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\begin{document}

\title{Phylogenetic relationship and biogeography of the genus Diestramima, a
cave cricket endemic to the Oriental realm (Rhaphidophoridae:
Aemodogryllinae)}



\author[1]{Jingsong Zong}%
\author[2]{Yanyan Qin}%
\author[3]{Chuze Shen}%
\author[1]{Zhu-Qing HE}%
\author[1]{Kai Li}%
\affil[1]{East China Normal University}%
\affil[2]{Hunan University}%
\affil[3]{Beijing Normal University}%


\vspace{-1em}



  
  \date{May 22, 2024}


\begingroup
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\selectlanguage{english}
\begin{abstract}
Diestramima is only distributed in the Oriental realm, and mainly in
China. Because they are apterous, these insects are good materials for
studying biogeography. In this study, we reconstruct the phylogeny tree
of Diestramima species based on maximum likelihood and Bayesian
inference based on six genes, and use fossil-based molecular dating and
ancestral range estimation to reconstruct the dispersal route. The
results of molecular system are highly consistent with the morphological
characteristics of these species. The distribution area of a species has
an important influence on their interspecific differentiation. Dating
and historical biogeography analyses suggest an early Miocene origin of
the genus. Some ancestors of the genus Diestramima have been distributed
in Guangxi, Yunnan province and other regions of China at this time. In
the late Miocene, the genus Diestramima dispersed from the low mountain
and hilly areas of Guizhou, Guizhou and Hunan to inland China. With the
uplift of the Qinghai-Tibet Plateau and the drop in temperature in the
Neogene leading to the divergence within this genus. Diestramima
gradually spread from Guangxi and Yunnan to Chinese inland area.
Quaternary glaciation determines their current geographical distribution
pattern. Southeast Asia is the most likely original place of the genus
Diestramima.%
\end{abstract}%



\sloppy


\textbf{Introduction}

Cave cricket (Rhaphidophoroidea Walker, 1869) as one of the oldest
species of Orthoptera, are widely distributed all over the world. It
considered to be the basis of the six families and one of the oldest
taxa in the Tettigoniidea clade (Song \emph{et al} ., 2020). They are
gregarious and typical apterous groups, which includes 10 subfamilies
and 87 genera (including 1 fossil subfamily and 2 fossil genera). They
mainly inhabit in dry leaves, stone crevices, caves and other dark humid
environments, therefore their speciation as well as population structure
have relatively strong dependence on geographical isolation
(Juan\emph{et al} ., 2010).

Dating estimates indicated that Rhaphidophoridae originated in the
Cretaceous period during the Mesozoic era with the ancestral area
located both in the northern and southern hemisphere. The current
distribution of Rhaphidophoridae might be explained by a combination of
both dispersal and vicariance events occurred especially in the
ancestral populations. The radiation of Rhaphidophoridae started within
the Pangaea, where the ancestor of Rhaphidophoridae occurred throughout
an ancestral area including Australia, North America, and the
Mediterranean region. The opening of the Atlantic Ocean promoted the
divergence of North American and Mediterranean lineages while the
differentiation of the southern lineages, spread from Australia, appears
to be related to the fragmentation of Gondwana land (Allegrucci \&
Sbordoni, 2019). The Indian subcontinent (including India and southeast
Asian countries north of the Indian river) was originally part of
Gondwana continent, and the initial collision between it and Eurasia
probably began in the late cretaceous, and the main collision period
occurred around 55--50Ma, which coincided with the isolation of the
Aemodogryllinae and Rhaphidophorinae. After the collision of the Indian
subcontinent and the Eurasian continent, insects from the two continents
met again here, which also provided rich raw material for later Oriental
insect species (Jia, 2010).

China straddles Palaearctic and Oriental realm, occupies an important
biogeographical position in the world. Recent years, many papers on
molecular biogeography of insects have been carried out in China. Based
on the phylogenetic relation and distribution of four grasshopper genera
of Catantopidae from the south of China, ten areas of endemism have been
recognized (Xu \emph{et al} ., 2003). The order of the formation of the
10 distribution areas of these insects was as follows: southern Yunnan
and Loess Plateau, Hainan Island, eastern Mountain and Taiwan Island;
The Yunnan-Guizhou plateau, northern Yunnan and Qinling-Daba mountains
formed at the same time, while the southern China and jiangnan hilly
region formed the last. Wang \emph{et al} . (2020) determined the
phylotaxonomic relationship and differentiation time of the
genus\emph{Sinocyrtaspis} endemic to China. Based on the geographic
dynamics and climatological data of its ancestors, it was concluded that
the centres of origin of \emph{Sinocyrtaspis} were Hunan and north-east
Guangxi. Their ancestors had two dispersal routes ending in Jiangxi and
Zhejiang, after the cold period of the Miocene, some of the ancestral
species remained in Guizhou while others moved out. And as a response to
climate change, species altered distribution areas by moving along
altitude gradients in mountain regions, whereas the anthropogenic global
warming trend has promoted some species moving to high-latitude areas or
caused population differentiation. However, there is still a lack of
biogeographic research on the widely distributed Oriental species,
especially the Rhaphidophoridae in Oriental realm.

