2 urn:lsid:arphahub.com:pub:45048D35-BB1D-5CE8-9668-537E44BD4C7E urn:lsid:zoobank.org:pub:91BD42D4-90F1-4B45-9350-EEF175B1727A ZooKeys ZK 1313-2989 1313-2970 Pensoft Publishers 10.3897/zookeys.911.38761 38761 Research Article Araneae Taxonomy Asia Asianopis gen. nov., a new genus of the spider family Deinopidae from Asia Lin Yejie https://orcid.org/0000-0002-6789-2731 1 Shao Lili 2 Hänggi Ambros 3 Caleb John T.D. https://orcid.org/0000-0002-9471-9467 4 Koh Joseph K.H. 5 Jäger Peter 6 Li Shuqiang lisq@ioz.ac.cn https://orcid.org/0000-0002-3290-5416 2 Hebei Key Laboratory of Animal Diversity, College of Life Science, Langfang Normal University, Langfang 065000, China Langfang Normal University Langfang China Institute of Zoology, Chinese Academy of Sciences, Beijing 100101, China Institute of Zoology, Chinese Academy of Sciences Beijing China Naturhistorisches Museum Basel, Augustinergasse 2, CH 4001 Basel, Switzerland Naturhistorisches Museum Basel Basel Switzerland Zoological Survey of India, Prani Vigyan Bhawan, M-Block, New Alipore, Kolkata 700053, West Bengal, India Zoological Survey of India Kolkata India National Biodiversity Centre, National Parks Board, 259598, Singapore National Biodi­versity Centre, National Parks Board Singapore Singapore Senckenberg Research Institute, Senckenberganlage 25, 60325 Frankfurt a. M., Germany Senckenberg Research Institute Frankfurt a. M. Germany

Corresponding author: Shuqiang Li (lisq@ioz.ac.cn)

Academic editor: Gergin Blagoev

 2020 12 02 2020 911 67 99 89C3F88E-7A03-5C24-A86E-3FA7EB401398 12B393ED-9CB7-4642-9127-B13BA1952BD3 3676178 02 08 2019 09 01 2020 Yejie Lin, Lili Shao, Ambros Hänggi, John T.D. Caleb, Joseph K.H. Koh, Peter Jäger, Shuqiang Li 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. http://zoobank.org/12B393ED-9CB7-4642-9127-B13BA1952BD3

A new genus of the spider family Deinopidae C.L. Koch, 1850 is described from Asia: Asianopis Lin & Li gen. nov., with A. zhuanghaoyuni Lin & Li sp. nov. as the type species. The new genus is divided into two species groups, of which the liukuensis-group includes two species: A. dumogae (Merian, 1911) sp. reval. comb. nov. (♀) and A. liukuensis (Yin, Griswold & Yan, 2002) comb. nov. (♂♀); and the zhuanghaoyuni-group comprises five species: A. celebensis (Merian, 1911) comb. nov. (♂), A. konplong (Logunov, 2018) comb. nov. (♂), A. wangi Lin & Li sp. nov. (♂♀), A. wuchaoi Lin & Li sp. nov. (♂♀), and A. zhuanghaoyuni Lin & Li sp. nov. All previously described species are transferred from Deinopis MacLeay, 1839. Deinopis scrubjunglei Caleb & Mathai, 2014 is treated as a junior synonym of Asianopis liukuensiscomb. nov.

 New combination new species species groups systematics taxonomy Citation

Lin Y, Shao L, Hänggi A, Caleb JTD, Koh JKH, Jäger P, Li S (2020) Asianopis gen. nov., a new genus of the spider family Deinopidae from Asia. ZooKeys 911: 67–99. https://doi.org/10.3897/zookeys.911.38761

 Introduction

The spider family Deinopidae C.L. Koch, 1850 (Araneae, Deinopoidea), known as net-casting or ogre-faced spiders, is a small family that consisted of two genera and 64 species prior to the current study (World Spider Catalog 2019). The genus Deinopis was established by MacLeay (1839) based on Deinopis lamia MacLeay, 1839 (♂♀) from Cuba. The other genus, Menneus, was established by Simon (1876) based on Menneus tetragnathoides Simon, 1876 (♂) from Angola.

Ten species of Deinopidae were known from Asia: Deinopis aruensis Roewer, 1938 (♀) and D. celebensis Merian, 1911 from Indonesia; D. fasciculigera Simon, 1909 (♀) and D. konplong Logunov, 2018 (♂) from Vietnam; D. scrubjunglei Caleb & Mathai, 2014 (♂♀) from India; D. gubatmakiling Barrion-Dupo & Barrion, 2018 (juvenile), D. labangan Barrion-Dupo & Barrion, 2018 (♀), and D. luzonensis Barrion-Dupo & Barrion, 2018 (♀) from the Philippines; D. kollari Doleschall, 1859 (♂) from Myanmar and Malaysia; D. liukuensis Yin, Griswold & Yan, 2002 (♂♀) from China. Here, we describe a new genus and three new species, and present a molecular phylogenetic analysis of these spiders.

 Material and methods

All specimens were preserved in 80% ethanol. Metatarsi and tarsi were removed for preservation in 100% ethanol for subsequent molecular work. Epigynes were cleared in proteinase K at 56 °C to dissolve non-chitinous tissues for three hours. Specimens were examined under a LEICA M205C stereomicroscope. Photomicroscope images were taken with an Olympus C7070 zoom digital camera (7.1 megapixels). Laboratory habitus photographs were taken with a Canon 5D Mark III digital camera equipped with a Canon MP-E 65 mm lens. Photos were stacked with Helicon Focus (version 6.7.1) or Zerene Stacker (version 1.04) and processed in Adobe Photoshop CC 2018. Photographs of Asianopis celebensis comb. nov. were taken by a KEYENCE. Photographs of Asianopis liukuensis comb. nov. from India (i.e., the type materials of D. scrubjunglei) were taken using a Leica DFC500 HD camera mounted on a Leica M205A stereomicroscope.

All measurements are in millimetres. Eye sizes are measured as the maximum diameter from either the dorsal or frontal view. Leg measurements are given as follows: total length (femur, patella+tibia, metatarsus, tarsus). Copulatory duct turns are defined by the number of apparent loops on the lateral margin of the copulatory/fertilization duct complex in dorsal view. The length of the embolic tip fold is measured as from the beginning of the fold to the embolic tip (Fig. 22D, E). The terminology used in the text and figures follows Coddington et al. (2012). Distribution maps were generated using ArcMap software (version 10.2).

