ABSTRACT Cave crickets (Rhaphidophoridae) are insects of an ancient and wingless lineage within Orthoptera that are distributed worldwide except in Antarctica, and each subfamily has a high

level of endemicity. Here, we show the comprehensive phylogeny of cave crickets using multi-gene datasets from mitochondrial and nuclear loci, including all extant subfamilies for the first

time. We reveal phylogenetic relationships between subfamilies, including the sister relationship between Anoplophilinae and Gammarotettiginae, based on which we suggest new synapomorphies.

Through biogeographic analyses based on divergence time estimations and ancestral range reconstruction, we propose novel hypotheses regarding the biogeographic history of cave crickets. We

the loss of wings in Rhaphidophoridae could be the result of their adaptation to low temperatures in the Mesozoic era. SIMILAR CONTENT BEING VIEWED BY OTHERS PHYLOGENY OF THE SYNLESTIDAE

(ODONATA: ZYGOPTERA), WITH AN EMPHASIS ON _CHLOROLESTES_ SELYS AND _ECCHLOROLESTES_ BARNARD Article Open access 15 September 2020 DIVERSIFICATION AND POST-GLACIAL RANGE EXPANSION OF GIANT

NORTH AMERICAN CAMEL SPIDERS IN GENUS _EREMOCOSTA_ (SOLIFUGAE: EREMOBATIDAE) Article Open access 11 November 2021 EARLY PLEISTOCENE ORIGIN AND EXTENSIVE INTRA-SPECIES DIVERSITY OF THE

EXTINCT CAVE LION Article Open access 28 July 2020 INTRODUCTION Rhaphidophoridae (Orthoptera: Ensifera), commonly known as cave crickets, cave wētā, land shrimp, sand treaders, jumping, and

camel crickets, are a wingless family consisting of nine extant subfamilies and one extinct subfamily with more than 1,100 described species1. They are considered the earliest diverging

lineage of six families in the infraorder Tettigoniidea, one of two major lineages within Ensifera2,3. These insects are usually found in caves, burrows, cellars, and under logs, preferring

dark and humid environments, and are characterized by their long legs and antennae, lack of wings, and often humped back4,5. Unlike most relatives in Ensifera, Rhaphidophoridae have no

stridulatory and auditory organs for acoustic communication, but some species are known to produce courtship signals by tapping the abdomen or vibrating the body6. Interestingly, some

species have been observed to visit flowers as potential plant pollinators in subarctic islands7. Rhaphidophoridae are widely distributed across all continents except Antarctica, with each

subfamily showing a geographically limited distribution1. For example, Aemodogryllinae and Rhaphidophorinae are distributed in Southeast Asia, with the latter family more widely distributed

to the south, including Oceania (Fig. 1a–e, g). Anoplophilinae are distributed only in Far East Asia (Fig. 1f). Ceuthophilinae are widely distributed across North America (Fig. 1h), while

the distribution of Tropidischinae and Gammarotettiginae is restricted to the west coast of North America (Fig. 1i). Dolichopodainae and Troglophilinae are distributed throughout the

Mediterranean Sea. Macropathinae show a Gondwanian pattern and are distributed in South America, South Africa, Australia, Tasmania, and New Zealand. An extinct fossil subfamily,

†Protroglophilinae, is found only in Baltic amber, which is estimated to have occurred 44 Mya in the Eocene epoch8,9,10. Because Rhaphidophoridae are primitively wingless, which potentially

limits their ability to disperse and colonize, it is reasonable to hypothesize that geological events could have significantly impacted lineage diversification, resulting in their current

distribution. Previous studies on cave-dwelling species showed that vicariant processes played an important role in shaping the complex paleogeographic histories of Dolichopodainae and

Troglophilinae in the Mediterranean and of Macropathinae in the Southern Hemisphere11,12,13,14,15,16,17,18,19,20. Numerous hypotheses on the origins of Rhaphidophoridae have been proposed

based on the morphological or molecular phylogeny of related ensiferan groups16,20,21,22,23,24,25. Karny26 and Ander21 hypothesized that Rhaphidophoridae originated in the Southern

Hemisphere and spread to the Northern Hemisphere, and Ander21 hypothesized that Dolichopodainae in the Mediterranean was the oldest representative subfamily. Hubbell and Norton25 stated that

