An evolutionary timescale for terrestrial isopods and a lack of molecular support for the monophyly of Oniscidea (Crustacea: Isopoda)

The marine metazoan fauna first diversified in the early Cambrian, but terrestrial environments were not colonized until at least 100 million years later. Among the groups of organisms that successfully colonized land is the crustacean order Isopoda. Of the 10,000 described isopod species, ~ 3,600 species from the suborder Oniscidea are terrestrial. Although it is widely thought that isopods colonized land only once, some studies have failed to confirm the monophyly of Oniscidea. To infer the evolutionary relationships among isopod lineages, we conducted phylogenetic analyses of nuclear 18S and 28S and mitochondrial COI genes using maximum-likelihood and Bayesian methods. We also analyzed a second data set comprising all of the mitochondrial protein-coding genes from a smaller sample of isopod taxa. Based on our analyses using a relaxed molecular clock, we dated the origin of terrestrial isopods at 289.5 million years ago (95% credibility interval 219.6–358.9 million years ago). These predate the known fossil record of these taxa and coincide with the formation of the supercontinent Pangaea and with the diversification of vascular plants on land. Our results suggest that the terrestrial environment has been colonized more than once by isopods. The monophyly of the suborder Oniscidea was not supported in any of our analyses, conflicting with classical views based on morphology. This draws attention to the need for further work on this group of isopods.     

Locomotory,ventilatory and metabolic responses of the subterranean Stenasellus virei (Crustacea,Isopoda) to severe hypoxia and subsequent recovery

The locomotory and ventilatory activities and the intermediary and energy metabolism modifications of the hypogean aquatic isopod crustacean Stenasellus virei were investigated in severe hypoxia (Po₂ < 0.03 kPa) and subsequent recovery. The aims of this study were i) to determine why the subterranean species displayed a greater tolerance of hypoxia than numerous other epigean crustaceans, ii) to confirm previous results obtained with four hypogean and epigean crustaceans, iii) to compare the responses to severe hypoxia in hypogean amphipods and isopods, and iv) to better understand the ecological problems of the hypogean organisms survival in subterranean habitats. S. virei responded to experimental long-term, severe hypoxia with classical anaerobic metabolism mainly characterized by a decrease in adenosine triphosphate (ATP) and phosphagen, utilization of glycogen and glutamate, and accumulation of lactate and alanine. Lactate was also largely excreted by this organism, which is unusual for crustaceans in general. Compared to most other epigean crustaceans, the isopod S. virei showed high amounts of stored glycogen and arginine phosphate. These differences in glycogen and phosphagen stores, and the ability to reduce energetic expenditures linked to locomotion and ventilation, extended the survival of S. virei under experimental anaerobiosis. During recovery, the isopod S. virei showed a higher capacity for glyconeogenesis from lactate and a faster and total replenishment of ATP and arginine phosphate levels than epigean crustaceans. Data concerning responses to hypoxia and subsequent recovery in S. virei are similar to those previously obtained with two other hypogean amphipods, except that this isopod did not synthesize succinate in anaerobiosis.     

Macrofaunal crustaceans in the benthic boundary layer from the shelf break to abyssal depths in the Ross Sea (Antarctica)

The New Zealand International Polar Year–Census of Antarctic Marine Life (NZ IPY-CAML) project added to previous benthic studies in the Ross Sea by extending sampling north from the continental shelf to previously unsampled areas of the shelf break, slope, abyssal plain and seamounts in the region. The aim of the current study is to give first insights into the deep-sea community structure of the Ross Sea focussing on a component of the benthic boundary layer that is macrofaunal crustaceans collected one metre above the seafloor. We assess changes in Ross Sea crustacean community composition from the shelf break (474 m) to the abyss (3,490 m) and compare the Ross Sea crustacean fauna to areas elsewhere in the Southern Ocean. Analyses reveal high relative abundances, suggesting an important role in the food web. Among the peracarid crustaceans, there was a decline in the proportion of amphipods with increasing depth. Three of 15 isopod families (Acanthaspidiidae, Nannoniscidae and Desmosomatidae) were identified to species level and about 72 % of the species were new to science. Isopod diversity in the Ross Sea abyss appears to be comparable to that in the highly speciose Weddell deep sea. Standardised sampling of these crustacean communities allows setting the biodiversity of the Ross Sea into a global context.     