China is the main distribution area of Rhaphidophorinae Walker, 1869 and
Aemodogryllinae Jacobson, 1905. \emph{Diestramima} Storozhenko, 1990 is
(Rhaphidophoroidea: Aemodogryllinae, Kevan, 1977) widely distributed in
the Oriental realm of China (with 70\% of known species) and adjacent
countries. Their range is limited to the Oriental realm (tropical and
subtropical mountainous hilly areas), because they usually choose wide
shrub leaves as mating sites. All species of this genus has long
protuberances on the seventh abdominal tergite, and the shape of the
protuberances is significantly different among different species. The
morphological classification system and the relationship between species
are relatively clear. Therefore, \emph{Diestramima} is a suitable
candidate group for studying the speciation and diversity patterns of
Oriental realm in China and the corresponding geological changes.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image1/image1}

\end{center}
\end{figure}

In this paper, we used an integrative approach to identify species,
including morphology and multiple methods for molecular identification,
which clarified the present taxonomic problems on \emph{Diestramima} .
We estimated the temporal and spatial divergence of this genus, and
present a historical framework for the phylogeny and biogeography of
these Oriental realm endemic cave crickets. And in the discussion, we
try to predict the historical dispersal pattern of pure Oriental species
in China from this groups.

\textbf{Fig. 1.} Distribution areas of the genus \emph{Diestramima} in
China.

\textbf{Material and methods}

\emph{Taxon sampling}

A total of 87 individuals from 19 species were collected by hand or
sweep net at night. Adults were preserved in 70\% ethanol in the field.
The morphological identifications prior to DNA extraction were made
depanding on the male apex of abdomen and paraproct (Gorochov \emph{et
al} ., 2015). Samples of genus \emph{Diestramima} were collected from
twelve provinces in China Oriental realm, including Anhui, Chongqing,
Fujian, Guangdong, Guangxi, Guizhou, Hubei, Hunan, Jiangxi, Sichuan,
Yunnan and Zhejiang (Fig. 1). All widespread species in China and a
cross-border species distributed in China and Vietnam have been
collected. Moreover, we examined a large of additional specimens and
relevant literature (Storozhenko, 1990; Gorochov, 1994, 2010; Liu \&
Zhang, 2001; Gorochov \& Storozhenko, 2015; Qin \emph{et al} ., 2016;
Zhu \& Shi, 2018; Wang \emph{et al} ., 2019) of Rhaphidophoroidea for
better understanding the morphological variations of species, to aid in
accurate species determination. Thus, the present sampling is the almost
comprehensive one for the phylogenetic analysis in terms of both
taxonomy and distribution. All diagnostic characters were photographed
by OLYMPUS DP27, and arranged using Adobe Photoshop CC (Fig. 2, Fig. 3).\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image2/image2}

\end{center}
\end{figure}

\textbf{Fig. 2.} Male abdomen apex of \emph{Diestramima} in dorsal view.
(A) \emph{D} . \emph{acutiapicis} , (B) \emph{D} . \emph{austrosinensis}
, (C) \emph{D} . \emph{beybienkoi} , (D) \emph{D} . \emph{brevis} ,
(E)\emph{D} . \emph{conica} , (F) \emph{D} . \emph{cycla} , (G) \emph{D}
.\emph{eurya} , (H) \emph{D} . \emph{excavate} , (I) \emph{D}
.\emph{guangxiensis} , (J) \emph{D} . \emph{intermedia} , (K) \emph{D}
.\emph{lamina} , (L) \emph{D} . \emph{major} , (M) \emph{D}
.\emph{pingmengensis} , (N) \emph{D} . \emph{gulinjingensis} , (O)
\emph{D} .\emph{subrectis} , (P) \emph{D} . \emph{subtilis} , (Q)
\emph{D} .\emph{triangulate} , (R) \emph{D} . \emph{truncate} , (S)
\emph{D} .\emph{yunnanensis} . Scale bar: 1.0 mm.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image3/image3}

\end{center}
\end{figure}

\textbf{Fig. 3.} Male abdomen apex of \emph{Diestramima} in lateral
view. (A) \emph{D} . \emph{acutiapicis} , (B) \emph{D}
.\emph{austrosinensis} , \emph{D} . (C) \emph{beybienkoi} , (D) \emph{D}
.\emph{brevis} , (E) \emph{D} . \emph{conica} , (F) \emph{D} .
\emph{cycla} , (G) \emph{D} . \emph{eurya} , (H) \emph{D} .
\emph{excavate} , (I) \emph{D} .\emph{guangxiensis} , (J) \emph{D} .
\emph{intermedia} , (K) \emph{D} .\emph{lamina} , (L) \emph{D} .
\emph{major} , (M) \emph{D} .\emph{pingmengensis} , (N) \emph{D} .
\emph{gulinjingensis} , (O) \emph{D} .\emph{subrectis} , (P) \emph{D} .
\emph{subtilis} , (Q) \emph{D} .\emph{triangulate} , (R) \emph{D} .
\emph{truncate} , (S) \emph{D} .\emph{yunnanensis} . Scale bar: 1.0 mm.

