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Research Article
The pseudoscorpion genus Nipponogarypus (Pseudoscorpiones, Olpiidae) found in seashore habitats in Japan and Korea
expand article infoKyung-Hoon Jeong‡§|, Danilo Harms#¤, Jung-Sun Yoo‡
‡ National Institute of Biological Resources, Incheon, Republic of Korea
§ Seoul National University, Seoul, Republic of Korea
| Jeonbuk National University, Jeonju, Republic of Korea
¶ Museum of Nature Hamburg – Zoology, Leibniz Institute for the Analysis of Biodiversity Change, Hamburg, Germany
# Murdoch University, Murdoch, Australia
¤ Australian Museum Research Institute, Sydney, Australia

Corresponding author: Kyung-Hoon Jeong ( ds16203@snu.ac.kr )

Academic editor: Martin Husemann
© 2024 Kyung-Hoon Jeong, Danilo Harms, Jung-Sun Yoo.
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: Jeong K-H, Harms D, Yoo J-S (2024) The pseudoscorpion genus Nipponogarypus (Pseudoscorpiones, Olpiidae) found in seashore habitats in Japan and Korea. Zoosystematics and Evolution 100(3): 1053-1060. https://doi.org/10.3897/zse.100.123213
ZooBank: urn:lsid:zoobank.org:pub:CF91D7F2-FC99-431C-AD78-6C5CB1DB1E9D
Open Access

Abstract

Some pseudoscorpions (Arachnida: Pseudoscorpions) occur in seashore habitats where they are typically found under driftwood or rocks. Here we review the genus Nipponogarypus Morikawa, 1955 from littoral habitats in Japan and South Korea and describe a new species, Nipponogarypus seosanensis sp. nov., from the Korean Peninsula. We also elevate two former subspecies to species rank: N. enoshimaensis enoshimaensis Morikawa, 1960 = N. enoshimaensis Morikawa, 1955, and N. enoshimaensis okinoerabensis Morikawa, 1960 = N. okinoerabensis Morikawa, 1960, stat. nov. The distribution of all Nipponogarypus species is mapped, and an identification key for the species is provided.

Key Words

False scorpions, morphology, South Korea, systematics, taxonomy

Introduction

Most arachnids are strictly terrestrial, but some lineages occur in seashore habitats or even under water. Some pseudoscorpions (Arachnida, Pseudoscorpiones) have also adapted to littoral or coastal habitats, and these include members of diverse families such as the Garypidae (Harvey et al. 2020), Parahyidae (Harvey et al. 2007), Neobisiidae (e.g., Glynne-Williams and Hobart 1952), and some species in the Olpiidae (Beier 1932; Sato 1994). The family Olpiidae, with its 24 genera and 211 described species (World Pseudoscorpiones Catalog 2022), is most diverse in xeric environments such as semideserts and deserts (Harvey and Leng 2008). However, in 1955, Japanese zoologist Kuniyasu Morikawa discovered several small blackish olpiid pseudoscorpions between rock crevices near the seashore on Enoshima Island (Kanagawa-ken, Honshū) in Japan. These specimens differed from all other Olpiidae genera and were described as a new monotypic genus, Nipponogarypus. In a subsequent publication, Morikawa (1960) split his nominate species, N. enoshimaensis, into two subspecies: N. enoshimaensis enoshimaensis Morikawa, 1955, originally described from Enoshima Island near Honshu, and N. enoshimaensis okinoerabensis Morikawa, 1960, originally described from Okinoerabu-jima in the Satsunan Islands. At least N. enoshimaensis seems to be widespread in supralittoral habitats, and Morikawa (1960) listed additional records from middle and southern Japan, but always in supralittoral habitats (Fig. 1). Nothing else has been published on this interesting genus ever since, and these two subspecies remain valid until today.

Figure 1. 

Distribution of the genus Nipponogarypus (Olpiidae) in Japan and Korea.

