Six new species of Rhaphidophora from China (Orthoptera: Rhaphidophoridae: Rhaphidophorinae)

. Six new species of the genus Rhaphidophora Serville, 1838 are described from China: R. hexagoniproctalis Wang, Di & He sp. nov., R . heterodentis Shen, Wang & He sp. nov., R . imbricofurca Shen, Wang & He sp. nov., R . glenoides Qin, Wang & He sp. nov., R . impressa Wang, Qin & He sp. nov., R . stenoterminata Zhang, Wang & He sp. nov. A key with previously described species from China is provided.


Introduction
The subfamily Rhaphidophorinae Walker, 1869 is distributed in East Asia, the Indo-Malayan and Papuan regions.Some species can be found in Australia and some Pacifi c islands (further than New Guinea), and west to the Seychelles.Up to now, 8 genera and 180 species and subspecies have been described worldwide.The genus Rhaphidophora was established by Serville in 1838.In recent decades, numerous authors described new species from China, including R. taiwana Shiraki, 1930 andR. sinica Bey-Bienko, 1962 from Yunnan, R. sichuanensis Liu & Zhang, 2002and R. minuolamella Liu & Zhang, 2002from South China, and R. xishuang Gorochov, 2012 from Yunnan.Furthermore, Bian et al. (2017) described six species: R. biprocera, R. brevispinula, R. incilis, R. longitabula, R. longispinula and R. qudarula from China.Among them, R. longispinula was transferred to the genus Neorhaphidophora Gorochov, 1999 by Qin et al. (2018).Qin et al. (2018) also described R. wuzhishanensis from Hainan.Additionally, R. quadrispina Liu & Bian, 2021, as well as seven other species (Lu et al. 2022)

Sampling
Specimens were collected by hand or sweep net at night.All the morphological structures necessary for diagnostic purposes were photographed by an Olympus E-M5 III camera and edited by using a combination of Adobe Photoshop plus Illustrator.The scale bars are equal to 1 millimeter unless otherwise indicated.All type specimens are deposited in the Shanghai Entomological Museum, Chinese Academy of Sciences (SEM CAS).
The measurements and indices taken are defi ned below: BL = body length from apex of fastigium of vertex to posterior margin of subgenital plate FFL = length of fore or hind femora HBL = length of hind basitarsus HFL = length of fore or hind femora HTL = length of hind tibiae OvL = straight distance from visible base corner of gonangulum to apex of ovipositor PL = length of pronotum

Diagnosis
This new species is similar to R. spinifera Gorochov, 2013 and R. complanatis, but especially to R. digitata and R. shii.It differs from R. spinifera by the main part of the epiproct compacted (Fig. 1D) instead of elongated; from R. complanatis by digitate processes of the male epiproct cylindrical and shorter (Fig. 1D-E); from R. digitata and R. shii by the distance between the processes of the male epiproct which is signifi cantly longer than the length of the process (Fig. 1D), while in the other species it is nearly equal.

Etymology
The new specifi c epithet is derived from 'epiproct' and the Latinised Greek prefi x 'hexagoni' due to the shape of the main part of the male epiproct in dorsal view.HEAD.Fastigium of vertex divided into pair of plate-like processes by longitudinal furrow (Fig. 1B).Eyes protruding outward, reniform, situated at upper part of external margin of antennal sockets (Fig. 1A); lateral ocelli almost occupying nearly three quarters of lateral margins of whole tubercles, suborbicular, situated at lateral base of tubercles (Fig. 1B); median ocellus situated between two antennal sockets, oval (Fig. 1A).Maxillary palpi stout, apical segment longer than subapical one, apex infl ated, almost spherical.
LEGS.Fore femur with 1 apical spine on internal genicular lobe (Fig. 1B); fore tibia ventrally with 1 internal spine, 2 external spines and 1 pair of apical spines.Middle femur with 1 apical spine on internal and external genicular lobe separately; middle tibia dorsally with 2 pairs of spines and 1 pair of apical spines, ventrally with 2 external spines and 1 pair of apical spines.Hind femur ventrally unarmed, inner genicular lobe with spinule; hind tibia dorsally with 17-18 spinules on both sides, subapical area dorsally with 1 pair of spines, apex with 1 pair of larger mediate spines and 1 pair of smaller ventral spines asymmetrically (always inner one longer), dorsal pair, which is largest one, is missing in the sole specimen (Fig. 1G-H) and seen as pair of circular pits.Hind tarsus laterally compressed, basitarsus dorsally with 5 spinules and 1 terminal spine exceeding ventral edge (Fig. 1G).
ABDOMEN.Abdominal tergite without distinct projection.Posterior margin of tenth abdominal tergite concave.Epiproct roughly hexagonal, with pair of digitative processes (Fig. 1D) in posterolateral corners, and with rather long lobe located between these processes and directed downwards/anteriorly as well as having bispinous apex (maybe due to shrinkage of dry specimen, Fig. 1E-F).Subgenital plate damaged, repaired structure as in Fig. 1H.
COLORATION.Body dark brown with yellowish spots, ventral surface brown, external surface of hind femora with stripes.

