scheme (dynamic vs. static notions of homology), resulting in a new phylogenetic proposal for Eupnoi and Dyspnoi. Ancestral state reconstruction suggests that the ancestral Palpatores was probably a tiny harvestman with an enlarged ocularium and glandular palpal setae in its enlarged and armed palps. We take the following taxonomic actions: Acropsopilionidae is removed from synonymy under Caddidae and its family status and membership in Dyspnoi are restored. Hesperopilio Shear, 1996 is removed from Caddoidea/Caddidae and transferred to Phalangioidea, but it is not assigned to any family.     

A morphometrics‐based phylogeny of the temperate Gondwanan mite harvestmen (Opiliones,Cyphophthalmi, Pettalidae)

A phylogenetic estimation of the temperate Gondwanan mite harvestman family Pettalidae (Arachnida, Opiliones, Cyphophthalmi) was conducted using 143 morphological variables (59 raw and 84 scaled measurements) from 37 ingroup and 15 outgroup terminals. We used custom algorithms to do pairwise comparisons between characters and identify sets of dependent characters, which were collapsed using principal components analysis. We analysed the resulting data without discretization under the parsimony criterion. Monophyly or paraphyly of most groups suspected from previous molecular and morphological phylogenetic studies were recovered. Trees were optimized for monophyly of 20 different focus clades by varying character phylogenetic independence. This yielded a final tree with monophyly of 15 out of 20 focus clades, including the South African pettalids, which contains the troglomorphic species Speleosiro argasiformis Lawrence, 1931. Two of the remaining five clades were found paraphyletic, with the genera Aoraki, Rakaia, and Siro always being found polyphyeletic.     

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.     

Skeletomuscular anatomy of the harvestman Leiobunum aldrichi (Weed, 1893) (Arachnida: Opiliones: Palpatores) and its evolutionary significance

Skeletal muscles of the North American harvestman Leiobunum aldrichi are exhaustively surveyed and compared with other chelicerates to clarify the evolutionary morphology and phylogenetic relationships of arachnids. Representatives of 104 muscle groups are described and illustrated, and their possible functions are proposed. Comparisons of the feeding apparatus of L. aldrichi with that of other opilions, especially Sim (Cyphophthalmi) and Acromares (Laniatores), and two scorpion genera ( Centruroides, Pandinus ) indicate that the pharyngeal apparatus in L. aldrichi is derived and that its ability to accommodate large food particles is a secondary rather than primitive condition. Comparisons reveal several possible synapomorphies between Opiliones and Scorpiones suggesting that these orders may be sister groups. Apparently unique synapomorphies include an extrinsic cheliceral muscle that arises from the carapace and inserts on the second cheliceral article (deutomerite); an epistome divided into distal and proximal parts by a transverse sulcus; pharyngeal dilator muscles supported by a peripharyngeal skeleton formed by one dorsomedial and two ventrolateral epistomal processes, the latter also with muscular attachments to the endosternite; a specialized preoral chamber (stomodieca) derived from extensions (coxapophyses) of the coxae of the pedipalp and first two leg pairs; internal processes associated with the coxapophyses that serve, in part, as an attachment for muscles operating the coxa-trochanter joints, and lateral endosternal suspensor muscles that insert on the arthrodial membrane between the leg coxae. These are the first observations providing explicit support for an Opiliones-Scorpiones clade.