In this paper, we establish the first record of Nipponogarypus on the Korean Peninsula and describe a new species, Nipponogarypus seosanensis sp. nov., from South Korea. We also provide adequate illustrations and images for the first time since the original description of this genus, which is now outdated, and diagnose the genus within a modern taxonomic concept for the order Pseudoscorpiones. We also take the liberty to elevate Morikawa’s original subspecies to species status because they are clearly diagnosable under the morphospecies concept. With this step, we recognize three species within Nipponogarypus and clarify the distribution of this genus in eastern Asia.

Materials and methods

All specimens used for this study are deposited in the National Institute for Biological Resources (NIBR) and were collected in Ganwoldo, Seosan-si, Chungcheongnam-Province, South Korea by the primary author. All specimens were preserved in 100% ethanol and examined using a Leica MSV266. Images were taken using a Leica Z16 AP0 attached to a Leica MSV266, and illustrations were created by hand, which were then enhanced using Adobe Illustrator 2023 and Adobe Photoshop 2023 (Adobe Inc.). Scanning electron micrographs were obtained using a Hitachi TM4000Plus scanning electron micrograph (SEM) system. Measurements and terminology follow Chamberlin (1931), Harvey (1992), Judson (2007), and Harvey et al. (2012). The distribution map was created using QGIS 3.22.10 (OGSeo). Abbreviations of chelal trichobothria: b – basal, sb – subbasal, st – subterminal, t – terminal, ib – internal basal, isb – internal subbasal, eb – external basal, esb – external subbasal, it – internal terminal, ist – internal subterminal, et – external terminal, est – external subterminal.

Systematics

Family Olpiidae Banks, 1895

Nipponogarypus Morikawa, 1955

Type species

Nipponogarypus enoshimaensis Morikawa, 1955, by original designation.

Diagnosis

Nipponogarypus can be distinguished from other olpiid genera known to occur in East Asia as follows: from Beierolpium Heurtault, 1977, by trichobothrium st positioned distal to sb in Nipponogarypus and dorsal to sb in Beierolpium (Harvey 1988; Harvey and Leng 2008); from Euryolpium Redikorzev, 1938, by trichobothria it, isb, esb, and eb not clustered in Nipponogarypus but clustered in the latter. Nipponogarypus also shows similar characteristics to Olpium Koch, 1873, and Indolpium Hoff, 1945. However, Nipponogarypus can be distinguished from Indolpium by the position of trichobothria st and isb. In Indolpium, trichobothrium isb is situated proximally to trichobohtorium st (Murthy and Ananthakrishnan 1977). However, in Nipponogarypus, trichobothrium isb is situated distally from st. Furthermore, Nipponogarypus an easily be distinguised from Olpium by the length of its venom ducts. Olpium has long venom ducts that extend to trichobothrium t, whereas Nipponogarypus’ venom ducts only extend to half of trichobothrium t (Mahnert 1991; Nassirkhani 2015). Nipponogarypus is morphologically most similar to Olpiolum Beier, 1931, and both have trichobothrium est positioned in the middle of the fixed finger; ist positioned between est and isb; isb, esb, and eb grouped together; and sb positioned closer to b than st. However, Nipponogarypus differs from Olpiolum by tergal chaetotaxy (four to six setae on the middle tergites in Nipponogarypus, always six setae on the middle tergites in Olpiolum) and the number of pseudotactile seta (two setae present in Nipponogarypus, one seta in Olpiolum) (Muchmore 1986).

