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RESEARCH ARTICLE (Open Access)
Contents Vol 39(2)

New Asian pseudoscorpions improve the phylogenetic resolution of Garypinoidea (Pseudoscorpiones)

Zhizhong Gao A B , Feng Zhang B * and Mark S. Harvey https://orcid.org/0000-0003-1482-0109 C D *
+ Author Affiliations
- Author Affiliations

A Department of Biology, Xinzhou Normal University, Xinzhou, Shanxi, 034000, PR China.

B The Key Laboratory of Invertebrate Systematics and Application, College of Life Sciences, Hebei University, Baoding, Hebei, 071002, PR China.

C Collections & Research, Western Australian Museum, 49 Kew Street, Welshpool, WA 6106, Australia.

D School of Biological Sciences, University of Western Australia, Crawley, WA 6009, Australia.

* Correspondence to: dudu06042001@163.com, mark.harvey@museum.wa.gov.au

Handling Editor: Gonzalo Giribet

Invertebrate Systematics 39, IS24098 https://doi.org/10.1071/IS24098
Submitted: 24 November 2024  Accepted: 3 January 2025  Published: 19 February 2025

© 2025 The Author(s) (or their employer(s)). Published by CSIRO Publishing. This is an open access article distributed under the Creative Commons Attribution-NonCommercial 4.0 International License (CC BY-NC)

Abstract

The recent subdivision of the pseudoscorpion family Garypinidae into three subfamilies, Garypininae, Amblyolpiinae and Protogarypininae, used a combination of molecular and morphological criteria. Newly obtained sequence data from several new garypinoid pseudoscorpions has helped clarify the relationships between various clades. Most importantly, we were able to include the type species of the family, Garypinus dimidiatus (L. Koch, 1873), and two additional species of Amblyolpium Simon, 1898, including A. shenzhou sp. nov. from southern China, which provided a better resolved phylogeny with Amblyolpium as sister to all other Garypinoidea. We raise the subfamily Amblyolpiinae to full family level, Amblyolpiidae stat. nov. In addition, we describe a new genus and species from the Himalayan Plateau, Absensus zhangi sp. nov., which has a morphological feature that allows placement in Amblyolpiidae.

ZooBank: urn:lsid:zoobank.org:pub:4BDE596B-6EFE-4AB8-932E-1D4379849677

Keywords: classification, identification key, molecular phylogeny, morphology, new genus, new species, new status, taxonomy.

Introduction

A recent molecular phylogeny of the pseudoscorpion superfamily Garypinoidea using two nuclear and one mitochondrial gene found evidence for some remarkable divergences yielding surprising results (Harvey 2023). Most notably, the family Garypinidae was not found to be monophyletic as it included the family Larcidae, a result that was also reported by Benavides et al. (2019) albeit with a much smaller sample size but greater gene sampling. Harvey (2023) proposed a novel classification for Garypinidae by recognising three subfamilies that were also supported by morphological data: Amblyolpiinae for Amblyolpium Simon, 1898, Neoamblyolpium Hoff, 1956 and the Eocene Baltamblyolpium Stanczak, Harvey, Harms, Hammel, Kotthoff & Loria, 2023; Protogarypininae for Neominniza Beier, 1930, Oreolpium Benedict & Malcolm 1978, Protogarypinus Beier, 1954, Teratolpium Beier, 1959 and Thaumatolpium Beier, 1931; and Garypininae for the remaining genera, Aldabrinus Chamberlin, 1930, Caecogarypinus Dashdamirov, 2007, Galapagodinus Beier, 1978, Garypinidius Beier, 1955, Garypinus Daday, 1888, Haplogarypinus Beier, 1959, Hemisolinus Beier, 1977, Nelsoninus Beier, 1967, Nobilipinus Harvey, 2023, Pseudogarypinus Beier, 1931, Serianus Chamberlin, 1930, Solinellus Muchmore, 1979, Solinus Chamberlin, 1930, and the Cretaceous Ajkagarypinus Novák, Harvey, Márton, Hammel, Harms, Kotthoff, Hörweg, Brazidec & Ősi, 2023.

We have studied specimens of a garypinid pseudoscorpion from the eastern Himalayan region that, at first glance, resembled a relatively standard garypinid with deeply divided arolia, divided tergites and sternites, and the sub-basal position of the trichobothria of the prolateral series. However, the presence of long venom ducts and the absence of sternal glandular setae posed a problem in finding a satisfactory subfamily placement, as long venom ducts are characteristic of Amblyolpiinae, but not of Garypininae or Protogarypininae, and the glandular setae have been reported for all garypinid taxa except the garypinine Pseudogarypinus (Harvey 2023). Although sequence data are lacking for the sequenced representative of this genus (P. cooperi Muchmore), we added newly obtained sequence data for several important garypinid taxa to the molecular dataset provided by Harvey (2023), including additional species of Amblyolpium from Australia and China, and Garypinus dimidiatus (L. Koch, 1873) from Greece. The inclusion of these taxa has vastly improved the resolution of the phylogenetic inferences that can be made of Garypinoidea. Most importantly, Amblyolpiinae was recovered as sister to the other garypinoids, confirming the importance of the venom duct length in the systematics of garypinoid pseudoscorpions, which allows us to reexamine the rank of this enigmatic clade, and to describe a new genus from China.

Materials and methods

The specimens are preserved in 75% ethanol and deposited in the Museum of Hebei University, Baoding City, China (MHBU). The methods used for the study follow Gao and Zhang (2019).

Terminology and mensuration mostly follow Chamberlin (1931), with the exception of the nomenclature of the pedipalps, legs and with some minor modifications to the terminology of the trichobothria (Harvey 1992), chelicera (Judson 2007) and faces of the appendages (Harvey et al. 2012). The following abbreviations are used: chelal trichobothria: fixed finger, eb, externo-basal trichobothrium; esb, externo-sub-basal trichobothrium; est, externo-subterminal trichobothrium; et, externo-terminal trichobothrium; ib, interno-basal trichobothrium; isb, interno-sub-basal trichobothrium; ist, interno-subterminal trichobothrium; it, interno-terminal trichobothrium; movable finger, b, basal trichobothrium; sb, sub-basal trichobothrium; st, subterminal trichobothrium; t, terminal trichobothrium; cheliceral setae: ebs, externobasal seta; es, external seta; gs, galeal seta; is, interior seta; ls, laminal seta; sbs, sub-basal seta.

