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Research Article
A taxonomic study on Epedanidae from Thailand including functional aspects of male genital morphology (Opiliones, Laniatores)
expand article infoChao Zhang, Jochen Martens§|
‡ Hebei University, Baoding, China
§ Institut für Organismische und Molekulare Evolutionsbiologie, Mainz, Germany
| Senckenberg Research Institute, Frankfurt am Main, Germany
Open Access

Abstract

The South-East Asian opilionid family Epedanidae Sørensen, 1886 has one of its strongholds in Thailand from where a multitude of genera and species have been described but the epedanid fauna of the country is still poorly known. This paper records four species from this country, three of which are new: Euepedanus dashdamirovi sp. nov. (male and female), Plistobunus jaegeri sp. nov. (male and female), and Toccolus kuryi sp. nov. (male and female). Toccolus globitarsis Suzuki, 1969 was previously known only from the type locality in Thailand and is redescribed here. Functional aspects of epedanid penial morphology are highlighted.

Keywords

Arachnida, harvestmen, male genitalia, taxonomy

Introduction

The opilionid family Epedanidae Sørensen, 1886 is widespread in the Indo-Malay Realm where it contains many genera, but the local species are often narrowly distributed and still poorly known. Thailand is one of the strongholds of the family. Many epedanid species display a bizarre peripheral morphology with heavily armed chelicerae, pedipalps, and a surprisingly long pointed spine or apophysis on top of the ocularium. In many cases, genera within the Epedanidae are not well characterized and probably do not represent monophyletic entities. This paper aims at increasing knowledge of the Thai fauna in re-defining three known genera and one known species, and in proposing three new species.

The two Epedanidae species first described from Thailand are Euepedanus orientalis (Hirst, 1912) and Caletorellus siamensis (Hirst, 1912). Originally, these were placed in genus Epedanus Thorell, 1876. Subsequently, E. orientalis was transferred to Mimepedanus Roewer, 1923 by Roewer (1923) and this genus was synonymized with Euepedanus Roewer, 1915 by Suzuki (1969b). The second species, E. siamensis was transferred to Euepedanus by Roewer (1923), and later to Caletorellus Roewer, 1938 by Roewer (1938) and is the type species of the latter genus.

Roewer (1938) systematically reorganized what is today known as Epedanidae (for him they were all included in Phalangodidae Simon, 1879) and he erected three subfamilies in it: Acrobuninae Roewer, 1912, Epedaninae Roewer, 1912 (incorrectly stated, author of subfamily is Sørensen 1886), and Sarasinicinae Roewer, 1912. Suzuki subsequently published three new Thai epedanid genera, and ten new species (Suzuki 1969a, 1969b, 1981, 1985; Suzuki and Stone 1986). All these descriptions are very accurately carried out including meticulous drawings of relevant characters of male genital traits.

The Naturmuseum Senckenberg, Frankfurt am Main, Germany houses most of Roewer's Epedanidae types which were described by him over several decades (Roewer 1912, 1915, 1927,1938), and this institution is continuously enlarging its collections. In this paper, we study a small epedanid collection from Thailand, which was gathered in 2010 by Selvin Dashdamirov. In addition, we pay attention not only to epedanid male genital morphology for species recognition but in addition try to disentangle functional morphology of the movable parts of the copulatory organs.

Materials and methods

Taxonomic methods follow the outline proposed by Acosta et al. (2007). The specimens were preserved in 70% ethanol, and examined and drawn under a Leica M205A stereomicroscope equipped with a drawing tube. Further details were studied using a compound microscope (Nikon YS100). Photographs were taken using a Leica M205A stereomicroscope equipped with a DFC450 CCD. The male genitalia were placed first in hot lactic acid, later transferred to distilled water to expand the movable parts of glans penis for observation (Schwendinger and Martens 2002). The terminology of genital structures follows Martens (1986) and Macías-Ordóñez et al. (2010), the macrosetae terminology follows Kury and Villarreal (2015). The type specimens are deposited in the Senckenberg Arachnology collections (SMF).

All measurements are given in mm. The following abbreviations are used in the text:

CI capsula interna;

CE capsula externa;

Pb pars basalis;

Pd pars distalis;

S stylus;

VP ventral plate.

Taxonomy

Euepedanus Roewer, 1915

Euepedanus Roewer 1915: 62; 1923: 206 (in part); 1938: 114. Suzuki 1969b: 83; 1985: 81. Zhu and Lian 2006: 64.

Mimepedanus Roewer 1923: 208; 1938: 105. Synonymized by Suzuki 1969b: 83.

Type species

Euepedanus trispinosus Roewer, 1915, subsequent designation by Suzuki (1969b: 83).

Diagnosis

Medium-sized epedanines (3.00–5.00 mm) with a long median spine on the ocularium. Area II with a pair of long spines. The basichelicerite of chelicerae with numerous tubercles. Pedipalpal femur ventrally with 5–7 seta-tipped tubercles, and distally with two tubercles on medial side. Pedipalpal patella with at least three seta-tipped tubercles. Distitarsus of leg I with two segments (cf. Zhu and Lian 2006: 64).

Included species

(nine species). Euepedanus trispinosus Roewer, 1915, Euepedanus orientalis (Hirst, 1912), Euepedanus chaiensis Suzuki, 1969b, Euepedanus dividuus Suzuki, 1969b, Euepedanus similis Suzuki, 1985, Euepedanus pentaspinulatus Suzuki, 1985, Euepedanus spinosus Suzuki, 1985, Euepedanus flavimaculatus Zhu and Lian, 2006, and Euepedanus dashdamirovi sp. nov.

Distribution

Thailand, China, Malaysia, Singapore.

Key to the currently known species of Euepedanus

1 Anterior margin of prosoma unarmed 2
Anterior margin of prosoma armed with tubercles 4
2 Basichelicerite of chelicerae laterally and medially with prominent spines E. trispinosus (Singapore and Malaysia)
Basichelicerite of chelicerae laterally and medially without prominent spines, with small tubercles or teeth 3
3 Pedipalpal femur ventrally with five seta-tipped tubercles E. similis (Thailand)
Pedipalpal femur ventrally with seven seta-tipped tubercles E. flavimaculatus (China)
4 Femora of all legs with ventral row of seta-tipped granules E. spinosus (Thailand)
Femora of all legs without a ventral row of seta-tipped granules 5
5 Scutal area I divided into three parts E. dividuus (Thailand)
Scutal area I entire .6
6 Posterolateral angle of scutum with one large tubercle E. orientalis (Thailand)
Posterolateral angle of scutum without one large tubercle 7
7 Conspicuous spines dorsally on the basichelicerite of chelicerae dispersed evenly E. dashdamirovi sp. nov. (Thailand)
Conspicuous spines dorsally on the basichelicerite of chelicerae dispersed unevenly, gathered near the base 8
8 Median tubercles of free tergites I and II enlarged, cheliceral hand with a ventral seta-tipped tubercle E. pentaspinulatus (Thailand)
Median tubercles of free tergites I and II small, cheliceral hand without a ventral seta-tipped tubercle E. chaiensis (Thailand)

Euepedanus dashdamirovi sp. nov.

Figures 1–8, 9–17, 18–25, 26–31

Type material

Holotype male (SMF-CJM7059-01): THAILAND: Sa Kaeo (Kaew) Province: Ta Phraya District, Ta Phraya NP (No.11), 14°8.37'N 102°40.19'E, alt. 164 m, 27 October 2010, S. Dashdamirov leg. Paratypes. one female (SMF-CJM7059-02), same collecting data as holotype; one female (SMF-CJM7051-01) and one juvenile (SMF-CJM7051-02), THAILAND: Sa Kaeo (Kaew) Province: Maeng, Srakaeo District, Wat Tham Khao Maka (No.8), 13°47.23'N 101°56.87'E, alt. 46m, 26 October 2010, S. Dashdamirov leg.

