Manon Bucher
email: manon.bucher@mnhn.fr

Fabien L Condamine
email: fabien.condamine@gmail.com

Yang Luo

Menglin Wang
email: wangmenglin123@126.com

Thierry Bourgoin
email: thierry.bourgoin@mnhn.fr

Affiliations

Phylogeny and diversification of planthoppers (Hemiptera: Fulgoromorpha) based on a comprehensive molecular dataset and large taxon sampling

Keywords: Fulgoromorpha, Molecular phylogeny, Dating analyses, Macroevolution, Diversification 1.Introduction

) the unexpected paraphyly of Delphacidae whose Protodelphacida seem more related to Cixiidae than to other Delphacidae; (2) the group Meenoplidae-Kinnaridae recovered sister to the remaining Fulgoroidea families; (3) the early branching node of Tettigometridae sister of all the other families; (4) the Achilidae-Derbidae clade with Achilidae Plectoderini including Achilixiidae recovered as monophyletic as well as the Fulgoridae-Dictyopharidae clade; and (5) the Tropiduchidae placed sister to the other so called 'higher' families (sec. Shcherbakov, 2006). Our divergence times analysis, calibrated with a set of duly verified fossils, suggests that the first diversification of planthoppers occurred in the Early Triassic around 240 Mya and those of the superfamilies Delphacoidea and Fulgoroidea in the Middle-Late Triassic around 210 Mya and 230 Mya, respectively. By the end of the Jurassic, all major planthopper lineages were originated, and all families, around 125 Mya, might have been driven in their distribution and evolution (in their first subfamilial divisions) by the geographical constraints of the Gondwanan break-up. Rapid evolutionary radiations occurred particularly in Fulgoridae around 125-130 Mya. Our results stress the importance of the good quality of the sequences used in the molecular analyzes and the primordial importance of a large sampling when analyzing the phylogeny of the group.

as the superfamily Delphacoidea Leach, 1815 sister clade to the remaining planthopper families all classified within the Fulgoroidea Latreille, 1807 (sec. Bourgoin & Szwedo, 2022); (2) a paraphyletic group of five families (Kinnaridae Muir, 1925+ Meenoplidae Fieber, 1872), (Achilidae Stål, 1866+ Achilixiidae Muir, 1923) and Derbidae Spinola, 1839; (3) a monophyletic group of so-called 'higher Fulgoroidea' families sec. [START_REF] Shcherbakov | The earliest find of Tropiduchidae (Homoptera: Auchenorrhyncha), representing a new tribe, from the Eocene of Green River, USA, with notes on the fossil record of higher Fulgoroidea[END_REF], with: Acanaloniidae Amyot & Audinet-Serville, 1843, Caliscelidae Amyot & Audinet-Serville, 1843, Flatidae Spinola, 1839, Issidae Spinola, 1839, Nogodinidae Melichar, 1898, Tettigometridae, Tropiduchidae Stål, 1866, Ricaniidae Amyot & Audinet-Serville, 1843 forming a clade sister to (4) an undefined group of the last four extant families (Fulgoridae Latreille, 1807 + Dictyopharidae Spinola, 1839) and (Lophopidae Stål, 1866+ Eurybrachidae Stål, 1862). This topology was updated by [START_REF] Brysz | Jeweled Achilidae -A new look at their systematics and relation to other Fulgoroidea (Hemiptera)[END_REF] with the fossil families, although before the recognition of Fulgoridoidea by Bourgoin et Szwedo (2022[START_REF] Bourgoin | FLOW (Fulgoromorpha Lists on The Web): a world knowledge base dedicated to Fulgoromorpha[END_REF].

However, behind this conventional agreement, many problems remain as these four divisions form a poorly useful classification not being based on monophyletic units that should offer strong frames of reference in addressing planthopper evolutionary history. Moreover, from the species to the family level, a consensus topology yet remains to emerge. Indeed, recent molecular phylogenies [START_REF] Song | A preliminary molecular phylogeny of planthoppers (Hemiptera: Fulgoroidea) based on nuclear and mitochondrial DNA sequences[END_REF][START_REF] Huang | Sequencing and analysis of the complete mitochondrial genome of Changeondelphax velitchkovskyi (Hemiptera: Fulgoroidea)[END_REF][START_REF] Yu | The complete mitochondrial genome of Ugyops sp. (Hemiptera: Delphacidae)[END_REF][START_REF] Wang | Characterization of the complete mitochondrial genomes of two species of the genus Aphaena Guérin-Méneville (Hemiptera : Fulgoridae) and its phylogenetic implications[END_REF][START_REF] Xu | Comparative analysis of the complete mitochondrial genomes of five Achilidae species (Hemiptera: Fulgoroidea) and other Fulgoroidea reveals conserved mitochondrial genome organization[END_REF][START_REF] Kim | Complete Mitochondrial Genomes of Metcalfa pruinosa and Salurnis marginella (Hemiptera: Flatidae) : Genomic Comparison and Phylogenetic Inference in Fulgoroidea[END_REF] have come to challenge the results of [START_REF] Urban | Evolution of the planthoppers (Insecta: hemiptera: Fulgoroidea)[END_REF], to the point that attempting a simple consensus of all the phylogenetic hypotheses would lead to retain only a few polytomous nodes, even for the seemingly well-established sister families. A review of these studies suggests that most of the issues pertain to taxa and characters sampling [START_REF] Bucher | Which biases still hamper a solid planthopper phylogeny?[END_REF]. Indeed, all molecular phylogenies of planthopper families were conducted either on a limited diversity of characters with only a few gene sequences in the pioneer studies [START_REF] Bourgoin | Molecular Phylogeny of Fulgoromorpha (Insecta, Hemiptera, Archaeorrhyncha). The Enigmatic Tettigometridae: Evolutionary Affiliations and Historical Biogeography[END_REF][START_REF] Yeh | Fulgoromorpha phylogeny based on 28S rDNA nucleotide sequence[END_REF][START_REF] Yeh | Phylogenetic Relationships of the Tropiduchidaegroup (Homoptera: Fulgoroidea) of Planthoppers Inferred through Nucleotide Sequences[END_REF][START_REF] Yeh | A Molecular Phylogeny of Planthoppers (Hemiptera : Fulgoroidea) Inferred from Mitochondrial 16S rDNA Sequences[END_REF], or later on a too restricted taxonomic sampling never exceeding ~130 taxa [START_REF] Urban | Evolution of the planthoppers (Insecta: hemiptera: Fulgoroidea)[END_REF], 2012;[START_REF] Song | A preliminary molecular phylogeny of planthoppers (Hemiptera: Fulgoroidea) based on nuclear and mitochondrial DNA sequences[END_REF][START_REF] Huang | Sequencing and analysis of the complete mitochondrial genome of Changeondelphax velitchkovskyi (Hemiptera: Fulgoroidea)[END_REF][START_REF] Yu | The complete mitochondrial genome of Ugyops sp. (Hemiptera: Delphacidae)[END_REF][START_REF] Wang | Characterization of the complete mitochondrial genomes of two species of the genus Aphaena Guérin-Méneville (Hemiptera : Fulgoridae) and its phylogenetic implications[END_REF][START_REF] Xu | Comparative analysis of the complete mitochondrial genomes of five Achilidae species (Hemiptera: Fulgoroidea) and other Fulgoroidea reveals conserved mitochondrial genome organization[END_REF][START_REF] Kim | Complete Mitochondrial Genomes of Metcalfa pruinosa and Salurnis marginella (Hemiptera: Flatidae) : Genomic Comparison and Phylogenetic Inference in Fulgoroidea[END_REF][START_REF] Michalik | Variable organization of symbiont-containing tissue across planthoppers hosting different heritable endosymbionts[END_REF], failing to adequately consider the high diversity and disparity of planthoppers. Altogether, this resulted in only partial or biased representations of the planthopper taxonomic diversity and phylogeny, leading to a weak congruence of the different hypotheses proposed until now.

Consequently, many questions about the phylogenetic relationships and evolutionary patterns of planthoppers, already formulated more than 20 years ago [START_REF] Bourgoin | Molecular Phylogeny of Fulgoromorpha (Insecta, Hemiptera, Archaeorrhyncha). The Enigmatic Tettigometridae: Evolutionary Affiliations and Historical Biogeography[END_REF], still continue to remain and can be summarized in five major issues as follows:

(1) What is the first cladogenetic event that splits extant planthoppers? For a long time, the hypothesis of Tettigometridae sister to the rest of planthoppers was consensual for the majority of morphological approaches [START_REF] Muir | On the classification of the Fulgoroidea (Homoptera)[END_REF][START_REF] Muir | On the classification of the Fulgoroidea (Homoptera). 1930. On the classification of the Fulgoroidea[END_REF][START_REF] Asche | Preliminary thoughts on the phylogeny of Fulgoromorpha (Homoptera: Auchenorrhyncha)[END_REF][START_REF] Emeljanov | An attempt of construction of the phylogenetic tree of the planthoppers (Homoptera, Cicadina)[END_REF][START_REF] Chen | The metatarsi of the Fulgoroidea (Homoptera: Auchenorrhyncha)[END_REF]. Later, the Meenoplidae sister to all other planthoppers were supported by some molecular approaches [START_REF] Yeh | Phylogenetic Relationships of the Tropiduchidaegroup (Homoptera: Fulgoroidea) of Planthoppers Inferred through Nucleotide Sequences[END_REF][START_REF] Yeh | Fulgoromorpha phylogeny based on 28S rDNA nucleotide sequence[END_REF], while the clade Cixiidae+Delphacidae, sister to all other families was finally proposed by most molecular analyses [START_REF] Bourgoin | Molecular Phylogeny of Fulgoromorpha (Insecta, Hemiptera, Archaeorrhyncha). The Enigmatic Tettigometridae: Evolutionary Affiliations and Historical Biogeography[END_REF][START_REF] Yeh | A Molecular Phylogeny of Planthoppers (Hemiptera : Fulgoroidea) Inferred from Mitochondrial 16S rDNA Sequences[END_REF][START_REF] Urban | Evolution of the planthoppers (Insecta: hemiptera: Fulgoroidea)[END_REF] Cryan & Urban, 2012;[START_REF] Song | A preliminary molecular phylogeny of planthoppers (Hemiptera: Fulgoroidea) based on nuclear and mitochondrial DNA sequences[END_REF][START_REF] Huang | Sequencing and analysis of the complete mitochondrial genome of Changeondelphax velitchkovskyi (Hemiptera: Fulgoroidea)[END_REF][START_REF] Yu | The complete mitochondrial genome of Ugyops sp. (Hemiptera: Delphacidae)[END_REF][START_REF] Kim | Complete Mitochondrial Genomes of Metcalfa pruinosa and Salurnis marginella (Hemiptera: Flatidae) : Genomic Comparison and Phylogenetic Inference in Fulgoroidea[END_REF]. Recent fossil-based analyses (Bourgoin & Szwedo, 2022[START_REF] Bourgoin | FLOW (Fulgoromorpha Lists on The Web): a world knowledge base dedicated to Fulgoromorpha[END_REF][START_REF] Zhao | The oldest genus and species of the fulgoroidian lineage in hemipteran Fulgoroidea from the Lower Cretaceous Laiyang Formation of China[END_REF] led to formalize this early split dividing extant planthopper families into Delphacoidea (Cixiidae+Delphacidae) and Fulgoroidea (all other families).