Diversity of isopods (Crustacea): new data from the Arctic and Atlantic Oceans

New data on the diversity pattern of isopods (Crustacea) from the northern most part of the North Atlantic and the Arctic Oceans is presented. The pattern of diversity with depth is similar at depths <1000m, but differs considerably below about 1000m. In the Arctic the diversity of isopods (expressed both as numbers of species per sled and expected number of species) increased with increased depth to a maximum at depths of about 320 to 1100m, but then declined towards deeper waters. There was a significant increase in numbers per sled and in the expected number of species with increased depth in the northernmost part of the North Atlantic Ocean. Additionally, changes occurred in the relative composition of the shallow and deep water fauna, with asellote isopods being relatively larger part of the isopod fauna in the Arctic than in the northern most part of the North Atlantic. This indicates major faunistic changes occurring at the Greenland-Iceland-Faeroe Ridge, possibly caused by rapid changes in the temperature. Furthermore, that the low diversity of the Arctic deep-sea is a regional phenomenon, and not a part of a large scale latitudinal pattern in the North Atlantic.     

Fossil isopods associated with a fish skeleton from the Lower Cretaceous of Queensland,Australia – direct evidence of a scavenging lifestyle in Mesozoic Cymothoida

Abstract: A dense assemblage of fossil isopod crustaceans (Brunnaega tomhurleyi Wilson, sp. nov.) from the Lower Cretaceous (Albian) Toolebuc Formation of Queensland, Australia, has been found within the carcass of a large actinopterygian fish, Pachyrhizodus marathonensis (Etheridge). Preservation of fine anatomical details supports referral to the genus Brunnaega Polz, which is herein reassigned to the family Cirolanidae. Furthermore, placement of this taxon within the cirolanid subfamily Conilerinae Kensley and Schotte is significant because the group includes modern species that are well known as voracious scavengers. This isopod–fish association represents the oldest unequivocal evidence of scavenging by Mesozoic cymothoidean isopods on a large vertebrate carcass.     

Ancient Origin of the Modern Deep-Sea Fauna

The origin and possible antiquity of the spectacularly diverse modern deep-sea fauna has been debated since the beginning of deep-sea research in the mid-nineteenth century. Recent hypotheses, based on biogeographic patterns and molecular clock estimates, support a latest Mesozoic or early Cenozoic date for the origin of key groups of the present deep-sea fauna (echinoids, octopods). This relatively young age is consistent with hypotheses that argue for extensive extinction during Jurassic and Cretaceous Oceanic Anoxic Events (OAEs) and the mid-Cenozoic cooling of deep-water masses, implying repeated re-colonization by immigration of taxa from shallow-water habitats. Here we report on a well-preserved echinoderm assemblage from deep-sea (1000–1500 m paleodepth) sediments of the NE-Atlantic of Early Cretaceous age (114 Ma). The assemblage is strikingly similar to that of extant bathyal echinoderm communities in composition, including families and genera found exclusively in modern deep-sea habitats. A number of taxa found in the assemblage have no fossil record at shelf depths postdating the assemblage, which precludes the possibility of deep-sea recolonization from shallow habitats following episodic extinction at least for those groups. Our discovery provides the first key fossil evidence that a significant part of the modern deep-sea fauna is considerably older than previously assumed. As a consequence, most major paleoceanographic events had far less impact on the diversity of deep-sea faunas than has been implied. It also suggests that deep-sea biota are more resilient to extinction events than shallow-water forms, and that the unusual deep-sea environment, indeed, provides evolutionary stability which is very rarely punctuated on macroevolutionary time scales.     