Remarks

Subspecies are a rare concept in pseudoscorpion taxonomy since recognizable morphological divergences between populations are usually associated with morphological species. Unfortunately, Morikawa had the habit of designating subspecies (and subgenera) when morphological divergences were seen by him as too minor to warrant species- or genus status for any given taxon (e.g., Morikawa 1960). Pseudoscorpion taxonomy has advanced significantly in the past decades, and we are now aware that minor morphological divergences in cryptic lineages such as pseudoscorpions are generally indicative of species status (e.g., Hlebec et al. 2024; Muster et al. 2024). Following the recent example set by You et al. (2022), who elevated all subterranean subspecies of the genus Spelaeochthonius (family Pseudotyrannochthoniidae) in Japan and Korea to species status, we also elevate Morikawa’s subspecies of Nipponogarypus enoshimaensis to species rank. Unfortunately, this taxonomic act needs to be done without reexamining the primary types that are held at Ehime University but are difficult to access and in poor condition (slide-mounted specimens in dried and contracted Hoyer’s solution; see You et al. 2022). However, Morikawa’s diagnoses are clear and reiterated here: N. enoshimaensis okinoerabensis is elevated to species rank as N. okinoerabensis stat. nov., and this name refers to specimens from the Ryuku and Satsunan Islands that have relatively short body appendages (pedipalpal femur length 0.48 mm, pedipalpal patella length 0.45 mm) and two pseudotacticle hairs that are equal in size on the palpal femur. The subspecies N. enoshimaensis enoshimaensis sensu Morikawa (1960) actually refers to N. enoshimaensis sensu Morikawa (1955) and is here recognized in its original form, N. enoshimaensis. This is a rather widespread morphospecies with records from Honshu and Shikoku (Fig. 1) that has slightly longer body appendages than N. okinoerabensis (pedipalpal femur length 0.52–0.59 mm, pedipalpal patella length 0.57 mm) and unequal pseudotactile hairs (proximal hair smaller than distal hairs). A full description of both species is available in Morikawa (1960). Following this taxonomic act, the genus Nipponogarypus now contains three morphospecies that are found along the coastlines of the Korean Peninsula and Japan.

Nipponogarypus seosanensis Jeong & Harms, sp. nov.

https://zoobank.org/2CFFDE30-7D05-4E5B-9006-E57EE69E780F

Type material

Holotype. Female (NUHGIV0000001225). Korea: Chungcheongnam-province: Ganwoldo 1–gil, Buseok-myeon, Seosan-si, 36°36'5.56"N, 128°10'28.54"E, 03, Jul 2022, KH Jeong leg.

Paratypes : One female and two males (NUHGIV0000001228), same data as holotype.

Etymology

This species is named after the type locality, Seosan-si, Chungcheongnam-do, in South Korea.

Habitat

This species was collected from cracks and fissures of moist rocks in the supralittoral zone, right next to the shoreline (Fig. 2).

Figure 2. 

Habitat of Nipponogarypus seosanensis sp. nov. in the littoral zone of Seosan-si, Chungcheongnam-do, South Korea.

Diagnosis

This species is most similar to N. enoshimaensis by having similar L/W ratios in the pedipalpal femur (0.56–0.61/0.18–0.20 mm in N. seosanensis sp. nov. and 0.52–0.59/0.18 mm in N. enoshimaensis) and pedipalpal patella (0.53–0.59/0.21–0.23 mm in N. seosanensis sp. nov. and 0.57/0.22 mm in N. enoshimaensis). Both species can easily be distinguished by the number of marginal teeth on the fixed chelal finger (40–43 in N. seosanensis sp. nov., 50 in N. enoshimaensis). N. seosanensis sp. nov. can further be distinguished from N. okinoerabensis stat. nov. by having a longer pedipalpal femur (0.56–0.61 mm in N. seosanensis sp. nov. and 0.48 mm in N. okinoerabensis stat. nov.) and patella (0.53–0.59 mm in N. seosanensis sp. nov. and 0.45 mm in N. okinoerabensis stat. nov.).

Description

Female, adult (holotype) (Fig. 3A, B).

Figure 3. 

Habitus of Nipponogarypus seosanensis sp. nov. A. Female holotype, dorsal view; B. Female holotype, ventral view; C. Male paratype, dorsal view; D. Male paratype, ventral view.

Color. Blackish-brown, glossy; the ventral surface darker than the end of the body appendages; the coxal region reddish-orange.

Cephalothorax (Fig. 4A, H). Carapace 1.18 times longer than the broad; carapace sub-rectangular; four conspicuous eyes; two transverse furrows on the carapace; ten lyrifissures; first furrow situated in medial position on the carapace, second furrow near the posterior margin of the carapace; carapacal chaetotaxy 4–2: 22; setae short and acuminate. Pedipalpal coxa with 10 setae; coxal chaetotaxy 5: 5: 6: 13; one lyrifissure on coxa I–III, two lyrifissures on coxa IV.

Figure 4. 

