Zoosyst. Evol. 101 (2) 2025, 533-550 | DOI 10.3897/zse.101.141741

Ree Ee
BERLIN

New species of 7ygarrup centipedes from Xizang, China
(Geophilomorpha, Mecistocephalidae)

Chao Jiang', Jiabo Fan'*, Chunxue You’, Weichun Li*, Lugi Huang!

1 State Key Laboratory for Quality Ensurance and Sustainable Use of Dao-di Herbs, National Resource Center for Chinese Materia Medica,
China Academy of Chinese Medical Sciences, Beijing 100700, China

2 Tianjin Key Laboratory of Agricultural Animal Breeding and Healthy Husbandry, College of Animal Science and Veterinary Medicine, Tianjin
Agricultural University, Tianjin 300392, China

3 College of Agronomy, Jiangxi Agricultural University, Nanchang 330045, China

https://zoobank. org/41 DEF 99E-4563-48E3-B7BB-59F8D3B19439

Corresponding authors: Chao Jiang (jiangchao0411@126.com); Luqi Huang (huangluqi01@126.com)

Academic editor: Martin Husemann @ Received 13 November 2024 # Accepted 21 January 2025 Published 26 February 2025

Abstract

Prior to this study, the genus 7ygarrup Chamberlin, 1914, is known to contain only two species in Xizang, China. New records
derived from multiple collections allow for the distribution ranges of six species to be refined: 7ygarrup tridentatus Jiang & Huang,
sp. nov., Zygarrup cerrus Jiang & Huang, sp. nov., Zygarrup multiporus Jiang & Huang, sp. nov., and Tygarrup fimbriatus Jiang &
Huang, sp. nov. are described as new. 7ygarrup muminabadicus Titova, 1965, was first reported in the fauna of Xizang. Tygarrup
poriger (Verhoeff, 1942) is rediscovered and redescribed, and Zygarrup nepalensis Shinohara, 1965 is synonymized with T! poriger.

Remarks are provided for all encountered species, and a key to all 7vgarrup species in China is presented.

Key Words

Biodiversity, key, new record, taxonomy

Introduction

Xizang, the second largest provincial administrative unit
of China, is located in the western and southern parts
of the Tibetan Plateau. This region, known as the “third
pole of the earth,” is the highest place in the world. Its
southern region, the Himalaya, exhibits a pronounced
vertical stratification of ecosystems and is recognized as
a biodiversity hotspot for the study of animal taxonomy
as well as biological evolution (Cai et al. 2018; Li et al.
2022). A century ago, seventeen centipede species have
been documented in Xizang, comprising eight species of
Scolopendromorpha and seven species of Lithobiomor-
pha (Lewis 2010; Song et al. 2010; Ma et al. 2014; Qiao
et al. 2019; Niu et al. 2021; Chen et al. 2023). However,
understanding of the soil centipede species diversity in
Xizang remains quite limited. To date, only two species
of Geophilomorpha have been described from the region,

both of which belong to the genus 7ygarrup Chamberlin,
1914 (Silvestri 1919; Verhoeff 1942). This includes the
incompletely described species 7: diversidens (Silves-
tri, 1919) from the Himalayas and 7: poriger (Verhoeff,
1942) from “the middle Himalaya,” near Shigatse. The
objective of the present study is to enhance our knowl-
edge of the species diversity within Geophilomorpha in
Xizang, with a focus on the discovery and description of
new species of 7ygarrup centipedes.

The genus 7ygarrup, characterized by its elongated
head capsule, clypeus with an entire plagula, spiculum
absent, clypeal pleurite without setae, coxosternum of the
first maxillae divided, whereas that of the second maxillae
is undivided, and the sternal typically with pores present
in males (Uliana et al. 2007). It is the second largest genus
in the family Mecistocephalidae Bollman, 1893, and com-
prises 15 known species (Uliana et al. 2007; Bonato et al.
2011; Chao et al. 2020). This genus is mainly distributed

Copyright Jiang, C. et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use,
distribution, and reproduction in any medium, provided the original author and source are credited.

534 Jiang, C. et al.: Tygarrup centipedes from Xizang, China (Geophilomorpha, Mecistocephalidae)

in the tropical and subtropical regions of East Asia and
Southeast Asia and can adapt to a variety of climates
ranging from tropical rainforests to cold high mountains
(Silvestri 1919; Verhoeff 1939, 1942: Paik 1961; Uliana
et al. 2007; Bonato et al. 2011; Chao et al. 2020). Before
this study, only three species of Zygarrup had been doc-
umented in China, with two species being restricted to
Xizang (Silvestri 1919; Verhoeff 1942; Chao et al. 2020).
During our examination of Zygarrup specimens from
Xizang, we identified six species within the genus, 1n-
cluding four new species. In this study, we describe these
species, with a particular focus on the new species.

Materials and methods

Specimens were collected by tweezers and preserved
in 75% ethanol. The type materials are deposited in the
National Resource Center for Chinese Materia Medica,
China Academy of Chinese Medical Sciences, Beijing,
China (CMMI).

Specimens were dissected, and their cephalic capsules,
forcipular segments, mandibles, maxillary complexes,
and remaining bodies were mounted on temporary slides
using 75% ethanol or lactic acid. Taxonomic characters
were examined and photographed (Figs 1, 2) using a
Leica M205 FCA stereomicroscope (7.8—160*) and an
Olympus BX51 microscope (100—-400x). Photos were
converted into hand-drawn illustrations using Sketchbook
6.0.6. Localities were mapped with ArcMap 10.7.1. The
terminology of morphology follows Bonato et al. (2010a).

Results
Taxonomy

Order Geophilomorpha Pocock, 1895
Family Mecistocephalidae Bollman, 1893

Genus 7ygarrup Chamberlin, 1914

Type species. Zygarrup intermedius Chamberlin, 1914,
by original designation.
Diagnosis. See Uliana et al. (2007).

Tygarrup tridentatus Jiang & Huang, sp. nov.
https://zoobank org/4201B13D-F951-465C-894D-450027BCB860
Figs 1A-C, 3

Material examined. Holotype. «© 2 (CMMI
20240725001D), Cuina, Xizang Autonomous Regions,
Yadong County, Xiasima town (27.4788°N, 88.9063°E),
2960 m asl., 25.vii.2024, leg. Chao Jiang.

Paratypes. © 429 343 3 juveniles (CMMI
20230720002D, CMMI 20230725002D, -003D), same as
holotype but collected at 20—25.vu1.2024.