Molecular methods

Taxon selection included all of the specimens studied by Harvey (2023) with the addition of two species of Amblyolpium, A. shenzhou sp. nov. from southern China, an undescribed species from South Australia, and Garypinus dimidiatus from Greece (Table 1). The analysis was rooted on Austrochthonius sp. of the superfamily Chthonioidea, and other outgroups included Neopseudogarypus scutellatus Morris, 1948 (Feaelloidea), Hya minuta Tullgren, 1907 and Neobisium carcinoides (Hermann, 1803) (Neobisioidea). For clarity, the phylogram (Fig. 1) omits the chthonioid and feaelloid outgroups.

Table 1.Specimens used in the molecular analysis.

FamilySpeciesLocalityRepository and numberCOI18S rRNA28S rRNA
Superfamily Chthonioidea
 ChthoniidaeAustrochthonius sp.AUSTRALIA: Western Australia: Millstream-Chichester National Park, George River, 2.2 km SE of Mt MontaguWAM T135835OR067318OR039602OR059142
Superfamily Feaelloidea
 PseudogarypidaeNeopseudogarypus scutellatus Morris, 1948AUSTRALIA: Tasmania: Launceston, Cataract GorgeWAM T104213OR067276OR039591OR059173
Superfamily Neobisioidea
 HyidaeHya minuta (Tullgren, 1905)BRUNEI DARUSSALAM: Mengkubau, off Jalan Penghubang mentiriWAM T146845OR067360OR039622OR059105
 NeobisiidaeNeobisium carcinoides (Hermann, 1804)DENMARK: Stampeskov at RaadvadWAM T143210OR067357OR039620OR059108
Superfamily Garypinoidea
 AmblyolpiidaeAmblyolpium sp. ‘PSE215’AUSTRALIA: Western Australia: Paraburdoo Range, ~12.0 km SW of ParaburdooWAM T146503OR359473OR359932OR372787
Amblyolpium sp. ‘PSE216’AUSTRALIA: Western Australia: ~44.4 km SSE of MenziesWAM T152658OR359474OR359933–
Amblyolpium sp. ‘SA’AUSTRALIA: South Australia: Brookfield Conservation ParkWAM T160786PQ834287*PQ834286*PQ834287*
Amblyolpium shenzhou sp. nov.CHINA: Guizhou Province, Tongren City, Jiangkou CountyGZ015PQ814591*PQ810648*PQ810644*
CHINA: Guizhou Province, Tongren City, Jiangkou CountyGZ016PQ814592*PQ810649*PQ810645*
CHINA: Guizhou Province, Tongren City, Jiangkou CountyGZ017PQ814593*PQ810650*PQ810646*
CHINA: Guizhou Province, Tongren City, Jiangkou CountyGZ018PQ814594*PQ810651*PQ810647*
 GarypinidaeAldabrinus rixi Harvey, 2023AUSTRALIA: Western Australia: Torndirrup National Park, Sharp Point at end of Eclipse Island RoadWAM T143175OR359471OR359931OR372788
AUSTRALIA: Western Australia: DenmarkWAM T160040OR359472––
Garypinus dimidiatus (L. Koch, 1873)GREECE: Naxos, 1.7 km ENE of Kastraki BeachWAM T162051PQ834815PQ834284PQ834288
GREECE: Naxos, 1.7 km ENE of Kastraki BeachWAM T162053PQ834816*PQ834285*PQ834289*
Garypinus sp. JA-2011[Unknown]MNHN-JAD71JN018179JN018296JN018393
Nobilipinus karenae Harvey, 2023BRUNEI DARUSSALAM: Ulu Temberong National Park, near Kuala Belalong Field Studies CentreBRUN001OR359476OR359936OR372784
Nobilipinus kohi Harvey, 2023BRUNEI DARUSSALAM: Mengkubau, off Jalan Penghubang mentiriBRUN002OR359477OR359934–
BRUNEI DARUSSALAM: Mengkubau, off Jalan Penghubang mentiriBRUN003OR359478OR359935–
Protogarypinus giganteus Beier, 1954AUSTRALIA: Western Australia: Walpole-Nornalup National Park, The Tingle TreeDNA103127EU559565EU559377EU559484
AUSTRALIA: Western Australia: Fitzgerald River National Park, summit of West Mt BarrenWAM T133897OR359479OR359937–
AUSTRALIA: Western Australia: Kuch RoadWAM T147599OR359480OR359938OR372783
AUSTRALIA: Western Australia: Walpole-Nornalup National Park, Cemetery RoadWAM T159155OR359481OR359939OR372782
Protogarypinus sp. ‘QLD’AUSTRALIA: Queensland: O’Reilly’s, Wishing Tree TrackWAM T145088OR359482OR359940OR372781
Protogarypinus sp. ‘SA’AUSTRALIA: South Australia: Burnside Quarry TrackWAMT157492OR359483OR359941OR372780
Pseudogarypinus cooperi Muchmore, 1980USA: California: Riverside County: James Reserve, Lake FulorDNA102463EU559566EU559423EU559485
Serianus sp.CHILE: La Araucania: Comunidad Indigena QuinquénWAM T152657OR359484OR359942OR372779
Solinus pingrup Harvey, 2023AUSTRALIA: Western Australia: Allanson Primary SchoolWAM T160525OR359490––
Solinus sp. ‘PSE214’AUSTRALIA: Western Australia: 22 km E of Mundrabilla Roadhouse, Eyre HighwayWAM T152660OR359485OR359943OR372777
AUSTRALIA: Western Australia: 24 km SSW of Eucla, Eyre HighwayWAM T152661OR359486OR359944OR372776
Solinus sp. ‘PSE222’AUSTRALIA: Western Australia: Strelley RiverWAM T136665OR359487OR359945OR372778
AUSTRALIA: Western Australia: Strelley RiverWAM T136683OR359488OR359946–
AUSTRALIA: Western Australia: Strelley RiverWAM T136684OR359489OR359947–
 LarcidaeLarca granulata (Banks, 1891)USA: New York: E.N. Huyck Preserve, near Research CenterWAM T143199–OR359930OR372789
Larca lata (Hansen, 1885)CZECHIA: Jihomoravský kraj: Lanzhot, Chanov ReserveDNA102460EU559563EU559425–
Superfamily Garypoidea
 GarypidaeAnchigarypus californicus (Banks, 1909)USA: California: Marin County: Bolinas PointWAM T92262MN058668MN065584MN065610
Anchigarypus japonicus (Beier, 1952)JAPAN: Ehime Prefecture: O-shima Island, 3 km E of MyakuboWAM T140775MN058687MN065603MN065628
Ammogarypus lawrencei Beier, 1962NAMIBIA: Erongo: GobabebWAM T132031MN058676MN065589MN065614
Anagarypus australianus Muchmore, 1982AUSTRALIA: Northern Territory: Bumtja Beach, GarrthalalaWAM T144963MN058692MN065607MN065632
Garypus beauvoisii (Audouin, 1826)CYPRUS: AkanasWAM T143813MN058691MN065606MN065631
Garypus latens Harvey, 2020AUSTRALIA: Western Australia: Barrow IslandWAM T143502MN058690MN065605MN065630
Synsphyronus gracilis Harvey, 1987AUSTRALIA: Western Australia: Mudlark, 92.3 km NW of NewmanWAM T122371MN058669MN065585MN065611
Synsphyronus xynus Cullen & HarveyAUSTRALIA: Western Australia: near Sulfur SpringsWAM T133129MN058679MN065595MN065620
Thaumastogarypus robustus Beier, 1947NAMIBIA: Hardap: Hardap Dam near MarieantalWAM T132035MN058678MN065592MN065617
Thaumastogarypus sp.NAMIBIA: Erongo: Uis (on the road C35)WAM T132030MN058675MN065588MN065613
 GeogarypidaeAfrogarypus deceptor Neethling & Haddad, 2016SOUTH AFRICA: KwaZuluNatal: Ndumo Game ReserveDNA102401EU559560EU559385EU559436
Afrogarypus purcelli (Ellingsen, 1912)SOUTH AFRICA: Western Cape Province: De Hoop Nature ReserveAfro_purcelliKP331817–KP297850
Geogarypus connatus Harvey, 1986AUSTRALIA: South Australia: 14 km WNW of YalataWAM T135409–OR359929OR372785
Geogarypus connatus Harvey, 1986AUSTRALIA: South Australia: 14 km WNW of YalataWAM T135408OR359475–OR372786
Geogarypus longidigitatus (Rainbow, 1897)SINGAPORE: Bukit Timah Nature Reserve, South view pathWAM T146858MN058693MN065608–
Geogarypus minor (L. Koch, 1873)[Unknown]MNHN-JAD10JN018180JN018297JN018394
 HesperolpiidaeApolpium parvum Hoff, 1945TRINIDAD & TOBAGO: Mt Saint BenedictDNA102467EU559541EU559380EU559489
Nanolpium sp.ZAMBIA: Kafue National Park, Chibila CampDNA102410EU559543EU559390EU559445
Pachyolpium sp.TRINIDAD & TOBAGO: Mt Saint BenedictDNA102466EU559542EU559421EU559488
Progarypus sp.CHILE: Concepción: Cerro CaracolDNA102468EU559538EU559420EU559490
 OlpiidaeAntillolpium sp. 772ACUBA: [no further data]sp. 772AKX263395–KX263356
Austrohorus sp.AUSTRALIA: Western Australia: Madura Pass, 800 m NE of MaduraWAM T135388MN058681MN065597MN065622
Beierolpium bornemisszai (Beier, 1966)AUSTRALIA: Western Australia: Gleneagle State ForestDNA102464EU559545EU559378EU559486
Calocheiridius cf. termitophilus Beier, 1964EQUATORIAL GUINEA: Bata District: near Rio Utonde, BataMCZIZ-130511EU559544EU559359EU559460
Euryolpium sp.AUSTRALIA: Western Australia: Steep Head IslandDNA102465MN058685MN065601MN065626
Indolpium sp.AUSTRALIA: Western Australia: 79.1 km SW of Tom PriceWAM T136397MN058683MN065599MN065624

