Review Article |
Corresponding author: Yucheng Lin ( linyucheng@scu.edu.cn ) Academic editor: Jeremy Miller
© 2019 Ya Li, Yucheng Lin.
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:
Li Y, Lin Y (2019) Taxonomic review of the Asian Trogloneta species (Araneae, Mysmenidae). ZooKeys 817: 41-60. https://doi.org/10.3897/zookeys.817.30468
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Five Trogloneta species from Southwest China and Japan are reviewed that two new combinations and a new synonymy are proposed in the current paper: T. nojimai (Ono, 2010), comb. n. is transferred from Mysmena, T. yunnanense (Song & Zhu, 1994), comb. n. (= T. denticocleari Lin & Li, 2008, syn. n.) is transferred from Pholcomma of the Theridiidae, T. speciosum Lin & Li, 2008, T. uncata Lin & Li, 2013, and T. yuensis Lin & Li, 2013. The female of T. yuensis is described for the first time. An identification key and diagnoses are provided for these species, as well as new photographs or illustrations of the genital organs and habitus of T. yuensis and T. yunnanense.
China, identification key, Japan, mysmenids, new combination, new synonymy, taxonomy
The genus Trogloneta Simon, 1922 (= Troglonata, lapsus calami) was erected by
Currently, Trogloneta consists of eleven described species (
In this paper we provide a brief revision of Trogloneta species from Asia. The female of T. yuensis Lin & Li, 2013 is described for the first time. An identification key is also provided for these Asian Trogloneta species.
Specimens were collected by hand and preserved in 95% ethanol. They were examined using a Leica M205 C stereomicroscope. Further details were studied under an Olympus BX43 compound microscope. The epigynes were removed and treated with lactic acid before being photographed. Photographs were taken with a Canon EOS 60D wide zoom digital camera (8.5 megapixels) mounted on an Olympus BX 43 compound microscope. The images were montaged using Helicon Focus 3.10 (
All measurements are in millimetres. Leg measurements are given as follow: total length (femur, patella, tibia, metatarsus, and tarsus). Abbreviations in figures or text are as follows:
AA apical apophysis
Acc accessory gland
AL apical lobe
ALE anterior lateral eyes
AME anterior median eyes
At atrium
CD copulatory ducts
CO copulatory opening
Co conductor
Cy cymbium
CyC cymbial conductor
CyF cymbial fold
CyFs setae on cymbial fold
CyP cymbial process
E embolus
Et embolic tip
FD fertilisation ducts
Pa patella
PC paracymbium
PLE posterior lateral eyes
PME posterior median eyes
S spermathecae
SD spermatic duct
Sp scape
St subtegulum
T tegulum
TA tegular apophysis
Ti tibia
Abbreviations of specimen depository institutions:
Trogloneta granulum Simon, 1922.
Trogloneta differs from other mysmenid genera by the following combination of features: AME (or absent) smaller than ALE, all eyes gathered at apex of carapace; one femoral spot on leg I on both sexes (none on leg II); carapace height dimorphism (male carapace higher than female); anterior booklungs reduced; males with shorter, but stout and straight setae comprising the tarsal prolateral row on leg I; male palp huge (at least as big as half carapace), embolus tubular, its tip simple; cymbium with intricate decorations, and cymbial terminal acting as conductor; tegulum broad, usually having a apophysis; epigynal area elevated ventrally, with a scape, accessory glands on valve, and smooth uniform proximal copulatory ducts of increased diameter (
T. canariensis Wunderlich, 1987, T. cantareira Brescovit & Lopardo, 2008, T. cariacica Brescovit & Lopardo, 2008, T. granulum Simon, 1922, T. madeirensis Wunderlich, 1987, T. mourai Brescovit & Lopardo, 2008, T. nojimai (Ono, 2010), comb. n., T. paradoxa Gertsch, 1960, T. speciosum Lin & Li, 2008, T. uncata Lin & Li, 2013, T. yuensis Lin & Li, 2013, T. yunnanense (Song & Zhu, 1994), comb. n. (= T. denticocleari Lin & Li, 2008, syn. n.).
China (Chongqing, Guizhou, Hunan, Yunnan), Japan (Honshu), Europe (Austria, Czech Republic, France, Germany, Italy, Poland, Slovakia), USA (Utah, Oregon, California), Brazil (Minas Gerais, Rio de Janeiro, San Paulo, Santa Catarina, Espirito Santo, Parana), Canary Island, Madeira.