The geological history of deep-sea colonization by echinoids: roles of surface productivity and deep-water ventilation

The origins and geological history of the modern fauna of deep-sea echinoids is explored using a combination of palaeontological and molecular data. We demonstrate that, whereas generalist omnivores have migrated into the deep sea in low numbers over the past 200 Myr, there was a short time-interval between approximately 75 and 55 Myr when the majority of specialist detritivore clades independently migrated off-shelf. This coincides with a marked increase in seasonality, continental run-off and surface water productivity, and suggests that increasing organic carbon delivery into ocean basins was an important controlling factor. Oceanic anoxic events, by contrast, appear to have played a subsidiary role in controlling deep-sea diversity.     

Trouble at the top? Restricted distribution and extreme population isolation in an alpine crustacean assemblage with unexpected lineage diversity

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Microplastic ingestion affects hydrogen production and microbiomes in the gut of the terrestrial isopod Porcellio scaber

Microplastic (MP) is an environmental burden and enters food webs via ingestion by macrofauna, including isopods (Porcellio scaber) in terrestrial ecosystems. Isopods represent ubiquitously abundant, ecologically important detritivores. However, MP-polymer specific effects on the host and its gut microbiota are unknown. We tested the hypothesis that biodegradable (polylactic acid [PLA]) and non-biodegradable (polyethylene terephthalate [PET]; polystyrene [PS]) MPs have contrasting effects on P. scaber mediated by changes of the gut microbiota. The isopod fitness after an 8-week MP-exposure was generally unaffected, although the isopods showed avoidance behaviour to PS-food. MP-polymer specific effects on gut microbes were detected, including a stimulation of microbial activity by PLA compared with MP-free controls. PLA stimulated hydrogen emission from isopod guts, while PET and PS were inhibitory. We roughly estimated 10⁷ kg year⁻¹ hydrogen emitted from the isopods globally and identified their guts as anoxic, significant mobile sources of reductant for soil microbes despite the absence of classical obligate anaerobes, likely due to Enterobacteriaceae-related fermentation activities that were stimulated by lactate generated during PLA-degradation. The findings suggest negative effects of PET and PS on gut fermentation, modulation of important isopod hydrogen emissions by MP pollution and the potential of MP to affect terrestrial food webs.     

Estimates of species diversity of free-living marine isopod crustaceans on coral reefs

 A core group of isopod crustacean genera appears to be present at many coral reef sites. Within these genera, however, species show high local endemicity. Based on the estimated percentage of endemism for the Atlantic (19% for individual sites, 90% for the tropical western Atlantic as a unit), the Indian Ocean (50%), the eastern-central Pacific (80%), and the western Pacific (40%), it is estimated that there are some 5,000 to 13,000 isopod species in the world’s coral reefs, and that some 2,000 to 6,000 of these are endemics. (At present, approximately 4,400 species of marine and 560 species of freshwater isopods have been described.) Based on the crudely estimated relative abundances of other peracaridan crustaceans in coral reefs (compared to isopods), the total diversity of reef amphipods, tanaidaceans, cumaceans, and mysidaceans is approximately 54,500 species. Accepted: 29 August 1997     

Adaptation to cadmium in three populations of the isopod Porcellio scaber

• 1.1. First-generation laboratory animals of three populations of the isopod Porcellio scaber, collected from a reference wood, a zinc smelter area and a lead mine site, were compared with respect to effects of Cd.
• 2.2. All populations reacted differently to Cd-contaminated food: increased Cd concentrations in the food resulted in decreased consumption and growth for the reference isopods; mine isopods were not affected by Cd and for the smelter animals Cd stimulated growth at an intermediate concentration.
• 3.3. Since Cd concentrations in the isopod did not differ between populations, adaptation is probably based on an increased detoxification capacity.
• 4.4. The assimilation of Cd did not affect the Cu or Zn content of the isopods although the adapted isopods regulated their Cu content on a lower level than reference isopods.