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Other materials. *> 299 1¢ 1 juvenile (CMMI
20240721001D, 20240724001D), China, Xizang Au-
tonomous Regions, Yadong County, Pangda village,
2100-2400 m asl., 21—24.vil.2024, leg. Chao Jiang &
Qing Li, 322 (CMMI 20240720001D), Upper Yadong
village, G562 National Road (27.5481°N, 88.9998°E),
3460 m asl., 20.vii1.2024, leg. Chao Jiang & Qing Li;
12 243 5 juveniles (CMMI 20240715001D—005D),
Cuona County, Lei Menba ethnic village, Lei Hydro-
electric-power station (27.8196°N, 91.7462°E), 2490 m
asl., 15.vii.2024, leg. Chao Jiang & Qing Li, 299 439
(CMMI 20240714001D), Entrance of leibugou valley
(27.9240°N, 91.8381°E), 3840 m asl., 14.vii.2024, leg.
Chao Jiang; * 19 14 (CMMI 20240714301D), Gon-
geri Menba ethnic village, Mengyu Sanshenghu Lake
(27.9267°N, 91.8728°E), 4410 m asl., 14.vii.2024, leg.
Chao Jiang & Qing L1.

Diagnosis. A Zygarrup species with 45 leg-bearing
segments. Body length of adult reaches 2.5 cm. Trans-
verse suture curved. Clypeal setae 9-12 pairs. Forcipular
trochanteroprefemur, tibia, and tarsungulum, each with
a well-developed tooth, article II without teeth. Sternal
pores present in males only.

Description. Maximum length of male 2.5 cm and fe-
male 2.3 cm.

Color (of preserved specimens in alcohol) morph 1:
head and forcipular segment reddish-brown, trunk and legs
homogeneously yellow, without dark patches (Fig. 1A-C,
in Yadong populations); morph 2: head and forcipular seg-
ment dark red, trunk and legs yellow, with dark patches
along the trunk, sternal sulci darker (in Cuona populations).

Cephalic plate 1.6 x longer than the widest. Transverse
suture rounded. Lateral margins sinuous to straight, conver-
gent backwards slightly, anterior margin convex, with a me-
dian incision, posterior margin slightly rounded (Fig. 3A).

Clypeus 1.9 x wider than long (Fig. 3B). Clypeus with
an entire plagula covering most of the clypeus, areola-
tion only present along the anterior margin of the head;
clypeal setae as follows: 1 pair on the areolate part; 7-10
pairs along the anterior margin of the clypeal plagula, 1
pair on the central part of plagula (Fig. 3C).

Labrum (Fig. 3C): Mid-piece ca. 2 x longer than wide,
posteriorly pointed and well sclerotized. Anterior ala tri-
angular, and medial margin reduced to a vertex; posterior
margin of each side-piece sinuous.

Cephalic pleurite without spiculum or setae; areolate
part present along anterior margin and paraclypeal suture;
stilus well-developed, nearly straight, with an anterior in-
cision on each side (Fig. 3B).

First maxillae (Fig. 3D): Coxosternite 2 < wider than
long, divided by mid-longitudinal suture, anterior corners
of coxosternum not projecting. Medial projection about
1.5 x wider than long, internal margin with several se-
tae, distal lobe curved inward; telopodite about 4 x longer
than wide, curved inward.

Second maxillae (Fig. 3D): Coxosternite undivided, ca.
1.5 x wider than long; anterior and posterior margins con-
cave, lateral margins concave medially, posterior corners
only slightly projecting externally; areolation on medial part,

Zoosyst. Evol. 101 (2) 2025, 533-550 535

uch

Figure 1. Cephalic plate and forcipular segments. A, D, G. Cephalic plates, dorsal; B, E, H. Cephalic plates, ventral; C, F, I. Forcipular
segments, dorsal. A-C. 7ygarrup tridentatus sp. nov., holotype; D-F. Tygarrup cerrus sp. nov., holotype; G—I. Tygarrup multiporus
sp. nov., holotype. Scale bar: 500 um.

with pores along the lateral margins, setae present along Telopodites of the second maxillae well developed, sur-
the posterior part. Foraminal process surrounding the meta- _ passing those of the first maxillae; article 1 of telopodite ca.
meric pore separated from lateral margin of coxosternite. 4.7 x longer than wide, curved outward, with distal setae on

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536 Jiang, C. et al.: Tygarrup centipedes from Xizang, China (Geophilomorpha, Mecistocephalidae)

—
a Ml

Figure 2. Cephalic plate and forcipular segments. A, D, G. Cephalic plates, dorsal; B, E, H. Cephalic plates, ventral; C, F, I. Forcip-
ular segments, dorsal. A-C. Zygarrup fimbriatus sp. nov., holotype; D-F. Tygarrup poriger (Verhoeff, 1942) (spm. CMMI
20240309001D); G-I. Tygarrup muminabadicus Titova, 1965 (spm. CMMI 20240310001D). Scale bar: 500 um.

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Zoosyst. Evol. 101 (2) 2025, 533-550 537

Vase e ers aa >
essessssireeze=ee

S5stss sitet:

==

ma

Figure 3. Zygarrup tridentatus sp. nov., holotype. A. Cephalic plate, dorsal; B. Cephalic plate, ventral (maxillary complex re-
moved); C. Clypeus, ventral (maxillary complex removed); D. Maxillary complex, ventral; E. Mandible, ventral; F. Forcipular seg-
ment, ventral; G. Forcipular segment, dorsal; H. Seventh sternum of leg-bearing segment, ventral; I. Ultimate leg-bearing segment
and left leg, ventral. Scale bars: 500 um (A, B, F, G, I); 100 um (C, D, H); 25 um (E).

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538 Jiang, C. et al.: Tygarrup centipedes from Xizang, China (Geophilomorpha, Mecistocephalidae)

the internal surface; article 2 of telopodite 1.3 x longer than
wide, with distal setae on the internal side; article 3 ca. 2.2
x longer than wide, with distal end densely covered with
setae; apical claw large and simple.

Mandible with ca. 5 well-developed and ca. 2 rudimen-
tary pectinate lamellae; first lamella with 3 teeth, similar
in size; intermediate lamellae with 6—7 teeth (Fig. 3E).

Forcipular segment: Coxosternite 1.2 <x wider than
long, anterior margin with a pair of small teeth (Fig. 3F);
scapular points of pleura surpass the anterior margin of
coxosternite. Forcipular trochanteroprefemur about 1.3 x
longer than wide, with a prominent distal tooth, femur
without teeth, tibia and tarsungulum each with a well-de-
veloped tooth (Fig. 3G).

Leg-bearing segments: A total of 45 leg-bearing seg-
ments. Sternal sulcus bifurcate, faintly impressed, with
an acute angle (Fig. 3H); sternal pores present on males,
absent on females.

Ultimate leg-bearing segment densely setose. Terg-
ite sub-rectangular, 1.6 x longer than wide. Coxopleuron
about 1.1 = longer than wide and 1.7 x longer than ultimate
sternite. Ca. 13—22 coxal pores on ventral and lateral sides
of each coxopleuron, the innermost two or three pores usu-
ally larger than others (Fig. 31). The ultimate leg telopodite
not swollen, with scattered setae, without pretarsus.

