Failed predation or transportation? Causes and consequences of phoretic behavior in the pseudoscorpionDinocheirus arizonensis (Pseudoscorpionida: Chernetidae)

The pseudoscorpion Dinocheirus arizonensisinhabits rotting saguaro cactus in the Sonoran Desert and has also been found attached to the legs of the cactophilic neriid fly, Odontoloxozus longicornis.Laboratory experiments demonstrated a higher incidence of phoresy on eclosing versus postteneral adult flies, a female bias in phoresy, and an increased rate in female phoresy through time. The pseudoscorpion may also prey on the fly, but predation rate was unaffected by fly category, pseudoscorpion gender, or food deprivation. A study of pseudoscorpion colonization in the field indicated that females were the first to colonize and the first to abandon the transient habitat of a saguaro rot and, thus, corroborated patterns of phoretic behavior in the laboratory. Taken together, these results establish that phoresy is a behavior functioning specifically for dispersal. The hypothesis that pseudoscorpion transport by other arthropods is accidental, motivated by hunger, and occurs because pseudoscorpions are incapable of consuming their hosts is rejected.     

<Emphasis Type="Italic">Roncus travuniensis</Emphasis> sp. n. (Neobisiidae,Pseudoscorpiones), a troglobitic false scorpion from Bosnia-Herzegovina

A new pseudoscorpion species, Roncus travuniensis sp. n. (Neobisiidae, Pseudoscorpiones), was collected in the Arenstorfova Pećina Cave, near Trebinje, Bosnia and Herzegovina. The main morphological traits of the specimen studied clearly demonstrate its affinities with Roncus vulcanius Beier, 1938. Additionally, some taxonomic and biogeographical traits of this new pseudoscorpion are also discussed.     

Soil and canopy Pseudoscorpiones (Arthropoda,Arachnida) in a monodominant forest of Attalea phalerata Mart. (Arecaceae) in the Brazilian Pantanal

We studied the occurrence of Pseudoscorpiones in the soil, leaf litter, and in canopies of a monodominant forest of Attalea phalerata at different seasons in the northern region of the Brazilian Pantanal. A total of 1197 pseudoscorpions from nine families and 16 species were sampled. Olpiidae, Chernetidae, and Geogarypidae predominated in soil and leaf litter. Chernetidae was the most abundant family in canopies. Soil and canopy corresponded to distinct habitats in relation to pseudoscorpion abundance and richness, with the canopies being the most diversified environment. These habitats are occupied in different ways by pseudoscorpion populations. Geogarypus sp. occurs in the edaphic environment during receding water and dry season, but can be found in canopies of A. phalerata exclusively during high water. This alternation in the use of the edaphic environments and canopies in the same area by pseudoscorpion species probably happens due to the strong seasonality of the Brazilian Pantanal.     

Predation ofApochthonius minimus (Pseudoscorpionida: Chthoniidae) onFolsomia canadida (Collembola: Isotomidae)

Summary We investigated the functional response of the predaceous pseudoscorpion,Apochthonius minimus, to density of the springtail,Folsomia candida, as well as the effects of temperature and prey size on handling time and attack rate. Temperature slightly altered the overall rise in response. Both the rise and shape of the response differed for two prey sizes. The changes in number of prey eaten over a 48-hour period were also examined, and the attack and feeding behaviour ofA. minimus was described and discussed in relation to the predation experiments. The pseudoscorpion preferred young adultF. candida, 0.8–1.2 mm in length, when offered prey in the range, 0.2–1.5 mm.     

Identification key to the species of Amblyolpium Simon (Pseudoscorpiones: Garypinidae), with first record of Amblyolpium bellum Chamberlin from Iran

A key based on bibliographic sources is given for identifying the 15 extant known pseudoscorpion species of the genus Amblyolpium recorded from different localities throughout the world. Species of Amblyolpium mostly occur in the Mediterranean basin, the Near to Far East, and the genus is only recorded with one species from South America. Amblyolpium bellum is redescribed and illustrated based on four males and two females collected for the first time from southern Iran.     

