# Out-of-Africa: Origin of the Disjunct Distribution of Paleotropical Eneopterinae Crickets (Gryllidae, Xenogryllini)

Zhe-yuan Yu, Jiajia Dong, Sylvain Hugel, Ranjana Jaiswara, Zhu-qing He, Tony Robillard, Gael J. Kergoat

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Zhe-yuan Yu, Jiajia Dong, Sylvain Hugel, Ranjana Jaiswara, Zhu-qing He, et al. Out-of-Africa: Origin of the Disjunct Distribution of Paleotropical Eneopterinae Crickets (Gryllidae, Xenogryllini). Zoologica Scripta, In press, 10.1111/zsc.70019. hal-05243558

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ORIGINAL ARTICLE

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# Out-of-Africa: Origin of the Disjunct Distribution of Paleotropical Eneopterinae Crickets (Gryllidae, Xenogryllini)

Zhe-Yuan Yu¹,² ID | Jiajia Dong³ ID | Sylvain Hugel⁴,⁵ ID | Ranjana Jaiswara⁶ ID | Zhu-Qing He⁷ ID | Tony Robillard¹ ID |
Gael J. Kergoat² ID

¹Institut de Systématique, Evolution, Biodiversité (ISYEB), Muséum National D'histoire Naturelle, CNRS, SU, EPHE-PSL, UA, Paris, France | ²CBGP, INRAE, IRD, CIRAD, Institut Agro, Université de Montpellier, Montpellier, France | ³School of Life Sciences, Jiangsu Key Laboratory of Brain Disease and Bioinformation, Research Center for Biochemistry and Molecular Biology, Xuzhou Medical University, Xuzhou, China | ⁴Madagascar Biodiversity Center, Antananarivo, Madagascar | ⁵Institut Des Neurosciences Cellulaires et Intégratives, UPR 3212 CNRS-Université de Strasbourg, Strasbourg, France | ⁶Department of Zoology, Panjab University, Chandigarh, India | ⁷School of Life Sciences, East China Normal University, Shanghai, China

**Correspondence:** Zhe-Yuan Yu (zheyuan.yu@edu.mnhn.fr) | Jiajia Dong (jiajia.dong@xzhmu.edu.cn) | Gael J. Kergoat (gael.kergoat@inrae.fr)

**Received:** 3 June 2025 | **Revised:** 1 August 2025 | **Accepted:** 9 August 2025

**Funding:** This work was supported by the National Natural Science Foundation of China (Grant #32100334, Grant #31801957), the Natural Science Foundation of Jiangsu Province (Grant #BK20210897, Grant #BK20181471), the Scientific Research Foundation for the Talents by the Xuzhou Medical University (Grant #D2020009), and the ANR Sociogenomics (Grant #ANR-19-CE02-0023-PI Frédéric Legendre, MNHN). Zhe-Yuan Yu's doctoral thesis is funded by the China Scholarship Council (Grant #202306140037). This work was supported by a DST Faculty Grant of the Government of India (project INSPIRE; Grant #DST/INSPIRE/04/2018/003866) awarded to Ranjana Jaiswara.

**Keywords:** Africa | Asia | biogeography | colonisation routes | India | Madagascar | paleotropical intercontinental disjunction

## ABSTRACT

Many groups of terrestrial plants and animals display a disjunct distribution pattern in the Paleotropics, being found almost exclusively in tropical Africa and Asia. The origin of such a pattern may be manifold, particularly for older lineages where plate tectonics potentially played an important role. Several potential colonisation routes may have been involved in dispersal events between the Afrotropics and tropical Asia, not to mention long-distance dispersal events through island hopping. To advance in the understanding of disjunct Paleotropical distributions, we focus on a tribe of Paleotropical crickets, the Xenogryllini. They supposedly originated in the Early Eocene, thus experiencing multiple past geological and environmental changes during their diversification. Resolving the evolutionary history of Xenogryllini is thus expected to shed light on the biogeographic processes that generated disjunct Paleotropical distributions. To tackle this, we assembled a multi-marker molecular dataset for the tribe encompassing all species. We carried out phylogenetic, molecular dating, and historical biogeography analyses. Our results provide a highly supported phylogenetic hypothesis for the tribe and suggest an out-of-Africa origin with several independent dispersal events towards Asia, likely involving distinct colonisation routes in the Neogene.

## 1 | Introduction

One of the main questions related to the biogeography of organisms is the origin of disjunct distributional ranges (Cox

and Moore 2010; Lomolino et al. 2010), where extant taxa consist of lineages that are found in distinct regions separated by gaps in geographic ranges. Many extant terrestrial lineages present disjunct distributions in the Paleotropics (distributed

Zhe-Yuan Yu and Jiajia Dong shared first authorship.

This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.

© 2025 The Author(s). Zoologica Scripta published by John Wiley & Sons Ltd on behalf of Royal Swedish Academy of Sciences.
in both tropical Africa and Asia), such as plants (Thorne 1972; Zhou et al. 2012; Yu et al. 2014), amphibians (Li et al. 2013; Oliver et al. 2015) and insects (Toussaint et al. 2021; Rossini et al. 2022). Several competing hypotheses have been proposed to explain this pattern of paleotropical intercontinental disjunction (PID) (e.g., see Thomas et al. 2015; Rossini et al. 2022), which account for past tectonic activities and environmental changes after the breakup of Gondwana (e.g., see the reviews by Sanmartín et al. 2001 for the Northern hemisphere and Sanmartín and Ronquist 2004 for the Southern hemisphere).