Postpedal segments densely setose in both sexes. Male
gonopods bi-articulate, narrow, and separated by a coni-
cal projection. Female gonopods inconspicuous bi-artic-
ulate, subtriangular (Fig. 31). Anal pores present on each
ventro-lateral side of postpedal segments.

Etymology. Latin: tridentatus = trident. The new spe-
cies name refers to its forcipular trochanteroprefemur,
tibia, and tarsungulum, each with a well-developed tooth.
We suggest the Chinese common name as “= HAR”,

Distribution. China (Xizang).

Remarks. This new species resembles Zygarrup
poriger (Verhoeff, 1942) in possessing a tooth on the dor-
sal base of the forcipular tarsungulum. However, it is dis-
tinct from 7: poriger by the presence of a well-developed
tooth on the forcipular femur (Figs 1C, 3F).

Tygarrup cerrus Jiang & Huang, sp. nov.
https://zoobank.org/35956858-705E-46F 1-A5D5-34D5477FD794
Figs 1D-F, 4

Material examined. Holotype. °2 (CMMI
20240307017D), Cuina, Xizang Autonomous Regions,
Bomi County, Qingduo town (30.0946°N, 95.7031°E),
2850 m asl., 7.111.2024, leg. Chao Jiang.

Paratypes. °69 2 73. (CMMI 20240307011 D—016D,
018D—024D), same as holotype.

Diagnosis. A 7ygarrup species with 45 leg-bearing
segments. Body length of adult reaches 3.6 cm. Trans-
verse suture curved. Clypeal setae 9-12 pairs. Forcipular
trochanteroprefemur and tibia, each with a small tuber-
cle. Cerrus present as two lateral groups of setae. Sternal
pores present in males only.

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Description. Maximum length of male 3.6 cm and fe-
male 3.4 cm.

Color (of preserved specimens in alcohol): head and
forcipular segment dark red, trunk and legs yellow, with
dark patches along the trunk (Fig. 1D-F).

Cephalic plate 1.5 x longer than the widest. Transverse
suture rounded. Lateral margins nearly straight and con-
vergent backwards, anterior margin convex, with a me-
dian incision, posterior margin rounded. Setae arranged
nearly symmetrically (Fig. 4A).

Clypeus 2.1 x wider than long (Fig. 4B). Clypeus with
an entire plagula covering most of the clypeus, areola-
tion only present along the anterior margin of the head;
clypeal setae as follows: 1 pair on the areolate part, 7-10
pairs along to the anterior margin of the clypeal plagula,
1 pair on the central part of plagula (Fig. 4C).

Labrum (Fig. 4C): Mid-piece ca. 2.4 = longer than
wide, posteriorly pointed and well sclerotized. Anterior
ala triangular, medial margin reduced to a vertex; posteri-
or margin of each side-piece sinuous.

Cephalic pleurite without spiculum or setae; areolate
part present along anterior margin and paraclypeal suture;
stilus well developed, nearly straight, with an anterior in-
cision on each side (Fig. 4B).

First maxillae (Fig. 4D): Coxosternite 1.7 =< wider
than long, divided by mid-longitudinal suture, the ante-
rior corners of coxosternum not projecting. Medial pro-
jection about 1.3 =< wider than long, internal margin with
several setae, distal lobe curved inward; telopodite about
3 x longer than wide, curved inward.

Second maxillae (Fig. 4D): Coxosternite undivided,
ca. 1.2 x wider than long; both the anterior and poste-
rior margins concave, lateral margins concave medially,
posterior corners only slightly projecting externally; are-
olation on medial part, setae present along the posterior
part; the foraminal process surrounding the metameric
pore separated from lateral margin of coxosternite. Te-
lopodites of the second maxillae well developed, surpass-
ing those of the first maxillae; article 1 of the telopodite
2.8—3.2 x longer than wide, curved outward; article 2 of
the telopodite ca. 1.1 x longer than wide; article 3 ca. 1.8
x longer than wide, with distal end densely covered with
setae; apical claw large and simple.

Mandible with ca. 6 well-developed and ca. 2 rudimen-
tary pectinate lamellae; the first lamella with 5 teeth, similar
in size; intermediate lamellae with 11—15 teeth (Fig. 4E).

Forcipular segment: Coxosternite 1.4 <x wider than
long, anterior margin with a pair of truncated teeth (Fig.
AF); scapular points of pleura not reaching the anterior
margin of coxosternite (Fig. 4G). Cerrus present as two lat-
eral groups of setae (Fig. 4H). Forcipular trochanteroprefe-
mur about 1.4 x longer than wide and has a small distal
tubercle, femur with or without tubercle, tibia with a small
pointed tubercle, tarsungulum without tubercle (Fig. 4G).

Leg-bearing segments. A total of 45 leg-bearing seg-
ments. Sternal sulcus bifurcate, faintly impressed, with
an acute angle (Fig. 41); sternal pores present on males,
absent on females.

Zoosyst. Evol. 101 (2) 2025, 533-550 539

Figure 4. 7ygarrup cerrus sp. nov., holotype. A. Cephalic plate, dorsal; B. Cephalic plate, ventral (maxillary complex removed);
C. Clypeus, ventral (maxillary complex removed); D. Maxillary complex, ventral; E. Mandible, ventral; F. Forcipular segment,
ventral; G. Forcipular segment, dorsal; H. Part of forcipular segment, dorsal; I. Seventh sternum of leg-bearing segment, ven-
tral; J. Ultimate leg-bearing segment and left leg, ventral. Abbreviations: ce — cerrus. Scale bars: 500 um (A, B, D, F, G, I, J):
100 um (C, H); 25 um (EF).

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540 Jiang, C. et al.: Tygarrup centipedes from Xizang, China (Geophilomorpha, Mecistocephalidae)

Ultimate leg-bearing segment densely setose. Terg-
ite sub-rectangular, 1.4 < longer than wide. Coxopleuron
about 0.9 x longer than wide and 1.5 x longer than ultimate
sternite. Ca. 30-60 coxal pores on ventral and lateral sides
of each coxopleuron, the innermost two or three pores usu-
ally larger than others (Fig. 4J). The ultimate leg telopodite
not swollen, with scattered setae, without pretarsus.

Postpedal segments densely setose for both sexes. Male
gonopods bi-articulate, narrow, and separated by a conical
projection. Female gonopods simple, subtriangular, and
touching each other at their bases (Fig. 4J). Anal pores
present on each ventro-lateral side of postpedal segments.

Etymology. The new species name refers to the presence
of a paired lateral group of setae on the dorsal side of the
forcipular coxosternite, termed ‘cerrus’ by Crabill (1970).
We suggest the Chinese common name as “24 HRW”,

Distribution. China (Xizang).