Diagnosis

Penis with a stereoscopic ventral plate (e.g., Fig. 132) and the stylus constricted apically. Anterior margin of prosoma armed with tubercles. The basichelicerite of chelicerae dorsally with five tubercles (Figs 2–4). Pedipalpal femur ventrally with six seta-tipped tubercles, and distally with two tubercles on medial side. Pedipalpal patella with three seta-tipped tubercles.

Etymology

Named in honor of the collector of the type material and our good friend, Dr. Selvin Dashdamirov.

Description

Male (holotype) habitus as in Figs 1, 9, 26, 27. Coloration (Figs 26, 27): entire body dorsally rusty yellow with black patches; median area of prosoma with black reticulations before and behind the ocularium; both lateral ridges of scuta with brown stripes; opisthosomal area I bordered posteriorly with brown and area II dark brown with paler interspaces; areae III and IV and free tergites each with a transverse dark band; venter concolorous with the dorsum; chelicerae and pedipalps reticulated; all leg segments other than trochanters and tarsi with black reticulations.

Figures 1–8. 

Euepedanus dashdamirovi sp. nov., male holotype 1 body, lateral view 2 right chelicera, lateral view 3 same, medial view 4 right basichelicerite, dorsal view 5 right cheliceral hand, frontal view 6 Right cheliceral fingers, frontal view 7 right pedipalp, lateral view 8 same, medial view. Scale bars: 1 mm (1–5, 7–8), 0.5 mm (6).

Dorsum (Figs 9, 26). Dorsal scutum more or less trapezoid in form, with sides slightly curved, proximal end of abdomen slightly enlarged and rounded. Anterior margin of prosoma with eight small tubercles at the lateral portion. Ocularium long oval, removed from anterior border of scutum by 0.50 mm, armed with an elongate, lightly curved spine. Posterior margin of prosoma bowed. Opisthosomal area I longer than other areae. Areae I and III each with a median pair of short spines. Area II with two median long spines. Area IV and all free tergites with a transverse row of seta-tipped tubercles, central ones larger than others, like slender spines. Lateral margins of the scutum with a longitudinal row of seta-tipped tubercles.

Figures 9–17. 

Euepedanus dashdamirovi sp. nov., male (Fig. 9, holotype), female (Figs 10–17, paratype) 9 male body, dorsal view 10 female body, dorsal view 11 right chelicera, medial view 12 same, lateral view 13 left basichelicerite, dorsal view 14 left cheliceral hand, frontal view 15 left pedipalp, medial view 16 ovipositor, ventral view 17 same, dorsal view. Scale bars: 1 mm (9, 10, 15), 0.5 mm (11–14, 16, 17).

Venter (Fig. 28). Coxae with tubercular surface, coxa I with enlarged seta-tipped tubercles. Coxae II, III, and IV with only minute similar tubercles. Genital operculum with a few seta-tipped granules. Free sternites with a row of minute tubercles, each with setae on top. Spiracles clearly visible.

Chelicerae (Figs 2–6). Basichelicerite elongate, dorsally with a slight bulla, and armed with four long spines and one seta-tipped tubercle dorsally; two similar spines and one seta-tipped tubercle medially on the ectal side; one seta-tipped tubercle on the lateral side; ten seta-tipped tubercles scattered over the ventral side; medial side with a basal protuberance. Cheliceral hand widened, pro-dorsally with some seta-tipped tubercles; two seta-tipped tubercles on the ventral side. Fingers relatively strong, inner edges toothed (Fig. 6): moveable finger and fixed finger each with five stump teeth.

Pedipalps (Figs 7, 8). Coxa dorsally with two seta-tipped tubercles; ventrally with one distal seta-tipped tubercle. Trochanter ventrally and dorsally each with one enlarged and one small seta-tipped tubercles. Femur ventrally with a row of six seta-tipped tubercles, of which the two longest at the base; dorsally with two longitudinal rows of conical tubercles; distally on medial side with two seta-tipped tubercles. Patella ventro-mesally with two setiferous tubercles, and ventro-ectally with one setiferous tubercles. Tibia ventro-mesally with three setiferous tubercles, and ventro-ectally with four setiferous tubercles. Tarsus with three setiferous tubercles on each side of ventral side. Tarsal claw curved, longer than tarsus.

Legs. All segments smooth, only with scarce short setae. Femur IV slightly curved. Distitarsus I with two, distitarsus II with three tarsomeres. Distitarsi III and IV without scopula. Tarsal claws smooth. Tarsal formula (I–IV): 9(2)/24(3)/7/8.

Penis (Figs 18–25). Shaft slender, elongate. The distal fourth of the shaft broadened and nearly parallel-sided till apical portion (pars distalis). Ventral plate nearly square (ventral view, Fig. 19), and slightly indented on the distal margin (Fig. 19). Glans partially sunken into dorsal depressed portion of pars distalis and not extending the distal margin of the ventral plate. Capsula externa and capsula interna cylindrical, and the inner side of capsula interna with dense cover of fur-like microtrichia. The ventral lobe of capsula interna conspicuous, somewhat triangular; the dorsal lobe inconspicuous. Stylus with irregular shape, constricted apically. Spination symmetrical. One pair of setae A, B, and C. Two pairs of setae D, E, and F (Figs 19–21).

Figures 18–25. 

Euepedanus dashdamirovi sp. nov., genitalia of male holotype 18 penis, lateral view 19 pars distalis of penis, ventral view (arrow indicates indented distal end) 20 same, lateral view 21 same, dorsal view 22 pars distalis of penis (everted), ventral view 23 same, dorsal view 24–25 same, lateral view. Abbreviations: CE capsula externa CI capsula interna DL dorsal lobe G glans Pb pars basalis Pd pars distalis S stylus VP ventral plate VL ventral lobe. Scale bars: 0.5 mm (18), 0.25 mm (19–25).

Female (Figs 10–17, 29–31). In general appearance similar to the male, abdomen more rounded posteriorly (Figs 10, 29). Basichelicerite dorsally with three and ectally with one long spines, other seta-tipped tubercles on chelicerae less stout than those of the male (Figs 11–13). Cheliceral hand prodorsally with two enlarged seta-tipped tubercles in the middle of inner portion (Fig. 11); one seta-tipped tubercle on the ventral side (Fig. 12). Pedipalp similar to that of male (Fig. 15). Tarsal formula (I–IV): 8(2)/19(3)/7/8.

Ovipositor (Figs 16, 17). Elongate. Ventral side with four and dorsal side with six setae of different lengths.

Measurements

Male holotype (female paratype): body 2.94 (2.36) long, 2.13 (2.48) wide at widest portion, scutum 2.36 (2.05) long. Ocularium 0.40 (0.30) long, 0.79 (0.69) wide. Proximal chelicerae 1.24 (1.08) long, 0.43 (0.38) wide; second 2.10 (1.72) long, 0.72 (0.59) wide; distal 0.91 (0.80) long, 0.21 (0.20) wide. Pedipalp claw 0.84 (0.80) long. Penis 1.67 long. Measurements of pedipalp and legs as in Tables 1 and 2.

Table 1.

Euepedanus dashdamirovi sp. nov. Measurements of the pedipalp and legs of the male holotype, as length/width.

Trochanter Femur Patella Tibia Metatarsus Tarsus Total
Pedipalp 0.59/0.39 1.65/0.31 0.85/0.30 0.92/0.35 0.57/0.33 4.58
Leg I 0.30/0.27 2.10/0.19 0.51/0.26 1.49/0.14 2.60/0.05 1.30/0.04 8.30
Leg II 0.32/0.33 3.54/0.18 0.64/0.27 3.02/0.15 4.24/0.05 3.22/0.04 14.98
Leg III 0.52/0.44 2.65/0.26 0.65/0.35 1.59/0.23 3.30/0.13 1.30/0.07 10.01
Leg IV 0.45/0.40 3.88/0.28 0.72/0.38 2.15/0.23 4.74/0.09 1.45/0.07 13.39
Table 2.