\emph{DNA extraction, amplification and sequencing}

The total genomic DNA was extracted from the muscles of one hind leg by
AxyPrep Genomic DNA Miniprep Kit (AXYGEN), according to the
manufacturer's instructions. We used previously published sequences from
the family Rhaphidophoroidea to jointly complete this sampling,
including six species of the genus \emph{Diestrammena} Brunner von
Wattenwyl, 1888\emph{,} four species of the genus
\emph{Rhaphidophora}Serville, 1838, twenty species of the subfamily
Dolichopodainae Brunner von Wattenwyl, 1888, nineteen species of the
subfamily Troglophilinae Krauss, 1879 (Allegrucci \& Sbordoni, 2019).
The samples and sequence information were listed in Table S1. The
following six gene regions were amplified: Mitochondrial ribosomal 16S,
nuclear ribosomal 28S, mitochondrial cytochrome oxidase I (COI), ATPase
catalytic subunit A-like (ACSL), cytoplasmic-like (CPL) and water
dikinase-like (WDL). PCR was performed using 2\selectlanguage{ngerman}×Es Taq Master Mix (Dye)
manufactured by Cowin Bio (China), with a final volume of 50\selectlanguage{greek}μ\selectlanguage{english}L, composed
of: 25\selectlanguage{greek}μ\selectlanguage{english}L of the 2\selectlanguage{ngerman}×Es Taq Master Mix (Dye), 19\selectlanguage{greek}μ\selectlanguage{english}L of the ddH2O, 2\selectlanguage{greek}μ\selectlanguage{english}L of
forward and reverse primer each and 2\selectlanguage{greek}μ\selectlanguage{english}L of the sample DNA extract. The
PCR annealing temperature for each gene was modified (Table S2). PCR
products were directly sequenced.

\emph{Phylogenetic analyses}

We used all \emph{Diestramima} samples as well as the samples of
adjacent genera \emph{Diestrammena} and \emph{Raphidophora} in the
phylogenetic tree construction. For each fragment, DNA alignment was
conducted by MEGA7 software (Loytynoja \& Goldman 2010; Kumar \emph{et
al} . 2016). Saturation level of sequences was verified by Xia's test
(Xia \emph{et al} ., 2003) in dambe 7 (Xia, 2018). In PhyloSuite
(Zhang\emph{et al} . 2020), the best-fit evolutionary model GTR was
derived from ModelFinder (Kalyaanamoorthy \emph{et al} . 2017), and the
Maximum likelihood (ML) analysis was performed in IQ-Tree (Nguyen
\emph{et al} . 2014) with 1000 ultrafast bootstrap replicates. Node
support is shown as posterior probabilities and bootstrap value, as
directly estimated from the majority rule consensus topology.

\emph{Divergence time estimation}

The phylogenetic of \emph{Diestramima} using BEAST v.1.8.4 (Drummond \&
Rambaut, 2007) to reconstruct by selecting the Birth-Death tree prior
and the Piecewise linear and constant root of population size model.
Given the absence of reliable fossil records for \emph{Diestramima} , we
relied on secondary calibrations based on the available worldwide dated
framework of Rhaphidophoroidea (Allegrucci \& Sbordoni, 2019), we used
representatives of four lineages (subfamily Dolichopodainae and
Troglophilinae; genus \emph{Diestrammena} and \emph{Rhaphidophora} )
that were sampled in this study, including 49 cave cricket species. In
their conclusion, the ancestor of the Aemodogryllinae and
Rhaphidophorinae spread from Australia to south-eastern Asia at 117Ma
(95\% HPD: 105--130 Ma), an estimate consistent with plate tectonics as
India separated from other parts of Gondwana at 130 Ma, prior to other
paleogeographic events (Chatterjee \& Bajhai, 2016). Therefore, in our
study, it was used as the first divergent calibration time to prove the
purity of these Oriental realm species.

The divergence times of major nodes were estimated under uncorrelated
relaxed clock by setting four fossil calibrations: (1) due to the plate
tectonics as India separated from other parts of Gondwana, the ancestor
of the Aemodogryllinae and Rhaphidophorinae spread from Australia to
south-eastern Asia at 117 Ma (Allegrucci \& Sbordoni, 2019); (2) the
opening of Mid-Aegean trench (9--12 Ma) separated the\emph{Troglophilus}
Krauss, 1879 species inhabiting the eastern Aegean islands and Asia
Minor from the Cyclades, Crete and mainland Greece (Allegrucci \emph{et
al} ., 2017); (3) the uplift of Anatolian Plateau happened 5--10 Ma due
to acceleration of northward movement of the Arabian Plate (Steininger
\& R\selectlanguage{ngerman}ögl, 1984) and separating the Anatolian species of
\emph{Troglophilus} from the Balkans ones; (4) the isolation of Crete
island, inhabited by\emph{Dolichopoda} \emph{paraskevi} Boudou-Saltet,
1973 (Aegean Sea, Greece) from neighboring mainland, happened at the end
of Messinian salinity crisis (5.33 Ma, Krijgsman \emph{et al} ., 1999).
Trees were sampled every 10000 generations for a total of 100 million
generations. We further checked whether the effective sample size (ESS)
was greater than 200 for most important parameters using Tracer v.1.7.1.
The consensus tree was exported using TreeAnnotator v.1.8.4
(Drummond\emph{et al} ., 2012), discarding the first 25\% of trees as
burn-in. The mean node heights and 95\% highest probability density
(95\% HPD) were displayed in FigTree v1.4.4.