For relatively old lineages originating during the Mesozoic and around the Mesozoic/Cenozoic boundary, PID patterns are associated with vicariance events, and the corresponding explanatory hypothesis is generally referred to as the 'Biotic ferry model', 'Out-of-India' or 'Indian raft' (Datta-Roy and Praveen Karanth 2009). Under its premise, vicariance events resulting from the split between Madagascar and the Indian Plate in the Late Cretaceous (see Figure 1A) ultimately led to disjunct distributions between tropical Africa and Asia. Such a pattern has been recovered in several groups such as insects (e.g., Toussaint et al. 2016), vertebrates (e.g., Bossuyt and Milinkovitch 2001; Okajima and Kumazawa 2010), or plants (e.g., Conti et al. 2002; Shukla et al. 2025). All remaining competing hypotheses always involve dispersal events, and they imply distinct colonisation routes that were generally opened for a given period of time. Starting in the Late Cretaceous, when the Indian Plate drifted northward (c. 90–85 million years ago, Mya; Storey et al. 1995; Ali and Aitchison 2008), a first colonisation route was identified (see Figure 1B) and is mainly associated with the 'Kohistan–Ladakh Island Arc' between India and Northeast Africa (Chatterjee and Bajpai 2016), acting as a pathway between India and North Africa along Arabia, and which lasted into the Paleocene (Chatterjee and Bajpai 2016; Smith et al. 2016; Morley 2018). This route was inferred to be an important pathway for several groups, such as plants (Morley 2018; Bansal et al. 2022).

For lineages originating from the Paleocene to the Eocene–Oligocene boundary, PID patterns have been hypothesized to originate from the crossing of the Tethyan sea, which separated Africa and Eurasia. These ‘trans-Tethyan dispersal’ events (see Figure 1C) were facilitated during low sea level episodes, initially through western routes (the Alboran and Apulian routes) opening in the Middle Eocene, and later on through an additional eastern route (the Iranian route) that was available until the Oligocene (Gheerbrant and Rage 2006). The terrestrial faunal exchanges between Afro–Arabia and Eurasia are well documented after the Paleocene, especially in mammals (Gheerbrant and Rage 2006; Sen 2013). In the mid-latitudes of the Northern hemisphere, the presence of a widespread boreotropical forest belt originating during the Paleocene and that persisted during the Early Eocene (Wolfe 1975; Wen 1999; Sanmartín et al. 2001) promoted the dispersal of organisms throughout Eurasia, therefore acting as a biotic bridge allowing boreotropical migrations between Africa and Eurasia (Toussaint et al. 2019; Couvreur et al. 2021). During the Eocene–Oligocene boundary (c. 34 Mya), the marked cooling of temperatures (Zachos et al. 2001) drove the contraction of the boreotropical forest and led to the southward dispersals

of multiple groups of organisms (e.g., Condamine et al. 2012; Thomas et al. 2015; Ye et al. 2017; Huang et al. 2019), ultimately resulting in PID patterns.

In the Middle Miocene a major colonisation route accounting for dispersal between Africa and Asia is the so-called 'Gomphotherium landbridge', which opened when the African and Arabian plates collided with Eurasia during the Burdigalian (20.44–15.97 Mya), and lasted till the Serravallian (13.82–11.63 Mya) (Rögl 1998). The timing of the 'hard' collision between Afro–Arabia and eastern Anatolia has been revised recently (Darin and Umhoefer 2022), with an age estimate of c. 25 to 122 Mya. This geological event is also associated with the permanent closure of the Neo-Tethys Ocean, which is also supported by the vicariance of the marine fauna (Harzhauser et al. 2007). This route (see Figure 1D) also promoted the dispersal of diverse groups of terrestrial animals and plants which dispersed in both directions through the Arabian Peninsula (e.g., Zhou et al. 2012; Sen 2013; Toussaint et al. 2019; Tejero-Cicuéndez et al. 2022).

Finally, the hypothesis of transoceanic long-distance dispersal (LDD) between India and Madagascar taking place any time during the Cenozoic cannot be ruled out (see Figure 1D). Patterns of transoceanic dispersal by birds, oceanic drift, rafting, or wind are not uncommon (see the review of Gillespie et al. 2012); transoceanic LDD events between the Indian Plate and Madagascar were also facilitated by the presence of the Seychelles–Mascareignes Plateau and the emergence of ephemeral islands that acted as stepping-stones for dispersal during periods of low sea level (Ali and Aitchison 2008; Warren et al. 2010). The onset of the winter monsoon from the Oligocene/Miocene also potentially promoted dispersal from the Indian subcontinent to Madagascar (Warren et al. 2010; Samonds et al. 2012). However, successful colonisation following transoceanic dispersal remains highly contingent on ecological factors. Some lineages may have reached Madagascar but failed to establish, as suggested by the presence of closely related species on neighbouring islands without representatives in Madagascar itself (Hugel 2022). The dispersal potential of Orthoptera, for example, highlights this pattern, with some groups successfully colonising Madagascar while others only transiently reaching the island without permanent establishment.

While vicariance and dispersal are primary explanations for disjunct distributions, extinction events have also played a critical role in shaping present-day biogeographic patterns. Lineages that were once more widespread may have disappeared from certain regions due to climate shifts, competition, or habitat loss, leading to apparent disjunctions. The 'Indian raft' hypothesis posits that multiple lineages were transported via the Madagascar–Seychelles–India landmass as it drifted northward following its separation from Madagascar. However, the current absence of many clades on one or more of these landmasses, despite their shared geological history, suggests that extinctions have played a crucial role in shaping modern distributions. This is exemplified by amphibians such as Nasikabatrachidae in India and Sooglossidae in the Seychelles, which appear to be relictual taxa, their present distribution likely shaped by historical extinction events (Biju and Bossuyt 2003). Similarly, the Kohistan–Ladakh Arc has been proposed as a corridor for faunal exchange between India and northern Africa via the Arabian
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<figure><img src="image_9.png" /><figcaption>**FIGURE 1** | Legend on next page.</figcaption></figure>
**FIGURE 1** | Paleomaps (with corresponding ages provided below each) illustrating competing biogeographic scenarios that account for PID patterns between Africa and Asia. (A) 'Indian raft' hypothesis; (B) 'Kohistan–Ladakh Island Arc' hypothesis; (C) 'Trans-Tethyan dispersal' hypothesis both western and eastern routes are depicted; (D) 'Gomphotherium landbridge' hypothesis and 'transoceanic LDD hypothesis'; (E) extant distribution of Xenogryllini with genus distribution highlighted using colours: *Indigryllus* (in magenta), *Pseudolebinthus* (in yellow) and *Xenogryllus* (in cyan). World map source from QGIS (https://docs.qgis.org/3.40/en/docs/server_manual/index.html). Paleomap reconstructions were made using GPlates v2.5 (Müller et al. 2018) with open-source data from the PALEOMAP PaleoAtlas for GPlates (Scotese 2016). Species illustrated: (F) *Xenogryllus marmoratus* (Haan, 1844) (picture by K.W. Chan; https://www.inaturalist.org/observations/245034087); (G) *Pseudolebinthus africanus* Robillard, 2006 (picture by B. Wursten; https://www.inaturalist.org/observations/42298851); (H) *Indigryllus sagani* Jaiswara and Robillard, 2022 (picture by S. Sreebin, with permission).

