Zoosyst. Evol. 100 (1) 2024, 1-8 | DO! 10.3897/zse.100.110020 > PENSUFT. Ge Ee BERLIN A new species of Ditha (Pseudoscorpiones, Chthoniidae, Tridenchthontinae) from the Western Ghats of India, with an identification key for the genus Kyung-Hoon Jeong!*, Danilo Harms**°, Jithin Johnson?®’ 1 Seoul National University, 1, Gwanak-ro, Gwanak-gu, Seoul, 08826, Republic of Korea 2 National Institute of Biological Resources, Species Diversity Research Division, Environmental Research Complex, Hwangyeong-ro 42, Seo-gu, Incheon, 22689, Republic of Korea 3 Museum of Nature Hamburg — Zoology, Leibniz Institute for the Analysis of Biodiversity Change, Martin-Luther-King-Platz 3, Hamburg, 20146, Germany 4 Harry Butler Institute, Murdoch University, Murdoch, Western Australia; Australian Museum Research Institute, Australian Museum, Sydney, Australia 5 SARCHI Chair on Biodiversity Value and Change, Centre for Invasion Biology, University of Venda, Thohoyandou, South Africa 6 Sacred Heart College (Autonomous), Thevara, Kerala, 682013, India 7 St. Joseph's College (Autonomous), Irinjalakuda, Kerala, 680121, India https://zoobank. org/EF0354EC-674D-43F9-B70A-BCB903 1SDF3C Corresponding author: Kyung-Hoon Jeong (ds16203@snu.ac.kr) Academic editor: Pavel Stoev # Received 25 July 2023 Accepted 7 September 2023 @ Published 26 January 2024 Abstract A new species of the pseudoscorpion genus Ditha Chamberlin, 1929, is described from Kerala State, India. A detailed morpholog- ical description, diagnostic features, and illustrations of Ditha (Paraditha) shivanparaensis Jeong, Harms & Johnson, sp. nov. are provided. The current distribution of all the known Ditha species is mapped, and an identification key for the genus is provided. Key Words Arachnida, Kerala, montane, morphology, taxonomy Introduction Tropical montane cloud forests or ‘sholas’ of the Western Ghats of India encompass unique ecosystems character- ized by their high elevations (>1000 m) and isolated na- ture. These sholas are considered as isolated ‘islands’ that are surrounded by the vast ocean of low-altitude forests or grasslands (Robin et al. 2015). The biota of these so- called ‘sky islands’ provide excellent templates for study- ing endemism and investigating the factors driving diver- sification and adaptation tn such specialized environments. Pseudoscorpions (order Pseudoscorpiones de Geer, 1778) are one such group of arachnids found in these ecosystems. Globally, ca. 4000 nominal species are described in 25 fam- ilies, with India hosting ca. 160 described species (WPC 2023). Due to their specific microhabitat preferences and poor dispersal abilities, most leaf-litter dwelling species, particularly those belonging to the family Chthoniidae Da- day, 1889, exhibit short-range distributions (Cosgrove et al. 2016; Johnson et al. 2022; Hlebec et al. 2023). Tridenchthoniinae Balzan, 1892 is a chthoniid subfamily with 71 extant species in 15 genera and one extinct species (Chelignathus kochii Menge, 1854). Until recently, it was regarded as a distinct family within the Chthonioidea super- family (Benavides et al. 2019). It differs from other chthoni- id subfamilies in the presence of granulate carapace, spines on both coxa I and II, trichobothria ib and isb located basal- ly or sub-basally on the chelal hand, and a galea with mul- Copyright Jeong, K-H. 