1 | Abdomen subglobose (Figs |
2 |
– | Abdomen pointed dorsally or posteriorly (Figs |
6 |
2 | Males | 3 |
– | Females | 5 |
3 | Embolus stubby, distal tip falcate (Fig. |
T. uncata |
– | Embolus long, distal tip spiculate (Figs |
4 |
4 | Cymbium strongly modified, with a huge cymbial process (Fig. |
T. yunnanense |
– | Cymbium moderately modified, cymbial process absent (Fig. |
T. nojimai |
5 |
Epigyne with a short scape, spermathecae globular (Fig. |
T. yunnanense |
– |
Epigyne with a long scape, spermathecae oviform (Fig. |
T. nojimai |
6 |
AME absent, embolus straight distally and epigynal scape short (Figs |
T. speciosum |
– |
AME present, embolus hooked distally and epigynal scape long (Figs |
T. yuensis |
Mysmena nojimai Ono, 2010: 2, figs 1–8.
Holotype: ♂ (
Trogloneta nojimai can be distinguished from other congeners but except of T. yuensis by the globular abdomen in both sexes (Figure
Trogloneta nojimai (Ono, 2010), comb. n., male holotype (A–F) and female paratype (G, H) (cited from Ono, 2010, slightly modified). A, B habitus (appendages omitted) C Prosoma D Endites, labium and sternum E, F Left palp G, H Epigyne A, F dorsal B, H lateral C anterior D, G ventral E retrolateral. Abbreviations: At atrium; CO copulatory opening; Cy cymbium; CyC cymbial conductor; CyFs setae on cymbial fold; E embolus; S spermathecae, SD spermatic duct; Sp scape; T tegulum, TA tegular apophysis; Ti tibia. Scale bars: 0.25 mm (A, B); 0.10 mm (C–H).
Although the type material of this species has not been examined for this study, the shape of palpal bulb, the configuration of epigyne, the patterns of eyes arrangement, and the distinctly elevated, conical carapace in male leave little doubt that it should be a member of the genus Trogloneta, but not Mysmena. The original illustrations of palp and epigyne of T. nojimai by (
Japan (Honshu).
Trogloneta speciosum Lin & Li, 2008: 514, figs 18A–E, 19A–I.
Holotype: ♂ (
This species differs from all species of Trogloneta by the absence of anterior median eyes and the posteriorly pointed abdomen in both sexes (Figure
Trogloneta speciosum Lin & Li, 2008, male (A–D) and female (E–G). A, B Left palp C, D Embolic division E, F Epigyne G Vulva. A prolateral B retrolateral C, G dorsal D, F ventral E lateral. Abbreviations: At atrium; CD copulatory ducts; CO copulatory opening; Co conductor; Cy cymbium; CyC cymbial conductor; CyF cymbial fold; CyFs setae on cymbial fold; CyP cymbial process; E embolus; Et embolic tip; S spermathecae, SD spermatic duct; Sp scape; T tegulum, TA tegular apophysis; Ti tibia. Scale bars: 0.20 mm (A, B, E–G); 0.10 mm (C, D).
This is the only species of Trogloneta spider ever found living in the tropical rainforest canopy, and is also a relatively rare six-eyed mysmenid species. We have tried to collect it again in the type locality, hoping to obtain some samples for molecular study but unfortunately, more material was not found.
China (Yunnan).
Trogloneta uncata Lin & Li, 2013b: 476, figs 23A–G, 24A–D, 25A–D.
Holotype ♂ (
Trogloneta uncata can be distinguished from Brazilian Trogloneta spp. (
Trogloneta uncata Lin & Li, 2013, male left pale (A–D). A prolateral B retrolateral C apical D ventral. Abbreviations: AA apical apophysis; AL apical lobe; Cy cymbium; CyC cymbial conductor; CyFs setae on cymbial fold; CyP cymbial process; E embolus; TA tegular apophysis; PC paracymbium; SD spermatic duct; St subtegulum; T tegulum; Ti tibia. Scale bars: 0.10 mm.
China (Yunnan).
Trogloneta yuensis Lin & Li, 2013a: 43, figs 8A–E, 9A–B, 10A–F, 11A–B, 12A–E.
Holotype ♂ (
5♂, 6♀ (
Trogloneta yuensis differs from most congeners except the Brazilian Trogloneta species and T. speciosum by the presence of pointed tubercle on abdomen dorso-posteriorly (Figure
Trogloneta yuensis Lin & Li, 2013, male left palp (A–D). A prolateral B retrolateral C apical D ventral. Abbreviations: Cy cymbium; CyC cymbial conductor; CyF cymbial fold; CyFs setae on cymbial fold; CyP cymbial process; E embolus; Et embolic tip; TA tegular apophysis; Pa patella; PC paracymbium; SD spermatic duct; T tegulum; Ti tibia. Scale bars: 0.20 mm.