<Emphasis Type="Italic">Neobisium babinzub</Emphasis> sp. n. and <Emphasis Type="Italic">Neobisium tarae</Emphasis> sp. n. (Neobisiidae,Pseudoscorpiones) from the central Balkan Peninsula (Serbia)

Two new pseudoscorpion species, Neobisium babinzub sp. n., and N. tarae sp. n. (Neobisiidae, Pseudoscorpiones), were collected in soil and leaf-litter on Mt. Stara Planina and Mt. Tara, Serbia, diagnosed and thoroughly illustrated. The main morphological characteristics of the two taxa clearly demonstrate their adaptation to life in soil.     

Prozercon iranensis sp. n., a new species of Zerconidae (Acari: Mesostigmata) from Iran

A new species of pseudoscorpion of the genus Calocheiridius Beier &; Turk, Calocheiridius iranicus n. sp., is described from Markazi province, Iran. Illustrations and measurements of the species are given. http://www.zoobank.org/urn:lsid:zoobank.org:pub:06332B5C-D43A-4955-AB6C- 5847D10C2BAC     

Phylogeography of the giant harlequin beetle (Acrocinus longimanus)

Aim The extent to which cryptic species contribute to neotropical diversity remains inadequately investigated. Based on its highly distinctive morphology, the giant harlequin beetle, Acrocinus longimanus, is currently described as a single species, ranging from southern Mexico to northern Argentina. However, the discovery of cryptic species in Cordylochernes scorpioides, a pseudoscorpion with obligate dependence on the harlequin beetle for dispersal, strongly suggests the existence of barriers to gene flow in A. longimanus. The aim of this study was therefore to determine whether levels of DNA divergence between geographical populations provided evidence of genetically distinct lineages in the harlequin beetle. Location Trinidad and Panamá. Methods Sequencing of 1245 bp of the mitochondrial cytochrome oxidase I (COI) gene of A. longimanus from seven locations in Trinidad and Panamá. Results Mitochondrial haplotype diversity in the harlequin beetle shows limited evidence of geographical structuring, with a maximum sequence divergence between populations of only 1.29%. This is an order of magnitude less than the level of COI divergence between harlequin beetle riding pseudoscorpions from the same geographical locations. Main conclusions The molecular data on populations from northern South America and Panamá are consistent with the current, morphologically based classification of A. longimanus as a single, pan‐neotropical species. In addition, the relatively low level of population divergence detected in this study indicates that speciation in the hitchhiking pseudoscorpion has occurred in the absence of significant barriers to gene flow in its beetle host. It is proposed that, in the harlequin beetle, the phylogenetic signal of colonization and vicariance associated with the formation of the Isthmus of Panamá has been obscured, although not fully erased, by historical and contemporary gene flow.     

Association of pseudoscorpions with different types of bird nests

The hypothesis of associating pseudoscorpions with bird nest types was tested on the basis of an analysis of 480 specimens. Eleven pseudoscorpion species were found in 171 nests of 28 different bird species collected in Slovakia, Austria and the Czech Republic. The frequent appearance of Cheiridium museorum, Dactylochelifer latreillii, Chernes hahnii, Dendrochernes cyrneus and Allochernes wideri was confirmed. High proportion and association of Pselaphochernes scorpioides in hoopoe hollow nests with decomposed substrate, D. cyrneus in the Eurasian tree sparrow nest boxes and A. wideri in the nests of the tawny owls, the European scops owls and the European roller was proved. In contrast, C. hahnii and D. latreillii were related to the nest fauna of blackbirds and song thrushes, C. museorum to the nests of white wagtails situated on the ground and on buildings and C. cancroides to the nests in synanthropic habitats. Until present, the occurrence of 22 pseudoscorpion species has been confirmed in the bird nests of Central Europe based on the obtained results and published resources. According to the association to bird nests these pseudoscorpions were classified as (1) nidixenous species — Chthonius fuscimanus, C. tetrachelatus, Mundochthonius styriacus, Neobisium carcinoides, N. crassifemoratum, N. inaequale, N. sylvaticum, Chernes cimicoides, C. similis, C. vicinus, Allochernes powelli, Lamprochernes chyzeri, L. nodosus and Larca lata; and (2) nidiphilous species — C. museorum, C. cancroides, A. wideri, D. cyrneus, D. latreillii, C. hahnii, D. panzeri and P. scorpioides.     