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. - tiple ramifications in the juvenile stage (Morikawa 1960; Kennedy 1989). Tridenchthoniinae is divided into two tribes, Tridenchthoniini and Verrucadithini, and they can be distinguished by the shape of the marginal chelal teeth, being relatively acute and distinctly spaced in Verrucadith- ini, while they are closely contiguous in Tridenchthoniini (Chamberlin and Chamberlin 1945). Tridenchthoniini com- prises nine genera; among these, the genus Ditha Chamber- lin, 1929 is distributed from East Asia to Africa. It includes two subgenera, Ditha (Ditha) and Ditha (Paraditha) Beier, 1931, with seven species recorded in each subgenus. The subgenus Paraditha comprises seven species: D. /aosana Beier, 1951, D. pahangica Beier, 1955, D. sumatraensis (Chamberlin, 1923) and D. tonkinensis (Beier, 1951) dis- tributed in Southeast Asia; D. /atimana (Beier, 1931) and D. sinuata (Tullgren, 1901) from Africa; and D. marcusen- sis (Morikawa, 1952) from East Asia (Fig. 1). Hitherto, only two species of Tridenchthoniinae be- longing to the genus Compsaditha Chamberlin, 1929 have been reported from India: Compsaditha indica Murthy, 1960, and C. camponota Sivaraman, 1980. Here, we present the first record of the genus Ditha (Paraditha) from India and describe a new species from the Southern Western Ghats (Kerala state) of India along with an iden- tification key for the genus Ditha. Materials and methods The specimens used for this study are lodged in the fol- lowing institutions: Museum of Nature, Hamburg — Zo- ology (ZMH, formerly Zoological Museum Hamburg) and the Arachnology Division of Sacred Heart College, Thevara, India (ADSH). All specimens were initially pre- served in 100% ethanol, with DNA extracted from the legs of one paratype female, while the other specimens are stored in 75% ethanol. All specimens were examined using a Leica M125C ste- reomicroscope in 75% ethanol. Images were captured using a BK Plus Lab System (Dun, Inc.) equipped with a Canon EOS 7D Mark II camera fitted with a microscopic lens (SX, 10X magnification) using the software CAPTURE ONE PRO 9.3 64 Bit (v.9.3.0.85) and stacked using Zerene Stacker, and a Leica DMC4500 digital camera attached to a Leica M205A stereomicroscope, using the program, LEICA APPLICATION SUITE X (LASX), ver. 3.0.1. LASX was also used for taking the measurements. Mensuration and terminology largely follow Chamberlin (1931), with some minor modifications to the terminology of the trichobothria (Harvey 1992), chelicera (Judson 2007), and appendages (Harvey et al. 2012). Scanning electron micrographs were obtained with a Hitachi TM4000Plus scanning electron micrograph (SEM) system. Illustrations were created using ADOBE ILLUSTRATOR 2023 and edited with ADOBE PHOTOSHOP 2023. The distribution map was made using QGIS 3.22.10. Abbreviations used for chelal trichobothria: b — basal, sb — subbasal, st — subterminal, ¢ — terminal, ib — internal basal, isb — internal subbasal, eb — external basal, esb — external subbasal, it — internal terminal, ist — internal zse.pensoft.net Jeong, K-H. et al: A new species of Ditha from the Western Ghats of India subterminal, ef — external terminal, est — external subtermi- nal, xs — duplex trichobothria; leg segments: tr — trochanter, fe—femur, pa — patella, i — tibia, mt — metatarsus, ta — tar- sus; cheliceral seta: as — accessory setae. Taxonomy Family Chthoniidae Daday, 1889 Subfamily Tridenchthoniinae (Balzan, 1892) Remarks. Balzan (1892) proposed the family Tridench- thoniidae to include the species 7ridenchthonius parvulus Balzan, 1887 from Paraguay which possessed triple galea in the chelicera. However, Hansen (1894) considered this species to be an unusual Chthonius and reduced the taxon to the