The majority of the data is derived from Harvey (2023), with the sequence data that is newly added for this project marked with an asterisk.

Fig. 1.

Maximum likelihood phylogeny of Garypoidea and Garypinoidea, based on alignment of concatenated COI, 18S and 28S. Bootstrap values are presented for nodes greater than 79%.


IS24098_F1.gif

We included fragments of the cytochrome c oxidase subunit I (COI), 18S ribosomal RNA (18S) and 28S ribosomal RNA (28S), which were sequenced using standard Sanger methodology.

In China, the DNeasy Blood & Tissue Kit (QIAGEN, Hilden, Germany) was used for total genomic DNA extraction. The sequences of primers and polymerase chain reaction (PCR) conditions for three gene regions followed (Folmer et al. 1994; Giribet et al. 1996; Whiting et al. 1997; Murienne et al. 2008). The 25-μL PCR reactions included 12.5 μL of 2×Taq MasterMix (KangWei Biotech, Beijing, China), 0.8 μL of each forward and reverse 10 μM of primer, 4 μL of genomic DNA, and 6.9 μL of double-distilled H2O. The PCR products were visualised by agarose gel electrophoresis (1% agarose). All PCR products were purified and sequenced at Sangon Biotech (Shanghai, China) Co., Ltd. Sequences of the fragment were obtained by using the Chromaseq package in Mesquite (ver. 3.81, W. P. Maddison and D. R. Maddison, see https://www.mesquiteproject.org/) from chromatograms by Phred (ver. 0.020425c, P. Green and B. Ewing, see https://www.phrap.com) and Phrap (ver. 1.090518, P. Green, see https://www.phrap.com). Sequence alignments were carried out using MAFFT (ver. 7.313, see https://mafft.cbrc.jp/alignment/software/; Katoh and Standley 2013) with the L-INS-I strategy, and checked for the presence of stop codons of COI by translating them into amino acid sequence using Geneious Prime (ver. 2024.0.7, Biomatters Ltd, see https://www.geneious.com/, accessed June 2021; Kearse et al. 2012). Ambiguously aligned positions were culled using trimAl (ver. 1.2, see http://trimal.cgenomics.org/; Capella-Gutiérrez et al. 2009) with default parameters.