     

Occurrence and assemblage composition of millipedes (Myriapoda, Diplopoda) and terrestrial isopods (Crustacea, Isopoda, Oniscidea) in urban areas of Switzerland

Terrestrial isopods and millipedes, members of the invertebrate macro-decomposer guild, were collected through pitfall traps in three Swiss cities (Zurich, Lucerne, Lugano). A total of 7,198 individuals of 17 isopod species (7093 ind.), and 10 millipede species (105 ind.) were captured. Besides the Alpine endemic isopod (Trichoniscus alemannicus) and millipede (Cylindroiulus verhoeffi), urban assemblages were mainly composed of widespread, native European and even cosmopolitan species, which are frequent in anthropogenic areas. Overall species richness (isopods and millipedes combined) was similar in Zurich (17 species) and Lucerne (16), while only 13 species were sampled in Lugano. According to the Sørensen index of similarity, species composition of Zurich and Lucerne were more alike, while the one of Lugano was more distinct from the other two cities. This result can be explained by the spatial proximity of Zurich and Lucerne in the north of the Alps compared to Lugano, which is located more distantly and in the south of the Alps. Dominant isopods and millipedes in Zurich and Lucerne were found to be widespread synanthropic species in temperate Europe(Porcellio scaber, Trachelipus rathkii and Ophyiulus pilosus) while the dominant isopod in Lugano (Trachelipus razzautii) is a species with a north-eastern Mediterranean distribution. Our study reveals that the urban millipede and isopod fauna in Swiss cities mainly consists of widespread species, but species of narrower distribution (e.g. Trichoniscus alemannicus, Cylindroiulus verhoeffi) may also find suitable habitats in cities. Despite some signs of biotic homogenization, our study also found compositional differences of millipede and isopod assemblages between northern and southern cities that suggest geographical effects of the regional species pool.     

Examination of functional morphology of dajiid isopods using Arthrophryxus sp. parasitising a mysid shrimp as an example

Parasitic isopods have become specialised to many different host species and therefore show a wide variety of attachment and feeding specialisations. Often such structures are difficult to examine due to their small sizes which makes a more complete understanding of their functional morphology difficult. Here we present a new report and a first time high-resolution, non-SEM documentation of a parasitic epicaridean isopod, a female of the dajiid Arthrophryxus sp. from the Sea of Okhotsk. It was found during the SokhoBio 2015 on its host, a mysid shrimp (Holmsiella sp.). Arthrophryxus has only one formally described species, Arthrophryxus beringanus Richardson, 1908. With high-resolution documentation methods, we reveal new details of the morphological structures of mouthparts and thoracopods of this dajiid. Furthermore, we discuss the functional morphology of attachment to the host and feeding according to our new findings. We suggest that the thoracopods are involved in the attachment process even more than formerly assumed for different dajiids. The piercing-sucking mouthparts and the additional attachment mechanisms strongly indicate a permanent parasitism of the dajiid isopod. Permanent parasites affect the fitness of their hosts; therefore, deep-sea forms of dajiid isopods have direct impact on the deep-sea crustacean communities.     

Exoskeletal cuticle of cavernicolous and epigean terrestrial isopods: A review and perspectives

Comparative ultrastructural studies of the integument in terrestrial isopod crustaceans show that specific environmental adaptations of different eco-morphotypes are reflected in cuticle structure. The biphasic molting in isopods is a valuable experimental model for studies of cuticular matrix secretion and degradation in the same animal. The aim of this review is to show structural and functional adaptations of the tergal cuticle in terrestrial isopods inhabiting cave habitats. Exoskeletal cuticle thickness, the number of cuticular layers, epicuticle structure, mineralization, pigmentation and complexity of sensory structures are compared, with greater focus on the well-studied cave trichoniscid Titanethes albus. A large number of thinner cuticular layers in cave isopods compared to fewer thicker cuticular layers in related epigean species of similar body-sizes is explained as a specific adaptation to the cavernicolous life style. The epicuticle structure and composition are compared in relation to their potential waterproofing capacity in different environments. Cuticle mineralization is described from the functional point of view as well as from the aspect of different calcium storage sites and calcium dynamics during the molt cycle. We also discuss the nature and reduction of pigmentation in the cave environment and outline perspectives for future research.     

