An Immature Dermapteran Misidentified as an Adult Zorapteran: The Case of Formosozoros newi Chao & Chen, 2000.

Zoraptera shows extreme uniformity in general body morphology, with the exception of Formosozoros (=Zorotypus) newi Chao & Chen, 2000, which stands out in terms of the shape and arrangement of its legs, the cerci, and several other morphological characters. After critical evaluation, we found that this species is not a zorapteran but is instead a nymph (1st instar) of an earwig; i.e., F. newi is a dermapteran. Because of the lack of morphological descriptions of Dermaptera nymphs that would allow species identification, and because the type material of F. newi is lost, a more detailed classification is not possible. We therefore propose that the genus name Formosozoros Chao & Chen, 2000 and the species name Formosozoros newi Chao & Chen, 2000 are nomina dubia.

Cryptic diversity in Zoraptera: Latinozoros barberi (Gurney, 1938) is a complex of at least three species (Zoraptera: Spiralizoridae).

The order Zoraptera contains relatively few species, but current molecular phylogenetic studies suggest an unexpectedly high level of cryptic diversity in the order with many overlooked species based on morphology alone. Latinozoros Kukalova-Peck & Peck, 1993 represents the only genus of monotypic Latinozorinae (Zoraptera: Spiralizoridae) with only one species described, L. barberi (Gurney, 1938), until now. Although this species has been repeatedly reported from a number of locations in South and Central America, it is likely a complex of unrecognized species. Here, we present a molecular phylogenetic reconstruction revealing three genetically distinct lineages in Latinozoros, and we also present detailed morphological comparisons that prove the species status of Latinozoros cacaoensis sp. nov. from French Guiana and L. gimmeli sp. nov. from the Dominican Republic, Trinidad and Panama. The results indicate that the species previously referred to L. barberi is actually a species complex that includes L. barberi, the new species described here, and perhaps other species.

Termitophily Documented in Earwigs (Dermaptera).

Based on behavioral observations, we report termitophily by the earwig Spirolabia kaja Kocárek, sp. nov. (Spongiphoridae: Labiinae). The new species was found in association with the wood-boring termite Schedorhinotermes sarawakensis (Holmgren, 1913) in a dipterocarp rain forest in Borneo; in addition to being observed in the galleries, termite-earwig interactions were subsequently documented in the laboratory. We found that earwigs and termites communicate by antennation, and we observed no form of targeted mutual or unilateral aggressive behavior. The earwigs responded to the proximity of an experimentally irritated termite soldier by conflict-avoidance behavior based on thanatosis, which seems to be a defensive reaction that may reduce the chance of being attacked by an irritated termite. Based on the analysis of gastrointestinal tract contents, we conclude that S. kaja sp. nov. is an omnivorous species that feeds mainly on plant tissues and fungi but occasionally on arthropod remains. The occurrence of S. kaja sp. nov. adults together with the nymphs (2nd to 4th instars) in the galleries of S. sarawakensis strongly suggests that the earwig can reproduce inside the termite colony. Spirolabia kaja Kocárek, sp. nov. is the first earwig species for which termitophily has been demonstrated.

Estimation of the Recent Expansion Rate of Ruspolia nitidula (Orthoptera) on a Regional and Landscape Scale.

Recent changes in insect distribution are consistent with the expected interacting effects of climate and habitat change. The orthopteran Ruspolia nitidula has expanded its area of distribution in Western and Central Europe in recent decades. Because males emit a sound that is easily detected at a distance of up to 40 m, it is possible to detect spreading individuals and to therefore document routes and rates of spread. Using occurrence data at the landscape scale and three methods, including least-cost path analysis with habitat suitability, we estimated the R. nitidula expansion rate from 2006 to 2020 in the Czech Republic; this involved estimating distances between two origin occurrences in 2006 and two occurrences on the area margin in 2020. For comparison, we directly monitored expansion based on detection of singing males at the regional scale at the areal margin in the Odra River basin (2016-2020). The estimated maximum expansion rate ranged from 13.8 to 16.2 km/year based on occurrence data at the landscape scale and from 11.1 to 11.7 km/year based on the monitoring of males in the Odra River basin. To our knowledge, this is the first report of the direct monitoring of individual spreading males to detect changes in the distribution of an orthopteran.