(2) Can we recognize large clades among these superfamilies? In addition of the splitting of Delphacoidea and Fulgoroidea, are the general divisions proposed by [START_REF] Bartlett | The Diversity of the True Hoppers (Hemiptera : Auchenorrhyncha)[END_REF] supported, and which families do they include? In particular, is the "issidoid group" proposed by Gnezdilov 2013 (grouping Issidae, Caliscelidae, Acanaloniidae, Tropiduchidae and Nogodinidae) phylogenetically supported? (3) Among the supposed monophyletic family pairs traditionally accepted, which ones can we retain: Cixiidae+Delphacidae, Kinnaridae+Meenoplidae, Achilidae+Achilixiidae, Fulgoridae+Dictyopharidae, Lophopidae+Eurybrachidae? (4) While the monophyly of most families remain to be confirmed with phylogenetic analyses [START_REF] Bartlett | The Diversity of the True Hoppers (Hemiptera : Auchenorrhyncha)[END_REF], should we reconsider the monophyly of Achilidae including Achilixiidae [START_REF] Liang | Morphology of antennal sensilla in Achilixius sandakanensis Muir (Hemiptera: Fulgoromorpha: Achilixiidae) with comments on the phylogenetic position of the Achilixiidae[END_REF]? Likewise, the relatedness of the different lineages within Cixiidae needs to be more closely investigated, as three incongruent evolutionary patterns are now proposed in recent molecular studies with (a) Cixiidae as a monophyletic unit sister to Delphacidae [START_REF] Song | A preliminary molecular phylogeny of planthoppers (Hemiptera: Fulgoroidea) based on nuclear and mitochondrial DNA sequences[END_REF][START_REF] Urban | Evolution of the planthoppers (Insecta: hemiptera: Fulgoroidea)[END_REF], 2012;[START_REF] Yu | The complete mitochondrial genome of Ugyops sp. (Hemiptera: Delphacidae)[END_REF]; or (b) Cixiidae as paraphyletic and including Delphacidae [START_REF] Ceotto | Molecular phylogenetics of cixiid planthoppers (Hemiptera : Fulgoromorpha): New insights from combined analyses of mitochondrial and nuclear genes[END_REF][START_REF] Urban | Evolution of Delphacidae (Hemiptera : Fulgoroidea): combined-evidence phylogenetics reveals importance of grass host shifts[END_REF], or (c) Delphacidae as paraphyletic inluding subfamily Asiracinae Motschulsky, 1863 sister to all Cixiidae [START_REF] Bucher | Which biases still hamper a solid planthopper phylogeny?[END_REF][START_REF] Kim | Complete Mitochondrial Genomes of Metcalfa pruinosa and Salurnis marginella (Hemiptera: Flatidae) : Genomic Comparison and Phylogenetic Inference in Fulgoroidea[END_REF]. In addition, a particular attention is needed for the taxonomic value of Nogodinidae that are now repeatedly found to be paraphyletic [START_REF] Yeh | Fulgoromorpha phylogeny based on 28S rDNA nucleotide sequence[END_REF][START_REF] Yeh | A Molecular Phylogeny of Planthoppers (Hemiptera : Fulgoroidea) Inferred from Mitochondrial 16S rDNA Sequences[END_REF][START_REF] Urban | Evolution of the planthoppers (Insecta: hemiptera: Fulgoroidea)[END_REF][START_REF] Song | A preliminary molecular phylogeny of planthoppers (Hemiptera: Fulgoroidea) based on nuclear and mitochondrial DNA sequences[END_REF][START_REF] Bucher | Which biases still hamper a solid planthopper phylogeny?[END_REF], as well as the issue to retain a separate paraphyletic family Kinnaridae including Meenoplidae [START_REF] Bourgoin | Female genitalia in Hemiptera Fulgoromorpha, morphological and phylogenetic data[END_REF].

(5) Lastly, questions remain on the placement of some families whose relationships were strongly not consensual in the different phylogenetic inferences i.e. Caliscelidae, Lophopidae, Tettigometridae and Tropiduchidae [START_REF] Bourgoin | Molecular Phylogeny of Fulgoromorpha (Insecta, Hemiptera, Archaeorrhyncha). The Enigmatic Tettigometridae: Evolutionary Affiliations and Historical Biogeography[END_REF][START_REF] Bucher | Which biases still hamper a solid planthopper phylogeny?[END_REF][START_REF] Song | A preliminary molecular phylogeny of planthoppers (Hemiptera: Fulgoroidea) based on nuclear and mitochondrial DNA sequences[END_REF][START_REF] Urban | Evolution of the planthoppers (Insecta: hemiptera: Fulgoroidea)[END_REF][START_REF] Yeh | A Molecular Phylogeny of Planthoppers (Hemiptera : Fulgoroidea) Inferred from Mitochondrial 16S rDNA Sequences[END_REF].

The weakness of current phylogenetic results is also attested by several molecular dating studies, which suggested conflicting hypotheses on the age of diversification of Fulgoromorpha (Fig. A). [START_REF] Song | A preliminary molecular phylogeny of planthoppers (Hemiptera: Fulgoroidea) based on nuclear and mitochondrial DNA sequences[END_REF] Fulgoromorpha (Bourgoin & Szwedo, 2022). With a difference that can reach more than 50 millions of years, the large time gap in the dating analyses underlines the need for a better understanding of planthopper evolutionary history, with more constrained divergence times associated with the great periods of diversification of the different lineages.

Addressing these questions, this study proposes a new investigation of the evolutionary history of extant Fulgoromorpha, through a phylogenetic analysis based on a more substantial molecular dataset, combining from previously published molecular works with newly published data. Second, a dating analysis was conducted to estimate divergence times of the different planthopper lineages. Using duly identified and classified fossils, a new estimate of their diversification timeframe is therefore proposed, aiming to refine the temporal dimension of the evolutionary framework of planthoppers. Accordingly, particular attention has been paid to the taxonomic sampling to better represent the diversity of the Fulgoromorpha, allowing to test the monophyly of the families and some subfamilies. This is the largest phylogenetic analysis integrating a comprehensive number of planthopper taxa. We take the opportunity to discuss the importance of sampling and data quality, and the theoretical and methodological biases responsible for the instability and lack of consensus on the evolutionary history of the Fulgoromorpha.

Material and methods

Taxonomic sampling

With the objective of providing a dataset that is as representative as possible of the diversity of Fulgoromorpha, the selection of terminal taxa was done with maximizing the coverage of the 299 taxonomic suprageneric taxa currently recognized in the planthopper classification available on FLOW [START_REF] Bourgoin | FLOW (Fulgoromorpha Lists on The Web): a world knowledge base dedicated to Fulgoromorpha[END_REF]. A preliminary taxonomic sampling was achieved with the selection of at least one to several species (with their molecular data sequences already available and published in previous studies) to represent each of these suprageneric taxonomic rank. From this first sampling of our ingroup taxa (572 sequenced specimens), a subsequent selection was made to discard doubtful molecular data yet previously used in published phylogenies, reducing the taxon sampling to 531. The process of verifying molecular data and excluding doubtful taxa from the sampling are detailed in the following section. In addition, 20 outgroup species were selected to represent the diversity of lineages within hemipteran suborders: Cicadomorpha Evans, 1946, whose taxonomic diversity was covered by 11 families distributed in 3 super-families with Cercopoidea Leach, 1815 (5 taxa), Cicadoidea Westwood, 1840 (2 taxa), Membracoidea Rafinesque, 1815 (4 taxa); Coleorrhyncha Myers & China, 1929 (2 taxa); Sternorrhyncha Duméril, 1806 (3 taxa); Heteroptera Latreille, 1810 (3 taxa), plus one species of Thysanoptera Haliday, 1836, sister-group to Hemiptera [START_REF] Johnson | Phylogenomics and the evolution of hemipteroid insects[END_REF].

Terminology and compatibility of molecular sequences

All DNA sequences were retrieved from data available on GenBank. Genes were selected to constitute a heterogeneous and varied dataset, including coding and non-coding fragments. Six molecular markers were included in our analysis with three non-coding nuclear genes (18S, 28S D3-D5, 28S D6-D10), the nuclear coding Wingless (Wg), and two coding mitochondrial genes: Cytochrome Oxydase I (COI) and Cytochrome b (Cytb). Wherever possible, sequences with the same voucher code were privileged to limit the use of chimeric taxa and get rid of potential misidentification. As these sequences came from different studies, a nomenclature work and delineation of molecular sequences had to be carried out to verify that they were comparable and could be considered as homologous markers. We used Geneious 2022.1.1 [START_REF] Kearse | Geneious Basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data[END_REF] to define the domains to which the sequences corresponded. For this purpose, all sequences for each RNA marker were mapped to a reference sequence of the complete gene of Brachyplatys subaeneus Westwood, 1837 for 18S (accession number: KY495690) and of Geisha distinctissima (Walker, 1858) for COI sequences (accession number: NC_012617.1). Delineation of 28S domains was performed by the same mapping process, following the secondary structure established by the annotated 28S guide of Eurydema maracandica Oshanin, 1871 [START_REF] Yu | Secondary structure models of 18S and 28S rRNAs of the true bugs based on complete rDNA sequences of Eurydema maracandica Oshanin, 1871 (Heteroptera, Pentatomidae)[END_REF] and by comparing primers used in the phylogenetic studies, which target specific domains for sequencing. The nomenclatural definition of the markers used in our study is summarized in Table S1. When genes were sequenced in multiple fragments (e.g. 18S; 28S D6-D10), we assembled the different regions to obtain continuous sequences with Geneious 2022.1.1. All assembled sequences from separate fragments are reported in Table S2.

Conversely, for continuously sequenced RNA genes, only fragments corresponding to the interest domains were then extracted. Size and location of extracted fragments from complete sequences are indicated in Table S2. For some taxa, COI and Cytb sequences were extracted from complete mitochondrial genomes. The mapping procedure revealed that the sequenced COI of several studies does not correspond to the true COI 5P barcode as usually recognized, but to the 3P terminal region of this gene. Two data blocks were thus identified for the gene COI. Complete COI sequences were also present in our dataset. Therefore, all COI sequences (3P, 5P and complete) were considered as a single marker with '?' for the corresponding missing parts in the concerned taxa.

Detection of erroneous data for the final dataset

Several erroneous data extracted from GenBank were identified during preliminary analyses.