Euepedanus dashdamirovi sp. nov. Measurements of the pedipalp and legs of the female paratype, as length/width.

Trochanter Femur Patella Tibia Metatarsus Tarsus Total
Pedipalp 0.46/0.32 1.32/0.30 0.72/0.29 0.76/0.34 0.45/0.31 3.71
Leg I 0.33/0.26 1.83/0.17 0.52/0.23 1.26/0.16 2.31/0.03 0.84/0.03 7.09
Leg II 0.34/0.29 2.77/0.16 0.68/0.25 2.82/0.15 3.37/0.08 2.44/0.06 12.42
Leg III 0.34/0.41 2.33/0.26 0.62/0.38 1.49/0.23 2.87/0.06 1.02/0.06 8.67
Leg IV 0.37/0.37 3.42/0.24 0.69/0.33 1.97/0.21 4.28/0.11 1.43/0.05 12.16

Habitat

The specimens were collected by hand under the decaying bark of various tree species.

Distribution

Known only from the type locality.

Figures 26–31. 

Photographs of male (Figs 26–28, holotype) and female (Figs 29–31, paratype) of Euepedanus dashdamirovi sp. nov. 26, 29 body and parts of appendages, dorsal view 27, 30 body and parts of appendages, lateral view 28, 31 body and parts of appendages, ventral view. Scale bars: 1 mm.

Plistobunus Pocock, 1903

Plistobunus Pocock 1903: 447; Roewer 1912: 232; 1923: 207; 1938: 124; Lian et al. 2011: 41.

Type species

Plistobunus rapax Pocock, 1903, by original designation.

Extended diagnosis

Medium-sized epedanines (2.40–3.57 mm) with a long median spine on the ocularium. Prosoma with a row of 4–6 setiferous tubercles on each side of the frontal margin. Areae II and IV with or without long spines. The basichelicerite of chelicerae elongate and armed with numerous tubercles, dorsally with three in males and with two in female (Lian et al. 2011: 42, figs 2, 3; 44: 9, 10; 46: 18, 19. Figs 33–35, 47–49). Pedipalpal femur of male ventrally with 9–13 seta-tipped tubercles, and distally with one or two tubercles on medial side. Pedipalpal patella with at least three seta-tipped tubercles. Distitarsus of leg I with two segments. The distal part of pars basalis of penis widened and with a stereoscopic ventral plate. Stylus columnar. Capsula interna ventrally and capsula externa dorsally surround the stylus.

Included species

Plistobunus rapax Pocock, 1903, Plistobunus columnarius Lian et al., 2011, and Plistobunus jaegeri sp. nov.

Distribution

China (Hong Kong and Hainan Island), Thailand (Ubon Ratchathani).

Sexual dimorphism

The species of this genus possess conspicuous sexual dimorphism in the armature of the appendages. The tubercles or spines in male on the basichelicerite of chelicerae, and those of the femur of pedipalps and legs are much more robust or conspicuous than those of in female; the shape of body is very similar in both sexes.

The numbers of seta-tipped tubercles distally on medial side of pedipalpal femur gives an impression of the differences between the species. Male specimens of P. rapax and P. columnarius are described with two (Lian et al. 2011: 42, fig. 6; 44, fig. 14) and those of P. jaegeri sp. nov. with only one such tubercle (Fig. 38). Females of P. columnarius (Lian et al. 2011: 46, fig. 22) and P. jaegeri sp. nov. (Fig. 52) possess no tubercles of this size or position (female of P. rapax unknown).

Key to the currently known species of Plistobunus

1 Distributed in Thailand, the scutal area without long spines in male P. jaegeri sp. nov.
Distributed in China, scutal area II with a pair of long spines and area IV with a median one in male 2
2 Distributed in Hong Kong, the anterior margin of prosoma with a row of nine tubercles, and the femur of pedipalp ventrally with nine seta-tipped tubercles. P. rapax
Distributed in Hainan Island, the anterior margin of prosoma with a row of 12 tubercles, and the femur of pedipalp ventrally with 13 seta-tipped tubercles P. columnarius

Plistobunus jaegeri sp. nov.

Figures 32–39, 40–54, 55–61, 62–67

Type material

Holotype male (SMF-CJM7060): THAILAND: Ubon Ratchathani Province: Khong Chiam District, Pha Taem NP (No.28), 15°23.94'N 105°30.49'E, alt. 185 m, 5 November 2010, S. Dashdamirov leg. Paratype. One female (SMF-CJM7062): THAILAND: Nong Khai Province: Sangkhom District, Tham Tip Waterfall (No. 23), 18°7.57'N 102°11.16'E, alt. 198 m, 3 November 2010, S. Dashdamirov leg.

Diagnosis

The dorso-distal margin (blue in Fig. 57a) of ventral plate much higher than the dorso-distal margin (purple in Fig. 57a) of pars basalis, the ventral and dorsal side of distal margin of pars basalis of penis not in the same level (cf. Fig. 57b). Dorsal opisthosomal areae II and IV without long spines in male (Figs 32, 63). The pedipalpal femur medio-distally with one seta-tipped tubercle, and the male patella with one short tubercle at base instead of unarmed in female (Figs 38, 52).

Etymology

The new species is dedicated to Dr. Peter Jäger, Germany, an esteemed arachnologist. He is a co-founder of the Asian Society of Arachnology (ASA) and due to his efforts, arachnology in Asia has taken a large step forward.

Description

Male (holotype) habitus as in Figs 32, 40, 62, 63. Coloration (Figs 62, 63): entire body dorsally rusty yellow with brown patches; median area of prosoma with brown reticulations in front of the ocularium; on each side behind ocularium brown outlines; both lateral ridges of the prosomal and opisthosomal scuta with dark brown stripes; opisthosomal areae I–III with light brown outlines; areae IV and V with a dark transverse band; free tergites with three large dark brown blotches; venter concolorous with the dorsum; chelicerae, pedipalps and legs rusty yellow, reticulated with light to dark brown.

Figures 32–39. 

Plistobunus jaegeri sp. nov., male holotype 32 body, lateral view 33 left chelicera, medial view 34 same, lateral view 35 left basichelicerite, dorsal view 36 left cheliceral hand, frontal view 37 left cheliceral fingers, frontal view 38 left pedipalp, medial view 39 same, lateral view. Scale bars: 1 mm (32–36, 38, 39), 0.5 mm (37).

Dorsum (Figs 40, 62). Dorsal scutum trapezoid in shape, widest portion of body at area IV, the abdomen broadly rounded posteriorly. Anterior margin of prosoma with six tubercles. Ocularium oval, removed from anterior border of scutum by 0.40 mm, armed with a long median upright spine. The anterior margin of opisthosomal area I strongly bowed and the borders of other areae slightly bowed. Opisthosomal area I with a median pair of seta-tipped tubercles; areae II and III with similar median tubercles in addition to a few smaller ones on each side; areae IV and V and free tergites with a transverse row of seta-tipped tubercles.

Venter (Fig. 64). Coxae with tubercular surface. Coxa I with enlarged seta-tipped tubercles. Coxae II–IV with minute seta-tipped tubercles. Genital operculum with a few seta-tipped granules. Free sternites with a row of small seta-tipped tubercles. Spiracles clearly visible.