In Australia, the molecular methods used for the polymerase chain reaction (PCR) amplification of each of these genes followed Harvey et al. (2015, 2016, 2020), Harvey (2023) with PCR purification and Sanger bidirectional sequencing conducted by the Australian Genome Research Facility (AGRF; Perth). Chromatograms were edited using the Geneious software package, and resulting nucleotide sequences for all taxa are deposited in GenBank (Table 1). The sequences were aligned using the MAFFT (ver. 7.490; Katoh et al. 2002; Katoh and Standley 2013) plug-in within Geneious with the default settings. The concatenated alignments were analysed using maximum likelihood (ML) methodology in the web version of IQ-TREE (ver. 2.2.0, see http://www.iqtree.org/; Trifinopoulos et al. 2016; Minh et al. 2020). The substitution model option was set at Auto, and the branch support analysis was performed with 5000 bootstrap alignments.

Results

The phylogeny (Fig. 1) resulting from the molecular analysis recovered a monophyletic Garypinoidea with moderate Bootstrap support (80%), and a monophyletic Garypoidea with poor support (48%). Whereas no differences in the Garypoidea topology were found in comparison to the phylogeny provided by Harvey (2023), there were significant differences in the topology of the garypinoid taxa. Garypinus dimidiatus was found to be the sister group to Garypinus sp. + Nobilipinus karenae + N. kohi (76%). The provenance of the specimen identified as Garypinus sp. was not stated by Arabi et al. (2012) but it seems likely to represent a species of Nobilipinus, a genus that was only recently recognised (Harvey 2023). This clade was sister to Larcidae + Pseudogarypinus + Protogarypinus + Serianus + Aldabrinus + Solinus. The low support value of 44% diminishes the confidence that can be placed on the monophyly of this clade, but internally there is strong support for Aldabrinus + Serianus + Solinus (100%) and a monophyletic Protogarypinus (98%). The four species of Amblyolpium included in the analysis formed a well-supported monophyletic clade (100%), and this clade was in turn sister to the remaining garypinoids but with low support (63%). In the earlier analysis, Amblyolpium and Larca formed a poorly supported clade that was sister to Nobilipinus + Garypinus sp., which was suggested to be due to long-branch attraction (Harvey 2023). The new analysis provides a better resolved phylogenetic scenario, at least in relation to the position of Amblyolpium. Indeed, the significant morphological differences shared by Amblyolpium, Neo-amblyolpium, Absensus and Baltamblyolpium when compared with the remaining garypinids allows us to elevate the subfamily Amblyolpiinae to full family level. Although this partly resolves the conundrum of the non-monophyly of Garypinoidea faced by Harvey (2023), the position of Larca, the sole genus of Larcidae (Harvey and Wynne 2014), still poses significant problems with the family level classification as currently perceived. Likewise, the subfamily Garypininae was not recovered as monophyletic as it also included Larcidae and the garypinid subfamily Protogarypininae (Fig. 1). The low support values suggest that further taxon sampling will provide better resolution to this part of the phylogeny, although it does not fully contradict the phylogeny recovered by Benavides et al. (2019) who used a 210-gene dataset and found Larcidae as sister to Pseudogarypinus, which in turn was sister to Protogarypinus.

Biogeography

The presence of a phylogenetically significant garypinoid pseudoscorpion highlights the importance of the Himalayan Plateau and southern China as a cradle for relictual taxa. The specimens of Amblyolpium shenzhou were collected from three provinces (Guangdong, Hunan and Guizhou) in southern China, and it may have a wide distribution in the Oriental Realm. Absensus zhangi was only found in the Xizang Autonomous region on the Himalayan Plateau, a region that also includes species of genus Centrochthonius Beier, 1931 (family Pseudotyrannochthoniidae), which is known from high-altitude regions of Asia (Harvey and Harms 2022; Kolesnikov et al. 2023), as well as from Eocene Bitterfeld Amber in northern Europe (Schwarze et al. 2022).

Taxonomy

Family AMBLYOLPIIDAE Harvey, 2023, stat. nov.

Amblyolpiinae Harvey, 2023, p. 631.

Diagnosis

The family Amblyolpiidae differs from Garypinidae by the long venom ducts, which are short in Garypinidae, and from Larcidae by the rectangular carapace, which is sub-triangular in Larcidae.

Remarks

As discussed above, the new molecular analysis (Fig. 1) recovered the species of Amblyolpium as sister to the remaining Garypinoidea. Owing to the presence of short femorae on legs I and II, and the positions of the trichobothria, these features must be excluded as diagnostic for Amblyolpiidae.

The inclusion of the Eocene genus Baltamblyolpium in Amblyolpiidae can be justified by the distal position of trichobothrium it that in Amblyolpium and Neoamblyolpium is situated medially (Stanczak et al. 2023). In all other garypinoids, including the Upper Cretaceous genus Ajkagarypinus (Novák et al. 2023), trichobothrium it is situated in a basal position alongside ib, isb and ist.