Peninsula. Yet, the absence of certain lineages in these regions today implies that extinctions have obscured former distributions. The role of extinction in shaping disjunct biogeographic patterns is further supported by the case of Rhaphidophoridae, a lineage of cave crickets conspicuously absent from Madagascar despite their presence in surrounding regions (Allegrucci and Sbordoni 2019; Hugel 2022).

To provide further understanding into the origin of PID patterns, we chose as a model system the Xenogryllini crickets, one of the five tribes in the subfamily Eneopterinae (Orthoptera: Gryllidae). The subfamily Eneopterinae comprises more than 300 species, which are widely distributed in tropical and subtropical areas worldwide; like most crickets (Huber et al. 1989), Eneopterinae are omnivorous, with a predominantly herbivorous diet. Species belonging to the tribe Xenogryllini are restricted to paleotropical areas, with a disjunct distribution in Africa and Asia (Vicente et al. 2017; see also Jaiswara, Dong, Ma, et al. 2019 for a more detailed distribution map). Their absence in other bioregions is well documented, as numerous orthopteran surveys have covered regions where Xenogryllini are absent; this is particularly true for Eneopterinae, which have been the focus of numerous studies on lineages found in regions that are devoid of Xenogryllini (e.g., Robillard and Desutter-Grandcolas 2004 for South America, or (Dong et al. 2018), and He et al. 2024 for Australasia). Currently, the tribe consists of three genera and 16 species: (i) the genus *Xenogryllus* Bolívar, 1890, which is distributed both in the Afrotropics and in Asia (mostly in tropical areas), (ii) the genus *Indigryllus* Robillard and Jaiswara, 2019, endemic to India, and (iii) the genus *Pseudolebinthus* Robillard and Desutter-Grandcolas (2006), endemic to East Africa (Vicente et al. 2017; Jaiswara et al. 2018; Jaiswara, Dong, and Robillard 2019; Salazar et al. 2020; Jaiswara et al. 2022). Given their disjunct distribution in the Afrotropics (including Madagascar) and in Asia (including India), Xenogryllini crickets represent a relevant group for investigating the origin of PID patterns. To date, two studies (Vicente et al. 2017; le Flanchec et al. 2025) have included Xenogryllini representatives to investigate biogeographical patterns of crickets at higher taxonomic levels. Both studies included only five Xenogryllini species and yielded contradictory conclusions, either inferring an origin in the Afrotropics (le Flanchec et al. 2025) or in India (Vicente et al. 2017).

Here, we generate a multi-molecular marker dataset (comprising five mitochondrial and three nuclear gene fragments) for all Xenogryllini species (41 specimens representing the 16 species, including *Xenogryllus lemia* Yu et al. (2025), a species recently described from Madagascar, which provides critical insight into

faunal exchanges between Africa and Asia; Yu et al. (2025)) and 22 outgroups. We reconstruct the evolutionary history of the tribe using both Bayesian inference (BI) and maximum likelihood (ML) approaches, and further leverage molecular dating and historical biogeography analyses to infer the ancestral range of the tribe and reconstruct its biogeographic history. The latter will allow us to assess the competing hypotheses (see Figure 1A–D) that led to the PID pattern in Xenogryllini and to determine the biogeographic processes at play while considering the ecological differences between lineages.

## 2 | Materials and Methods

### 2.1 | Taxon Sampling and Molecular Datasets

Based on recent taxonomic studies (Jaiswara, Dong, Ma, et al. 2019; Jaiswara, Dong, and Robillard 2019; Robillard and Murphy 2020; Jaiswara et al. 2022), the tribe Xenogryllini is composed of three genera and 16 valid species (Cigliano et al. 2025; Yu et al. 2025): *Xenogryllus* (nine species), *Pseudolebinthus* (five species) and *Indigryllus* (two species). To reconstruct the phylogeny of the tribe Xenogryllini, a total of 41 individuals representing the 16 species belonging to the three xenogrylline genera were sampled (see Table S1), by assembling recent data produced by our research group (Jaiswara, Dong, and Robillard 2019; Dong et al. 2024) and generating sequences for 13 new specimens (including sequences for six Xenogryllini species that have never been sequenced before). New specimens were obtained from the following insect collections: Muséum national d'Histoire naturelle, Paris (MNHN), Academy of Natural Sciences of Drexel University, Philadelphia, USA (ANSP), personal collection of Ranjana Jaiswara, India (RJINS) and Museum of Biology, East China Normal University, Shanghai, China (ECNU). For specimens collected relatively recently (< 20 years old), we obtained DNA sequences following the protocol described by (Dong et al. 2018). Molecular experimentations were carried out in the 'Service de Systématique Moléculaire', UMS 2700 2 AD, MNHN, CNRS. For two older specimens corresponding to the paratype of *P. africanus* and the only known specimen of an undescribed species from Madagascar, we used a non-destructive protocol for DNA extraction as described in Dong et al. (2024), and used a genome skimming approach (Straub et al. 2012) to sequence molecular markers by following the protocol described by Salazar et al. (2020). The selection of outgroups was based on the phylogeny of Eneopterinae inferred by Vicente et al. (2017): we selected 13 species (represented by 20 individuals) belonging to the four other Eneopterinae tribes along with two Gryllinae species as the most distant outgroups.
Eight molecular markers were selected, consisting of five mito-
chondrial (cytochrome c oxidase subunit I 'COI', cytochrome c
oxidase subunit II 'COII', cytochrome b 'Cytb', 12S and 16S ri-
ribosomal RNA) and three nuclear (histone H3 'H3', 18S and 28S
ribosomal RNA) gene fragments. All individual gene fragments
were aligned in MAFFT v7 (Katoh et al. 2019) with default
parameters and further concatenated under Mesquite v3.83
(Maddison and Maddison 2023). The concatenated molecular
matrix comprises 3664 base pairs (bp) for 63 terminals (here-
after named 'specimen-level dataset'): 414 bp for 12S, 506 bp for
16S; 652 bp for 18S, 372 bp for 28S, 708 bp for COI, 338 bp for
COII, 346 bp for Cytb, 328 bp for H3. Based on the latter dataset,
we also defined a reduced dataset, with only one specimen per
species ('species level dataset', with 33 species); such a dataset
is better suited for dating and historical biogeography analyses,
which typically require one individual per species in order to
properly infer speciation times and dispersal and vicariance
processes.

