A survey of the spider genus Lipocrea Thorell, 1878 (Araneae, Araneidae) from Guiyang City, Southwest China: An integrated morphological and molecular approach

207 A survey of the spider genus Lipocrea Thorell, 1878 (Araneae, Araneidae) from Guiyang City, Southwest China: An integrated morphological and molecular approach Jianshuang Zhang1* , Chengwen Zhang1* , Yuanqian Xing1, Hao Yu1, Xiaoqi Mi2 1 The State Key Laboratory of Southwest Karst Mountain Biodiversity Conservation of Forestry Administration, School of Life Sciences, Guizhou Normal University, Guiyang, Guizhou, China 2 College of Agriculture and Forestry Engineering and Planning, Guizhou Provincial Key Laboratory of Biodiversity Conservation and Utilization in the Fanjing Mountain Region, Tongren University, Tongren, Guizhou, China Corresponding author: Xiaoqi Mi ([email protected]) Copyright: © Jianshuang Zhang et al. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International – CC BY 4.0). Research Article Abstract A survey was undertaken to study the spider genus Lipocrea Thorell, 1878, from Guiyang City, Guizhou Province, southwest China. A total of two species is here addressed based on morphology and five methods of molecular species delimitation, comprising L. guiyang J. Zhang, Yu & Mi, sp. nov. and L. fusiformis (Thorell, 1877), the type species of the genus as well as a new record for mainland China. These two species are distributed in Huaxi District and Kaiyang County of Guiyang, respectively, providing the first formal record of this genus from mainland China. Key words: Biodiversity, DNA barcoding, molecular species delimitation, morphology, new species, orb-weaver Introduction Lipocrea Thorell, 1878 is a small spider genus that was originally established to include three species: L. phthisica (L. Koch, 1871) and L. tabida (L. Koch, 1872) from the Australasian Region, and the type species L. fusiformis (Thorell, 1877) from the Oriental Region. An additional Oriental species, L. diluta Thorell, 1887 was subsequently described. However, the validity of Lipocrea as a distinct genus was not recognized by Simon (1887: 187), who treated these species under the genus Larinia Simon, 1874. Of the aforementioned species, L. phthisica has since been transferred to Larinia sensu stricto (Grasshoff 1970). The remaining species, together with Larinia longissima (Simon, 1881) from the Ethiopian Region, were subsequently removed from Larinia and placed in a newly erected genus, Larinopa Grasshoff (1970), by Grasshoff (1970: 226). Although the name Lipocrea remains available under the rules of zoological nomenclature, it has been largely overlooked (Levy 1986). The genus Lipocrea, as redefined by Grasshoff under the name Larinopa, is primarily distinguished from Larinia by the structure of the genitalia, especially the morphology of the male palpal organ. Academic editor: Cristina Rheims Received: 12 May 2025 Accepted: 25 August 2025 Published: 10 October 2025 ZooBank: https://zoobank. org/07FB18BD-9211-413B-A66005B73FF82A0F Citation: Zhang J, Zhang C, Xing Y, Yu H, Mi X (2025) A survey of the spider genus Lipocrea Thorell, 1878 (Araneae, Araneidae) from Guiyang City, Southwest China: An integrated morphological and molecular approach. ZooKeys 1255: 207–237. https://doi.org/10.3897/ zookeys.1255.158340 ZooKeys 1255: 207–237 (2025) DOI: 10.3897/zookeys.1255.158340 * These authors contributed equally to this work. 208 ZooKeys 1255: 207–237 (2025), DOI: 10.3897/zookeys.1255.158340 Jianshuang Zhang et al.: A survey of Lipocrea from Guiyang Currently, the WSC (2025) lists five species under Lipocrea, none of which are known from the Chinese mainland (WSC 2025). However, in several publications, some of these species are considered to belong to Larinia. For example, L. phosop (Tanikawa, Into & Petcharad, 2023) was originally described as Larinia has not yet been formally transferred to Lipocrea. Nevertheless, due to its strong