A New Darwin Wasp (Hymenoptera: Ichneumonidae) and New Records of Behavioral Manipulation of the Host Spider Leucauge volupis (Araneae: Tetragnathidae).

Some ichneumonid wasps of the Polysphincta group of genera (Hymenoptera: Ichneumonidae: Pimplinae) induce behavioral modifications in their host spiders during a specific moment of their development, resulting in the construction of webs that differ in several aspects from those constructed by unparasitized individuals. In this study, we describe the parasitoid wasp Hymenoepimecis pinheirensis sp. n. (Ichneumonidae: Pimplinae) and present information on behavioral modifications in the orb-web structure of its host, the spider Leucauge volupis (Keyserling 1893). Previously, reported observation on this host/parasitoid interaction was restricted to one locality, and the wasp species was misidentified as Hymenoepimecis jordanensis Loffredo and Penteado-Dias 2009. Modified webs built by parasitized spiders lack adhesive spirals and have several radii that converge to the web hub. The cocoon built by the wasp larvae is attached to the web hub, suspended by horizontal radial lines, and surrounded by a tridimensional tangle positioned below the hub. This modified web structure is similar to the most frequent architecture of webs constructed by individuals of Leucauge mariana (Taczanowski 1881) parasitized by Hymenoepimecis tedfordi Gauld 1991. However, cocoon webs built by L. volupis parasitized by H. pinheirensis sp. n. differ from the cocoon webs described for the other Leucauge species parasitized by Hymenoepimecis wasps. This evidence suggests that the modified web pattern in Leucauge species is determined by specific responses of each spider species to the behavioral manipulation mechanism displayed by the wasps.

TARANTOBELUS JEFFDANIELSI N. SP. (PANAGROLAIMOMORPHA; PANAGROLAIMIDAE), A NEMATODE PARASITE OF TARANTULAS.

Multiple tarantula deaths for a wholesale breeder were reported in 2018. The breeder noticed white discharge in the oral cavities of the tarantulas. Upon inspection, it was discovered that the white discharge was a large group of nematodes intertwined inside the tarantula's oral cavity. We examined the nematodes and propose a new species, Tarantobelus jeffdanielsi n. sp., in the currently monotypic genus Tarantobelus based on a combination of morphological and morphometrical data and unique nuclear rDNA 28S and 18S sequences. Based on phylogenetic analyses, the previously described Tarantobelus arachnicida was relocated, along with T. jeffdanielsi, into the family Panagrolaimidae. We also provide evidence of the ability of T. jeffdanielsi to parasitize Galleria mellonella larvae and the tarantula Grammostola pulchra. The life span and fecundity of the new species were also assessed, resulting in an 11.2-d average life span, and a total fertility rate of 158 nematodes/adult.

Tachinobia repanda (Hymenoptera: Eulophidae) From Egg Sacs of a Colonial Spider, Cyrtophora moluccensis (Araneae: Araneidae) in Papua New Guinea.

We report the discovery of the wasp Tachinobia repanda Boucek collected from egg sacs of the colonial spider Cyrtophora moluccensis (Doleschall) in Morobe Province, Papua New Guinea (PNG) by Lubin, Y.D. in 1980. This is the first record of T. repanda from egg sacs of a colonial spider. The likely host of this eulophid wasp was the larvae of a sarcophagid fly that parasitizes the egg sacs of these spiders. The 67 T. repanda collected were all females and varied little in body size. We suggest that this species is a gregarious hyperparasitoid.

Prey and Venom Efficacy of Male and Female Wandering Spider, Phoneutria boliviensis (Araneae: Ctenidae).