Chelicerae (Figs 33–37). Basichelicerite elongate, its dorsal side centrally with three spines, distal one the longest and proximal one the shortest; the lateral side with a row of four medium-sized spines; on the medial side four seta-tipped tubercles spirally arranged, in addition with a basal protuberance; ventral side with eight seta-tipped tubercles. Cheliceral hand considerably broad, pro-dorsally with some seta-tipped tubercles; three seta-tipped tubercles on the ventral side. Fingers strong, inner edges toothed (Fig. 37): moveable finger and fixed finger with five stumpy teeth.

Pedipalps (Figs 38, 39). Coxa dorsally and ventrally each with three seta-tipped tubercles. Trochanter ventrally with three, dorsally with one seta-tipped tubercles. Femur ventrally with a row of nine; dorsally with a row of nine seta-tipped tubercles; medio-distally with one seta-tipped tubercle. Patella ventro-mesally and ventro-ectally each with two seta-tipped tubercles. Tibia ventro-mesally with three and ventro-ectally with four and ventrally with one seta-tipped basal tubercle. Tarsus with four seta-tipped tubercles on ventral side. Tarsal claw curved, longer than tarsus.

Legs (Figs 41–44). Trochanters I–IV ventrally with a few small seta-tipped granules. Femora I–IV ventrally with longitudinal row of many seta-tipped tubercles; size on femur I enlarged and on femora II–IV reduced, on femur IV inconspicuous. Patella I ventrally with two seta-tipped granules. Tarsi III and IV with smooth double claws, without scapulae. Distitarsi I and II two-jointed. The remaining leg segments unarmed. Tarsal formula (I–IV): 6(2)/15(2)/6/7.

Figures 40–54. 

Plistobunus jaegeri sp. nov. male (Figs 40–44, holotype) and female (Figs 45–54, paratype) 40, 45 body, dorsal view 41 right leg I, retrolateral view 42–44, 46 femur of right legs, retrolateral view 42 leg II 43 leg III 44 leg IV 46 leg I 47 left chelicera, medial view 48 same, lateral view 49 left basichelicerite, dorsal view 50 left cheliceral hand, frontal view 51 left cheliceral fingers, frontal view 52 left pedipalp, medial view 53 ovipositor, ventral view 54 same, dorsal view. Scale bars: 1 mm (40–46, 52), 0.5 mm (47–51), 0.25 mm (53, 54).

Penis (Figs 55–61). The shaft slender, parallel-sided, enlarged towards glans, distal end broadest. Distal part of truncus, i.e., glans area, almost trapezoid (ventral view, Figs 55, 56, 59), and dorsally without an indentation on the distal margin (Fig. 56, cf. Fig. 19). Glans partially sunken into dorsally depressed portion of pars distalis of penis, its tip slightly extending the distal margin (blue in Fig. 57a) of the ventral plate. The distal extension of capsula externa of semi-circular shape (Fig. 60). The distal extension of capsula interna fused into a V-shaped structure (ventral view, Fig. 59). Stylus columnar, flat distally (lateral view, Fig. 60). Spination asymmetrical, in position a single seta D; spination symmetrical, one pair of setae B, two pairs of setae C, E, and F, and three pairs of setae A.

Female (Figs 45–54, 65–67). In general appearance similar to the male. Scutum more widely trapezoid and wider than that of male in the posterior margin. Opisthosomal area II with a median pair of spines, area IV with a median spine (Fig. 66). Chelicera smaller and with reduced tubercles, the dorsal surface of basichelicerite centrally with one spine, the ventral surface of cheliceral hand with one seta-tipped tubercle, inner edges of finger toothed as illustrated (Fig. 51). Pedipalpal (Fig. 52) femur ventrally with a row of six, dorsally with three conspicuous seta-tipped tubercles, medio-distally without any seta-tipped tubercle; patella with three seta-tipped tubercles. The seta-tipped tubercles on femur I (Fig. 46), as well as femora II–IV inconspicuous. Tarsal formula (I–IV): 7(2)/18(2)/7/8.

Figures 55–61. 

Plistobunus jaegeri sp. nov., genitalia of male holotype (not included Fig. 57b) 55 penis, ventral view 56 pars distalis of penis, ventral view 57a same, lateral view 58 same, dorsal view 57b pars distalis of penis, lateral view (Plistobunus columnarius Lian et al., 2011) 59 pars distalis of penis (everted), ventral view 60 same, lateral view 61 same, dorsal view. Abbreviations: CE capsula externa CI capsula interna G glans Pb pars basalis Pd pars distalis S stylus VP ventral plate. Scale bars: 0.25 mm.

Ovipositor (Figs 53, 54). On ventral side with four and on dorsal side with six setae.

Measurements

Male holotype (female paratype). Body 2.40 (3.53) long, 1.94 (2.08) wide at the widest portion, scutum1.68 (2.23) long. Ocularium 0.33 (0.32) long, 0.61 (0.56) wide. Proximal chelicerae 1.24 (1.11) long, 0.35 (0.30) wide; second 1.80 (1.50) long, 0.60 (0.42) wide; distal 0.76 (0.76) long, 0.19 (0.16) wide. Pedipalpus claw 0.60 (0.79) long. Penis 0.94 long. Measurements of pedipalpus and legs as in Tables 3 and 4.

Table 3.

Plistobunus jaegeri sp. nov. Measurements of the pedipalp and legs of the male holotype, as length/width.

Trochanter Femur Patella Tibia Metatarsus Tarsus Total
Pedipalp 0.43/0.25 1.44/0.20 0.68/0.23 0.70/0.27 0.51/0.31 3.76
Leg I 0..27/0.23 1.33/0.16 0.48/0.23 0.91/0.16 1.53/0.08 0.87/0.06 5.39
Leg II 0.30/0.26 1.80/0.15 0.56/0.22 1.43/0.15 2.03/0.06 1.71/0.05 7.83
Leg III 0.36/0.33 1.33/0.23 0.44/0.29 1.00/0.20 1.72/0.11 0.99/0.07 5.84
Leg IV 0.35/0.31 1.95/0.20 0.61/0.23 1.34/0.21 2.28/0.12 1.18/0.09 6.71
Table 4.

Plistobunus jaegeri sp. nov. Measurements of the pedipalp and legs of the female paratype, as length/width.

Trochanter Femur Patella Tibia Metatarsus Tarsus Total
Pedipalp 0.46/0.26 2.19/0.17 1.00/0.25 0.91/0.26 0.82/0.24 5.38
Leg I 0.27/0.21 1.46/0.15 0.51/0.22 1.29/0.11 2.28/0.06 1.03/0.04 5.84
Leg II 0.33/0.23 2.64/0.13 0.55/0.22 2.42/0.13 3.20/0.06 2.75/0.06 11.89
Leg III 0.34/0.36 2.08/0.18 0.56/0.31 1.44/0.19 2.91/0.10 1.14/0.06 8.47
Leg IV 0.41/0.33 3.24/0.21 0.63/0.30 1.86/0.20 4.10/0.12 1.49/0.08 11.73

Habitat

The specimens were collected by hand under stones in forest.

Distribution

Known only from the type locality, the Pha Taem National Park in the Ubon Ratchathanl Province, Thailand.

Notes. The genus Plistobunus was known by two species restricted to China, i.e., P. rapax Pocock, 1903 (type species, Hong Kong) and P. columnarius Lian et al., 2011 (Hainan Island). The most distinctive characters of these two species are a pair of long dorsal spines on opisthosomal area II and a median long spine on area IV. However, these characters are not conspicuous in the male of P. jaegeri sp. nov. (Figs 32, 63).

Figures 62–67. 

Photographs of male (Figs 62–64, holotype) and female (Figs 65–67, paratype) of Plistobunus jaegeri sp. nov. 62, 65 body and parts of appendages, dorsal view 63, 66 body and parts of appendages, lateral view 64, 67 body and parts of appendages, ventral view. Scale bars: 1 mm.

