Research Article |
Corresponding author: Chao Zhang ( opiliones@163.com ) Academic editor: Shahan Derkarabetian
© 2024 Xiaoru Qi, Adriano B. Kury, Chao Zhang.
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.
Citation:
Qi X, Kury AB, Zhang C (2024) A new genus of Assamiidae (Opiliones, Grassatores) from Xizang, China. ZooKeys 1215: 27-52. https://doi.org/10.3897/zookeys.1215.132189
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Knowledge of the family Assamiidae in the Eastern Himalayas is primarily concentrated from Nepal, while Bhutan, Xizang, and northeastern India remain much less studied. Herein, a new genus, Linzhiassamia gen. nov., is described from Xizang, China, along with two new species, Linzhiassamia medogensis sp. nov. and Linzhiassamia zayuensis sp. nov. These represents the first records of the family Assamiidae from Xizang. A comparative analysis with other continental Asian Assamiidae is undertaken, with a focus on genital morphology. Potential closest relatives are identified, including the genus Dhaulagirius Martens, 1977 from Nepal.
Arachnida, Eastern Himalayas, genitalia, harvestmen, taxonomy
The Paleotropical family Assamiidae encompasses more than 450 valid species (
Modern descriptions of assamiid species are sparse and even combined, they fail to encompass their entire taxonomic and geographic range. Specifically,
In this article, two new species from Xizang are described. Unable to find an existing genus that matches their characters, we have erected the new genus Linzhiassamia gen. nov. Additionally, we describe the intraspecific variation of L. zayuensis sp. nov., providing detailed morphological descriptions, observations of genitalia, and discussions on the taxonomic position of the new species.
Assamiidae was initially a monogeneric family for Assamia Sørensen, 1884 from the Indian highlands.
Recent trends in taxonomy have seen a dismissal of subfamilies that are not clades. For instance,
The specimens were preserved in 75% ethanol, examined under a Leica M205A stereomicroscope, and the overall drawings were made using a drawing tube-equipped Leica M205A stereomicroscope, while the detailed drawings were created using Inscape v. 1.3. Photographs were taken using a Leica M205A stereomicroscope, equipped with a DFC 450 CCD. The male genitalia were initially placed in hot lactic acid (40–50 °C) for about 1–2 min, then transferred to distilled water; the movable parts of the glans will mostly expand within 1 min (
The terminology of genital structures follows
In the diagnosis given here, Linzhiassamia sp. nov. is assessed in relation to genera that possess some features in common, such as the Nepalese highlanders Dhaulagirius Martens, 1977; Micrassamula Martens, 1977; Nepalsia Martens, 1977; and Nepalsioides Martens, 1977, as well as Nilgirius Roewer, 1915, which occurs in Yunnan and India. It is also compared with Paktongius Suzuki, 1969, from the lowlands of Laos, Malaysia, and Thailand.
Linzhiassamia medogensis sp. nov. (type species) and Linzhiassamia zayuensis sp. nov.
The genus name is based on 林芝 (Linzhi), an alternative name for Nyingchi. This is associated with the pre-existing genus name Assamia. The gender is feminine.
Linzhiassamia is similar to Paktongius for the sexually dimorphic coxa IV, which in males may reach areas III, IV or even V of the dorsal scutum, while in females it is much shorter, reaching area II, and not projected laterally (Nilgirius and the Nepalese genera treated here, all possess coxa IV monomorphic).
Linzhiassamia is similar to Dhaulagirius due to the sexually dimorphic chelicerae; however, in Dhaulagirius, the cheliceral hand is much more exaggeratedly developed. In the other genera compared here, the chelicerae are monomorphic.
Linzhiassamia is similar to Nilgirius and Paktongius for having a “pseudonychium” (tarsal process) in the tarsi of legs III and IV.
Linzhiassamia is similar to Dhaulagirius, Micrassamula, and Nepalsia for lacking a sharp annular joint-like constriction in the distal third of the truncus penis, between the pars basalis and pars distalis (in contrast to Nepalsioides and Nilgirius).
Linzhiassamia is similar to Dhaulagirius, Nepalsia, and Nepalsioides for having MS A organized in a triangle (in contrast to Micrassamula, which has MS A extended into a line, and Nilgirius, which has a ring-girdle formed by macrosetae D1, A1–A3, and B1).
Linzhiassamia is similar to Dhaulagirius, Nepalsia, and Nilgirius for not having a wide dorsal concavity, giving the distal truncus the aspect of an ice-cream scoop (in contrast to Micrassamula and Nepalsioides).
