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Background Dispersal is essential for terrestrial organisms living in disjunct habitats and constitutes a significant challenge for the evolution of wingless taxa. Springtails (Collembola), the sister-group of all insects (with Diplura), are reported since the Lower Devonian and are thought to have originally been subterranean. The order Symphypleona is reported since the early Cretaceous with genera distributed on every continent. This distribution implies an ability to disperse over oceans, however symphypleonan Collembola have never been reported in marine water contrary to other springtail orders. Despite being highly widespread, modern springtails are rarely reported in any kind of biotic association. Interestingly, the fossil record has provided occasional occurrences of Symphypleona attached by the antennae onto the bodies of larger arthropods. Results Here, we document the case of a ~ 16 Ma old fossil association: a winged termite and ant displaying not some, but 25 springtails attached or in close proximity to the body. The collembola exhibit rare features for fossils, reflecting their courtship and phoretic behaviours. By observing the modes of attachment of springtails on different arthropods, the sex representation and ratios in springtail antennal anatomies in new and previously reported cases, we infer a likely mechanism for dispersal in Symphypleona. By revealing hidden evidence of modern springtail associations with other invertebrates such as ants and termites, new compelling assemblages of fossil springtails, and the drastic increase of eusocial insects’ abundance during the Cenozoic (ants/termites comprising more than a third of insects in Miocene amber), we stress that attachment on winged castes of ants and termites may have been a mechanism for the worldwide dispersal of this significant springtail lineage. Moreover, by comparing the general constraints applying to the other wingless soil-dwelling arthropods known to disperse through phoresy, we suggest biases in the collection and observation of phoretic Symphypleona related to their reflexive detachment and infer that this behaviour continues today. Conclusions The specific case of tree resin entrapment represents the (so far) only condition uncovering the phoretic dispersal mechanism of springtails - one of the oldest terrestrial arthropod lineages living today.

Exceptional and extremely rare preservation of soft parts, eyes, or syn-vivo associations provide crucial palaeoecological information on fossil-rich deposits. Here we present exceptionally preserved specimens of the polychelidan lobster Voulteryon parvulus , from the Jurassic of La Voulte-sur-Rhône Fossil-Lagerstätte, France, bearing eyes with hexagonal and square facets, ovaries, and a unique association with epibiont thecideoid brachiopods, giving insights onto the palaeoenvironment of this Lagerstätte. The eyes, mostly covered in hexagonal facets are interpreted as either apposition eyes (poorly adapted to low-light environment) or, less likely, as refractive or parabolic superposition eyes (compatible with dysphotic palaeoenvironments). The interpretation that V. parvulus had apposition eyes suggests an allochthonous, shallow water origin. However, the presence of thecideoid brachiopod ectosymbionts on its carapace, usually associated to dim-light paleoenvironments and/or rock crevices, suggests that V. parvulus lived in a dim-light setting. This would support the less parsimonious interpretation that V. parvulus had superposition eyes. If we accept the hypothesis that V. parvulus had apposition eyes, since the La Voulte palaeoenvironment is considered deep water and had a soft substrate, V. parvulus could have moved into the La Voulte Lagerstätte setting. If this is the case, La Voulte biota would record a combination of multiple palaeoenvironments.

Here, we report fossil isopods preserved in laminated oil-shale mudstone (dysodile) from the Lower Cretaceous of Lebanon (Lower Barremian, 125 Ma, Grès du Liban Alloformation, Jezzine District). Based on a variety of proxies, their palaeoenvironments are determined to have been a shallow freshwater lake. The fossil isopods were studied using modern imaging techniques, such as multispectral imaging and photometric stereo, allowing for a detailed comparison of these specimens with comparable extant and fossil taxa. The conspecific fossils are herein recognized as remains of a new species — † Dysopodus gezei gen. et sp. nov.—of uncertain affinity within Cymothoida and bearing a strong resemblance to its non-parasitic lineages (Cirolanidae). A conspicuous pleotelson and uropod morphology set it apart from most species, with the notable exception of † Pseudoplakolana chiapaneca gen. nov. et comb. nov. from the Cretaceous of Mexico, originally attributed to an Australasian lineage (herein disputed). So far, the biogeographical distribution of the peri-Mediterranean underground fauna has predominantly been explained through a passive isolation process of former marine species, driven by regressing coastlines. Stemming from a freshwater lake environment, the 125 million-year-old fossils from Lebanon provide an unconventional perspective on the evolutionary origin of extant cave- and groundwater-dwelling cymothoidans.

