Comment on "The lower Cambrian lobopodian Cardiodictyon resolves the origin of euarthropod brains".

Strausfeld et al. (Report, 24 Nov 2022, p. 905) claim that Cambrian fossilized nervous tissue supports the interpretation that the ancestral panarthropod brain was tripartite and unsegmented. We argue that this conclusion is unsupported, and developmental data from living onychophorans contradict it.

Response to Comment on "The lower Cambrian lobopodian Cardiodictyon resolves the origin of euarthropod brains".

Budd et al. challenge the identity of neural traces reported for the Cambrian lobopodian Cardiodictyon catenulum. Their argumentation is unsupported, as are objections with reference to living Onychophora that misinterpret established genomic, genetic, developmental, and neuroanatomical evidence. Instead, phylogenetic data corroborate the finding that the ancestral panarthropod head and brain is unsegmented, as in C. catenulum.

Distribution pattern of arthropods on the leaf surfaces of Acacia auriculiformis saplings / Padrão de distribuição de artrópodes nas superfícies foliares de plantas de Acacia auriculiformis

Acacia auriculiformis A. Cunn. Ex Benth. (Fabaceae), a non-native pioneer species in Brazil with fast growth and rusticity, is used in restoration programs. Our goal was to assess during a 24-month survey the pattern of arthropods (phytophagous insects, bees, spiders, and predator insects) on the leaf surfaces of A. auriculiformis saplings. Fourteen species of phytophagous, two of bees and eleven of predators were most abundant on the adaxial surface. The values of the ecological indexes (abundance, diversity, and species richness) and the rarefaction, and k-dominance curves of phytophagous, bees and arthropod predators were highest on the adaxial leaf surface of A. auriculiformis. The k-dominance and abundance of Aleyrodidae (Hemiptera) (both leaf surfaces), the native stingless bee Tetragonisca angustula Latreille (Hymenoptera: Apidae) (both leaf surfaces) and the ant Brachymyrmex sp. (adaxial surface) and Pheidole sp. (Hymenoptera: Formicidae) (abaxial surface) were the highest between the taxonomic groups of phytophagous, bees, and predators, respectively on A. auriculiformis saplings. The ecological indexes and rarefaction, abundance, and k-dominance curves of phytophagous insects, bees, and predators were highest on the adaxial leaf surface. The preference of phytophagous insects for the adaxial leaf surface is probably due to the lower effort required to move on this surface. Understanding the arthropod preferences between leaf surfaces may help to develop sampling and pest management plans for the most abundant phytophagous insects on A. auriculiformis saplings. Also, knowledge on the preference pattern of bees and predators may be used to favour their conservation.

Acacia auriculiformis A. Cunn. Ex Benth. (Fabaceae), espécie pioneira com rápido crescimento e rusticidade, é utilizada em programas de recuperação de áreas degradadas. O objetivo deste trabalho foi avaliar, durante 24 meses, o padrão de distribuição de artrópodes (insetos fitófagos, abelhas, aranhas e insetos predadores) nas superfícies foliares de A. auriculiformis. Quatorze espécies de fitófagos, duas de abelhas e onze de predadores foram mais abundantes na superfície adaxial. Índices ecológicos (abundância, diversidade e riqueza de espécies) e curvas de rarefação e dominância-k de fitófagos, abelhas e artrópodes predadores foram maiores na face adaxial de folhas de A. auriculiformis. A dominância-k e a abundância de Aleyrodidae (Hemiptera) (ambas as superfícies foliares), da abelha nativa sem ferrão Tetragonisca angustula Latreille (Hymenoptera: Apidae) (ambas as superfícies foliares) e das formigas Brachymyrmex sp. (superfície adaxial) e Pheidole sp. (Hymenoptera: Formicidae) (superfície abaxial) foram as maiores entre os grupos taxonômicos de fitófagos polinizadores e predadores, respectivamente, em plantas jovens de A. auriculiformis. A abundância, diversidade e riqueza e as curvas de rarefação e dominância-k de artrópodes fitófagos, abelhas e predadores foram maiores nas superfícies adaxiais das folhas dessa árvore. A preferência pela superfície adaxial da folha se deve, provavelmente, ao menor esforço para se movimentarem na mesma. Compreender as preferências dos artrópodes pelas superfícies foliares pode auxiliar no desenvolvimento de planos de amostragem e manejo de pragas em A. auriculiformis. Além disso, o conhecimento da distribuição de abelhas e predadores pode favorecer a conservação desses insetos.