All sequences selected in our preliminary sampling have therefore been rechecked by: (1) BLAST; (2) Detection of sequence divergence within the same family during alignment; [START_REF] Muir | On the classification of the Fulgoroidea (Homoptera)[END_REF] comparison of sequences divergence from same taxa with different specimens when it was possible; and (4) Phylogenetic inference to detect artefact and to double-check contaminations and reconstruction artefacts. Details on sequences that are contaminated or qualified as doubtful -and therefore that should be avoided in future molecular analyses -is appended in Supplementary Table S3 and will be reported in FLOW in the 'molecular data' section (Bourgoin 2023, in preparation). Our final taxonomic sampling contains 551 taxa including ingroup and outgroup (Supplementary Table S2). Eleven COI sequences are not published on GenBank but attached as supplementary data (Dataset S1), whose specimens are indicated in Table S2. New sequences produced are newly published in this study with 181 newly generated sequences for five families (Achilidae, Cixiidae, Derbidae, Issidae, Nogodinidae) corresponding to five genes. All references to these new sequences are specified in Table S2.

The suprageneric taxonomic diversity covered by the final dataset is illustrated in Table A.

Alignment and phylogenetic inferences

Multiple sequence alignment was carried out in MAFFT 7.450 as implemented in Geneious. coding sequences (generated with a various set of primers in the different studies), could appear highly fragmented and not always congruent in coverage in the outer regions. We corrected these problems by slightly refining the alignment manually. Secondary structure for 18S and 28S genes was not considered in the alignment process and highly variables internal regions have been kept with the automatic alignment to limit eye-driven homology hypotheses. We checked that alignments for protein-coding genes were congruent with codon reading frame using Geneious. In our study, the supermatrix approach (de Queiroz & Gatesy, 2007) was preferred to the supertree method [START_REF] Gordon | Consensus supertrees: the synthesis of rooted trees containing overlapping sets of labeled leaves[END_REF]. We used the software SequenceMatrix 1.7.7 [START_REF] Vaidya | SequenceMatrix: concatenation software for the fast assembly of multi-gene datasets with character set and codon information[END_REF] to concatenate the dataset.

We performed maximum likelihood analyses using IQ-TREE 2.1.2 [START_REF] Nguyen | IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies[END_REF] with a dataset partitioned by genes and codon positions. For each partition, best-fitting substitution models were selected by PartitionFinder 2.2.1 [START_REF] Lanfear | PartitionFinder 2: new methods for selecting partitioned models of evolution for molecular and morphological phylogenetic analyses[END_REF] under the Akaike Information Criterion [START_REF] Akaike | 2nd International Symposium on Information Theory[END_REF]. Details on optimizing partition schemes and substitution models are shown in Table S4. Phylogenetic analyses were carried out with the more thorough and slower nearest neighbor interchange (NNI) search. The perturbation strength was defined to 0.2 as recommended by IQ-TREE with datasets containing short sequences and the number of unsuccessful iterations to stop was set to 10,000. The final analysis was repeat 10 times to test the robustness and stability of the results and to avoid the tree search getting stuck in a local optimum. Support for nodes was evaluated with 1,000 ultrafast bootstrap (UFBS) approximations (considering UFBS values ≥95 as strong node support). All phylogenetic analyses were performed on the computer cluster CIPRES Science Gateway 3.3 [START_REF] Miller | A RESTful API for Access to Phylogenetic Tools via the CIPRES Science Gateway[END_REF].

Fossil calibrations

To calibrate the tree, we selected 10 fossils (Table B) according to the best practices suggested by [START_REF] Parham | Best Practices for Justifying Fossil Calibrations[END_REF] for justifying fossil calibrations. As none of the present fossils were previously included in a phylogenetic analysis, attribution to a phylogenetic lineage was carried out according to the apomorphy-based method. Fossils that did not fulfill these specific conditions were not included in the dating analysis. As a result, the selected fossils are not necessarily the oldest known for a given group but are those whose attribution to a systematic position and stratigraphic age have been verified and justified. All fossils were used as minimum age constraints for their specific nodes: minimum ages of fossils correspond to the youngest limit of the stratigraphic interval in which the fossil was found or to the precise age revealed by isotopic studies. We considered here that the oldest boundary of node constraints by Fulgoromorpha fossils (related to extant lineage) in our analysis corresponds to the minimum age of †Aviorrhyncha magnifica Nel et al., 2013 (310 Mya). Due to the limited taxonomic sampling and computational time, outgroup taxa were not included in the dataset for dating analyses [START_REF] Drummond | Bayesian evolutionary analysis with BEAST[END_REF]. We choose to constrain the age of the root as an additional calibration. In our study, tree root height corresponds to the diversification of Fulgoromorpha with the divergence of Delphacoidea and Fulgoroidea. To calibrate this node, we used the age provided by the fossil †Fulgoridiella raetica Becker-Migdisova, 1962 dated from the Early Jurassic (199.6 Mya) as the minimum age, and the maximum age was set with the age of Insecta (406 Mya) according to [START_REF] Misof | Phylogenomics resolves the timing and pattern of insect evolution[END_REF] (with a 95% confidence interval of 451-373 Mya).

Bayesian relaxed-clock analyses

Molecular dating analyses were carried out with BEAST 1.10.4 [START_REF] Suchard | Bayesian phylogenetic and phylodynamic data integration using BEAST 1.10[END_REF]. BEAUti 1.10.4 was used to create the input for BEAST. We used a reduced partitioned dataset without partitions by coding positions to limit the number of estimated parameters for improving convergence of the analyses. Best-fitting substitution models (Supplementary Table S4) were selected by PartitionFinder 2.2.1 with a constrained partition as an input parameter and all the parameters were estimated in BEAST. We implemented 3 partitioned clock models with 18S + 28S D3-D5 + Wg; 28S D6-D10; COI + Cytb under an uncorrelated lognormal distribution (UCLD). We defined a uniform prior distribution between 0 and 1, with a starting value of 0.1 on the mean of the UCLD model. We set the tree prior to the birth-death process with the following priors: a uniform prior between 0 and 10 with a starting value of 0.1 for the mean growth rate, and a uniform prior between 0 and 2 with a starting value of 0.5 for the relative death rate. We used the most conservative approach of node-dating, in which fossil constraints were applied on the stem-node as proposed by [START_REF] Parham | Best Practices for Justifying Fossil Calibrations[END_REF]. Here, morphological support for assigning a fossil to an extant lineage within a clade was limited to avoid overestimating divergence times, the parent node was therefore chosen to define the minimum appearance age of a group. Table S5 summarizes fossil clade assignment and priors used in the analysis with distribution and shape. We generated a starting tree with the topology resulting from the phylogenetic inference: the topology was ultrametricized to fit our calibration scheme in BEAST using the chronos function of the R package ape [START_REF] Paradis | ape 5.0: an environment for modern phylogenetics and evolutionary analyses in R[END_REF]. The tree model operators were turned off to keep the topology fixed so that only branch lengths were estimated in the dating analysis. All others parameters were kept to their default prior distribution. BEAST analysis was run for 300 million generations and sampled every 30,000 generations. All analyses were performed on the computer cluster CIPRES Science Gateway 3.3. We used Tracer 1.7.1 to assess the convergence of the run by examining the log files to check the ESS values for all parameters. Maximum clade credibility tree was reconstructed with TreeAnnotator 1.10.4 after removing a burn-in of 20%.

Results

Dataset characterization and phylogenetic analyses

Taxonomic sampling covers 80% of the suprageneric taxonomic rank within the extant Fulgoromorpha. Even when combining all data, there was still great heterogeneity in the representation of intra-familial taxonomic diversity for some families included in our study, particularly for Flatidae, Tropiduchidae and Nogodinidae. The combined dataset of 6 genes contains 8,159 characters for 551 taxa (including outgroups) with lengths per locus of: 2,145 pb (18S); 786 pb (28S D3D5); 3,169 pb (28S D6D10); 1,250 pb (COI); 491 pb (Cytb); 318 pb (Wg). The alignment contains 3,729 parsimony informative sites with 53.9% of missing data (?) according to the nucleotide statistics estimated in Geneious. Same topology was recovered in the 10 independent phylogenetic analyses. The most likely tree (ln L= -258,102.81) is fully resolved and most of the nodes are supported with high UFBS: 25.7% of nodes have UFBS=100 and 40.4% of nodes supported with UFBS≥95 with a median of node support of 87. The least supported nodes generally involve nodes reflecting infra-family relationships.

Planthopper phylogeny

The simplified phylogeny with major lineages collapsed is presented in for Delphacidae and Cixiidae, respectively), suggests here that Delphacidae is paraphyletic, and splits into two separated groups: Eudelphacida sec. [START_REF] Asche | Vizcayinae, a new subfamily of Delphacidae with revision of Vizcaya Muir (Homoptera: Fulgoroidea), a significant phylogenetic link[END_REF](= Delphacinae Leach, 1815 sec. [START_REF] Emeljanov | On the question of the classification and phylogeny of the Delphacidae (Homoptera, Cicadina), with reference to larval characters[END_REF] on one side (Clade 2, UFBS=100), and clade 3 (UFBS=68) including Protodelphacida sec. Emeljanov, 1996 (= Asiracinae sec. [START_REF] Asche | Zur Phylogenie der Delphacidae Leach, 1815 (Homoptera Cicadina Fulgoromorpha)[END_REF][START_REF] Asche | Vizcayinae, a new subfamily of Delphacidae with revision of Vizcaya Muir (Homoptera: Fulgoroidea), a significant phylogenetic link[END_REF], as a single monophyletic lineage (versus [START_REF] Asche | Zur Phylogenie der Delphacidae Leach, 1815 (Homoptera Cicadina Fulgoromorpha)[END_REF][START_REF] Asche | Vizcayinae, a new subfamily of Delphacidae with revision of Vizcaya Muir (Homoptera: Fulgoroidea), a significant phylogenetic link[END_REF][START_REF] Emeljanov | On the question of the classification and phylogeny of the Delphacidae (Homoptera, Cicadina), with reference to larval characters[END_REF], sister-group to Cixiidae on the other side. In our preliminary analyses, computed with a reduced taxonomic sampling, Delphacidae was found monophyletic only once, with low support (no stability of topologies in 10 independent analyses). Other phylogenetic units among Eudelphacida are congruent with the hypotheses proposed by [START_REF] Asche | Zur Phylogenie der Delphacidae Leach, 1815 (Homoptera Cicadina Fulgoromorpha)[END_REF][START_REF] Asche | Vizcayinae, a new subfamily of Delphacidae with revision of Vizcaya Muir (Homoptera: Fulgoroidea), a significant phylogenetic link[END_REF], [START_REF] Emeljanov | On the question of the classification and phylogeny of the Delphacidae (Homoptera, Cicadina), with reference to larval characters[END_REF], [START_REF] Urban | Evolution of Delphacidae (Hemiptera : Fulgoroidea): combined-evidence phylogenetics reveals importance of grass host shifts[END_REF] and [START_REF] Huang | Resolving phylogenetic relationships of Delphacini and Tropidocephalini (Hemiptera: Delphacidae: Delphacinae) as inferred from four genetic loci[END_REF], as all of the previously recognized lineages either as subfamilies or tribes (Kelisiini/ae Wagner, 1963, Stenocranini/ae Wagner, 1963, Delphacinae Leach, 1815) are monophyletic (BS ≥ 99), as well as Saccharosydnini, Tropidocephalini, Delphacini with UFBS ≥ 94). However, a strong support for a sister relationship between Kelisiini/ae and Stenocranini/ae is observed (versus Emeljanov, 1996). clade showing an instable basal branching with any of the two families according to our preliminary analyses, always introducing a low UFBS at the corresponding node. In Dictyopharidae, all tribes included in our sampling are retrieved as monophyletic, with the exception of Orthopagini Emeljanov, 1983. In Fulgoridae, intra-familial relationships within the family are not congruent with the current classification -based on the suggestions of [START_REF] Lallemand | Contribution à l'etude des Fulgoridae (Hemiptera) (1re note)[END_REF][START_REF] Lallemand | Description de nouvelles espèces de Fulgorides d'Asie et d'Afrique[END_REF][START_REF] Lallemand | De quibusdam Fulgoris. Entomologische Mitteilungen aus dem Zoologischen Staatsinstitut u[END_REF][START_REF] Lallemand | Nouvelles espèces d'Homoptères[END_REF], [START_REF] Emeljanov | Problema razgranichenhiya semeïstv Fulgoridae i Dictyopharidae (Homoptera, Auchenorrhyncha)[END_REF][START_REF] Emeljanov | A new genus and tribe of Strongylodematinae (Homoptera: Fulgoridae)[END_REF]) -and the phylogenetic results of [START_REF] Urban | Entomologically famous, evolutionarily unexplored : The first phylogeny of the lanternfly family Fulgoridae (Insecta: Hemiptera: Fulgoroidea)[END_REF].