morphological similarity to the type species of Lipocrea, L. phosop has been provisionally placed in Lipocrea by the WSC (2025). Even the type species, L. fusiformis, has frequently been placed in Larinia and redescribed as such by various authors, including Tanikawa (1989, 2007, 2009), Okuma et al. (1993), Barrion and Litsinger (1995), and Chang and Tso (2004). In view of the above-mentioned, the validity of the genus Lipocrea and the current generic placement of its constituent species remains in dispute. A preliminary genus-level taxonomic molecular analysis of Grasshoff’s (1970) ‘Larinia group’ was carried out based on all available COI sequences of the Larinia group and related genera (69 from NCBI and 9 newly sequenced here) (Suppl. material 1: table S1). According to the results (Suppl. material 1: fig. S1): (1) the monophyly of the genus Lipocrea is well supported; (2) Larinia is polyphyletic, which is consistent with the results of Scharff et al. (2020). Based on this preliminary result and considering that a formal comprehensive revision of the Larinia group is lacking, we agree with Framenau and Castanheira (2022) that the Larinia group requires further systematic study that includes Larinia lineata Lucas, 1846 (the type species of the genus). A review of the Larinia group is not within the scope of this work. Consequently, the present study assigns the two species treated here to the monophyletic genus Lipocrea, rather than to the polyphyletic Larinia sensu lato. Guiyang, the provincial capital of Guizhou Province (Fig. 1A), is the first city to receive the accolade of ‘the national forest city’ in China, famous for more than 55% forest coverage (Zhong 2018). Guiyang is also known as the ‘city of a thousand gardens’, with 1025 urban parks, and is considered one of China’s most biodiverse provincial capitals (Yang 2020; Cheng and Chen 2022). However, spiders can be regarded as being poorly represented in Guiyang, with only 64 species from 25 families recorded or described to date (JZ and HY, unpubl. data). Of these, 21 species are endemic, 16 were newly described, and 15 were reported as new records in recent years (Yu et al. 2018; Yu and Zhang 2019; Xin et al. 2020; Yan et al. 2021; Long et al. 2022; Zhang et al. 2022a, b, 2023; He et al. 2023; Li et al. 2023; Yang et al. 2023a, b, c; Ding et al. 2024; Pan et al. 2024; Jiang et al. 2025; Zhang et al. 2025). This estimate of spider diversity is assumed to be far from the true diversity within this city. As mentioned above, spiders are poorly studied in Guiyang, including Araneidae. Only five species have been recorded: Cyrtarachne bufo (Bösenberg & Strand, 1906), Hypsosinga pygmaea (Sundevall, 1831), Neoscona xishanensis Yin, Wang, Xie & Peng, 1990, Nephila pilipes (Fabricius, 1793) and Pronoides brunneus Schenkel, 1936. Recently, short but intensive field collections in Guiyang have been conducted by the staff of the Guizhou Normal University. During these surveys, we have found some Lipocrea specimens that belong to at least two morphospecies (Figs 1–3): one is new to science, and the other one has been identified as L. fusiformis, a new record for mainland China. The sympatric distribution with L. fusiformis and the high intraspecific morphological variation in females of the new species pose significant challenges for sexual pairing and species identification. We therefore generated DNA barcode data 209 ZooKeys 1255: 207–237 (2025), DOI: 10.3897/zookeys.1255.158340 Jianshuang Zhang et al.: A survey of Lipocrea from Guiyang Figure 1. Locality of Guiyang City (A) and distribution records of Lipocrea species in Guiyang (B). A B 210 ZooKeys 1255: 207–237 (2025), DOI: 10.3897/zookeys.1255.158340 Jianshuang Zhang et al.: A survey of Lipocrea from Guiyang Figure 2. Living specimens of Lipocrea guiyang J. Zhang, Yu & Mi, sp. nov. A. Male; B. Female. Photographs by Q Lu (Shenzhen). A B 211 ZooKeys 1255: 207–237 (2025), DOI: 