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Concept: Rove beetle


Agaricomycetes, or mushrooms, are familiar, conspicuous and morphologically diverse Fungi. Most Agaricomycete fruiting bodies are ephemeral, and their fossil record is limited. Here we report diverse gilled mushrooms (Agaricales) and mycophagous rove beetles (Staphylinidae) from mid-Cretaceous Burmese amber, the latter belonging to Oxyporinae, modern members of which exhibit an obligate association with soft-textured mushrooms. The discovery of four mushroom forms, most with a complete intact cap containing distinct gills and a stalk, suggests evolutionary stasis of body form for ∼99 Myr and highlights the palaeodiversity of Agaricomycetes. The mouthparts of early oxyporines, including enlarged mandibles and greatly enlarged apical labial palpomeres with dense specialized sensory organs, match those of modern taxa and suggest that they had a mushroom feeding biology. Diverse and morphologically specialized oxyporines from the Early Cretaceous suggests the existence of diverse Agaricomycetes and a specialized trophic interaction and ecological community structure by this early date.

Concepts: Evolution, Insect, Fungus, Beetle, Mushroom, Agaricales, Agaricomycetes, Rove beetle


Some flying beetles have peculiar functional properties of their elytra, if compared with the vast majority of beetles. A “typical” beetle covers its pterothorax and the abdomen from above with closed elytra and links closed elytra together along the sutural edges. In the open state during flight, the sutural edges diverge much more than by 90°. Several beetles of unrelated taxa spread wings through lateral incisions on the elytra and turn the elytron during opening about 10-12° (Cetoniini, Scarabaeus, Gymnopleurus) or elevate their elytra without partition (Sisyphus, Tragocerus). The number of campaniform sensilla in their elytral sensory field is diminished in comparison with beetles of closely related taxa lacking that incision. Elytra are very short in rove beetles and in long-horn beetles Necydalini. The abundance of sensilla in brachyelytrous long-horn beetles Necydalini does not decrease in comparison with macroelytrous Cerambycinae. The strong reduction of the sensory field was found in brachyelytrous Staphylinidae. Lastly, there are beetles lacking the linkage of the elytra down the sutural edge (stenoelytry). Effects of stenoelytry were also not uniform: Oedemera and flying Meloidae have the normal amount of sensilla with respect to their body size, whereas the sensory field in the stenoelytrous Eulosia bombyliformis is 5-6 times less than in chafers of the same size but with normally linking broad elytra.

Concepts: Normal distribution, Beetle, Elytron, Polyphaga, Aleochara, Rove beetle, Blister beetle, Staphylinidae


Recent adaptive radiations provide striking examples of convergence [1-4], but the predictability of evolution over much deeper timescales is controversial, with a scarcity of ancient clades exhibiting repetitive patterns of phenotypic evolution [5, 6]. Army ants are ecologically dominant arthropod predators of the world’s tropics, with large nomadic colonies housing diverse communities of socially parasitic myrmecophiles [7]. Remarkable among these are many species of rove beetle (Staphylinidae) that exhibit ant-mimicking “myrmecoid” body forms and are behaviorally accepted into their aggressive hosts' societies: emigrating with colonies and inhabiting temporary nest bivouacs, grooming and feeding with workers, but also consuming the brood [8-11]. Here, we demonstrate that myrmecoid rove beetles are strongly polyphyletic, with this adaptive morphological and behavioral syndrome having evolved at least 12 times during the evolution of a single staphylinid subfamily, Aleocharinae. Each independent myrmecoid clade is restricted to one zoogeographic region and highly host specific on a single army ant genus. Dating estimates reveal that myrmecoid clades are separated by substantial phylogenetic distances-as much as 105 million years. All such groups arose in parallel during the Cenozoic, when army ants diversified into modern genera [12] and rose to ecological dominance [13, 14]. This work uncovers a rare example of an ancient system of complex morphological and behavioral convergence, with replicate beetle lineages following a predictable phenotypic trajectory during their parasitic adaptation to host colonies.