Egg production and life history of Alona guttata Sars, 1862 (Cladocera, Chydoridae): implications for colonization of temporary ponds / Produção de ovos e história de vida de Alona guttata Sars, 1862 (Cladocera, Chydoridae): implicações para a colonização de lagoas temporárias

Cladocerans are a diverse group of species that show rapid responses to changes in environmental conditions. This adaptive capacity has important implications for egg production and life cycle, especially in transitory environments such as temporary waterbodies. The present study investigated the life history and egg production of Alona gutatta Sars, 1862 (Crustacea, Cladocera), an abundant and frequent species from a high-altitude temporary pond (Lagoa Seca, Minas Gerais, Brazil). Newly hatched neonates were monitored in relation to time of maturation, number of eggs produced per female and time of survival. Neonates required a mean of 8 days to mature. A. guttata survived for a mean of 30.9 ± 8.1 days and produced 2 eggs per brood, generating a mean of 10.95 ± 6.41 neonates during the entire life cycle. The rapid development, short time to produce eggs and long life cycle are important adaptations to the adverse environmental conditions of temporary aquatic environments, which can contribute to the rapid colonization of Alona guttata in transitory ecosystems.

Os cladóceros formam um grupo diverso de espécies que apresentam respostas rápidas às mudanças nas condições ambientais. Essa capacidade adaptativa tem implicações importantes para a produção de ovos e para o ciclo de vida, especialmente em ambientes transitórios, como corpos d’água temporários. O presente estudo investigou a história de vida e a produção de ovos da espécie Alona guttata Sars, 1862 (Crustacea, Cladocera) coletada em uma lagoa temporária de altitude (Minas Gerais, Brasil). Organismos recém eclodidos foram observados (em condições de laboratório) em relação ao tempo de maturação, número de ovos produzidos por fêmea e tempo de sobrevivência. Os neonatos levaram em média 8 dias para atingir o primeiro estágio de maturação. As fêmeas produziram 2 ovos por ninhada e geraram uma média de 10,95 ± 6,41 neonatos durante todo o ciclo de vida. A. guttata apresentou uma média de 30,9 ± 8,1 dias de sobrevivência. O rápido desenvolvimento, o pouco tempo para a produção de ovos e o ciclo de vida longo são adaptações importantes às condições ambientais adversas de ambientes aquáticos temporários, que podem contribuir para a rápida colonização de Alona guttata em ecossistemas transitórios.

Field efficacy of Bt cotton containing events DAS-21023-5 × DAS-24236-5 × SYN-IR102-7 against lepidopteran pests and impact on the non-target arthropod community in Brazil.

The efficacy and non-target arthropod effects of transgenic DAS-21023-5 × DAS-24236-5 × SYN-IR102-7 Bt cotton, expressing proteins Cry1Ac, Cry1F and Vip3Aa19, was examined through field trials in Brazil. Fifteen field efficacy experiments were conducted from 2014 through the 2020 growing season across six different states in Brazil to evaluate performance against key lepidopteran pests through artificial infestations of Chrysodeixis includens (Walker), Spodoptera frugiperda (J.E. Smith,1797), Spodoptera cosmioides (Walker, 1858) and Chloridea virescens (F., 1781), and natural infestations of Alabama argillacea (Hübner) and S. frugiperda. The impact of this Bt cotton technology on the non-target arthropod community in Brazilian cotton production systems was also assessed in a multi-site experiment. DAS-21023-5 × DAS-24236-5 × SYN-IR102-7 cotton significantly reduced the feeding damage caused by S. frugiperda, S. cosmioides, C. includens, C. virescens and A. argillacea, causing high levels of mortality (greater than 99%) to all target lepidopteran pests evaluated during vegetative and/or reproductive stages of crop development. Non-target arthropod community-level analyses confirmed no unintended effects on the arthropod groups monitored. These results demonstrate the value of transgenic Bt cotton containing event DAS-21023-5 × DAS-24236-5 × SYN-IR102-7 for consideration as part of an integrated approach for managing key lepidopteran pests in Brazilian cotton production systems.