Macropathinae could be the subfamily most related to Ceuthophilinae in North America based on their morphological characteristics. However, Allegrucci et al.16 and Allegrucci and Sbordoni20

provided an alternative hypothesis based on molecular evidence, in which they proposed that Rhaphidophorinae and Aemodogryllinae in Southeast Asia would be the most closely related

subfamilies to Macropathinae. In addition, they estimated the origin of Rhaphidophoridae in the Cretaceous period (117 Mya; 95% HPD: 105–130 Mya) and suggested that ancestors of

Rhaphidophoridae must have been distributed in both the Southern and Northern Hemispheres since Pangaea. However, a recent phylogenomic study of Orthoptera3 led authors to estimate that the

crown Tettigoniidea originated in the Permian (268 Mya; CI, 308.1–227.7 Mya) and Rhaphidophoridae in the Jurassic, both of which were considerably older estimates than those of previous

studies. Despite a long evolutionary history and a relatively large number of species, the morphological uniformity and lack of diagnostic characteristics within Rhaphidophoridae have made

it difficult to robustly classify this family. For example, Anoplophilinae, which is found only in Far East Asia (Fig. 1a), has been a particularly enigmatic group. Ichikawa27 and Ishikawa28

first used the subfamily name Anoplophilinae, but it was rendered invalid due to the apparent lack of proper diagnosis and description. Otte29 considered each of the two genera of

Anoplophilinae to be placed in two different extant subfamilies: the genus _Anoplophilus_ Karny in the Mediterranean subfamily Troglophilinae and the genus _Alpinanoplophilus_ Ishikawa in

the North American Tropidischiinae. Kim and Kim30 also treated _Anoplophilus_ as a member of Troglophilinae. On the other hand, Gorochov31 first proposed the species of Anoplophilinae to be

closely related to the extinct subfamily †Protroglophilinae found in Baltic amber, followed by Ishikawa32 and Sugimoto and Ichikawa33, who placed the species of Anoplophilinae in

†Protroglophilinae and considered them to be related to Ceuthophilinae in North America based on their morphological features. Later, Storozhenko and Paik34 established Anoplophilinae as a

separate subfamily based on its morphological features, such as hind tibiae, a male subgenital plate, and ovipositor, but its relationships with other subfamilies remain unresolved. Despite

these controversies, the systematics of the Asian Rhaphidophoridae have never been examined using molecular data. In this study, we present the most comprehensive phylogeny of

Rhaphidophoridae, in which all extant subfamilies, including the controversial subfamily Anoplophilinae, were sampled for the first time. The subfamilies Tropidischinae and Gammarotettiginae

from western North America were also included for the first time. Based on our divergence time estimation and ancestral range reconstruction with time-stratified analyses, we propose new

biogeographical hypotheses that are in line with the lineage diversification history within Rhaphidophoridae and major geological events. We also suggest a hypothesis regarding wing loss in

Rhaphidophoridae, which will provide new insights into the origin and evolution of this interesting family. RESULTS PHYLOGENY OF RHAPHIDOPHORIDAE A total of 3,151 bp nucleotide sequences

were used for phylogenetic reconstruction, including 951 bp of COI, 463 bp of 12 S rRNA, 590 bp of 16 S rRNA, 519 bp of 18 S rRNA, and 628 bp of 28 S rRNA. We recovered monophyletic

Rhaphidophoridae in the dataset containing 112 species, with strong nodal support in both the ML and BI analyses (Fig. 2). All subfamilies were recovered as monophyletic except

Tropidischiinae, which was a monotypic subfamily, and Gammarotettiginae, which included only one species in the analyzed dataset. Rhaphidophoridae was largely divided into two lineages:

(Macropathinae + ((Gammarotettiginae + Anoplophilinae) + (Rhaphidophorinae + Aemodogryllinae))) and ((Ceuthophilinae + Tropidischiinae) + (Dolichopodainae + Troglophilinae)). Macropathinae,

which includes species from Australia, New Zealand, South Africa, and South America, was recovered as monophyletic, confirming a Gondwanan origin of the subfamily. Within Macropathinae, the

species from South America and those from New Zealand each formed clades, whereas the species from Australia did not form a monophyletic group. The earliest diverging lineage within