A total of 31 specimens of Deinopidae were collected for phylogenetic analysis (Suppl. material 1: Table S1). Sequences of seven specimens were from the National Center for Biotechnology Information (NCBI) public data, and the other 24 were from recent field collections. Whole genomic DNA was extracted from 2–4 legs using a TIANamp Genomic DNA kit (TIANGEN Inc., Beijing, China) following the manufacturer’s protocol. Seven gene fragments were amplified in 20-μL reactions: COI (~640 bp), 12S (~330 bp), 16S (~470 bp), 18S (~1700 bp), 28S (~1200 bp), H3 (~310 bp) and wnt (~330 bp). Primers and PCR conditions for each locus are listed in Suppl. material 1: Table S2. Sequence chromatograms were proofed and edited using Sequencher version 4.2 Demo (Gene Codes Corporation, Ann Arbor, MI USA). The COI, H3 and wnt fragments were translated in MEGA version 7 (Kumar et al. 2016) to check for the presence of stop codons. A representative of the family Uloboridae was used as the outgroup, with the corresponding sequences downloaded from NCBI. The complete list of 32 taxa and GenBank accession numbers are provided in Suppl. material 1: Table S1.

Multiple sequence alignments were carried out with MAFFT version 7.243 (Katoh and Standley 2013). Alignments of the protein-coding COI, H3 and wnt genes were produced using the L-INS-i method. As for the highly variable ribosomal genes, the E-INS-i method was used to generate alignments of 12S, 16S, 18S, and 28S. To exclude the ambiguously aligned regions, alignments of the ribosomal genes were processed with the program trimAl version 1.3 (Capella-Gutiérrez et al. 2009). The alignments are shown in the supplementary data.

The concatenated gene matrix was partitioned by gene using PartitionFinder version 1.1.1 (Lanfear et al. 2012). The best partitioning scheme was selected based on the Akaike information criterion (AIC) (Suppl. material 1: Table S3). Maximum likelihood (ML) analysis was performed using RAxML version 8.2.9 with a GTR + Γ + I model applied to each partition (Stamatakis 2014). One thousand non-parametric bootstrap replicates were conducted to obtain the best-scoring ML tree.

Bayesian analysis was performed using MrBayes version 3.2.6 (Ronquist et al. 2012). Two independent runs, each with four independent chains, were carried out for 20,000,000 generations and were sampled every 1,000 generations with a burn-in of 25%. Partitions and models followed the result of PartitionFinder. Convergence of the runs was determined with the standard deviation of split frequencies (<0.01). Effective sampling sizes (>200) of all parameters were checked in Tracer version 1.6 (Rambaut et al. 2014). A 50% majority-rule consensus tree was then constructed from the post-burnin sampled trees to estimate posterior probabilities (PP).

Abbreviations

ALE anterior lateral eye

AME anterior median eye

CD copulatory duct

CO copulatory opening

E embolus

EMA embolic middle apophysis

EO embolic opening

ETA embolic terminal apophysis

FD fertilization duct

MA median apophysis

MABL median apophysis–basal lobe

MADL median apophysis–distal lobe

MP median plate

PLE posterior lateral eye

PME posterior median eye

S spermatheca

SD sperm duct

SpD spermathecal duct

T tegulum.

 Museum abbreviations

HNU Hunan Normal University, Changsha, China

IZCAS Institute of Zoology, Chinese Academy of Sciences, Beijing, China

MMUE Manchester Museum of the University of Manchester, UK

NMB Naturhistorisches Museum Basel, Basel, Switzerland

SRC-ZSI Southern Regional Centre, Zoological Survey of India, Kolkata, India

Taxonomy Family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Deinopidae</tp:taxon-name-part></tp:taxon-name> C.L. Koch, 1850 Animalia Araneae Deinopidae 2DF06D83-9CC6-5EF2-90BE-9E565F1FE4B5 Asianopis http://zoobank.org/C8CA3BB7-776C-4BB9-9E19-F819587E87AB Lin & Li gen. nov. Type species.

Asianopis zhuanghaoyuni Lin & Li, sp. nov.

 Etymology.

The generic name is a combination of the word “Asia”, referring to the distribution of the genus, and the generic name Deinopis. The gender is feminine.

 Diagnosis.

Asianopis gen. nov. can be easily distinguished from Deinopis by the following characters: a prominent setal fringe can be found above the posterior median eyes in both sexes of Asianopis species (Fig. 4A, B), which is absent in Deinopis (Coddington et al. 2012: fig. 3a); the embolic tip of male Asianopis has an embolic middle apophysis (liukuensis-group, Fig. 21A), an embolic terminal apophysis or is weakly folded apically (zhuanghaoyuni-group, Fig. 21B–E), whereas none of these characters is present in Deinopis (Coddington et al. 2012: fig. 11m); the MADL in Asianopis is small and has a basal lobe, while in Deinopis, the median apophysis is larger than the MABL and covers the entire base (Coddington et al. 2012: fig. 11m); female chelicerae with many denticles between the promarginal and retromarginal teeth (Fig. 2F) or female chelicerae without denticles (Fig. 2H), in contrast, denticles are only at the center of any two adjoining retromarginal teeth in Deinopis (Coddington et al. 2012: fig. 5c); femora I enlarged proximally in Asianopis gen. nov. (liukuensis group, Fig. 2I) or not enlarged (zhuanghaoyuni-group, Fig. 2J), but they are enlarged distally in Deinopis (Coddington et al. 2012: fig. 3b); epigynal median plate lateral margins anchor-shaped in Asianopis gen. nov. (Figs 3A, 6A), but ellipsoid in Deinopis (Coddington et al. 2012: fig. 9b); SpD is consistently narrow in Asianopis gen. nov. (Figs 3B, 6B) but tapering in Deinopis (Coddington et al. 2012: fig. 9d).