The strongly supported Tropiduchidae family (Clade 18, UFBS=100) was retrieved as sister lineage to remaining Fulgoroidea families (Fig. C.4, Clade 17, UFBS=59). However, this placement remains questionable due to its low support at node. In preliminary analyses the placement of this family was not stable, moving within the topology with always a low support, especially as sister lineage to (Dictyopharidae + Fulgoridae). Mongolianina. As expected, Nogodinidae appear as a polyphyletic grouping, split according to the genera in various place within the 'higher' planthoppers. Mya). Our dating results indicate that all major lineages of extant Fulgoromorpha were already present by the Late Jurassic. Nevertheless, 'higher' planthopper families diverged during the Early Cretaceous with rapid diversification during middle to Late Cretaceous. Because of our conservative strategy of uniform distributions for node calibrations, the 95% CIs of our analysis were relatively wide (Fig. S1). All divergence time estimates for each family are reported in Table C.

Within clade 19 (

Dating analysis

Discussion

Our study revealed unsurprisingly that Fulgoromorpha is composed of a monophyletic superfamily Delphacoidea sister to a monophyletic superfamily Fulgoroidea. This hypothesis is now considered as consensual, being both supported by morphological analyses [START_REF] Asche | Preliminary thoughts on the phylogeny of Fulgoromorpha (Homoptera: Auchenorrhyncha)[END_REF][START_REF] Emeljanov | An attempt of construction of the phylogenetic tree of the planthoppers (Homoptera, Cicadina)[END_REF][START_REF] Bourgoin | Female genitalia in Hemiptera Fulgoromorpha, morphological and phylogenetic data[END_REF] including from a fossil perspective (Bourgoin & Szwedo, 2022[START_REF] Bourgoin | FLOW (Fulgoromorpha Lists on The Web): a world knowledge base dedicated to Fulgoromorpha[END_REF], and most molecular phylogenies (Cryan & Urban, 2012;[START_REF] Kim | Complete Mitochondrial Genomes of Metcalfa pruinosa and Salurnis marginella (Hemiptera: Flatidae) : Genomic Comparison and Phylogenetic Inference in Fulgoroidea[END_REF][START_REF] Song | A preliminary molecular phylogeny of planthoppers (Hemiptera: Fulgoroidea) based on nuclear and mitochondrial DNA sequences[END_REF][START_REF] Urban | Evolution of the planthoppers (Insecta: hemiptera: Fulgoroidea)[END_REF], 2012;[START_REF] Yu | The complete mitochondrial genome of Ugyops sp. (Hemiptera: Delphacidae)[END_REF][START_REF] Michalik | Variable organization of symbiont-containing tissue across planthoppers hosting different heritable endosymbionts[END_REF].

Paraphyly of Delphacidae

In most phylogenetic studies, Cixiidae and Delphacidae were considered as sister monophyletic families. However, this result was challenged by [START_REF] Urban | Evolution of the planthoppers (Insecta: hemiptera: Fulgoroidea)[END_REF], which depicted a paraphyletic Cixiidae including the Delphacidae (a scenario already suggested by [START_REF] Asche | Zur Phylogenie der Delphacidae Leach, 1815 (Homoptera Cicadina Fulgoromorpha)[END_REF] with a maximum parsimony analysis based on four selected species in both Cixiidae and Delphacidae, although a Bayesian analysis of the same sampling recovered two monophyletic sister taxa. [START_REF] Ceotto | Molecular phylogenetics of cixiid planthoppers (Hemiptera : Fulgoromorpha): New insights from combined analyses of mitochondrial and nuclear genes[END_REF] ). Delphacidae was found monophyletic only once in preliminary analyses under conditions of taxonomic under-sampling within these two families and with low support (no stability of topologies in 10 independent analyses). These successive results reveal that increasing the molecular sampling of delphacid taxa with a better sampling at the tribal level, even confronted to a large cixiid sampling, disclose the paraphyly of the family as a stable characteristic.

From a morphological perspective this result is counter intuitive, rejecting the most significant and unique apomorphic character of Delphacidae with the presence of a movable and apical metatibial spur exposing a gradual transformation series of the spur conformation [START_REF] Markevich | Morphological reassessment of the movable calcar of delphacid planthoppers (Hemiptera: Fulgoromorpha: Delphacidae)[END_REF]. It suggests two possible scenarios: (1) the spur appeared twice, independently in Protodelphacida in which it remained quite simple, more or less conical, never denticulated, versus flattened and always dentated in Eudelphacida [START_REF] Asche | Vizcayinae, a new subfamily of Delphacidae with revision of Vizcaya Muir (Homoptera: Fulgoroidea), a significant phylogenetic link[END_REF][START_REF] Markevich | Morphological reassessment of the movable calcar of delphacid planthoppers (Hemiptera: Fulgoromorpha: Delphacidae)[END_REF]; or (2) the spur appeared once in Delphacoidea much probably unarmed, secondly disappearing in Cixiidae therefore miming the plesiomorphic pattern of other planthoppers, leading to consider them as 'cryptic' delphacids. While this second scenario would apparently be more parsimonious (one apomorphic acquisition secondarily absent in Cixiidae), we would favor the first one (two independent acquisitions) because up to now, all Cretaceous fossils observed exhibit a cixioid pattern, without any metatibial spur, which should have been observed in their common ancestor [START_REF] Luo | Acrotiarini trib. nov., in the Cixiidae (Insecta, Hemiptera, Fulgoromorpha) from mid-Cretaceous amber of northern Myanmar, with new insights in the classification of the family[END_REF]. Testing further this paraphyly of Delphacidae is now necessary and an even more representative sampling of Protodelphacida is strongly needed: only Asiracini Motschulsky, 1863, Idiosystatini Emeljanov, 1995, Ugyopini Fennah, 1979 were involved in our analysis, but Eodelphacini Emeljanov, 1995, Platysystatini Emeljanov, 1995, Tetrasteirini Emeljanov, 1995 should be included in future analyses. Very unfortunately we had to discard Neopunanini Emeljanov, 1995, the only available sequences (Urban et al., 2011) appearing to be corrupted. According to our results, relationships between Ugyopinae Fennah, 1969 and Asiracinae Motschulsky, 1863 suggested by [START_REF] Emeljanov | On the question of the classification and phylogeny of the Delphacidae (Homoptera, Cicadina), with reference to larval characters[END_REF] is not supported, the former being included in the later. Paraphyletic Asiracinae, as suggested by [START_REF] Emeljanov | On the question of the classification and phylogeny of the Delphacidae (Homoptera, Cicadina), with reference to larval characters[END_REF] including all other Delphacidae or as sister clade to all other Delphacidae by [START_REF] Asche | Zur Phylogenie der Delphacidae Leach, 1815 (Homoptera Cicadina Fulgoromorpha)[END_REF][START_REF] Asche | Vizcayinae, a new subfamily of Delphacidae with revision of Vizcaya Muir (Homoptera: Fulgoroidea), a significant phylogenetic link[END_REF] is also not supported.

Systematics of Cixiidae

The three recently identified monophyletic lineages [START_REF] Luo | Acrotiarini trib. nov., in the Cixiidae (Insecta, Hemiptera, Fulgoromorpha) from mid-Cretaceous amber of northern Myanmar, with new insights in the classification of the family[END_REF] are recovered particularly with the oeclinian lineage including Bothriocerini [START_REF] Muir | On the classification of the Fulgoroidea (Homoptera)[END_REF], as recently published (Le Cesne et al., 2021). Based on only one sequenced taxon, Mnemosysini Emeljanov, 1992 is retrieved as sister to the oeclinian clade, although it was previously placed in the pentastrinian lineage [START_REF] Le Cesne | Coframalaxius bletteryi gen. et sp. nov. from subterranean habitat in Southern France (Hemiptera, Fulgoromorpha, Cixiidae, Oecleini)[END_REF]. Nevertheless, we suggest that it might be an artefact in our study and it needs to be consolidated with additional sequenced taxa of the tribe. Branching of Borysthenini Emeljanov, 1989 as a sister clade to the pentastirinian lineage (already evocated in Luo et al.

(2021)) needs confirmation with a more consistent sampling of the tribe. In the cixiinian lineage, the split of Cixiini Spinola, 1839 into two different independent lineages, separating 'Australian cixiini' (related to Pintaliini Metcalf, 1938) as sister clade Andini Emeljanov, 2002 to the other Cixiini taxa is strongly supported and a tribal recognition of this group should be proposed, although morphological-based evidences are yet to be found. The exact placement of Gelastocephalini Emeljanov, 2000 andEucarpiini Emeljanov, 2000 needs to be confirmed with a larger sampling as well as the placement of the genus Achaemenes Stål, 1866.