## 2.2 | Molecular Phylogenetic Analyses

The phylogenetic analyses were carried out on the ‘specimen-
level dataset’ under ML using IQ-TREE v2.0.7 (Minh et al. 2020)
and under BI with MrBayes v3.2.6 (Ronquist et al. 2012). The
concatenated dataset was split into 16 partitions a priori, with
three partitions (one per codon position) defined for each cod-
ing gene fragment (COI, COII, Cytb and H3) and one partition
defined for each non-coding gene fragment (12S, 16S, 18S and
28S). The Bayesian Information Criterion implemented through
ModelFinder (Kalyaanamoorthy et al. 2017) was used to select
best-fit substitution models and partition schemes under ML.
For BI analyses, instead of relying on a single substitution model
per partition, we used the ‘mixed’ option of MrBayes to sample
across substitution models with reversible-jump Markov Chains
Monte Carlo (rj-MCMC; Huelsenbeck et al. 2004).

Maximum likelihood trees were obtained using heuristic
searches implementing 20 random-addition replicates with the
following settings: parsimony-starting tree, hill-climbing nearest
neighbour interchange (NNI) search (–*allnni* option), partition-
resampling strategy (–*sampling GENE* option), best partition
scheme allowing the merging of partitions (–*m MFP* + *MERGE*
option). The perturbation strength was set to a default value of
0.2. Clade support for all analyses was assessed using 1000 rep-
licates for both SH-like approximate likelihood ratio tests (SH-
aLRT; Guindon et al. 2010) and ultrafast bootstraps (uBV; Minh
et al. 2013). Following the authors’ recommendations, branches
with SH-aLRT values ≥ 80% and uBV values ≥ 95% were consid-
ered highly supported.

For BI, two independent runs with eight MCMC (one cold and
seven incrementally heated chains) were conducted: they ran for
20 million generations, with trees sampled every 1000 genera-
tions. A conservative 25% burn-in was applied after checking for
stability on the log-likelihood curves and the split frequencies of
the runs in Tracer v1.7 (Rambaut et al. 2018). Support of nodes
for MrBayes analyses was provided by clade posterior probabil-
ities (PP) directly estimated from the majority-rule consensus
topology. Following Erixon et al. (2003), branches supported by
PP ≥ 0.95 were considered highly supported.

## 2.3 | Molecular Dating Analyses

Divergence times were estimated on the ‘species-level dataset’ using Bayesian relaxed clocks with BEAST v1.10.4 (Suchard et al. 2018). Given the absence of well-assigned fossils for the Xenogryllini or closely related eneopterine groups, we relied on secondary calibrations based on a study on Eneopterinae implementing external fossil calibrations (Vicente et al. 2017). Based on the taxa shared between our dataset and theirs, we used uniform distributions to constrain the ages of the most recent common ancestors (MRCA) of the following clades, using the 95% highest posterior densities (HPD) of the estimated ages from the Vicente et al. (2017) study as a reference: (i) MRCA of crown Eneopterinae (minimum age of 63.28 Mya and maximum age of 96.76 Mya), (ii) MRCA of crown Lebinthini (minimum age of 44.43 Mya and maximum age of 69.53 Mya), (iii) MRCA of crown Nisitrini (minimum age of 40.05 Mya and maximum age of 67.15 Mya), and (iv) MRCA of crown *Eneoptera* Burmeister, 1838 (minimum age of 16.25 Mya and maximum age of 30.66 Mya). The best-fit partition scheme inferred by ModelFinder was used to determine the set of uncorrelated lognormal clocks for the dating analyses. The tree model was set to a birth-death speciation process (Gernhard 2008) to account more accurately for extinct and missing lineages. To limit the risk of over-parameterisation, we enforced a fixed topology corresponding to the tree with the best support (measured by the number of highly supported nodes for the BI and ML trees); here it is the best-score tree obtained under ML (see the Results section). This tree was further pruned under Mesquite (‘prune clade’ tool) to include just one specimen per species, and the molecular dataset was modified in the same way (specimens with the greatest gene coverage were given preference). Two independent BEAST analyses were carried out with analyses consisting of 50 million generations of MCMC, with the parameters and trees sampled every 5000 generations. The results of the two independent runs were combined with LogCombiner v.1.10.4 (part of the BEAST software package), after applying a burn-in period of 25%. The maximum credibility tree, median ages and their 95% HPD were generated with TreeAnnotator v.1.10.4 (part of the BEAST software package). Convergence of runs was assessed graphically and by examining the effective sampling size (ESS) of relevant parameters under Tracer v.1.7 (Rambaut et al. 2018).