Toccolus Roewer, 1927

Toccolus Roewer 1927: 203; 1938: 102; Kury 2008: 62.

Type species

Toccolus minimus Roewer, 1927, by monotypy.

Extended diagnosis

Medium-sized epedanines (1.40–3.21 mm) with a median spine on the ocularium. Opisthosomal areae unarmed except for some seta-tipped tubercles. The basichelicerite of chelicerae armed with small or enlarged tubercles. Pedipalpal femur ventrally with five or six seta-tipped tubercles, and distally with two or three tubercles on medial side. Pedipalpal patella ventro-mesally with two tubercles, and ventro-ectally with one tubercle. Distitarsus of leg I with two segments.

Included species

Toccolus minimus Roewer, 1927, Toccolus chibai Suzuki, 1976, Toccolus globitarsis Suzuki, 1969, Toccolus javanensis Kury, 2008, and Toccolus kuryi sp. nov.

Distribution

Thailand, Java, Malaysia, Vietnam.

Key to the currently known species of Toccolus

1 Basichelicerite with small tubercles (the height almost as long as its width) T. minimus (Vietnam)
Basichelicerite with enlarged tubercles (the height much longer than its width) 2
2 Ocularium with a long median spine T. chibai (Malaysia)
Ocularium with a short median spine 3
3 Basichelicerite dorsally with four conspicuous spines T. javanensis (Java)
Basichelicerite dorsally with three conspicuous spines 4
4 Pedipalpal femur ventrally with five seta-tipped tubercl T. globitarsis (Thailand)
Pedipalpal femur ventrally with six seta-tipped tubercles T. kuryi sp. nov. (Thailand)

Toccolus globitarsis Suzuki, 1969

Figures 68–72, 73–86, 87–95, 96–101

Toccolus globitarsis Suzuki 1969b: 91–96, fig. 9–11, pls 5–6.

Type specimens

(not examined). Holotype, male: THAILAND: Paktong Cha, Nakhon Ratchasima Province: 24 November – 10 December 1963. Paratypes: three females, same collection data as the holotype. All specimens were collected by H. Watanabe.

The type-specimens are all deposited in the Zoological Laboratory, Faculty of Science, Hiroshima University.

Additional material examined

One male, one female and one juvenile (SMF-CJM7053), THAILAND: Chaiyaphum Province: Phakdi Chumphon Distr., near cave Wat Tham Kaeo (No. 15), 15°58.50'N 101°24.66'E, 350 m alt. 1 November 2010, S. Dashdamirov leg.

Redescription

Male habitus as in Figs 68, 73, 96, 97. Coloration (Figs 96, 97): body light yellow with brown patches; median area of prosoma with brown reticulations; both lateral ridges of scutum with brown stripes; opisthosomal areae I–III each with a transverse band of brown markings; area IV and all free tergites with a cross series of three brown flecks; all coxae yellow with brown reticulations distally; free sternites each with brown bands, bands somewhat paler on the central portion; chelicerae and pedipalps with reticulated irregular markings; trochanters pale yellow, femur, patella, tibia, and metatarsus with black reticulations, tarsus lighter.

Dorsum (Figs 73, 96). Scutum nearly parallel-sided, slightly trapezoid, the widest portion at scutal area IV, and abdomen bluntly pointed posteriorly. Prosoma with a row of nine sharp pointed tubercles along anterior margin (Fig. 73); otherwise whole dorsum smooth. Ocularium oval, removed from anterior border of scutum by 0.33 mm, armed with a conspicuously shorter median spine than the height of ocularium (lateral view), and two small tubercles above the eyes. The posterior area of ocularium on prosoma raised and broadly rounded. The anterior margin of opisthosomal area I bowed and the borders of other areae slightly bowed. Opisthosomal areae I–IV with a row of seta-tipped tubercles and a longitudinal row of similar tubercles on the lateral margins. Free tergites with a transverse row of seta-tipped tubercles each, anal operculum with scattered granules.

Venter (Fig. 98). Coxae with tubercular surface, coxa I with enlarged seta-tipped tubercles. Coxae II–IV with minute tubercles. Genital operculum and free sternites with seta-tipped granules. Spiracles clearly visible.

Chelicerae (Figs 69–70, 74–77). Basichelicerite elongate and armed with three long, curved spines dorsally (Figs 69, 70); one similar spine distally on the ectal side (Figs 70, 74); six seta-tipped tubercles scattered over the ventral side (Fig. 69); medial side with a basal protuberance (Fig. 69). Tubercle on left basichelicerite more medially than on right one (arrowed in Fig. 75). Cheliceral hand considerably widened, pro-dorsally with some seta-tipped tubercles, the inner ones much larger than the outer ones (Fig. 76); ectally with two seta-tipped tubercles at the subdistal portion (Figs 70, 76); one enlarged seta-tipped tubercle and a smaller one on the ventral side (Fig. 70). Fingers strong, inner edges toothed (Fig. 77): moveable finger with five teeth, the proximal one square, the other four pointed; fixed finger with six crested teeth.

Figures 68–72. 

Toccolus globitarsis Suzuki, 1969. Male 68 body, lateral view 69 left chelicera, medial view 70 same, lateral view 71 left pedipalp, medial view 72 same, lateral view. Scale bars: 1 mm.

Pedipalps (Figs 71, 72). Coxa dorsally with three seta-tipped tubercles, one strong distal, one subdistal more exteriorly placed, one blunt proximal; ventrally with two seta-tipped tubercles. Trochanter ventrally with two setiferous tubercles, dorsally with an enlarged one and a small one. Femur ventrally with a row of five setiferous tubercles of which the longest one close to the base; dorsally with a row of ten setiferous tubercles of which the distal two inconspicuous; distally on medial side with two setiferous tubercles. Patella ventro-mesally with two setiferous and ventro-ectally with one setiferous tubercles. Tibia ventro-mesally with three and ventro-ectally with four setiferous tubercles. Tarsus on ventral side para-median with three setiferous tubercles each. Tarsal claw curved, longer than tarsus.

Legs. All segments unarmed, nearly smooth. Legs I–II slender, legs III–IV much stronger. Femora I–III not curved, almost straight, femur IV slightly curved. Distitarsi I and II with two tarsomeres. Distitarsi III and IV without scopula, only two bare claws present. Tarsal formula (I–IV): 7(2)/16(2)/6/7.

Penis (Figs 87–95). Shaft slender, nearly parallel-sided and only slightly distended, then more pronounced towards apical portion (pars distalis). Ventral surface of pars distalis formed by ventral plate conspicuously widened distally, and at its distal end with a deep and rounded incision (Fig. 88). Glans partially sunken into dorsally depressed portion of pars distalis and extending the distal margin of the ventral plate (Figs 88–90). Capsula externa cylindrical. Capsula interna inside with a dense coat of fur-like microtrichia, including ventral lobe and dorsal lobe; everted ventral lobe ventrally extended beyond the ventral plate, and folded on both sides; everted dorsal lobe embracing the capsula externa, and with two small triangular projections on the sides of the distal margin of pars distalis. Stylus curled by the membrane forming irregular shape. Spination symmetrical. One pair of setae A, B, and F. Two pairs of setae C1–2, D1–2, and E1–2 (Figs 88–92).

Figures 73–86. 

Toccolus globitarsis Suzuki, 1969, male (Figs 73–77) and female (Figs 78–86) 73, 78 body, dorsal view 74, 75 basichelicerite, dorsal view 74 left 75 right 76 left cheliceral hand, frontal view 77 left cheliceral fingers, frontal view 79 left chelicera, medial view 80 same, lateral view 81 left basichelicerite, dorsal view 82 left cheliceral hand, frontal view 83 left pedipalp, medial view 84 same, lateral view 85 ovipositor, ventral view 86 same, dorsal view. Scale bars: 1 mm (73–78, 83–84), 0.5 mm (79–82), 0.25 mm (85–86).