Tettigometridae within the Fulgoroidea

Another interesting result of the study is the relatively early branching of Tettigometridae as sister clade to all Fulgoroidea families (except Kinnaridae-Meenoplidae having already diverged). Tettigometridae were traditionally placed at the first branching lineage of planthoppers according to morphological characters [START_REF] Muir | On the classification of the Fulgoroidea (Homoptera)[END_REF][START_REF] Muir | On the classification of the Fulgoroidea (Homoptera). 1930. On the classification of the Fulgoroidea[END_REF][START_REF] Asche | Preliminary thoughts on the phylogeny of Fulgoromorpha (Homoptera: Auchenorrhyncha)[END_REF][START_REF] Emeljanov | An attempt of construction of the phylogenetic tree of the planthoppers (Homoptera, Cicadina)[END_REF][START_REF] Chen | The metatarsi of the Fulgoroidea (Homoptera: Auchenorrhyncha)[END_REF]. Challenged by Bourgoin (summarized and reviewed in [START_REF] Bourgoin | Molecular Phylogeny of Fulgoromorpha (Insecta, Hemiptera, Archaeorrhyncha). The Enigmatic Tettigometridae: Evolutionary Affiliations and Historical Biogeography[END_REF], Tettigometridae were later constantly placed in the 'higher' families group, often closely related to Tropiduchidae and Flatidae [START_REF] Bourgoin | Molecular Phylogeny of Fulgoromorpha (Insecta, Hemiptera, Archaeorrhyncha). The Enigmatic Tettigometridae: Evolutionary Affiliations and Historical Biogeography[END_REF][START_REF] Yeh | A Molecular Phylogeny of Planthoppers (Hemiptera : Fulgoroidea) Inferred from Mitochondrial 16S rDNA Sequences[END_REF][START_REF] Urban | Evolution of the planthoppers (Insecta: hemiptera: Fulgoroidea)[END_REF]Cryan & Urban, 2012;[START_REF] Song | A preliminary molecular phylogeny of planthoppers (Hemiptera: Fulgoroidea) based on nuclear and mitochondrial DNA sequences[END_REF][START_REF] Huang | Sequencing and analysis of the complete mitochondrial genome of Changeondelphax velitchkovskyi (Hemiptera: Fulgoroidea)[END_REF][START_REF] Yu | The complete mitochondrial genome of Ugyops sp. (Hemiptera: Delphacidae)[END_REF][START_REF] Kim | Complete Mitochondrial Genomes of Metcalfa pruinosa and Salurnis marginella (Hemiptera: Flatidae) : Genomic Comparison and Phylogenetic Inference in Fulgoroidea[END_REF]. The new position of Tettigometridae within Fulgoroidea might be due to our larger sampling, more representative of the diversity of the group than in previous studies in which the family was often summarized to just one or two strongly related species. However, the Tettigometridae initial long branch also suggests a possible long branch attraction effect. According to the recent revision of the higher classification of the family (Mozzafarian et al., 2019), still some key tettigometrid lineages are missing for an optimal representation in a molecular analysis (particularly Asian Egropinae Baker, 1924 and Afrotropical Plesiometrini Bourgoin 2018) to confirm this new placement. While Meenoplidae-Kinnaridae and Tettigometridae are undoubtedly monophyletic groups with their own strong morphological apomorphies, it is interesting to note that both share fully reduced female genitalia, although in strongly different abdominal terminalia conformations.

The clade Derbidae-Achilidae

Already proposed [START_REF] Bourgoin | Female genitalia in Hemiptera Fulgoromorpha, morphological and phylogenetic data[END_REF][START_REF] Urban | Evolution of the planthoppers (Insecta: hemiptera: Fulgoroidea)[END_REF], the Achilidae+Derbidae clade (Clade 11;Fig. C.3,UFBS=63) remains relatively weak in our analysis, despite strong support for the two families. It likely reflects the breadth of other phylogenetic results, which often find them successively placed separately [START_REF] Asche | Preliminary thoughts on the phylogeny of Fulgoromorpha (Homoptera: Auchenorrhyncha)[END_REF][START_REF] Emeljanov | An attempt of construction of the phylogenetic tree of the planthoppers (Homoptera, Cicadina)[END_REF][START_REF] Chen | The metatarsi of the Fulgoroidea (Homoptera: Auchenorrhyncha)[END_REF][START_REF] Yeh | A Molecular Phylogeny of Planthoppers (Hemiptera : Fulgoroidea) Inferred from Mitochondrial 16S rDNA Sequences[END_REF][START_REF] Kim | Complete Mitochondrial Genomes of Metcalfa pruinosa and Salurnis marginella (Hemiptera: Flatidae) : Genomic Comparison and Phylogenetic Inference in Fulgoroidea[END_REF], often with Derbidae (sometimes including some achilid taxa, the latter appearing paraphyletic [START_REF] Urban | Evolution of the planthoppers (Insecta: hemiptera: Fulgoroidea)[END_REF] as sister clade to Achilidae [START_REF] Emeljanov | An attempt of construction of the phylogenetic tree of the planthoppers (Homoptera, Cicadina)[END_REF][START_REF] Huang | Sequencing and analysis of the complete mitochondrial genome of Changeondelphax velitchkovskyi (Hemiptera: Fulgoroidea)[END_REF]. The already mentioned issue of a too short sampling probably explain that, also for these two families. Unfortunately, Derbidae remain difficult to place, probably due to their higher rate of divergence within this family as shown by its particular long branches. While wider than in any other molecular analysis, our own sampling remains far from optimal for these families and their precise relationships as well as their internal ordering of tribes still need to be investigated more carefully. The genus Bebaiotes Muir, 1924 is strongly nested within Achilidae, definitively excluding its placement within Cixiidae [START_REF] Liang | Do the Achilixiidae belong to the Cixiidae?-evidence from antennal sensilla (Hemiptera: Fulgoromorpha: Fulgoroidea)[END_REF][START_REF] Liang | Morphology of antennal sensilla in Achilixius sandakanensis Muir (Hemiptera: Fulgoromorpha: Achilixiidae) with comments on the phylogenetic position of the Achilixiidae[END_REF]. Achilixiidae currently contain two subfamilies: Achilixiinae [START_REF] Muir | On the classification of the Fulgoroidea (Homoptera)[END_REF]Bebaiotinae Emeljanov, 1991 both classified within Achilidae [START_REF] Emeljanov | K voprosu ob obeme i podrazdeleniyakh sem. Achlidae (Homoptera, Cicadina)[END_REF] or in a separate family Achilixiidae [START_REF] Urban | Evolution of the planthoppers (Insecta: hemiptera: Fulgoroidea)[END_REF], Bartlett et al., 2018). Exact positioning of these subfamilies related to Achilidae as a monophyletic group or not, still need to be solved with a wider sampling analysis of the clade (Achilidae+Derbidae) including representatives of genus Achilixius Muir, 1923.

Phylogeny and classification of Dictyopharidae + Fulgoridae

Clade 14 (Fig. C.3, UFBS=100) fully supports the monophyly of this taxonomic unit, as all previously published results. However, the division between the two families remains uncertain due to the uncertain placement of Zanninae Metcalf, 1938 (either retrieved within Fulgoridae sister group of the remaining taxa or included in the Dictyopharidae as sister clade to the other taxa). This makes the monophyly of the two families weakly supported with UFBS= 93 and 50 for the Dictyopharidae and the Fulgoridae, respectively. Although originating mainly from [START_REF] Urban | Entomologically famous, evolutionarily unexplored : The first phylogeny of the lanternfly family Fulgoridae (Insecta: Hemiptera: Fulgoroidea)[END_REF] data, our resulting topology for this clade differs markedly from those proposed by these authors not only in suggesting paraphyletic Fulgoridae in two modalities:

(Zanninae + (Dictyopharidae + other Fulgoridae)) under ML analysis or ((Zanninae + Dictyopharidae) + Fulgoridae) under Bayesian analysis, but also in the distribution of genera within the Fulgoridae. Despite this, we confirm that the current classification of Fulgoridae in subfamilies and tribes largely recognizes non-monophyletic taxa [START_REF] Urban | Entomologically famous, evolutionarily unexplored : The first phylogeny of the lanternfly family Fulgoridae (Insecta: Hemiptera: Fulgoroidea)[END_REF], and that a revised classification is strongly needed for this emblematic family. Of course, alleviating the geographical bias in the sampling of data concerning the almost exclusively Neotropical Fulgoridae is obvious, and Afrotropical, Australian and especially Asian taxa would have to be added to the dataset.

For Dictyopharidae, our dataset resulted in a more structured result consistent with the tribal units already recognized, but differently arranged compared to the topology obtained in the morphological analysis by [START_REF] Song | Morphological phylogeny of Dictyopharidae (Hemiptera: Fulgoromorpha)[END_REF]. In our analysis, Orgeriinae represent an early branching position (considered as sister-clade to all other Dictyopharidae) in line with their recognition as a distinct subfamily, versus in the study by [START_REF] Song | Morphological phylogeny of Dictyopharidae (Hemiptera: Fulgoromorpha)[END_REF] in a group of taxa which was probably artificial, bringing together genera with reduced and leathery wings (Scoloptini, Fernandea sp., Phylloscelini, Ranconini, Capenini, Orgeriini). To resolve both this uncertainty in the delineation of the respective families and their inner classification, a larger sampling is again necessary. It needs focusing on the hinge taxa between the two families distinguished by [START_REF] Emeljanov | Problema razgranichenhiya semeïstv Fulgoridae i Dictyopharidae (Homoptera, Auchenorrhyncha)[END_REF] with representatives of Dichopterinae Melichar, 1912, Lyncidinae Schmidt, 1915, but also on the early branching groups recognized by [START_REF] Song | Morphological phylogeny of Dictyopharidae (Hemiptera: Fulgoromorpha)[END_REF] in their morphological phylogeny (e.g. Aluntiini [START_REF] Emeljanov | Problema razgranichenhiya semeïstv Fulgoridae i Dictyopharidae (Homoptera, Auchenorrhyncha)[END_REF]Cleotychini Emeljanov, 1997, Arjunini Song & Szwedo, 2016).

Tropiduchidae

With our available sampling, the family Tropiduchidae appears as a moderately supported clade sister to the rest of the 'higher' Fulgoroidea families ( tribes (four of them fossils) are described within Tropiduchidae [START_REF] Stroiński | Laberiini, a new tribe of Tropiduchidae planthoppers from Madagascar (Hemiptera: Fulgoroidea)[END_REF], which are represented by only 10 taxa in the present dataset: how relevant is the current Tropiduchidae [START_REF] Fennah | A tribal classification of the Tropiduchidae (Homoptera: Fulgoroidea), with the description of a new species on tea in Malaysia[END_REF], Gnezdilov, 2013) classification remains to be assessed. Approaching the family with only one species and mitogenomic data only probably explains its unusual branching just after Meenoplidae in [START_REF] Michalik | Variable organization of symbiont-containing tissue across planthoppers hosting different heritable endosymbionts[END_REF]. This should be regarded as artefactual.