Female
Unknown.

Diagnosis
The new species is rather similar to R. brevispinula Bian, Zhu & Shi, 2017, R. spinita and R. quadrispina, but differs from them by the upper apical half of the epiproct lamellated and irregularly incised at the terminal end, instead of becoming a pair of separated spines (Fig. 2D).

Etymology
The new specifi c epithet is derived from the Latin 'dens' and the Latinised Greek prefi x 'hetero-', referring to the irregular tooth of the upper apex of the male epiproct.HEAD.Fastigium of vertex divided into pair of plate-like processes by longitudinal furrow (Fig. 2B).Eyes protruding outward, reniform, situated at upper part of external margin of antennal sockets (Fig. 2A); lateral ocelli suborbicular, situated at lateral base of tubercles (Fig. 2B); median ocellus situated between two antennal sockets, oval (Fig. 2A).Maxillary palpi strong, apical segment longer than subapical one, apex infl ated, almost spherical (Fig. 2C).
LEGS.Fore femur with 1 apical spine on internal genicular lobe (Fig. 2B); fore tibia ventrally with 1 internal spine, 2 external spines and 1 pair of apical spines.Middle femur with 1 apical spine on internal and external genicular lobe separately; middle tibia dorsally with 2 pairs of spines and 1 pair of apical spines, ventrally with 2 external spines, 1 internal spine and 1 pair of apical spines.Hind femur ventrally unarmed, internal genicular lobe with 1 spinule; hind tibia dorsally with 21 external spinules and 20 internal spinules, subapical area dorsally with 1 pair of spines; apex with 1 pair of long dorsal spines, 1 pair of short ventral spines and 1 pair of medium long mediate spines asymmetrically, tip of the longest apical spine exceeded tip of metatarsal terminal spine (Fig. 2G-H).Hind tarsus laterally compressed, basitarsus dorsally with 5 spinules and 1 terminal spine exceeding ventral edge (Fig. 2G).
ABDOMEN.Posterior margin of tenth abdominal almost straight.Basal portion of epiproct subquadrate, upheaved aside middle (Fig. 2D-E); its more distal portion curved downwards and consisting of 2 parts: upper (posterior) part lamellar, elongate, narrowing to irregularly notched apex having pair of rather narrow and angular lateral lobules; lower (anterior) part wider than upper one and widely divided into pair of decurved hooks (Fig. 2F).Posterior margin of subgenital plate wide, widely concave in middle, subapical with 1 pair of styli (Fig. 2E); styli almost conical, basal half wide, apical half slender, apex acute.COLORATION.Body reddish brown, ventral surface yellowish brown.Scape and pedicel of antennae pale brown, fl agellum darkish in dorsal surface.Eyes darkish brown, ocelli pale.Legs pale brown, external surface hind femur with darkish stripes and pale spots.Spines of hind tibiae darkish.

Female
Unknown.

Diagnosis
The new species is similar to R. spinita and the previous species, but differs from R. spinita by the upper apical part of the epiproct lamellated then diverged to a pair of short spines (Fig. 3D) instead of becoming a pair of long spines directly; and differs from the previous species in the upper apex of the epiproct notched symmetrically (Fig. 3D).