Effects of Essential Oils from Eucalyptus globulus Leaves on Soil Organisms Involved in Leaf Degradation

The replacement of native Portuguese forests by Eucalyptus globulus is often associated with deleterious effects on terrestrial and aquatic communities. Several studies have suggested that such a phenomenon is linked with the leaf essential oils released into the environment during the Eucalyptus leaf degradation process. However, to date, the way these compounds affect leaf degradation in terrestrial systems i.e. by direct toxic effects to soil invertebrates or indirectly by affecting food of soil fauna, is still unknown. In order to explore this question, the effect of essential oils extracted from E. globulus leaves on terrestrial systems was investigated. Fungal growth tests with species known as leaf colonizers (Mucor hiemalis, Alternaria alternata, Penicillium sp., Penicillium glabrum and Fusarium roseum) were performed to evaluate the antifungal effect of essential oils. In addition, a reproduction test with the collembolans Folsomia candida was done using a gradient of eucalyptus essential oils in artificial soil. The influence of essential oils on feeding behaviour of F. candida and the isopods Porcellio dilatatus was also investigated through food avoidance and consumption tests. Eucalyptus essential oils were lethal at concentrations between 2.5–20 µL/mL and inhibited growth of all fungal species between 1.25–5 µL/mL. The collembolan reproduction EC₅₀ value was 35.0 (28.6–41.2) mg/kg and both collembola and isopods preferred leaves without oils. Results suggested that the effect of essential oils in leaf processing is related to direct toxic effects on fungi and soil fauna and to indirect effects on the quality and availability of food to soil invertebrates.     

How Deep-Sea Wood Falls Sustain Chemosynthetic Life

Large organic food falls to the deep sea – such as whale carcasses and wood logs – are known to serve as stepping stones for the dispersal of highly adapted chemosynthetic organisms inhabiting hot vents and cold seeps. Here we investigated the biogeochemical and microbiological processes leading to the development of sulfidic niches by deploying wood colonization experiments at a depth of 1690 m in the Eastern Mediterranean for one year. Wood-boring bivalves of the genus Xylophaga played a key role in the degradation of the wood logs, facilitating the development of anoxic zones and anaerobic microbial processes such as sulfate reduction. Fauna and bacteria associated with the wood included types reported from other deep-sea habitats including chemosynthetic ecosystems, confirming the potential role of large organic food falls as biodiversity hot spots and stepping stones for vent and seep communities. Specific bacterial communities developed on and around the wood falls within one year and were distinct from freshly submerged wood and background sediments. These included sulfate-reducing and cellulolytic bacterial taxa, which are likely to play an important role in the utilization of wood by chemosynthetic life and other deep-sea animals.     

First glimpse into Lower Jurassic deep-sea biodiversity: in situ diversification and resilience against extinction

Owing to the assumed lack of deep-sea macrofossils older than the Late Cretaceous, very little is known about the geological history of deep-sea communities, and most inference-based hypotheses argue for repeated recolonizations of the deep sea from shelf habitats following major palaeoceanographic perturbations. We present a fossil deep-sea assemblage of echinoderms, gastropods, brachiopods and ostracods, from the Early Jurassic of the Glasenbach Gorge, Austria, which includes the oldest known representatives of a number of extant deep-sea groups, and thus implies that in situ diversification, in contrast to immigration from shelf habitats, played a much greater role in shaping modern deep-sea biodiversity than previously thought. A comparison with coeval shelf assemblages reveals that, at least in some of the analysed groups, significantly more extant families/superfamilies have endured in the deep sea since the Early Jurassic than in the shelf seas, which suggests that deep-sea biota are more resilient against extinction than shallow-water ones. In addition, a number of extant deep-sea families/superfamilies found in the Glasenbach assemblage lack post-Jurassic shelf occurrences, implying that if there was a complete extinction of the deep-sea fauna followed by replacement from the shelf, it must have happened before the Late Jurassic.     