Linzhiassamia is similar to Nepalsia and Nilgirius for having all macrosetae concentrated distally (in contrast to Dhaulagirius, Micrassamula, and Nepalsioides).
Linzhiassamia is similar to Dhaulagirius for having the distal part of the truncus with constrictions, flaring to form a pyriform structure (in contrast to Micrassamula, Nepalsia, Nepalsioides, and Nilgirius, which have more or less continuous widening of the truncus, getting rounded apically).
Holotype • male (MHBU-Opi-24ZC011501): China: Xizang, Nyingchi, Medog County, 29°33'N, 95°33'E, alt. 1116 m, 22 May 2019, H. Wang, L.Y. Wang leg. Paratypes: • one male (MHBU-Opi-24ZC011502) and two females (MHBU-Opi-24ZC011503-04), same collecting data as holotype • one male (MHBU-Opi-24ZC011601): China: Xizang, Nyingchi, Medog County, 29°33'N, 95°33'E, alt. 1116 m, 23 May 2019, H. Wang leg.
Distal section of penis (pars distalis) markedly enlarged: ventral plate nearly triangle and frontal rim with median crevice (Figs
The species name originates from the specimen collection site: Medog County, Nyingchi, Xizang.
Male (holotype and paratype). Habitus as in Figs
Linzhiassamia medogensis sp. nov., male (2–6 holotype), male (1 paratype) 1 male body, lateral view 2 left chelicera of male, ental view 3 left chelicera of male, ectal view 4 left pedipalp of male, dorsal view 5 left pedipalp of male, ental view 6 left pedipalp of male, ectal view. Scale bars: 1 mm (1); 0.5 mm (2–6).
Linzhiassamia medogensis sp. nov., male (9, 11 holotype), male (7 paratype), female (8, 10, 12 paratype) 7 male body, dorsal view 8 female body, dorsal view 9 left cheliceral fingers of male, frontal view 10 left cheliceral fingers of female, frontal view 11 right tarsal claw IV of male, lateral view 12 right tarsal claw IV of female, lateral view. Scale bars: 1 mm (7, 8); 0.5 mm (11–12); 0.25 mm (9–10).
Linzhiassamia medogensis sp. nov., genitalia of male holotype (13–17) and female paratype (18–19) 13 penis, ventral view 14 distal part of penis, lateral view 15 same, dorsal view 16 distal part of penis (expanded), lateral view 17 same, dorsal view 18 ovipositor, dorsal view 19 same, ventral view. Pb, pars basalis, Pd pars distalis. Scale bars: 0.25 mm.
Dorsum
(Figs
Venter (Fig.
Chelicera
(Figs
Pedipalpus
(Figs
Legs. Slender and elongated. Trochanters I–IV with small, hair-tipped granules on the ventral surface. All femora with hair-tipped granules, femora III and IV curved. Tarsi III–IV with a pseudonychium and two bare claws (Fig.
Penis
(Figs
Female (Figs
Ovipositor
(Figs
Male holotype (female paratype): Body 3.53 (3.53) long, 2.29 (2.00) wide at the widest portion. Scutum 1.85 (1.64) long. Interocular mound 0.63 (0.57) long, 0.34 (0.34) wide, 0.24 (0.20) high, 0.13 (0.12) far from the anterior border of the scutum. Pedipalpal claw 0.39 (0.45) long. Penis 1.02 long. Measurements of left pedipalpus and legs as in Tables
Linzhiassamia medogensis sp. nov. Measurements of the pedipalp and legs of the male holotype (MHBU-Opi-24ZC011501), as length/width.
Trochanter | Femur | Patella | Tibia | Metatarsus | Tarsus | Total | |
---|---|---|---|---|---|---|---|
Pedipalp | 0.47/0.22 | 1.09/0.32 | 0.78/0.24 | 0.67/0.29 | 0.58/0.28 | 3.59 | |
Leg I | 0.36/0.25 | 1.73/0.21 | 0.63/0.28 | 1.26/0.18 | 1.98/0.09 | 1.18/0.05 | 7.14 |
Leg II | 0.39/0.30 | 3.22/0.20 | 0.94/0.29 | 2.72/0.16 | 3.24/0.10 | 2.52/0.07 | 13.03 |
Leg III | 0.46/0.37 | 2.30/0.24 | 0.81/0.35 | 1.51/0.21 | 2.68/0.16 | 1.40/0.11 | 9.16 |
Leg IV | 0.60/0.36 | 3.75/0.26 | 1.02/0.42 | 2.23/0.21 | 4.32/0.18 | 1.89/0.12 | 13.81 |
Linzhiassamia medogensis sp. nov. Measurements of the pedipalp and legs of the female paratype (MHBU-Opi-24ZC011503), as length/width.