10.3897/zookeys.911.38761.figure1 3676180 6310C386-12F3-52AC-80DC-C94A582DE604

Phylogenetic tree of Deinopidae spiders based on 31 specimens. Numbers on nodes indicate Maximum Likelihood bootstrap values and Bayesian posterior probabilities.

https://binary.pensoft.net/fig/380013 10.3897/zookeys.911.38761.figure2 3676204 FE228848-15E5-5997-93CA-A24347A452EB

Prosoma (frontal view, upper ♂, lower ♀) (A–D), chelicerae (E–H) and leg I (I–J). Figures A and C modified from Coddington et al. (2012). ADeinopis spinosaBAsianopis liukuensis comb. nov. CMenneus dromedariusDAsianopis zhuanghaoyuni sp. nov. E Chelicerae of male A. liukuensis comb. nov. F Chelicerae of female A. liukuensis comb. nov. (Arrows indicate the denticles) G Chelicerae of male A. zhuanghaoyuni sp. nov. H Chelicerae of female A. zhuanghaoyuni sp. nov. I Left leg I of female A. liukuensis comb. nov. Arrow shows enlarged femur J Left leg I of female A. zhuanghaoyuni sp. nov.

https://binary.pensoft.net/fig/380014 Description.

Male. Total length 12.14–16.10 (n = 8), carapace pear-shaped, yellow-brown (liukuensis-group) or brown (zhuanghaoyuni-group) with white edge, white line extending from cephalic area to posterior margin and small spines sparsely distributed; fovea longitudinal, indistinct. Chelicerae with a promarginal tooth and one or two retromarginal teeth (liukuensis-group) or with four promarginal teeth and 2–6 retromarginal teeth (zhuanghaoyuni-group), no denticles. Endites and labium brown, distally white; sternum diamond-shaped, brown with median light band and few small spines. Legs brown, ventrally with black pattern and short spines, leg formula 1243. Opisthosoma cylindrical, brown or dark-brown with small black spots and irregular pattern. Cribellum entire, spinnerets brown (Figs 4, 10, 13, 16).

Female. Total length 14–24 (n = 13). Chelicerae with four promarginal teeth and seven retromarginal teeth, many denticles in between the promarginal and retromarginal teeth (liukuensis-group) or four promarginal teeth and 8–13 retromarginal teeth, without denticles (zhuanghaoyuni-group). Appearance of carapace, opisthosoma and legs as in male but femora of legs I enlarged basally (liukuensis-group) (Fig. 2I).

Male palpal tibia longer than cymbium; cymbium almost round; tegulum distinctly wider than the diameter of embolic coil (liukuensis-group) or tegulum obscured by embolic coil (zhuanghaoyuni-group) (Figs 17, 18); embolus long and strongly coiled around MA, embolic base beginning at 7–8 o’clock position, coiled 1200° (liukuensis-group) or more than 1500° (zhuanghaoyuni-group), embolic tip straight (liukuensis-group) or widened subapically, folded and without apophysis (zhuanghaoyuni-group); MA small, directed at 7–8 o’clock position, with two lobes, a small lobe at the base, and a narrow distal lobe with two apophyses (liukuensis-group) or large, with two lobes, a large lobe at the base and a kidney-shaped distal lobe (zhuanghaoyuni-group).

Epigyne with anchor-shaped median plate, CO distinct, CD with three turns, S oval, SpD consistently wide (liukuensis-group) or with a well-developed MP, obscuring CO, CD with 7–8 turns, S oval, SpD consistently thin (zhuanghaoyuni-group).

 Molecular phylogeny.

The molecular phylogenetic analysis indicates with strong support that all the species in this study do not belong to Deinopis. Based on the 4893 bp-aligned sequences of seven gene fragments, the ML and Bayesian analyses produced the same topology, showing a split of a Southwest China clade from other clades and is strongly supported (Bootstrap value: 88; PP: 0.98) (Fig. 1). Our results are consistent with the results of Chamberland et al. (2018) who conducted a global phylogenetic analysis of Deinopis. Therefore, the Southwest China clade can be classified as a new genus with strong support (Bootstrap value: 100; PP: 1). Although intraspecific support values are low in both ML and Bayesian analyses results, basal nodes are strongly supported, including the sister relationship of A. wangi Lin & Li, sp. nov. & A. zhuanghaoyuni Lin & Li, sp. nov. (Bootstrap value: 95; PP: 1).

 Natural habitat.

All the species of Asianopis gen. nov. were collected from bushes in low-elevation forests.

 Composition.

This new genus comprises two species groups: the liukuensis-group with two species: A. dumogae (Merian, 1911) sp. reval. comb. nov. and A. liukuensis (Yin, Griswold & Yan, 2002) comb. nov. and the zhuanghaoyuni-group with five species: A. celebensis (Merian, 1911) comb. nov., A. konplong (Logunov, 2018) comb. nov., A. wangi sp. nov., A. wuchaoi sp. nov., and A. zhuanghaoyuni sp. nov.

 Distribution.

China (Fujian, Yunnan, Hong Kong, Guangxi, Hainan), India, Indonesia, and Vietnam.

 The <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Asianopis"/><tp:taxon-name-part taxon-name-part-type="species" reg="liukuensis">liukuensis</tp:taxon-name-part></tp:taxon-name></italic>-group Animalia Araneae Deinopidae AB4BA7F0-0635-5CC3-BBA6-9EAC7114A56E Asianopis dumogae (Merian, 1911), sp. reval. comb. nov. Fig. 3 Dinopis dumogae Merian, 1911: 171 (♀ only, ♂ mismatched). Type material examined.

1♀ (NMB-ARAN-00514a), “Wald bei Duluduo”, Sulawesi Utara, forest near Duluduo, 00°31'33"N, 123°57'10"E, Sulawesi, Indonesia.

 Diagnosis.

This species can be distinguished from A. liukuensis comb. nov. by the MP nearly covering the CO, S round, and the overall equal thickness of the CD (Figs 4, 6).