Etymology
The new specifi c epithet is derived from the Latin 'imbrico' and 'furca', and refers to the overlapped forks of the male epiproctal apex.HEAD.Fastigium of vertex divided into pair of plate-like processes by longitudinal furrow (Fig. 3B).Eyes reniform, situated near upper part of external margin of antennal sockets (Fig. 3A); lateral ocelli almost occupying nearly three quarters of lateral margins of whole tubercles, situated at lateral base of tubercles, oval (Fig. 3C); median ocellus situated between antennal sockets, oval (Fig. 3A).Apical segment of maxillary palpi longer than subapical segment, apex infl ated, almost spherical.
LEGS.Fore coxa dorsally with spine; fore femur with 1 apical spine on internal genicular lobe (Fig. 3A); fore tibia ventrally with 2 external spines, 1 internal spine and 1 pair of apical spines.Anterior margin of middle coxa dorsally with 1 spinule; middle femur with 1 long apical spine on internal and external genicular lobe separately; middle tibia dorsally with 3 external and 2 internal spines, ventral surface with 2 external and 1 internal spine, apex with 1 pair of spines dorsally and ventrally separately.Hind femur ventrally unarmed, inner genicular lobe with 1 spinule; hind tibia dorsally with 22 spinules on both sides, subapical area dorsally with 1 pair of spines, apex with 1 pair of long dorsal spines, 1 pair of short ventral spines, and 1 pair of medium long mediate spines asymmetrically, largest apical spine broken (Fig. 3G); hind tarsus laterally compressed, basitarsus dorsally with 4 spinules and 1 long terminal spine longer than ventral edge (Fig. 3G).
ABDOMEN.Each abdominal tergite without processes.Base of epiproct almost trapezoidal; its more distal portion directed downwards, divided into lamellar upper (posterior) and lower (anterior) parts: upper part narrower, narrowest in distal third, with apex deeply and angularly notched as well as forming pair of diverged spines (Fig. 3D); lower part wider, apically with more rounded notch and pair of lateral (shorter than above-mentioned spines) hooks (Fig. 3E-F).Apex of subgenital plate almost angular, sub apex with 1 pair of styli almost cylindrical (Fig. 3F).
COLORATION.Body darkish brown.Head dorsally darkish, face pale, within antennal sockets with 1 pair of blackish patches, below eyes with 1 pair of darkish patches almost circular; eyes blackish brown, ocelli pale.Legs yellowish brown, genicular lobes darkish; basal half of hind femora with oblique yellowish stripes on external margin, dorsal surface with pale patches; spines of hind tibiae blackish.

Female
Unknown.

Diagnosis
This species differs from R. sichuanensis in the dorsal surface of the male epiproct with 2 crooked carinae, surface between carinae with oval concave (Figs 4D, 5A), while R. sichuanensis has almost straight but unparallel carinae, surface between carinae with narrower, shallower and longer concave.
The new species differs from R. xishuang in having few basitarsal spines and basal one third of male epiproct with broader concave.

Etymology
The new specifi c epithet is derived from the Latinised Greek 'glene' and refers to the small pit of the male epiproctal terminal.HEAD.Fastigium of vertex divided into pair of plate-like processes by longitudinal furrow (Fig. 4B).Eyes protruding outward, reniform, situated near upper part of external margin of antennal sockets (Fig. 4A); lateral ocelli almost occupying half of lateral margins of whole tubercles, situated at lateral base of tubercles, suborbicular (Fig. 4C); median ocellus situated between antennal sockets, oval (Fig. 4A).
COLORATION.Body reddish-brown, face light, eyes dark, ocelli light.Legs yellowish-brown, external surface of hind femora with oblique stripes.

Female
Unknown.Chinese name

Diagnosis
The new species is similar to those species bearing a simple male epiproct.It differs from the previous species by the apex of the male epiproct only plump without being vertically expanded (Fig. 8B), and bears a circular imprint at the bottom of the epiproctal concavity (Fig. 8A); from R. setiformis Qin, Jiang, Liu & Li, 2018 by the apex of the male epiproct swell not depressed (Figs 8B, 9A), the concavity of the male epiproct being not narrow and prolonged from base to apex; and from R. duxiu in the shape of the male epiproctal concavity and the length of the female subgenital plate.