Female

(Figs 78–86, 99–101). Generally similar to male except abdomen slightly wider than in male (Figs 78, 99). The median spine on ocularium greatly enlarged and ocularium with two reduced tubercles above each eye. Chelicerae not enlarged but of normal shape, with a slight difference in inner edges of cheliceral finger (Fig. 82). Femora of pedipalpi dorsally with a row of five setiferous tubercles, distally on medial side with three setiferous tubercles. Tarsal formula (I–IV): 6(2)/15(2)/6/7.

Ovipositor (Figs 85, 86). Ventral surface with four setae and dorsal surface with six setae.

Figures 87–95. 

Toccolus globitarsis Suzuki, 1969, genitalia of male 87 penis, dorsal view 88 pars distalis of penis, ventral view 89, 90 same, lateral view 91 same, dorsal view 92 pars distalis of penis(everted), ventral view 93, 94 same, lateral view 95 same, dorsal view. Abbreviations: CE capsula externa CI capsula interna DL dorsal lobe G glans Pb pars basalis Pd pars distalis S stylus VP ventral plate VL ventral lobe. Scale bars: 0.25 mm.

Measurements

Male (female): body 2.89 (2.41) long, 2.35 (2.10) wide at widest portion, scutum 2.27 (1.80) long. Ocularium 0.45 (0.32) long, 0.77 (0.62) wide. Proximal chelicerae 1.60 (1.01) long, 0.50 (0.36) wide; second 2.56 (1.52) long, 0.92 (0.51) wide; distal 1.15 (0.74) long, 0.25 (0.14) wide. Pedipalpal claw 0.95 (0.87) long. Penis 1.15 long. Measurements of pedipalp and legs as in Tables 5 and 6.

Table 5.

Toccolus globitarsis Suzuki, 1969. Measurements of the pedipalp and legs of the male, as length/width.

Trochanter Femur Patella Tibia Metatarsus Tarsus Total
Pedipalp 0.60/0.41 1.68/0.34 0.81/0.39 1.07/0.40 0.83/0.50 4.99
Leg I 0.30/0.24 1.68/0.18 0.57/0.23 1.13/0.15 1.97/0.06 1.09/0.04 6.74
Leg II 0.34/0.27 2.31/0.15 0.67/0.23 1.89/0.13 2.88/0.08 1.97/0.05 10.06
Leg III 0.39/0.37 1.97/0.25 0.62/0.37 1.30/0.23 2.23/0.11 0.83/0.06 7.34
Leg IV 0.41/0.39 2.40/0.27 0.69/0.36 1.64/0.24 2.86/0.15 1.17/0.08 9.17
Table 6.

Toccolus globitarsis Suzuki, 1969. Measurements of the pedipalp and legs of the female, as length/width.

Trochanter Femur Patella Tibia Metatarsus Tarsus Total
Pedipalp 0.48/0.34 1.23/0.28 0.77/0.30 0.80/0.34 0.64/0.34 3.92
Leg I 0.28/0.21 1.25/014 0.52/0.20 1.04/0.12 1.66/0.06 0.91/0.05 5.66
Leg II 0.33/0.24 1.71/0.13 0.56/0.21 1.07/0.11 2.45/0.06 1.82/0.05 8.57
Leg III 0.31/0.34 1.47/0.19 0.55/0.31 1.15/0.20 1.90/0.11 0.65/0.06 6.03
Leg IV 0.33/0.34 1.45/0.20 0.63/0.30 1.51/0.19 2.62/0.08 1.19/0.06 7.73

Habitat

The specimens were collected in litter and under stones.

Distribution

Thailand (Nakhon Ratchasima Province, Chieng Mai Province, Chaiyaphum Province).

Figures 96–101. 

Photographs of male (Figs 96–98) and female (Figs 99–101) of Toccolus globitarsis Suzuki, 1969. 96, 99 body and parts of appendages, dorsal view 97, 100 body and parts of appendages, lateral view 98, 101 body and parts of appendages, ventral view. Scale bars: 1 mm.

Toccolus kuryi sp. nov.

Figs 102–109, 110–117, 118–123, 124–129

Type material

Holotype male (SMF-CJM7065): THAILAND: Loel Province: Nang Hin District, near cave Wat Tham Malohan (No.18a), 17°06.47'N 101°52.69'E, alt. 362 m, 2 November 2010, S. Dashdamirov leg. Paratype. One female (SMF-CJM7063): THAILAND: Nong Khai Province: Sangkhom District, Tham Tip Waterfall (No. 23), 18°07.57'N 102°11.16'E, alt. 198 m, 3 November 2010, S. Dashdamirov leg.

Diagnosis

The distal part of glans nearly triangular (ventral and dorsal views, Figs 119, 121) and the tip curved ventrally (lateral view, Fig. 120). Basichelicerite dorsally with three spines (Figs 103, 104, 112, 113). Pedipalpal femur ventrally with a row of six setiferous tubercles, the longest one at the base (Figs 108, 116).

Etymology

The specific name is a patronym in honor of Dr. Adriano B. Kury, a well-known arachnologist from Brazil. A. Kury (2000) created a website of Opiliones and built a platform for acquiring knowledge accessible for everyone. He redefined the family Epedanidae to include four subfamilies (Kury 1993).

Description

Male (holotype) habitus as in Figs 102, 110, 124–126. Coloration (Figs 124–126): body light yellow with brown patches; median area of prosoma with brown reticulations; both lateral ridges of scutum with brown stripes; opisthosomal areae I–III each with a transverse brownish band; area IV with a series of three brown flecks; all free tergites brown; all coxae yellow with brown reticulations distally; free sternites each with brown bands, these somewhat paler on the central portion; chelicerae and pedipalps reticulated; trochanters pale yellow, femur, patella, tibia and metatarsus with black reticulations, tarsus lighter.

Figures 102–109. 

Toccolus kuryi sp. nov., male holotype 102 body, lateral view 103 left chelicera, medial view 104 same, lateral view 105 left basichelicerite, dorsal view 106 left cheliceral hand, frontal view 107 left pedipalp, medial view 108 same, lateral view 109 right pedipalp, medial view. Scale bars: 1 mm.

Dorsum (Figs 110, 124). Scutum trapezoid in appearance, both parts nearly parallel-sided, the widest portion of body at opisthosomal area IV, posterior end of opisthosoma bluntly rounded. Prosoma with a row of ten sharply pointed tubercles along anterior margin. Dorsal scutum smooth. Ocularium oval, removed from anterior border of prosoma scutum by 0.38 mm, armed with a conspicuous short median spine of about the height of ocularium (lateral view), four small tubercles above both eyes. The area behind ocularium raised, and the posterior margin of prosoma smoothly rounded. The anterior margin of opisthosomal area I markedly bowed out, borders of other scutal areae slightly bowed out. Opisthosomal areae I–IV with a row of seta-tipped tubercles each and a longitudinal row of similar tubercles on left and right lateral margins. Free tergites with a transverse row of seta-tipped tubercles and anal operculum with scattered granules.

Venter (Fig. 126). Coxae with tubercular surface, coxa I additionally with enlarged seta-tipped tubercles. Coxae II–IV with minute tubercles. Genital operculum and free sternites with seta-tipped granules. Spiracles clearly visible.

Chelicerae (Figs 103–106). Basichelicerite elongate, armed with two long, curved spines, one short spine dorsally; another one distally on the ectal side; six seta-tipped tubercles scattered over the ventral side; medial side with a basal protuberance (Fig. 103). Cheliceral hand considerably widened, pro-dorsally with few seta-tipped tubercles, the inner ones larger than the outer ones; ectally with two seta-tipped tubercles at the subdistal portion; one enlarged seta-tipped tubercle and a smaller one on the ventral side. Fingers strong, inner edges toothed (Fig. 106): moveable finger with five teeth, the proximal one cut rectangularly, the other four pointed; fixed finger with five pointed teeth.