With approximately 1,500 extant species, freshwater crabs (Decapoda: Brachyura) are among the most diverse decapod crustaceans. Nevertheless, their fossil record is extremely limited: only Potamidae, Potamonautidae and Trichodactylidae are reported up to the Eocene of the Neotropics so far. This work documents unusually large decapod claws from the Upper Cretaceous (Campanian) continental deposits of Velaux and vicinity (southern France), in close association with large vertebrate remains. In addition to (1) the systematic assignment of these claws, the study addresses (2) the salinity trends in the deposit environment from its faunal assemblage and the elementary chemical patterns of fossils, and (3) the likely scenario for their auto/allochthony in the Velaux fluvial system. These claws belong to a new taxon, Dinocarcinus velauciensis n. gen. n. sp., referred to as Portunoidea sensu lato, a group of “true” crabs nowadays linked to marine systems. However, the faunal assemblage, the claw taphonomy and the carbonates Y/Ho signatures support their ancient freshwater/terrestrial ecology, making them the oldest reported continental brachyurans and extending the presence of crabs in freshwater environments by 40 Ma. Either as primary or as secondary freshwater crabs, the occurrence of these portunoids in Velaux is an evidence for the independent colonizations of continental environments by multiple brachyuran clades over time, as early as the Campanian.

Background Among minute-sized and wingless arthropods, astigmatid mites stand out for their diverse range of symbiotic associations (parasitic, neutral and mutualistic), with both invertebrate and vertebrate hosts. When inhabiting discontinuous and ephemeral environments, astigmatid mites adapt their life cycle to produce a phoretic heteromorphic nymph. When feeding resources are depleted, phoretic nymphs disperse to new habitats through phoresy, attaching to a larger animal which transports them to new locations. This dispersal strategy is crucial for accessing patchy resources, otherwise beyond the reach of these minute arthropods. In Astigmata, the phoretic nymph is highly specialized for dispersal, equipped with an attachment organ and lacking a mouth and pharynx. Despite the common occurrence of phoretic associations in modern mites, their evolutionary origins remain poorly understood. Among Astigmata, the family Schizoglyphidae represents an early derivative lineage with phoretic tritonymphs; however, our knowledge of this family is limited to a single observation. Results Here, we report the oldest biotic association of arthropods fossilised in amber (~ 130 Ma, Lebanon): an alate termite with 16 phoretic tritonymphs of Schizoglyphidae ( Plesioglyphus lebanotermi gen. et sp. n.). The mites are primarily attached to the membranes of the host’s hindwings, using their attachment organs, pretarsal claws and tarsal setae. Additionally, we report new modern phoretic tritonymphs of this same family, on one of the earliest lineages of termites. These data collectively indicate that schizoglyphid-termite associations represent the oldest continuous mite-host associations. Notably, phoretic schizoglyphids retain a distinct mouth and pharynx, whereas these structures are absent in the modern phoretic stages of non-schizoglyphid Astigmata. Conclusion The discovery of Schizoglyphidae mites in Lebanese amber represents the oldest known continuous association between acariform mites and their hosts. This finding demonstrates the long-term evolutionary significance of phoresy in Astigmata, evidencing a relationship sustained for over 130 Ma. It indicates that these early mites lived inside termite nests as inquilines and used alate termites for dispersal. This ancient association offers key insights into the coevolution of both mites and termites, highlighting a potential for the future discoveries of similar mites. This fossil —a stem-group Astigmata— is important for the accurate calibration of acariform mite phylogenies, advancing our understanding of these mites evolutionary history.

The earliest known crabs are of Early and Middle Jurassic age; in general, they are rare. Here we describe a new species of homolodromioid from the late Bathonian of Sarthe (France), based on a single dorsal carapace, Tanidromites raboeufi n. sp. This specimen has mostly well-preserved cuticle, and shows two episkeletozoans (bryozoan colonies) on its dorsal side, which is unique amongst early brachyurans (Early and Middle Jurassic). These bryozoans are identified as juvenile colonies of the morphogenus Berenicea (sheet-like tubuluporines), typical encrusters of hard substrates. Their presence directly on the carapace crab shows that they were closely cohabitating with this early brachyuran in its palaeoenvironment. One of these colonies appears to have attached to the internal carapace surface, indicative of post-mortem settlement and growth. Colony sizes suggest a growth phase of at least several months on the sea floor, implying a certain resistance to decomposition of the crab carapace. This is of note in the light of extant homolodromioids which are known to have extremely fragile carapaces. This association demonstrates that post-mortem associations may yield palaeoecological insights into both substrate and colonising organisms.