Spiders rely on venom to catch prey and few species are even capable of capturing vertebrates. The majority of spiders are generalist predators, possessing complex venom, in which different toxins seem to target different types of prey. In this study, we focused on the trophic ecology and venom toxicity of Phoneutria boliviensis F. O. Pickard-Cambridge, 1897, a Central American spider of medical importance. We tested the hypothesis that its venom is adapted to catch vertebrate prey by studying its trophic ecology and venom toxicity against selected vertebrate and invertebrate prey. We compared both trophic ecology (based on acceptance experiments) and toxicity (based on bioassays) among sexes of this species. We found that P. boliviensis accepted geckos, spiders, and cockroaches as prey, but rejected frogs. There was no difference in acceptance between males and females. The venom of P. boliviensis was far more efficient against vertebrate (geckos) than invertebrate (spiders) prey in both immobilization time and LD50. Surprisingly, venom of males was more efficient than that of females. Our results suggest that P. boliviensis has adapted its venom to catch vertebrates, which may explain its toxicity to humans.

What's gotten into you?: a review of recent research on parasitoid manipulation of host behavior.

Some parasitoids modify the behavior of their hosts, benefiting themselves at the host's expense. This phenomenon is called 'manipulation', and current research on parasitoid manipulation of host behavior tends to fall into one of three categories. First, the frequency of manipulation and the magnitude of its benefits to the parasitoid remains unclear. Basic documentation of manipulations is thus a major research focus, with especially valuable recent data coming from spiders manipulated by Polysphincta wasps. Second, for a handful of systems, we now have sufficient phylogenetic and behavioral data to begin asking questions about how manipulation evolved. Finally, the field continues to probe the mechanisms through which parasitoids manipulate host behavior, and now examines the role of parasitoid symbionts in this interaction.

Tarantobelus arachnicida (Nematoda: Rhabditida) invasion of exotic spiders in Poland

Nematodes of the Brevibuccidae family were stated among spiders of the Theraphosidae family (the South American species), which were bred in Poland. The first sign of infection was anorexia which led to gradually increasing lethargy progressed to a huddled posture. Additionally, a white discharge with nematodes between mouth and chelicerae was noted. All of the derived nematodes were morphologically identified and determined to the species Tarantobelus arachnicida. A molecular analysis covered amplification and sequencing of small subunit ribosomal RNA (18S rRNA). A post mortem examination demonstrated the presence of nematodes not only near the chelicerae, but also inside the intestine, hence the source of infection might be insects used as a food. The research showed that such kind of infection is an important disease, which poses a serious risk to the breeding spiders. To date there is no effective treatment, however, we demonstrated that usage of the Lugol's solution seems to be promising.

The legs of "spider associated" parasitic primary larvae of Mantispa aphavexelte (Mantispidae, Neuroptera) - Attachment devices and phylogenetic implications.

The legs of the primary larva of Mantispa aphavexelte, parasite in egg sacks of spiders, were examined using scanning electron microscopy (SEM), histology and confocal laser scanning microscopy (CLSM). The leg morphology is described in detail, including intrinsic muscles. Functional adaptations of the leg attachment devices are discussed, especially regarding the material composition. For example, a sole-like flexible ventral tarsal surface containing resilin is combined with sclerotized pseudo-claws. This likely enables the larvae to cope with surface structures on the spider's body, with substrates on the ground, and also with various structural elements in the spider's nest. The leg morphology is evaluated with respect to phylogenetic affinities. A trumpet-shaped, elongated empodium has likely evolved early in the evolution of Neuroptera and may consequently belong to the groundplan of a large subgroup of the order. It characterizes most groups of the hemerobiform lineage and is also present in the myrmeleontiform Psychopsidae. The presence of a tarsal protrusion resembling a pretarsus confirms the monophyletic origin of Mantispoidea. A single fixed tooth and a specific surface structure are potential autapomorphies of Mantispidae. A distal tibial subunit partly separated from the main part of the leg segment is an apomorphy only described for larvae of M. aphavexelte.

The changing use of the ovipositor in host shifts by ichneumonid ectoparasitoids of spiders (Hymenoptera, Ichneumonidae, Pimplinae).