Key to genera of Amblyolpiidae
1.Trichobothrium it situated in basal half of finger (Fig. 40); trichobothrium est situated much closer to et than esb (Fig. 40); sternites without glandular setae (Fig. 44)...Absensus gen. nov. (China)
Trichobothrium it situated in medial or distal half of finger (Fig. 18); trichobothrium est situated either much closer to esb than et, or midway between esb and et (Fig. 18); sternites VI, VII and sometimes VIII with glandular setae (Fig. 19)...2
2.Trichobothrium st closer to sb than t; trichobothrium it situated subdistally; legs I and II with femur approximately same length as patella; sternites VI–VIII with glandular setae...Baltamblyolpium Stanczak, Harvey, Harms, Hammel, Kotthoff & Loria (Eocene)
Trichobothrium st closer to t than sb (Fig. 18); trichobothrium it situated sub-medially (Fig. 18); legs I and II with femur longer than patella (Fig. 22); sternites VI and VII, but not VIII, with glandular setae (Fig. 19)...3
3.Trichobothrium ib situated on same level as, or slightly anterior to level of, eb and esb (Fig. 19)...Amblyolpium Simon (cosmopolitan)
Trichobothrium ib situated posterior to level of eb and esb...Neoamblyolpium Hoff (western USA)

Genus Amblyolpium Simon, 1898

Type species: Amblyolpium dollfusi Simon, 1898, by monotypy.

Remarks

The genus Amblyolpium is widely distributed around the world, with 16 named species known from Asia, southern Europe, northern Africa and South America (World Pseudoscorpiones Catalog 2024), and several unnamed species from Australia (Harvey 2023).

Amblyolpium shenzhou sp. nov.

(Fig. 2–24.)

ZooBank: urn:lsid:zoobank.org:act:818D25B7-D2A6-410F-9C6A-365E37AEE02F

Material examined

Holotype. CHINA: Guangdong Province: ♂, Meizhou City, Mt Qingliangshan: [24°14′28.46″N, 116°08′22.61″E], elev. 232 m, 10 April 2018, under bark, Xiangbo Guo (Ps.–MHBU–GD18041001).

Paratypes. CHINA: Guangdong Province: 1 ♂, 1 ♀, same data as holotype (Ps.–MHBU–GD18041002 and GD18041003). Hunan Province: 1 ♂, Huaihua City, Yuanling County [28°34′52.25″N, 110°27′28.04″E], elev. 139 m, 9 July 2022, under bark, Yanmeng Hou, Nana Zhan and Jianzhou Sun (Ps.–MHBU–HN22070901). Guizhou Province: 3 ♂, 1 ♀, Tongren City, Jiangkou County [27°49′35.79″N, 108°51′42.32″E], elev. 403 m, 4 July 2022, under bark, Yanmeng Hou, Nana Zhan and Jianzhou Sun (Ps.–MHBU–GZ22070401–03 and GZ22070404).

Etymology

The specific name, shenzhou, is derived from the Chinese mandarin phrase ‘Shénzhōu’ (神舟), referring to the Shenzhou Crewed Spacecraft, a Chinese spacecraft developed for the nation’s crewed space program. Shenzhou-19 manned spacecraft was successfully launched in 2024. The name is a noun in apposition.

Diagnosis

Amblyolpium shenzhou closely resembles A. japonicum, but differs by having stouter pedipalps (patella 2.5× in A. japonicum), and by the different carapaceal chaetotaxy (4 in each row, with a total of 28 setae in A. japonicum) (Morikawa 1960); it differs from A. biaroliatum (Tömösváry, 1884) and A. birmanicum (With, 1906) by the stout pedipalps (femur 4.3, patella 2.9× in A. biaroliatum and A. birmanicum) (Beier 1932); it differs from A. bellum Chamberlin, 1930 in the shape of the marginal teeth on the fixed chelal finger (retrorse pointed teeth in A. shenzhou, normal teeth in A. bellum) and the chaetotaxy of tergites I–III (4: 4: 4–6 v. 4–6–6), also by the slender pedipalps (femur 3.1× in A. bellum) (Chamberlin 1930a, 1930b; Harvey 1988); A. shenzhou shares a similar trichobothrial pattern with A. atropatesi Nassirkhani & Doustaresharaf, 2019 but can be distinguished by the different carapaceal chaetotaxy (4: 6: 4: 4–6: 4 (22–24) in the latter) and the stout chelal patella (2.07–2.14 v. 2.47–2.67×) (Nassirkhani and Doustaresharaf 2019); A. shenzhou resembles A. goldastehae Nassirkhani, Shoushtari & Abadi, 2016, but differs by the different trichobothrial pattern (est at the same level as t in A. goldastehae, distal to t in A. shenzhou) and the stouter pedipalpal patella (2.07–2.14 v. 2.69×) (Nassirkhani et al. 2016).

Description (adult)
Male

Body moderately flattened. Colour with sclerotised portions generally yellowish-brown, pedipalps, carapace and tergites darker.

Chelicera

With 5 setae on hand (Fig. 14), all setae acuminate, ls and is situated adjacent to each other; movable finger with one subdistal seta; subterminal tooth of movable finger not bifurcate and not enlarged; with two lyrifissures on dorsal face; galea with bifurcate tip (Fig. 15); rallum composed of 4 blades, distal blade with serrations on anterior margin, basal blade 1–3 smooth (Fig. 16); serrula exterior with 10–13 blades; lamina exterior present, moderately broad.

Pedipalps (Fig. 5, 6, 17, 18)

All segments completely smooth; setae very long and acicular; trochanter elongate, without small and round tubercles; trochanter 1.73, femur 3.73–3.91, patella 2.07–2.14, chela (with pedicel) 3.57–3.79, chela (without pedicel) 3.39–3.58, hand (with pedicel) 1.70–1.74, hand (without pedicel) 1.48–1.53× longer than broad, movable finger 1.12–1.15× longer than hand with pedicel. Femur without tactile seta on dorso-basal face. Fixed chelal finger with eight trichobothria, movable chelal finger with four trichobothria (Fig. 18): eb, esb and ib situated basally; esb in the midway of eb and isb; isb, ist and it grouped sub-medially; trichobothrium est in the midway of fixed chelal finger, and situated closer to et than to esb; ist slightly distal to isb; et situated near sub-distal end of finger; sb situated closer to b than to st; t situated closer to st, both in the midway of movable chelal finger, away from the tip of movable finger; no microsetae present on either finger. Venom apparatus present in both chelal fingers, venom ducts long and slender, terminating in nodus ramosus near one-third of the way from the tip of the finger. Fixed finger with 30–34 retrorse pointed teeth; movable finger with 21–24 teeth, distal five pointed and straight, the remainder retrorse and obtuse; without accessory teeth.