subfamily Tridenchthoniinae. Since then, the taxo- nomic status of the group has undergone several changes. Chamberlin (1929) in his revised classification scheme for pseudoscorpions, proposed a new subfamily Dith- inae Chamberlin, 1929 to include the new genus Ditha Chamberlin, 1929. Dithinae chthoniids closely resemble Tridenchthoniinae, except that they include non-gale- ate forms, but they differ from members of Chthontinae proper in the characters such as the presence of coxal spines (of single simple type) on coxa I and II and the patella of leg IV being distinctly longer in relation to the femur. Chamberlin (1931) later raised this group to the family status. However, without providing proper justifi- cation, Beier (1932) reduced the family to the subfamily status and later to Dithiidae again. However, recogniz- ing Balzan’s species T. parvulus as a Dithiidae member, Chamberlin and Chamberlin (1945) proposed the family Dithiidae as a junior synonym of Tridenchthoniidae. Fi- nally, in a recent study on pseudoscorpion transcriptom- ics (Benavides et al. 2019), the group was again reduced to a subfamily within Chthoniidae that now includes Tri- denchthoniinae as a subfamily, alongside the Chthoniinae and Lechytiinae. Genus Ditha Chamberlin, 1929 Type species. Ditha elegans Chamberlin, 1929. Diagnosis. The genus Ditha can be diagnosed by the following combination of characters: tergites with biseri- ate setae; carapace with more than 50 setae; trichobothria st and ¢ more than one areolar diameter apart, and triple galea in the juvenile stage. Ditha is divided into two sub- genera, Ditha (Ditha) and Ditha (Paraditha). The sub- genera Paraditha and Ditha can be differentiated based on the intercoxal tubercle and the number of accessory setae on the cheliceral palm (Beier, 1955). In Ditha, the intercoxal tubercle is strongly distinct and 10—12 acces- sory setae are present on the cheliceral palm, while the intercoxal tubercle is indistinct or even absent in Para- ditha and only 1—2 accessory seta present on the cheli- ceral palm. Zoosyst. Evol. 100 (1) 2024, 1-8 Distribution of the species D (D). elegans D (D). loricata D (D). novaeguieneae D (D). ogasawarensis D (D). palauensis D (D). philippinensis D (PD). proxima D (P). laosana D (P). latimana D (P). marcusensis D (P). pahangica D (P). sinuata D (P). sumatraensis D (P). tokinensis Ditha (Paraditha) shivanparaensis sp. nov FHP PrPrPPrPPrPepPpeeecedes ®@ Pee Figure 1. Distribution of the Ditha (Ditha) and Ditha (Paraditha) species. Ditha (Paraditha) shivanparaensis sp. nov. https://zoobank. org/879135E3-C951-4A67-9FF8-4233D 198E34B Figs 1-4 Type material. Holotype (Fig. 2A, B). Female (ADSH PSTOOO1. INDIA: Kerala: Shivanpara, Mathiket- tan Shola National Park, Kerala, Idukki, 9°58'05.1"N, 77°13'43.5"E, altitude 1072 m, litter sample (sifting and Berlese trap), J. Johnson leg. Paratypes (Fig. 2C, D): 3 females (ADSH PSTO0003, ADSH PST0004, ZMH-A0013527) and one male (ADSH PST0002), same data as holotype. Etymology. This species is named after the hill ‘Shivanpara’ in the Mathiketan Shola National Park, where all specimens were collected. Diagnosis. Within the subgenus Paraditha, Ditha shivanparaensis sp. nov. closely resembles D. tonkinen- sis Beier, 1951 found in Vietnam in having an indistinct intercoxal tubercle between coxa III and IV, and presence of eight setae on the posterior margin of the carapace. However, the new species differs from D. tonkinensis and from all other species within the subgenus in having two accessory setae on the cheliceral