Accurate egg placement into or onto a living host is an essential ability for many parasitoids, and changes in associated phenotypes, such as ovipositor morphology and behaviour, correlate with significant host shifts. Here, we report that in the ichneumonid group of koinobiont spider-ectoparasitoids ("polysphinctines"), several putatively ancestral taxa (clade I here), parasitic on ground-dwelling RTA-spiders (a group characterised by retrolateral tibial apophysis on male palpal tibiae), lay their eggs in a specific way. They tightly bend their metasoma above the spider's cephalothorax, touching the carapace with the dorsal side of the ovipositor apically ("dorsal-press"). The egg slips out from the middle part of the ventral side of the ovipositor and moves towards its apex with the parted lower valves acting as rails. Deposition occurs as the parasitoid draws the ovipositor backwards from under the egg. Oviposition upon the tough carapace of the cephalothorax, presumably less palatable than the abdomen, is conserved in these taxa, and presumed adaptive through avoiding physical damage to the developing parasitoid. This specific way of oviposition is reversed in the putatively derived clade of polysphinctines (clade II here) parasitic on Araneoidea spiders with aerial webs, which is already known. They bend their metasoma along the spider's abdomen, grasping the abdomen with their fore/mid legs, pressing the ventral tip of the metasoma and the lower valves of the ovipositor against the abdomen ("ventral-press"). The egg is expelled through an expansion of the lower valves, which is developed only in this clade and evident in most species, onto the softer and presumably more nutritious abdomen.

Morphological and molecular characterization of Tarantobelus arachnicida gen. n., sp. n. (Nematoda, Rhabditida, Brevibuccidae), a parasitic nematode of tarantulas.

A new genus and new species, Tarantobelus arachnicida, was found in the oral opening of tarantula spiders bred in captivity in Poland. The new species is characterized by having a small body (0.77-0.95 mm long in females and 0.66-0.84 mm in males), cuticle poorly annulated by transverse incisures, lateral field inconspicuous, lips separated with small cuticular flaps topping each lip, stoma panagrolaimoid with gymnostom well developed with robust and refringent rhabdia, pharynx panagrolaimoid with isthmus slightly longer than the basal bulb, intestine with cardiac (anterior) and rectal (posterior) areas with narrower walls. Mature females with intestinal cells including needle crystal packs, excretory pore at isthmus level, female reproductive system panagrolaimoid with post-vulval sac 0.4-0.8 times the length of the corresponding body diameter and having very thick walls, vulva very prominent, female rectum 0.8-1.3 times the length of the anal body diameter, female tail conical with acute tip with phasmids at 58-62% of its length. Male tail conical with long and thin mucro, spicules ventrad bent having rounded manubrium and thick gubernaculum. Description, measurements and illustrations of the new species are provided. Molecular analyses show its relationship with Brevibucca and Cuticonema. On the other hand, Medibulla and its corresponding subfamily Medibullinae, previously in Osstellidae, are transferred to Panagrolaimidae, being Shahnematinae, the junior synonym of Medibullinae. Indocephalobus, recently proposed and located in the family Panagrolaimidae, is considered a junior synonym of Diplogastrellus (Diplogasteromorpha), and its only species, I. zebrae, is considered a junior synonym of D. gracilis. In addition, a key to identification of panagrolaimoid genera is included.

The effect of fight cost structure on fighting behaviour involving simultaneous decisions and variable investment levels.