Carapace (Fig. 4, 13)

Longer than broad (1.37–1.39); sub-rectangular; with two pairs of eyes, each with developed lenses and situated near anterior margin of carapace; with 20 setae, anterior and posterior margin respectively with four setae; without transverse furrows; with four pairs of lyrifissures. Manducatory process with two simple and acute setae; remainder of maxilla with 7–8 setae. Chaetotaxy of coxae I–IV: 5–6: 6–7: 3–4: 5–6.

Abdomen

Pleural membrane longitudinally striate, without setae on membrane. Tergites (some incompletely divided, see Fig. 2) and sternites with narrow medial suture. Tergal chaetotaxy: 4: 4: 4-6: 6: 6: 6: 6–7: 7: 7: 8–9: 4; uniseriate; all setae acicular. Sternal chaetotaxy (IV–XI): 6–8: 8: 8–9: 10: 9–10: 8–9: 7–8: 4; both males and females without patches of chemosensorial setae in the middle of sternites VI–VIII (Fig. 19); setae uniseriate and acuminate; glandular setae absent; anus not surrounded by sternite XI.

Legs (Fig. 10–11, 22–23)

Junction between femora and patellae I and II broad and apparently sub-mobile; patella I distinctly shorter than femur I; femur + patella of leg IV 2.11× longer than broad; metatarsus and tarsus IV see Fig. 12, 24; subterminal tarsal setae arcuate and acute; arolium much longer than claws, distinctly divided into two branches.

Measurements (length/breadth or depth in millimetres, ratios in parentheses)

Male. Body length 1.90–1.99. Pedipalps: trochanter 0.19/0.11 (1.73), femur 0.41–0.43/0.11 (3.73–3.91), patella 0.29–0.30/0.14 (2.07–2.14), chela (with pedicel) 0.72–0.83/0.19–0.23 (3.57–3.79), chela (without pedicel) 0.68–0.78 (3.39–3.58), hand length (with pedicel) 0.33–0.39 (1.70–1.74), hand length (without pedicel) 0.29–34 (1.48–1.53); movable finger length 0.38–0.39. Carapace 0.46–0.48/0.33–0.35. Leg I trochanter 0.08–0.10/0.08 (1.00–1.25), femur 0.28–0.20/0.07 (2.57–2.86), patella 0.10/0.07 (1.43), tibia 0.17/0.05 (3.40), metatarsus 0.07/0.03 (2.33), tarsus 0.12/0.03 (4.00); leg IV trochanter 0.13–0.14/0.09 (1.44–1.56), femur + patella 0.38/0.18 (2.11), tibia 0.25/0.08 (3.13), metatarsus 0.10/0.02 (2.00), tarsus 0.16/0.04 (4.00).

Brief description of females

Mostly same as holotype, Body length 2.06–2.49. Carapace 0.51–0.53/0.37–0.38 (1.38–1.39). Pedipalps: trochanter 0.22–0.24/0.11 (2.00–2.18), femur 0.47–0.48/0.12–0.13 (3.69–3.92), patella 0.34–0.35/0.15 (2.27–2.37), chela (with pedicel) 0.83/0.24 (3.46), chela (without pedicel) 0.78 (3.25), hand length (with pedicel) 0.39–0.40, hand length (without pedicel) 0.34–0.35, movable finger length 0.43–0.44. Leg I femur 0.20–0.21/0.08 (2.50–2.63), patella 0.11/0.08 (1.38), tibia 0.18–0.20/0.05-0.06 (3.33–3.60), metatarsus 0.07–0.08/0.04 (1.75–2.00), tarsus 0.13/0.03 (4.33); leg IV trochanter 0.15/0.10 (1.50), femur + patella 0.42/0.19–0.20 (2.10–2.21), tibia 0.27/0.09–0.10 (2.70–3.00), metatarsus 0.11/0.06 (1.83), tarsus 0.17/0.05 (3.40).

Distribution

Amblyolpium shenzhou has been collected from three provinces (Guizhou, Hunan and Guangdong) in southern China, including the Yunnan–Guizhou Plateau.

Fig. 2–3.

Amblyolpium shenzhou sp. nov., habitus, dorsal: 2, male holotype; 3, female paratype.


IS24098_F2_3.gif
Fig. 4–9.

Amblyolpium shenzhou sp. nov.: male holotype except Fig. 9: 4, carapace, dorsal; 5, left pedipalp, dorsal; 6, left chela, retrolateral; 7, left chelicera, dorsal; 8, genital area, ventral; 9, genital area of female paratype, ventral.


IS24098_F4_9.gif
Fig. 10–12.

Amblyolpium shenzhou sp. nov.: holotype: 10, left leg I, lateral; 11, left leg IV, lateral; 12, left leg IV, metatarsus and tarsus, lateral.


IS24098_F10_12.gif
Fig. 13–24.

Amblyolpium shenzhou sp. nov.: holotype except Fig. 21: 13, carapace, position of lyrifissures indicated by black bars, dorsal; 14, right chelicerae, dorsal; 15, right galea; 16, right rallum; 17, left pedipalp (minus chela), dorsal; 18, left chela, retrolateral; 19, sternites VI-VIII, ventral; 20, genital area, ventral; 21, genital area of female paratype, ventral; 22, left leg I, lateral; 23, left leg IV, lateral; 24, left leg IV, metatarsus and tarsus. Scale bars: 0.10 mm.


IS24098_F13_24.gif

Genus Absensus gen. nov.

ZooBank: urn:lsid:zoobank.org:act:4F082DC6-7B5C-4C74-842C-EBECB04C61C5

Type species: Absensus zhangi sp. nov.

Etymology

The genus name is derived from the Latin word ‘absens’, means absence, referring to the lack of glandular setae on the abdominal sternites. It is to be treated as masculine.

