DOI: 10.5061/dryad.pk0p2ngq5
title: Unraveling the evolutionary history of the snakefly family Inocelliidae (Insecta: Raphidioptera) through integrative phylogenetics

description: Inocelliidae is one of the two extant families of the holometabolan order
 Raphidioptera (snakeflies), with the modern fauna represented
 by seven genera and 44 species. The evolutionary history
 of the family is little known. Here we present the first phylogenetic and
 biogeographic analyses based on a worldwide sampling of taxa and datasets
 combined with morphological characters and mitochondrial genomes, aiming
 to investigate the intergeneric phylogeny and historical biogeography of
 Inocelliidae. The phylogenetic inference from the combined analysis of
 morphological and molecular data recovered the sister-group relationship
 between a clade of (Negha + Indianoinocellia) +
 Sininocellia and a clade of Fibla + the
 Inocellia clade (interiorly nested by Amurinocellia and
 Parainocellia). Amurinocellia stat. rev. and
 Parainocellia stat. rev. et emend. nov. are relegated to
 subgeneric status within Inocellia, while a newly erected
 subgenus of Inocellia, Epinocellia subgen.
 nov., accommodates the former Parainocellia
 burmana (U. Aspöck and H. Aspöck, 1968) plus a new species
 Inocellia (Epinocellia) weii sp. nov. Further,
 the Inocellia crassicornis group constitutes the
 nominate subgenus Inocellia stat. nov., but the Inocellia
 fulvostigmata group is paraphyletic. Diversification within
 Inocelliidae is distinguished by an Eocene divergence leading to extant
 genera and a Miocene radiation of species. A biogeographic scenario
 depicts how the diverse inocelliid fauna from East Asia could have
 originated from western North America via dispersal across the Beringia
 during the early Tertiary, and how the Miocene ancestors of
 Inocellia could have accomplished long-distance dispersals via
 the Tibet‐Himalayan corridor or eastern Palaearctic to western
 Palaearctic. Our results shed new light specifically on the evolution of
 Inocelliidae and, in general, the Raphidioptera.
description: Data Matrix
description: File S1. Checklist of extant species of Inocelliidae.
 File S2. Specimens examined for the present study.
 File S3. Nexus file of Maximum parsimony tree based on
 morphological data. File S4. TNT file of Maximum
 parsimony tree based on morphological data. File
 S5. Nexus file of Bayesian inference tree based on dataset PCG123
 (nucleotide data of 13 protein-coding genes). File
 S6. Phylip file of Maximum likelihood tree based on dataset PCG123
 (nucleotide data of 13 protein-coding genes). File S7.
 Fasta file of Maximum parsimony tree based on dataset PCG123 (nucleotide
 data of 13 protein-coding genes). File S8. Nexus file
 of Bayesian inference tree based on dataset PCG123rRNA (nucleotide data of
 the 13 protein-coding genes and rRNA genes). File
 S9. Phylip file of Maximum likelihood tree based on dataset
 PCG123rRNA (nucleotide data of the 13 protein-coding genes and rRNA
 genes). File S10. Fasta file
 of Maximum parsimony tree based on dataset PCG123rRNA (nucleotide data of
 the 13 protein-coding genes and rRNA genes). File S11. Nexus file of Bayesian inference
 tree based on dataset PCG12 (nucleotide data of the 13 protein-coding
 genes without the third position of PCGs) File S12. Phylip file of Maximum likelihood
 tree based on dataset PCG12 (nucleotide data of the 13 protein-coding
 genes without the third position of PCGs) File
 S13. Fasta file of Maximum parsimony tree based on dataset PCG12
 (nucleotide data of the 13 protein-coding genes without the third position
 of PCGs) File S14. Nexus file of Bayesian inference
 tree based on dataset PCG_AA (amino acid sequences of the 13
 protein-coding genes). File S15. Phylip file of Maximum
 likelihood tree based on dataset PCG_AA (amino acid sequences of the 13
 protein-coding genes). File S16. Fasta file of Maximum
 parsimony tree based on dataset PCG_AA (amino acid sequences of the 13
 protein-coding genes). File S17. Nexus file containing
 the molecular and morphological alignments. File
 S18. TNT file containing the molecular and morphological
 alignments. Table S1. List of species sampled in the
 present phylogenetic analysis and GenBank accession number.
 Table S2. The partitioning schemes and selected amino acid
 substitution models for phylogenetic analysis. Table
 S3. Saturation test of the gene PCG and its different codon sites and gene
 rRNA. Table S4. Divergence time estimates of
 Inocelliidae. Nodal code corresponds to that in Fig. 4. Time-scale units
 are in millions of years (Ma). Table S5. Likelihood
 scores and model comparison for ancestral area reconstruction using
 BioGeoBEARS. Nodal code corresponds to that in Fig. 4.
 Table S6. Major biogeographic events of Inocelliidae estimated
 with BioGeoBEARS. Nodal code corresponds to that in Fig. 4.