In the "producer-scrounger" model, a producer discovers a resource and is in turn discovered by a second individual, the scrounger, who attempts to steal it. This resource can be food or a territory, and in some situations, potentially divisible. In a previous paper we considered a producer and scrounger competing for an indivisible resource, where each individual could choose the level of energy that they would invest in the contest. The higher the investment, the higher the probability of success, but also the higher the costs incurred in the contest. In that paper decisions were sequential with the scrounger choosing their strategy before the producer. In this paper we consider a version of the game where decisions are made simultaneously. For the same cost functions as before, we analyse this case in detail, and then make comparisons between the two cases. Finally we discuss some real examples with potentially variable and asymmetric energetic investments, including intraspecific contests amongst spiders and amongst parasitoid wasps. In the case of the spiders, detailed estimates of energetic expenditure are available which demonstrate the asymmetric values assumed in our models. For the wasps the value of the resource can affect the probabilities of success of the defender and attacker, and differential energetic investment can be inferred. In general for real populations energy usage varies markedly depending upon crucial parameters extrinsic to the individual such as resource value and intrinsic ones such as age, and is thus an important factor to consider when modelling.

Molecular phylogeny of Pompilinae (Hymenoptera: Pompilidae): Evidence for rapid diversification and host shifts in spider wasps.

Pompilinae is one of the largest subfamilies of spider wasps (Pompilidae). Most pompilines are generalist spider predators at the family level, but some taxa exhibit ecological specificity (i.e., to spider-host guild). Here we present the first molecular phylogenetic analysis of Pompilinae, toward the aim of evaluating the monophyly of tribes and genera. We further test whether changes in the rate of diversification are associated with host-guild shifts. Molecular data were collected from five nuclear loci (28S, EF1-F2, LWRh, Wg, Pol2) for 76 taxa in 39 genera. Data were analyzed using maximum likelihood (ML) and Bayesian inference (BI). The phylogenetic results were compared with previous hypotheses of subfamilial and tribal classification, as well as generic relationships in the subfamily. The classification of Pompilus and Agenioideus is also discussed. A Bayesian relaxed molecular clock analysis was used to examine divergence times. Diversification rate-shift tests accounted for taxon-sampling bias using ML and BI approaches. Ancestral host family and host guild were reconstructed using MP and ML methods. Ancestral host guild for all Pompilinae, for the ancestor at the node where a diversification rate-shift was detected, and two more nodes back in time was inferred using BI. In the resulting phylogenies, Aporini was the only previously proposed monophyletic tribe. Several genera (e.g., Pompilus, Microphadnus and Schistonyx) are also not monophyletic. Dating analyses produced a well-supported chronogram consistent with topologies from BI and ML results. The BI ancestral host-use reconstruction inferred the use of spiders belonging to the guild "other hunters" (frequenting the ground and vegetation) as the ancestral state for Pompilinae. This guild had the highest probability for the ML reconstruction and was equivocal for the MP reconstruction; various switching events to other guilds occurred throughout the evolution of the group. The diversification of Pompilinae shows one main rate-shift coinciding with a shift to ground-hunter spiders, as reconstructed by the BI ancestral character-state analysis.

Clistopyga caramba sp. nov. (Hymenoptera: Ichneumonidae; Pimplinae), an astonishing example of mimicry in spider-attacking parasitoid wasps.

Clistopyga caramba sp. nov. Castillo & Sääksjärvi, collected from the tropical Andean-Amazonian interface in Peru, is described and illustrated. The new species is characterized by highly modified posterior metasomal tergites and a rare colour pattern of the metasoma. Two possible explanations for the bizarre metasomal morphology and colouration are proposed.

Establishment of the Brown Widow Spider (Araneae: Theridiidae) and Infestation of its Egg Sacs by a Parasitoid, Philolema latrodecti (Hymenoptera: Eurytomidae), in French Polynesia and the Cook Islands.