Diagnosis

Absensus can be assigned to Amblyolpiidae due to the presence of long venom ducts (Fig. 40). It differs from the other three genera, Amblyolpium, Neoamblyolpium and Baltamblyolpium, by the lack of glandular setae on the abdominal sternites (Fig. 44), and the basal position of trichobothrium it, which is situated medially in Amblyolpium and Neoamblyolpium and distally in Baltamblyolpium. It further differs from Amblyolpium and Neoamblyolpium by the femorae of the anterior legs being shorter than the patellae (femorae much longer than patellae in Amblyolpium and Neoamblyolpium).

Description
Adult
Setae

Long, straight and acicular.

Chelicera

Hand with 5 long, acuminate setae, ls and is situated adjacent to each other; movable finger with 1 long subdistal seta; rallum of 4 blades, distal blade with several spinules on anterior margin, others smooth; with 2 dorsal and 1 ventral lyrifissures; lamina exterior present; serrula interior modified to form velum; galea of ♀ with 2 long distal rami and a long sub-basal ramus, of ♂ with two distal rami.

Pedipalps

Femur with a single sub-medial tactile seta; patella without disto-prolateral excavation; chelal hand ovate; pedicel prolaterally medially inserted. Fixed chelal finger with 8 trichobothria, movable chelal finger with 4 trichobothria: eb and esb situated at base of fingers; est situated sub-distally, closer to et than to esb; et situated sub-distally; ib, isb, ist and it situated sub-basally, with ib separated from the others; b, sb and st situated sub-basally, with st situated dorso-distally to sb; t situated sub-medially. Venom apparatus present in both chelal fingers, venom ducts long, terminating in nodus ramosus near tip of fingers. Chelal teeth juxtadentate; accessory teeth absent; chelal hand with retrolateral condyle rounded.

Carapace

Sub-rectangular; with 4 large eyes situated close to anterior margin of carapace; epistome absent; without furrows.

Coxal region

Manducatory process with 1–2 long distal setae; maxillary shoulder absent; median maxillary lyrifissure present and situated sub-medially; all coxae approximately same width.

Legs

Femora I and II shorter than patellae I and II; junction between anterior femora and patellae perpendicular; oblique suture line between femora III and IV and patella III and IV; tarsi slightly longer than metatarsi; metatarsi III and IV with a long basal tactile seta; tibia III and IV with a long medial tactile seta; subterminal tarsal setae acuminate; arolium deeply divided and longer than claws; claws slender and simple.

Abdomen

Most tergites and sternites strongly divided. Pleural membrane longitudinally striate, and lacking setae. Sternites lacking glandular setae. Stigmatic sclerites with a single seta; spiracles with spiracular helix. Setae of anterior genital operculum (sternite II) approximately same size as other sternal setae. Posterior tergites and sternites with tactile setae. Anus (tergite XII and sternite XII) without raised rim; anus situated between tergite XI and sternite XI.

Absensus zhangi sp. nov.

(Fig. 25–45.)

ZooBank: urn:lsid:zoobank.org:act:C1C02372-6719-4812-B67B-A9F9E0D30414

Material examined

Holotype. CHINA: Xizang Autonomous Region: ♂, Luozha County [28°12.752′N, 91°00.770′E], elev. 3225 m, 7 August 2014, Chao Zhang (Ps.–MHBU–XZ14080701).

Paratypes. CHINA: Xizang Autonomous Region: 1 ♂, 2 ♀, same data as holotype (Ps.–MHBU–XZ14080702–04).

Etymology

The specific epithet is a patronym in honour of Dr Chao Zhang, who collected the specimens.

Diagnosis

As for the genus Absensus.

Description (adult)
Male

Body moderately flattened. Colour with sclerotised portions generally yellowish-brown, pedipalps, carapace and tergites darker.

Chelicera

With 5 setae on hand (Fig. 36), all setae acuminate, ls and is situated adjacent to each other; movable finger with one subdistal seta; subterminal tooth of movable finger not bifurcate and not enlarged; with two lyrifissures on dorsal face; galea with bifurcate tip and one sub-medial ramus (Fig. 37); rallum composed of 3–4 blades, distal blade with 1–2 basal serrations on anterior margin, basal blade smooth (Fig. 38); serrula exterior with 18–19 blades; lamina exterior present, moderately broad.

Pedipalps

All segments completely smooth; setae very long and acicular; trochanter elongate, without small and round tubercles; trochanter 1.75, femur 2.71–2.88, patella 2.25–2.35, chela (with pedicel) 3.19–3.27, chela (without pedicel) 2.96–3.00, hand (with pedicel) 1.50–1.52, hand (without pedicel) 1.23–1.30× longer than broad, movable finger 1.12–1.15× longer than hand with pedicel. Femur with one tactile seta on dorso-basal face (Fig. 39). Fixed chelal finger with eight trichobothria, movable chelal finger with four trichobothria (Fig. 40): eb, esb and ib situated basally; esb closer to eb than to isb; isb, ist and it grouped sub-basally; trichobothrium est situated closer to et than to esb; ist distal to isb; et situated near sub-distal end of finger; sb situated closer to b than to st; t situated slightly closer to st than to tip of movable finger; no microsetae present on either finger. Venom apparatus present in both chelal fingers, venom ducts long and slender, terminating in nodus ramosus near one-third of the way from the fingertip. Fixed finger with 26–28 pointed teeth; movable finger with 29–30 obtuse teeth; without accessory teeth.

Carapace (Fig. 27, 35)

Longer than broad (1.37–1.54); sub-rectangular; with two pairs of eyes, each with developed lenses and situated near anterior margin of carapace; with 22 setae, anterior and posterior margin respectively with four setae; without transverse furrows; with 6 pairs of lyrifissures. Manducatory process with one long distal, and one long sub-distal setae; remainder of maxilla with eight setae. Chaetotaxy of coxae I–IV: 6: 7: 5: 5.