Estimating the effect of tracking tag weight on insect movement using video analysis: A case study with a flightless orthopteran.

In this study, we describe an inexpensive and rapid method of using video analysis and identity tracking to measure the effects of tag weight on insect movement. In a laboratory experiment, we assessed the tag weight and associated context-dependent effects on movement, choosing temperature as a factor known to affect insect movement and behavior. We recorded the movements of groups of flightless adult crickets Gryllus locorojo (Orthoptera:Gryllidae) as affected by no tag (control); by light, medium, or heavy tags (198.7, 549.2, and 758.6 mg, respectively); and by low, intermediate, or high temperatures (19.5, 24.0, and 28.3°C, respectively). Each individual in each group was weighed before recording and was recorded for 3 consecutive days. The mean (± SD) tag mass expressed as a percentage of body mass before the first recording was 26.8 ± 3.7% with light tags, 72 ± 11.2% with medium tags, and 101.9 ± 13.5% with heavy tags. We found that the influence of tag weight strongly depended on temperature, and that the negative effects on movement generally increased with tag weight. At the low temperature, nearly all movement properties were negatively influenced. At the intermediate and high temperatures, the light and medium tags did not affect any of the movement properties. The continuous 3-day tag load reduced the average movement speed only for crickets with heavy tags. Based on our results, we recommend that researchers consider or investigate the possible effects of tags before conducting any experiment with tags in order to avoid obtaining biased results.

Energy balance of food in a detrito-bryophagous groundhopper (Orthoptera: Tetrigidae).

Detritus (decaying organic matter) and phyllodes of mosses are two main components in the diet of groundhoppers (Orthoptera: Tetrigidae). We studied the energy balance of consumed food under laboratory conditions in the detrito-bryophagous groundhopper, Tetrix subulata (Linnaeus, 1758). The results indicated that the energy food budget of this detrito-bryophagous groundhopper was comparable to those of small herbivorous grasshoppers (Acrididae: Gomphocerinae, Melanoplinae), which have a similar energy food budget of approximately 800-1,100 J/g. T. subulata consumed four times more detritus than mosses, although both components provided similar amounts of energy (ca. 15-16 kJ/g). However, in contrast with detritus, moss fragments passed through the digestive tract without a distinct change in their mass or a loss in their energy value. We assume that moss may cause the longer retention of semifluid mass of partly digested food in the alimentary tract; hence, the digestion and efficiency of nutrient absorption from detritus could be more effective.

Molecular Phylogeny and Infraordinal Classification of Zoraptera (Insecta).

Zoraptera is a small and predominantly tropical insect order with an unresolved higher classification due to the extremely uniform external body morphology. We, therefore, conducted a multigene molecular phylogeny of extant Zoraptera and critically re-evaluated their morphological characters in order to propose a natural infraordinal classification. We recovered a highly-resolved phylogeny with two main clades representing major evolutionary lineages in Zoraptera, for which we propose family ranks. The two families exhibit striking differences in male genitalia and reproductive strategies. Each family contains two subclades (subfamilies) supported by several morphological synapomorphies including the relative lengths of the basal antennomeres, the number and position of metatibial spurs, and the structure of male genitalia. The newly proposed higher classification of Zoraptera includes the family Zorotypidae stat. revid. with Zorotypinae Silvestri, 1913 (Zorotypus stat. revid., Usazoros Kukalova-Peck and Peck, 1993 stat. restit.) and Spermozorinae subfam. nov. (Spermozoros gen. nov.), and Spriralizoridae fam. nov. with Spiralizorinae subfam. nov. (Spiralizoros gen. nov., Scapulizoros gen. nov., Cordezoros gen. nov., Centrozoros Kukalova-Peck and Peck, 1993, stat. restit., Brazilozoros Kukalova-Peck and Peck, 1993, stat. restit.), and Latinozorinae subfam. nov. (Latinozoros Kukalova-Peck and Peck, 1993, stat. restit.). An identification key and morphological diagnoses for all supraspecific taxa are provided.