10.3897/zookeys.911.38761.figure3 3676214 F8B20ED6-F4D1-5AAA-91AD-44FC64E4AFE0

Asianopis dumogae sp. reval. comb. nov., female type. A Epigyne B Vulva, dorsal view.

https://binary.pensoft.net/fig/380015 10.3897/zookeys.911.38761.figure4 3676216 C5B9A284-755D-5688-A503-5310066607BC

Asianopis liukuensis comb. nov., male from Xishuangbanna and female from Jianfengling. A Male prosoma, frontal view B Female prosoma, frontal view C Male habitus, dorsal view D Male habitus, ventral view E Female habitus, dorsal view F Female habitus, ventral view.

https://binary.pensoft.net/fig/380016 10.3897/zookeys.911.38761.figure5 3676218 208CC294-D3F6-5841-B6DC-8FBFC62384F7

Asianopis liukuensis comb. nov., left palp, male from Xishuangbanna. A Prolateral view B Retrolateral view.

https://binary.pensoft.net/fig/380017 Description.

See Merian (1911). Photos of the epigyne of the syntype are given in Figure 6.

10.3897/zookeys.911.38761.figure6 3676220 C8F5717E-3D9F-5BC8-A01B-CC1B2DB948BB

Asianopis liukuensis comb. nov., female from Jianfengling. A Epigyne B Vulva, dorsal view.

https://binary.pensoft.net/fig/380018 Distribution.

Indonesia (North Sulawesi).

 Comments.

Merian (1911) reported D. celebensis based on three specimens from different localities in Sulawesi, Indonesia. One male (NMB-ARAN-00514b, “Zentral-Celebes, nördlich vom Golf von Bone”, South Sulawesi, north of the Gulf of Boni (precise locality not known), one female from North Sulawesi (NMB-ARAN-00514a, “Wald bei Duluduo”, Sulawesi Utara, forest near Duluduo, 00°31'33"N, 123°57'10"E and one female from Central Sulawesi (NMB-ARAN-00514c, Larga, südlich vom Posso-See, unterhalb Patiro Rano, bei 900 m, Central Sulawesi, south of Lake Poso at an elevation of 900 m (the localities “Larga” and “Patiro Rano” could not be located on maps; the epigyne of this specimen is missing, but the specimen is clearly larger than the others).

Merian (1911) stated that the male and the females may not represent the same species and suggested the name D. celebensis for the male, and D. dumogae for the female. According to the International Code of Zoological Nomenclature (International Commission on Zoological Nomenclature 1999: Article 11.5.1), such conditionally proposed species names are potentially available as valid names if published before 1961. The species has not been listed in any of the catalogues. We examined the types and concluded the male and the two females are indeed three different species. The palp of the male D. celebensis exhibits features of the zhuanghaoyuni group: the tegulum is obscured by the embolic coil, and the embolus is long and strongly coiled around the MA. The female from North Sulawesi (Doloduo) has features of the liukuensis group: an anchor-shaped median plate, CO distinct, CD with three turns. Thus, we revalidated the female D. dumogae as Asianopis dumogae (Merian, 1911), sp. reval. comb. nov.

 Animalia Araneae Deinopidae 0B61E438-BC43-5540-8A8D-04906837BEB3 Asianopis liukuensis (Yin, Griswold & Yan, 2002) comb. nov. Figs 2B, E, F, I , 4 , 5 , 6 , 7 , 8 , 19 , 21A , 22A, G , 23 Deinopis liukuensis Yin et al., 2002: 610, figs 1–7 (♂♀) Deinopis liukuensis Zhang & Wang, 2017: 238 (♂♀) Deinopis scrubjunglei Caleb & Mathai, 2014: 2, figs 1–20 (♂♀) syn. nov. Type.

Holotype. ♂ (HNU, no. 00-LK-1, lost), China, Yunnan Province, Liuku, Mt Gaoligong, 25°30'48"N, 98°30'36"E, elevation ca 800 m, 26.VI.2000, Heng-Mei Yan leg.

 Type materials of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Deinopis">Deinopis</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="scrubjunglei">scrubjunglei</tp:taxon-name-part></tp:taxon-name></italic> examined.

♂ (SRC-ZSI I/SP 19), Madras Christian College, Chennai, Tamil Nadu, 12°55'12.7"N, 80°07'24.6"E, elevation ca 32 m, 5.XII.2013, John Caleb T.D. leg.; ♀ (SRC-ZSI I/SP 20), 22.IV.2014, same location, John Caleb T.D and Karthy leg.

 Other material examined.

2♂, China, Yunnan Province, Xishuangbanna Dai Autonomous Region, rubber tree plantation near Jinghong City, 28.IV.2016, Chaotai Wei leg.; 1♀, China, Hainan Island, Ledong County, Jianfengling National Park, 13.VII.2019, Zixuan Lin leg.

 Diagnosis.

This species can be distinguished from other congeners by the distinct female copulatory opening, oval S, and CD tapering from the copulatory opening to spermatheca (Figs 6, 8).

 Description.

See Yin et al. (2002) and Caleb and Mathai (2014).

 Distribution.

China (Yunnan, Guangxi, Hainan), India.

 Comments.

Type materials of D. scrubjunglei syn. nov. were examined and no differences between A. liukuensis and D. scrubjunglei were observed. Thus, we consider D. scrubjunglei to be a synonym of A. liukuensis, and the figures of D. scrubjunglei are given for comparison (Figs 7, 8, 19C).

10.3897/zookeys.911.38761.figure7 3676222 FE3ADF08-3DBC-5D97-9F23-B6FF6EFC054B

Asianopis liukuensis comb. nov., left palp, holotype male of Deinopis scrubjunglei syn. nov. A Prolateral view B Retrolateral view.

https://binary.pensoft.net/fig/380019 10.3897/zookeys.911.38761.figure8 3676224 82F1DF54-580B-56EE-B1E4-C6661719BBCA

Asianopis liukuensis comb. nov., paratype female of Deinopis scrubjunglei syn. nov. A Epigyne B Vulva, dorsal view.

https://binary.pensoft.net/fig/380020 The <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Asianopis"/><tp:taxon-name-part taxon-name-part-type="species" reg="zhuanghaoyuni">zhuanghaoyuni</tp:taxon-name-part></tp:taxon-name></italic>-group Animalia Araneae Deinopidae 08ADBAE9-C3A2-5D4F-A2FF-133634EEA3BD Asianopis celebensis (Merian, 1911) comb. nov. Fig. 9A–F Dinopis celebensis Merian, 1911: 167, figs A, B (♂only, ♀ mismatched). Type material examined.