A terrestrial micro-ecosystem for measuring effects of pollutants on isopod-mediated litter decomposition

Soil fauna-mediated litter decomposition was simulated in micro-ecosystems, using oak and poplar litter, with the isopod Porcellio scaber (Crustacea) as a representative soil fauna species. The aim was to identify the conditions under which decomposition rate and influence of isopods could be established accurately, yet keeping the system complex enough to be relevant for the field situation. The results showed that it was possible to divide total carbon dioxide production of a micro-ecosystem into three components: microbial respiration, isopod respiration and extra microbial, isopod-mediated respiration. The isopods stimulated the microbial respiration by 10–50%, depending on litter type and experimental design. It appears that this stimulation is achieved in two ways: 1. by increasing litter moisture content and 2. by production of faecal pellets. It is concluded that the micro-ecosystem described in this paper combines ecological relevance with reproducibility and simplicity of operation and thus provides a useful tool for measuring effects of soil contaminants.     

Ben Franklin temperate reef and deep sea `Agassiz Coral Hills' in the Blake Plateau off North Carolina

This paper deals with two deep water coral ecosystems off the North Carolina coast, both under the depositional, erosional, and biological influence of the Gulf Stream and its eddies and episodic upwelling processes. The first coral community, Ben Franklin temperate reef, is located at 20 meters in Onslow Bay and is characterized by the ahermatypic coral Oculina arbuscula Verrill and unusually high abundance of the small predatory isopod Eurydice bowmani George and Longerbeam. The deep sea coral community, Agassiz Coral Hills, is located over the Blake Plateau at 650 meters, dominated by the fossilized, dead and living coral Bathypsammia tintinnabulum, with rare occurrence of the deep sea solitary coral Flabellum goodei. The deep sea coral sites were sampled with an otter trawl, outfitted with `Benthos' flotation spheres. The results suggest that the fish and shrimp fauna exhibits high species richness in the Agassiz Coral Hills off North Carolina in comparison with a control site over the Blake Plateau off Florida. Species richness may be linked to episodic upwelling events along the Gulf Stream meanders. The fish fauna in the Agassiz Coral Hills includes the following two commercially important deep sea fish species: the wreck fish Polyprion americanus and the eel Synaphobranchus koupi. The Ben Franklin reef and the Agassiz Coral Hills are recommended as Marine Protected Areas.     

Evolutionary transition to XY sex chromosomes associated with Y-linked duplication of a male hormone gene in a terrestrial isopod

Sex chromosomes are highly variable in some taxonomic groups, but the evolutionary mechanisms underlying this diversity are not well understood. In terrestrial isopod crustaceans, evolutionary turnovers in sex chromosomes are frequent, possibly caused by Wolbachia, a vertically-transmitted endosymbiont causing male-to-female sex reversal. Here, we use surgical manipulations and genetic crosses, plus genome sequencing, to examine sex chromosomes in the terrestrial isopod Trachelipus rathkei. Although an earlier cytogenetics study suggested a ZZ/ZW sex chromosome system in this species, we surprisingly find multiple lines of evidence that in our study population, sex is determined by an XX/XY system. Consistent with a recent evolutionary origin for this XX/XY system, the putative male-specific region of the genome is small. The genome shows evidence of Y-linked duplications of the gene encoding the androgenic gland hormone, a major component of male sexual differentiation in isopods. Our analyses also uncover sequences horizontally acquired from past Wolbachia infections, consistent with the hypothesis that Wolbachia may have interfered with the evolution of sex determination in T. rathkei. Overall, these results provide evidence for the co-occurrence of multiple sex chromosome systems within T. rathkei, further highlighting the relevance of terrestrial isopods as models for the study of sex chromosome evolution.Subject terms: Evolutionary genetics, Genome evolution