skip to main content
US FlagAn official website of the United States government
dot gov icon
Official websites use .gov
A .gov website belongs to an official government organization in the United States.
https lock icon
Secure .gov websites use HTTPS
A lock ( lock ) or https:// means you've safely connected to the .gov website. Share sensitive information only on official, secure websites.


Title: A molecular phylogeny of the cicadas (Hemiptera: Cicadidae) with a review of tribe and subfamily classification
A molecular phylogeny and a review of family-group classification are presented for 137 species (ca. 125 genera) of the insect family Cicadidae, the true cicadas, plus two species of hairy cicadas (Tettigarctidae) and two outgroup species from Cercopidae. Five genes, two of them mitochondrial, comprise the 4992 base-pair molecular dataset. Maximum-likelihood and Bayesian phylogenetic results are shown, including analyses to address potential base composition bias. Tettigarcta is confirmed as the sister-clade of the Cicadidae and support is found for three subfamilies identified in an earlier morphological cladistic analysis. A set of paraphyletic deep-level clades formed by African genera are together named as Tettigomyiinae n. stat. Taxonomic reassignments of genera and tribes are made where morphological examination confirms incorrect placements suggested by the molecular tree, and 11 new tribes are defined (Arenopsaltriini n. tribe, Durangonini n. tribe, Katoini n. tribe, Lacetasini n. tribe, Macrotristriini n. tribe, Malagasiini n. tribe, Nelcyndanini n. tribe, Pagiphorini n. tribe, Pictilini n. tribe, Psaltodini n. tribe, and Selymbriini n. tribe). Tribe Tacuini n. syn. is synonymized with Cryptotympanini, and Tryellina n. syn. is synonymized with an expanded Tribe Lamotialnini. Tribe Hyantiini n. syn. is synonymized with Fidicinini. Tribe Sinosenini is transferred to Cicadinae from Cicadettinae, Cicadatrini is moved to Cicadettinae from Cicadinae, and Ydiellini and Tettigomyiini are transferred to Tettigomyiinae n. stat from Cicadettinae. While the subfamily Cicadinae, historically defined by the presence of timbal covers, is weakly supported in the molecular tree, high taxonomic rank is not supported for several earlier clades based on unique morphology associated with sound production.  more » « less
Award ID(s):
1655891
PAR ID:
10063712
Author(s) / Creator(s):
; ; ; ; ; ; ; ; ; ; ; ; ;
Date Published:
Journal Name:
Zootaxa
Volume:
4424
Issue:
1
ISSN:
1175-5326
Page Range / eLocation ID:
1; 64
Format(s):
Medium: X
Sponsoring Org:
National Science Foundation
More Like this
  1. null (Ed.)
    A recent paper by Ruschel & Campos (2019) on “leaf-winged” cicadas proposed a significant reorganization of the cicada tribe Hemidictyini Distant, 1905g, including synonymization of the monogeneric tribe Lacetasini Moulds & Marshall, 2018 following the results of a cladistic parsimony analysis of morphological characters. In this study, we reconsider and revise the morphological analysis of Ruschel & Campos and obtain new genetic data for Hemidictya. We find that their study suffers from a limited taxon sample, inappropriate outgroup selection, and misinterpretation of genitalic characters (uncus vs. claspers). We show that Hemidictyini sensu Ruschel & Campos includes members of multiple tribes and subfamilies, and we conclude that some of the taxonomic transfers by Ruschel & Campos are not supported. The two most similar and leaf-like cicadas, Hemidictya Burmeister, 1835 (South America) and Hovana Distant, 1905g (Madagascar), are probably not closely related but rather an excellent example of convergent evolution. Lacetasini is not a junior synonym of the Hemidictyini but a distinct part of the Tettigomyiinae Distant, 1905g as originally classified. We return or transfer the genera Lacetas Karsch, 1890, Iruana Distant, 1905g, Bafutalna Boulard, 1993, and Murphyalna Boulard, 2012 to the Lacetasini. With the transfer of all genera of Iruanina Boulard, 1993 and Bafutalnina Boulard, 1993 to Lacetasini and with Lacetas transferred to the Iruanina, Lacetasini n. syn. becomes a subjective junior synonym of Iruanini rev. stat. in the Tettigomyiinae. We assign Hovana to Hovanini n. tribe in the Tettigomyiinae and Sapantanga Distant, 1905g to Sapantangini n. tribe in the Tibicininae Distant, 1905b. We propose that Hemidictyini sensu novo contains only the genus Hemidictya and we assign the tribe to Tibicininae with a revised diagnosis. 