palm. Ditha shivanpar- aensis sp. nov. also bears similarity to Ditha (Ditha) prox- ima (Beier, 1951) from Vietnam in terms of having two accessory setae on the cheliceral palm and 45-47 margin- al teeth on the movable chelal finger. However, it differs from the latter in possessing 14 setae on the anterior mar- gin of the carapace (as opposed to 10 setae in D. proxima) and by the positioning of sz, 1.e., halfway between sb and ¢ (whereas st is much closer to sb in D. proxima). Description. Female, adult (holotype, Fig. 2A, B). Colour. Uniformly orange-brownish, the legs lighter than the body. Chelicera (Figs 3F, H, 4D, E). Cheliceral palm coarse- ly granulate and with seven setae (including two accesso- ry setae), movable finger with one seta in medial position and seven marginal teeth, inner margin granulate; fixed finger with seven marginal teeth, the terminal one larger than the others (Fig. 3H); with two dorsal lyrifissures; ral- lum with ten blades (Figs 3F, 4E); serrula exterior with 16 blades of similar size. Pedipalp (Figs 3G, 4B, C). Trochanter 1.44 times, fe- mur 3.54 times, patella 1.83 times, chela (with pedicel) 3.79 times, hand 1.51 times longer than broad, movable finger 1.54 times longer than hand. Femur, patella, chela smooth. Fixed chelal finger with six trichobothria, mov- able finger with four trichobothria, ib and isb situated basally on the dorsum of the chelal hand; sb, st, and ¢ lo- cated separately from b; sb, st and ¢ all spaced more than one areolar distance; st slightly closer to sb than £, eb, esb and ist forming a group at basal to subbasal position; esb slightly closer to ist than eb; est and it positioned at the median on the fixed finger; et closely located with xs, almost less than one areolar distance (Figs 3G, 4B). Four lyrifissures on the dorsum of fixed chelal finger, one on the basal dorsum of chelal hand. Both fingers with small juxtadentate teeth; fixed finger with 55 triangular and ret- rorse teeth; movable finger with 47 rounded teeth. Cephalothorax (Figs 3A, D, E, 4F, G). Carapace 1.04 times longer than broad sub-rectangular; lateral margins almost parallel but slightly wide at the base; coarsely zse.pensoft.net Jeong, K-H. et al: A new species of Ditha from the Western Ghats of India Figure 2. Habitus of Ditha shivanparaensis sp. nov. A. Female holotype, dorsal view; B. Female holotype, ventral view; C. Male paratype, dorsal view; D. Male paratype, ventral view. Scale bars: 1 mm. granulate, without furrows; four corneate eyes; ante- rior margin serrated; epistome small and serrated; with 94 setae; 14 setae on the anterior margin, eight setae on posterior margin; setae short and acuminate; with ten lyr- ifissures, three each on the sides near the anterior margin, one between each eye, and one on each side situated near the posterior margin (Fig. 3A). Manducatory process with one long, acuminate setae, remainder of maxilla with ten setae; four lyrifissures on the maxilla. Coxal chaetotaxy: 10: 11: 11: 20 (Fig. 3D). Coxa I with ca. 5-6 spines, each spine serrated terminally, and ca. 2-3 spines from one base; coxa II with ca. 5-6 spines (Figs 3E, 4E, F); each spine serrated from the medial. Intercoxal tubercle indistinctly present between coxa III and IV, seta weakly present (Fig. 4F). zse.pensoft.net Abdomen. Pleural membrane papillostriate, tergites un- divided, but sternites III-IV partially divided; setae biseriate and acuminate. Tergal chaetotaxy, 14: 23: 23: 25: 26: 24: 24: 25:21: 17: 13: 0. Sternal chaetotaxy, 15: 10: 8: 8: 8: 8: 8: 9: 12: 4: 2. Sternite