Abdomen

Pleural membrane longitudinally striate, without setae on membrane. Tergites (some incompletely divided, see Fig. 25) and sternites with narrow medial suture. Tergal chaetotaxy: 3–3: 2–2: 2–2: 2–2: 2–2: 2–2: 2–2: 2–2: 2–2: 5–5: 10; uniseriate; all setae acicular. Sternal chaetotaxy (IV–XI): 1–1: 3–3: 3–3: 3–3: 3–3: 3–3: 4–5: 10; both males and females without patches of chemosensorial setae in the middle of sternites VI–VIII; setae uniseriate and acuminate; glandular setae absent; anus not surrounded by sternite XI.

Legs

Junction between femora and patellae I and II broad and apparently sub-mobile; femur I distinctly shorter than patella I; femur + patella of leg IV 2.61–2.67× longer than broad; metatarsus and tarsus IV see Fig. 43; subterminal tarsal setae arcuate and acute; arolium much longer than claws, distinctly divided into two branches.

Measurements (length/breadth or depth in millimetres, ratios in parentheses)

Measurements (length/breadth or depth in millimetres, ratios in parentheses). Male. Body length 2.61–2.68. Pedipalps: trochanter 0.28/0.16 (1.75), femur 0.46–0.49/0.17 (2.71–2.88), patella 0.45–0.47/0.20 (2.25–2.35), chela (with pedicel) 0.85–0.86/0.26–0.27 (3.19–3.27), chela (without pedicel) 0.78–0.80 (2.96–3.00), hand length (with pedicel) 0.39–0.41 (1.50–1.52), hand length (without pedicel) 0.32–0.35 (1.23–1.30); movable finger length 0.45–0.46. Carapace 0.56-0.60/0.39–0.41. Leg I femur 0.12–0.13/0.10–0.11 (1.18–1.20), patella 0.21–0.22/0.10–0.11 (2.00–2.10), tibia 0.25–0.26/0.06 (4.17–4.33), metatarsus 0.08–0.09/0.04–0.05 (1.80–2.00), tarsus 0.13–0.14/0.04 (3.25–3.50); leg IV femur + patella 0.47–0.48/0.18 (2.61–2.67), tibia 0.34–0.35/0.09–0.10 (3.50–3.78), metatarsus 0.11–0.12/0.06 (3.20–3.40), tarsus 0.16–0.17/0.05 (3.20–3.40).

Brief description of females

Mostly same as holotype, Body length 2.94–3.17. Carapace 0.59–0.64/0.43–0.45 (1.37–1.42). Pedipalps: trochanter 0.31/0.16 (1.94), femur 0.48–0.54/0.18–0.19 (2.67–2.84), patella 0.47–0.53/0.21–0.23 (2.24–2.30), chela (with pedicel) 0.87–0.96/0.27–0.32 (3.00–3.22), chela (without pedicel) 0.81–0.90 (2.81–3.00), hand length (with pedicel) 0.43–0.46, hand length (without pedicel) 0.36–0.38, movable finger length 0.45–0.51. Leg I femur 0.13–0.14/0.12 (1.08–1.17), patella 0.22–0.24/0.11–0.12 (2.00), tibia 0.25–0.28/0.07 (3.58–4.00), metatarsus 0.08–0.09/0.05 (1.60–1.80), tarsus 0.12–0.14/0.04 (3.00–3.50); leg IV trochanter 0.17–0.18/0.09–0.11 (1.64–1.89), femur + patella 0.48–0.52/0.17–0.19 (2.74–2.82), tibia 0.36–0.39/0.09–0.11 (3.55–4.00), metatarsus 0.11–0.12/0.06 (1.83–2.00), tarsus 0.13–0.15/0.05 (2.60–3.00).

Distribution

Specimens of Absensus zhangi have been collected from a single locality in southern China on the Himalayan Plateau.

Fig. 25–26.

Absensus zhangi sp. nov., habitus, dorsal: 25, male holotype; 26, female paratype.


IS24098_F25_26.gif
Fig. 27–34.

Absensus zhangi sp. nov.: holotype except 31: 27, carapace, dorsal; 28, left pedipalp, dorsal; 29, left chela, retrolateral; 30, genital area, ventral; 31, genital area of female paratype, ventral; 32, left leg I, lateral; 33, left leg IV, lateral; 34, left leg IV, metatarsus and tarsus, lateral.


IS24098_F27_34.gif
Fig. 35–45.

Absensus zhangi sp. nov.: holotype: 35, carapace, position of lyrifissures indicated by black bars, dorsal; 36, right chelicerae, dorsal; 37, right galea; 38, right rallum; 39, left pedipalp (minus chela), dorsal; 40, left chela, retrolateral; 41, left leg I, lateral; 42, left leg IV, lateral; 43, left leg IV, metatarsus and tarsus; 44, sternites VI–VIII, ventral; 45, genital area, ventral. Scale bars: 0.25 mm.


IS24098_F35_45.gif

Data availability

All genetic data have been deposited in GenBank.

Conflicts of interest

M. Harvey is an editor for Invertebrate Systematics but did not at any stage have editor-level access to this manuscript while in peer review, as is the standard practice when handling manuscripts submitted by an editor to this journal. The authors have no further conflicts of interest to declare.

Declaration of funding

This work was supported by Fundamental Research Program of Shanxi Province (202403021211040), the Special Project of ‘1331 Project’ to Wutai Mountain Cultural Ecological Collaborative Innovation Center in 2022, and Xinzhou Science and Technology Plan Project (20230214), all awarded to Zhizhong Gao.

Acknowledgements

Z. Gao and F. Zhang thank Prof. Chao Zhang, Yanmeng Hou, Nana Zhan and Jianzhou Sun (College of Life Science, Hebei University) for collecting the specimens. We express our gratitude to Nana Zhan for the assistance in obtaining the molecular data, and to the editor and anonymous reviewers for providing helpful comments on a draft of the manuscript. M. S. Harvey is grateful to Dr Bruno Buzatto (Flinders University) for supplying the specimen of Amblyolpium from South Australia, and the Western Australian Museum’s Molecular Systematics Unit for supplying some of the new sequence data.

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