10.3897/zookeys.1255.158340 Jianshuang Zhang et al.: A survey of Lipocrea from Guiyang Figure 3. Living specimens of Lipocrea fusiformis (Thorell, 1877). A. Male; B, C. Female. Photographs by Q Lu (Shenzhen). AB C 212 ZooKeys 1255: 207–237 (2025), DOI: 10.3897/zookeys.1255.158340 Jianshuang Zhang et al.: A survey of Lipocrea from Guiyang to devise a specimen phylogeny and used five molecular species delimitation methods to test morphology-based species identification. The goal of this paper is to: 1) use the consensus results of an integrated morphological and molecular approach to delimit Lipocrea spiders from Guiyang; 2) describe the new species under the name of Lipocrea guiyang J. Zhang, Yu & Mi, sp. nov.; 3) re-illustrate L. fusiformis based on new material from Guiyang, including supplementary micrographs; and 4) provide a distribution map of Lipocrea species in Guiyang City. Materials and methods Taxon sampling Specimens in this study were collected by hand and by beating vegetation, and directly fixed in absolute ethanol. The right legs were removed and stored at −80 °C for subsequent DNA extraction. The remainder of the specimens was preserved in 80% ethanol for identification and morphological examination. A total of 11 adults were obtained, examined, and processed for DNA extraction but only nine individuals yielded useable DNA (Table 1). All voucher specimens (including types of the new species) are deposited in the Museum of Guizhou Normal University, Guiyang, China (MGNU). Molecular protocols Total genomic DNA was extracted using the Cell & Tissue Genomic DNA Isolation Kit (Bioteke, Beijing, China) following the manufacturer’s protocols. We amplified cytochrome c oxidase subunit I (COI) using the primer pairs LCO1490/ HCO2198 (Folmer et al. 1994) and standard polymerase chain reaction (PCR) settings (Wheeler et al. 2016). PCR products were transported to the Beijing Tsingke Biotech Co., Ltd. (Chongqing, China) for sequencing using the same PCR primers. We manually edited the sequences using Geneious Prime 2024 (Kearse et al. 2012), translated nucleotide sequences into amino acids to check for stop codons, and ensured the proper configuration of codon positions. Phylogenetic analyses For phylogenetic analyses, the COI data of 17 specimens of Lipocrea were used as the in-group, including eight sequences of L. phosop downloaded from GenBank (Tanikawa et al. 2023; NCBI 2025). To root the tree, we included two specimens of Larinia joei Tanikawa & Petcharad, 2021 from NCBI (2025) as the outgroup (Table 1). We performed maximum-likelihood (ML) analyses using IQ-TREE v. 2.3.1 (Minh et al. 2020) based on the best COI substitution model (GTR+I+G) in jMODELTEST v. 2.1.10 (Darriba et al. 2012). Branch support was estimated with ultrafast bootstrapping with 1000 replicates (Hoang et al. 2018). Bayesian-inference (BI) was performed with MrBayes v. 3.2.1 (Ronquist et al. 2012) using one independent chain for 50 million generations. The first 10% of trees from each run were discarded as burn-in. Finally, we used FigTree v. 1.4.4 (Rambaut 2012) to visualize and manipulate trees and used Photoshop CC 2018 to summarize them. 213 ZooKeys 1255: 207–237 (2025), DOI: 10.3897/zookeys.1255.158340 Jianshuang Zhang et al.: A survey of Lipocrea from Guiyang Table 1. Samples used in species delimitation: specimen label, taxon name, sample collection locality with coordinates, and GenBank accession numbers. Specimen code Genus/Species Sex Locality Country Coordinates Elevation (m a.s.l.) COI GenBank accession YHGY208 Lipocrea guiyang sp. nov. ♂Nanjiang Grand Canyon, Kaiyang County, Guiyang City, Guizhou Prov. China 26.94°N, 106.97°E 861 PX230067 YHGY209 Lipocrea guiyang sp. nov. ♀Nanjiang Grand Canyon, Kaiyang County, Guiyang City, Guizhou Prov. China 26.94°N, 106.97°E 861 PX230068 YHGY428 Lipocrea guiyang sp. nov. ♀Nanjiang Grand Canyon, Kaiyang County, Guiyang City, Guizhou Prov. China 26.94°N, 106.97°E 861 