I with 4 setae in the genital opening area. Legs (Fig. 3B, C). Leg I: trochanter 1.18 times, femur 3.87 times, patella 2.98 times, tibia 2.56 times, tarsus 5.93 times longer than broad, leg IV: trochanter 1.48 times, fe- mur 2.75 times, tibia 3.44 times, metatarsus 2.57 times, tarsus 7.00 times longer than broad; leg IV: trochanter 1.67 times, femur 2.39 times, tibia 3.47 times, metatarsus 2.57 times, tarsus 5.49 times longer than broad. Tarsus of leg IV longer than the metatarsus; arolium undivided and shorter than the claws; pseudotactile seta located basally on leg I patella and medially on leg IV metatarsus. Zoosyst. Evol. 100 (1) 2024, 1-8 Figure 3. Drawings of Ditha shivanparaensis sp. nov. (all paratypes unless mentioned otherwise) A. Carapace, dorsal view; B. Leg IV; C. leg I; D. Coxa; E. Coxal spines; F. Rallum; G. Left chela from the lateral; H. Cheliceral, dorsal view, two accessory setae in the box; I. Male genital area external view; J. Female genital area external view (holotype). Scale bars: 0.5 mm (A—D, G); 1 mm (E-F); 2 mm (H—J). Genitalia (Fig. 3J). Typical shape of the genus Ditha. 15 setae in the genital opening area, including nine setae on the center of the opening and three setae each on either side of the opening; ten setae on the sternite III; six setae on the anterior part of sternite HII; four setae on the poste- rior region, and two setae each on the part of sternite III. Dimensions (in mm). Body length 1.47; Pedipalp: trochanter 0.22/0.15, femur 0.48/0.13, patella 0.27/0.15, chela (with pedicel) 0.72/0.19, movable finger 0.45, hand 0.29/0.19; Chelicera: total 0.32/0.18, movable finger 0.17; Cephalothorax: Carapace 0.47/0.45, anterior eye 0.005, posterior eye 0.004; Leg I: trochanter 0.13/0.11, femur 0.27/0.07, patella 0.19/0.06, tibia 0.15/0.06, tarsus 0.27/0.05; Leg IV: trochanter 0.16/0.11, femur + patella 0.47/0.17, tibia 0.33/0.10, metatarsus 0.15/0.06, tarsus 0.28/0.04. Male, adult (paratype, Fig. 2 C, D). Same as the holotype, except slightly smaller (body length 1.291 mm). Pedipalp. Trochanter 1.46 times, 3.54 times, patella 1.87 times, chela (with pedicel) 3.85 times, hand 1.47 times longer than broad, movable finger 1.56 times lon- zse.pensoft.net Jeong, K-H. et al: A new species of Ditha from the Western Ghats of India Figure 4. Ditha shivanparaensis sp. nov. A. Carapace, dorsal view; B. Left chela, lateral view; C. Left chela, dorsal view; D. Cheliceral, dorsal view; E. Rallum; F. Coxa (intercoxal tubercle in the circle); G Coxal spine. Scale bars: 200 um (A—C); 100 um (D, F); 30 um (E); 50 um (G). ger than hand; fixed finger with 50, movable finger with 48 teeth. Cephalothorax. Carapace 1.06 times longer than broad; with 92 setae; 14 setae on the anterior margin, eight setae on the posterior margin; Coxal chaetotaxy: 10: 11: 11: 20. Coxa I with ca. 5—6 spines. Abdomen. Tergal chaetotaxy, 10: 14: 15: 16: 19: 19: 19: 18: 16: 15: 13: 0. Sternal chaetotaxy, 10: 36: 6: 8: 10: 8: 13: 13: 11: 5: 2. Sternite I with nine total setae includ- ing six setae near the genital opening, sternite III with 38 total setae including 22 setae near the genital opening. zse.pensoft.net Legs. Leg I: trochanter 1.33 times, femur 3.65 times, patella 2.92 times, tibia 2.58 times, tarsus 5.80 times lon- ger than broad; leg IV: trochanter 1.67 times, femur 2.39 times, tibia 3.47 times, metatarsus 2.57 times, tarsus 5.49 times longer than broad. Genitalia (Fig. 31). Typical shape of the genus Ditha. 