skip to main content

Title: Morphology-based Phylogenetic Analysis of Membracoidea (Hemiptera: Cicadomorpha) With Placement of Fossil Taxa and Description of a New Subfamily

Recently discovered amber-preserved fossil Cicadellidae exhibit combinations of morphological traits not observed in the modern fauna and have the potential to shed new light on the evolution of this highly diverse family. To place the fossils explicitly within a phylogenetic context, representatives of five extinct genera from Cretaceous Myanmar amber, and one from Eocene Baltic amber were incorporated into a matrix comprising 229 discrete morphological characters and representatives of all modern subfamilies. Phylogenetic analyses yielded well resolved and largely congruent estimates that support the monophyly of most previously recognized cicadellid subfamilies and indicate that the treehoppers are derived from a lineage of Cicadellidae. Instability in the morphology-based phylogenies is mainly confined to deep internal splits that received low branch support in one or more analyses and also were not consistently resolved by recent phylogenomic analyses. Placement of fossil taxa is mostly stable across analyses. Three new Cretaceous leafhopper genera, Burmotettix gen. nov., Kachinella gen nov., and Viraktamathus gen. nov., consistently form a monophyletic group distinct from extant leafhopper subfamilies and are placed in Burmotettiginae subfam. nov. Extinct Cretaceous fossils previously placed in Ledrinae and Signoretiinae are recovered as sister to modern representatives of these groups. Eomegophthalmus Dietrich and Gonçalves from Baltic amber consistently groups with a lineage comprising treehoppers, Megophthalminae, Ulopinae, and Eurymelinae but its position is unstable. Overall, the morphology-based phylogenetic estimates agree with recent phylogenies based on molecular data alone suggesting that morphological traits recently used to diagnose subfamilies are generally informative of phylogenetic relationships within this group.

more » « less
Award ID(s):
Author(s) / Creator(s):
; ; ; ; ; ; ;
Publisher / Repository:
Oxford University Press
Date Published:
Journal Name:
Insect Systematics and Diversity
Medium: X
Sponsoring Org:
National Science Foundation
More Like this
  1. null (Ed.)
    Xiphosurans are aquatic chelicerates with a fossil record extending into the Early Ordovician and known from a total of 88 described species, four of which are extant. Known for their apparent morphological conservatism, for which they have gained notoriety as supposed ‘living fossils’, recent analyses have demonstrated xiphosurans to have an ecologically diverse evolutionary history, with several groups moving into non-marine environments and developing morphologies markedly different from those of the modern species. The combination of their long evolutionary and complex ecological history along with their paradoxical patterns of morphological stasis in some clades and experimentation among others has resulted in Xiphosura being of particular interest for macroevolutionary study. Phylogenetic analyses have shown the current taxonomic framework for Xiphosura—set out in the Treatise of Invertebrate Paleontology in 1955—to be outdated and in need of revision, with several common genera such as Paleolimulus Dunbar, 1923 and Limulitella Størmer, 1952 acting as wastebasket taxa. Here, an expanded xiphosuran phylogeny is presented, comprising 58 xiphosuran species as part of a 158 taxon chelicerate matrix coded for 259 characters. Analysing the matrix under both Bayesian inference and parsimony optimisation criteria retrieves a concordant tree topology that forms the basis of a genus-level systematic revision of xiphosuran taxonomy. The genera Euproops Meek, 1867, Belinurus König, 1820, Paleolimulus , Limulitella , and Limulus are demonstrated to be non-monophyletic and the previously synonymized genera Koenigiella Raymond, 1944 and Prestwichianella Cockerell, 1905 are shown to be valid. In addition, nine new genera ( Andersoniella gen. nov. , Macrobelinurus gen. nov. , and Parabelinurus gen. nov. in Belinurina; Norilimulus gen. nov. in Paleolimulidae; Batracholimulus gen. nov. and Boeotiaspis gen. nov. in Austrolimulidae; and Allolimulus gen. nov., Keuperlimulus gen. nov., and Volanalimulus gen. nov. in Limulidae) are erected to accommodate xiphosuran species not encompassed by existing genera. One new species, Volanalimulus madagascarensis gen. et sp. nov., is also described. Three putative xiphosuran genera— Elleria Raymond, 1944, Archeolimulus Chlupáč, 1963, and Drabovaspis Chlupáč, 1963—are determined to be non-xiphosuran arthropods and as such are removed from Xiphosura. The priority of Belinurus König, 1820 over Bellinurus Pictet, 1846 is also confirmed. This work is critical for facilitating the study of the xiphosuran fossil record and is the first step in resolving longstanding questions regarding the geographic distribution of the modern horseshoe crab species and whether they truly represent ‘living fossils’. Understanding the long evolutionary history of Xiphosura is vital for interpreting how the modern species may respond to environmental change and in guiding conservation efforts. 
    more » « less
  2. Hyaenodonta is a diverse, extinct group of carnivorous mammals that included weasel- to rhinoceros-sized species. The oldest-known hyaenodont fossils are from the middle Paleocene of North Africa and the antiquity of the group in Afro-Arabia led to the hypothesis that it originated there and dispersed to Asia, Europe, and North America. Here we describe two new hyaenodont species based on the oldest hyaenodont cranial specimens known from Afro-Arabia. The material was collected from the latest Eocene Locality 41 (L-41, ∼34 Ma) in the Fayum Depression, Egypt.Akhnatenavus nefertiticyonsp. nov. has specialized, hypercarnivorous molars and an elongate cranial vault. InA. nefertiticyonthe tallest, piercing cusp on M1–M2is the paracone.Brychotherium ephalmosgen. et sp. nov. has more generalized molars that retain the metacone and complex talonids. InB. ephalmosthe tallest, piercing cusp on M1–M2is the metacone. We incorporate this new material into a series of phylogenetic analyses using a character-taxon matrix that includes novel dental, cranial, and postcranial characters, and samples extensively from the global record of the group. The phylogenetic analysis includes the first application of Bayesian methods to hyaenodont relationships.B. ephalmosis consistently placed within Teratodontinae, an Afro-Arabian clade with several generalist and hypercarnivorous forms, andAkhnatenavusis consistently recovered in Hyainailourinae as part of an Afro-Arabian radiation. The phylogenetic results suggest that hypercarnivory evolved independently three times within Hyaenodonta: in Teratodontinae, in Hyainailourinae, and in Hyaenodontinae. Teratodontines are consistently placed in a close relationship with Hyainailouridae (Hyainailourinae + Apterodontinae) to the exclusion of “proviverrines,” hyaenodontines, and several North American clades, and we propose that the superfamily Hyainailouroidea be used to describe this relationship. Using the topologies recovered from each phylogenetic method, we reconstructed the biogeographic history of Hyaenodonta using parsimony optimization (PO), likelihood optimization (LO), and Bayesian Binary Markov chain Monte Carlo (MCMC) to examine support for the Afro-Arabian origin of Hyaenodonta. Across all analyses, we found that Hyaenodonta most likely originated in Europe, rather than Afro-Arabia. The clade is estimated by tip-dating analysis to have undergone a rapid radiation in the Late Cretaceous and Paleocene; a radiation currently not documented by fossil evidence. During the Paleocene, lineages are reconstructed as dispersing to Asia, Afro-Arabia, and North America. The place of origin of Hyainailouroidea is likely Afro-Arabia according to the Bayesian topologies but it is ambiguous using parsimony. All topologies support the constituent clades–Hyainailourinae, Apterodontinae, and Teratodontinae–as Afro-Arabian and tip-dating estimates that each clade is established in Afro-Arabia by the middle Eocene.