PX230069 YHGY431 Lipocrea fusiformis ♂Dangwu Town, Huaxi District, Guiyang City, Guizhou Prov. China 26.38°N, 106.60°E 1138 PX230061 YHGY432 Lipocrea fusiformis ♀University Town, Huaxi District, Guiyang City, Guizhou Prov. China 26.38°N, 106.65°E 1173 PX230062 YHGY433 Lipocrea fusiformis ♀Dangwu Town, Huaxi District, Guiyang City, Guizhou Prov. China 26.38°N, 106.60°E 1138 PX230063 YHGY495 Lipocrea fusiformis ♂University Town, Huaxi District, Guiyang City, Guizhou Prov. China 26.38°N, 106.65°E 1173 PX230065 YHGY507 Lipocrea fusiformis ♀University Town, Huaxi District, Guiyang City, Guizhou Prov. China 26.38°N, 106.65°E 1173 PX230066 YHGY508 Lipocrea fusiformis ♀University Town, Huaxi District, Guiyang City, Guizhou Prov. China 26.38°N, 106.65°E 1173 PX230064 AT5343 Lipocrea phosop ♂Mai Khet, Mueang Prachinburi District, Prachinburi Prov. Thailand 14.10°N, 101.33°E LC756453 AT5337 Lipocrea phosop ♀Mai Khet, Mueang Prachinburi District, Prachinburi Prov. Thailand 14.10°N, 101.33°E LC756454 AT5338 Lipocrea phosop ♀Mai Khet, Mueang Prachinburi District, Prachinburi Prov. Thailand 14.10°N, 101.33°E LC756455 AT5339 Lipocrea phosop ♀Mai Khet, Mueang Prachinburi District, Prachinburi Prov. Thailand 14.10°N, 101.33°E LC756456 AT5340 Lipocrea phosop ♀Mai Khet, Mueang Prachinburi District, Prachinburi Prov. Thailand 14.10°N, 101.33°E LC756457 AT5341 Lipocrea phosop ♀Mai Khet, Mueang Prachinburi District, Prachinburi Prov. Thailand 14.10°N, 101.33°E LC756458 AT5342 Lipocrea phosop ♂Mai Khet, Mueang Prachinburi District, Prachinburi Prov. Thailand 14.10°N, 101.33°E LC756459 AT5345 Lipocrea phosop ♂Ban Pathum, Sam Khok District, Pathum Thani Prov. Thailand 14.08°N, 100.58°E LC756460 LJO01 Larinia joei ♂Khlong Sam, Pathum Thani Prov. Thailand 14.16°N, 100.66°E LC597525 LJO02 Larinia joei ♀Khlong Sam, Pathum Thani Prov. Thailand 14.16°N, 100.66°E LC597526 Molecular species delimitation To delimit three morphospecies of Lipocrea based on an accompanying morphological study of the genus, we used two genetic distance-based methods: the DNA barcoding gap (Barrett and Hebert 2005) and ABGD (Puillandre et al. 2012), as well as three methods based on the inferred tree, GMYC (Pons et al. 2006), P ID (Liberal), and mPTP (Kapli et al. 2017). Because the P ID (Liberal) and DNA barcoding gap (Barrett and Hebert 2005) methods require a priori designation, we assigned 17 Lipocrea individuals to three putative species based on a combination of phylogenetic topology and morphological characteristics. With the DNA barcoding gap, we used the overlap between the interspecies and intraspecies Kimura two-parameter (K2P) and uncorrected p-distance for each candidate species calculated in MEGA X (Kumar et al. 2018). The P ID (Liberal) method tests species delimitation by 214 ZooKeys 1255: 207–237 (2025), DOI: 10.3897/zookeys.1255.158340 Jianshuang Zhang et al.: A survey of Lipocrea from Guiyang relying on defining the putative species groups. We used the BI tree as a guide to test species hypothesis (Xu et al. 2017). The other three methods that we used do not require terminals to be a priori assigned to putative species. ABGD calculates all pairwise distances in the data set, evaluates intraspecific divergences, and then sorts the terminals into candidate species with calculated P values. We performed the ABGD analysis on a web server (https://bioinfo.mnhn.fr/abi/public/abgd/) using three different models: Jukes-Cantor (JC69; Jukes and Cantor 1969), K2P (Kimura 1980), and simple distance (p-distance; Nei and Kumar 2000). We analyzed the data using two different values for the parameters Pmin (0.001 and 0.0001), Pmax (0.1 and 0.2), and relative gap width (X = 1.5 or 2), with the other parameters set to default values. We used the BI tree as a guide to test the species hypothesis (Xu et al. 2017). Two runs of 100 million steps were used for the mPTP logging every 1 million steps, discarding the first 2 million steps. Each run was