Macropathinae was _Parvotettix_ sp. from Tasmania, and the next branching lineage was _Spelaeiacris tabulae_ Péringuey in South Africa. However, internal relationships between the clades of

species from South America, New Zealand, and several Australian species were incongruent between the BI and ML trees. In addition, the three Asian subfamilies did not form monophyletic

groups. Rhaphidophorinae was found to be sister to Aemodogryllinae, but Anoplophilinae, whose phylogenetic placement had been previously unknown, was recovered as sister to Gammarotettiginae

from the west coast of North America in both the ML and BI analyses. In Aemodogryllinae, the genus _Diestrammena_ was recovered as paraphyletic because _Atachycines_, _Paratachycines_, and

_Tachycines_ were nested within _Diestrammena. Tachycines coreana_ Yamasaki, which was previously synonymized under _T. asynamorus_ Adelung, was separated from _T. asynamorus_ as a distinct

species with a relatively long branch length. Tropidischiinae was recovered as sister to Ceuthophilinae from North America. Troglophilinae was recovered as sister to Dolichopodainae from the

Mediterranean region in the ML analysis, but in the BI analysis, it was recovered as sister to (Ceuthophilinae + Tropidischiinae). In Troglophilinae, the species of _Troglophilus_ from

insular Greece and Anatolia were recovered as monophyletic. _Gammarotettix genitalis_ Caudell and _Comicus campestris_ (one of the outgroups) each had a relatively long branch length in the

ML tree. DIVERGENCE TIME ESTIMATE AND BIOGEOGRAPHY The estimated divergence time indicated that Rhaphidophoridae originated at approximately 138 Mya (early Cretaceous), and lineage

diversification to subfamilies occurred during the Cretaceous period, radiating until the Cenozoic (Fig. 3). Both DIVALIKE + J and BAYAREALIKE + J were recommended as best-fit models in the

BioGeoBEARS analysis with the same LnL scores (−45.92) (Figs. S2–S7; Table S4). The results of the two analyses were highly similar, and the result of DIVALIKE + J was selected for

describing biogeographic events. The ancestral distribution of the most recent common ancestor (MRCA) of extant Rhaphidophoridae was proposed to various probabilities. All regions except for

South America were believed to contribute to the ancestral distribution of the MRCA of extant Rhaphidophoridae around 138 Mya. The MRCA of Rhaphidophoridae was largely divided into two

lineages: (Macropathinae + ((Anoplophilinae + Gammarotettiginae) + (Aemodogryllinae + Rhaphidophorinae))) lineage in Gondwana and Laurasia (Asia and West Coast of North America) and

((Ceuthophilinae + Tropidischiinae) + (Troglophilinae + Dolichopodainae)) lineage in western Laurasia (North America and the Mediterranean region). Macropathinae shows the clearest

distribution pattern that is characteristic of the ancient radiation in Gondwana, after having diverged at approximately 133 Mya. The ancestral distribution of MRCA of Macropathinae was

proposed in Tasmania, New Zealand, and Africa. The earliest diverging lineage within this subfamily was _Parvotettix_ species found in southern Australia (Tasmania), followed by a South

African lineage, _Spelaeiacris tabulae_. The remaining species within the subfamily are closely related to each other, including those currently found in New Zealand and those found in South

America. The lineage distributed in Asia and California was proposed to have diverged from the common ancestor with Macropathinae to 139 Mya and recolonized to Laurasia (Asia and

California). The Laurasian lineage was divided into the Anoplophilinae + Gammarotettiginae lineage in eastern Asia and the west coast of North America and the Rhaphidophorinae +

Aemodogryllinae lineage in Southeast Asia at 106 Mya. Among them, Gammarotettiginae diverged from Anoplophilinae at 96 Mya in eastern Asia and the west coast of North America. In Southeast

Asia, Aemodogryllinae diverged from Rhaphidophorinae. Within Aemodogryllinae, _Diestramima_ Storozhenko + _Atachycines_ Furukawa remained in Southeast Asia, whereas _Paratachycines_ +