Remarks. In mecistocephalids, the presence of a cer-
rus is a consistent character observed in several species,
particularly within the genera Dicellophilus Cook, 1896,
Takashimaia Miyosi, 1955, Proterotaiwanella Bonato,
Foddai & Minelli, 2002, Krateraspis Lignau, 1929, and
some species of Mecistocephalus Newport, 1843 (Bonato
et al. 2002, 2003, 2010b; Uliana et al. 2007; Dyachkov
and Bonato 2022; Pan et al. 2024). However, there have
been no reports of the presence of a cerrus in any species
of Zygarrup to date. The discovery of cerrus in the newly
described species 7’ cerrus sp. nov. offers a further under-
standing of the evolutionary development of this struc-
ture within the Mecistocephalidae as well as phylogenetic
relationships among the mecistocephalid genera.

Tygarrup multiporus Jiang & Huang, sp. nov.
https://zoobank.org/5A9F733C-5880-4647-A 377-706428584E8D
Figs 1G-I, 5

Material examined. Holotype. «© 2 (CMMI
20240312003D), Cuina, Xizang Autonomous Regions,
Medog County, near Deergong village (29.1810°N,
95.1439°E), 1710 masl., 12.111.2024, leg. Chao Jiang.

Paratypes. «© 2292 (CMMI = 20240309012D,
20240311003D), CHINA, Xizang Autonomous Regions,
Medog County, near Renqingbensi Temple (29.3075°N,
95.3573°E), 1920 m asl., 9-11 .111.2024, leg. Chao Jiang.

Diagnosis. A 7ygarrup species with 45 leg-bearing
segments. Body length of adult reaches 3.4 cm. Cephalic
frontal plate postero-lateral corner bears 10—20 pore-like
sensilla. Clypeal plagula with 6 pairs of setae on ante-
ro-medial and several pore-like sensilla on antero-lateral
comers. Forcipular trochanteroprefemur and tibia, each
with a tubercle, femur and tarsungulum without tuber-
cles. Sternal pores present in males only.

Description. Maximum length 3.4 cm.

Color (of preserved specimens in alcohol): Head and
forcipular segment reddish-brown, trunk and legs homo-
geneously yellow, without dark patches (Fig. 1G—I).

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Cephalic plate (Fig. 5A) 1.5 x longer than the widest.
Transverse suture curved, forms a backward-directed an-
gle medially. Lateral margins sinuous, convergent back-
wards strongly, anterior margin convex, with a median
incision, posterior margin rounded. Frontal plate poste-
ro-lateral corner bears 10—20 pore-like sensilla. Setae ar-
ranged nearly symmetrically.

Clypeus 2 x wider than long. Clypeus with an entire
plagula covering most of the clypeus, areolation only
present along the anterior margin of the head; clypeal se-
tae as follows: 1 pair on the areolate part, 2 pairs along the
anterior margin of the clypeal plagula, 2 pairs on the an-
tero-medial part inside a large insula, | pair on the central
part of plagula. Five to eight pore-like sensilla on each
antero-lateral corner (Fig. 5B).

Labrum: Mid-piece ca. 2.1 x longer than wide, poste-
riorly pointed and well sclerotized. Anterior ala triangular,
medial margin reduced, distinct shorter than lateral mar-
gin; posterior margin of each side-piece sinuous (Fig. 5B).

Cephalic pleurite without spiculum or setae; areolate
part present along anterior margin and paraclypeal suture;
stilus well developed, nearly straight, with an anterior in-
cision on each side (Fig. 5B).

First maxillae (Fig. 5C): Coxosternite 1.9 < wider
than long, divided by mid-longitudinal suture, anterior
corners of coxosternum not projecting. Medial projection
about 1.3 x wider than long, internal margin with several
setae, distal lobe curved inward; telopodite about 2.3 x
longer than wide, curved inward.

Second maxillae (Fig. 5C): Coxosternite undivided,
ca. 1.4 x wider than long; both the anterior and poste-
rior margins concave, lateral margins concave medially,
posterior corners only slightly projecting externally; are-
olation on medial part, setae present along the posterior
part; the foraminal process surrounding the metameric
pore separated from lateral margin of coxosternite. Te-
lopodites of the second maxillae well developed, surpass-
ing those of the first maxillae; article 1 of the telopodite
2.9-3.5 x longer than wide, curved outward; article 2 of
the telopodite ca. 1.4 x longer than wide; article 3 ca. 2.3
x longer than wide, with distal end densely covered with
setae; apical claw large and simple.

Mandible with ca. 6—7 well-developed and ca. 2 ru-
dimentary pectinate lamellae; the first lamella with 4
teeth, similar in size; intermediate lamellae with 9-13
teeth (Fig. 5D).

Forcipular segment: Coxosternite 1.4 <x wider than
long, anterior margin with a pair of truncated teeth (Fig.
SE); scapular points of pleura not reaching the anterior
margin of coxosternite (Fig. 5F). Forcipular trochantero-
prefemur about 1.3 x longer than wide and has a conical
distal tooth, intermediate articles each with a small tooth,
tarsungulum without tooth.

Leg-bearing segments. A total of 45 leg-bearing
segments. Sternal sulcus bifurcate faintly impressed,
with an acute angle (Fig. 5G); sternal pores present on
males, absent on females.

Zoosyst. Evol. 101 (2) 2025, 533-550 541

Figure 5. Zygarrup multiporus sp. nov., holotype. A. Cephalic plate, dorsal; B. Cephalic plate, ventral (maxillary complex removed);
C. Maxillary complex, ventral; D. Mandible, ventral; E. Forcipular segment, ventral; F. Forcipular segment, dorsal; G. Seventh ster-
num of leg-bearing segment, ventral; H. Ultimate leg-bearing segment and left leg, ventral. Abbreviations: pls — pore-like sensilla.
Scale bars: 500 um (A—C, E—H); 25 um (D).

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542 Jiang, C. et al.: Tygarrup centipedes from Xizang, China (Geophilomorpha, Mecistocephalidae)

Ultimate leg-bearing segment densely setose. Tergite
sub-rectangular, 1.1 =< longer than wide. Coxopleuron
about 0.8 x longer than wide and 1.7 x longer than ulti-
mate sternite. Ca. 20—28 coxal pores on ventral and lateral
sides of each coxopleuron, the innermost two pores usu-
ally larger than others (Fig. 5H). The ultimate leg telopo-
dite not swollen, with scattered setae, without pretarsus.

Postpedal segments densely setose for both sexes. Male
gonopods bi-articulate, narrow, and separated by a conic
projection. Female gonopods simple, subtriangular, and
touching each other at their bases (Fig. 5H). Anal pores
present on each ventro-lateral side of postpedal segments.

Etymology. The new species name refers to the pres-
ence of many pore-like sensilla on cephalic capsule and
clypeus plagula. We suggest the Chinese common name
as “2% FL tea”,

Distribution. China (Xizang).