started from a random delimitation. The GMYC methodology (Pons et al. 2006) analysis was conducted using the single-threshold model in the “splits” package (Ezard et al. 2009) for R 4.2.2 (R Development Core Team 2024). BEAST 2.6.7 (Bouckaert et al. 2014) was used to produce an ultrametric tree for the GMYC analysis. Analyses were run for 50 million steps with 10% of the trees in each chain discarded as burn-in. Morphological protocols Specimens were examined using an Olympus SZX7 stereomicroscope. Further details were studied under a CX41 compound microscope. Male and female copulatory organs were examined and illustrated after dissection. Epigynes were removed and cleared in lactic acid or a warm 10% potassium hydroxide (KOH) solution. Images were captured with a Canon EOS 70D digital camera (20.2 megapixels) mounted on an Olympus CX41 compound microscope and assembled using Helicon Focus v. 6.80 image-stacking software. All measurements were obtained using an Olympus SZX7 stereomicroscope and are given in millimeters. Eye diameters were measured at the widest part. The total body length does not include the chelicerae or spinnerets. Leg lengths are given as total length (femur, patella+tibia, metatarsus, tarsus). The terminology used in the text and figure legends follows Grasshoff (1970), Framenau and Scharff (2008), Framenau and Castanheira (2022), Tanikawa and Petcharad (2021), and Tanikawa et al. (2023). References to figures in the cited papers are listed in lowercase (fig. or figs); figures from this paper are noted with an initial capital (Fig. or Figs). The abbreviations used in the text are: AER anterior eye row; ALE anterior lateral eyes; AME anterior median eyes; C conductor; CD copulatory duct; CO copulatory opening; Cy cymbium; Em embolus; FD fertilisation duct; HP hook-shaped process of MA; KP knob-like projection; MA median apophysis; MS median septum; MOQ median ocular quadrangle; Pc paracymbium; PLE posterior lateral eyes; 215 ZooKeys 1255: 207–237 (2025), DOI: 10.3897/zookeys.1255.158340 Jianshuang Zhang et al.: A survey of Lipocrea from Guiyang PME posterior median eyes; R radix; RA radix apophysis; RER posterior eye row; Sc scape; Sp spermatheca; St subtegulum; T tegulum; TA terminal apophysis; TA I terminal apophysis I; TA II terminal apophysis II; TAA terminal apophysis appendix; TE tegular extension. The distribution map was generated with ArcGIS v. 10.5 (Environmental Systems Research Institute, Inc.). Results and discussion Based on traditional morphological characters and experience (matching of males and females we had hypothesized mainly on the basis of co-occurrence and compatibility of epigynes with male pedipalpal structures), all examined materials could be identified as at least two morphospecies: one belongs to an undescribed species new to science: L. guiyang sp. nov.; the other one was identified as L. fusiformis (Grasshoff 1970; Chang and Tso 2004; Tanikawa et al. 2023). However, some morphological variation is exhibited in females of L. guiyang sp. nov. Five molecular species delimitation methods were employed to test the validity of the morphology-based identification of the three Lipocrea species and the accuracy of the proposed sex matching. The COI matrix of 17 Lipocrea individuals analyzed in this study had a sequence length of 629 bp, with 132 variable and 116 parsimony-informative sites. For COI, phylogenetic inference from BI and ML analyses yielded similar topologies with high support (Fig. 4; posterior probability, PP = 1; bootstrap value, BS = 100). The trees clearly divided the samples into four deeply divergent clades (Fig. 4). When considering three species of Lipocrea, interspecific distances were higher than intraspecific distances. Interspecific distances range from 3.27 to 11.69% for K2P (Fig. 5A) and from 3.18 to 10.65% for uncorrected p-distance (Fig. 5B). The lowest mean interspecific distance was 12.75% / 11.51% (K2P / uncorrected