\emph{Ancestral range estimation}

We defined the following six biogeographical areas of
\emph{Diestramima}according to their known distributions and the
zoogeographic regionalization of China (Zhang \emph{et al} ., 2011): A.
mountainous region of southern Yunnan (Baoshan city, Xishuangbanna Dai
autonomous prefecture, south of Ailaoshan in Yuxi city, Puer city,
Honghe Hani \& Yi autonomous prefecture, west of Wenshan Zhuang \& Miao
autonomous prefecture); B. mountainous hilly region of Yunnan and
Guangxi (east of Wenshan Zhuang \& Miao autonomous prefecture, Baise
city, Chongzuo city, Fangchenggang city, Nanning city); C. low
mountainous hilly region of Guangxi, Guizhou and Hunan (Hechi city,
Liuzhou city, Guilin city, south of Qiannan Buyi \& Miao autonomous
prefecture, east of Qiandongnan Miao \& Dong autonomous prefecture, east
of Hunan province); D. Sichuan basin (most of Chongqing city and east of
Sichuan province); E. Jiangnan hilly region (southeast of Hunan,
Xianning city in Hubei, most of Jiangxi, north of Fujian, most of
Zhejiang); F. mountainous hilly region of Fujian and Guangdong (east of
Guangxi province, north of Guangdong province, south of Fujian
province).

Ancestral ranges were reconstructed using RASP 4.0 (Yu \emph{et al} .,
2020) beta. As a guide tree for statistical
dispersal-extinction-cladogenesis (S-DEC; Beaulieu \emph{et al} ., 2013)
analyses, we removed all outgroups and constructed a phylogenetic tree
for the genus \emph{Diestramima} . Modify and guide this tree according
to the maximum credibility tree from the BEAST analysis. Ancestral areas
were limited to no more than three areas. Statistical
dispersal-vicariance (S-DIVA; Yu \emph{et al} ., 2010) analysis were
performed based on a sample of 1000 random post burn-in RASP trees and
use the final tree. The maximum number of ancestral areas was limited to
three.

\textbf{Result}

\emph{Species delimitation and phylogeny}\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image4/image4}

\end{center}
\end{figure}

Phylogenetic trees (Fig. 4) constructed either by BI or ML analyses had
the same topology. Between species, Posterior probability (PP) of all
nodes is 1 and bootstrap support values (UFB) are all above 98
except\emph{D} . (\emph{D} ). \emph{eurya} is 94. Based on different
tree-constructing methods as well as single or combined dataset, the
monophyly of the species in \emph{Diestramima} was well-supported. The
membership of each of these major clades were not strongly correlated
with the distribution of their subgenera (\emph{Baculitettix}
,\emph{Diestramima} and \emph{Excisotettix} , Gorochov \emph{et al} .,
2019). The species within each clade live in roughly the same area,
especially in the southwestern border region of China. \emph{D}
.\emph{conica} , \emph{D} . \emph{yunnanensis} . \emph{D.} sp. 2,
\emph{D} .\emph{major} , \emph{D} . \emph{subtilis} , \emph{D} .
\emph{cycla} and\emph{D} . \emph{triangulata} are in a clade, although
they contain two subgenera. In the southwestern border region of China,
even within the same species, samples from different regions showed
great variability. We collected \emph{D} . \emph{conica} samples from
Taiyanghe Reserve, Gaoligongshan and Laifengshan, and the samples from
these three regions in Yunnan showed a high divergence. Similarly,
\emph{D} . \emph{subtilis}samples from three regions (Xiaojie town,
Gulinjing town and Ailaoshan) in Yunnan also produced this high
divergence. However, widely distributed species in inland areas of China
always tend together. We collected \emph{D. beybienkoi} samples from
seven regions (Jinyunshan, Emeishan, Zhangjiajie, Dujiangyan,
Leigongshan, Simianshan and Yangmeiao) in five provinces (Sichuan,
Chongqing, Hunan, Guizhou and Guangxi) of China, but they only
divergence two clades. This divergence phenomenon is more prominent in
\emph{D} . \emph{intermedia} and\emph{D} . \emph{austrosinensis} .
Although the samples we collected spanned several provinces and regions
in China, there were basically no divergence within their species.