15 setae in the genital opening area, including nine on the center of the opening and three each on either side of the Opening; ten setae on the sternite III; six setae on the ante- rior part of sternite III; four setae on the posterior region, and two setae each on the part of sternite III. Zoosyst. Evol. 100 (1) 2024, 1-8 Dimensions (mm). Body length 1.29. Pedipalp: trochan- ter 0.19/0.13, femur 0.44/0.12, patella 0.24/0.13, chela (with pedicel) 0.64/0.17, movable finger 0.38, hand 0.24/0.17; Chelicera: total 0.30/0.16, movable finger 0.17; Cephalo- thorax: Carapace 0.42/0.39, anterior eye 0.005, posterior eye 0.004; Leg I: trochanter 0.12/0.09, femur 0.24/0.07, patella 0.16/0.06, tibia 0.39/0.09, tarsus 0.24/0.04; Leg IV: trochanter 0.18/0.11, femur + patella 0.40/0.17, tibia 0.31/0.09, metatarsus 0.14/0.05, tarsus 0.26/0.05. Variation of female paratypes (n=2). Pedipalp. Tro- chanter 1.44—-1.46 times, femur 3.46—3.68 times, patella 1.85—1.98 times, chela (with pedicel) 3.65—3.74 times, hand 1.48—1.52 times longer than broad, movable finger 1.43—1.44 times longer than hand. Movable finger with A8, fixed finger with 53-54 marginal teeth. Cephalothorax. Carapace 1.04 times longer than broad. 96—100 setae. Abdomen. TYergal chaetotaxy, 13-14: 21-23: 22-24: 24-25: 24-25: 25: 25-26: 23-25: 18: 15-16: 14-15: 0. Sternal chaetotaxy, 10: 10: 8: 7-8: 9-11: 8-10: 8-9: 10: 11-13: 4: 2. Legs. Leg I: trochanter 1.20—1.34 times, femur 3.60— 3.93 times, patella 2.61—2.64 times, tibia 2.47—2.77 times, tarsus 5.88—6.08 times longer than broad, leg IV: trochan- ter 1.59-1.66 times, femur 2.38—2.53 times, tibia 3.40— Key to the species of Ditha 7 3.55 times, metatarsus 2.17—2.29 times, tarsus 5.39-6.14 times longer than broad. Dimensions (mm): Body length 1.33—1.41; Pedipalp: trochanter 0.21-0.22/0.15, femur 0.48—0.51/0.13-0.15, patella 0.28-0.30/0.15, chela 0.71-0.73/0.19-0.20, movable finger 0.41-0.43, hand 0.29—0.30/0.19-0.20; Chelicera: total 0.29-0.31/0.18—-0.19, movable finger 0.18—0.19; Cephalothorax: carapace 0.46—0.47/0.45, an- terior eye 0.005—0.006, posterior eye 0.004; Leg I: tro- chanter 0.12—0.13/0.10, femur 0.28/0.07-0.08, patella 0.17—0.19/0.07, tibia 0.14—0.16/0.06, tarsus 0.27/0.04— 0.05; Leg IV: trochanter 0.21/0.13, femur + patella 0.43- 0.44/0.17-0.18, tibia 0.35—0.37/0.10—0.11, metatarsus 0.14—0.15/0.06—0.07, tarsus 0.27—0.30/0.05. Distribution. Currently known only from the type locality. Habitat. The species was collected from moist leaf litter of montane ‘shola’ forests of the Mathiketan Shola National Park and is probably endemic to this mountain, although more field sampling 1s required to verify this. Erratum. The sequences with GenBank accession numbers OM792092.1, OM832661.1 and OM876917.1, previously attributed to the genus Compsaditha by John- son et al. (2022), actually belong to the newly discovered species described herein as Ditha shivanparaensis sp. nov. i Intercoxal tubercle distinct; 10-12 accessory seta positioned on the cheliceral palM...............:.:::006: 2, subgenus Ditha NM | Cheliceral palm with less than six accessory setae.......... - Cheliceral palm with more than six accessory setae........ S Carapace witht less than lO@sseta es ret ccc s2l centtey tole - Carapacewithrmore thar lOO Seta ni. ieodeey sane sseesewes Ae Catapace witit léss-thanyl SO'Setae 24.5 eco sree hehe - Carapace withimore that PSG setae. os auuieteesganntsiteeaun 5 12 setae on the anterior margin of the carapace............. — 14 setae on the anterior margin of the carapace............. Or —“Sereite witht, de Serae rae). i ss5 tenn cetercqarpentcsooMereneeass - Tergite | with 21 setae in male..................ccccsccseesseeeeeeeees ch 14 setae on the anterior margin of the carapace............. - 16 setae on the anterior margin of the carapace............. 