p-distance) found between L. phosop and L. fusiformis, and the highest mean intraspecific distance (1.81% / 1.77% K2P / uncorrected p-distance) was estimated for L. fusiformis. The barcoding gap range identified three species, one of which is new (Fig. 4). The ABGD results varied based on different parameter combinations of both the initial and recursive partitions. Based on the barcoding gap results, a distinct gap was observed among the three Lipocrea species, demonstrating substantial genetic divergence. Therefore, the value of X=2 was selected as the relative gap width (Table 2). The initial three species partition was consistent with the three morphospecies, while the recursive partition regime yielded more species (Fig. 4; Table 2). However, all analyses under different assumptions revealed three hypothetical species which is entirely consistent with the morphological identification (Fig. 4). Our P ID (Liberal) results revealed high P ID (Liberal) values > 0.95 (0.86–1.0) (Table 3), thereby also supporting the taxonomy of three putative species (Fig. 4). The results of our mPTP analysis indicated that, when only monophyletic species are considered, our three hypothetical species were identified as antic- 222 ZooKeys 1255: 207–237 (2025), DOI: 10.3897/zookeys.1255.158340 Jianshuang Zhang et al.: A survey of Lipocrea from Guiyang Figure 7. Lipocrea guiyang J. Zhang, Yu & Mi, sp. nov., female paratypes, YHGY428 (A–C) and YHGY209 (D–F), habitus. A, D. Dorsal; B, E. Ventral; C, F. Lateral. Scale bars: 1 mm. A B C D E F 223 ZooKeys 1255: 207–237 (2025), DOI: 10.3897/zookeys.1255.158340 Jianshuang Zhang et al.: A survey of Lipocrea from Guiyang dered with yellow lines and bearing a prominent large black spot anteriorly; laterally with two distinct yellow longitudinal lines, each line accompanied by approximately four small black spots; venter grayish, without distinct pattern; spinnerets yellow. Palp (Figs 8A–D, 9A–E). Cymbium (Cy) navicular, ~2.2× longer than wide, dorsally with sparse, long setae (all detached in ethanol), basoretrolaterally with a thumb-like paracymbium (Pc). Pc moderately large, about 1/5 length of cymbium, apex blunt, slightly curved and pointing retrolatero-distally. Tegulum (T) disc-shaped, slightly wider than cymbium, with distinct sperm duct along anterior margin, proximally covered by broad subtegulum (St). Tegular extension (TE) laminar, extending dorsally, almost completely concealed by conductor (C) in ventral view. St ~1/2 cymbium length, partly membranous, surface wrinkled and ribbed, with numerous diagonal ridges. Radix (R) leaf shaped, ~½ the width of the subtegulum length, distally with a triangular apophysis (RA). RA hyaline, nearly as long as radix, apex sharp and pointing distally. Median apophysis (MA) heavily sclerotized, located prolaterally to tegulum, consisting of a broad base and a hook-shaped process (HP); base navicular, ~2/5 the width of the subtegulum in length; HP nearly as long as base, apex sharp, distinctly curved and pointing retrolaterally. Terminal apophysis (TA) hidden behind tegulum, extending distally, apex surpassing the tegulum and bifurcating into two apophyses, forming a C-shape in anterior view; both terminal apophysis I (TA I) and terminal apophysis II (TA II) heavily sclerotized, with blunt apices pointing prolaterally; TA I relatively large, its length nearly equal to the width of the tegulum; TA II smaller and humble, ~½ the length of TAI. Terminal apophysis appendix (TAA) membranous, digitiform, accompanied by terminal apophysis, hidden behind tegulum, extending distally. C originating from dorsal-anterior portion of tegulum, proximally fused to weakly sclerotized TE; tip distinctly curved, shaped like an ox horn, with a sharp apex pointing dorso-distally. Embolus (Em) spine-shaped, nearly as long as the hook-shaped process of the median apophysis, originating centrally in anterior view, extending distally, surrounded