Fleas are one of the major lineages of ectoparasitic insects and are now highly specialized for feeding on the blood of birds or mammals. This has isolated them among holometabolan insect orders, although they derive from the Antliophora (scorpionflies and true flies). Like most ectoparasitic lineages, their fossil record is meagre and confined to Cenozoic-era representatives of modern families, so that we lack evidence of the origins of fleas in the Mesozoic era. The origins of the first recognized Cretaceous stem-group flea, Tarwinia, remains highly controversial. Here we report fossils of the oldest definitive fleas--giant forms from the Middle Jurassic and Early Cretaceous periods of China. They exhibit many defining features of fleas but retain primitive traits such as non-jumping hindlegs. More importantly, all have stout and elongate sucking siphons for piercing the hides of their hosts, implying that these fleas may be rooted among the pollinating 'long siphonate' scorpionflies of the Mesozoic. Their special morphology suggests that their earliest hosts were hairy or feathered 'reptilians', and that they radiated to mammalian and bird hosts later in the Cenozoic.
Staphylinoidea (Insecta: Coleoptera) is one of the most species-rich groups in animals, but its huge diversity can hardly be explained by the popular hypothesis (co-radiation with angiosperms) that applies to phytophagous beetles. We estimated the evolutionary mode of staphylinoid beetles and investigated the relationship between the evolutionary mode and palaeoclimate change, and thus the factors underlying the current biodiversity pattern of staphylinoid beetles. Our results demonstrate that staphylinoid beetles originated at around the Triassic–Jurassic bound and the current higher level clades underwent rapid evolution (indicated by increased diversification rate and decreased body size disparity) in the Jurassic and in the Cenozoic, both with low-energy climate, and they evolved much slower during the Cretaceous with high-energy climate. Climate factors, especially low O2 and high CO2, promoted the diversification rate and among-clade body size disparification in the Jurassic. In the Cenozoic, however, climate factors had negative associations with diversification rate but little with body size disparification. Our present study does not support the explosion of staphylinoid beetles as a direct outcome of the Cretaceous Terrestrial Revolution (KTR). We suppose that occupying and diversifying in refuge niches associated with litter may elucidate rapid radiations of staphylinoid beetles in low-energy conditions.
Stick and leaf insects (Phasmatodea) are a distinctive insect order whose members are characterized by mimicking various plant tissues such as twigs, foliage and bark. Unfortunately, the phylogenetic relationships among phasmatodean subfamilies and the timescale of their evolution remain uncertain. Recent molecular clock analyses have suggested a Cretaceous–Palaeogene origin of crown Phasmatodea and a subsequent Cenozoic radiation, contrasting with fossil evidence. Here, we analysed transcriptomic data from a broad diversity of phasmatodeans and, combined with the assembly of a new suite of fossil calibrations, we elucidate the evolutionary history of stick and leaf insects. Our results differ from recent studies in the position of the leaf insects (Phylliinae), which are recovered as sister to a clade comprising Clitumninae, Lancerocercata, Lonchodinae, Necrosciinae and Xenophasmina . We recover a Permian to Triassic origin of crown Phasmatodea coinciding with the radiation of early insectivorous parareptiles, amphibians and synapsids. Aschiphasmatinae and Neophasmatodea diverged in the Jurassic–Early Cretaceous. A second spur in origination occurred in the Late Cretaceous, coinciding with the Cretaceous Terrestrial Revolution, and was probably driven by visual predators such as stem birds (Enantiornithes) and the radiation of angiosperms.
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 . 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.
Cycads, unlike modern wind-pollinated conifers and Ginkgo, are unusual in that they are an ancient group of gymnosperms pollinated by insects [1-3]. Although it is well documented that cycads were diverse and abundant during the mid-Mesozoic, little is known about their biogeography and pollination before the rise of angiosperms. Direct fossil evidence illuminating the evolutionary history of cycads is extremely rare [4, 5]. Here we report a specialized beetle-mediated pollination mode from the mid-Cretaceous of Myanmar, wherein a new boganiid beetle, Cretoparacucujus cycadophilus, with specialized pollen-feeding adaptations in its mouthparts and legs, was associated with many pollen grains of Cycadopites. Phylogenetic analyses indicate Cretoparacucujus as a sister group to the extant Australian Paracucujus, which pollinate the cycad Macrozamia riedlei. Our discovery, along with the current disjunct distribution of related beetle-herbivore (tribe Paracucujini) and cycad-host (tribe Encephalarteae) pairs in South Africa and Australia, indicate a probable ancient origin of beetle pollination of cycads at least in the Early Jurassic, long before angiosperm dominance and the radiation of flowering-plant pollinators later in the Cretaceous.
Significance We report on the unique discovery of Jurassic and Cretaceous carrion beetles (Silphidae) from China and Myanmar, early relatives of one of the most protected of beetle species in North America, and which clearly preserve evidence indicative of complex parental care. This finding represents the earliest evidence of parental care, a behavioral repertoire that is the first step in the development of truly social behavior and one that is intensely studied by ecologists, ethologists, and evolutionary biologists alike. Our fossils clearly span the origins of parent–offspring communication and allow us to provide a robust estimate of the time of origin for this complex behavior.
Burmese amber represents the world’s most diverse biota in the Mesozoic. Previous studies have focused on the biodiversity of its inclusions, as well as pholadid borings. Here we report a variety of marine animals symbiotic with or adhere to Burmese amber or the amber deposits, including crinoid columns, corals and oysters. We propose that there is no distinct evidence indicating the secondary transportation of Burmese amber over long distances. The ancient sedimentary environment was likely located in the coastal area. The hardening time of the resin was not long after secretion. The resin has been mixed with fragments of marine organisms in the ancient sediments, and has been deposited for a long time. The zircon age in the sediments surrounding amber approximately represents the age of Burmese amber, but due to limits of the method, the current zircon U-Pb SIMS age may be younger. Therefore, as far as the situation is concerned, the age of Burmese amber may be close to the boundary between the Albian and Cenomanian, or even late Albian. We suggest that it is plausible to generally refer to the age of Burmese amber as mid-Cretaceous, and a precise age requires further biostratigraphic and chronological studies.
Staphylininae is the third largest subfamily of the enormous family Staphylinidae. Monophyly of Staphylininae and its sister relationship to the subfamily Paederinae have been broadly accepted according to both conventional morphology- and molecular-based phylogenies until the last three years. Recent molecular phylogenies rejected monophyly of Staphylininae and regarded Paederinae as a clade within it. This paper re-evaluates the recent molecular work, aiming to clarify the relationship between Staphylininae and Paederinae and resolve intertribal relationships within Staphylininae. Based on a new six-gene data set (5707 bp) for 92 taxa including Oxyporinae (outgroup), representatives of Paederinae, and members of all extant tribes of Staphylininae from published DNA data in GenBank, we generated a well-resolved phylogeny of Staphylininae with all deep nodes (intertribal relationships) strongly supported, and reassert the hypothesis that Staphylininae is monophyletic and indeed the sister group to Paederinae using both Bayesian and maximum likelihood inference. Additionally, our study is a case-study to show that both outgroup selection and completeness of nucleotide data can influence the outcome of a molecular phylogeny. With an increasing number of staphylinid fossils being discovered, the robust phylogeny of Staphylininae inferred by our research will provide a good framework for understanding the early evolution of this group.
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