Anatomical changes in postembryonic development of Pycnogonum litorale.

Sea spiders (Pycnogonida) are a small group of arthropods, sister to other chelicerates. They have an unusual adult bauplan, oligosegmented larvae, and a protracted postembryonic development. Pycnogonum litorale (Strøm, 1762) is an uncommonly long-lived sea spider with a distinctive protonymphon and adult anatomy. Although it was described ~250 years ago, little is known about its internal organization and development. We examined the anamorphic and early epimorphic development of this species using histology, light microscopy, and SEM, and provide the first comprehensive anatomical study of its many instars. Postembryonic development of P. litorale includes transformations typical of pycnogonids: reorganization of the larval organs (digestive, nervous, secretory), formation of the abdomen, trunk segments (+ appendages), primary body cavity and reproductive system. Specific traits include the accelerated articulation of the walking legs, formation of the subesophageal and posterior synganglia, and the system of twin midgut diverticula. In addition, P. litorale simultaneously lose the spinning apparatus and all larval appendages. We found that developmental changes occur in synchrony with changes in ecology and food sources. The transition from the anamorphic to the epimorphic period in particular is marked by considerable anatomical and lifestyle shifts. HIGHLIGHTS: Postembryonic development of P. litorale includes numerous anamorphic and epimorphic stages. The instars acquire abdomen, trunk segments, body cavity, and gonads, while losing all larval appendages. Developmental changes are synchronized with changes in lifestyle and food sources.

Impacts of thermal fluctuations on heat tolerance and its metabolomic basis in Arabidopsis thaliana, Drosophila melanogaster, and Orchesella cincta.

Temperature varies on a daily and seasonal scale and thermal fluctuations are predicted to become even more pronounced under future climate changes. Studies suggest that plastic responses are crucial for species' ability to cope with thermal stress including variability in temperature, but most often laboratory studies on thermal adaptation in plant and ectotherm organisms are performed at constant temperatures and few species included. Recent studies using fluctuating thermal regimes find that thermal performance is affected by both temperature mean and fluctuations, and that plastic responses likely will differ between species according to life strategy and selective past. Here we investigate how acclimation to fluctuating or constant temperature regimes, but with the same mean temperature, impact on heat stress tolerance across a plant (Arabidopsis thaliana) and two arthropod species (Orchesella cincta and Drosophila melanogaster) inhabiting widely different thermal microhabitats and with varying capability for behavioral stress avoidance. Moreover, we investigate the underlying metabolic responses of acclimation using NMR metabolomics. We find increased heat tolerance for D. melanogaster and A. thaliana exposed to fluctuating acclimation temperatures, but not for O. cincta. The response was most pronounced for A. thaliana, which also showed a stronger metabolome response to thermal fluctuations than both arthropods. Generally, sugars were more abundant across A. thaliana and D. melanogaster when exposed to fluctuating compared to constant temperature, whereas amino acids were less abundant. This pattern was not evident for O. cincta, and generally we do not find much evidence for similar metabolomics responses to fluctuating temperature acclimation across species. Differences between the investigated species' ecology and different ability to behaviorally thermoregulate may have shaped their physiological responses to thermal fluctuations.

Inferring Evolutionary Timescales without Independent Timing Information: An Assessment of "Universal" Insect Rates to Calibrate a Collembola (Hexapoda) Molecular Clock.