    more » « less
  3. Portunoidea is a diverse lineage of ecologically and economically important marine crabs comprising 8 families and 14 subfamilies. Closely related portunid subfamilies Caphyrinae and Thalamitinae constitute some of this group’s greatest morphological and taxonomic diversity, and are the only known lineages to include symbiotic taxa. Emergence of symbiosis in decapods remains poorly studied and portunoid crabs provide an interesting, but often overlooked example. Yet the paucity of molecular phylogenetic data available for Portunoidea makes it challenging to investigate the evolution and systematics of the group. Phylogenetic analyses, though limited, suggest that many putative portunoid taxa are para- or polyphyletic. Here I augment existing molecular data—significantly increasing taxon sampling of Caphyrinae, Thalamitinae, and several disparate portunoid lineages—to investigate the phylogenetic origin of symbiosis within Portunoidea and reevaluate higher- and lower-level portunoid classifications. Phylogenetic analyses were carried out on sequences of H3, 28S rRNA, 16S rRNA, and CO1 for up to 168 portunoid taxa; this included, for the first time, molecular data from the genera Atoportunus , Brusinia , Caphyra , Coelocarcinus , Gonioinfradens , Raymanninus , and Thalamonyx . Results support the placement of all symbiotic taxa ( Caphyra , Lissocarcinus , and two Thalamita ) in a single clade derived within the thalamitine genus Thalamita . Caphyrina Paulson, 1875, nom. trans. is recognized here as a subtribe within the subfamily Thalamitinae. Results also support the following taxonomic actions: Cronius is reclassified as a thalamitine genus; Thalamonyx is reestablished as a valid genus; Goniosupradens is raised to the generic rank; and three new genera ( Zygita gen. nov., Thranita gen. nov., and Trierarchus gen. nov.) are described to accommodate some Thalamita s.l. taxa rendered paraphyletic by Caphyrina. A new diagnosis of Thalamitinae is provided. Results also support a more conservative classification of Portunoidea comprising three instead of eight extant families: Geryonidae (Geryonidae + Ovalipidae; new diagnosis provided), Carcinidae (Carcinidae + Pirimelidae + Polybiidae + Thiidae + Coelocarcinus ; new diagnosis provided) and Portunidae. Finally, 16s rRNA data suggests family Brusiniidae might not be a portunoid lineage. 
    more » « less
  4. Abstract