The cephalo-thoracic apparatus of Caputoraptor elegans may have been used to squeeze prey.

Alienoptera is an insect order recently described from mid-Cretaceous amber [1] and is phylogenetically nested in the Dictyoptera lineage. Alienoptera currently comprises three species: Alienopterus brachyelytrus[1], Alienopterella stigmatica[2] and Caputoraptor elegans[3]. The most interesting is Caputoraptor elegans, which was recently described in Current Biology by Bai and colleagues [3] and which has an unusual cephalo-thoracic device formed by wing-like extensions of the genae and the corresponding edges of the pronotum. Bai and colleagues [3] suggested that the cephalo-thoracic apparatus may have been used to hold the female and male together during copulation. According to this possible function, the cephalo-thoracic apparatus of the female would fit together with the spread forewings of the male while the female was on the back of the male during copulation. This function was proposed based on examination of females and nymphs, and the authors stated that it could be falsified if a male with a similar apparatus were discovered. After examining a male nymph of this species (Figure 1), I here suggest that the cephalo-thoracic apparatus was not used for copulation but was instead used for predation and feeding.

Influence of surface flattening on biodiversity of terrestrial arthropods during early stages of brown coal spoil heap restoration.

Heterogeneity of environmental conditions is the crucial factor supporting biodiversity in various habitats, including post-mining sites. The effects of micro-topographic heterogeneity on biodiversity and conservation potential of arthropod communities in post-industrial habitats had not been studied before now. At one of the largest European brown coal spoil heaps, we sampled eight groups of terrestrial arthropods with different life strategies (moths, spiders, ground beetles, ants, orthopteroids, centipedes, millipedes, and woodlice), in successionally young plots (5-18 y), with a heterogeneous wavy surface after heaping, and compared the communities with plots flattened by dozing. A combination of the standardized quantitative sampling, using two different methods, and a paired design of the plot selection enabled a robust analysis. Altogether, we recorded 380 species of the focal arthropods, 15 of them nationally threatened. We revealed the importance of the micro-topographic heterogeneity for the formation of the biodiversity of arthropods in their secondary refuges. The communities with higher biodiversity and conservation value were detected in the plots with heterogeneous surfaces; exceptions were ground beetles and millipedes. The surface flattening, often the first step of technical reclamation projects, thus suppress biodiversity of most terrestrial arthropods during the restoration of post-mining sites. Since the communities of both surface types differed, the proportional presence on both surfaces could be more efficient in supporting the local biodiversity. We suggest reducing the surface dozing for the cases with other concerns only, to achieve a proportional representation of both surface types. Such a combination of different restoration approaches would, thus, efficiently support high biodiversity of groups with various needs.

Weather-dependent microhabitat use by Tetrix tenuicornis (Orthoptera: Tetrigidae).

For ectothermic animals, selection of a suitable microhabitat is affected by a combination of abiotic and biotic factors. Also important is the trade-off between those microhabitats with optimal microclimatic conditions and food availability vs. those with the lowest level of competition and lowest risk of predation. Central European species of groundhoppers (Orthoptera: Tetrigidae) live in locations with small-scale mosaics of patches formed by bare ground, moss cushions and vascular plants (grasses and forbs). Our research focused on the effects of selected weather components (current temperature, humidity, atmospheric pressure and sunlight) on specific microhabitat selection by adults (during the reproductive season) and by the last-instar nymphs (during the non-reproductive season) of the groundhopper Tetrix tenuicornis. Using experimental conditions, we determined that microhabitat use by T. tenuicornis is sex-specific and that microhabitat preference differs between adults and nymphs. We suppose that microhabitats are used according to groundhopper current needs in relation to each habitat's suitability for maintaining body temperature, food intake and reproductive behaviour. Microhabitat preferences were significantly associated with temperature and atmospheric pressure. Changes in atmospheric pressure signal changes in weather, and insects respond to increases or decreases in pressure by adjusting their behaviour in order to enhance survival. We propose that, under low atmospheric pressure, T. tenuicornis actively seeks microhabitats that provide increased protection from adverse weather.