Trochanter | Femur | Patella | Tibia | Metatarsus | Tarsus | Total | |
---|---|---|---|---|---|---|---|
Pedipalp | 0.44/0.22 | 0.97/0.28 | 0.70/0.24 | 0.62/0.28 | 0.54/0.24 | 3.27 | |
Leg I | 0.45/0.28 | 2.45/0.16 | 0.72/0.25 | 2.35/0.15 | 2.83/0.08 | 2.08/0.08 | 10.88 |
Leg II | 0.46/0.29 | 2.75/0.18 | 0.74/0.24 | 2.44/0.15 | 2.71/0.08 | 2.04/0.08 | 11.14 |
Leg III | 0.42/0.35 | 2.03/0.23 | 0.58/0.31 | 1.32/0.22 | 2.21/0.13 | 1.21/0.11 | 7.77 |
Leg IV | 0.54/0.33 | 3.09/0.23 | 0.78/0.32 | 1.92/0.23 | 3.45/0.14 | 1.52/0.08 | 11.30 |
The specimens were collected under stones and on the leaves of the shrubbery.
Known only from the type locality, the Medog County, Nyingchi City, Xizang Autonomous Region, China.
Specimens examined included three males and two females, the number of tarsal segments on the second legs was not constant, which varied from eleven to fourteen segments. On the fourth leg, the number of tarsal segments varies from seven to eight. The number of tarsal segments on the second leg and third leg is constant, with five segments on the second and six segments on the third. Another variation is the number of setiferous tubercles on the pedipalpus trochanter. For example, the male holotype (MHBU-Opi-24ZC011501) has only one setiferous tubercle on the pedipalpus trochanter (Figs
Holotype • male (MHBU-Opi-24ZC011801): China: Xizang, Nyingchi, Zayu County, 28°29'N, 97°30'E, alt. 1405 m, 13 July 2020, L.Y. Wang leg. Paratypes: • one female (MHBU-Opi-24ZC011802), China: Xizang, Nyingchi, Zayu County, 28°77'N, 96°72'E, alt. 1945 m, 27 May 2019, H. Wang leg • one male and one female (MHBU-Opi-24ZC011803-04), China: Xizang, Nyingchi, Zayu County, 28°53'N, 96°99'E, alt. 1509 m, 11 May 2023, Y. M. Hou, Z. Y. Yang leg • one male (MHBU-Opi-24ZC011901), China: Xizang, Nyingchi, Lulang Town, 29°96'N, 94°82'E, alt. 2472 m, 21 May 2019, H. Wang leg • one female (MHBU-Opi-24ZC011902), China: Xizang, Nyingchi, Bome County, 30°10'N, 95°07'E, alt. 2037 m, 02 June 2022, B. Liu leg • one male and one female (MHBU-Opi-24ZC011903-04), China: Xizang, Nyingchi, Bome County, 30°04'N, 95°02'E, alt. 2051 m, 17 July 2020, L. Y. Wang, Y. M. Hou, leg.
The distal margin of the ventral plate is smooth and without any indentation (Figs
The external morphological differences of this species are significant. The interocular have either a short spine or are smooth, and at the lateral portion there are two or three spines on the anterior margin of the carapace. However, the seta-tipped tubercles on the opisthosomal region of scutum are relatively small, which allows for a preliminary differentiation from the other three species. Additionally, by examining the expanded structure of the genitalia together with external morphological characteristics, this species can be accurately distinguished from others.
The name of this species is derived from its collection locality in Zayu County, Nyingchi, Xizang.
Male (holotype and paratype). Habitus as in Figs
Dorsum
(Figs
Linzhiassamia zayuensis sp. nov., male (27–31 holotype), male (26 paratype) 26 male body, lateral view 27 left chelicera of male, ental view 28 left chelicera of male, ectal view 29 left pedipalp of male, dorsal view 30 left pedipalp of male, ental view 31 left pedipalp of male, ectal view. Scale bars: 1 mm (26); 0.5 mm (27–31).
Linzhiassamia zayuensis sp. nov., male (34, 36 holotype), male (32 paratype), female (33, 35, 37 paratype) 32 male body, dorsal view 33 female body, dorsal view 34 left cheliceral fingers of male, frontal view 35 left cheliceral fingers of female, frontal view 36 right tarsal claw IV of male, lateral view 37 right tarsal claw IV of female, lateral view. Scale bars: 1 mm (32–33); 0.5 mm (36–37); 0.25 mm (34–35).