Etymology
The new specifi c epithet is derived from the Latin 'imprimo' and stands for the circular imprint of the epiproctal concavity in the male.HEAD.Fastigium of vertex divided into pair of plate-like processes by longitudinal furrow (Fig. 6B).Eyes protruding outward, reniform, situated near upper part of external margin of antennal sockets (Fig. 6A); lateral ocelli situated at lateral base of rostral tubercles, suborbicular, nearly occupying three fourths of lateral margin of rostral tubercles (Fig. 6C); median ocellus situated between antennal sockets, oval (Fig. 6A).Maxillary palpi strong, apical segment longer than subapical segment, apex infl ated, almost spherical.
LEGS.Fore coxa infl ated, dorsally with 1 spinule; fore femur with 1 apical spine on internal genicular lobe (Fig. 6B); ventral surface of fore tibia with 2 external and 1 internal spine, apex with 1 pair of spines on ventral surface.Anterior margin of middle coxa dorsally with 1 pair of spiniform processes; mid femur with 1 apical spine on internal and external genicular lobe separately; mid tibia dorsally with 1 pair of spines and 1 pair of apical spines, ventrally with 2 pairs of external spines and 1 pair of apical spines.Hind femur ventrally unarmed, with 1 spinule on internal genicular lobe; hind tibia dorsally with 17-18 spinules on both dorsal margins, subapical area dorsally with 1 pair of spines, apex with 1 pair of dorsal long spines and 2 pairs of shorter ventral spines asymmetrically, the largest apical spine slightly surpassing tip of terminal basitarsal spine (Fig. 6G); hind tarsus laterally compressed, basitarsus with 2 spinules and a terminal spine on dorsal surface, dorsal tip longer than ventral edge (Fig. 6G).
ABDOMEN.Abdominal tergite without processes, posterior margin of tenth abdominal tergite emarginate (Fig. 6D).Epiproct similar to that of previous species but with basal portion trapezoidal, more distal portion elongately triangular, concavity between dorsolateral carinae drop-shaped, base of concavity widened and expanding to middle forming circular imprint, apex of concavity constricted and narrowing to epiproctal end (Figs 6D,8A).Subgenital plate roughly semicircular, sub apex with pair of stout styli with apex enlarged (Fig. 6E-F).Cerci elongated, basal part wider than apical part, nearly conical, apex obtuse (Fig. 6E).

Female
Similar to male.Hind basitarsus with 4-5 spinules and a terminal spine on dorsal surface.Subgenital plate transverse and short, near semicircular, hind margin pointed in middle (Figs 7A,8C).Ovipositor longer than half length of hind femora, narrower from base to apex gradually, evenly upcurved, ventral margin serrated near apex (Fig. 7B).Chinese name

Diagnosis
The new species differs from species with a simple male epiproct by the unique undulated bilateral carinae at the basal part of the male epiproct, for other differences see the key to all Chinese species below.HEAD.Fastigium of vertex divided into pair of plate-like processes by longitudinal furrow (Fig. 10C).
THORAX.Anterior margin of pronotum straight, posterior margin of pronotum projected (Fig. 10B), length of lateral lobes nearly equal to width, posterior margin of ventral surface arched; posterior margin emarginate; posterior margin of mesotergum obviously projected, posterior margin of metanotum straight (Fig. 10C).
LEGS.Fore coxa dorsally with 1 apical spine; fore femur with 1 apical spine on internal genicular lobe; fore tibia with 2 external and 1 internal spine on ventral surface, apex with 1 pair of spines on ventral surface, dorsal surface unarmed.Anterior margin of middle coxa dorsally with 1 spinule; middle femur with 1 pair of apical spines on dorsal surface and ventral surface separately; middle tibia with 2 pairs of spines and 1 pair of apical spines on dorsal surface, ventral surfaces with 2 external spines and 1 pair of apical spines.Hind femur ventrally unarmed, inner genicular lobe with 1 spinule; hind tibia dorsally with 20-21 spinules on both sides, subapical area dorsally with 1 pair of spines, apex with 1 pair of dorsal long spines and 2 pairs of shorter spines asymmetrically, largest apical spine protruding over tip of terminal basitarsal spine (Fig. 10G); hind tarsus laterally compressed, basitarsus with 3-5 spinules and 1 large terminal spine on dorsal surface (Fig. 10G).