In search for a compromise between biodiversity conservation and human health protection in restoration of fly ash deposits: effect of anti-dust treatments on five groups of arthropods.

Recently, fly ash deposits have been revealed as a secondary refuge of critically endangered arthropods specialised on aeolian sands in Central Europe. Simultaneously, these anthropogenic habitats are well known for their negative impact on human health and the surrounding environment. The overwhelming majority of these risks are caused by wind erosion, the substantial decreasing of which is thus necessary. But, any effects of anti-dust treatments on endangered arthropods have never been studied. We surveyed communities of five arthropod groups (wild bees and wasps, leafhoppers, spiders, hoverflies and orthopteroid insects) colonising three fly ash deposits in the western Czech Republic. We focused on two different anti-dust treatments (~70 and 100 % cover of fly ash by barren soil) and their comparison with a control of bare fly ash. Altogether, we recorded 495 species, including 132 nationally threatened species (eight of them were considered to be extinct in the country) and/or 30 species strictly specialised to drift sands. Bees and wasps and leafhoppers contained the overwhelming majority of species of the highest conservation interest; a few other important records were also in spiders and orthopteroids. Total soil cover depleted the unique environment of fly ash and thus destroyed the high conservation potential of the deposits. On the other hand, partial coverage (with ~30 % of bare fly ash) still offered habitats for many of the most threatened species, as we showed by both regression and multivariate analyses, with a decrease of wind erosion. This topic still needs much more research interest, but we consider mosaic-like preservation of smaller spots of fly ash as one of the possible compromises between biodiversity and human health.

Synonymy of Discotettix adenanii Mahmood, Idris & Salmah, 2007 with D. belzebuth (Serville, 1838) (Orthoptera: Tetrigidae).

According to a study of type specimens of Discotettix adenanii Mahmood, Idris & Salmah, 2007 and copious specimens of D. belzebuth (Serville, 1838) collected in different parts of Borneo, we found that all species-specific morphological characters of D. adenanii, according to its original description, fall into the morphological variability of D. belzebuth. Thus, we synonymize D. adenanii with D. belzebuth. The sequence of a segment of the mitochondrial 16S ribosomal RNA of D. belzebuth was also evaluated and added to GenBank as a DNA barcode.

Description and biology of Euborellia arcanum sp. nov., an alien earwig occupying greenhouses in Germany and Austria (Dermaptera: Anisolabididae).

Greenhouses in botanical or zoological gardens are home to dozens of species of invertebrates that were introduced alongside plants or potting soil. Our study presents the description of an alien species of earwig, Euborellia arcanum sp. nov., found in tropical greenhouses in Leipzig and Potsdam (Germany) and in Vienna (Austria), including information about its biology in breeding culture. The species was most likely introduced into Europe by way of plants or plant matter from Florida, but the region of its natural habitat is unknown. The sequence of the mitochondrial gene cytochrome c oxidase subunit I (COI) was also evaluated and added to GenBank as a DNA barcode for further identification.

Age-related mandible abrasion in the groundhopper Tetrix tenuicornis (Tetrigidae, Orthoptera).

A study was conducted to determine whether the mandibles of the detrito-/bryophagous groundhopper Tetrix tenuicornis are subject to mechanical wear as a result of feeding, as is the case for grasshoppers that feed on silica-rich grasses. Abrasion was evaluated by measuring the length and width of the 3rd incisor and length of the 4th incisor in adults of different ages collected under natural conditions during one season. Although T. tenuicornis and other groundhoppers avoid feeding on grasses, we found that mandible abrasion increased with T. tenuicornis age. Age-related abrasion of the incisors of left and right mandibles was statistically significant in both sexes but the degree of abrasion was greater for females than males, apparently reflecting differences in the frequency and magnitude of feeding. Degree of abrasion also differed between right and left mandibles, probably because of differences in how each mandible is used during food processing. Abrasion of cuticular mandible structures may reduce the effectiveness of food processing late in the season.

When the body hides the ancestry: phylogeny of morphologically modified epizoic earwigs based on molecular evidence.