8 [MISKCOXa| TUBSECIS AlSSIiE er Ue rnc osee reed ry 5 weak aod tes SPE - Posterior margin of carapace with 14 setae IG heCarapace more tna: /Oesetae.. Mie. tert cot) enor ee sie - Carapace fewer than 70 Setae..............::cccceeeeeeeeeeeeseeeseees 12 Carapace withemore than WO0-setaea esos sccrs28 hee noes - Carapace with fewer than 100 setae 13-« Carapace with Moré than-70 Selaes. ce cccccs:sssteacesurensseceess - Carapace with fewer than 70 setae ................2.::::cceeeeeees 14 Pedipalpal femur at least 4 times longer than broad....... - Pedipalpal femur at least 4 times longer than broad....... b> JCheliceranwithisl accessory Selae + 154.22.csb ye oletertsh cet - Chelicera with 2 accessory setae 10 Posterior margin of carapace with 8 setae..............:::0065 Intercoxal tubercle indistinct or absent; 1-2 accessory seta positioned on the cheliceral palm.....9, subgenus Paraditha occ, bee hag COMES tS Sa, Mie M attri D. proxima (Beier, 1951) (Vietnam) nat hx: ee CRRA TED pee ae PE D. ogasawarensis Sato, 1981 (Japan) ree eee D. novaeguineae Beier, 1965 (Papua New Guinea) apne Fees {net een cciadt chy Rte oe D. palauensis Beier, 1957 (Palau) Py, USS. eereee elt eee D. elegans Chamberlin, 1929 (Indonesia) Lisette ns seek D. philippinensis Chamberlin, 1929 (Philippines) Eb eh rh SAR ne RE hs EE D. loricata Beier, 1965 (Indonesia) ghee ie ce atte wet De oe ee karen § Fes oan hee Cok ery ens Pg St ee EE 10 Weck angel. Attest Seal D. marcusensis (Morikawa, 1952) (Japan) Od Ort est ere a0 Serre ee ves D. pahangica Beier, 1955 (Malaysia) oceeey Pee ete soo rey D. sumatraensis (Chamberlin, 1923) (Indonesia) Wee cg Boneh ied aries on 28 eee em nk oe D. laosana Beier, 1951 (Laos) feet Meme accel et D. sinuata (Tullgren, 1901) (Cameroon) Nags diesen psaeenn Sica etang tease D. latimana (Beier, 1931) (Tanzania) heath det oS atinRe Ob uid na uae Bs D. tonkinensis Beier, 1951 (Vietnam) ik chk SEARERARNORDET La 5 SEM BAR TS D. shivanparaensis sp. nov. (India) zse.pensoft.net Acknowledgements KHJ thanks Jung-sun Yoo (National Institute for Bio- logical Resources, Korea) for introducing him to Danilo Harms and facilitating his research stay at the Museum of Nature, Hamburg — Zoology. KHJ also extends deep gratitude to his parents, Aekyung Lim and Hyuncheol Jeong for their wholehearted support of his research stay in Hamburg, Germany. JJ thanks the Deutscher Akade- mischer Austauschdienst (DAAD), Germany for funding his research stay in Germany (No. 57440919, 2019/20). The authors are also grateful to Stephanie F. Loria and Nadine Dupérré (both from Hamburg) for their help with distribution mapping and imaging, respectively, and the forest department of the Indian state of Kerala and the National Biodiversity Authority, Govt. of India (No. NBA/Tech Appl/9/Form B-92/19/19-20/676 dat- ed 20.06.2019) for providing necessary permits for the study. Furthermore, the authors also thank the academic editor and the two reviewers, one anonymous and the other Mark Harvey, for their valuable comments and suggestions that have improved this manuscript. References Balzan L (1892) Voyage de M. E. Simon au Venezuela (Décembre 1887—Avril Annales de la Société Entomologique de France 60: 513-552. 1888). Arachnides. Chernetes (Pseudoscorpiones). Beier M (1932) Pseudoscorpionidea I. Subord. Chthoniinea et Neobisiinea. 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