Previous estimates of nucleotide substitution rates are routinely applied as secondary or "universal" molecular clock calibrations for estimating evolutionary timescales in groups that lack independent timing information. A major limitation of this approach is that rates can vary considerably among taxonomic groups, but the assumption of rate constancy is rarely evaluated prior to using secondary rate calibrations. Here I evaluate whether an insect mitochondrial DNA clock is appropriate for estimating timescales in Collembola-a group of insect-like arthropods characterized by high levels of cryptic diversity. Relative rates of substitution in cytochrome oxidase subunit 1 (COI) were inferred via Bayesian analysis across a topologically constrained Hexapod phylogeny using a relaxed molecular clock model. Rates for Collembola did not differ significantly from the average rate or from the rates estimated for most other groups (25 of 30), suggesting that (1) their apparent cryptic diversity cannot be explained by accelerated rates of molecular evolution and (2) clocks calibrated using "universal" insect rates may be appropriate for estimating evolutionary timescales in this group. However, of the 31 groups investigated, 10 had rates that deviated significantly from the average (6 higher, 4 lower), underscoring the need for caution and careful consideration when applying secondary insect rate calibrations. Lastly, this study exemplifies a relatively simple approach for evaluating rate constancy within a taxonomic group to determine whether the use of secondary rates are appropriate for molecular clock calibrations.

Life after the mother's hug: Late post-embryonic development of Cryptops parisi (Chilopoda: Scolopendromorpha: Cryptopidae).

Here we describe in detail the late post-embryonic development of the common European scolopendromorph centipede Cryptops parisi. Canonical variate analyses of two groups of external morphological characters, viz., cephalic capsule characters (head length, length of the anterior and posterior paramedian cephalic sutures) and coxopleuron surface characters (number of pores in the coxal pore-field, number of setae on the posterior coxopleuron edge, their number on the coxal pore-field, and their number posterior to the coxal pore-field) were conducted on a large sample of specimens collected from two localities in Serbia. Ten free-living stages are recognized: three pre-adult stages (adolescens I, II, and III) and seven adult stages (one maturus junior stage, four maturus, and two maturus senior stages). The fourth late post-embryonic stage is the first mature stage in both sexes. Sexual dimorphism in the aforementioned characters was not observed. Morphological variation of coxopleuron characters was more informative for the discrimination of developmental stages in Cryptops than the morphological variation of cephalic capsule characters.

Evolutionary dynamics of gene regulation.

The long controversy over the importance of changes in the regulatory genome has been resolved with the recognition that such changes are a fundamental component of evolutionary dynamics. Comparative studies have revealed four dominant modes of change as the regulatory genome evolved: (1) the origin of regulatory novelties such as distal enhancers and new types of promoters at the origin of Metazoa; (2) the expansion of regulatory capacity, most notably with diversification of transcription factors. Together these changes expanded the available combinatoric complexity of regulatory interactions and allow an increase in the variety of cell types. There are two more common modes of regulatory evolution: (3) Repatterning of gene regulatory networks. Such repatterning largely involves the introduction of transposons, promoter switching, co-option of regulatory genes or subcircuits, recombination, and the de novo generation of new regulatory sequences. Finally, (4) changes in enhancer and promoter specificity enable fine-scale adaptive changes. One of the outstanding issues at the intersection of evolutionary and developmental biology is how these various modes of regulatory evolution translate to morphological change, and particularly macro- and microevolutionary patterns and whether evolutionary novelties are associated with distinctive patterns of regulatory change.

Responses of soil microarthropod taxon (Hexapoda: Protura) to natural disturbances and management practices in forest-dominated subalpine lake catchment areas.

Disturbances are intrinsic drivers of structure and function in ecosystems, hence predicting their effects in forest ecosystems is essential for forest conservation and/or management practices. Yet, knowledge regarding belowground impacts of disturbance events still remains little understood and can greatly vary by taxonomic and functional identity, disturbance type and local environmental conditions. To address this gap in knowledge, we conducted a survey of soil-dwelling Protura, across forests subjected to different disturbance regimes (i.e. windstorms, insect pest outbreaks and clear-cut logging). We expected that the soil proturan assemblages would differ among disturbance regimes. We also hypothesized that these differences would be driven primarily by variation in soil physicochemical properties thus the impacts of forest disturbances would be indirect and related to changes in food resources. To verify that sampling included two geographically distant subalpine glacial lake catchments that differed in underlying geology, each having four different types of forest disturbance, i.e. control, bark beetle outbreak (BB), windthrow + BB (wind + BB) and clear-cut. As expected, forest disturbance had negative effects on proturan diversity and abundance, with multiple disturbances having the greatest impacts. However, differences in edaphic factors constituted a stronger driver of variability in distribution and abundance of proturans assemblages. These results imply that soil biogeochemistry and resource availability can have much stronger effects on proturan assemblages than forest disturbances.