♂ (NMB), NMB-ARAN-00514b, “Zentral-Celebes, nördlich vom Golf von Bone”, South Sulawesi, north of the Gulf of Boni (precise locality not known).

 Diagnosis.

The male can be distinguished from other congeners by having the distal lobe of the MA distinctly smaller than the basal lobe; in other Asianopis spp., the distal lobe is slightly smaller than the basal lobe (Fig. 9A, C).

10.3897/zookeys.911.38761.figure9 3676226 16CE0FE6-2721-5A01-AB59-0D83F448A1CD

Asianopis celebensis comb. nov., male type. A Male right palp, prolateral view B Embolic tip C male left palp (embolic tip detached), retrolateral view D Male prosoma, lateral view E Male opisthosoma, dorsal view F Male prosoma, frontal view.

https://binary.pensoft.net/fig/380021 Description.

See Merian (1911). Photos of holotype male habitus and palps are shown in Fig. 9A–F.

 Distribution.

Indonesia (Sulawesi).

 Comments.

One male and two females were types for Asianopis celebensis (Merian, 1911) comb. nov. after Merian (1911). Based on the current study, one type female from North Sulawesi is Asianopis dumogae (Merian, 1911) sp. reval. comb. nov., and the other type female from South Sulawesi is a member of the zhuanghaoyuni-group, but its status at the species level is uncertain because of the missing epigyne.

 Animalia Araneae Deinopidae 884BE3FF-059D-511D-BD11-8D1BBC06B116 Asianopis konplong (Logunov, 2018) comb. nov. Deinopis konplong Logunov, 2018: 141, figs 1–7 (♂). Type.

Holotype ♂ (MMUE, G7579.37) from Vietnam, Kon Tum Province, Kon Plong District, 14 km north of Kon Plong, 14°43'20"N, 108°18'59"E, elevation ca 1030 m, 3–12.VI.2016, A.A. Abramov leg. Not examined.

 Diagnosis.

This species can be distinguished from other Asianopis species by the short palp (ratio of the length of the palpal tarsus to the length of the cymbium: 1:1) and upturned embolic tip (Logunov 2018: fig. 4).

 Description.

See Logunov (2018).

 Distribution.

Vietnam (Kon Tum).

 Animalia Araneae Deinopidae CCC7E087-4B38-54C6-BFC6-F70A71B74D98 Asianopis wangi http://zoobank.org/64A4C3D1-03A5-4D7A-B2E6-E30EA28DC41C Lin & Li sp. nov. Figs 10 , 11 , 12 , 20B , 21C , 22C, D, H , 23 Type.

Holotype. ♂ (IZCAS-Ar39681), China, Hainan Province, Wuzhishan City, Wuzhishan Nature Reserve, Diewupo, 17.V.2019, Dongdong Wang leg.

Paratypes. 1♂1♀ (IZCAS-Ar39682-Ar39683), same data as holotype; 1♂2♀ (IZCAS-Ar39684-Ar39686) China, Hainan Province, Wuzhishan City, Nansheng Town, Maoxiang Village, 18.V.2019, Dongdong Wang leg.

 Etymology.

The species is named after Mr Dongdong Wang, the collector of the holotype; noun (name) in genitive case.

 Diagnosis.

The males resemble A. zhuanghaoyuni sp. nov. but can be distinguished from other species by the ratio of the length of the embolic opening to the length of the embolic tip fold, which is 1:6 in A. wangi sp. nov. and 1:8 in A. zhuanghaoyuni sp. nov. The fold is more developed in A. wangi sp. nov. (Fig. 21C, D). The median plate is triangular in A. wangi sp. nov. and subtriangular in A. zhuanghaoyuni sp. nov. (Figs 12, 19).

 Description.

Male holotype (Figs 10A, C, D, 11, 20B, 21D, 22C). Total length 15.31, carapace 6.22 long, 4.60 wide, opisthosoma 9.32 long, 2.10 wide. Eye sizes and interdistances: AME 0.30, ALE 0.38, PME 0.65, PLE 0.34, AME–AME 0.30, AME–ALE 0.97, PME–PME 0.23, PME–PLE 0.69, AME–PME 0.24, ALE–PLE 1.82. Clypeus height 0.10. Chelicerae with four promarginal and 10–13 retromarginal teeth. Leg measurements: leg I: 84.08 (21.13 + 26.50 + 29.53 + 6.92), leg II: 59.70 (18.39 + 19.55 + 15.80 + 5.96), leg III: 36.14 (12.05 + 11.79 + 10.26 + 2.04), leg IV: 37.23 (11.92 + 12.37 + 11.28 + 1.66). Leg formula: 1243.

Male palp (Figs 11, 20B, 21D). Cymbium hemispherical; tegulum flat, obscured by embolic coils; embolus long and strongly coiled, originating at 10 o’clock and coiling 1500° around MA; embolic tip widened subapically, strongly folded and without apophysis. MA large, with two lobes.

Female paratype (Figs 10B, E, F, 12, 22D). Total length 24.04, carapace 7.56 long, 5.32 wide, opisthosoma 16.28 long, 6.86 wide. Eye sizes and interdistances: AME 0.28, ALE 0.38, PME 1.34, PLE 0.42, AME–AME 0.13, AME–ALE 1.03, PME–PME 0.39, PME–PLE 1.30, AME–PME 0.22, ALE–PLE 1.92. Clypeus height 0.34 (n = 1). Chelicerae with four promarginal and 8–13 retromarginal teeth (8(n = 1), 13(n = 1)). Leg measurements: Leg I: 54.24 (16.22 + 16.83 + 17.63 + 3.56), leg II: 50.59 (15.90 + 16.41 + 15.00 + 3.28), leg III: 30.84 (10.96 + 10.38 + 7.88 + 1.62), leg IV: 30.28 (10.13 + 10.58 + 8.27 + 1.30). Leg formula: 1234.