_Diestrammena_ + _Tachycine_s migrated to East Asia and diversified. Except _Paratachycines_, which first branched out in East Asia, _Diestrammena_ and _Tachycines_ had multiple dispersal

events and recolonized Southeast Asia several times. The divergence time estimation without the constraint was different in detail, but the results of the biogeographic analyses consistently

showed that Gammarotettiginae diverged in eastern Asia and the west coast of North America (Figs. S8–S13). In the unconstrained timetree, Gammarotettiginae was separated from the common

ancestor of Anoplophilinae + Rhaphidophorinae + Aemodogryllinae at 122 Mya in Asia, including Beringia. The lineage that diverged from the MRCA of Rhaphidophoridae in western Laurasia was

divided into two clades at 108 Mya: (Ceuthophilinae + Tropidischiinae) in North America and (Troglophilinae + Dolichopodainae) in the Mediterranean region. The Mediterranean lineage was

further divided into Dolichopodainae and Troglophilinae at 91 mya. The extant species of Dolichopodainae and Troglophilinae were estimated to have radiated at 30 Mya and 36 Mya,

respectively. Tropidischiinae was split from its common ancestor with Ceuthophilinae at 89 Mya and recolonized the west coast of North America. The west coast of North America was

recolonized twice independently by Gammarotettiginae and Tropidischiinae, lineages with an ancient divergence history of 138 Mya. DISCUSSION This study represents the first molecular

phylogenetic analysis that included all known subfamilies of Rhaphidophoridae. The monophyly of Rhaphidophoridae is supported based on molecular data. Our recovered topology was congruent

with that described in previous studies20, and our results clarified the phylogenetic position of Anoplophilinae, Gammarotettiginae, and Tropidischiinae, which were previously unresolved.

Our study also supports the establishment of Anoplophilinae as an independent subfamily34. As such, the previous hypotheses that placed the East Asian genus _Anoplophilus_ in Troglophilinae

in the Mediterranean29,30 are refuted based on our results. However, our study did not include the genus _Alpinoplophilus_ in Anoplophilinae, which inhabits only Japan and Far East Russia,

and therefore, the hypothesis that placed the genus _Alpinoplophilus_ in Tropidischiinae in North America29 remains untested. In addition, the hypothesis that Anoplophilinae species were

part of the extinct †Protroglophilinae found in Baltic amber31,32,33 could not be evaluated with molecular phylogeny. Unlike another hypothesis, which considered Gammarotettiginae to be

sister to Ceuthophilinae31, our results revealed Gammarotettiginae as sister to Anoplophilinae. Although Gammarotettiginae was analyzed using one gene from public data, we were also able to

present morphological evidence to support the relationship between the two groups. We propose the synapomorphies that unite Gammarotettiginae and Anoplophilinae as a monophyletic group

include a straight dorsal profile and upper margin of the upper valve of the ovipositor with denticles at the apex (Fig. 2b). The denticles on the upper valve of the ovipositor of

Anoplophilinae are weakly pronounced compared to those of Gammarotettiginae. The members of Gammarotettiginae, known as arboreal camel crickets, have tree-dwelling habitats35, and the

members of Anoplophilinae are often found on trees (personal observation). This similarity in ecological behavior can also be considered a trait derived from the same ancestor, and the

weakness of denticles on the upper valve of the ovipositor can be considered a result of the behavioral shift from arboreal to terrestrial habitats. Tropidischiinae, whose phylogenetic

position was uncertain based on morphology alone31, has been found as sister to Ceuthophilinae from North America. However, we were unable to clearly define a synapomorphy that unites

Tropidischiinae and Ceuthophilinae due to the wide range of morphological diversity in Ceuthophilinae. Macropathinae had been considered to be the earliest diverging lineage within

Rhaphidophoridae25,31, but our study, as well as that by Allegrucci and Sbordoni20, refuted this hypothesis, as Rhaphidophoridae was found to be broadly divided into two major clades.

Macropathinae is characterized by a few unique characters that were considered primitive by previous authors, but here, we consider them to be autapomorphic, such as the distal part of the

male genital plate divided into upper and lower parts and the upper margin of the first segment of the hind tarsi with paired apical spines. Based on our phylogeny, we consider morphological

synapomorphy that unites Rhaphidophorinae and Aemodogryllinae to be the inner apical spine on the fore femora (Fig. 2c). The ML tree showed Dolichopodainae and Troglophilinae to form a

monophyletic group, which is supported by the transverse epiphallic sclerite in male genitalia (Fig. 2d). Rhaphidophoridae, found on all continents except Antarctica, exhibit geographic

endemism influenced by paleogeological events11,12,13,14,15,16,17,18,19,20. Using the taxonomically comprehensive dataset for the first time, we evaluated and reviewed the biogeographic

hypothesis with estimated divergence time (Fig. 4). We found a new concurrence between our estimated divergence in Rhaphidophoridae and the age of specific paleogeological events, indicating

that dispersal and vicariance occurred when the region was connected and separated in biogeographic history. We propose a new biogeographic hypothesis suggesting that Gammarotettiginae in