    more » « less
  2. The cicadas (Hemiptera: Cicadidae) related to tribe Cicadini exhibit some of the most remarkable phenotypes in the family, with many genera possessing striking colour patterns and unusual morphological features. This largely Asian group of 13 tribes has proven challenging for cicada taxonomists, in part because of likely convergent evolution or losses of these phenotypes. We present the first focused molecular phylogeny of this clade, including ~60 described genera. The genetic dataset contains 839 ingroup-informative sites (out of 2575) from mitochondrial cytochrome c oxidase subunit I, nuclear elongation factor-1 α, and nuclear acetyltransferase. We use Bayesian and maximum likelihood trees to test recent changes in tribe- and subtribe-level classification, and we reconstruct ancestral character states for potentially convergent traits influencing tribe descriptions. We use fossil and molecular clock calibrations to estimate the temporal and geographic context of the radiation. The tribes Gaeanini, Leptopsaltriini, Platypleurini, Psithyristriini, and Tosenini appear polyphyletic and in need of revision, in part because of convergent evolution of opaque wings and multiple convergent gains or losses of abdominal tubercles. Kalabita Moulton, 1923 is transferred from Platypleurini to Leptopsaltriini. Vittagaeana gen. nov. is established for Vittagaeana paviei comb. nov. and Vittagaeana dives comb. nov., formerly in Tosena. Sinosenini syn. nov. is synonymised with Dundubiina. Ayuthiini trib. nov. is established with two new subtribes for Ayuthia Distant, 1919 and Distantalna Boulard, 2009, formerly in Tosenini. For the earliest split in the tree, one common ancestor appears to have been Indian + Asian in geographic distribution and the other Asian. We estimate that the radiation began in the middle Cenozoic Era, possibly as recently as the early Miocene. The recent and steady pattern of diversification suggests that refinement of tribe diagnoses will prove challenging. 
    more » « less
  3. Abstract Contamination of a genetic sample with DNA from one or more nontarget species is a continuing concern of molecular phylogenetic studies, both Sanger sequencing studies and next-generation sequencing studies. We developed an automated pipeline for identifying and excluding likely cross-contaminated loci based on the detection of bimodal distributions of patristic distances across gene trees. When contamination occurs between samples within a data set, a comparison between a contaminated sample and its contaminant taxon will yield bimodal distributions with one peak close to zero patristic distance. This new method does not rely on a priori knowledge of taxon relatedness nor does it determine the causes(s) of the contamination. Exclusion of putatively contaminated loci from a data set generated for the insect family Cicadidae showed that these sequences were affecting some topological patterns and branch supports, although the effects were sometimes subtle, with some contamination-influenced relationships exhibiting strong bootstrap support. Long tip branches and outlier values for one anchored phylogenomic pipeline statistic (AvgNHomologs) were correlated with the presence of contamination. While the anchored hybrid enrichment markers used here, which target hemipteroid taxa, proved effective in resolving deep and shallow level Cicadidae relationships in aggregate, individual markers contained inadequate phylogenetic signal, in part probably due to short length. The cleaned data set, consisting of 429 loci, from 90 genera representing 44 of 56 current Cicadidae tribes, supported three of the four sampled Cicadidae subfamilies in concatenated-matrix maximum likelihood (ML) and multispecies coalescent-based species tree analyses, with the fourth subfamily weakly supported in the ML trees. No well-supported patterns from previous family-level Sanger sequencing studies of Cicadidae phylogeny were contradicted. One taxon (Aragualna plenalinea) did not fall with its current subfamily in the genetic tree, and this genus and its tribe Aragualnini is reclassified to Tibicininae following morphological re-examination. Only subtle differences were observed in trees after the removal of loci for which divergent base frequencies were detected. Greater success may be achieved by increased taxon sampling and developing a probe set targeting a more recent common ancestor and longer loci. Searches for contamination are an essential step in phylogenomic analyses of all kinds and our pipeline is an effective solution. [Auchenorrhyncha; base-composition bias; Cicadidae; Cicadoidea; Hemiptera; phylogenetic conflict.] 