 Table S7. Summary of ancestral area reconstructions based on
 DIVALIKE + J model. Node for nodes in Fig. 4. Table
 S8. Beast median divergence times for major nodes in Fig. 4, with 95%
 credibility intervals and marginal likelihoods. Fig.
 S1. Images of antennae of inocelliids. (a) Indianoinocellia mayana U.
 Aspöck, H. Aspöck, & H. Rausch; (b) Inocellia elegans Liu, H.
 Aspöck, Yang & U. Aspöck. Fig. S2. Images of
 head and pronotum of snakeflies. (a) Amurinocellia sinensis (Navás), male;
 (b) Inocellia cheni Liu, H. Aspöck, Yang & U. Aspöck, male; (c)
 Inocellia elegans Liu, H. Aspöck, Yang & U. Aspöck, male; (d)
 Negha inflata (Hagen), male; (e) Sininocellia gigantos Yang, male; (f)
 Inocellia elegans Liu, H. Aspöck, Yang & U. Aspöck, female; (g)
 Amurinocellia sinensis (Navás), female; (h) Negha inflata (Hagen), female;
 (i) Agulla arizonica (Banks), female; (j) Sininocellia chikun Liu, H.
 Aspöck, Zhan, & U. Aspöck, female; (k) Fibla pasiphae (H. Aspöck
 & U. Aspöck), female; (l) Inocellia fulvostigmata fulvostigmata U.
 Aspöck & H. Aspöck, female; (m) Inocellia crassicornis (Schummel),
 female. Arrows indicate character state used in the phylogenetic
 analysis of the morphology-only dataset as indicated in the
 text. Fig. S3. Images of wings of snakeflies. Arrows
 indicate character state used in the phylogenetic analysis of the
 morphology-only dataset as indicated in the text. Fig.
 S4. Drawings of male fused gonocoxites 11 (gonarcus) of inocelliids. (a),
 (d), (e) and (g)–(i) modified from H. Aspöck et al. (1991); (b) modified
 from Shen et al. (2019); (f), (k) (l) and (o) modified from Liu et al.
 (2010a); (c) modified from Liu et al. (2010b); (j) modified from Liu
 & Hajong. (2015); (m) modified from Liu et al. (2012a); (n)
 modified from Liu et al. (2009b). Scale bar = 0.5 mm.
 Fig. S5. Drawings of male fused gonocoxites 11 (gonarcus) of
 inocelliids. (a)–(n) modified from H. Aspöck et al. (1991); (o) modified
 from Liu et al. (2012b). Scale bar = 0.5 mm. Fig.
 S6. Drawings of male fused gonocoxites 11 (gonarcus) of inocelliids.
 (e)–(h) modified from U. Aspöck et al. (2011); (b) modified from Liu et
 al. (2010a); (a) (j) (c) and (d) modified from Liu et al. (2010b); (i)
 modified from U. Aspöck et al. (2009); (k) modified from Liu et al.
 (2014a). Scale bar = 0.5 mm. Fig. S7. Maximum
 likelihood tree based on PCG123 dataset.
 Fig. S8. Bayesian inference tree based on
 PCG123 dataset. Fig. S9. Maximum parsimony tree based
 on PCG123 dataset. Fig. S10. Maximum likelihood tree
 based on PCG123rRNA dataset. Fig. S11. Bayesian
 inference tree based on PCG123rRNA dataset. Fig.
 S12. Maximum parsimony tree based on PCG123rRNA dataset.
 Fig. S13. Maximum likelihood tree based on PCG12
 dataset. Fig. S14. Bayesian inference tree based on
 PCG12 dataset. Fig. S15. Maximum parsimony tree based
 on PCG12 dataset. Fig. S16. Bayesian inference tree
 based on PCG_AA dataset. Fig. S17. Maximum parsimony
 tree based on PCG_AA dataset. Fig. S18. Bayesian
 inference tree based on the combined analysis of morphological and
 molecular data. The genera in colored taxa are inocelliids, and the grey
 colored taxa are outgroups in Raphidiidae. Fig.
 S19. Maximum parsimony tree based on the combined analysis of
 morphological and molecular data. The genera in colored taxa are
 inocelliids, and the grey colored taxa are outgroups in
 Raphidiidae. Fig. S20. BEAST divergence times estimates
 for key nodes in inocelliid cladogenesis (node codes correspond to those
 in Fig. 4), under different priors Violin plots presenting mean ages
 (white dots) along with the posterior distribution (blue area) of the 95%
 credibility intervals (black bars) inferred in the different BEAST
 analyses. (a) node Ino4: divergence between Sininocellia and Negha, (b)
 node Ino8: crown Inocelliidae), (c) node Ino9: divergence between
 Fibla and Inocellia clade, (d) node Ino21: crown I. crassicornis group,
 (e) node Ino23: divergence between western Palaearctic and eastern
 Palaearctic I. crassicornis, (f) node Ino24: divergence between
 Parainocellia and Amurinocellia. YU, Yule Tree model; BD, birth-death Tree
 model; Exp, exponential prior distribution for fossil calibrations; LogN;
 lognormal prior distribution for fossil calibrations; Uni, uniform prior
 distribution for fossil calibrations.