California originated near Beringia when Asia and America were connected by the Bering land bridge (Fig. 5). The connection between Asia and America by the Bering land bridge during the

Cretaceous Period is supported by multiple trans-Beringia dispersals of dinosaurs such as ceratopsids, hadrosaurids and theropods (Fig. 4c)36,37,38,39. During the late Cretaceous Period,

when Tropidischiinae diverged from Ceuthophilinae on the west coast of North America, the region was separated by the opening of the western interior seaway40,41. At 133 Mya, Macropathinae

and the Asia-Beringian lineage were divided between Gondwana and eastern Laurasia, even though the two continents were already separated. However, fossil records suggest the possibility of a

connection and exchange of biota between the two continents42,43. Since the distribution of the MRCA of Rhaphidophoridae was proposed to encompass all regions except South America in the

ancestral state reconstruction, it is challenging to specify its exact origin. Our divergence time estimates sometimes deviate from previously considered paleogeological events. For

instance, the divergence of _Spelaeiacris tabulae_ from South Africa (63 Mya) postdates the separation of South Africa from Gondwana (117–96 Mya)44,45. In our biogeographic analysis, we

confirmed that some areas have been recolonized several times by various lineages. The west coast of North America was recolonized by both Gammarotettiginae from Beringia and

Tropidischiinae, splitting from the common ancestor of the North American lineage. Southeast Asia also experienced multiple recolonizations by Rhaphidophorinae and some Aemodogryllinae

lineages. The extant Mediterranean lineage radiated recently, and before this radiation, †Protroglophilinae, a fossil found in Baltic amber, could have occupied the ecological niche of the

European region. The phylogenetic position of †Protroglophilinae remains unresolved, but based on the endemic distribution of Rhaphidophoridae, it could be hypothesized that

†Protroglophilinae was closely related to western Laurasia lineages. We inferred that the ancestral lineages that gave rise to Rhaphidophoridae could have been distributed throughout Pangaea

around 138 Mya. The recent molecular phylogeny by Allegrucci and Sbordoni20 suggested a similar distribution in which Rhaphidophoridae originated in the Mesozoic Cretaceous period of 117

Mya, but our estimated age was older. The previous study, based on morphology, suggested the following existing hypothesis for the origin of Rhaphidophoridae: Ander21, who proposed

Dolichopodainae as the oldest representative of Rhaphidophoridae; Hubbell and Norton25, who considered Macropathinae as an ancestor lineage with Ceuthophilinae; Gorochov31, who suggested

Macropathinae as the most basal lineage of Rhaphidophoridae, and Ceuthophilinae and Gammarotettiginae is a direct descendant of Macropathinae, and Aemodogryllinae and Rhaphidophorinae sister

to Dolichopodainae and Troglophilinae. Our study found that there is no single basal lineage, but the family consists of two major lineages that diverged early on. Dispersal ability plays a

key role in genetic differentiation to speciation46,47. In the same context, the endemic distribution and biogeographic history of Rhaphidophoridae are caused by their common trait, which

is flightless with loss of wings. Although flightlessness and the loss of wings are common in Orthoptera48,49,50,51, Rhaphidophoridae is significantly older than other wingless

orthopterans3,52,53. Flightlessness has long influenced their evolutionary history, and furthermore, the loss of wings in their common ancestor would imply the origin of Rhaphidophoridae.