Remarks. This new species is distinguished from all
the hitherto known 7Jygarrup species by the presence of
pore-like sensilla on its clypeal plagula and cephalic plate,
a character not previously documented in this genus.
However, when Titova (1983) described 7: crassignathus
Titova, 1983, and 7’ singaporiensis Verhoeff, 1937, the il-
lustrations of the clypeal plagula (Titova 1983: figs 5, 10)
showed numerous dense small spots, which are inferred
herein to possibly be pore-like sensilla.

The new species is further distinguished from T° crassig-
nathus by the absence of an incrassate forcipular trochantero-
prefemurand a large distal tooth, features present tn the latter
(Titova 1983: fig. 7). Instead, the new species possesses a
forcipular trochanteroprefemurtooth of typical dimensions.
The new species can also be distinguished from 7) singa-
poriensis by the number of clypeus plagula pore-like sensilla
(more than 5—10 on each side, confined to the antero-lateral
corner vs. more than 50 on each side, extending from the an-
tero-lateral corner to the paramedian setae) and the clypeus
setae (2 lateral and 6 paramedian setae vs. 10 or 11 lateral
and 6 or fewer paramedian setae) (Titova 1983: fig. 9).

Tygarrup fimbriatus Jiang & Huang, sp. nov.
https://zoobank.org/60EDCB5D-4A74-48 10-9394-60 1036025430
Figs 2A-C, 6

Material examined. Holotype. «© 2 (CMMI
20240311004D), China, Xizang Autonomous Regions,
Medog County, Hanmi (29.3619°N, 95.1337°E), 2000 m
asl., 11.111.2024, leg. Chao Jiang.

Paratypes.*39 2 443.3,,(CMMI20240311005D—007D,
009D-012D), same as holotype.

Other materials. * 12 (CMMI 20240311013D), Cui-
NA, Xizang Autonomous Regions, Medog County, Beibeng
village (29.2488°N, 95.1901°E), 920 m asl., 11.11.2024,
leg. Chao Jiang, 12 (CMMI 20240313011D), Damu vil-
lage (29.4864°N, 95.4547°E), 1350 m asl., 8.111.2024, leg.
Chao Jiang; * 999 444, (CMMI 20240307001D-009D,
20240313001D—004D), Bomi County, Yigong village, near

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Tongmai Bridge (30.0989°N, 95.0673°E), 2050 m asl.,
7-13.111.2024, leg. Chao Jiang, 1 2 (CMMI 20240308004D),
Gu village, G319 National Road (29.9973°N, 95.3092°B),
2405 masl., 13.111.2024, leg. Chao Jiang.

Diagnosis. A 7Zygarrup species with 45 leg-bearing
segments. Body length of adult reaches 5.5 cm. Cephalic
frontal plate postero-lateral corner bears 10—20 pore-like
sensilla. Clypeal plagula with 9-12 pairs of setae along
the anterior margin and several pore-like sensilla on ante-
ro-lateral corners. Labral posterior alae fringed with hair-
like projections along the posterior margins. Forcipular
trochanteroprefemur with a tooth, tarsungulum without
tubercle. Sternal pores present in males only.

Description. Maximum length of male 4.1 cm, and of
females, 5.5 cm.

Color (of preserved specimens in alcohol): Head and
forcipular segment reddish-brown, trunk and legs homo-
geneously yellow, without dark patches (Fig. 2A—C).

Cephalic plate (Fig. 6A) 1.4 < longer than the wid-
est. Transverse suture rounded. Lateral margins sinuous,
convergent backwards, anterior margin convex, with a
median incision, posterior margin rounded. Frontal plate
postero-lateral corner bears 10—20 pore-like sensilla. Se-
tae arranged nearly symmetrically.

Clypeus (Fig. 6B) 2.1 < wider than long. Clypeus with
an entire plagula covering most of the clypeus, areolation
only present along the anterior margin of the head; clypeal
setae as follows: | pair on the areolate part, 4 (considered
as damaged and repaired in left side of holotype)—10 pairs
along the anterior margin of the clypeal plagula, 1 pair on
the central part of plagula. Six to twelve pore-like sensilla
on each antero-lateral corner.

Labrum: Mid-piece ca. 1.8 x longer than wide, pos-
teriorly pointed and well sclerotized. Anterior ala trian-
gular; medial margin reduced, distinct shorter than lat-
eral margin; posterior margin of each side-piece sinuous
(Fig. 6B); posterior alae fringed with hair-like projections
along the posterior margins (Fig. 6C).

Cephalic pleurite without spiculum or setae; areolate
part present along anterior margin and paraclypeal suture;
stilus well developed, nearly straight, with an anterior in-
cision on each side (Fig. 6B).

First maxillae (Fig. 6D): Coxosternite 1.7 < wider
than long, divided by mid-longitudinal suture, anterior
corners of coxosternum not projecting. Medial projection
about 1.4 < wider than long, internal margin with several
setae, distal lobe curved inward; telopodite about 2.3 x
longer than wide, curved inward.

Second maxillae (Fig. 6D): Coxosternite undivided,
ca. 1.4 x wider than long; both the anterior and poste-
rior margins concave, lateral margins concave medi-
ally, posterior corners only slightly projecting exter-
nally; areolation on medial part, setae present along
the posterior part; the foraminal process surround-
ing the metameric pore separated from lateral margin
of coxosternite. Telopodites of the second maxillae
well developed, surpassing those of the first maxillae;

Zoosyst. Evol. 101 (2) 2025, 533-550 543

Figure 6. 7ygarrup fimbriatus sp. nov., holotype. A. Cephalic plate, dorsal; B. Cephalic plate, ventral (maxillary complex removed);
C. Labrum, dorsal; D. Maxillary complex, ventral; E. Mandible, ventral; F. Forcipular segment, ventral; G. Forcipular segment,
dorsal; H. Seventh sternum of leg-bearing segment, ventral; I. Ultimate leg-bearing segment and left leg, ventral. Abbreviations:
pls — pore-like sensilla, hlp — hair-like projections. Scale bars: 500 um (A, B, D, F—I; 250 um (C); 25 um (E).

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544 Jiang, C. et al.: Tygarrup centipedes from Xizang, China (Geophilomorpha, Mecistocephalidae)

article 1 of the telopodite 3—3.3 x longer than wide,
curved outward; article 2 of the telopodite ca. 1.1 x lon-
ger than wide; article 3 ca. 1.8 x longer than wide, with
distal end densely covered with setae; apical claw simple.

Mandible with ca. 10—12 well-developed and ca. 2 ru-
dimentary pectinate lamellae; the first lamella with 4—5
teeth, similar in size; intermediate lamellae with 9-12
teeth (Fig. 6E).