\textbf{Fig. 4.} Phylogenetic relationships of the
genus\emph{Diestramima} . Maximum likelihood (ML) phylogenetic tree
based on the concatenated 12S, 16S, \emph{COI} , ACSL, CPL and WDL
matrix sequences. Topology supports of all major nodes are indicated
above branches in this order: the posterior probability and the
bootstrap resampling. Vertical lines of different colors represent
different subgenera: \emph{Diestramima} in dark blue;
\emph{Baculitettix} in dark green; \emph{Excisotettix} in dark red. The
color bar represents the six major biogeographical areas, letter
abbreviations represent collection areas: Taiyanghe Reserve (TYH),
Gligongshan (GLGS), Laifengshan (LFS), Xishuangbanna Tropical Botanical
Garden (XTBG), Gulinjing (GLJ), Daqingshan (DQS), Xiaojie (XJ),
Ailaoshan (ALS), Daweishan (DWS), Fenshuiling (FSL), Leigongshan (LGS),
Dujiangyan (DJY), Jiugongshan (JGS), Fanjingshan (FJS), Zhangjiajie
(ZJJ), Maoershan (MES), Emeishan (EMS), Wuyishan (WYS), Wangdongyang
(WDY), Yangmeiao (YMA), Lianhuashan (LHS), Jinyunshan (JYS), Simianshan
(SMS), Dapanshan (DPS), Ganjiangyuan (GJY), Fushoushan (FSS), Huangshan
(HS), Tiantongshan (TTS), Gutianshan (GTS), Baishanzu (BSZ), Nanling
(NL), Nonggang (NG), Pingmeng (PM), Hekou town (HKT).

\emph{Divergence time estimation}

The divergence time tree is divided into two branches: Aemodogryllinae
and Rhaphidophorinae from Asia, Dolichopodainae and Troglophilinae from
Europe. We analyze the Asian taxa separately (Fig. 5). The BEAST
analysis indicated that the subfamily Aemodogryllinae and
Rhaphidophorinae have been diverged since Cretaceous. The speciation
events in genus \emph{Diestramima} occurred from late Miocene to the
early Pliocene, and most species was completed in or before early
Quaternary. Following the speciation of four Yunnan and Guangxi lineages
(\emph{D} . \emph{conica} , \emph{D} . \emph{major} , \emph{D}
.\emph{subtilis} and \emph{D} . \emph{yunnanensis} ) diverged at 8.32 Ma
(95\% HDP: 4.84--12.88 Ma), two clades split at 3.66 to 8.96 Ma. Both
clades include species from Yunnan and Guangxi (border lineages) as well
as those from inland China (inland lineages), and they further diverged
at 5.84 Ma (95\% HDP: 3.66--8.96 Ma) and 5.85 Ma (95\% HDP: 3.67--9.10
Ma) respectively. In one clade, \emph{D} . sp. 2, a new species we
discovered on the China-Vietnam border, was diverged at 4.02 Ma (95\%
HDP: 2.42--6.31 Ma). At this point the border and inland lineages began
to diverge. The widely distributed species \emph{D} .
\emph{intermedia}was diverged at 1.02 Ma (95\% HDP: 0.44--1.86 Ma). In
the other clade, border and inland lineages was diverged with \emph{D} .
sp. 1 and\emph{D} . \emph{lamina} at 3.52 Ma (95\% HDP: 2.18--5.29 Ma).
Two widely distributed species, \emph{D} . \emph{austrosinensis} and
\emph{D} .\emph{beybienkoi} , was diverged at 2.04 Ma (95\% HDP:
1.08--3.24 Ma) and 2.54 Ma (95\% HDP: 1.49--3.84 Ma) respectively.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image5/image5}

\end{center}
\end{figure}

\textbf{Fig. 5.} Divergence time estimated in BEAST. Blue horizontal
bars show the 95\% highest posterior density interval (HPD) of the
estimated node age. Black circles on the nodes indicate the calibration
points. Vertical bars show relevant events in that transition period of
time. K, Cretaceous; N2, Pliocene; Q, Quaternary.

\emph{Ancestral range estimation}

S-DEC and S-DIVA analyses showed a highly similar biogeographic patterns
for \emph{Diestramima} species and the results shown in Fig. 6. The
reconstructed topological tree is highly consistent with the BEAST tree,
the lineages located in Guangxi and Yunnan are still the first to be
diverged. The ancestral range for \emph{Diestramima} in China covers
mountainous region of southern Yunnan (A) and mountainous hilly region
of Yunnan and Guangxi (B). As for the two clades that followed, the
ancestral distribution changed from mountainous region of southern
Yunnan (A, point 152) to mountainous region of southern Yunnan and low
mountainous hilly region of Guangxi, Guizhou and Hunan (AC, point 145).
The ancestral distribution regions of all reconstructed species in both
clades include the low mountainous hilly region of Guangxi, Guizhou and
Hunan (C). The species in Sichuan basin (D) came from low mountainous
hilly region of Guangxi, Guizhou and Hunan (C). Jiangnan hilly region
(E) and mountainous hilly region of Fujian and Guangdong (F) greatly
promoted the change of the distribution of widespread species in inland
China. The main dispersal routes are indicated by arrows in Fig. 7.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image6/image6}

\end{center}
\end{figure}

\textbf{Fig. 6.} Historical biogeography of \emph{Diestramima} as
inferred by ancestral area reconstructions carried out with RASP under
S-DEC and S-DIVA model. Showing estimations in color charts. The number
represents the optimal distributions at each node.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image7/image7}

\end{center}
\end{figure}

\textbf{Fig. 7.} The map represents the six major biogeographical areas
and dispersal routes of \emph{Diestramima} : mountainous region of
southern Yunnan (A), mountainous hilly region of Yunnan and Guangxi (B),
low mountainous hilly region of Guangxi, Guizhou and Hunan (C), Sichuan
basin (D), Jiangnan hilly region (E) and mountainous hilly region of
Fujian and Guangdong (F).