Linzhiassamia zayuensis sp. nov., genitalia of male holotype (38–42) and female paratype (43, 44) 38 penis, ventral view 39 distal part of penis, lateral view 40 same, dorsal view 41 distal part of penis (expanded), lateral view 42 same, dorsal view 43 ovipositor, dorsal view 44 same, ventral view. Pb, pars basalis, Pd pars distalis. Scale bars: 0.5 mm (38); 0.25 mm (39–44).
Linzhiassamia zayuensis sp. nov., male (51–56 paratype) 51 male body, lateral view 52 left chelicera of male, ental view 53 left chelicera of male, ectal view 54 left pedipalp of male, dorsal view 55 left pedipalp of male, ental view 56 left pedipalp of male, ectal view. Scale bars: 1 mm (51); 0.5 mm (52–56).
Linzhiassamia zayuensis sp. nov., male (59, 61, 57 paratype), female (58, 60, 62 paratype) 57 male body, dorsal view 58 female body, dorsal view 59 left cheliceral fingers of male, frontal view 60 left cheliceral fingers of female, frontal view 61 right tarsal claw IV of male, lateral view 62 right tarsal claw IV of female, lateral view. Scale bars: 1 mm (57, 58); 0.5 mm (61–62), 0.25 mm (59–60).
Linzhiassamia zayuensis sp. nov., genitalia of male paratype (63–67) and female paratype (68–69) 63 penis, ventral view 64 distal part of penis, lateral view 65 same, dorsal view 66 distal part of penis (expanded), lateral view 67 same, dorsal view 68 ovipositor, dorsal view 69 same, ventral view. Pb, pars basalis, Pd pars distalis. Scale bars: 0.5 mm (63); 0.25 mm (64–69).
Venter
(Fig.
Chelicera
(Figs
Pedipalpus
(Figs
Legs. Slender and elongated. Trochanters I–IV with small hair-tipped granules on the ventral surface. All femora with hair-tipped granules, femora III and IV curved. Tarsi III–IV with a pseudonychium and two bare claws (Fig.
Penis
(Figs
Female (Figs
Ovipositor
(Figs
Male holotype (female paratype): body 3.42 (3.88) long, 2.33 (2.30) wide at the widest portion. Scutum 2.60 (1.74) long. Interocular mound 0.50 (0.54) long, 0.30 (0.23) wide, 0.19 (0.13) high, 0.19 (0.26) far from the anterior border of the scutum. Pedipalpal claw 0.43 (0.33) long. Penis 1.34 long. Measurements of left pedipalpus and legs as in Tables
Linzhiassamia zayuensis sp. nov. Measurements of the pedipalp and legs of the male holotype (MHBU-Opi-24ZC011801), as length/width.
Trochanter | Femur | Patella | Tibia | Metatarsus | Tarsus | Total | |
---|---|---|---|---|---|---|---|
Pedipalp | 0.37/0.21 | 0.93/0.26 | 0.72/0.20 | 0.60/0.25 | 0.51/0.26 | 3.13 | |
Leg I | 0.31/0.23 | 1.32/0.18 | 0.59/0.24 | 1.06/0.17 | 1.49/0.07 | 0.71/0.07 | 5.48 |
Leg II | 0.35/0.27 | 2.27/0.18 | 0.73/0.23 | 1.96/0.19 | 2.13/0.07 | 2.21/0.07 | 9.65 |
Leg III | 0.43/0.34 | 1.69/0.19 | 0.62/0.31 | 1.22/0.24 | 1.91/0.12 | 1.14/0.08 | 7.01 |
Leg IV | 0.52/0.36 | 2.41/0.19 | 0.79/0.36 | 1.74/0.26 | 2.78/0.16 | 1.54/0.10 | 9.78 |
Linzhiassamia zayuensis sp. nov. Measurements of the pedipalp and legs of the female paratype (MHBU-Opi-24ZC011802), as length/width.1
Trochanter | Femur | Patella | Tibia | Metatarsus | Tarsus | Total | |
---|---|---|---|---|---|---|---|
Pedipalp | 0.39/0.19 | 0.89/0.25 | 0.67/0.20 | 0.59/0.22 | 0.53/0.21 | 3.07 | |
Leg I | 0.29/0.21 | 1.30/0.19 | 0.58/0.20 | 0.98/0.17 | 1.40/0.07 | 1.02/0.06 | 5.57 |
Leg II | 0.39/0.25 | 2.17/0.17 | 0.64/0.23 | 1.97/0.17 | 2.16/0.08 | 2.06/0.08 | 9.39 |
Leg III | 0.37/0.27 | 1.71/0.20 | 0.57/0.26 | 1.11/0.21 | 1.86/0.08 | 1.19/0.07 | 6.81 |
Leg IV | 0.43/0.26 | 2.27/0.22 | 0.66/0.27 | 1.55/0.21 | 2.82/0.09 | 1.52/0.07 | 9.25 |
These specimens were collected by sifting through the fallen leaves in the dark and humid undergrowth of the forest, as well as under stones and on the leaves of the shrubbery.