Forcipular segment: Coxosternite 1.4 < wider than
long, anterior margin with a pair of truncated teeth (Fig.
6F); scapular points of pleura reaching the anterior mar-
gin of coxosternite (Fig. 6G). Forcipular trochantero-
prefemur about 1.4 = longer than wide and has a conical
distal tooth (Bomi population) or a truncated distal tooth
(Medog populations), intermediate articles with or with-
out tubercle, tarsungulum without tubercle.

Leg-bearing segments: A total of 45 leg-bearing seg-
ments. Sternal sulcus bifurcate faintly impressed, with an
acute angle (Fig. 6H); sternal pores present on males, ab-
sent on females.

Ultimate leg-bearing segment densely setose. Tergite
sub-rectangular, 1.1 x longer than wide. Coxopleuron as
long as wide and 1.7 x longer than sternite. Ca. 50-80
coxal pores on ventral and lateral sides of each coxopleu-
ron, the innermost two to four pores usually larger than
others (Fig. 61). The ultimate leg telopodite not swollen,
with scattered setae, without pretarsus.

Postpedal segments densely setose for both sexes.
Male gonopods bi-articulate, narrow, and separated by
a conical projection. Female gonopods simple, sub-
triangular, and touching each other at their bases (Fig.
61). Anal pores present on each ventro-lateral side of
postpedal segments.

Etymology. Latin fimbr = fiber; the new species name
refers to the presence of hair-like projections along the
posterior margins of the labral posterior alae. We suggest
the Chinese common name as “Wi 7p HARM”.

Distribution. China (Xizang).

Remarks. This new species resembles 7) multiporus
Sp. nov., especially in the presence of pore-like sensilla on
the clypeal plagula and cephalic plate. However, it can be
distinguished from the latter by several distinct features:
it possesses 9-12 pairs of setae along the anterior margin
of the clypeal plagula, has hair-like projections along the
posterior margins of the labral posterior alae, trunk along
with dark patches, and exhibits a larger body size and el-
evated coxopleural pore numbers. In contrast, 7. multipo-
rus sp. nov. has only 3{+4 pairs of setae along the anterior
margin of the clypeal plagula, smooth posterior margins
on the labral posterior alae, and a homogeneously yellow
trunk without dark patches.

The presence of hair-like projections along the posteri-
or margins of the labral posterior alae, a character typical-
ly associated with the genus Dicellophilus Cook, 1896, 1s
reported in the genus 7ygarrup for the first time. These
projections are clearly shorter in 7ygarrup than in Di-
cellophilus (Bonato et al. 2010b; Tsukamoto and Eguchi
2024). This distinction highlights the unique adaptation
of the new species within the 7ygarrup genus.

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Tygarrup poriger (Verhoeff, 1942)
Figs 2D-F, 7

Brahmaputrus poriger Verhoeff, 1942: 51-52, figs 15-21.
Tygarrup poriger: Crabill 1968: 286-287.
Tygarrup nepalensis Shinohara, 1965: 303-306, figs 1-11. syn. nov.

Material examined. * 12 spms (CMMI 20240309001D,
-003D, -005D, -008D, -013D, -014D, -016D,
-018D—-020D, 20240311001D, 20240311002D), Cui-
NA, Xizang Autonomous Regions, Medog County,
near Renqingbensi Temple (29.3075°N, 95.3573°E),
1920 m asl., 9-11.111.2024, leg. Chao Jiang, * 10 spms
(CMMI 20240310002D, -O06D-—O009D, -011D, -014D,
20240312005D, -007D, -013D) near Deergong village
(29.1810°N, 95.1439°E), 1710 m asl., 10—12.111.2024,
leg. Chao Jiang, * 3 soms (CMMI 20240310016D, -017D,
-019D), Shuwang Forestry (29.1946°N, 95.1920°E),
1780 masl., 10.11.2024, leg. Chao Jiang.

Diagnosis. A Zygarrup species with 45 leg-bearing
segments. Clypeal setae 7-9 pairs. Forcipular trochan-
teroprefemur and tibia, each with a tooth, femur without,
tarsungulum with a triangular tooth on the dorsal base.
Sternal pores present in males only.

Re-description. Maximum length of male 3.6 cm and
female 3.3 cm.

Color (of preserved specimens in alcohol): head and
forcipular segment dark red, trunk and legs yellow, with
dark patches along the trunk (Fig. 2D-F).

Cephalic plate (Fig. 7A) 1.5 < longer than the wid-
est. Transverse suture rounded. Lateral margins nearly
straight and convergent backwards, anterior margin con-
vex, with a median incision, posterior margin rounded.
Setae arranged nearly symmetrically.

Clypeus 2 x wider than long (Fig. 7B). Clypeus with
an entire plagula covering most of the clypeus, areola-
tion only present along the anterior margin of the head;
clypeal setae as follows: 1 pair on the areolate part, 3—5
pairs along the anterior margin of the clypeal plagula, 2
pairs on the antero-medial part inside a large insula, and 1
pair on the central part of plagula (Fig. 7C).

Labrum (Fig. 7C): Mid-piece ca. 1.7 < longer than
wide, posteriorly pointed and well sclerotized. anterior
ala triangular, medial margin reduced to a vertex; posteri-
or margin of each side-piece sinuous.

Cephalic pleurite without spiculum or setae; areolate
part present along anterior margin and paraclypeal suture;
stilus well developed, nearly straight, with an anterior in-
cision on each side (Fig. 7B).

First maxillae (Fig. 7D): Coxosternite 1.8 < wider
than long, divided by mid-longitudinal suture, anterior
corners of coxosternum not projecting. Medial projec-
tion about 1.2 x wider than long, internal margin with
several setae, distal lobe curved inward; telopodite about
2.1 x longer than wide, curved inward.

Second maxillae (Fig. 7D): Coxosternite undivided,
ca. 1.4 x wider than long; both the anterior and poste-
rior margins concave, lateral margins concave medial-
ly, posterior corners only slightly projecting externally;

Zoosyst. Evol. 101 (2) 2025, 533-550 545

Figure 7. 7vgarrup poriger (Verhoeff, 1942) (spm. CMMI 20240309001D). A. Cephalic plate, dorsal; B. Cephalic plate, ventral
(maxillary complex removed); C. Clypeus, ventral (maxillary complex removed); D. Maxillary complex, ventral; E. Mandible, ven-
tral; F. Forcipular segment, ventral; G. Forcipular segment, dorsal; H. Seventh sternum of leg-bearing segment, ventral; I. Ultimate
leg-bearing segment and left leg, ventral. Scale bars: 500 um (A, B, D, F—I); 100 um (C); 25 um (E).

areolation on medial part, setae present along the pos- Telopodites of the second maxillae developed, surpassing
terior part; the foraminal process surrounding the meta- __ those of the first maxillae when stretched; article 1 of the
meric pore separated from lateral margin of coxosternite. _telopodite ca. 3.1 < longer than wide, curved outward;

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546 Jiang, C. et al.: Tygarrup centipedes from Xizang, China (Geophilomorpha, Mecistocephalidae)

article 2 of the telopodite ca. 1.6 x longer than wide; ar-
ticle 3 ca. 2.2 x longer than wide, with distal end densely
covered with setae; apical claw large and simple.