10.3897/zookeys.911.38761.figure10 3676182 FF4347E8-34E3-5232-B962-D05B846EA712

Asianopis wangi sp. nov., male holotype and female paratype. A Male prosoma, frontal view B Female prosoma, frontal view C Male habitus, dorsal view D Male habitus, ventral view E Female habitus, dorsal view F Female habitus, ventral view.

https://binary.pensoft.net/fig/380022 10.3897/zookeys.911.38761.figure11 3676184 5A8DB6B9-A124-5AF9-87BE-A449D6F5F4E3

Asianopis wangi sp. nov., left palp, male holotype. A Prolateral view B Retrolateral view.

https://binary.pensoft.net/fig/380023 10.3897/zookeys.911.38761.figure12 3676186 0906786B-E462-50D6-9E4F-2C2AA43FBECD

Asianopis wangi sp. nov., female paratype. A Epigyne B Vulva, dorsal view.

https://binary.pensoft.net/fig/380024

Epigyne (Fig. 12) with a median plate, CD with 7 or 8 turns, S oval, SpD consistently narrow.

 Distribution.

China (Hainan).

 Animalia Araneae Deinopidae 4E6A07E0-5DE1-56A3-BD97-D1F0919F2CA0 Asianopis wuchaoi http://zoobank.org/F05E46B7-98E7-4DA1-B7DF-AD440C2E05B6 Lin & Li sp. nov. Figs 13 , 14 , 15 , 21B , 22B , 23 Type.

Holotype. ♂ (IZCAS-Ar39687), China, Yunnan Province, Jinghong City, Mount Jinuo, 10.V.2019, Chao Wu leg.

Paratypes. 2♀ (IZCAS-Ar39688-Ar39689), China, Yunnan Province, Jinghong City, Mengla County, Mengxing Village, 16.VI.2019, Yi Li leg.; 1♀ (IZCAS-Ar39690), China, Yunnan Province, Jinghong City, Situlaozhai Village, 20.V.2019, Chaotai Wei leg.

 Etymology.

The species is named after Mr Chao Wu, the collector of the holotype male; noun (name) in genitive case.

 Diagnosis.

The males can be easily distinguished by the length of the palpal tibia which is almost equal to the length of the cymbium; simple embolic tip with ETA (Fig. 21B); embolus coiling almost 3300° around MA. Epigyne with a well-developed, subtriangular median plate, obscureing CO, and CD with 9 turns (Fig. 14).

 Description.

Male holotype (Figs 13A, C, D, 14, 21A). Total length 12.14, carapace 4.00 long, 3.40 wide, opisthosoma 8.14 long, 2.4 wide. Eye sizes and interdistances: AME 0.15, ALE 0.26, PME 0.52, PLE 0.29, AME–AME 0.17, AME–ALE 0.70, PME–PME 0.16, PME–PLE 0.61, AME–PME 0.11, ALE–PLE 0.95. Clypeus height 0.05. Chelicerae with four promarginal and six retromarginal teeth. Leg measurements: leg I: damaged, leg II: damaged, leg III: (6.92 + 6.86 + ? + 1.44), leg IV: 21.82 (6.91 + 7.18 + 6.35 + 1.38).

10.3897/zookeys.911.38761.figure13 3676190 A6EAA07B-0A86-5575-B87D-8A0C5BDED61C

Asianopis wuchaoi sp. nov., male holotype and female paratype. A Male prosoma, frontal view B Female prosoma, frontal view C Male habitus, dorsal view D Male habitus, ventral view E Female habitus, dorsal view F Female habitus, ventral view.

https://binary.pensoft.net/fig/380025 10.3897/zookeys.911.38761.figure14 3676192 D62ED350-D48F-5A9E-9C51-49F196030E9F

Asianopis wuchaoi sp. nov., male holotype. A Right palp (flipped horizontally), prolateral view B Right palp (flipped horizontally), retrolateral view C Left palp, prolateral view D Left palp, prolateral view.

https://binary.pensoft.net/fig/380026

Male palp (Figs 14, 21A). Cymbium hemispherical; tegulum flat, obscured by embolic coils; embolus long and strongly coiled, originating at five o’clock and coiling 3300° around MA. MA large, with two lobes.

Female paratype (Figs 13B, E, F, 15). Total length 14.60, carapace 6.28 long, 4.10 wide, opisthosoma 9.29 long, 3.72 wide. Eye sizes and interdistances: AME 0.11, ALE 0.34, PME 0.94, PLE 0.29, AME–AME 0.30, AME–ALE 1.03, PME–PME 0.06, PME–PLE 0.64, AME–PME 0.14, ALE–PLE 1.33. Clypeus height 0.13 (n = 1). Chelicerae with four promarginal and 8–13 retromarginal teeth (8(n = 1), 10(n = 1), 13(n = 1)). Leg measurements: Leg I: 39.82 (12.11 + 11.67 + 13.01 + 3.03), leg II: 36.81 (11.47 + 11.79 + 10.83 + 2.72), leg III: 23.53 (9.47 + 6.79 + 5.83 + 1.44), leg IV: 21.71 (7.18 + 7.76 + 5.70 + 1.07). Leg formula: 1234.

10.3897/zookeys.911.38761.figure15 3676194 1825FBB0-0B43-5C7E-A960-D107E3469101

Asianopis wuchaoi sp. nov., female paratype. A Epigyne B Vulva, dorsal view.

https://binary.pensoft.net/fig/380027

Epigyne (Fig. 15) with a median plate, obscuring CO, CD with 9 turns, S oval, SpD is consistently thin.

 Distribution.

China (Yunnan).

 Note.

The male died during ecdysis so some legs are damaged or curled, and the palps are expanded.

 Animalia Araneae Deinopidae 5184F46B-9ADD-58D6-AAAD-2A7212722DC1 Asianopis zhuanghaoyuni http://zoobank.org/21A5E514-F8EE-4479-9338-51D419AA6E4A Lin & Li sp. nov. Figs 2D, G, H, J , 16 , 17 , 18 , 20 , 21D , 22E, F, H , 23 Type.

Holotype. ♂ (IZCAS-Ar39691), China, Fujian Province, Fuzhou City, Minhou County, Xiyuan Reservoir, 26°03'15.5"N, 119°06'05.4"E, elevation ca 102 m, 25.VI.2018, Haoyun Zhuang and Zhuoheng Jiang leg.

Paratypes. 1♀ (IZCAS-Ar39692), same data as holotype, Haoyun Zhuang leg.; 1♂1♀ (IZCAS-Ar39693-Ar39694), same locality data as holotype, but 15.V.2018, Haoyun Zhuang leg.; 1♂4♀ (IZCAS-Ar39695-Ar39699), same locality data as holotype, but 19.VI.2019, Haoyun Zhuang leg.; 1♂1♀ (IZCAS-Ar39700-Ar39701), same locality data as holotype, but 26.V.2019, Haoyun Zhuang leg.