## 2.4 | Historical Biogeography

Historical biogeography analyses were carried out using para-
metric methods (Ree and Sanmartín 2009), as implemented
in the *R* package BioGeoBEARS (Matzke 2014). The following
models were assessed: (i) dispersal– extinction cladogenesis
(DEC) model (Ree and Smith 2008), (ii) the likelihood imple-
mentation of the dispersal–vicariance analysis (DIVA) model
(Ronquist 1997; i.e., DIVAlike in BioGeoBEARS), and (iii) the
Bayesian Analysis of Biogeography (BayArea) model (Landis
et al. 2013; i.e., BAYAREAlike in BioGeoBEARS). All analy-
ses were carried out with and without the founder-event spe-
ciation parameter (+j) sensu Matzke (2014). The dated tree
from the BEAST analyses was used as a guide tree and pruned
to include only the 16 Xenogryllini species. Five geographical
areas were defined, which account for the five competing hy-
potheses presented in the Introduction: (A) Africa (excluding
Madagascar), (I) India, (P) Palearctic, (M) Madagascar, and (S) Southeast Asia (S). To better reflect extant distribution patterns, a maximum number of two areas was allowed (see Table S2). A time-stratified model was then defined, implementing a matrix of scaling factors for dispersal rates (DR) between areas (from 0 to 1) to account for the respective positions of the geographic areas through time while also accounting for potential geographical barriers to dispersal: (i) 1.0: adjacent areas with no barrier, (ii) 0.75: areas separated by minor barriers, (iii) 0.5: two areas separated by one area (without a barrier), (iv) 0.25: two areas separated by one area (with minor barriers), and (v) 0.1: two areas separated by one area (with major barriers). A time-stratified model was defined (see Table S3), with three time slices. Importantly, this model did not a priori favour an origin in Africa or in Asia. The first time slice (t1) runs from 56 to 33.9 Mya (Eocene), the second (t2) from 33.9 to 23.03 Mya (Oligocene), and the third (t3) from 23.03 to 0.1 Mya (from Miocene onwards). Following Kawahara et al. (2023), we added a last time slice (t0) from 0.1 to 0 Mya where all combinations of areas were allowed so that the adjacency matrix was less strict.

# 3 | Results

## 3.1 | Phylogenetic Relationships

The best-fit partition schemes and models are detailed in the Table S4. Both ML and BI analyses recovered all tribes as monophyletic with high support values (SH-aLRT ≥ 80%, uBV ≥ 95%, and PP ≥ 0.95), with the following relationships: ((Xenogryllini + Lebinthini) (Eneopterini + Nisitrini) Eurepini) (Figure 2; see Figure S1 for the BI tree). Within Xenogryllini, all species and genera are found to be monophyletic, generally with high support. *Xenogryllus* is sister to *Indigryllus*, with high support (SH-aLRT of 99.1%, uBV of 96%, and PP of 1.0); *Pseudolebinthus* is sister to the clade formed by *Xenogryllus* and *Indigryllus*, with high support (SH-aLRT of 97.8%, uBV of 100%, and PP of 1.0). Within *Xenogryllus*, the Asian species *X. maichauensis* Gorochov, 1992, *X. marmoratus*, *X. transversus* (Walker, 1869) and *X. ululiu* Gorochov, 1990 form a highly supported clade (SH-aLRT of 97.2%, uBV of 100%, and PP of 1.0), with *X. lamottei* Robillard, 2009 from West Africa found to be sister to this clade, with high support (SH-aLRT of 93.91%, uBV of 100%, and PP of 1.0). The African species, *X. eneopteroides* Bolívar, 1890, *X. maniema* Robillard and Jaiswara, 2009, *X. mozambicus* Robillard, 2009 form a clade along with *X. lemuria* from Madagascar (SH-aLRT and uBV of 100%, and PP of 1.0). The relationships within the genus *Pseudolebinthus* differ slightly between ML and BI analyses: *P. ntchisi* Robillard and Murphy (2020) is found sister to *P. whellani* Robillard and Desutter-Grandcolas (2006) under ML (SH-aLRT of 62% and uBV of 86%), while it is sister to the clade formed by *P. gorochovi* Robillard (2018), *P. lunipterus* Salazar et al. (2020), and *P. whellani* under BI analysis (PP of 0.52).

## 3.2 | Molecular Dating

The post-burn-in parameters of the combined runs from the BEAST analyses show ESS ≥ 200 for all relevant parameters. The age of the subfamily Eneopterinae is estimated to be

at c. 84.76 Mya (95% HPD: 96.75–74.88 Mya) during the Late Cretaceous (see Figure S2). The tribe Xenogryllini started diversifying in the Early Eocene c. 51.62 Mya (95% HPD: 60.16–43.92 Mya) (Figure 3). The crown age of *Pseudolebinthus* was dated in the Late Oligocene c. 23.4 Mya (95% HPD: 33.3–15.25 Mya). The clade formed by *Indigryllus* and *Xenogryllus* originated in the Early Oligocene c. 32.46 Mya (95% HPD: 38.8–27.01 Mya). The crown ages of *Xenogryllus* and *Indigryllus* were estimated to be at c. 26.03 Ma (95% HPD: 31.27–21.39 Mya) and c. 3.02 Mya (6.75–0.59 Mya), respectively.

## 3.3 | Historical Biogeography

Likelihood scores and weighted Akaike information criterion values indicate that +j models provide the best fit across all BioGeoBEARS analyses. The DIVALIKE+j model is recovered as the best-fitting model (LnL = -21.05; AIC weight = 0.61); the second-best model (LnL = -21.5; AIC weight = 0.39) is the BAYAREALIKE+j model, and yields results that are almost identical to the best model. Both models unequivocally support an origin in Africa for the MRCA of Xenogryllini (Figure 3; see also Figures S3–S5 and Table S5, for the raw outputs of all BioGeoBEARS analyses). From there, we infer a pattern of in situ diversification in Africa for the clade corresponding to the genus *Pseudolebinthus*, while a more dynamic pattern is inferred in the other clade, with two out-of-Africa dispersal events towards Asia. A first split led to the colonisation of Asia by a lineage that is nowadays represented by the two known species of *Indigryllus*. For this lineage, it is not possible to precisely determine the timeframe of colonisation of Asia due to the large difference between the crown (median age of 3.02 Mya) and the stem (median age of 32.46 Mya) age of the MRCA of *Indigryllus*. The other lineage is made up of representatives of the genus *Xenogryllus*, which in part pursued its diversification within Africa. In the Early Miocene (c. 19.95 Mya; 95% HPD: 24.4–15.95 Mya), a lineage of *Xenogryllus* colonised the Palearctic region from Africa, while *X. marmoratus* further expanded its range in Southeast Asia. Another dispersal to Asia is inferred for the clade including *X. maichauensis*, *X. ululiu*, and *X. transversus*, which diverged from *X. marmoratus* and colonised Southeast Asia during the Middle Miocene (c. 15.64 Mya; 95% HPD: 19.7–12.03 Mya), with *X. transversus* further expanding its range into India.