'Higher' planthopper families

This well-supported node and generally well found in previous studies [START_REF] Urban | Evolution of the planthoppers (Insecta: hemiptera: Fulgoroidea)[END_REF][START_REF] Yeh | A Molecular Phylogeny of Planthoppers (Hemiptera : Fulgoroidea) Inferred from Mitochondrial 16S rDNA Sequences[END_REF][START_REF] Huang | Sequencing and analysis of the complete mitochondrial genome of Changeondelphax velitchkovskyi (Hemiptera: Fulgoroidea)[END_REF][START_REF] Kim | Complete Mitochondrial Genomes of Metcalfa pruinosa and Salurnis marginella (Hemiptera: Flatidae) : Genomic Comparison and Phylogenetic Inference in Fulgoroidea[END_REF] includes eight families including nearly 4,100 species for nearly 30% of planthopper diversity. However, all the groupings between families found below this node appear weakly supported and cannot be reasonably discussed further. Having already shown the significant impact of sampling throughout this work, we consider it very likely that a sample that is more representative of the diversity of these families in the future will have significant effects on the grouping of taxonomic units observed. A good example of this instability is the Nogodinidae issue, regularly found polyphyletic and in different ways, in molecular analyses [START_REF] Yeh | A Molecular Phylogeny of Planthoppers (Hemiptera : Fulgoroidea) Inferred from Mitochondrial 16S rDNA Sequences[END_REF][START_REF] Urban | Evolution of the planthoppers (Insecta: hemiptera: Fulgoroidea)[END_REF][START_REF] Song | A preliminary molecular phylogeny of planthoppers (Hemiptera: Fulgoroidea) based on nuclear and mitochondrial DNA sequences[END_REF][START_REF] Bucher | Which biases still hamper a solid planthopper phylogeny?[END_REF]this work). It is clear that the Nogodinidae do not form a monophyletic group and that the family cannot be maintained: its non-related subunits belong to several of these related families whose diagnoses will have to be reviewed in order to accommodate these taxonomic subunits of the Nogodinidae.

What will be the impact of the huge family Flatidae on the taxonomic arrangement of all these families when it will be better sampled? Except for polyphyletic Nogodinidae, all these families seem to be good monophyletic groups, although the monophyly of Issidae remains questionable. The "issidoid group" of families proposed by Gnezdilov, 2013 (grouping Issidae, Caliscelidae, Acanaloniidae, Tropiduchidae and Nogodinidae) is not supported. Here also it is too early to discuss further how these families should be classify because of the reduced sampling of our dataset regarding the diversity of these taxa.

Issidae phylogeny and classification

Issidae molecular phylogeny was first addressed by [START_REF] Wang | Planthopper family Issidae (Insecta: Hemiptera: Fulgoromorpha): linking molecular phylogeny with classification[END_REF], dividing the family into Thioninae, Issinae -including Issini and Hysteropterini which upgraded later to subfamily [START_REF] Bourgoin | The oldest fossil of the family Issidae (Hemiptera, Fulgoromorpha) from the Paleocene of Menat (France)[END_REF]) -and Hemisphaerinae. Gneznilov et al. (2020) downgraded Thioninae to subtribe Thionina sister to Issina, both upgraded to tribe later [START_REF] Gnezdilov | From modern to classic: Classification of the planthopper family Issidae (Hemiptera, Auchenorrhyncha, Fulgoroidea) derived from a total-evidence phylogeny[END_REF]. In ). These results address the origin of Nearctic Issidae as paraphyletic group, which draw totally different diversification events: an older vicariance event splitting the family into two major lineages, and one to several more recent New World dispersals. As for the other planthopper families, enrichment of the Issidae dataset with new sequenced taxa since [START_REF] Wang | Planthopper family Issidae (Insecta: Hemiptera: Fulgoromorpha): linking molecular phylogeny with classification[END_REF] has progressively modified the topology the tree for this family. However, as several Neotropical taxa from early branching lineages of the Issidae are still missing from all these analyses (which also suffer from the same bias as [START_REF] Gnezdilov | From modern to classic: Classification of the planthopper family Issidae (Hemiptera, Auchenorrhyncha, Fulgoroidea) derived from a total-evidence phylogeny[END_REF]'s study of insufficient representation of neighboring taxa with Flatidae, Nogodinidae in particular), all this generates taxonomic instability at the major node of the family, and we should expect further changes in the classification of Issidae in the future. Genus Picumna Stål, 1864 was not included in our analysis because of the absence of gene in common for both taxa available on GenBank and on the too short sequences. In [START_REF] Gnezdilov | From modern to classic: Classification of the planthopper family Issidae (Hemiptera, Auchenorrhyncha, Fulgoroidea) derived from a total-evidence phylogeny[END_REF], this Nearctic genus was also basal to all Issidae and he even suggested that it might represent a tribal rank. Although not discussed in [START_REF] Gnezdilov | From modern to classic: Classification of the planthopper family Issidae (Hemiptera, Auchenorrhyncha, Fulgoroidea) derived from a total-evidence phylogeny[END_REF], all recent results based on a wider Nearctic sampling agree with an early split of a Nearctic lineage at this stage, fully compatible with a Gondwanan origin of the family as suggested in [START_REF] Bourgoin | Issidae molecular K-libration[END_REF], an age of the lineage also recovered here.

Timing of diversification in extant Fulgoromorpha

Our dating analysis suggests that the diversification of extant Fulgoromorpha occurred in the Early Triassic (median age = 238.06 Mya) and that their origin within Hemiptera likely occurred in the Permian [START_REF] Misof | Phylogenomics resolves the timing and pattern of insect evolution[END_REF]. We found that the diversification of Delphacoidea and Fulgoroidea sec. Bourgoin & Szwedo, 2022 took place in the Middle Triassic, 213.28 Mya and 227.15 Mya, respectively. All major lineages of extant Fulgoromorpha emerged and were already present by the end of the Jurassic. Nevertheless, the 'higher' families clade diverged during the Early Cretaceous with a rapid diversification during the middle to Late Cretaceous, a period that witnessed the Angiosperm Radiation [START_REF] Benton | The Angiosperm Terrestrial Revolution and the origins of modern biodiversity[END_REF]. According to our results, all planthopper families where already present around 125 Mya. Without excluding dispersal events, it is therefore likely that all families have been driven in their distribution and evolution in subsequent subfamiliales linages by the geographical constraints of the Gondwanan break-up. Evolutionary history of all extant planthopper families might have therefore been shaped by early vicariance events during the middle to Late Cretaceous period. Such a scenario was already suggested in a first molecular dating analysis for the Issidae that was reported from the Cretaceous between 116.67 and 131.58 Mya according to different probability densities of priors on node ages [START_REF] Bourgoin | Issidae molecular K-libration[END_REF]. These previous results are confirmed here, versus Gnezdilov (2022)'s statement considering Issidae as a younger lineage of Oriental origin (Eocene, Gnezdilov, 2013b), already challenged in this 2022 analysis by the sister position of the Nearctic genus Picumna to all other Issidae. Interestingly, all internal branches of Fulgoridae are very short, suggesting a rapid diversification processes. It appears that an explosive evolutionary radiation took place for the family around 135-125 Mya (Supplementary Fig. S1). Such a rapid radiation can probably explain the difficulties in establishing a reliable classification of the family and in reaching a clear delimitation between Dictyopharidae and Fulgoridae.

A similar rapid evolutionary radiation event seems to have occurred concurrently in their sister group of the 'higher' planthopper families. For instance, it is clear in the sister group of the Tropiduchidae, where the successive short-branched basal nodes gave rise to nearly a third of the current diversity of the Fulgoromorpha, albeit with various evolutionary successes according to these higher families. At this stage, Gondwana was already separated from Laurasia, which is congruent with the current distribution of all these families still mainly located in the tropical belt. Diversification through early vicariance events, linked with the break-up of Gondwana into smaller continents and leading to the emergence of subsequent lineages during the Late Cretaceous, is a hypothesis that should be tested in the future once phylogenies of these families will be better advanced.

Limitations of the study

Using all relevant molecular data combined into a single phylogenetic analysis for the first time, our study highlights the fundamental impact of sampling on the resulting tree topologies. If the question "More characters or more taxa for a robust phylogeny?" has been addressed many times (reviewed in [START_REF] Nabhan | The impact of taxon sampling on phylogenetic inference: a review of two decades of controversy[END_REF], our study provides another example of the impact of adding many taxa, all carrying their evolutionary history, on the resolution and robustness of the nodes, especially at the backbone level of the tree. Increasing the taxon sampling (here defined a priori based on the taxonomic diversity currently recognized in the group) likely increases the opportunity of including key taxa that diverged early, which makes it possible to reconstruct a more consistent evolutionary history at deep nodes. This issue of homogeneous representation of diversity was nevertheless a limitation for outgroup sampling. If we tried to represent the maximum number of lineages within Hemiptera for the phylogenetic analyses, the under sampling of taxa within these lineages led us to very high divergence rates between our ingroup and outgroup taxa, which did not allow us to reach convergence for the divergence time estimates. Having identified contaminated, corrupted molecular sequences used in previous phylogenetic studies, we also underline the crucial importance of their good quality systematically rechecked by a blast and ideally confirmed with good congruence of molecular sequences for related species.

The importance of the quality of a fossil dataset for the calibration of internal nodes has also been highlighted. In this study, a revision of the morphological characters justifying the relatedness of a fossil taxon to an extant lineage led us to modify the sampling of the taxa commonly used in the dating analyses of Fulgoromorpha, as well as their stratigraphic age that are updated. Nonetheless, it is important to note that none of the fossils used here have been included in a phylogenetic analysis and therefore the character sets are not stated to be taken as phylogenetically justified. It is thus quite likely that the phylogenetic placement of these fossils will be rearranged because of a future analysis grouping together extant and fossil taxa. We have no doubt that the subsequent results that would bring together the paleontological and neontological data will challenge their use as calibration points for node-dating analyses, which would also have a consequent impact on divergence times and diversification for Fulgoromorpha.

Conclusions

This study proposes a new phylogenetic framework and timeline for Fulgoromorpha. Although challenged by studies biased by too small sampling to infer the planthopper phylogeny, we generally confirm the results of [START_REF] Urban | Evolution of the planthoppers (Insecta: hemiptera: Fulgoroidea)[END_REF], with Delphacoidea including Cixiidae and Delphacidae being sister to Fulgoroidea (sec. Bourgoin & Szwedo, 2022). However, some unexpected results also emerge: (1) the possible paraphyly of Delphacidae whose Protodelphacida seems more related to Cixiidae than to other delphacids; (2) the Tettigometridae moving out from the group of the 'higher' families to be considered as sister clade to all others Fulgoroidea families (except for Meenoplidae-Kinnaridae); and ( 3 Reconstructed families are color coded (Delphacidae: blue, Cixiidae: green, Issidae: purple, others families: brown shading). This color code is reused in the complete tree (Fig. 3) and the dated phylogeny (Fig. 4). Color code is reused in the simplified phylogeny (Fig. 2) and complete tree (Fig. 3). The red circles correspond to the fossil calibration of the tree. The complete tree with confidence intervals associated with each node age is presented in supplementary data. (Nel et al., 1999). Date of this deposit was re-evaluated as equivalent to Clarkforkian by Smith et al. (2014).