Therefore, we propose a narrative hypothesis about wing loss in Rhaphidophoridae, which could be a result of adaptation to low temperatures in the Mesozoic era. Although there are some

exceptions54, the loss of wings in cold-specialized insects is common since they reduce their activity and metabolism for adaptation to low temperatures, and the frequency of encountering

predators and flying away from them is low in cold environments49,55. Prior to the first divergence of the MRCA of Rhaphidophoridae, there was global climate cooling at the Middle–Late

Jurassic transition (Late Callovian–Middle Oxfordian) (Fig. 3a)56,57,58,59. In addition, the dispersal path and distribution were restricted to near the polar regions (Fig. 4c and d) with

relatively low temperatures. The extant cold-specialized insects, Boreidae (Mecoptera) and Grylloblattodea (Notoptera), were derived at a roughly similar age to Rhaphidophoridae, supported

by fossil evidence60,61 and molecular estimation62. Moreover, extant Rhaphidophoridae species show a preference for low temperatures, such as alpine species that are found close to permanent

ice63, subantarctic species7, and vast troglophile species, which can be the result of cold adaptation, relating to our hypothesis about the loss of wings. To support this hypothesis,

further research is needed to precisely estimate divergence times and obtain genomic evidence of cold adaptation in wingless insects, including cave crickets. CONCLUSION Our phylogenic

analysis, which included all known subfamilies, revealed a unique and novel placement of the Asian subfamily Anoplophilinae. We also confirmed that the endemic distribution of

Rhaphidophoridae as a result of winglessness is valid. Beringia, which connected Asia and North America, and the opening of the western interior seaway during the Cretaceous period coincide

with the estimated divergence time of the Rhaphidophoridae lineages. The difference between the geological events and the molecular clock can be explained by several hypotheses regarding the

dispersal capabilities of Rhaphidophoridae, but it can also be caused by a limitation of the dataset. Here, we suggest a hypothesis that global temperature and climate changes have affected

lineage diversification and propose a narrative hypothesis that adaptation to low temperatures caused the loss of wings, leading to the endemic distribution of Rhaphidophoridae. Further

research is needed to fully test these interesting hypotheses and gain a deeper understanding of the evolution and diversification of the cave cricket. MATERIAL AND METHODS TAXON AND

CHARACTER SAMPLING In total, 112 species from all nine extant subfamilies within Rhaphidophoridae were used for our ingroup taxon sampling process. We sampled one species of Anoplophilinae,

represented by _Anoplophilus koreanus_ Storozhenko and Paik, collected from Korea, and five species of Aemodogryllinae, collected from Korea and Russia. For Aemodogryllinae, we included

Asian representatives of _Paratachycines_ Storozhenko, _Tachycines_ Adelung, and _Diestramena_ Brunner von Wattenwyl. In addition, for this study, we incorporated sequence data from

Tropidischinae and Gammarotettiginae and other species belonging to different subfamilies that were not previously analyzed in phylogenetic analyses. The sequence data for the remaining

taxa, which were part of previous studies14,15,16,17,18,19,20 were retrieved from GenBank. For outgroups, we included three ensiferan species representing Schizodactylidae, Gryllacrididae,

and Tettigoniidae, all of which belong to the infraorder Tettigoniidea (Table S1). Genomic DNA was extracted from specimens’ legs preserved in 99% EtOH or dry-mounted using the DNeasy Blood

and Tissue kit (QIAGEN, Inc.) according to the manufacturer’s guidelines. The voucher specimens were preserved in 100% ethanol and stored in a −80 °C deep freezer with matching extracted DNA

in the School of Biological Sciences, Seoul National University, Seoul, South Korea (SNUE). Mitochondrial cytochrome _c_ oxidase I (COI), small ribosomal subunit (12 S), large ribosomal

subunit (16 S), and nuclear ribosomal RNA 18 S and 28 S gene fragments were selected for multigene phylogenetic analysis based on previous studies14,15,16,17,18,19,20. Fragments of the genes

were amplified using AccuPower PCR primers (Bioneer, Korea), and information on the primers and PCR conditions for each gene is listed in Table S2. PCR products were visualized and

confirmed by electrophoresis on a 2% agarose gel and sequenced using the Sanger method. The sequence data of each gene from both directions were assembled using SeqMan Pro v. 7.1.0 (DNASTAR,

Inc., U.S.A). MOLECULAR PHYLOGENETIC ANALYSES Protein-coding genes and ribosomal RNA genes were aligned using different methods. The sequence of mitochondrial COI was translated to its

amino acid sequence for conservation of reading frames, aligned using MUSCLE64, and back-translated to nucleotides in MEGA X65. The ribosomal RNA sequences (12 S, 16 S, 18 S, and 28 S) were

aligned using MAFFT ver.766 with the E-INS-i method. All individual gene alignments were concatenated into a single matrix using FASconCAT-G67. The nucleotide substitution model for each

gene partition was estimated using PartitionFinder 268 with a greedy algorithm. We performed maximum likelihood (ML) and Bayesian inference (BI) analyses on the concatenated dataset using a

cluster computer on SNUE, and both analyses were performed by applying the recommended substitution model for each partition by PartitionFinder2. The ML analysis was performed using IQ-tree