Mandible with ca. 7 well-developed and ca. 2 rudimen-
tary pectinate lamellae; the first lamella with 5 teeth, simi-
lar in size; intermediate lamellae with 9-12 teeth (Fig. 7E).

Forcipular segment: Coxosternite 1.3 x wider than long,
anterior margin with a pair of truncated teeth (Fig. 7F); scap-
ular points of pleura reaching the anterior margin of cox-
osternite (Fig. 7G). Forcipular trochanteroprefemur about
1.3 x longer than wide and has a prominent distal tooth,
femur with or without tubercle, tibia with a small pointed
tubercle, tarsungulum with a triangular tooth on dorsal base.

Leg-bearing segments: A total of 45 leg-bearing seg-
ments. Sternal sulcus bifurcate faintly impressed, with an
acute angle (Fig. 7H); sternal pores present on males, ab-
sent on females.

Ultimate leg-bearing segment densely setose. Tergite
sub-rectangular, 1.1 =< longer than wide. Coxopleuron
about 0.9 x longer than wide and 1.6 x longer than stern-
ite. Ca. 20-35 coxal pores on ventral and lateral sides of
each coxopleuron, the innermost two pores usually larger
than others (Fig. 71). The ultimate leg telopoditenot swol-
len, with scattered setae, without pretarsus.

Postpedal segments densely setose for both sexes.
Male gonopods bi-articulate, narrow, and separated by
a conical projection. Female gonopods inconspicuous
bi-articulate, subtriangular, and touching each other at
their bases (Fig. 71). Anal pores present on each ven-
tro-lateral side of postpedal segments.

Distribution. China (Xizang), Nepal.

Remarks. Verhoeff (1942) originally described a new
genus, Brahmaputrus, and type species, B. poriger, from
Shigatse, Xizang, of China, with 45 leg-bearing segments
and males with sternal pores. Crabill (1968) considered
the genus Brahmaputrus Verhoeff, 1942, as a synonym
of Zygarrup Chamberlin, 1914. This species differs from
other 7Zygarrup species for its forcipular trochantero-
prefemur with a tooth, femur without, and tarsungulum
with a tooth on dorsal base.

Shinohara (1965) originally described a Tygarrup spe-
cies, 7’ nepalensis, based on 28 specimens from Ralwaling
Himal, East Nepal. However, illustrations given by Shino-
hara (1965: figs 5-11) show minor variation with 7° poriger
sensu Verhoeff (1942: figs 15—21) and the present study (Fig.
7) in the clypeal setae, forcipular articles, maxillae, shape of
the ultimate leg-bearing segment, and coxal pore numbers,
particularly the dorsal tooth on the forcipular tarsungulum.
Therefore, Zygarrup nepalensis Shinohara, 1965, is treated
as a junior synonym of 7ygarrup poriger (Verhoeff, 1942).

Tygarrup muminabadicus Titova, 1965
Figs 2G-I, 8

Tygarrup muminabadicus Titova, 1965: 871-876, figs 1, 2; Bonato et al.
2003: figs 8C, 9E, 10A, 13B; Dyachkov 2020: 80, 2022:75, figs 5-11.

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Material examined. ¢ 3 soms, (CMMI 20220525124, -126,
-128), CuiNa, Xizang Autonomous Regions, Jilong County,
near Rukaer(28.4481°N, 85.2467°E),2970masl.,25.v.2022,
leg. Xiaoxia Tian; * 2 soms (CMMI 20210705142, CMMI
20210705144), Luozha County, Lakang town, (28.1125°N,
91.1251°E), 3230 m asl., 5.vil.2022, leg. Chao Jiang; « 1
spms (CMMI 20210708102), Jiacha County, Anrao town
(29.1420°N, 92.5829°E), 3240 m asl., 8.vii.2021, leg. Chao
Jiang; 1 som (CMMI 20220620103), Linzhi, Bayi district,
Sejilashan Mt. (29.6213°N, 94.6514°E), 20.vi.2022, leg.
Xiaoxia Tian, * 5 spms (CMMI 20201013119D-123D),
Fuqing Road, 13.x.2020, leg. Chao Jiang, * 2 spms
(CMMI 20201013137, CMMI 20201013142), Birish-
enshan Mt., 13.x.2020, leg. Chao Jiang; 1 spm (CMMI
20240310018D), Medog County, Shuwang Forestry
(29.1946°N, 95.1920°E), 1780 m asl., 10.11.2024, leg.
Chao Jiang, * 7 spms (CMMI 20240310001D, -003D,
-004D, -015D, CMMI 20240312001D, -008D, -009D),
near Deergong village (29.1810°N, 95.1439°E), 1710 m
asl., 10—12.111.2024, leg. Chao Jiang.

Diagnosis. A Zygarrup species with 45 leg-bearing
segments. Body length of adult reaches 5 cm. Clypeal
plagula with 6—9 pairs of setae along the anterior margin;
plagula without pore-like sensilla. Labral posterior alae
posterior margins smooth, without hair-like projection.
Forcipular trochanteroprefemur with a tooth, femur, and
tarsungulum without tooth. Sternal pores present in males
only. Coxopleuron with ca. 40-70 coxal pores. Female
gonopods unarticulated.

Distribution. China (Xizang), Tajikistan, Kashmir.

Remarks. Specimens from Xizang could be assigned
only tentatively to 7’ muminabadicus, as they are largely
consistent with the original description (Titova 1965) and
topotypes (Dyachkov 2022) of this species, only differing
for a more inward-curved first maxillary telopodite (clav-
ate in the original illustration) and for the elevated number
of coxal pores on coxopleuron. This species has been pre-
viously documented in Tajikistan (Titova 1965; Dyachkov
2020, 2022) and Kashmir (Bonato et al. 2003) and is now
reported for the first time in the Chinese fauna. The mor-
phological characters of Chinese specimens are delineated
through the accompanying illustrations (Figs 2G—I, 8).

T. diversidens, a Himalayan species that was original-
ly described by Silvestri (1919) as Lamnonyx diversidens.
This description was based on numerous specimens across
a wide area of the Himalayas, extending from Assam in
the east to Kashmir in the west. These distributions also in-
clude two localities in Xizang (Rotung and Renging) and
are considered Xizang’s Tygarrup species. The species
also has the inwardly curved first maxillary telopodite (Sil-
vestr1 1919: fig. XX5) but is distinguished from 7? mumin-
abadicus by having bi-articulate female gonopods (Silves-
tri 1919: fig. XX8). However, the original description of
T: diversidens is very inadequate, leaving the taxonomic
relationship of this nominal species with other 7yvgarrup
species from the Himalayas undetermined, necessitating
further investigation (Bonato and Minelli 2004).