Known only from the type locality, the Zayu County, Bome County, and Lulang Town, Nyingchi City, Xizang Autonomous Region, China.
Five male specimens were examined, displaying two distinct external morphologies, with a male paratype (MHBU-Opi-24ZC011901) chosen for discussion due to its differences from the male holotype (MHBU-Opi-24ZC011801). Compared to MHBU-Opi-24ZC011801, MHBU-Opi-24ZC011901 exhibits darker body coloration, there are no dark brown patches located on the sides of the anterior margin of the prosoma and larger dark brown patches on the opisthosomal areas I–IV (Figs
Females also exhibit variation in morphological characteristics. Similar to the differences observed in males, females also display slight differences in body coloration (Figs
The observed external morphological differences initially led us to consider these as potentially separate species. However, upon dissecting the genitalia, we found remarkable similarities both before and after expansion, with only slight differences in the shapes of the stylus and prickly funnel (Figs
The Qinghai-Xizang Plateau is characterized by high altitude, thin atmospheric layers, and unique geographical and climatic conditions. These factors likely contribute to the observed morphological variations among specimens, particularly given the restricted gene flow between populations at different altitudes due to the limited dispersal capacity of harvestmen. While these variations may suggest the presence of more than one species, the current sample size and geographic coverage do not provide enough evidence to definitively separate these populations into distinct species. A more extensive collection of specimens from a broader range of localities is necessary before any formal taxonomic decisions can be made.
The phylogenetic analysis by
We may endeavor to categorize the existing diversity of continental Asian Assamiidae into coherent groups of species, disregarding the Roewerian subfamilies which rely on superficial traits. This way, evaluating the affinities of any genus within Assamiidae becomes challenging due to the often concise nature of descriptions. To accurately categorize subfamilies, we should reduce each to their type genus, and ideally, to their type species if we wish to designate a subfamily by name.
L. medogensis sp. nov. and L. zayuensis sp. nov. exhibit pseudonychiate tarsal claws, prompting comparisons with Indian Trionyxellidae such as Nilgirius. However, a preliminary examination of the genitalia of Nilgirius scaber Roewer, 1915 from southern India (Kury unpubl. data), reveals a markedly distinct genital structure. Our Tibetan species of Linzhiassamia also feature a pyriform distal truncus, which is not as depressed, with elongated macrosetae similar to those observed in Dhaulagirius altitudinalis Martens, 1977. However, the sexual dimorphism in the chelicerae is much more subtle.
While a comparison with Dhaulagirius is not out of the question, defining suprageneric groups for these highlanders remains a distant prospect. Knowledge of the highlands of the Eastern Himalaya remains fragmentary. There are no assamiids known from Bhutan, and the few species known from Arunachal Pradesh come from modest altitudes (up to 600 m) with only limited descriptions by Roewer.
Many thanks are due to Peter Jäger for providing literature and to Luyu Wang, Hui Wang, Zhiyong Yang, and Yanmeng Hou for collecting the specimens. Special thanks are given to Willians Porto and Shahan Derkarabetian for insightful comments of the manuscript.
The authors have declared that no competing interests exist.
No ethical statement was reported.
This work was supported by (1) the Survey of Wildlife Resources in Key Areas of Tibet (ZL202203601) (2) # E-26/200.085/2019 (Apoio Emergencial ao Museu Nacional) from Fundação de Amparo à Pesquisa do Estado do Rio de Janeiro (FAPERJ) (3) # E-26/210.148/2019 (249116) (APQ1-Auxílio à Pesquisa básica- 2019) from Fundação de Amparo à Pesquisa do Estado do Rio de Janeiro (FAPERJ) (4) #311531/2019-9 (Produtividade em Pesquisa) from the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) (5) #430748/2018-3 (Chamada MCTIC/CNPqN° 28/2018-Universal) from the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) to ABK.
All authors have contributed equally.
Xiaoru Qi https://orcid.org/0009-0006-9458-2532
Adriano B. Kury https://orcid.org/0000-0002-8334-6204
Chao Zhang https://orcid.org/0000-0003-1702-1206
All of the data that support the findings of this study are available in the main text.