In the 19th century, natural mummies of amphibians were discovered in the Quercy Phosphorites. The specific collection site was never formally reported, which hampers precise dating of these specimens. Still, the name Bufo servatus was erected based on the external morphology of one of the mummified specimens. A tomography of a similarly preserved specimen revealed a preserved skeleton, soft tissues and gut contents. We analyze here the holotype of Bufo servatus using CT-scanning in order to investigate its potentially preserved internal features. Like the previous specimen, a subcomplete articulated skeleton was identified in the B. servatus holotype. Surprisingly, this skeleton is almost identical to that of Thaumastosaurus gezei, an Eocene anuran from Western Europe to which other specimens from this mummy series were previously assigned. The few differences between the specimen skeletons highlight ontogenetic and intraspecific variations, making T. gezei a junior synonym of B. servatus and creating the new combination Thaumastosaurus servatus. Given its association with previously described Quercy specimens, this redescribed anuran is probably from the same time interval as T. gezei. Previous phylogenetic analyses assigned T. servatus to Ranoides, with natatanuran affinities. Using data from this newly described specimen, we tested here further its taxonomic affinities. Our analyses confirm this position, and formally identify T. servatus as a Natatanuran member of Pyxicephalidae (currently endemic of equatorial Africa) and more precisely, a stem-Pyxicephalinae. This result confirms the origin of Thaumastosaurus, a member of the African herpetofauna occupying Western Europe before the Grande Coupure at the Eocene/Oligocene transition.

Tardigrades are a diverse group of charismatic microscopic invertebrates that are best known for their ability to survive extreme conditions. Despite their long evolutionary history and global distribution in both aquatic and terrestrial environments, the tardigrade fossil record is exceedingly sparse. Molecular clocks estimate that tardigrades diverged from other panarthropod lineages before the Cambrian, but only two definitive crown-group representatives have been described to date, both from Cretaceous fossil deposits in North America. Here, we report a third fossil tardigrade from Miocene age Dominican amber. Paradoryphoribius chronocaribbeus gen. et sp. nov. is the first unambiguous fossil representative of the diverse superfamily Isohypsibioidea, as well as the first tardigrade fossil described from the Cenozoic. We propose that the patchy tardigrade fossil record can be explained by the preferential preservation of these microinvertebrates as amber inclusions, coupled with the scarcity of fossiliferous amber deposits before the Cretaceous.

Here, we report fossil isopods preserved in laminated oilshale mudstone (dysodile) from the Lower Cretaceous of Lebanon (Lower Barremian, 125 Ma, Gres du Liban Alloformation, Jezzine District). Based on a variety of proxies, their palaeoenvironments are determined to have been a shallow freshwater lake. The fossil isopods were studied using modern imaging techniques, such as multispectral imaging and photometric stereo, allowing for a detailed comparison of these specimens with comparable extant and fossil taxa. The conspecific fossils are herein recognized as remains of a new species - +Dysopodus gezei gen. et sp. nov.-of uncertain affinity within Cymothoida and bearing a strong resemblance to its non-parasitic lineages (Cirolanidae). A conspicuous pleotelson and uropod morphology set it apart from most species, with the notable exception of +Pseudoplakolana chiapaneca gen. nov. et comb. nov. from the Cretaceous of Mexico, originally attributed to an Australasian lineage (herein disputed). So far, the biogeographical distribution of the peri-Mediterranean underground fauna has predominantly been explained through a passive isolation process of former marine species, driven by regressing coastlines. Stemming from a freshwater lake environment, the 125 million-year-old fossils from Lebanon provide an unconventional perspective on the evolutionary origin of extant cave- and groundwater-dwelling cymothoidans.

Epibionts are uniquely valuable in their ability to constrain paleoecological hypotheses about their own as well as their host's behavior and environment. Rarely preserved epizoic bryozoans are here reported on fossil crabs from the Miocene Mishan Formation in the Zagros Basin of southwestern Iran. One-hundred-thirty-eight decapod crustaceans were recovered from the upper marly member of the Mishan Formation. Of those, seven decapods (5%) were fouled by bryozoans. Of these seven decapods, five had bryozoans growing attached to the cuticle of the hard outer surface of the exoskeleton of the host crabs and thus fouled while the host crab was potentially alive. Forty percent of the bryozoan colonies occurred on the host crab's dorsal carapace, and 60% were found on chelipeds. On average, 30% of the surface area of the host crabs' fouled skeletal components were covered by bryozoan colonies. The brachyurans were mostly leucosiids, including Leucosia persica and Myra sp. The bryozoans were all cheilostomes and included Acanthodesia sp., Thalamoporella sp., and an indeterminate ascophoran. These bryozoans are all the first reported occurrences in the Mishan Formation. The low incidence of crabs fouled by bryozoans is attributed to preservational bias. The results from this study are compared to those of fossil and extant host crabs reported in the literature. The bryozoan-crab relationship documented here is best described as commensalism.