    Hatching is a pivotal moment in the life of most animals. Diverse chemical, behavioural and mechanical methods have evolved in metazoans to break the egg membranes. Among them, many arthropod and vertebrate embryos hatch using ephemeral, frequently convergent structures known as egg bursters. However, the evolutionary processes by which hatching mechanisms and related embryonic structures became established in deep time are poorly understood due to a nearly complete absence from the fossil record. Herein we describe an exceptionalc. 130‐million‐year‐old association in Lebanese amber composed of multiple neonate green lacewing larvae,Tragichrysa ovoruptoragen. et sp. nov. (Neuroptera, Chrysopoidea), and conspecific egg remains. Egg bursters with a serrated blade bearing a short process are attached to three longitudinally split egg shells. Embryos of extant green lacewing relatives (Chrysopidae) utilize this egg burster morphotype to open a vertical slit on the egg, after which the burster is moulted and left joined to the empty egg shell. Additionally, the new larval species has extremely elongate dorsal tubercles, an adaptation to carry exogenous debris for protection and camouflage also known from other Cretaceous chrysopoids but absent in modern relatives. The present discovery demonstrates that the hatching mechanism of modern green lacewings was established in the chrysopoid lineage by the Early Cretaceous and proves through direct fossil evidence how some morphological traits related to hatching and linked behaviours, at least in insect embryos, have been subject to a high degree of evolutionary conservatism.

    more » « less
  5. INTRODUCTION Resolving the role that different environmental forces may have played in the apparent explosive diversification of modern placental mammals is crucial to understanding the evolutionary context of their living and extinct morphological and genomic diversity. RATIONALE Limited access to whole-genome sequence alignments that sample living mammalian biodiversity has hampered phylogenomic inference, which until now has been limited to relatively small, highly constrained sequence matrices often representing <2% of a typical mammalian genome. To eliminate this sampling bias, we used an alignment of 241 whole genomes to comprehensively identify and rigorously analyze noncoding, neutrally evolving sequence variation in coalescent and concatenation-based phylogenetic frameworks. These analyses were followed by validation with multiple classes of phylogenetically informative structural variation. This approach enabled the generation of a robust time tree for placental mammals that evaluated age variation across hundreds of genomic loci that are not restricted by protein coding annotations. RESULTS Coalescent and concatenation phylogenies inferred from multiple treatments of the data were highly congruent, including support for higher-level taxonomic groupings that unite primates+colugos with treeshrews (Euarchonta), bats+cetartiodactyls+perissodactyls+carnivorans+pangolins (Scrotifera), all scrotiferans excluding bats (Fereuungulata), and carnivorans+pangolins with perissodactyls (Zooamata). However, because these approaches infer a single best tree, they mask signatures of phylogenetic conflict that result from incomplete lineage sorting and historical hybridization. Accordingly, we also inferred phylogenies from thousands of noncoding loci distributed across chromosomes with historically contrasting recombination rates. Throughout the radiation of modern orders (such as rodents, primates, bats, and carnivores), we observed notable differences between locus trees inferred from the autosomes and the X chromosome, a pattern typical of speciation with gene flow. We show that in many cases, previously controversial phylogenetic relationships can be reconciled by examining the distribution of conflicting phylogenetic signals along chromosomes with variable historical recombination rates. Lineage divergence time estimates were notably uniform across genomic loci and robust to extensive sensitivity analyses in which the underlying data, fossil constraints, and clock models were varied. The earliest branching events in the placental phylogeny coincide with the breakup of continental landmasses and rising sea levels in the Late Cretaceous. This signature of allopatric speciation is congruent with the low genomic conflict inferred for most superordinal relationships. By contrast, we observed a second pulse of diversification immediately after the Cretaceous-Paleogene (K-Pg) extinction event superimposed on an episode of rapid land emergence. Greater geographic continuity coupled with tumultuous climatic changes and increased ecological landscape at this time provided enhanced opportunities for mammalian diversification, as depicted in the fossil record. These observations dovetail with increased phylogenetic conflict observed within clades that diversified in the Cenozoic. CONCLUSION Our genome-wide analysis of multiple classes of sequence variation provides the most comprehensive assessment of placental mammal phylogeny, resolves controversial relationships, and clarifies the timing of mammalian diversification. We propose that the combination of Cretaceous continental fragmentation and lineage isolation, followed by the direct and indirect effects of the K-Pg extinction at a time of rapid land emergence, synergistically contributed to the accelerated diversification rate of placental mammals during the early Cenozoic. The timing of placental mammal evolution. Superordinal mammalian diversification took place in the Cretaceous during periods of continental fragmentation and sea level rise with little phylogenomic discordance (pie charts: left, autosomes; right, X chromosome), which is consistent with allopatric speciation. By contrast, the Paleogene hosted intraordinal diversification in the aftermath of the K-Pg mass extinction event, when clades exhibited higher phylogenomic discordance consistent with speciation with gene flow and incomplete lineage sorting. 
    more » « less