Springtail postmolt vulnerability to pseudoscorpion predation: Mechanisms and implications

Arthropod prey are expected to be more vulnerable to their predators immediately following molt. The effects of springtail (Isotoma carpenteri) postmolt vulnerability on interactions with a pseudoscorpion predator were examined in the laboratory. Springtails exposed to vials pretreated with pseudoscorpions (Apochthonius minimus) delayed molting compared to those prey that were exposed to vials pretreated only with springtails. Although their escape ability (measured as distance jumped) was unaffected by molt condition, postmolt springtails were more profitable in terms of reduced predator handling time following capture. Despite this,A. minimus did not distinguish between postmolt and intermolt prey presented at either end of a T-maze.     

Adaptations for matrotrophy in the female reproductive system in the pseudoscorpion Chelifer cancroides (Chelicerata: Pseudoscorpiones,Cheliferidae)

Pseudoscorpiones (pseudoscorpions, false scorpions) is an order of small terrestrial chelicerates. While most chelicerates are lecithotrophic, that is, embryos develop due to nutrients (mostly yolk) deposited in the oocyte cytoplasm, pseudoscorpions are matrotrophic, that is, embryos are nourished by the female. Pseudoscorpion oocytes contain only a small amount of yolk. The embryos develop within a brood sac carried on the abdominal site of the female and absorb nutrients by a pumping organ. It is believed that in pseudoscorpions nutrients for developing embryos are produced in the ovary during a postovulatory (secretory) phase of the ovarian cycle. The goal of our study was to analyze the structure of the female reproductive system during the secretory phase in the pseudoscorpion Chelifer cancroides, a representative of the family Cheliferidae, considered to be one of the most advanced pseudoscorpion taxa. We use diverse microscopic techniques to document that the nutritive fluid is produced not only in the ovaries but also by the epithelial cells in the oviducts. The secretory active epithelial cells are hypertrophic and polyploid and release their content by fragmentation of apical parts. Our observations also indicate that fertilization occurs in the oviducts. Moreover, in contrast to previous findings, we show that secretion of the nutritive material starts when the fertilized oocytes reach the brood sac and thus precedes formation of the pumping organ. Summing up, we show that C. cancroides exhibits traits of advanced adaptations for matrotrophy due to coordinated secretion of the nutritive fluid by the ovarian and oviductal epithelial cells, which substantially increases the efficiency of nutritive fluid formation. Since the secretion of nutrients starts before formation of the pumping organ, we suggest that the embryos are able to absorb the nutritive fluid also in the early embryonic stages.     

WHEN MORPHOLOGY MISLEADS: INTERPOPULATION UNIFORMITY IN SEXUAL SELECTION MASKS GENETIC DIVERGENCE IN HARLEQUIN BEETLE-RIDING PSEUDOSCORPION POPULATIONS

Differences in secondary sexual characteristics of males often provide the most conspicuous means of distinguishing between closely related species. Does this therefore imply that the absence of differentiation in exaggerated male traits between allopatric populations provides evidence of a single, genetically cohesive species? We addressed this question with a comprehensive investigation of two populations (French Guiana and Panama) of the harlequin beetle-riding pseudoscorpion, Cordylochernes scorpioides. This highly sexually dimorphic pseudoscorpion is currently described as a single species, ranging throughout the Neotropics. Our morphometric analyses detected minimal differentiation between the two populations in all nine external morphological characters measured, including sexually dimorphic traits in males. Only in traits of the spermatophore was there any appreciable level of differentiation. Behavior differentiation and prezygotic reproductive isolation were also limited: 78.3% of males successfully transferred sperm to “foreign” females, and in 63.9% of these cases, females' eggs were successfully fertilized. By contrast, extensive divergence existed in two of nine electrophoretic loci, including an essentially fixed-allele difference at the Ldh locus. Most significantly, postzygotic reproductive isolation was complete, with heteropopulation zygotes invariably aborting early in development. These results strongly suggest that the two populations are, in fact, sibling species, a conclusion supported by our recently published findings on their marked divergence in minisatellite DNA. How can such interpopulation homogeneity in male sexually dimorphic traits exist in the face of strong genetic divergence? We propose that sexual selection, oscillating between favoring small and then large males, maintains such high levels of male variability within each population that it has obscured a speciation event in which genetic divergence and postzygotic incompatibility have clearly outpaced the evolution of prezygotic reproductive isolation.     