\textbf{Discussion}

\emph{Phylogenetic relationships and systematics}

Our phylogenetic results (Fig. 4), based on the combined analysis, are
highly similar to the morphological analysis (Fig. 2, Fig. 3). Based on
the results from our phylogenetic analyses, species in
genus\emph{Diestramima} that are morphologically similar and
geographically area similar tend to be grouped together. Due to the lack
of samples from Southeast Asian countries, we compared our data with the
descriptions of some species of \emph{Diestramima} by Gorochov \emph{et
al} . (2015) (Fig. 8) and proposed some species that may be closely
related. There is currently no description of the male \emph{D}
.\emph{yunnanensis} . We compared the genital plate of female
individuals (Fig. 8. F, by sample No. 2204) collected in Xishuangbanna
Tropical Botanical Garden Chinese Academy of Sciences with the
description of this site sample by Gorochov \emph{et al} . (Fig. 8. A46)
and found morphological consistency. Therefore, we supplemented the
abdominal apex of male \emph{D} . \emph{yunnanensis} (Fig. 8. E, by
sample No. 2201) based on the results of collection region and
phylogenetic tree.\emph{D} . \emph{yunnansis} is highly similar to
\emph{D} . \emph{propria}(F. 8. A31, located in Laos), except that the
male has a shorter abdomen apex and paraproct. Laos borders
Xishuangbanna Dai autonomous prefecture of Yunnan province in China.
Similarly, Myanmar borders Baoshan city of Yunnan province in China. The
abdomen apex of male \emph{D} .\emph{conica} (Fig. 8. B) is highly
similar to \emph{D} . \emph{minor}(F. 8. A36, located in Myanmar), but
has completely different shape of paraproct.

In Fig. 2, Fig. 3, Fig. 8, species in Vietnam and the area bordering
China can be divided into three groups based on the shape of the male
abdominal apex, which basically conclude the morphological
characteristics of all species in the genus \emph{Diestramima} . Most
males have similar short and simple abdomen apex, but their paraproct
are completely different in shape and length (\emph{D} .\emph{bispinosa}
, \emph{D} . \emph{cycla} , \emph{D} . \emph{hainanensis} ,\emph{D} .
\emph{hamata} , \emph{D} . \emph{palpata} , \emph{D} . sp. 1 and\emph{D}
. \emph{subtilis} ). The second group of males always has a wide and
flat abdomen apex, and some of their paraproct tend to be short and
blunt round, some tend to be long and pointed (\emph{D} . \emph{eurya}
,\emph{D} . \emph{lamina} and \emph{D} . \emph{vietnamensis} ). The male
abdomen apex of the last group is significantly longer than the first
two groups, and most of them are curved downwards (\emph{D}
.\emph{major} and \emph{D} . sp. 2). This male species with a more
slender abdomen apex (\emph{D} . \emph{acutiapicis} , \emph{D}
.\emph{austrosinensis} , \emph{D} . \emph{brevis} , \emph{D}
.\emph{intermedia} ) is widely distributed in the Oriental realm of
inland China.Gorochov \emph{et al} .(2019) divided three subgenera of
the genus\emph{Diestramima} according to the male paraproct and the
female genital plate. Paraproct of the subgenus \emph{Baculitettix} are
stick-like (rather long and thin) but with slightly or moderately higher
proximal part. In the subgenus \emph{Diestramima} and\emph{Excisotettix}
, paraproct rather short and usually high (plate-like), with distal part
truncate, angular or having hooks and/or teeth (spinules); or paraproct
moderately short, almost S-shaped in profile, with proximal and distal
parts more or less equal to each other in height, and with angular or
almost spinose apex. The difference is that the female genital plate of
the subgenus \emph{Excisotettix} with rather large posteromedian notch
and angular projections around this notch. But in our current
phylogenetic tree, we have not found a reasonable explanation for
distinguishing these subgenera. In all our samples, we found small
differences in the length and curvature of the male abdomen apex even
within the same species. But the length of the paraproct, especially the
shape always remains the same. Combining with the strong similarities
between the male abdomen apex of many species mentioned earlier, perhaps
the specialization of paraproct occupies a more dominant position in the
morphological taxonomic research of the genus \emph{Diestramima} .\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image8/image8}