This paper presents two newly established species for French Polynesia: the invasive brown widow spider, Latrodectus geometricus C. L. Koch, and its potential biocontrol agent, the parasitoid wasp, Philolema latrodecti (Fullaway). The brown widow spider was recorded from the island of Moorea in 2006 and, since that discovery, the occurrence of this species has expanded to two of the five archipelagos of French Polynesia including the main island of Tahiti and four of the Cook Islands. Although the tropical climate contributes to the establishment of L. geometricus, a biotic factor, P. latrodecti, may restrain population from demographic explosion. This eurytomid wasp is present in French Polynesia and is a parasitoid that has been used in biological control of the southern black widow Latrodectus mactans (F.) in Hawaii. This wasp could become a significant limiting factor for L. geometricus distribution on these islands, as it was found in 31% of the Tahitian brown widow spider egg sacs that were dissected. However, thus far, the wasp was only found on Tahiti in association with the brown widow spider. Although the brown widow is generally considered to be less toxic than its black widow relatives, it remains of medical concern in French Polynesia because reactions to its bites can, at times, be severe. The spider remains of public concern because it is a novel species; it has the word widow in its name and dark morphs are mistaken as black widows.

Host manipulation by an ichneumonid spider ectoparasitoid that takes advantage of preprogrammed web-building behaviour for its cocoon protection.

Host manipulation by parasites and parasitoids is a fascinating phenomenon within evolutionary ecology, representing an example of extended phenotypes. To elucidate the mechanism of host manipulation, revealing the origin and function of the invoked actions is essential. Our study focused on the ichneumonid spider ectoparasitoid Reclinervellus nielseni, which turns its host spider (Cyclosa argenteoalba) into a drugged navvy, to modify the web structure into a more persistent cocoon web so that the wasp can pupate safely on this web after the spider's death. We focused on whether the cocoon web originated from the resting web that an unparasitized spider builds before moulting, by comparing web structures, building behaviour and silk spectral/tensile properties. We found that both resting and cocoon webs have reduced numbers of radii decorated by numerous fibrous threads and specific decorating behaviour was identical, suggesting that the cocoon web in this system has roots in the innate resting web and ecdysteroid-related components may be responsible for the manipulation. We also show that these decorations reflect UV light, possibly to prevent damage by flying web-destroyers such as birds or large insects. Furthermore, the tensile test revealed that the spider is induced to repeat certain behavioural steps in addition to resting web construction so that many more threads are laid down for web reinforcement.

Parasitism of Trombidium brevimanum larvae on agrobiont linyphiid spiders from Germany.

An experiment on three differently-managed agricultural fields in Ahlum, Germany, which aimed at establishing the impact of different management systems on the biodiversity of predators and decomposers, yielded a significant number of spiders parasitized by larvae of Trombidium brevimanum (Actinotrichida, Parasitengona, Trombidiidae). Spider data from the whole sampling period (September 2010-July 2012), indicated that ectoparasitic larvae were recorded only on spiders in pitfall traps in the period of June-July 2011. In this period, only eight species of Linyphiidae--out of 42 species assigned to nine spider families recorded from the study area--were parasitized by mites; considerable levels of parasitism were recorded on Erigone atra, E. dentipalpis, and Oedothorax apicatus. The highest prevalence of parasitism was recorded on the organic field for E. atra (29%), while on the integrated and conventional fields significantly fewer parasitized spiders were observed. The preferred attachment sites on the spider host were regions with softer cuticle, especially regions on the carapace and on the abdomen, adjacent to the pedicel.

Two new species of the Polysphincta genus-group (Hymenoptera: Ichneumonidae: Pimplinae) reared from their spider (Araneae) hosts in Europe.

Flacopimpla barathrica sp. n. and Zatypota flamma sp. n., belonging to the Polysphincta genus-group, are described, the former from Fennoscandia and the latter from Hungary. Like other species of this group for which the natural history is known, the new species are koinobiont ectoparasitoids of spiders. They have been reared from the theridiid spiders Thymoites bellissimus (L. Koch) and Parasteatoda sp. Archer respectively. The host records support the hypothesis of Flacopimpla being parasitoids of the spider family Theridiidae. The results also indicate that F. barathrica sp. n. has a very narrow host range consisting of a single species in Fennoscandia. The genus Flacopimpla was previously only known from the New World, and F. barathrica sp. n. is thus the first species of its genus in the Palaearctic. It is also the first troglophilic species of the Polysphincta genus-group. Zatypota albicoxa (Walker) is reported as new to Hungary. 