The ecology of marine colonization by terrestrial arthropods.

Terrestrial arthropods often colonized and became important in freshwater ecosystems, but did so less often and with little consequence in marine habitats. This pattern cannot be explained by the physical properties of water alone or by limitations of the terrestrial arthropod body plan alone. One hypothesis is that transitions among terrestrial, aquatic and marine ecosystems are unlikely when well-adapted incumbent species in the recipient realm collectively resist entry by initially less well adapted newcomers. I evaluated and modified this hypothesis by examining the properties of donor and recipient ecosystems and the roles that insects play or do not play in each. I argue that the insularity and diminished competitiveness of most freshwater ecosystems makes them vulnerable to invasion from land and sea, and largely prevent transitions from freshwater to terrestrial and marine habitats by arthropods. Small terrestrial arthropods emphasize high locomotor performance and long-distance communication, traits that work less well in the denser, more viscous medium of water. These limitations pose particular challenges for insects colonizing highly escalated marine ecosystems, where small incumbent species rely more on passive than on active defences. Predatory insects are less constrained than herbivores, wood-borers, filter-feeders, sediment burrowers and social species.

Ecdysteroids are present in the blood of wild passerine birds.

Ecdysteroids (arthropod molting hormones) play an important role in the development and sexual maturation of arthropods, and they have been shown to have anabolic and "energizing" effect in higher vertebrates. The aim of this study was to assess ecdysteroid diversity, levels according to bird species and months, as well as to observe the molting status of hard ticks (Acari: Ixodidae) infesting the birds. Therefore, blood samples and ticks were collected from 245 birds (244 songbirds and a quail). Mass spectrometric analyses showed that 15 ecdysteroids were regularly present in the blood samples. Molting hormones biologically most active in insects (including 20-hydroxyecdysone [20E], 2deoxy-20E, ajugasterone C and dacryhainansterone) reached different levels of concentration according to bird species and season. Similarly to ecdysteroids, the seasonal presence of affected, apolytic ticks peaked in July and August. In conclusion, this study demonstrates the presence of a broad range and high concentrations of ecdysteroids in the blood stream of wild-living passerine birds. These biologically active, anabolic compounds might possibly contribute to the known high metabolic rate of songbirds.

Developing an integrated understanding of the evolution of arthropod segmentation using fossils and evo-devo.

Segmentation is fundamental to the arthropod body plan. Understanding the evolutionary steps by which arthropods became segmented is being transformed by the integration of data from evolutionary developmental biology (evo-devo), Cambrian fossils that allow the stepwise acquisition of segmental characters to be traced in the arthropod stem-group, and the incorporation of fossils into an increasingly well-supported phylogenetic framework for extant arthropods based on genomic-scale datasets. Both evo-devo and palaeontology make novel predictions about the evolution of segmentation that serve as testable hypotheses for the other, complementary data source. Fossils underpin such hypotheses as arthropodization originating in a frontal appendage and then being co-opted into other segments, and segmentation of the endodermal midgut in the arthropod stem-group. Insights from development, such as tagmatization being associated with different modes of segment generation in different body regions, and a distinct patterning of the anterior head segments, are complemented by palaeontological evidence for the pattern of tagmatization during ontogeny of exceptionally preserved fossils. Fossil and developmental data together provide evidence for a short head in stem-group arthropods and the mechanism of its formation and retention. Future breakthroughs are expected from identification of molecular signatures of developmental innovations within a phylogenetic framework, and from a focus on later developmental stages to identify the differentiation of repeated units of different systems within segmental precursors.

Antimicrobial activity and mechanism of action of a novel peptide present in the ecdysis process of centipede Scolopendra subspinipes subspinipes.