Acixiites immodesta

Hamilton, 1990

AMNH 43679 Compression Crato Fm, Ceará, Brazil

Stem Achilidae

For Hamilton (1990) the flattened body of the described specimen, the overlapping tegmina and the "spurious" claval suture allow us to assign this fossil to the Achilidae without the possibility of assignment to a modern lineage. This specimen doesn't possess the veinlet connecting the posterior branch of Cua with the tegmen margin, which is very characteristic to Achilidae representents (Martill et al., 2007). Nevertheless, the vertex, the pronotom and the lack of subapical setae on the hind tarsomere could related this species to more basal Derbidae. Nevertheless, the long rostrum extended beyond the apex of the hind femora excludes this species from this group.

-113

The age of the Nova Olinda Member of the Crato Formation (Araripe Group) has been interpreted as Aptian (Maisey, 1990;Martill, 2007;Staniczek et al., 2022Staniczek et al., , paleobiodb.org, 2022)).

Acriotiara multigranulata

Luo & Bourgoin, 2021 HO221990 Amber inclusion Hukawng Valley, Kachin State, Manmar

Cixiidae

Stem Pentastirinian lineage

This fossil was recently described and all assigned to the new tribe Acrotiarini [START_REF] Luo | Acrotiarini trib. nov., in the Cixiidae (Insecta, Hemiptera, Fulgoromorpha) from mid-Cretaceous amber of northern Myanmar, with new insights in the classification of the family[END_REF]. This taxon is excluded from the Oecleinian lineage by the absence of a metatibial diastema and share with the Cixiinian + Pentasterinian (Pentastirini + Mnemosynini) lineages the distal displacement of the icu vein distal to the apex of clavus. The presence of a pentacarinated mesonotum in the Acrotiarini species allows to place them within the pentastirinian lineage based on this only apomorphic character but as a separate lineage by the unique presence of the paired sublaterofrontal carinae and the arched RA venation on tegmen.

99.41 -98.17

The Hukawng Valleyamber deposit has been radiometrically dated using zircons to 98.79 ± 0.62 Ma, in earliest Cenomanian (Shi et al., 2012). 98.17 Ma should be taken as a conservative minimum age of the deposit (Kania et al., 2015;Mao et al., 2018).

Barremixius petrinus (Fennah, 1961)

BMNH In. 59335 Compression

Lamb's Brickworks, locality 170/348485,

Cixiidae

Stem Cixiinian lineage Fennah first assigned B. petrinus 'with confidence' to the Cixiina subtribe and uncertainly to the genus Cixius Latreille, 1804. Inded, the fossil lacks all apomorphies characters of both oecleidian and pentastiridian lineages.

-125

Deposits from Lamb's Brickworks, Weald Clay Formation (Upper Members) are dated from Barremian (paleobiodb.org, 2022).

  placed the divergence of Delphacoidea+Fulgoroidea in the Late Permian, contradicting Johnson et al. (2018) who propose a younger diversification of the two lineages, dated from the Late Triassic. Both studies nevertheless propose a relatively congruent age of origination, placing the divergence Fulgoromorpha from other Hemiptera during the Carboniferous (mid-Carboniferous for Song et al. (2013), Late Carboniferous for Johnson et al. (2018)), which is in line with the recent addition of Aviorrhynchidae to

  For the full 18S, 28S D3D5 and 28S D6D10 the following parameters were used: algorithm: E-INS-I; Scoring-matrix: 200 PAM / k=2; Gap open penalty: 1.5; COI, Cytb and Wg sequences were aligned as follows: Algorithm: G-INS-I; Scoring-matrix: 200 PAM / k=2; Gap open penalty: 1.5. Because of their length polymorphism, initial and terminal regions of the non-

  Fig. B. The complete phylogeny (Fig. C) is presented here in 4 parts (1-4) in which main clades are numbered from 1 to 26. Our study recovers extant Fulgoromorpha (UFBS=100) as monophyletic and strong UFBS is found across deep nodes (i.e. inter-familial relationships). Clade 1 (Fig. C.1, UFBS=100) supports families Cixiidae and Delphacidae as a clade sister to the remaining planthopper families, corresponding to the recently recognized superfamily Delphacoidea. Our wider taxonomic sampling for both families (122 and 126 taxa

  Clade 4 (Fig. C.2, UFBS=71) represents Cixiidae and comprises three previously identified monophyletic lineages (Luo et al., 2021): the oeclinian lineage (Fig. C.2, Clade 5, UFBS=92) with Oeclini including Bothriocerini, sister to the pentastrinian lineage including Borysthenini sister to Pentastrini with low support (Fig. C.2, Clade 6, UFBS=22), and the cixiinian lineage (Fig. C.2, Clade 7, UFBS=93) with all other sampled tribes of our study according to the following sequence: (Gelastocephalini + ((Pintalini + Australian Cixiini) + (Andini + (Eucarpini + (Bennini + other Cixiini including Semonini))))). Mnemosysini, previously placed in the pentastrinian lineage (Luo et al., 2021), is here moved at the basal node of the oeclinian lineage. Clade 9 (UFBS=99), corresponding to all Fulgoroidea families (sec. Bourgoin & Szwedo, 2022), separates basally into clade 8 (Fig. C.3, UFBS=100) Meenoplidae included in Kinnaridae sister to all other extant Fulgoroidea families in clade 10 (UFBS=100). This strong clade 10 separates the Tettigometridae (Fig. C.3, Clade 10, UFBS=100) sister to the remaining planthopper families. Clade 11 (Fig. C.3, UFBS=63) is weakly supported, although it suggests a sister relationship between the strongly supported monophyletic units of Derbidae (Clade 12) and Achilidae including Achilixidae (Clade 13) (UFBS=100 and 95, respectively). In Derbidae, taxa show longer internal branches suggesting a higher rate of divergence within this family. Following Clade 11, the strongly supported Clade 14 (Fig. C.3, UFBS=100) separates sister families Dictyopharidae and Fulgoridae from the remaining planthoppers. If Dictyopharidae monophyly is well supported (Fig. C.3, Clade 15, UFBS=93), monophyly of Fulgoridae is not (Clade 16, UFBS = 50). This weakness is potentially due to the Zanna sp.

  Fig. C.4, UFBS = 98), monophyly of most families is recovered with strong support value for Acanaloniidae (Clade 22, UFBS=100), Caliscelidae (Clade 23, UFBS=100), Flatidae (Clade 24, UFBS=99), Ricaniidae (Clade 25, UFBS=100). Surprisingly, monophyly of Lophopidae was not recovered due to the genus Serida Walker, 1857 moved within Eurybrachidae, while these families and their sister relationships (Clade 20, UFBS=98) remain strongly supported monophyletic units. Issidae (Clade 26, UFBS=74) is recovered as monophyletic, but with a low basal bootstrap support. Within this family, all tribal units reviewed by Wang et al. (2016) also recovered as monophyletic (Hysteropterini, Kodaianellini, Issini, Sarimini, Parahiraciini, Hemisphaeriini), with the exception of Thioniini and

  Convergence of dating analysis was ensured by ESS values and plots indicated in Tracer (The .log file resulting from the analysis is appended in supplementary data). Results of analysis are presented as a simplified chronogram in Fig.Dwhere only major clades are presented. The complete tree, with node age and 95% CI, is presented in Supplementary Data as a tree file.The dating analysis suggests that the diversification of extant Fulgoromorpha occurred in the Early Triassic with a median age of 238.06 Mya (95% CI: 199.6-352.9 Mya). Divergence of Fulgoromorpha with the rest of Hemiptera was not estimated because outgroups were excluded from the analysis to solve convergence issues in dating analyses. The age of the early lineages of Delphacoidea, with the Cixiidae and Delphacidae, is estimated in the Late Triassic at 213.28 Mya (95% CI: 131.16-262.25 Mya), while all other families in Fulgoroidea sec.Bourgoin & Szwedo, 2022 would date back to the Middle Triassic at 227.15 Mya (95% CI: 180.21-344.97

  recovered a similar paraphyletic Cixiidae during an analysis based on four delphacid species for 55 cixiid ones. Yet, Kim et al. (2021), in a strongly disequilibrated sampling (11 and 1 species for Delphacidae and Cixiidae, respectively) produced either two monophyletic sister families, or refuted the Delphacoidea clade with Cixiidae sister clade to all planthoppers, or a paraphyletic Delphacidae: their unique cixiid taxon Pentastiridius grouping with their single protodelphacid taxon Ugyops, both sister to the rest of delphacids. This last unconventional pattern was already proposed by Bucher & Bourgoin (2019) based on a wider sampling of respectively 16 and 19 delphacid and cixiid species. Our study based on a much wider taxonomic sampling for both families (122 and 126 taxa for Delphacidae and Cixiidae, respectively) seems to confirm paraphyly of Delphacidae, splitting into Eudelphacida sec. Asche 1990 and monophyletic Protodelphacida sec. Emeljanov, 1996, sister group to Cixiidae on the other side (Fig. C.1, Clade 2, UFBS=80

  Fig. C.4, Clade 19, UFBS=98). However, their sister relationship remains questionable (Fig. C.4, Clade 17, UFBS=59). Currently 25

Gnezdilov

  et al. (2022), Hemisphaerinae sec. Wang et al., 2016 remains a valid taxon in a new trichotomic node Issinae sec. Gnezdilov et al., 2022 with Issini and Thioniini, both sister to Hysteropterinae (op. cit. Fig. 3, 50% majority-rule consensus tree). A result rather similar is depicted in our new analysis but Kodaianellini moved as sister clade to a paraphyletic Thioniini group, sister to Issini. 2 Thioniini taxa are found as sister clade to all other Issidae (Fig. C.4, Clade 26