Here, we present a study regarding the phylogenetic positions of two enigmatic earwig lineages whose unique phenotypic traits evolved in connection with ectoparasitic relationships with mammals. Extant earwigs (Dermaptera) have traditionally been divided into three suborders: the Hemimerina, Arixeniina, and Forficulina. While the Forficulina are typical, well-known, free-living earwigs, the Hemimerina and Arixeniina are unusual epizoic groups living on molossid bats (Arixeniina) or murid rodents (Hemimerina). The monophyly of both epizoic lineages is well established, but their relationship to the remainder of the Dermaptera is controversial because of their extremely modified morphology with paedomorphic features. We present phylogenetic analyses that include molecular data (18S and 28S ribosomal DNA and histone-3) for both Arixeniina and Hemimerina for the first time. This data set enabled us to apply a rigorous cladistics approach and to test competing hypotheses that were previously scattered in the literature. Our results demonstrate that Arixeniidae and Hemimeridae belong in the dermapteran suborder Neodermaptera, infraorder Epidermaptera, and superfamily Forficuloidea. The results support the sister group relationships of Arixeniidae+Chelisochidae and Hemimeridae+Forficulidae. This study demonstrates the potential for rapid and substantial macroevolutionary changes at the morphological level as related to adaptive evolution, in this case linked to the utilization of a novel trophic niche based on an epizoic life strategy. Our results also indicate that the evolutionary consequences of the transition to an ectoparazitic mode of living, which is extremely rare in earwigs, have biased previous morphology-based hypotheses regarding the phylogeny of this insect group.

Annotated checklist of the grasshoppers and crickets (Orthoptera) of the Czech Republic.

A checklist is presented of the Orthoptera of the Czech Republic. Based on the critical revision of published data and previous lists of species, which often contained only general or questionable data and which often inaccurately cited each other, we found 96 species of Orthoptera belonging to the fauna of the Czech Republic. We emphasize those changes that are based on comparison with previous checklists. We provide information on missing, unclear, and extinct species and on newly detected species, and we confirm the status of species that have been missing for a long time (Leptophyes boscii, Polysarcus denticauda, Ruspolia nitidula, Eumodicogryllus bordigalensis, Tetrix bolivari, Mecostethus parapleurus). We also note those species for which only several individuals have been detected (Pteronemobius heydenii) or those survived only at a single locality (Platycleis montana, Aiolopus thalassinus, Dociostaurus brevicollis, Omocestus petraeus) or at two localities (Poecilimon intermedius, Platycleis veyseli, Pseudopodisma nagyi). Phaneroptera nana is recorded as new for Bohemia.

Ovaries and germline cysts and their evolution in Dermaptera (Insecta).

We studied the ovary structure and initial stages of oogenesis in 15 representatives of several dermapteran taxa, including the epizoic Arixeniina. In all examined species, the ovaries are meroistic-polytrophic. The ovaries of the basal taxa ('Pygidicranidae', 'Diplatyidae', and Labiduridae) are composed of elongated ovarioles, attached to short lateral oviducts. In these groups, ovarioles contain several (more than 30) ovarian follicles in a linear arrangement. In the Eudermaptera, the ovaries are composed of 1-6 (Spongiphoridae) or 20-40 (Forficulidae, Chelisochidae) short ovarioles (containing 2 ovarian follicles only) that open to strongly elongated lateral oviducts. In all investigated dermapterans, the ovarian follicles are composed of two germline cells only: an oocyte and a polyploid nurse cell that are covered by a simple follicular epithelium. Our studies indicate that despite a rather unique morphology of the ovarian follicles in the examined species, the processes leading to the formation of the oocyte and nurse cell units are significantly different in basal versus derived taxa. The ovaries of Arixenia esau are composed of 3 short ovarioles attached to a strongly dilated lateral oviduct, 'the uterus', containing developing embryos. Histological analysis suggests that the origin of the oocyte and nurse cell units in this species follows the pattern described in eudermapterans. The interpretation of our results in an evolutionary context supports the monophyly of the Dermaptera and Eudermaptera, and the inclusion of the Arixeniina and Hemimerina in the latter taxon.