Concepts: Evolution, Insect, Beetle, Ant, Polyphaga, Aleochara, Rove beetle, Staphylinidae


Foldable wings of insects are the ultimate deployable structures and have attracted the interest of aerospace engineering scientists as well as entomologists. Rove beetles are known to fold their wings in the most sophisticated ways that have right-left asymmetric patterns. However, the specific folding process and the reason for this asymmetry remain unclear. This study reveals how these asymmetric patterns emerge as a result of the folding process of rove beetles. A high-speed camera was used to reveal the details of the wing-folding movement. The results show that these characteristic asymmetrical patterns emerge as a result of simultaneous folding of overlapped wings. The revealed folding mechanisms can achieve not only highly compact wing storage but also immediate deployment. In addition, the right and left crease patterns are interchangeable, and thus each wing internalizes two crease patterns and can be folded in two different ways. This two-way folding gives freedom of choice for the folding direction to a rove beetle. The use of asymmetric patterns and the capability of two-way folding are unique features not found in artificial structures. These features have great potential to extend the design possibilities for all deployable structures, from space structures to articles of daily use.

Concepts: Insect, Beetle, Wing, Insect wing, Polyphaga, Aleochara, Rove beetle, Aerospace engineering


Myrmecophiles-species that depend on ant societies-include some of the most morphologically and behaviorally specialized animals known [1]. Remarkable adaptive characters enable these creatures to bypass fortress-like security, integrate into colony life, and exploit abundant resources and protection inside ant nests [2, 3]. Such innovations must result from intimate coevolution with hosts, but a scarcity of definitive fossil myrmecophiles obscures when and how this lifestyle arose. Here, we report the earliest known morphologically specialized and apparently obligate myrmecophile, in Early Eocene (∼52 million years old) Cambay amber from India. Protoclaviger trichodens gen. et sp. nov. is a stem-group member of Clavigeritae, a speciose supertribe of pselaphine rove beetles (Coleoptera: Staphylinidae) heavily modified for myrmecophily via reduced mouthparts for trophallaxis with worker ants, brush-like trichomes that exude appeasement compounds, and fusions of many body and antennal segments [4, 5]. Protoclaviger captures a transitional stage in the evolutionary development of this novel body plan, most evident in its still-distinct abdominal tergites. The Cambay paleobiota marks one of the first occurrences in the fossil record of a significant presence of modern ants [6]. Protoclaviger reveals that sophisticated social parasites were nest intruders throughout, and probably before, the ascent of ants to ecological dominance, with ancient groups such as Clavigeritae primed to radiate as their hosts became increasingly ubiquitous.

Concepts: Evolution, Ecology, Beetle, Ant, Ant colony, Aleochara, Rove beetle, Myrmecophile


Joe Parker introduces the staphylinids or ‘rove beetles’, the most species-rich groups of insect on Earth.

Concepts: Beetle, Polyphaga, Aleochara, Rove beetle


Army ant colonies host a diverse community of arthropod symbionts. Among the best-studied symbiont communities are those of Neotropical army ants of the genus Eciton. It is clear, however, that even in these comparatively well studied systems, a large proportion of symbiont biodiversity remains unknown. Even more striking is our lack of knowledge regarding the nature and specificity of these host-symbiont interactions. Here we surveyed the diversity and host specificity of rove beetles of the genus Tetradonia Wasmann, 1894 (Staphylinidae: Aleocharinae). Systematic community sampling of 58 colonies of the six local Eciton species at La Selva Biological Station, Costa Rica, combined with an integrative taxonomic approach, allowed us to uncover species diversity, host specificity, and co-occurrence patterns of symbionts in unprecedented detail. We used an integrative taxonomic approach combining morphological and genetic analyses, to delineate species boundaries. Mitochondrial DNA barcodes were analyzed for 362 Tetradonia specimens, and additional nuclear markers for a subset of 88 specimens. All analyses supported the presence of five Tetradonia species, including two species new to science. Host specificity is highly variable across species, ranging from generalists such as T. laticeps, which parasitizes all six local Eciton species, to specialists such as T. lizonae, which primarily parasitizes a single species, E. hamatum. Here we provide a dichotomous key along with diagnostic molecular characters for identification of Tetradonia species at La Selva Biological Station. By reliably assessing biodiversity and providing tools for species identification, we hope to set the baseline for future studies of the ecological and evolutionary dynamics in these species-rich host-symbiont networks.