Mating Behavior and Spermatophore Morphology of the Cave Pseudoscorpion Maxchernes iporangae (Arachnida: Pseudoscorpiones: Chernetidae)

The mating behavior of the cave pseudoscorpion Maxchernes iporangae Mahnert & Andrade, 1998 (Chernetidae), was studied under laboratory conditions. A total of 24 pairings was observed. Mating was divided into three phases: predeposition (dance), spermatophore deposition and transfer, and posttransfer phase. The mean mating duration was 67.3 min based on 10 observations. Spermatophore morphology is described and consists of a stalk, a drop of fluid, and an -shaped sperm package. Males and females can mate several times in the same reproductive period. However, consecutive mating was not observed between a female and the same or another male.     

Description of a new pseudoscorpion species of the genus Amblyolpium Simon (Pseudoscorpiones: Garypinidae) from north-western Iran

A new species of pseudoscorpion, Amblyolpium atropatesi n. sp. is described and illustrated on the basis of the adult specimens extracted from soil by Berlese funnel.

http://www.zoobank.org/urn:lsid:zoobank.org:pub:0150B850-E80D-46F5-983C-7E958FE69B87     

The population metabolism and life history tactics of Neobisium muscorum (Leach) (Arachnida: Pseudoscorpiones)

Summary The population metabolism of N. muscorum at a beechwood site in southern England was determined for two separate year periods by two methods; a detailed computer analysis and a best estimate. These methods, which are based on data of biomass, population dynamics, field temperatures, and respiration rates of all life stages with temperature, are compared. The validity of extrapolating laboratory determined basal metabolic rates to field activity is discussed; and the population metabolism of N. muscorum is compared with that reported for other soil and litter invertebrate species, both predatory and non-predatory.The metabolic rates of the adults of a second species of pseudoscorpion found at the beechwood site (Chthonius orthodactylus (Leach)), were determined, and the life cycle and metabolism of this species is compared to N. muscorum. The influence of low environmental temperatures on poikilothermic predators, with reference to life cycle strategies, basal metabolism and geographic distribution, is discussed.     

Cooperation in the neotropical pseudoscorpion, <Emphasis Type="Italic">Paratemnoides nidificator</Emphasis> (Balzan, 1888): feeding and dispersal behavior

Social behavior is rare among arachnids, and “permanent-sociality” is a strategy documented only in a few spider species. Here we describe the feeding and dispersal behavior of Paratemnoides nidificator, a Neotropical pseudoscorpion presenting “non-territorial permanent-sociality”. Field and laboratory observations (100 hours and 200 sessions over four years) and manipulative experiments revealed that P. nidificator is a generalist predator that lives in rough bark of trees, in groups of 2 to 175 individuals (42.62 ± 28.27; N=35 colonies; solitary individuals were never found), including reproductive adults and nymphs. Individuals in groups cooperatively capture prey four times larger than solitary individuals. Adult hunters share food with siblings. Reproduction in natural conditions occurs during the hot/wet season, when food availability is high. Prey diversity and abundance seem to be the main factor limiting reproduction in this species. Phoresy (attachment to a transporter animal) is the basic mode of dispersion in P. nidificator. Grouped individuals use large animals as vectors for dispersal, and once established they use the vector as the first food item of the new colony. However, they can also disperse by colony fission, newly documented herein for this taxon. The elaborate, cooperative behavior in P. nidificator is unique among pseudoscorpions and represents an excellent model for studies on the evolution of sociality in invertebrates. Received 10 August 2006; revised 17 November 2006; accepted 23 January 2007.     