\end{center}
\end{figure}

\textbf{Fig. 8.}A. Descriptions of some species in \emph{Diestramima} by
Gorochov \emph{et al} . (2015): 1, 2, \emph{D} . \emph{vietnamensis
vietnamensis} (Northern Vietnam); 3--5, \emph{D} . \emph{vietnamensis
saturata} (Northern Vietnam); 6--8, \emph{D} . \emph{austrosinensis}
(China); 9--16,\emph{D} . \emph{palpata} (Northern Vietnam); 17, 18,
\emph{D} .\emph{distincta} (China); 19--22, \emph{D} .
\emph{hainanensis} (China); 23--26, \emph{D} . \emph{bispinosa}
(Northern Vietnam); 27--30, \emph{D} .\emph{hamata} (Northern Vietnam);
31--33, \emph{D} . \emph{propria}(Laos); 34, 35, \emph{D} . \emph{major}
(China \& Vietnam); 36--38,\emph{D} . \emph{minor} (Myanmar); 39, 40,
\emph{D} . \emph{cryptopygia}(India); 41, 42, \emph{D} .
\emph{intermedia} (China); 43--45, \emph{D} .\emph{tsongkhapa} (Bhutan);
46, \emph{D} . \emph{yunnanensis} (China); 47, \emph{D} .
\emph{champasak} (Laos). Male abdominal apex from side (1, 6, 17, 34,
36, 39) and from above (18, 32, 35, 37, 40); same without lower part
from above / behind (2, 7) and from side (9, 19, 23, 27, 31); female
abdominal apex from above (4); female genital plate from below (5, 8,
12, 22, 26, 30, 38, 42, 45--47); seventh abdominal tergite and hind part
of sixth one in male from above (10, 20, 24, 28); apical part of
posteromedian process of male seventh abdominal tergite from behind and
slightly below (11, 21, 25, 29, 33); hind part of female seventh
abdominal tergite (13) and its posteromedian part (14, 15) from above;
female paraproct from side (16); posteromedian process of seventh
abdominal tergite (43) and paraproct (44) in male from side. B--E. apex
of male abdomen in lateral view: B. \emph{Diestramima conica} ;
C.\emph{Diestramima subtilis} ; D. \emph{Diestramima} sp. 1;
E.\emph{Diestramima yunnanensis} . F. female genital plate of \emph{D}
.\emph{yunnanensis} , ventral view. Scale bar: 1.0 mm.

\emph{Time and space frame of Diestramima evolution}

The results of biogeographic analysis of all genetic samples (Fig. 5)
are basically consistent with the conclusions drawn by Allegrucci \&
Sbordoni (2019). With the gradual disintegration of Pangea,
Rhaphidophoroidea is divided into two clades. The ancestors of
Dolichopodainae and Troglophilinae are mainly distributed in Laurasia
and the ancestors of Aemodogryllinae and Rhaphidophorinae are mainly
distributed in Gondwana. With the Indian subcontinent drift and
collision with Eurasia, the separation of the two clades Aemodogryllinae
and Rhaphidophorinae were formed gradually.

In the Paleogene period, we found that the separation of
genus\emph{Diestramima} from \emph{Diestrammena} diverged at 33.90Ma
(95\% HDP: 18.85--54.36 Ma) and can also be regarded as the separation
of tribe Diestramimini and Aemodogryllini. During this epoch, one of the
most significant climate changes on Earth since the Cenozoic era
occurred: the Eocene/ Oligocene boundary cooling event. It is
characterized by the rapid change of the Earth's climate from the
previous Greenhouse to Icehouse, and the emergence of Ice sheets in
Antarctica is accompanied by a significant drop in global temperature
(Sun \emph{et al} ., 2014). A drop in global temperatures may eventually
lead to the divergence of Aemodogryllini, which is more resilient to
cold conditions and can be dispersed to a wider area. The extreme
temperature rise of the Paleocene-Eocene thermal maximum did not affect
the diverged of the Aemodogryllinae and Rhaphidophorinae ancestors, and
may have only intensified their dispersal in the Oriental realm.

In the Neogene period, the first major separation time of the
genus\emph{Diestramima} diverged at 8.32Ma (95\% HDP: 4.84--12.88 Ma).
At this time, temperatures began to warm during the Miocene, and various
tectonic processes occurred due to the collision and compression of
continental plate drift and the intensification of mountain building.
Among them, the Himalayan orogeny had the most profound impact on China
(Li \emph{et al} ., 2011). It transformed the planetary wind system in
mainland China from early Miocene to monsoon wind system; the arid
inland areas in northwest China and the monsoon areas in east China
began to form. Until the late Miocene, the climate pattern of humid
southeast and arid northwest was basically formed. Paleoclimate changes
peaked during the Miocene and gradually shaped the modern climate
patterns that prevail today (Zachos \emph{et al} ., 2001). The humid
climate in the eastern area and the sea level caused by warming
temperatures may cause some species in \emph{Diestramima} to gradually
migrate east and north, and eventually become isolated due to the
frequent mountain building activities. In Fig. 5, the differentiation
in\emph{Diestramima} has a certain lag in this gradually humid and warm
climate, a reasonable explanation is that their mating behavior is
dependent on broadleaf shrubs or dwarf trees. Due to the monsoon climate
change, the relationship between the plant communities and vegetation
types in Eastern China during this period is also undergoing changes
(Jacques \emph{et al} ., 2011; Su \emph{et al} ., 2013; Li \emph{et al}
., 2015; Zhang \emph{et al} ., 2015). We found that in the males of the
genus \emph{Diestramima} , they will stay in place after the ejaculation
is over and continue to lift and press the female genitalia, and the
frequency and method of this behavior vary according to the species. We
believe that this behavior may speed up the sperm entry into the female
body while also preventing or slowing down the rate at which the female
eats the spermatophore. Correspondingly, all species
in\emph{Diestramima} we have observed will choose dwarf trees, shrubs
and ferns with large leaves as their venues for this mating behavior.
This behavior has not been found in other species in the family
Aemodogryllinae.