Concepts: Evolution, Biology, Life, Beetle, Ant, Rove beetle, Eciton, Eciton burchellii


Termitophiles, symbionts that live in termite nests, include a wide range of morphologically and behaviorally specialized organisms. Complex adaptive mechanisms permit these animals to integrate into societies and to exploit their controlled physical conditions and plentiful resources, as well as to garner protection inside termite nests. An understanding of the early evolution of termitophily is challenging owing to a scarcity of fossil termitophiles, with all known reliable records occurring from the Miocene (approximately 19 million years ago [mya]) [1-6], and an equivocal termitophile belonging to the largely free-living Mesoporini from the mid-Cretaceous [7]. Here we report the oldest, morphologically specialized, and obligate termitophiles from mid-Cretaceous Burmese amber (99 mya). Cretotrichopsenius burmiticus gen. et sp. nov. belongs to Trichopseniini, a group of distinctive termitophilous aleocharine rove beetles, all of which possess specialized swollen or horseshoe-crab-shaped body plans. Cretotrichopsenius display the protective horseshoe-crab-shaped body form typical of many modern termitophiles, with concealed head and antennae and strong posteriorly directed abdominal setae. Cretotrichopsenius represent the earliest definitive termitophiles, shedding light on host associations in the early evolution of termite societies. The fossil reveals that ancient termite societies were quickly invaded by beetles and by multiple independent lineages of social parasites over the subsequent eons.

Concepts: Evolution, Insect, Symbiosis, Beetle, Ant, Termite, Polyphaga, Rove beetle


Host-symbiont interactions are embedded in ecological communities and range from unspecific to highly specific relationships. Army ants and their arthropod guests represent a fascinating example of species-rich host-symbiont associations where host specificity ranges across the entire generalist - specialist continuum. In the present study, we compared the behavioral and chemical integration mechanisms of two extremes of the generalist - specialist continuum: generalist ant-predators in the genusTetradonia(Staphylinidae: Aleocharinae: Athetini), and specialist ant-mimics in the generaEcitomorphaandEcitophya(Staphylinidae: Aleocharinae: Ecitocharini). Similar to a previous study ofTetradoniabeetles, we combined DNA barcoding with morphological studies to define species boundaries in ant-mimicking beetles. This approach found four ant-mimicking species at our study site at La Selva Biological Station in Costa Rica. Community sampling ofEcitonarmy ant parasites revealed that ant-mimicking beetles were perfect host specialists, each beetle species being associated with a singleEcitonspecies. These specialists were seamlessly integrated into the host colony, while generalists avoided physical contact to host ants in behavioral assays. Analysis of the ants' nestmate recognition cues, i.e. cuticular hydrocarbons (CHCs), showed close similarity in CHC composition and CHC concentration between specialists andEciton burchellii forelihost ants. On the contrary, the chemical profiles of generalists matched host profiles less well, indicating that high accuracy in chemical host resemblance is only accomplished by socially integrated species. Considering the interplay between behavior, morphology, and cuticular chemistry, specialists but not generalists have cracked the ants' social code with respect to various sensory modalities. Our results support the long-standing idea that the evolution of host-specialization in parasites is a trade-off between the range of potential host species and the level of specialization on any particular host.

Concepts: Evolution, Species, Sensory system, Beetle, DNA barcoding, Ant, Costa Rica, Rove beetle


Myrmecophiles (i.e. organisms that associate with ants) use a variety of ecological niches and employ different strategies to survive encounters with ants. Because ants are typically excellent defenders, myrmecophiles may choose moments of weakness to take advantage of their ant associates. This hypothesis was studied in the rove beetle, Myrmedonota xipe, which associates with Azteca sericeasur ants in the presence of parasitoid flies. A combination of laboratory and field experiments show that M. xipe beetles selectively locate and prey upon parasitized ants. These parasitized ants are less aggressive towards beetles than healthy ants, allowing beetles to eat the parasitized ants alive without interruption. Moreover, behavioural assays and chemical analysis reveal that M. xipe are attracted to the ant’s alarm pheromone, the same secretion used by the phorid fly parasitoids in host location. This strategy allows beetles access to an abundant but otherwise inaccessible resource, as A. sericeasur ants are typically highly aggressive. These results are the first, to our knowledge, to demonstrate a predator sharing cues with a parasitoid to gain access to an otherwise unavailable prey item. Furthermore, this work highlights the importance of studying ant-myrmecophile interactions beyond just their pairwise context.

Concepts: Insect, Ecology, Beetle, Ant, Biological pest control, Hymenoptera, Aleochara, Rove beetle