# 4 | Discussion

## 4.1 | Phylogeny and Systematics of Xenogryllini

Our study presents the molecular phylogeny of the tribe Xenogryllini based on a comprehensive sampling of all the extant species, including the recently described species from Madagascar (Yu et al. 2025). The relationships among the five eneopterine tribes are consistent with the results of the recent study focusing on Lebinthini by le Flanchec et al. (2025) and are highly supported (SH-aLRT ≥ 80%, uBV ≥ 95%, and PP ≥ 0.95), albeit for the grouping of Eneopterini and Nistrini for which the support is low (SH-aLRT of 4.3%, uBV of 42%, and PP of 0.73). This relationship between Eneopterini and Nistrini was also recovered with weak support in le Flanchec
<figure><img src="image_10.png" /><figcaption>**FIGURE 2** | Legend on next page.</figcaption></figure>
**FIGURE 2** | Maximum likelihood phylogeny of the tribe Xenogryllini based on the analysis of the 'specimen level dataset'. Branch support values for both ML and BI analyses are given at nodes (SH-aLRT/uBV/PP, in that order). Nodes with maximum support values (100% for SH-aLRT or uBV and 1.0 for PP) are highlighted by asterisks. Clades not recovered by the BI analysis are indicated by '.'. On the left, dorsal views of male specimens are presented for illustrative purposes (labels correspond to the species or the tribe in the phylogenetic tree).

<figure><img src="image_11.png" /></figure>

**FIGURE 3** | Reconstruction of the historical biogeography for tribe Xenogryllini based on the BEAST tree with outgroups removed, and estimated in BioGeoBEARS using the time-stratified DIVALIKE+j model with time slices divided by dashed lines (t1-t3). Geographic distributions are indicated by a colour scheme consistent with the map shown in Figure 1E. Coloured squares at the tips of terminal branches represent present-day distributions, while coloured rectangles at internal nodes denote inferred ancestral ranges based on the most likely reconstruction. Median ages and 95% HPD of ages are given at each node. The time period for all alternative hypotheses is given below the tree.

et al. (2025) (SH-aLRT of 24.8%, uBV of 82%, and PP of 0.82) and Jaiswara, Dong, and Robillard (2019) (standard bootstrap value ‘BV’ of 59% and PP of 0.56). Vicente et al. (2017) did infer an alternative placement for these two tribes with Eneopterini sister to a clade grouping Lebinthini, Nistrini, and Xenogryllini, and the support for it was slightly better (BV of 62% and PP of 1.0), which suggests that future studies on Eneopterinae with a denser sampling for all tribes and the inclusion of more markers will likely clarify this issue. Within Xenogryllini, the relationships we inferred between the three genera (*Pseudolebinthus* sister to *Indigryllus* + *Xenogryllus*) are highly supported and congruent with the results of previous phylogenetic studies that sampled fewer species (Jaiswara, Dong, and Robillard 2019; eight species; Vicente et al. 2017; le Flanchec et al. 2025: five species).

Within *Xenogryllus*, in a clade of three Asian species (*X. maichauensis*, *X. transversus*, and *X. ululiu*), we found *X.*

*transversus* sister (SH-aLRT of 37.3%, uBV of 72%, and PP of 0.71) to *X. ululiu*, consistent with the results of the ML analysis of Jaiswara, Dong, and Robillard (2019) which has a higher support (BV of 80%); however, it is noteworthy that their BI analyses gave a different topology, placing *X. transversus* sister to *X. maichauensis* and *X. ululiu* with a low support (PP of 0.73). Interestingly, our study unravels with a high support (SH-aLRT of 93.9%, uBV of 100%, and PP of 1.0) that the sister species to this Asian clade is *X. lamottei*, which is endemic to Guinea in West Africa. This relationship is supported by multiple morphological characters such as the shape of male genitalia and the absence of lateral carina on the pronotum (Jaiswara, Dong, Ma, et al. 2019).

The *Xenogryllus* species from Madagascar is recovered with high support (SH-aLRT and uBV of 100%, and PP of 1.0) sister to a clade of three African species (*X. eneopteroides*, *X. maniema*, and *X. mozambicus*). *Xenogryllus leuria* shares some similar
characters of pronotum and male genitalia compared with other African species in this clade, but is distinctly different from *X. lamottei* and the Asian *Xenogryllus* species (see Yu et al. 2025 for details).

## 4.2 | Origin of Xenogryllini in Africa: Where and When?

Our historical biogeography analyses unequivocally support an origin of the Xenogryllini tribe in Africa, dating back to the Middle Paleocene to the Middle Eocene (c. 60.16–43.92 Mya). Back then, the African continent was markedly different from what it is today. The latitudinal paleo-position of Africa was 15° further south (the African plate was tilted and moving counterclockwise) and, in northern Africa, parts of the continent were also largely submerged under the epicontinental Sahara Sea (Couvreur et al. 2021). At that time, most of continental Africa was low in elevation, and at mid-latitudes, the climate was warm and humid, especially during three major climatic optimums: (i) the Paleocene–Eocene Thermal Maximum (PETM) (c. 56 Mya), (ii) the Early Eocene Climatic Optimum (EECO; 53–51 Mya), and (iii) the Mid–Eocene Climatic Optimum (MECO; c. 40 Mya) (Couvreur et al. 2021). Although it has been hypothesized that a pan-African rain-forest extended from western to eastern Africa during this period (Axelrod and Raven 1978; Morley 2000, 2007; Tolley et al. 2013), in eastern Africa, fossil evidence is more consistent with the presence of more arid environments with open woodlands (Jacobs 2004; Jacobs and Herendeen 2004; Linder 2017). In southern Africa, paleoclimate reconstructions based on palynofloras support the hypothesis that environmental conditions were largely unaffected by the successive global warming events, and that the local climate was more temperate and drier (Korasidis et al. 2022). Assuming that the ecological preferences of Xenogryllini have been conserved over time (phylogenetic niche conservatism; Harvey and Pagel 1991; Wiens and Graham 2005) and are similar to those of extant species (e.g., Jaiswara, Dong, and Robillard 2019; Salazar et al. 2020), we can posit that early representatives of the tribe likely inhabited open habitats (as extant *Xenogryllus* spp.; Jaiswara, Dong, Ma, et al. 2019) or in forest edges (as extant *Indigryllus* and *Pseudolebinthus* spp.; e.g., Robillard and Murphy 2020; Salazar et al. 2020; Jaiswara et al. 2022), the latter is thus more consistent with an origin in southern or eastern Africa, where open or mixed habitats were available and possibly prevalent. A parallel can be drawn here with the results of several studies on Afrotropical insect groups, which have inferred an origin in southern and/or eastern Africa for lineages found predominantly in open habitats (e.g., Eberle et al. 2017; Kamiński et al. 2022; Hévin et al. 2023, 2024).