One of the most important cellular events in arthropods is the moulting of the cuticle (ecdysis). This process allows them to grow until they reach sexual maturity. Nevertheless, during this stage, the animals are highly exposed to pathogens. Consequently, it can be assumed that arthropods counter with an efficient anti-infective strategy that facilitates their survival during ecdysis. Herein, we characterized a novel antimicrobial peptide called Pinipesin, present in the exuviae extract of the centipede Scolopendra subspinipes subspinipes. The antimicrobial activity of Pinipesin was tested. The haemolytic activity of the peptide was evaluated and its possible mechanism of action was investigated. Identification was carried out by mass spectrometry analysis. Pinipesin displayed potent antimicrobial effects against different microorganisms and showed low haemolytic effects against human erythrocytes at high concentrations. It has a monoisotopic mass of 1213.57 Da, its sequence exhibited high similarity with some cuticular proteins, and it might act intracellularly by interfering with protein synthesis. Our data suggest that Pinipesin might be part of a prophylactic immune response during the ecdysis process of centipedes. Therefore, it is a promising candidate for the development of non-conventional antibiotics that could help fight infectious diseases and represents an exciting discovery for this taxon.

Ancient origins of arthropod moulting pathway components.

Ecdysis (moulting) is the defining character of Ecdysoza (arthropods, nematodes and related phyla). Despite superficial similarities, the signalling cascade underlying moulting differs between Panarthropoda and the remaining ecdysozoans. Here, we reconstruct the evolution of major components of the ecdysis pathway. Its key elements evolved much earlier than previously thought and are present in non-moulting lophotrochozoans and deuterostomes. Eclosion hormone (EH) and bursicon originated prior to the cnidarian-bilaterian split, whereas ecdysis-triggering hormone (ETH) and crustacean cardioactive peptide (CCAP) evolved in the bilaterian last common ancestor (LCA). Identification of EH, CCAP and bursicon in Onychophora and EH, ETH and CCAP in Tardigrada suggests that the pathway was present in the panarthropod LCA. Trunk, an ancient extracellular signalling molecule and a well-established paralog of the insect peptide prothoracicotropic hormone (PTTH), is present in the non-bilaterian ctenophore Mnemiopsis leidyi. This constitutes the first case of a ctenophore signalling peptide with homology to a neuropeptide.

Lethal and Sublethal Toxicity of Thiamethoxam and Clothianidin Commercial Formulations to Soil Invertebrates in a Natural Soil.

The use of neonicotinoids in agriculture is a critical environmental protection issue. Although there has been considerable research on pollinator exposure and aquatic toxicological effects, few studies have investigated the chronic impacts on soil-dwelling species. Given the application of neonicotinoids into soil systems, there is the potential for risk to soil invertebrates. The toxicity of 2 commercial formulations containing the active ingredients (a.i.) thiamethoxam (Actara® 240SC) or clothianidin (Titan™) was investigated using 3 soil invertebrate species: Oppia nitens, Eisenia andrei, and Folsomia candida. No adverse effects were observed for O. nitens at the highest tested concentrations (≥92 mg a.i./kg dry soil) after a 28-d exposure. Exposure to clothianidin resulted in a 28-d median inhibitory concentration (IC50) of 0.069 (95% confidence limits: 0.039-0.12) mg/kg dry soil for F. candida, and a 56-d IC50 of 0.26 (0.22-3.2) mg a.i./kg dry soil for E. andrei. Exposure to thiamethoxam was less toxic, with IC50s of 0.36 (0.19-0.66) and 3.0 (2.2-4.0) mg a.i./kg dry soil for F. candida and E. andrei reproduction, respectively. The observed toxicity for F. candida adult survival and reproduction and for E. andrei reproduction occurred at environmentally relevant concentrations. However, because clothianidin is a degradation product of thiamethoxam, and detection of clothianidin rose to levels of concern in the thiamethoxam-amended soils over time, the observed toxicity may be partly attributed to the presence of clothianidin. Environ Toxicol Chem 2019;38:2111-2120. © 2019 Crown in the right of Canada. Published by Wiley Periodicals Inc. on behalf of SETAC.