Appendages of Nipponogarypus seosanensis sp. nov., holotype female. A. Carapace; B. Chelicera in dorsal view; C. Left pedipalp in dorsal view; D. Right chela from lateral view; E. Coxa; F. Right leg I; G. Right leg II; H. Female genital sternites; I. Right leg IV; J. Male genital sternites. Scale bars: 0.5 mm (A, C–G, I); 0.2 mm (H, J); 0.1 mm (B).

Chelicera (Figs 4C, 5A). Cheliceral margin smooth; five setae on the cheliceral hand, one seta on the movable finger; galea long and shortly three-branched at the tip in both sexes; serrula exterior with 19 blades, rallum with three blades. Two lyrifissures on the hand.

Pedipalp (Figs 4B, 5B, C). Trochanter 1.60, femur 2.95, patella 2.46, chela 3.24 times longer than the broad, movable finger 0.99 times longer than the hand. Two pseudotactile setae on the femur. Sensory setae present between et and isb. b and sb on the basal of the finger; st on the middle of the t and b. ib, esb, and eb grouped at the base of the finger; est on the middle of et and esb; it and ib on the dorsal surface of the fixed finger, it on the middle of the finger, and ib on the basal of the finger; ist on the paraxial surface of the chela, between isb and est, but closer to isb; short venom ducts in both fingers, not extending past trichobothrium et.

Marginal teeth small and contiguous; triangular teeth on both fingers; fixed finger with 40, movable finger with 43 marginal teeth.

Legs (Figs 4D, E, 5D). Leg I: trochanter 1.56, femur 2.51, patella 1.80, tibia 3.06, metatarsus 2.08, tarsus 2.33 times longer than the broad; leg IV: trochanter 1.40, femur + patella 3.26, tibia 3.93, metatarsus 2.69, tarsus 2.98 times longer than the broad. Typical for the genus: femur and patella of leg I freely mobile; metatarsus of leg IV shorter than the tarsus; metatarsus of leg I longer than the tarsus; one each pseudotactile seta in the distal position of the tarsus; basal position of the metatarsus; arolium undivided, longer than claws, and simple.

Abdomen (Fig. 5F). Pleural membrane striate; all tergites and sternites both undivided; all setae short and acuminate; tergal chaetotaxy 2: 4: 4: 4: 4: 4: 6: 5: 6: 4: 4: 2. Sternal chaetotaxy 10: 4: 6: 8: 6: 4: 6: 6: 4: 4: 2.

Figure 5. 

Appendages of Nipponogarypus seosanensis sp. nov., paratype female. A. Galea; B. Left chela, lateral view; C. Left chelal fingers, lateral view, sensory setae in the white box; D. Right leg I; E. Claws and arolium of right leg I; F. Pleural membrane. Scale bars: 0.02 mm (A); 0.3 mm (B); 0.1 mm (C, D, F); 0.05 mm (E).

Dimensions (in mm). Body length: 1.93 cephalothorax: carapace 0.63/0.54, anterior eye 0.07, posterior eye 0.05; chelicera: total 0.22/0.12, movable finger 0.16; pedipalp: trochanter 0.31/0.19, femur 0.59/0.20, patella 0.58/0.23, chela 1.08/0.33, movable finger 0.53, hand 0.54; leg I: trochanter 0.15/0.10, femur 0.23/0.09, patella 0.18/0.10, tibia 0.21/0.07, metatarsus 0.10/0.05, tarsus 0.11/0.05; leg IV: trochanter 0.16/0.11, femur + patella 0.55/0.17, tibia 0.38/0.10, metatarsus 0.17/0.06, tarsus 0.16/0.05.

Variation (one female, paratype)

Cephalothorax. Carapace 1.16 times longer than broad.

Pedipalp. Trochanter 1.65, femur 3.13, patella 2.64, chela 3.29 times longer than broad, movable finger 0.88 times longer than hand. Fixed finger with 42, movable finger with 43 marginal teeth.

Legs. Leg I: trochanter 1.28, femur 3.29, tibia 3.34, metatarsus 2.38, tarsus 2.89 times longer than broad; leg IV: trochanter 1.50, femur + patella 3.06, tibia 3.08, metatarsus 2.25, tarsus 2.82 times longer than broad.