Discussion
Up to now, there are 27 species of the genus Rhaphidophora distributed in China including the six species described in this study (Fig. 14).Most species are distributed in the southern regions of the Yangtze River, with Yunnan being the hotspot, consisting of eleven species.The current classifi cation of these species is primarily based on morphological features, especially the male epiproct.However, it is worth noting that many species were described based on few samples, and these morphological features should be reevaluated in future studies.During our fi eld investigations, we have encountered species of Rhaphidophorinae many times at night in central, east and south China.However, their population sizes are small, and they can be challenging to capture due to their swift movement and ability to clim high on tree branches, which distinguishes them from species of Aemodogryllinae Jacobson, 1905.Therefore, more effi cient collecting methods are needed, and it is expected that there are undescribed species in those areas.Qin (2020) erroneously synonymized R. setiformis with R. xishuang in her doctoral dissertation, because the male epiproct apex of the former species is not vertically incrassated and without a conical tip as in the latter species (Fig. 9A-B).Additionally, the hind basitarsus of R. setiformis has 2 to 3 spinules, while that of R. xishuang has 5 to 6.
Camel crickets, as wingless orthopterans with limited migratory capabilities, are ideal organisms for biogeographic research.Zhu et al. (2022) conducted phylogeographic studies on the genus Diestramima Storozhenko, 1990 in China, which revealed that Yunnan is the likely origin of this genus, consistent with the biodiversity of Rhaphidophora in China.In future studies, it is encouraged to collect DNA samples, as molecular studies can provide insights into the phylogeny of the genus Rhaphidophora.

Fig. 1 .
Fig. 1.Rhaphidophora hexagoniproctalis Wang, Di & He sp.nov., holotype, ♂ (SEM CAS 14098450).A. Head in frontal view.B-C.Head and thorax in dorsal and lateral view, respectively.D-F.Apex of abdomen in rear dorsal, ventral and lateral view, respectively.G-H.Right hind tarsus in externally lateral and dorsal view, respectively.I. Body in lateral view.

Fig. 2 .
Fig. 2. Rhaphidophora heterodentis Shen, Wang & He sp.nov., holotype, ♂ (SEM CAS14098451).A. Head in frontal view.B-C.Head and thorax in dorsal and lateral view, respectively.D-F.Apex of abdomen in rear dorsal, ventral and lateral view, respectively.G-H.Left hind tarsus in inner lateral and dorsal view.I. Body in lateral view.

Fig. 3 .
Fig. 3. Rhaphidophora imbricofurca Shen, Wang & He sp.nov., holotype, ♂ (SEM CAS 14098466).A. Head in frontal view.B-C.Head and thorax in lateral and dorsal view, respectively.D-F.Apex of abdomen in right rear dorsal, lateral and ventral view, respectively.G. Left hind tarsus in inner lateral view.H. Body in lateral view.

Fig. 4 .
Fig. 4. Rhaphidophora glenoides Qin, Wang & He sp.nov., holotype, ♂ (SEM CAS 14098452).A. Head in frontal view.B-C.Head and thorax in dorsal and lateral view, respectively.D-F.Apex of abdomen in left dorsal, lateral and rear ventral view, respectively.G. Right hind tarsus in outer lateral view.H. Body in lateral view.

Fig. 6 .
Fig. 6.Rhaphidophora impressa Wang, Qin & He sp.nov., holotype, ♂ (SEM CAS 14098455).A. Head in frontal view.B-C.Head and thorax in dorsal and lateral view, respectively.D-F.Apex of abdomen in rear dorsal, lateral and ventral view, respectively.G. Left hind tarsus in outer lateral view.H. body in lateral view.

Fig. 8 .
Fig. 8. Line drawing of Rhaphidophora impressa Wang, Qin & He sp.nov. A. Male epiproct in dorsal view.B. Male epiproct in lateral view.C. Renatured hind margin of female subgenital plate in rear ventral view.

Fig. 12 .
Fig. 12. Line drawing of Rhaphidophora stenoterminata Zhang, Wang & He sp.nov. A. Male epiproct in dorsal view.B. Male epiproct in lateral view.C. Female subgenital plate in rear ventral view.