Effects of Soil Properties on the Toxicity and Bioaccumulation of Lead in Soil Invertebrates.

The present study examined the effects of soil physical and chemical properties on the toxicity of lead (Pb) to earthworms (Eisenia fetida) and collembolans (Folsomia candida), and on bioaccumulation of Pb by earthworms, in soils amended with Pb salts. Toxicity tests were conducted in 7 soils varying in soil properties (pH 4.7-7.4, effective cation exchange capacity [eCEC] 4-42 cmolc /kg, organic carbon 10-50 g C/kg) that were leached and pH corrected after spiking with PbCl2 . The median effect concentrations (EC50s) based on total soil Pb concentrations ranged from 35 to 5080 mg Pb/kg for earthworms and 389 to >7190 mg/kg for Collembola. Significant positive correlations were observed between log (EC50) for earthworm reproduction and log (eCEC, total C, exchangeable Ca and Mg, or clay content), but no significant correlations were observed between Pb toxicity to Collembola and soil properties. Expressing Pb dose as either the free ion (Pb2+ ) activity in porewater or as the measured dissolved porewater concentration of Pb did not explain differences in toxicity among soils. The bioaccumulation factors (BAFs) for Pb in earthworms ranged up to >10-fold across 6 soil treatments, with a median of 0.16, and the BAF was significantly correlated with eCEC (p = 0.038, r = -0.84), but not with any other soil properties. Soil properties related to eCEC (total C, exchangeable Ca and Mg, clay content) had a significant effect on Pb toxicity and bioaccumulation in earthworms, but no relationship was found for Collembola. As a major soil property affecting the bioavailability of Pb, CEC should be incorporated into any soil hazard assessment of Pb as a modifying factor of toxicity and bioaccumulation for earthworms. Environ Toxicol Chem 2019;38:1486-1494. © 2019 SETAC.

Developmental expression and evolution of hexamerin and haemocyanin from Folsomia candida (Collembola).

Haemocyanins constitute a group of copper-containing respiratory proteins, and hexamerins were derived from hexapod haemocyanin but lost the ability to transport oxygen and serve as storage proteins. Although hexamerins have been reported in most insect species, none of them has been identified in Collembola, one of the most primitive hexapod lineages, thereby preventing us from exploring relevant evolutionary scenarios regarding the origin and evolution of hexamerins in hexapods. Here we report on collembolan hexamerins for the first time, and investigated the temporal expression profiles of hexamerin and haemocyanin in the collembolan Folsomia candida. Haemocyanin was expressed over the entire life cycle, with higher expression at the embryonic stage than at other stages, whereas hexamerin expression was restricted to embryos, unlike insect hexamerins, which are generally expressed from larval to adult stages. A phylogenetic analysis and molecular clock estimation suggested that all investigated hexapod hexamerins have a single and ancient origin (~423 Ma), coincident with the rise of atmospheric oxygen levels in the Silurian-Devonian period, indicating a physiological link between molecular evolution and Palaeozoic oxygen changes.

Ecdysis in a stem-group euarthropod from the early Cambrian of China.

Moulting is a fundamental component of the ecdysozoan life cycle, but the fossil record of this strategy is susceptible to preservation biases, making evidence of ecdysis in soft-bodied organisms extremely rare. Here, we report an exceptional specimen of the fuxianhuiid Alacaris mirabilis preserved in the act of moulting from the Cambrian (Stage 3) Xiaoshiba Lagerstätte, South China. The specimen displays a flattened and wrinkled head shield, inverted overlap of the trunk tergites over the head shield, and duplication of exoskeletal elements including the posterior body margins and telson. We interpret this fossil as a discarded exoskeleton overlying the carcass of an emerging individual. The moulting behaviour of A. mirabilis evokes that of decapods, in which the carapace is separated posteriorly and rotated forward from the body, forming a wide gape for the emerging individual. A. mirabilis illuminates the moult strategy of stem-group Euarthropoda, offers the stratigraphically and phylogenetically earliest direct evidence of ecdysis within total-group Euarthropoda, and represents one of the oldest examples of this growth strategy in the evolution of Ecdysozoa.