Thermal biology and immersion tolerance of the Beringian pseudoscorpion <Emphasis Type="Italic">Wyochernes asiaticus</Emphasis>

Wyochernes asiaticus (Arachnida: Pseudoscorpiones: Chernetidae) is a pseudoscorpion distributed across Beringia, the areas of Yukon, Alaska and Siberia that remained unglaciated at the last glacial maximum. Along with low temperatures, its streamside habitat suggests that submergence during flood events is an important physiological challenge for this species. We collected W. asiaticus in midsummer from 66.8°N Yukon Territory, Canada, and measured thermal and immersion tolerance. Wyochernes asiaticus is freeze-avoidant, with a mean supercooling point of ?6.9 °C. It remains active at low temperatures (mean critical thermal minimum, CT_min, is ?3.6 °C) and has a critical thermal maximum (CT_max) of 37.8 °C, which is lower than other arachnids and consistent with its restriction to high latitudes. Fifty per cent of W. asiaticus individuals survived immersion in oxygen-depleted water for 17 days, suggesting that this species has high tolerance to immersion during flooding events. To our knowledge, these are the first data on the environmental physiology of any pseudoscorpion and a new addition to our understanding of the biology of polar microarthropods.     

On the threshold of dispersal: hitchhiking on a giant fly favours exaggerated male traits in a male‐dimorphic pseudoscorpion

The evolution of exaggerated male traits is frequently driven by competition between males to control resources critical for female survival and/or reproductive success. For flightless arthropods specializing on patchy habitats, dispersal agents may represent one such critical resource. The Neotropical pseudoscorpion, Semeiochernes armiger, disperses to new habitats by attaching to the giant timber fly, Pantophthalmus tabaninus, as it ecloses from pupal boreholes within decaying Ficus trees. In a study that combined field observations of mating with experimental removal of individuals from a large, pre‐dispersal population, our morphometric analyses revealed that S. armiger is among the most highly sexually dimorphic pseudoscorpions known, with males possessing unusual, triangular‐shaped pedipalpal chelae (hands) and a male‐specific, dimorphic chela peg that exhibits threshold trait expression. Several lines of evidence indicate that extreme sexual dimorphism in S. armiger results from male competition to monopolize pantophthalmid bores as strategic sites for inseminating females on the verge of dispersal. Sexually dimorphic pedipalpal characters were significantly larger in males located in and around pantophthalmid boreholes, compared with males collected at the periphery of the pantophthalmid emergence zone. Removal of pseudoscorpions resulted in a significant decline in pedipalpal size of males associated with pantophthalmid bores, followed by a rebound in size after collected individuals were returned to the tree. Most significantly, field observations of mating indicate that this competition translates into intense selection for exaggerated male traits, with all traits of the sexually dimorphic chelae exhibiting highly significant sexual selection differentials in males. © 2013 The Linnean Society of London     

Predation ofApochthonius minimus (Pseudoscorpionida: Chthoniidae) onFolsomia Candida (Collembola: Isotomidae) II. Effects of predation on prey populations

This study utilized the preference the pseudoscorpion, A. minimus, displays for certain sizes of the springtail, F. candida, in order to assess the population consequences of this selective predation. Although predation drove half of the treated collembolan populations to extinction, the populations which survived were not reduced in size and were in fact larger than most of the control populations (Figs. 1 and 2). During the first two weeks of the experiment, the predators apparently fed mainly on the original adults and the larger individuals of the second generation, thus increasing available space and improving the prey's chances for successful reproduction. The resulting demographic shift accounts for the significant decrease in mean length that occurred in the experimental populations without any accompanying significant change in their total numbers. Some effects of density, food, and age-structure on the fecundity of F. candida, acting in concert with the effects of size-selective predation, may lead to a variety of population consequences. In some circumstances, predation may cause increased production with decreased mean size, and no drastic change in standing crop. In different circumstances, a local population may become extinct. These results should further our understanding of the sporadic record of establishment and growth displayed byspringtail populations in the small, moist litter patches where they tend to aggregate.