Regarding the origin of the extant PID pattern, both the ‘Indian raft’ and ‘Kohistan–Ladakh Island Arc’ hypotheses—associated with biogeographic events occurring in the Late Cretaceous or in the Paleocene—can be rejected with some certainty, given that Xenogryllini only began their diversification between the Middle Paleocene to the Middle Eocene, and that ancestral area reconstructions indicate that, from their origin until at least the Oligocene, Xenogryllini were distributed exclusively in continental Africa.

## 4.3 | Out-of-Africa Pattern for a Lineage of *Xenogryllus*

Of the three extant Xenogryllini genera, *Xenogryllus* has by far the widest distribution, being present in both Africa and Asia. This is possibly because *Xenogryllus* species have a greater dispersal capacity, as it is the only genus of Xenogryllini with long wings and living in open habitats such as savannas, unlike *Indigryllus* and *Pseudolebinthus*, which are brachypterous and live in forest edges. Within *Xenogryllus*, we identified one split leading to a lineage that is nowadays exclusively distributed in Asia (including India and parts of Eurasia in the East Palearctic; see Figure 1A for details).

Our inferred temporal framework (dispersal occurring between 24.4 and 12.03 Mya, based on 95% HPD intervals) is consistent with the ‘Gomphotherium landbridge’ hypothesis, which posits that the collision of the African and Arabian plates with Eurasia resulted in the establishment of a dispersal corridor that opened in the Burdigalian (20.44–15.97 Mya) and lasted until the Serravallian (13.82–11.63 Mya) (Rögl 1998). Following this period, the spread of hyperaridity in northern Arabia impeded the dispersal of organisms not adapted to hyperarid conditions, although evidence of recurrent humid intervals in Arabia has been recently unravelled (Markowska et al. 2025). For organisms that are more adapted to open habitats, the effectiveness of this landbridge was also enhanced by the marked environmental changes that followed the Middle Miocene Climatic Optimum (MMCO; 17–15 Mya; Zachos et al. 2001). Indeed, the sudden drop of temperatures in association with the eastern Africa uplift led to aridification and the progressive opening of habitats (Trauth et al. 2009; Couvreur et al. 2021), which facilitated dispersal between Africa and Asia in numerous groups of organisms (e.g., Zhou et al. 2012; Yu et al. 2014; Pokorny et al. 2015; Thomas et al. 2015; Bourguignon et al. 2017; Rossini et al. 2022; Hévin et al. 2024). Here, it should also be added that the hypothesis of a transoceanic LDD colonisation event, although theoretically conceivable, is not supported by our biogeographical reconstructions.

Our biogeographic reconstructions suggest that this out-of-Africa pattern first involved dispersal to the Palearctic in the Early Miocene (Figure 4: 1A), followed by eastward dispersal to Southeast Asia during the Miocene (Figure 4: 1B), and a final westward range expansion towards India for *X. transversus* (Figure 4: 1C). The inferred eastward dispersal towards Southeast Asia instead of a southward dispersal towards India is probably explained by the orogenesis of the Himalayas, which would have constituted a major barrier to dispersal. This is consistent with the fact that the Himalayas uplifted rapidly from the Middle Miocene onwards (c. 20 Mya; Ding et al. 2022), thus limiting biotic exchanges towards lower latitudes (e.g., Zhou et al. 2012; Nie et al. 2013). Open habitats were also spreading in Asia during the Late Miocene to the Pliocene (Morley 2018), which probably promoted the eastward dispersal of *Xenogryllus*. The final range expansion that is inferred towards northern India occurred relatively recently (either in the Pliocene or Pleistocene), when tropical forests receded in northern India (Morley and Morley 2022), which may have promoted the range expansion of *Xenogryllus* there.
<figure><img src="image_12.png" /><figcaption>**FIGURE 4** | Putative colonisation routes based on the results of the historical biogeography analyses within *Xenogryllus*: 1A, initial dispersal from Africa to the Palearctic through the *Gomphotherium* landbridge in the Early Miocene; 1B, further colonisation of Southeast Asia in the Miocene; 1C, late range expansion in Northern India for *X. transversus*. For the lineage leading to *Indigryllus*: 2, Trans-Tethyan dispersal event at the end of the Eocene followed by southward dispersal. 2', Transoceanic LDD event (Africa-to-India) that could have occurred at any time between the Early Oligocene to the Pliocene. For the lineage leading to *Xenogryllus lemuria*: 3, dispersal to Madagascar that could have occurred at any time since the Early Miocene. Paleomap reconstruction was made using GPlates (with an age set at 15.97 Mya and apparent plate boundaries) with open-source data from the EarthByte alternative plate reconstructions (Merdith et al. 2021). An adult of the brachypterous species *Pseudolebinthus lunipterus* is shown on the bottom right (picture by K. Salazar).</figcaption></figure>

## 4.4 | Out-of-Africa: The Case of *Indigryllus*

The other out-of-Africa dispersal event inferred by our historical biogeography analyses concerns the lineage that is nowadays represented by the genus *Indigryllus*. This genus has been only described recently (Jaiswara, Dong, and Robillard 2019), and it consists of at least two species (another potential new species was also spotted through citizen science in the Northwest coast of India; Jaiswara, Dong, and Robillard 2019). There is much more uncertainty for this colonisation event, as it could have occurred at any time between the inferred crown (median age of 3.02 Mya; 95% HPD: 6.75–0.59 Mya) and stem (median age of 32.36 Mya; 95% HPD: 38.8–27.01 Mya) ages of *Indigryllus*. That being said, the estimated crown age of *Indigryllus* may also be underestimated if we consider the possibility that more species have yet to be discovered, not to mention the fact that extinct lineages could have been stem lineages, both of which could make the age of the MRCA of the genus much older.