Zoosyst. Evol. 101 (2) 2025, 533-550 547

Figure 8. 7ygarrup muminabadicus Titova, 1965 (spm. CMMI 20240310001D). A. Cephalic plate, dorsal; B. Cephalic plate, ven-
tral (maxillary complex removed); C. Maxillary complex, ventral; D. Mandible, ventral; E. Forcipular segment, ventral; F. Forcip-
ular segment, dorsal; G. Seventh sternum of leg-bearing segment, ventral; H. Ultimate leg-bearing segment and left leg, ventral.
Scale bars: 500 um (A—C, E—H); 25 um (D).

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548 Jiang, C. et al.: Tygarrup centipedes from Xizang, China (Geophilomorpha, Mecistocephalidae)

Discussion

In the present study, we investigated the genus 7ygar-
rup from Xizang and identified six species within the
genus (Fig. 9). Certain morphological characters, such
as pore-like sensilla, cerrus, and the hair-like projec-
tions along the posterior margins of the labral posterior
alae, were discovered for the first time to be effective
in distinguishing species within the 7ygarrup genus for
the first time. The introduction of these new morpho-
logical characters allows for a more accurate differen-
tiation of Zygarrup species in Xizang, contributing to a
better understanding of the diversity of soil centipede
species in this region.

Tygarrup species from Xizang dwell in both tropical
dry forests and alpine environments. The lowest record-
ed altitude for their habitat is 920 meters (T. fimbriatus
sp. nov.), while the highest extends up to 4400 meters
(Tf. tridentatus sp. nov.). Among them, 7: tridentatus
sp. nov. and 7. poriger can inhabit both cold mountain
regions and low-altitude warmer forests. In contrast, 7.
cerrus sp. nov. is found exclusively in high mountain
coniferous forests at around 3000 meters in altitude.
Based on our available specimens and historical re-
cords (Verhoeff 1942; Shinohara 1965), most 7ygarrup

84°E 86°E 88°E
36°N
7. poriger

T. muminabadicus

T. cerrus sp. nov.

T. multiporus sp. nov.

T’ tridentatus sp. nov.

T: fimbriatus sp. nov.

26°N

‘
A

84°E 86°F 88°E

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90°E

90°E

species in Xizang live at altitudes above 1700 meters,
while Zygarrup species in other regions predominant-
ly inhabit forests or greenhouses below 1500 meters
(Silvestri 1919; Attems 1929; Titova 1983; Bonato et
al. 2004; Uliana et al. 2007; Bonato and Minelli 2010;
Dyachkov 2020, 2022). Historical literatures and our
results also show that the number of Zygarrup spe-
cies in the Himalayan mountainous regions within the
same scale is higher than in the northern Qinghai-Tibet
Plateau region as well as in the southern tropical rain-
forests and Gangetic Plains (Silvestri 1919). The com-
plexity of mountains is usually tightly associated with
high biodiversity (Perrigo et al. 2020). In multiple an-
imal and plant groups, the uplift of the Himalayas has
promoted biological diversification (Xia et al. 2022;
Liu et al. 2024). Therefore, the Himalayas, with their
complete tropical monsoon mountain vertical natural
zones ranging from lowland tropical monsoon rain-
forests to alpine snow zones, may have facilitated the
diversification of the 7ygarrup species in Xizang. Con-
sidering the diversity of the Himalayan ecosystem, the
sampled regions available are still only fragmentary
and insufficient for the diversity research of 7ygarrup.
Therefore, the faunal diversity in this region is likely
still underestimated.

92°E 94°E 96°E

36°N

28°N

26°N

92°R

Figure 9. Collected localities of species of 7ygarrup in Xizang of China. Stars represent type localities.

94° 96°E

Zoosyst. Evol. 101 (2) 2025, 533-550 549
Key to the six confined 7ygarrup species of China

1 Cerrus composed ot we lateral*eroups of Setde- (Rig. AO. ...9.ctatecisscccstefschs des Sees. caeesslidessdedgredsacere sect T. cerrus Sp. nov.
- COnAIS: GUSCUE Jak nare zeae at ameere sine! tek en tases oom genius, wee etn se de bles nated oR octal hai ele rose te Mink ual aad on Melb 2
2 ~-Antéro-lateralcorners.ofsplagula eviciently.bear pore-like: sensilla. cus. essen ee AS estos Ws ee ea OG Se 3
- Anterolateral corners-ot plaguila absent oore-iIke-sensi lla cnr. seers sees ees ee erg as Po ea es Se es ere en eee che Sens 4
3 Posterior margins of labral posterior alae bear hair-like projections (Fig. OC) ............:ceceeeeeeeeeeeeee ees T. fimbriatus sp. nov.
- Posterior margins of labral posterior alae absent hair-like projection (Fig. 5B).............::::eeeeeeeeeeee es T. multiporus sp. nov.
4 Central part of plagula with ca. 15 paired setae arranged tn three irregular rows (Chao et al. 2020: fig. 3)... 7. daliensis
- Central part of plagula with 6-11 paired setae arranged in two irregular rOWS............cccccccceceeeeeeeeceeeeeeeaeeeeeeeseeeseeeeaes 5
5 Forced ay tarsey ee) Maret NceehOOUNe. oo cage. < com States ns cerechitchac.cqosaplacder exkat bye on sem Wadedacsne sch teh «doraprneedae cbt lgarack -ecbedadetabet seen 6

- Forcipular tarsungulum without tooth (Fig. 8F)...............

hase, Shon WENT «0 hot aagl anteater Pha Ne in I, Un OPIS T. muminabadicus

6 Forcipular article Ill and tarsungulum with a prominent pointed tooth (Fig. 3F, G).............::: eee T. tridentatus sp. nov.
- Forcipular article Ill with a small tubercle, tarsungulum with a triangular tooth on dorsal base (Fig. 7G).......... T. poriger
Acknowledgements Bonato L, Edgecombe GD, Lewis JG, Minelli A, Pereira LA, Shelley

We are grateful to Dr. Huigin Ma (Hengshui Universi-
ty, China) for providing important literature. We wish
to thank Ms. Jiaxin Dong and Ms. Yaru Zhang for their
assistance in preparing the line drawings of the new spe-
cies. We sincerely thank Dr. Huiqin Ma and the anony-
mous reviewer for providing valuable comments that sig-
nificantly improved this paper.

The research was supported by the Key Project at
Central Government Level: the Ability Establishment of
Sustainable Use for Valuable Chinese Medicine Resourc-
es (nos. 2060302), the CACMS Innovation Fund (nos.
CI2023E002), and the Survey of Wildlife Resources in
Key Areas of Xizang (Phase IT) (nos. ZL202303601).

References

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