1.6.269 with 2,000 bootstrap replications. BI analysis was performed using MrBayes 3.2.670. Except for the applied substitution models for each partition followed by PartitionFinder2,

default priors were used for all other parameters. The posterior distribution was estimated using Markov chain Monte Carlo (MCMC) with four chains for 100 million generations and sampling

every 5,000 generations. Using Tracer 1.771, we determined that convergence of the run and all effective sample sizes (ESSs) of parameters were over 200. The trees were summarized in MrBayes

3.2.670 by discarding the first 25% of the results as burn-in. In summarizing trees, the contype was set to ‘Allcompat’, which adds all compatible groups to the tree, and the minimum

probability of partitions was assigned a value of 10. DIVERGENCE TIME ESTIMATION The divergence time of Rhaphidophoridae was estimated using MrBayes 3.2.670 in a Bayesian framework. Because

this program requires only one outgroup, we kept _Comicus campestris_ Irish, and two other outgroup taxa were removed from the dataset. The nucleotide substitution models proposed by

PartitionFinder 2 were applied to each gene. Since a recent study showed that the birth-death prior produced stable results on molecular dating across all scenarios72, we used the

birth–death process for a tree model. The posterior distribution was estimated using MCMC with four chains for 100 million generations and sampling every 10,000 generations. The process of

examining the results and summarizing trees was performed with the same methods as the BI analysis using Tracer 1.771 and MrBayes 3.2.670. It was confirmed that all ESSs of the model

parameters exceeded 200. A topological constraint was applied to the clade between Anoplophilinae and Gammarotettiginae for consistency in topologies of the molecular clock dating tree with

the ML tree and BI tree. Although fossil records provide reliable calibrations for molecular clock and lineage diversification, we were unable to use fossil calibrations in this study.

†Protroglophilinae, the extinct Rhaphidophoridae subfamily found in Baltic amber, could not be used for calibration since its phylogenetic position was ambiguous. Gorochov31 proposed the

phylogenetic position of †Protroglophilinae, but his proposal was challenging to confirm with recent molecular phylogenetic studies16,20 and our own phylogenetic analysis due to differing

relationships between the subfamilies. In addition, while previous studies have utilized paleogeographic events to calibrate their time tree20, we opted not to use such events for

calibration to re-evaluate biogeographic hypotheses based on the divergence time estimated by only molecular clocks. Instead, we relied on divergence time estimates from the most recent

phylogenomic study3 of Orthoptera as our secondary calibration points. In this study, we used two calibration points: (1) 268 Mya as the origin of crown-Tettigoniidea, at the base of the

phylogeny, and (2) 138 Mya, at the node where there was a division of Macropathinae and Aemodogryllinae + Rhaphidophorinae + Anoplophilinae + Gammarotettiginae. ANCESTRAL RANGE ESTIMATION

FOR BIOGEOGRAPHIC ANALYSES The biogeographical history of Rhaphidophoridae was analyzed using the package BioGeoBEARS in R 4.1.073. Based on the divergence time estimation, ancestral range

estimation was performed excluding an outgroup. We defined the distribution of Rhaphidophoridae in nine areas: West Coast of North America (California), North America, Mediterranean Region,

South America (including Falkland Islands), South Africa, Tasmania (Australia), New Zealand, Eastern Asia (Korea, Japan, and Far East Russia), and Southeast Asia (South China, Vietnam,

Bhutan, Indonesia, and Philippines). The distribution of taxa was obtained from previous studies that retrieved data19,20 or the Orthoptera Species File1. A time-stratified analysis with

dispersal probabilities specified for each period was conducted to consider the junction and separation of biographic areas according to continental drift during the diversification time of