    more » « less
  4. Abstract The Platypleurini is a large group of charismatic cicadas distributed from Cape Agulhas in South Africa, through tropical Africa, Madagascar, India and eastern Asia to Japan, with generic diversity concentrated in equatorial and southern Africa. This distribution suggests the possibility of a Gondwanan origin and dispersal to eastern Asia from Africa or India. We used a four‐gene (three mitochondrial) molecular dataset, fossil calibrations and molecular clock information to explore the phylogenetic relationships of the platypleurine cicadas and the timing and geography of their diversification. The earliest splits in the tribe were found to separate forest genera in Madagascar and equatorial Africa from the main radiation, and all of the Asian/Indian species sampled formed a younger clade nested well within the African taxa. The tribe appears to have diversified during the Cenozoic, beginningc. 50–32 Ma, with most extant African lineages originating in the Miocene or later, well after the breakup of the Gondwanan landmass. Biogeographical analysis suggests an African origin for the tribe and a single dispersal event founding the Asian platypleurines, although additional taxon sampling and genetic data will be needed to confirm this pattern because key nodes in the tree are still weakly supported. Two Platypleurini genera from Madagascar (PycnaAmyot & Audinet‐Serville,YangaDistant) are found to have originated by late Miocene dispersal of a single lineage from Africa. The genusPlatypleurais recovered as polyphyletic, withPlatypleura signiferaWalker from South Africa and many Asian/Indian species apparently requiring assignment to different genera, and a newPlatypleuraconcept is proposed with the synonymization ofAzanicadaVilletsyn.n.The generaOrapaDistant andHamzaDistant, currently listed within separate tribes but suspected of platypleurine affinity, are nested deeply within the Platypleurini radiation. The tribe Orapinisyn.n. is here synonymized while the tribe Hamzini is pending a decision of the ICZN to preserve nomenclatorial stability. 
    more » « less
  5. The Camponotus subgenus Myrmosaga subgen. rev. from the Malagasy region is revised based on analysis of both qualitative morphological characters and morphometric traits. The multivariate analysis used the Nest Centroid (NC)-clustering method to generate species hypotheses based on 19 continuous morphological traits of minor workers. The proposed species hypotheses were confirmed by cumulative Linear Discriminant Analysis (LDA). Morphometric ratios for the subsets of minor and major workers were used in species descriptions and redefinitions. The present study places the subgenus Myrmopytia syn. nov. in synonymy to Myrmosaga . It recognizes 38 species, of which 19 are newly described: C. aina sp. nov. , C. aro sp. nov. , C. asara sp. nov. , C. atimo sp. nov. , C. bemaheva sp. nov. , C. bozaka sp. nov. , C. daraina sp. nov. , C. harenarum sp. nov. , C. joany sp. nov. , C. karsti sp. nov. , C. kelimaso sp. nov. , C. lokobe sp. nov. , C. mahafaly sp. nov. , C. niavo sp. nov. , C. rotrae sp. nov. , C. sambiranoensis sp. nov. , C. tapia sp. nov. , C. tendryi sp. nov. , C. vano sp. nov. Eleven species are redescribed: C. aurosus Roger, C. cervicalis Roger, C. dufouri Forel, C. gibber Forel, C. hagensii Forel, C. hova Forel, C. hovahovoides Forel, C. immaculatus Forel, C. quadrimaculatus Forel, C. roeseli Forel, C. strangulatus Santschi. The following are raised to species and redescribed: C. becki Santschi stat. nov. , C. boivini Forel stat. rev. , C. cemeryi Özdikmen stat. rev. , C. mixtellus Forel stat. nov. , C. radamae Forel stat. nov. Camponotus maculatus st. fairmairei Santschi syn. nov. , is synonymized under C. boivini . The following are synonymized under C. cervicalis : Camponotus cervicalis gaullei Santschi, syn. nov. ; Camponotus perroti Forel, syn. nov. ; Camponotus perroti aeschylus Forel, syn. nov. ; Camponotus gerberti Donisthorpe, syn. nov. Camponotus dufouri imerinensis Forel, syn. nov. is a synonym of C. dufouri , Camponotus hova var. obscuratus Emery, syn. nov. is a synonym of C. hova , Camponotus quadrimaculatus opacata Emery, syn. nov. is a synonym of C. immaculatus , Camponotus maculatus st. legionarium Santschi, syn. nov. is a synonym of C. roeseli , Camponotus hova maculatoides Emery, syn. nov. is a synonym of C. strangulatus . The following are synonymized under C. quadrimaculatus : Camponotus kelleri Forel, syn. nov. , Camponotus kelleri var. invalidus Forel, syn. nov. , Camponotus quadrimaculatus sellaris Emery, syn. nov. As C. imitator Forel, C. liandia Rakotonirina & Fisher, and C. lubbocki Forel have been recently described and redescribed, only diagnoses and taxonomic discussions are provided. This revision also includes an illustrated species identification key, taxonomic discussions, images, and distribution maps for each species superimposed on the ecoregions of Madagascar. 
    more » « less