Our estimated timeline is partially compatible with a colonisation of Asia during the Middle to Late Eocene, in line with the Trans-Tethyan hypothesis. However, this scenario implies an

intermediate colonisation of the Palearctic (Figure 4: 2A), which we did not infer, followed by a southward dispersal (Figure 4: 2B), which could only have occurred in a very short time window given that the split between *Xenogryllus* and *Indigryllus* occurred during the Eocene/Oligocene boundary.

The alternative *Gomphotherium* landbridge* hypothesis falls within the wide time interval leading to *Indigryllus*, but it is not supported by the biogeographic reconstructions. This hypothesis is only conceivable if we postulate that extinct lineages have followed the same colonisation path as the one we have inferred for the Asian *Xenogryllus* lineage. However, such a hypothesis seems rather unlikely, given that the two groups differ in their ability to disperse and present a very distinct distribution pattern in India, with, on the one hand, a wide distribution in Northern India for *X. transversus* compared to a very localised distribution in Southwest India, near the coast, for *Indigryllus*.

Lastly, the colonisation of Asia by *Indigryllus* could correspond to a transoceanic LDD event between East Africa and the western coast of the Indian subcontinent (Figure 4: 2'). Transoceanic LDD dispersal was hypothesised as an explanatory hypothesis for PID patterns in the Cenozoic, for several plant (e.g., Renner and Meyer 2001; Malcomber 2002; Zhou et al. 2012) vertebrate (e.g., Jansa et al. 1999), and insect (e.g., Condamine et al. 2013; Toussaint et al. 2019) lineages. Although one may think that such an event is implausible due to the fact that *Indigryllus* individuals are brachypterous, it is worth highlighting that one of the largest radiations of eneopterine crickets is those of Lebinthina, a subtribe composed of only brachypterous species (Tan et al. 2021) that managed to colonise most of the archipelagos in Asia and in the Southwest Pacific region (e.g., New Caledonia, New Guinea, Philippines, Solomon Islands, and Vanuatu). Considering the latter, the results of our biogeographic reconstructions and also the fact that *Indigryllus* is only known from the coastal areas of Southwest India, it is suggested that this hypothesis is currently the best explanation for this particular PID pattern.

## 4.5 | Colonisation of Madagascar

Madagascar is a well-known biodiversity hotspot, with high levels of endemicity (Myers et al. 2000). Numerous studies have investigated the origins of Madagascar's flora and fauna, and reviews of this question (Yoder and Nowak 2006; Buerki et al. 2013; Antonelli et al. 2022) have revealed that most of its extant biodiversity was built upon Cenozoic dispersers originating mainly from Africa. In insects, the inferred timing and the colonisation routes of Madagascar are consistent with this general finding (e.g., Eberle et al. 2017; Eliassen and Jordal 2021; Kamiński et al. 2022; Wang et al. 2023; Hévin et al. 2023, 2024), with most studies unravelling the Africa to Madagascar pattern (but see also Condamine et al. (2013) or Forthman and Weirauch (2016) for counterexamples of India-to-Madagascar colonizations).

The results of biogeographic analyses recover a continental Africa origin for the colonisation of Madagascar (Figure 4: 3), with dispersal occurring from 13.45 Mya onwards (based on 95% HPD intervals of age estimates). This timeline is in line with the potential existence of shallow landbridges in the Mozambique
Channel between the Late Miocene and the Early Pliocene (12–5 Mya; Masters et al. 2021) and the presence of ephemeral islands acting as potential stepping-stones from the Pliocene onwards (Warren et al. 2010), both of which may have facilitated the colonisation of Madagascar.

## 5 | Conclusion

This study constitutes the most comprehensive phylogenetic study on Xenogryllini, encompassing all known species of the tribe. Our historical biogeography analyses allow us to determine that the tribe originated in Africa between the Middle Paleocene and the Middle Eocene, and that the extant PID pattern for Xenogryllini can be explained by two independent out-of-Africa dispersal events. Although it seems more likely to postulate that one lineage relied on the *Gomphotherium* land-bridge* route and that the other colonised Asia via transoceanic LDD, it is worth stressing that the inferred patterns may have been obfuscated by extinction events, and that—as highlighted by the recent discoveries of the genus *Indigryllus* and of a new *Xenogryllus* species in Madagascar—the possible discovery of new Xenogryllini species may well change our comprehension of their biogeographical history.

### Acknowledgements

We want to thank the editor Dr. Hege Värdal and two anonymous reviewers for their constructive comments on a previous version of the manuscript. We thank G. Cowper (ANSP) for his help during the visit of T. Robillard in Philadelphia. We are also indebted to K. Salazar for the picture of *Pseudolebinthus lunipterus* and to the following persons who gave us permission to use pictures posted on iNaturalist: K.W. Chan, S. Sreebin, and B. Wursten. We thank the personnel of the molecular platform of the 'Service de Systématique Moléculaire', UMS 2700 2 AD, MNHN, CNRS for their help during experimentation.

### Conflicts of Interest

The authors declare no conflicts of interest.

### Data Availability Statement

The data that supports the findings of this study are available in the Supporting Information—S1 of this article.

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**Supporting Information**

Additional supporting information can be found online in the Supporting Information section. **Data S1:** zsc70019-sup-0001-Figures.docx. **Data S2:** zsc70019-sup-0002-Tables.xlsx.