 Etymology.

The species is named after Mr Haoyun Zhuang, the collector of the type specimens; noun (name) in genitive case.

 Diagnosis.

The males resemble A. konplong (Logunov, 2018) comb. nov. but can be distinguished by the embolus originating at five o’clock in A. zhuanghaoyuni sp. nov. (9 o’clock in A. konplong (Logunov, 2018) comb. nov.); the ratio of the length of the palpal tarsus to the length of the cymbium is 11:9 in A. zhuanghaoyuni sp. nov., while in A. konplong (Logunov, 2018), comb. nov. it is 1:1 (Figs 18, 22A; Logunov 2018, figs 4–6).

 Description.

Male holotype (Figs 2G, 16A, C, D, 17, 20A, 21E, 22E). Total length 16.54, carapace 5.58 long, 3.84 wide, opisthosoma 11.40 long, 1.90 wide. Eye sizes and interdistances: AME 0.25, ALE 0.30, PME 0.59, PLE 0.30, AME–AME 0.25, AME–ALE 0.85, PME–PME 0.23, PME–PLE 0.59, AME–PME 0.19, ALE–PLE 1.28. Clypeus height 0.20. Chelicerae with four promarginal teeth and a retromarginal tooth. Leg measurements: leg I: 66.35 (18.50 + 22.55 + 18.95 + 6.35), leg II: 52.87 (16.54 + 17.65 + 13.10 + 5.58), leg III: 30.39 (10.78 + 10.83 + 7.18 + 1.60), leg IV: 30.06 (10.42 + 11.12 + 7.18 + 1.34). Leg formula: 1234.

10.3897/zookeys.911.38761.figure16 3676196 12F00672-15A4-58A6-9419-4A6194138A58

Asianopis zhuanghaoyuni sp. nov., male holotype and female paratype. A Male prosoma, frontal view B Female prosoma, frontal view C Male habitus, dorsal view D Male habitus, ventral view E Female habitus, dorsal view F Female habitus, ventral view.

https://binary.pensoft.net/fig/380028 10.3897/zookeys.911.38761.figure17 3676198 B3849819-6C7C-5CF3-BB0B-FC7225759B61

Asianopis zhuanghaoyuni sp. nov., male holotype, left palp. A Prolateral view B Retrolateral view.

https://binary.pensoft.net/fig/380029

Male palp (Figs 18, 22A). Cymbium hemispherical; tegulum flat, obscured by embolus coils; originating at five o’clock, coiling 1500° around MA, embolic tip widened subapically, folded and without apophysis. MA large, with two lobes.

10.3897/zookeys.911.38761.figure18 3676200 77313048-6C4B-50D4-B4C2-AC05327F0C25

Asianopis zhuanghaoyuni sp. nov., female paratype. A Epigyne B Vulva, dorsal view.

https://binary.pensoft.net/fig/380030 10.3897/zookeys.911.38761.figure19 3676202 3BFD452E-27FE-5AF9-95DE-F5CE5D08C521

Asianopis liukuensis comb. nov., left palp, ventral view. A, B Male from Xishuangbanna C Male from India, type of Deinopis scrubjunglei syn. nov.

https://binary.pensoft.net/fig/380031 10.3897/zookeys.911.38761.figure20 3676206 888AAAB9-7F26-5369-8705-3FECCAE0B248

Ventral view of left palp, holotype males. AA. zhuanghaoyuni sp. nov. BA. wangi sp. nov.

https://binary.pensoft.net/fig/380032 10.3897/zookeys.911.38761.figure21 3676208 CF38CE57-1BE0-52AC-92E5-BCBAD8135160

Embolic tips of four species of Asianopis gen. nov. AA. liukuensis (Yin, Griswold & Yan, 2002) comb. nov. BA. wuchaoi sp. nov. CA. zhuanghaoyuni sp. nov. DA. wangi sp. nov.

https://binary.pensoft.net/fig/380033 10.3897/zookeys.911.38761.figure22 3676210 739D710A-C438-57A9-8523-C01C72598308

Photos of four live spiders of Asianopis gen. nov., including webs of two species of Asianopis gen. nov. AA. liukuensis comb. nov., female BA. wuchaoi sp. nov., female CA. wangi sp. nov., male DA. wangi sp. nov., female EA. zhuanghaoyuni sp. nov., male FA. zhuanghaoyuni sp. nov., female G Web of A. liukuensis comb. nov. H Web of A. wangi sp. nov.

https://binary.pensoft.net/fig/380034 10.3897/zookeys.911.38761.figure23 3676212 26DB2995-A035-5569-A8D4-3C584173BB07

Distribution records of seven species of Asianopis gen. nov. in Asia. 1A. liukuensis comb. nov. 2A. dumogae sp. reval. comb. nov. 3A. celebensis comb. nov. 4A. konplong comb. nov. 5A. wangi sp. nov. 6A. wuchaoi sp. nov. 7A. zhuanghaoyuni sp. nov.

https://binary.pensoft.net/fig/380035

Female paratype (Figs 2H, J, 16B, E, F, 18, 22F, H). Total length 22.60, carapace 5.90 long, 4.55 wide, opisthosoma 15.40 long, 5.90 wide. Eye sizes and interdistances: AME 0.22, ALE 0.35, PME 1.08, PLE 0.33, AME–AME 0.37, AME–ALE 1.22, PME–PME 0.16, PME–PLE 0.98, AME–PME 0.081, ALE–PLE 1.61. Clypeus height 0.59. (n = 1). Chelicerae with four promarginal and 10 or 11 (10 (n = 2), 11 (n = 1)) retromarginal teeth. Leg measurements: Leg I: 49.68 (14.80 + 15.83 + 16.02 + 3.03), leg II: 46.08 (14.71 + 15.20 + 13.33 + 2.84), leg III: 27.79 (9.73 + 9.41 + 7.18 + 1.47), leg IV: 26.78 (9.02 + 9.61 + 6.86 + 1.29). Leg formula: 1234.

Epigyne (Fig. 18) with a median plate, CD with 7–8 turns, S oval, SpD consistently thin.