The genus \emph{Diestramima} diverge into two clades in the late
Miocene. A clade species are all from the southeast border of China and
countries in Southeast Asia, including \emph{D} . \emph{conica} (similar
to the species of Myanmar), \emph{D} . \emph{yunnanensis} (similar to
the species of Laos), \emph{D} . \emph{subtilis} (similar to the species
of Vietnam) and \emph{D} . \emph{major} (distributed in Vietnam and
China), and another clade species are almost all from inland China.
During this period, the uplift of the Himalayan-Tibetan Plateau led to
increased mountain folds in southwestern China and further changes in
the Asian seasonal pattern (An \emph{et al} ., 2001). The ancestors of
the genus\emph{Diestramima} have dispersal to Guizhou, Hunan and deeper
inland areas of China from the end of the Pliocene to the early
Pleistocene. Three most widely distributed species of the genus
\emph{Diestramima}(\emph{D} . \emph{austrosinensis} , \emph{D} .
\emph{beybienkoi} , \emph{D} .\emph{intermedia} ) in inland areas of
China diverged almost at the same time (2.54--1.02 Ma). The temperature
drop and ice sheet increase brought about by the Quaternary glaciation
also affected their spread at this time. The distribution and ancestral
proliferation characteristics of the genus \emph{Diestramima} in China
are very similar to the ancestral distribution characteristics of the
genus \emph{Sinocyrtaspis}(Wang \emph{et al} ., 2020) and the four
genera of Catantopidae (Xu\emph{et al} ., 2003) in China. The
mountainous region of southern Yunnan is one of the important origins of
these grasshopper genera of Catantopidae ancestors. The centres of
origin of \emph{Sinocyrtaspis}were Hunan and north-east Guangxi, and
they spread with this place as the center. This may be one of the
dispersal paths of Orthoptera in China.

\textbf{Conclusions}

Here we according to the morphological characteristics and the
phylogenetic tree constructed based on the 6 genes, phylogenetic
relationship of the genus \emph{Diestramima} is shown. At present, the
interspecies morphological characteristics and molecular phylogeny
relationships of all the samples have maintained extremely high
consistency. However, at the subgenus level, it is difficult to give a
reasonable explanation on the phylogenetic tree although we found that
some groups share some common characteristics. In phylogenetic clades
of\emph{Diestramima} , similarity in geographic distribution is as
important as similarity in morphological characteristics. This
phenomenon may be related to their lack of wings and weak mobility.
Combined with the results of biogeographic analysis, the ancestors of
the genus \emph{Diestramima} had already arrived in Guangxi and Yunnan
provinces in China before the Miocene. The two temperature rises in the
Palacocene and Miocene may have intensified the dispersal of the
ancestors of the genus \emph{Diestramima} in the Oriental realm. The
temperature drop and the uplift of the Qinghai-Tibet Plateau were the
main reasons for the divergence within the genus \emph{Diestramima} . In
addition, changes in humidity and changes in surface plant communities
are also important factors affecting their divergence. The low
mountainous hilly region of Guangxi, Guizhou and Hunan is the area where
the insects of this genus are most differentiated. At the early
Pliocene, they may have reached most of the areas where they are now.
The arrival of the Quaternary Ice Age has caused these species to
undergo violent and complex evolution in inland China.

\textbf{Acknowledgements}

We thank Dr. Hanqiang Wang (Shanghai Entomological Museum, Chinese
Academy of Sciences) and Huicong Xie (College of Life Sciences, East
China Normal University) for helping with providing specimens data. We
especially thank Jiazhi Zhang, Sainan Zhang, Tianxu Gao, Wenkai Kou,
Ziqi Jin for assistance in specimen access. We thank Emeishan National
Nature Reserve, Fenshuiling National Nature Reserve, Guangxi Forestry
Bureau, Jinyunshan National Nature Reserve, Jiuwanshan National Nature
Reserve, Laojunshan Nature Reserve, Leigongshan National Nature Reserve,
Maoershan National Nature Reserve, Nonggang National Nature Reserve,
Ruijin Ganjiangyuan Nature Reserve Administration, Shiwandashan National
Nature Reserve, Taiyanghe Provincial Nature Reserve, Wenshan Prefecture
Forestry and Grassland Bureau and Xishuangbanna Tropical Botanical
Garden Chinese Academy of Sciences for their cooperation. This study was
sponsored by the Natural Science Foundation of Shanghai (19ZR1416100).
The authors declare no conflicts of interest.

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\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/Fig1/Fig1}

\end{center}
\end{figure}\selectlanguage{english}
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