  ) the Tropiduchidae placed in a sister group of the other 'higher' families including the Lophopidae-Eurybrachidae. Addressing more precisely the other issues listed in the introduction, the couples of previously paired families are well recovered with: (1) the Meenoplidae-Kinnaridae with the paraphyletic Kinnaridae; (2) the Achilidae-Derbidae clade with the Achilidae Plectoderini including the Achilixiidae (represented by the genus Bebaiotes); (3) the Fulgoridae-Dictyopharidae clades, with Zanninae retrieved as the early branching lineage of the Fulgoridae in a weakly supported node; and (4) the Lophopidae-Eurybrachidae. Four families are now documented enough to start looking to their main internal phylogenetical structuration. In Delphacidae Eudelphacida, all major subfamilies/tribes are fully recovered with a sister relationship between Kelisiinae/i and Stenocraninae/i. Protodelphacida is a separate sister to Cixiidae in a monophytetic clade including Ugyopini, Asiracini and Idiosystatini grouped in a sister taxon. In Cixiidae, three main clades are recovered corresponding to the oeclinian, pentastirianian and cixiinian lineages suggested by Luo et al. (2021) but with Mnemosynini placed sister to the oeclenian. In Fulgoridae, the current classification of subfamilies and tribes does not recover at all the phylogeny. All tribal groups are recovered for Issidae excepted Thioniini splitting into a group sister to all other issids and another sister to Issini, both sister to Kodaianellini. Fossil-calibrated molecular dating of current families estimates the first diversification of planthoppers in Early Triassic around 240 Mya and those of the superfamilies Delphacoidea and Fulgoroidea around 220 Mya. By the end of the Jurassic, all major planthoppers lineageswere originated and all families diversified around 125 Mya. In some parts of the tree topology, the dating analyses point to rapid evolutionary radiations, which occurred particularly in Fulgoridae around 125-130 Mya. However, both our taxon and character samplings remain under-represented to draw more definitive conclusions. The Tropiduchidae and the Flatidae, rich and disparate groups, seem to be the most critical clades from this point of view, and the addition of new sequences for these families is a priority for testing the present results. The paraphyly found in Delphacidae represents a very interesting point to study if we can complete the dataset with new sequences of Protodelphacida; the same applies to the Tettigometridae and the Achilixiidae whose new phylogenetic position must be confirmed. In other words, more reliable molecular sequences are critically needed since many nodes remain unstable at the mercy of adding a new taxon in the analysis.

Fig. 1 :

 1 Fig.1:A. Hypotheses on the origin, diversification dating and phylogenetic relationship of extant Fulgoromorpha lineages. Phylogeny is based on synthesis of few molecular studies and is summarized in[START_REF] Bartlett | The Diversity of the True Hoppers (Hemiptera : Auchenorrhyncha)[END_REF]. Branch lengths are not to scale. B. Live representatives of Fulgoromorpha a. Megadelphax sordidula (Stål, 1853) (Delphacidae) b. Reptalus quinquecostatus (Dufour, 1833) (Cixiidae) c. Nisia sp. (Meenoplidae) d. Anotia sp. (Derbidae) e. Plectoderes scapularis Metcalf, 1938 (Achilidae) f. Dictyophara europaea (Linné, 1767) (Dictyopharidae) g. Phrictus quinquepartitus Distant, 1883 (Fulgoridae) h. Paracorethrura iocnemis (Jacobi, 1905) (Lophopidae) i. Trienopa typica (Distant, 1911) (Tropiduchidae) j. Tettigometra laeta Herrich-Schäffer, 1835 (Tettigometridae) k. Acanalonia conica (Say, 1900) (Acanaloniidae) l. Ommatidiotus dissimilis (Fallén, 1806) (Caliscelidae) m. Nephesini sp. (Flatidae) n. Biolleyana costalis (Fowler, 1900) (Nogodinidae) o. Ricania speculum (Walker, 1851) (Ricaniidae) p. Hemisphaerius stali Banks, 1910 (Issidae). All photographies belong to © Gernot Kunz.

Fig. 2 :

 2 Fig. 2: Summary phylogeny of extant Fulgoromorpha with major clades (families) collapsed.Reconstructed families are color coded (Delphacidae: blue, Cixiidae: green, Issidae: purple, others families: brown shading). This color code is reused in the complete tree (Fig.3) and the dated phylogeny (Fig.4).

Fig. 3 :Figure 4 :

 34 Fig. 3: Complete phylogeny of extant Fulgoromorpha reconstructed with Maximum Likelihood inference. Branch support is indicated by ML ultrafast bootstraps (1000 iterations). The tree is presented in four parts (A-D): arrows correspond to the location where the branches have been cut. Reconstructed families are color coded (Delphacidae: blue, Cixiidae: green, Issidae: purple, others families: brown shading) with subfamilies and other lineages indicated. A. Phylogeny of Delphacidae. B. Phylogeny of Cixiidae. C-D. Phylogeny of Fulgoroidea families

Table A : Taxonomic diversity represented by taxa included in the final analysis (in-group).

 A Taxonomic classification follows FLOW[START_REF] Bourgoin | FLOW (Fulgoromorpha Lists on The Web): a world knowledge base dedicated to Fulgoromorpha[END_REF], which is based on the Metcalf (1958) catalog and recent taxonomic revisions. "." denotes a lack of classificatory rank. Not all terminal taxa are identified to genus and specie level. Several specimens may represent the same taxonomic rank.

	Families	Subfamillies sampled	Tribes sampled	Sub-tribe sampled	Total number of terminal taxa a
	Acanaloniidae	.	.	.	6
	Achilidae	2/3	2/11	2/3	20
	Achilixiidae	1/2	.	.	1
	Caliscelidae	1/2	2/5	0/4	7
	Cixiidae	1/1	11/16	2/3	126
	Delphacidae	5/6	6/10	.	122
	Derbidae	3/3	4/20	2/9	12
	Dictyopharidae	2/2	5/17	1/2	17
	Eurybrachidae	2/2	2/6	.	7
	Flatidae	2/2	3/8	2/9	7
	Fulgoridae	8/11	9/19	4/4	67
	Issidae	4/4	6/9	4/9	95
	Kinnaridae	2/2	3/5	.	4
	Lophopidae	2/2	3/6	.	5
	Meenoplidae	1/2	.	.	3
	Nogodinidae	2/4	3/10	2/4	6
	Ricaniidae	.	.	.	10
	Tettigometridae	2/4	1/4	.	5
	Tropiduchidae	2/2	7/20	2/7	11

a

Table B : Fossils used in the analyses for estimating divergence dates and calibration point prior settings.

 B Fossil selection followed the best practices suggested by[START_REF] Parham | Best Practices for Justifying Fossil Calibrations[END_REF] to justify fossil calibrations. As none of the present fossils were included in a phylogenetic analysis, attribution to a phylogenetic lineage was carried out according to the apomorphy-based method. All dates based on stratigraphic intervals have been updated following the latest version of the International Commission on Stratigraphy 2022/10 and the latest information published. Here we use calibration points in the sense of most representatives of early-branching lineages (stem group) which only give minimum ages. Szwedo (2012), the specimen possesses all combinations of characters forming the venation pattern of Ricaniidae as follows: costal area narrower than costal cell, costal cell with numerous transverse veinlets, basal cell strongly rounded, longitudinal venation wavy, absence of nodal line. A. curvivenatus is not supposed to be related to an extant lineage within our Ricaniidae sampling.Ambers from deposit of Le Quesnoy, Oise River area of the Paris basin, France were aged Lowermost Eocene, Sparnacien 55-53 Ma

		Fossil identity				Systematic assignment		Geological age (Ma)
	Fossil taxon	Museum specimen number	Mode of preservation	Locality	Taxonomic group	References	Age (Mya)	References
	Abraracourcix curvivenatus	MNHN PA5155	Amber	Le Quesnoy,	Stem Ricaniidae	According to Stroiński & 56.8
	Stroinski & Szwedo, 2012		impression	Oise, France			-
							55.4

Acknowledgements

For providing additional sequences, data and discussions, the authors are strongly indebted to Birgit Loecker, Roslyn Mirrington, Maxime Le Cesne, Adeline Soulier-Perkins and Jacek Szwedo. All photographs included in this publication were taken by Gernot Kunz and we thank him for their use. This research was partly funded by National Natural Science Foundation of China (32100369).

South Godstone, Tandridge, Surrey, England, UK [START_REF] Luo | Acrotiarini trib. nov., in the Cixiidae (Insecta, Hemiptera, Fulgoromorpha) from mid-Cretaceous amber of northern Myanmar, with new insights in the classification of the family[END_REF] erected a new genus for this fossil and provisionaly maintained it into Cixiini but in the cixiinian lineage with which it share the usual plesiomorphic set of characters of the lineage.

Bolbossus bervoetsi [START_REF] Gnezdilov | On the taxonomic position of Issus reticulatus Bervoets, 1910 (Hemiptera: Fulgoroidea: Issidae) from Baltic amber[END_REF] According to J. Szwedo (pers. comm.), the location of the type material is unknown. , 1939, (Bervoets, 1910)). Nevertheless, [START_REF] Gnezdilov | On the taxonomic position of Issus reticulatus Bervoets, 1910 (Hemiptera: Fulgoroidea: Issidae) from Baltic amber[END_REF], erected it in the tribe Parahiraciini based on the venation pattern with the very embossed longitudinal veins as well as the particular shape of metope very convex. B. bervoetsi is not assigned to a modern lineage within the tribe.

Amber inclusion

37.8 -33.9

Initially the material from the Pr. R. Kleb's collection was dated from Oligocene by Bervoets (1910). Material was re-examined by Szwedo et al., 2004 and date was reevaluated from Eocene and later from Priabonian according to Gnezdilov & Bourgoin (2016, Szwedo comm.).

Cubicostissus palaeocaeni [START_REF] Bourgoin | The oldest fossil of the family Issidae (Hemiptera, Fulgoromorpha) from the Paleocene of Menat (France)[END_REF] The presence of a calcar spur on hindtibia of the specimen places it without doubt within Delphacidae. Gebicki & Szwedo (2000) proposed its placement within 47.8 -43.9 Baltic amber is a secondary deposit found mainly in glauconitic marine sediments of middle Eocene age (Lutetian Stage: 41.3-47.8 Mya), deposited along deposit: "blue earth" formation)

Ugyopini Fennah, 1979 based on the general shape of the head and the position of carinae on the vertex which suggests that this specimen is closely related to some Pacific genera (Ugyopana, Melanesia, Melanugyops, Livatiella). The absence of characters for justifying its inclusion within the genus Ugyops Guérin-Méneville, 1834 makes us exclude this fossil from the Crown Ugyopini of our sampling.

the paleo-North Sea margin. Problems to date a precise for Baltic amber come from amber deposits being reworked and deposited in sediments that are millions of years younger than the original time of production (Weitschat and Wichard, 2002). Therefore, several ages for of Baltic amber were proposed (Mertz et al., 2000;Drohojowska et al., 2015;Szadziewski, 2018;Sadowski et al., 2022). We retain here the age proposed by Mertz et al., (2000) who consider a middle Eocene age of Baltic amber which comes from correlations with limnic sediments within Eckfeld Maar. They defined 44.3 ± 0.4 Ma as the age of Baltic amber. Accordingly, minimum age of S. perperunae has been choose to 43.9 Ma.