 Distribution.

China (Fujian).

 Acknowledgements

The manuscript benefitted greatly from comments by Gergin Blagoev, Alireza Zamani, Marc Milne, Jie Liu, and an anonymous referee. Sarah Crews kindly checked the English of the manuscript. Jishen Wang, Chaotai Wei, Yi Li, Haoyun Zhuang, Dongdong Wang, Yuanping Wang, Chao Wu, Pinmin Li, and Qianle Lu rendered effective assistance in fieldwork. Zhuoheng Jiang, Linrui Yu and colleagues from the College of Bioscience & Engineering of Fuzhou University helped in collecting the type materials of Asianopis zhuanghaoyuni Lin & Li, sp. nov. Dongdong Wang, Zixuan Lin, Haoyun Zhuang, and Zhengzhong Huang took photos of the live spiders. This study was supported by the National Natural Science Foundation of China (NSFC- 31530067) to Shuqiang Li.

 References CalebJTDMathaiMT (2014) A new species of Deinopis MacLeay (Araneae: Deinopidae) from India.Indian Journal of Arachnology3(1): 1–7. Capella-GutiérrezSSilla-MartínezJMGabaldónT (2009) TrimAl: a tool for automated alignment trimming in large-scale phylogenetic analyses.Bioinformatics25: 1972–1973. https://doi.org/10.1093/bioinformatics/btp348 ChamberlandLMcHughAKechejianSBinfordGJBondJECoddingtonJADolmanGHamiltonCAHarveyMSKuntnerMAgnarssonI (2018) From Gondwana to GAARlandia: evolutionary history and biogeography of ogre-faced spiders (Deinopis).Journal of Biogeography45: 2442–2457. https://doi.org/10.1111/jbi.13431 CoddingtonJAKuntnerMOpellBD (2012) Systematics of the spider family Deinopidae with a revision of the genus Menneus.Smithsonian Contributions to Zoology636: 1–61. https://doi.org/10.5479/si.00810282.636.1 DoleschallL (1859) Tweede Bijdrage tot de kennis der Arachniden van den Indischen Archipel.Acta Societatis Scientiarum Indica-Neerlandica5: 1–60. GriswoldCERamírezMJCoddingtonJAPlatnickNI (2005) Atlas of phylogenetic data for entelegyne spiders (Araneae: Araneomorphae: Entelegynae) with comments on their phylogeny. Proceedings of the California Academy of Sciences 56 (Supplement II): 1–324. KatohKStandleyDM (2013) MAFFT. Multiple sequence alignment software version 7: improvements in performance and usability.Molecular Phylogenetics and Evolution30: 772–780. https://doi.org/10.1093/molbev/mst010 KumarSStecherGTamuraK (2016) MEGA7: Molecular Evolutionary Genetics Analysis version 7.0 for bigger datasets.Molecular Biology and Evolution33(7): 1870–1874. https://doi.org/10.1093/molbev/msw054 LanfearRCalcottBHoSYGuindonS (2012) Partitionfinder: combined selection of partitioning schemes and substitution models for phylogenetic analyses.Molecular Biology and Evolution29: 1695–1701. https://doi.org/10.1093/molbev/mss020 LogunovDV (2018) A new ogre-faced spider species of the genus Deinopis MacLeay, 1839 from Vietnam (Aranei: Deinopidae).Arthropoda Selecta27(2): 139–142. https://doi.org/10.15298/arthsel.27.2.05 MacLeayWS (1839) On some new forms of Arachnida. Annals of Natural History (Series 1) 2(7): 1–14. [pls 1, 2] https://doi.org/10.1080/00222933809496646 MerianP (1911) Die Spinnenfauna von Celebes. Beiträge zur Tiergeographie im Indoaustralischen Archipel.Zoologische Jahrbücher, Abteilung für Systematik, Geographie und Biologie der Tiere31: 165–354. RambautASuchardMAXieDDrummondAJ (2014) Tracer v1.6. http://tree.bio.ed.ac.uk/software/tracer [Accessed on: 2019–12–31] RoewerCF (1938) Araneae. Résultats scientifiques du Voyage aux indes orientales néerlandaises de la SS. AA. RR. le Prince et la Princesse Leopold de Belgique.Mémoires du Musée Royal d’Histoire Naturelle de Belgique3(19): 1–94. RonquistFTeslenkoMvan der MarkPAyresDLDarlingAHohnaSLargetBLiuLSuchardMAHuelsenbeckJP (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space.Systematic Biology61: 539–542. https://doi.org/10.1093/sysbio/sys029 SimonE (1876) Etude sur le arachnides du Congo. Bulletin de la Société Zoologique de France 1: 12–15, 215–224. SimonE (1909) Etude sur les arachnides du Tonkin (1re partie).Bulletin Scientifique de la France et de la Belgique42: 69–147. https://doi.org/10.5962/bhl.part.24151 StamatakisA (2014) RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies.Bioinformatics30: 1312–1313. https://doi.org/10.1093/bioinformatics/btu033 World Spider Catalog (2019) World Spider Catalog, version 20.5. Natural History Museum, Bern. http://wsc.nmbe.ch [Accessed on: 2019–12–21] YinCMGriswoldCEYanHM (2002) A new ogre-faced spider (Deinopis) from the Gaoligong Mountains, Yunnan, China (Araneae, Deinopidae). Journal of Arachnology 30: 610–612. https://doi.org/10.1636/0161-8202(2002)030[0610:ANOFSD]2.0.CO;2 Supplementary materials 10.3897/zookeys.911.38761.suppl1 3676228 2BE549ED-34FF-53B2-ADDD-0B62AF65B1A3

Asianopis gen. nov., a new genus of the spider family Deinopidae from Asia

specimen/primer/DNA sequence

Table S1. List of voucher information and GenBank accession numbers. Table S2. Primers and PCR conditions for the genetic markers used in this study (modified after Zhao and Li unpublished). Table S3. Sequence characteristics and models of DNA evolution selected for the seven sequence regions analyzed.References cited in supplementary tables.

 https://binary.pensoft.net/file/380036 This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Yejie Lin, Lili Shao, Ambros Hänggi, John T.D. Caleb, Joseph K.H. Koh, Peter Jäger, Shuqiang Li