Rhaphidophoridae, which is nearly 150 Mya. The time scale was stratified into 30 million-year slices74: 0–30 Mya, 30–60 Mya, 60–90 Mya, 90–120 Mya, and 120–150 Mya (Fig. S1). The dispersal

probabilities were scored by the following categories according to the connectivity of the biogeographic areas74,75: 0.01 for well-separated areas by water, 0.1 for moderately separated or

connected areas, but the other area was inserted between the areas, or the areas were distant and separated by two or more land masses, and 1.0 for contiguous areas (Table S3). The

connectivity of the biogeographic area over time was determined according to Scotese76. We used six biogeography models: (1) DEC (dispersal-extinction-cladogenesis)77; (2) DEC + J (including

founder-event speciation); (3) DIVALIKE, a likelihood version of DIVA (dispersal-vicariance)78; (4) DIVALIKE + J (including founder-event speciation); (5) BAYAREALIKE, a likelihood version

of BayArea (Bayesian inference of historical biogeography for discrete areas)79; and (6) BAYAREALIKE + J (including founder-event speciation). Two parameters, d = dispersal and e =

extinction, were included in these six models. We compared the likelihood values of the models using the likelihood ratio test, and the most likely model was selected using the Akaike

information criterion (AIC)80,81. REPORTING SUMMARY Further information on research design is available in the Nature Portfolio Reporting Summary linked to this article. DATA AVAILABILITY

The additional supporting information of this study are available in the supplementary material of this article. Datasets are archived at the Zenodo Digital Repository at

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PubMed  Google Scholar  Download references ACKNOWLEDGEMENTS The first author expresses his special gratitude to Jaedong Gim (Chungnam National University) and Min Kim (Kangwon National

University) for their cooperation on taxon sampling and field observation. He also thanks to Sangjin Han (Seoul National University) and Jihoo Kim (Coconut Rhinoceros Beetle Response Hawaii)

for their cooperation on taking photograph of _Gammarotettix genitalis_ in California. This research was supported by the National Research Foundation of Korea (NRF) grants funded by the

Korean government (MEST, No. 2019R1A6A1A10073437; MSIT, No. 2021R1C1C1003452; Comparative medicine Disease Research Center, SRC, No. 2021R1A5A1033157). This work is also supported by Seoul

National University (Creative-Pioneering Researchers Program; the New Faculty Startup Fund; Laying the Groundwork for Peace and Unification, the Institute for Peace and Unification Studies),

the U.S. National Science Foundation (No. DEB-1937815) and the United State Department of Agriculture (Hatch Grant, No. TEX0-1-6584 to H.S). AUTHOR INFORMATION Author notes * These authors

contributed equally: Do-Yoon Kim, Sangil Kim. AUTHORS AND AFFILIATIONS * School of Biological Sciences, Seoul National University, Seoul, 08826, Republic of Korea Do-Yoon Kim, Sangil Kim 

& Seunggwan Shin * Comparative Medicine Disease Research Center, Seoul National University, Seoul, 08826, Republic of Korea Do-Yoon Kim & Seunggwan Shin * Research Institute of Basic

Sciences, Seoul National University, Seoul, 08826, Republic of Korea Sangil Kim & Seunggwan Shin * Museum of Comparative Zoology and Department of Organismic and Evolutionary Biology,

Harvard University, Cambridge, MA, 02138, USA Sangil Kim * Department of Entomology, Texas A&M University, College Station, TX, USA Hojun Song Authors * Do-Yoon Kim View author

publications You can also search for this author inPubMed Google Scholar * Sangil Kim View author publications You can also search for this author inPubMed Google Scholar * Hojun Song View

author publications You can also search for this author inPubMed Google Scholar * Seunggwan Shin View author publications You can also search for this author inPubMed Google Scholar

CONTRIBUTIONS D.K. designed the study and collected samples and data. D.K., S.K. and S.S. performed phylogenetic analyses. D.K., S.K. and H.S. performed divergence time estimation. D.K. and

S.K. performed biogeographic analyses. All authors contributed to the writing of the manuscript. CORRESPONDING AUTHOR Correspondence to Seunggwan Shin. ETHICS DECLARATIONS COMPETING

INTERESTS The authors declare no competing interests. PEER REVIEW PEER REVIEW INFORMATION : _Communications Biology_ thanks Zhuqing HE and the other, anonymous, reviewer(s) for their

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Phylogeny and biogeography of the wingless orthopteran family Rhaphidophoridae. _Commun Biol_ 7, 401 (2024). https://doi.org/10.1038/s42003-024-06068-x Download citation * Received: 16 June

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