Abdomen. Tergal chaetotaxy 2: 4: 4: 4: 4: 6: 6: 6: 4: 2: 2. Sternal chaetotaxy 8: 4: 6: 6: 6: 6: 6: 4: 4: 4: 2.

Dimensions (in mm). Body length 1.90; cephalothorax: carapace 0.58/0.50, anterior eye 0.06, posterior eye 0.05; chelicera: total 0.20/0.12, movable finger 0.16; pedipalp: trochanter 0.33/0.20, femur 0.61/0.20, patella 0.59/0.22, chela 1.04/0.32, movable finger 0.49, hand 0.56; leg I: trochanter 0.14/0.11, femur 0.24/0.10, patella 0.19/0.10, tibia 0.25/0.08, metatarsus 0.12/0.05, tarsus 0.13/0.05; leg IV: trochanter 0.19/0.13, femur + patella 0.60/0.20, tibia 0.34/0.11, metatarsus 0.15/0.07, tarsus 0.18/0.06.

Variation (two males, paratype)

Cephalothorax. Carapace 1.20–1.23 times longer than broad.

Pedipalp. Trochanter 1.34–1.59, femur 3.07–3.13, patella 2.6, chela 3.05–3.21 times longer than the broad, movable finger 0.81–0.89 times longer than the hand. Fixed finger with 40–41, movable finger with 41–43 marginal teeth.

Legs. Leg I: trochanter 1.20–1.67, femur 2.08–2.45, patella 1.66–1.70, tibia 2.67–3.22, metatarsus 2.62–2.67, tarsus 2.86–3 times longer than broad; leg IV: trochanter 1.25–1.39, femur + patella 2.67–2.74, tibia 3.54–3.63, metatarsus 2.77–3.16, tarsus 2.5–2.7 times longer than broad.

Abdomen. Tergal chaetotaxy 2: 4: 4: 4: 4: 6: 6: 6: 6: 4: 4: 2. Sternal chaetotaxy 11: 6: 4: 6: 6: 6: 6: 5: 4: 4: 2.

Male genital sternites (Fig. 4G). Nine lyrifissures on sternites II and III; 11 setae on the genital opening area; six setae on the sternite III.

Dimensions (in mm). Body length: 1.83–1.84; cephalothorax: carapace 0.61/0.50–0.51, anterior eye 0.06, posterior eye: 0.04–0.05; chelicera: total 0.20/0.10–0.11 movable finger 0.14; pedipalp: trochanter 0.25–0.32/0.19–0.20, femur 0.56–0.60/0.18–0.19, patella 0.53–0.57/0.21–0.22, chela 0.91–0.92/0.21–0.29, movable finger 0.41–0.43, z 0.49–0.50; leg I: trochanter 0.12/0.07–0.10, femur 0.18–0.24/0.09–0.10, patella 0.15–0.16/0.09–0.10, tibia 0.17–0.23/0.06–0.07, metatarsus 0.12–0.13/0.05, tarsus 0.13/0.04; leg IV: trochanter 0.17–0.18/0.13, femur + patella 0.51/0.19, tibia 0.37/0.10, metatarsus 0.15–0.16/0.05, tarsus 0.12–0.16/0.05–0.06.

Key to the species of Nipponogarypus

1 Length of pedipalpal femur and patella more than 0.5 mm 2
– Length of pedipalpal femur and patella less than 0.5 mm Nipponogarypus okinoerabensis stat. nov.
2 Fixed chelal finger with more than 45 marginal teeth N. enoshimaensis
– Fixed chelal finger with less than 45 marginal teeth N. seosanensis sp. nov.

Acknowledgements

KHJ would like to thank Mark Harvey (Western Australian Museum) for sending valuable reference papers and Sora Kim (Jeonbuk National University) for accessing the microscope and camera systems at Jeonbuk National University. This work has been supported by a grant from the National Institute of Biological Resources (NIBR), funded by the Ministry of Environment (MOE) of the Republic of Korea (NIBR202304103).

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