Male. Measurements: total length 1.08. Prosoma 0.46 long, 0.45 wide, 0.60 high. Clypeus 0.32 high. Sternum 0.31 long, 0.30 wide. Opisthosoma 0.57 long, 0.55 wide, 0.98 high. Length of legs: I 1.45 (0.44, 0.17, 0.33, 0.29, 0.22); II 1.17 (0.39, 0.16, 0.23, 0.23, 0.16); III 1.00 (0.30, 0.13, 0.22, 0.18, 0.17); IV 1.18 (0.37, 0.15, 0.26, 0.22, 0.18). Legs formula: I-II-IV-III.
Somatic characters (Figure
Female. Measurements: Total length 1.32. Prosoma 0.52 long, 0.49 wide, 0.46 high. Clypeus 0.15 high. Sternum 0.34 long, 0.33 wide. Opisthosoma 0.92 long, 0.88 wide, 1.13 high. Length of legs: I 1.64 (0.53, 0.19, 0.36, 0.31, 0.25); II 1.39 (0.44, 0.18, 0.30, 0.25, 0.22); III 1.15 (0.35, 0.15, 0.25, 0.21, 0.19); IV 1.32 (0.42, 0.17, 0.28, 0.24, 0.21). Somatic characters (Figure
The species is original described on the basis of only a male specimen from Jinyun Mt. of Chongqing City that was donated by Prof Zhisheng Zhang (Southwest University in Chongqing, China) (
China (Chongqing, Hunan).
Pholcomma yunnanense Song & Zhu, 1994: 38, fig. 4A–C; Song, Zhu & Chen, 1999: 127, fig. 66A–B; Li & Lin, 2016: 320.
Trogloneta denticocleari Lin & Li, 2008: 513, figs 16A–E, 17A–E. Syn. n.
Holotype ♀ (of Pholcomma yunnanense) (
(Types of T. denticocleari): Holotype ♂, and paratypes 6♂, 29♀ (
2♂, 18♀ (
Trogloneta yunnanense can be distinguished from those Trogloneta species with a pointed abdominal tubercle (T. cantareira, T. cariacica, and T. mourai in
Trogloneta yunnanense (Song & Zhu, 1994) comb. n., male left palp (A–D). A prolateral B retrolateral C dorsal D ventral. Abbreviations: Cy cymbium; CyC cymbial conductor; CyF cymbial fold; CyFs setae on cymbial fold; CyP cymbial process; E embolus; Et embolic tip; TA tegular apophysis; Pa patella; PC paracymbium; SD spermatic duct; T tegulum; Ti tibia. Scale bars: 0.20 mm.
The type material of Pholcomma yunnanense has been examined as well as its related literatures in this study, the habitus features, the body size, the shape of protruded epigyne with an inflexed space, the broad epigynal plate, the configuration of vulva with a pair of far apart, globular spermathecae depicted in the type and original illustrations (
China (Guizhou, Yunnan).
After this study, the genus Trogloneta contains twelve nominal mysmenid species. Among its members, three species live in caves (T. yunnanense, T. granulum, and T. uncata), and the other nine are found in surface leaf litter (T. canariensis, T. cantareira, T. cariacica, T. madeirensis, T. mourai, T. nojimai, T. paradoxa, and T. yuensis) or in forest canopy (T. speciosum). Although the genus Trogloneta is widely distributed in Europe, Asia, North to South America, and in parts of the Atlantic islands, its many members are clearly endemic species according to the original literature. However, there are two exceptions, and they are T. granulum and T. yunnanense. The former as the type species of this genus was first found in caves of France, and later reported to be widespread on the surface Beech forest floor and in caves of many European countries, such as Austria, Czech Republic, France, Germany, Italy, Poland, and Slovakia. The latter as a new combination proposed in the current paper is also widely distributed in the isolated limestone caves in Southwest China. So far it has not yet been found on the surface. We know that the caves are relatively closed and isolated habitats: why they have such a distribution pattern, how do they get into caves to eventually colonise them, how does the isolation mechanism of the population work, and other questions are worth further study.
The manuscript benefitted greatly from comments by Lara Lopardo (Lufthansa, Germany) and Hirotsugu Ono (Tokyo, Japan). We are especially grateful to Jeremy A Miller (Leiden, Netherlands), the subject editor of this manuscript, for his editorial efforts. Thanks to Mr Guchun Zhou for donating part specimen materials of Trogloneta yuensis. This study was supported by the National Natural Science Foundation of China (NSFC-31772410, 31750002).