Figures 110–117. 

Toccolus kuryi sp. nov., male (Fig. 110, holotype) and female (Figs 111–117, paratype) 110, 111 body, dorsal view 112 right chelicera, lateral view 113 same, medial view 114 right basichelicerite, dorsal view 115 right cheliceral hand, frontal view 116 right pedipalp, lateral view 117 same, medial view. Scale bars: 1 mm (110–113, 116, 117), 0.5 mm (114, 115).

Pedipalps (Figs 107–109). Coxa dorsally with three seta-tipped tubercles, one strong distal, one sub-distal exterior to it, and a blunt one situated proximally; ventrally with two seta-tipped tubercles. Trochanter ventrally with two setiferous tubercles and dorsally with an enlarge one and a small one. Femur ventrally with a row of five (right pedipalp, Fig. 102) or six (left pedipalp, Fig. 108) setiferous tubercles, the longest one at the base of the row; dorsally with a row of ten setiferous tubercles, distal one and proximal one inconspicuous; disto-medially side with two setiferous tubercles. Patella ventro-mesally with two and ventro-ectally with one setiferous tubercles. Tibia ventro-mesally with three and ventro-ectally with four setiferous tubercles. Tarsus with three setiferous tubercles each ventro-ectally and ventro-medially. Tarsal claw massive, curved, longer than tarsus.

Legs. All segments unarmed, nearly smooth. Legs I–II slender, legs III–IV strong. Femora I–III not curved, almost straight, femur IV slightly curved. Distitarsi I and II with two tarsomeres. Distitarsi III and IV without scopula and with two unpectinated claws. Tarsal formula (I–IV): 7(2)/16(2)/7/7.

Penis (Figs 118–121). Shaft slender, nearly parallel-sided, apical part (pars distalis) distended. Ventral plate conspicuously constricted apically, and dorsally with deep indented near the glans. Glans partially sunken into dorsal depressed portion of pars distalis and extended the distal margin of the ventral plate (Fig. 120). The distal part of glans almost triangular (ventral and dorsal views, Figs 119, 121), the tip curved ventrally. Spination symmetrical. One pair of setae A, B, and F. Two pairs of setae C1–2, D1–2, and E1–2 (Figs 120, 121).

Female

(SMF-CJM7063) (Figs 111–117, 122, 123, 127–129). Body nearly oval. Granulation and spination of body similar to the male (Figs 111, 127). Prosoma with a row of eight sharp pointed tubercles along anterior margin (Fig. 111). Surface of dorsum smooth. Ocularium with only one conspicuous tubercle near the left eye except for the long median spine. Chelicerae of normal shape, not enlarged, slight difference in inner edges of cheliceral finger (Fig. 115). Femora of pedipalpi ventrally with six and dorsally with a row of five setiferous tubercles, disto-medially with three setiferous tubercles. Tarsal formula (I–IV): 6(2)/15(2)/7/7.

Ovipositor (Figs 122, 123). Ventral side with four, dorsal side with six setae.

Measurements. Male (female): body 2.27 (2.76) long, 2.08 (1.96) wide at widest portion, scutum 1.95 (1.79) long. Ocularium 0.41 (0.30) long, 0.69 (0.51) wide. Proximal chelicerae 1.24 (0.84) long, 0.47 (0.31) wide; second 2.20 (1.37) long, 0.78 (0.45) wide; distal 0.97 (0.62) long, 0.22 (0.14) wide. Pedipalp claw 0.82 (0.79) long. Penis 1.02 long. Measurements of pedipalp and legs as in Tables 7 and 8.

Table 7.

Toccolus kuryi sp. nov. Measurements of the pedipalp and legs of the male holotype, as length/width.

Trochanter Femur Patella Tibia Metatarsus Tarsus Total
Pedipalp 0.56/0.43 1.56/0.30 0.98/0.32 0.95/0.35 0.70/0.38 4.75
Leg I 0.35/0.24 1.60/0.17 0.52/0.23 1.12/0.13 1.94/0.06 1.06/0.04 6.59
Leg II 0.34/0.25 2.15/0.14 0.59/0.21 1.86/0.12 2.74/0.05 1.94/0.04 9.62
Leg III 0.42/0.29 1.86/0.22 0.61/.032 1.30/0.21 2.21/0.12 1.03/0.07 7.43
Leg IV 0.48/0.35 2.32/0.24 0.63/0.32 1.76/0.21 2.71/0.11 1.15/0.05 9.05
Table 8.

Toccolus kuryi sp. nov. Measurements of the pedipalp and legs of the female paratype, as length/width.

Trochanter Femur Patella Tibia Metatarsus Tarsus Total
Pedipalp 0.42/0.34 1.28/0.23 0.78/0.26 0.87/0.27 0.66/0.29 4.01
Leg I 0.24/0.22 1.24/0.15 0.48/0.19 0.93/0.11 1.54/0.06 0.64/0.04 5.71
Leg II 0.22/0.19 1.72/0.13 0.52/0.19 1.51/0.10 2.27/0.05 2.46/0.04 8.70
Leg III 0.26/0.30 1.35/0.18 0.53/0.27 1.07/0.19 1.79/0.09 0.76/0.08 5.76
Leg IV 0.32/0.28 1.67/0.19 0.56/0.25 1.39/0.17 2.48/0.08 1.13/0.07 7.55

Habitat

The specimens were collected under stones (SMF-CJM7065) and under Acacia bark and rotten stumps (SMF-CJM7063).

Distribution

Thailand (Loel Province, Nong Khai Province).

Figures 118–123. 

Toccolus kuryi sp. nov., genitalia of male holotype 118 penis, dorsal view 119 pars distalis of penis, dorsal view 120 same, lateral view 121 same, ventral view 122 ovipositor, ventral view 123 same, dorsal view. Abbreviations: G glans Pb pars basalis Pd pars distalis VP ventral plate. Scale bars: 0.5 mm (118), 0.25 mm (119–123).

Figures 124–129. 

Photographs of male (Figs 124–126, holotype) and female (Figs 127–129, paratype) of Toccolus kuryi sp. nov. 124, 127 body and parts of appendages, dorsal view 125, 128 body and parts of appendages, lateral view 126, 129 body and parts of appendages, ventral view. Scale bars: 1 mm.

Discussion

According to the present results, the typical members of Epedanidae (Acrobuninae Roewer, 1912; Sarasinicinae Roewer, 1923; Epedaninae Sørensen, 1886; and Dibuninae Roewer, 1912) possess similar external morphology, e.g., “the greatly elongate pedipalps, high erect spine on eye tubercle, fused scutal areas I–II etc” (Lian et al. 2008: 58). However, the morphology of the male genitalia of epedanids is in fact highly diverse.

The penis can be classified at least into two different types by the shape of the ventral plate. One type of ventral plate is well defined by a single plate (Figs 130, 131) which is shovel-shaped (Fig. 130) and usually is C-shaped in cross-section. Its distal border greatly extends the penis by more than the length of the glans (Fig. 130).

The other type of ventral plate is called a ventral frame (Zhang and Martens 2018) and is not sharply defined by a three-dimensional structure (Figs 132–137). This plate is circularly shaped in cross-section (Fig. 135) and is almost completely covered by the glans (Fig. 132). The distal margin of ventral plate is straight or concave.

The structures of the capsula externa and the capsula interna fall into at least two groups. One group is characterized by a saccular and circular capsula externa and capsula interna (Figs 134, 135a, b). The other group is defined by an arc-shaped capsula externa and capsula interna (Figs 136, 137a, b). Until now it is not clear if these differences may have an impact on epedanid taxonomy and systematics.