New host records for European Acroceridae (Diptera), with discussion of species limits of Acrocera orbiculus (Fabricius) based on DNA-barcoding.

New European host records for the Acroceridae species Acrocera orbiculus (Fabricius) and Ogcodes reginae Trojan are reported. Acrocera orbiculus was reared from Amaurobius erberi (Keyserling), and O. reginae from Clubiona leucaspis (Simon) and Evarcha jucunda (Lucas). Where possible, DNA-barcodes are presented for reared endoparasitoids and their host specimens. Based on mitochondrial COI, the intraspecific genetic variability of 15 western Palaearctic A. orbiculus is discussed. Maximum likelihood analysis reveals two clades, though they have low statistical support and no distinct barcoding gap. Therefore, we consider all barcoded specimens of A. orbiculus to be a single biological species with a high degree of phenotypic plasticity regarding body size and coloration. Based on molecular and morphological evidence, Paracrocera kaszabi Majer, Paracrocera manevali Séguy and Paracrocera minuscula Séguy are placed in synonymy with A. orbiculus. The male of the Canary Islands endemic Acrocera cabrerae Frey is described for the first time. 

Description and host interactions of a new species of Exetasis Walker (Diptera: Acroceridae), with a key to species of the genus.

Exetasis jujuyensis Gillung sp. nov. (Acroceridae) is described from Argentina and a dichotomous key to species of Exetasis Walker is provided. Detailed observations are presented on the effect of fly larval development on the behavior of the host spider (Acanthoscurria sternalis Pocock (Theraphosidae)).

Description of the male Hymenoepimecis japi Sobczak et al. 2009 (Hymenoptera, Ichneumonidae) parasitoid of Leucauge roseosignata Mello-Leitão 1943 (Araneae: Tetragnathidae) / Descrição do macho de Hymenoepimecis japi Sobczak et al. 2009 (Hymenoptera, Ichneumonidae) parasitoide de Leucauge roseosignata Mello-Leitão 1943 (Araneae: Tetragnathidae)

The male of Hymenoepimecis japi (Hymenoptera, Ichneumonidae, Pimplinae) is described and illustrated. The specimen was collected in a modified web (cocoon web) of Leucauge roseosignata (Araneae, Tetragnathidae) made in a laboratory. Both, host and parasitoid were collected in Reserva Biológica Serra do Japi, located in Jundiaí, São Paulo, Brazil.

O macho de Hymenoepimecis japi (Hymenoptera, Ichneumonidae, Pimplinae) é descrito e ilustrado. O espécime foi coletado em teia modificada "cocoon web" de Leucauge roseosignata (Araneae, Tetragnathidae), construída no laboratório. Ambos, hospedeiro e parasitoide, foram coletados na Reserva Biológica Serra do Japi, localizada em Jundiaí, Estado de São Paulo, Brasil.

Mites parasitic on Australasian and African spiders found in the pet trade; a redescription of Ljunghia pulleinei Womersley.

Parasitic mites associated with spiders are spreading world-wide through the trade in tarantulas and other pet species. Ljunghia pulleinei Womersley, a mesostigmatic laelapid mite originally found in association with the mygalomorph spider Selenocosmia stirlingi Hogg (Theraphosidae) in Australia, is redescribed and illustrated on the basis of specimens from the African theraphosid spider Pterinochilus chordatus (Gerstäcker) kept in captivity in the British Isles (Wales). The mite is known from older original descriptions of Womersley in 1956; the subsequent redescription of Domrow in 1975 seems to be questionable in conspecificity of treated specimens with the type material. Some inconsistencies in both descriptions are recognised here as intraspecific variability of the studied specimens. The genus Arachnyssus Ma, with species A. guangxiensis (type) and A. huwenae, is not considered to be a valid genus, and is included in synonymy with Ljunghia Oudemans. A new key to world species of the genus Ljunghia is provided.