We assume that during copulation glans and stylus are exposed by internal hemolymph pressure. The movement causes the glans sclerites to expand under the internal pressure in two different ways. The trajectory of saccular and circular capsula externa and capsula interna is unidirectional and they are extruded in straight prolongation of the entire penis cane (Figs 138, 139a, b). In contrast, the movement of penes with the arc-shaped capsula externa and capsula interna appears opposite and they move up and down according of the internal pressure applied (Figs 60, 140, 141a, b; e.g., Lian et al. 2011: 48, fig. 31).

Figures 130–141. 

Schematic drawings contrasting the morphology of pars distalis (Pd) of male genitalia in different representatives of the Epedanidae 130, 131 Epedanellus tuberculatus Roewer, 1911 (arrow indicates the cross-section of ventral plate at the marked point, adapted from Suzuki, 1973) 130 ventral view 131 lateral view 132, 133 Lobonychium palpiplus Roewer, 1938 (arrow indicates the cross-section of ventral plateat the marked point, adapted from Zhang & Martens, 2018) 132 dorsal view 133, 134 lateral view 135a, b cross-section through ventral plate and glans at the marked point 136, 137, 140, 141 Plistobunus jaegeri sp. nov. 136 lateral view 137 cross-section through ventral plate and glans at the marked point 138, 139 Toccolus globitarsis Suzuki, 1969 138 lateral view 139a, b expanded, lateral view (arrow indicates the movement of capsula interna in 139b) 140 lateral view 141a, b expanded, lateral view (arrow indicates the movement of capsula interna and capsula externa in 141b).

Acknowledgements

We are grateful to Peter Jäger for providing access to the opilionid collection of Senckenberg Research Museum, Frankfurt am Main, Germany, and to Julia Altmann for her very kind attention during Chao Zhang’s visit to the SMF. Cordial thanks are due to Selvin Dashdamirov who put material at our disposal. This work was supported by the Researcher Alumni Network Senckenberg, Frankfurt am Main, Germany (CZ) and by Feldbausch Foundation and Wagner Foundation of Fachbereich Biologie of Johannes Gutenberg University, Mainz for fieldwork in Asia (JM).

References

  • Acosta LE, Pérez-González A, Tourinho AL (2007) Methods for taxonomic study. In: Pinto-da-Rocha R, Machado G, Giribet G (Eds) Harvestmen: The biology of Opiliones. Harvard University Press, Cambridge and London, 494–510.
  • Kury AB (1993) Análise filogenètica de Gonyleptoidea (Arachnida, Opiliones, Laniatores). PhD Thesis, Universidade de São Paulo, São Paulo.
  • Kury AB (2008) On the systematic position of Dino Loman and Toccolus Roewer (Opiliones, Laniatores, Epedanidae), with the description of a new species from western Java, Indonesia. Zootaxa 1932: 61–68. https://doi.org/10.11646/zootaxa.1932.1.6
  • Kury AB, Villarreal MO (2015) The prickly blade mapped: establishing homologies and a chaetotaxy for macrosetae of penis ventral plate in Gonyleptoidea (Arachnida, Opiliones, Laniatores). Zoological Journal of the Linnean Society 174: 1–46. https://doi.org/10.1111/zoj.12225
  • Lian WG, Zhang C, Zhang F (2011) Review of the genus Plistobunus Pocock, 1903, with description of a new species from Hainan Island, China (Opiliones, Laniatores, Epedanidae). ZooKeys 112: 39–52. https://doi.org/10.3897/zookeys.112.1110
  • Macías-Ordóñez R, Machado G, Pérez-González A, Shultz J (2010) Genitalic Evolution in Opiliones. In: Leonard J, Córdoba-Aguilar A (Eds) The Evolution of Primary Sexual Characters in Animals. Oxford University Press, New York, 285–306.
  • Martens J (1986) Die Großgliederung der Opiliones und die Evolution der Ordnung (Arachnida). In: Barrientos JA (Ed.) Actas del X Congreso Internacional de Aracnología. Editorial Juvenil, Barcelona, 289–310.
  • Roewer CF (1912) Die Familien der Assamiiden und Phalangodiden der Opiliones-Laniatores. (= Assamiden, Dampetriden, Phalangodiden, Epedaniden, Biantiden, Zalmoxiden, Samoiden, Palpipediden anderer Autoren). Archiv für Naturgeschichte, Berlin, Abt. A, Original-Arbeiten 78(3): 1–242.
  • Roewer CF (1915) 106 neue Opilioniden. Archiv für Naturgeschichte, Berlin, Abt. A, Original-Arbeiten 81(3): 1–152.
  • Roewer CF (1923) Die Weberknechte der Erde. Systematische Bearbeitung der bisher bekannten Opiliones. Gustav Fischer, Jena, 1116 pp.
  • Roewer CF (1927) Ostasiatische Opiliones, von Herrn Prof. F. Silvestri im Jahre 1925 erbeutet. Bollettino del Laboratorio di Zoologia Generale e Agraria della Facoltà Agraria in Portici 20: 192–210.
  • Roewer CF (1938) Über Acrobuninae, Epedaninae und Sarasinicinae. Weitere Weberknechte IX. (9. Ergänzung der "Weberknechte der Erde" 1923). Veröffentlichungen aus dem Deutschen Kolonial- und Übersee-Museum in Bremen 2(2): 81–169.
  • Schwendinger PJ, Martens J (2002) A taxonomic revision of the family Oncopodidae III: Further new species of Gnomulus Thorell (Opiliones, Laniatores). Revue suisse de zoologie 109: 47–113. https://doi.org/10.5962/bhl.part.79580
  • Simon E (1879) Les Arachnides de France. Tome 7. Contenant les ordres des Chernetes, Scorpiones et Opiliones. Librairie Encyclopédique de Roret, Paris, 332 pp.
  • Sørensen WE (1886) Opiliones. In: Koch L, Keyserling E (Eds) Die Arachniden Australiens nach der Natur beschrieben und abgebildet, vol. [Theil] 2 of two volumes & atlas, 1871–1889, fascicle [Lieferung] 33. Bauer & Raspe, Nürnberg, 53–86.
  • Suzuki S (1969a) On a collection of opilionids from Southeast Asia. Journal of Science of the Hiroshima University 22(2): 11–77.
  • Suzuki S (1969b) Some phalangids from Thailand. Journal of Science of the Hiroshima University 22(3): 79–101.
  • Suzuki S (1973) Opiliones from the South-west Islands, Japan. Journal of Science of the Hiroshima University 24: 205–279.
  • Suzuki S (1976) Report on a collection of opilionids from Pasoh Forest Reserve, West Malaysia. Nature and Life in Southeast Asia 7: 9–38.
  • Suzuki S (1981) Three opilionids from Thailand. Annotationes Zoologicae Japonenses 54(4): 267–272.
  • Suzuki S (1985) A synopsis of the Opiliones of Thailand (Arachnida) I. Cyphophthalmi and Laniatores. Steenstrupia 11(3): 69–110.
  • Suzuki S, Stone FD (1986) The fauna of Thai caves. 1. Three phalangids from Thailand (Arachnida). Occasional Papers Bernice P. Bishop Museum 26: 123–127.
  • Thorell TTT (1876) Descrizione di alcune specie di Opilioni dell' Arcipelago Malese appartenenti al Museo Civico di Genova. Annali del Museo Civico di Storia Naturale di Genova 9: 111–138.
  • Zhu MS, Lian WG (2006) First record of the genus Euepedanus from China, with the description of a new species (Opiliones: Laniatores: Epedanidae). Zootaxa 1367: 63–68. https://doi.org/10.11646/zootaxa.1367.1.4