by the RA, MA, TA, and C. Female (YHGY428). Total length 7.30. Carapace 2.83 long, 1.81 wide. Abdomen 4.96 long, 2.67 wide. Sternum 1.25 long and 0.84 wide. Labium 0.28 long and 0.49 wide. Endites 0.61 long and 0.43wide. Clypeus height 0.10. Both margins of chelicerae with four teeth. Eye sizes and interdistances: AME 0.15, ALE 0.13, PME 0.13, PLE 0.11, AME–AME 0.23, ALE–AME 0.19, PME–PME 0.02, PME–PLE 0.35. MOQ 0.44 long, anterior width 0.48, posterior width 0.27. Leg measurements: I 14.53 (3.33, 5.36, 4.54, 1.30), II 13.15 (3.26, 4.89, 3.86, 1.14), III 6.96 (2.31, 2.27, 1.58, 0.80), IV 11.80 (3.28, 4.39, 3.14, 0.99). Habitus (Fig. 7A–C, 10A). Similar to males, but the dorsum of the abdomen lacks the prominent anterior black spot. Genitalia (Fig. 10B–F). Epigyne strongly sclerotized with large postero-lateral lobes, distinctly wider than long, nearly trapeziform in ventral view and inverted triangular in lateral view. Knob-shaped projection (KP) represented by a small, partly membranous tubercle, located at anterior portion of epigynal plate. Copulatory openings (CO) large, located on the comma-shaped window (or pockets with chitinous posterior margins) is at the postero-lateral portion of the epigynal plate, separated by indistinct median septum (MS). Copulatory ducts (CD) short, diverging and ascending obliquely, forming V-shaped course 224 ZooKeys 1255: 207–237 (2025), DOI: 10.3897/zookeys.1255.158340 Jianshuang Zhang et al.: A survey of Lipocrea from Guiyang Figure 8. Male palp of the holotype of Lipocrea guiyang J. Zhang, Yu & Mi, sp. nov. A. Dorsal; B. Ventral; C. Prolateral; D. Retrolateral. Abbreviations: C = conductor; Cy = cymbium; Em = embolus; HP = hook-shaped process of MA; MA = median apophysis; Pc = paracymbium; R = radix; RA = radix apophysis; St = subtegulum; T = tegulum; TA I = terminal apophysis I; TA II = terminal apophysis II; TAA = terminal apophysis appendix; TE = tegular extension. Scale bar: 0.2 mm. A B C D 225 ZooKeys 1255: 207–237 (2025), DOI: 10.3897/zookeys.1255.158340 Jianshuang Zhang et al.: A survey of Lipocrea from Guiyang Figure 9. Male palpal bulb of the holotype of Lipocrea guiyang J. Zhang, Yu & Mi, sp. nov. A. Dorsal; B. Ventral; C. Prolateral; D. Retrolateral; E. Anterior. Abbreviations: C = conductor; Em = embolus; HP = hook-shaped process of MA; MA = median apophysis; R = radix; RA = radix apophysis; St = subtegulum; T = tegulum; TA I = terminal apophysis I; TA II = terminal apophysis II; TAA = terminal apophysis appendix; TE = tegular extension. Scale bar: 0.2 mm. A B C D E 226 ZooKeys 1255: 207–237 (2025), DOI: 10.3897/zookeys.1255.158340 Jianshuang Zhang et al.: A survey of Lipocrea from Guiyang Figure 10. Frontal view of cephalothorax (A.) and macerated epigyne (B–F) of the paratype (YHGY428) of Lipocrea guiyang J. Zhang, Yu & Mi, sp. nov. A. Female; B. Lateral; C. Ventral (blue dashed line showing the lateral and posterior margins of the epigynal base); D. Dorsal; E. Ventro-posterior; F. Dorso-anterior. Abbreviations: CD = copulatory duct; CO = copulatory opening; FD = fertilization duct; KP = knob-like projection; MS = median septum; Sp = spermatheca. Scale bars: 1 mm (A), 0.2 mm (B–F). A B C D E F 227 ZooKeys 1255: 207–237 (2025), DOI: 10.3897/zookeys.1255.158340 Jianshuang Zhang et al.: A survey of Lipocrea from Guiyang Figure 11. Epigyne of the paratype (YHGY209) of Lipocrea guiyang J. Zhang, Yu & Mi, sp. nov. A. Intact, ventral; B. Macerated, lateral; C. Macerated, ventral; D. Macerated, dorsal; E. Macerated, ventro-posterior; F. Macerated, dorso-anterior. Abbreviations: CD = copulatory duct; CO = copulatory opening; FD = fertilization duct; KP = knob-like projection; MS = median septum; Sp = spermatheca. Scale bar: 0.2 mm. A B C D E F 228 ZooKeys 1255: 207–237 (2025), DOI: 10.3897/zookeys.1255.158340 Jianshuang Zhang et al.: A survey of Lipocrea from Guiyang in dorsal view, finally entering anteriorly located spermathecae. Spermathecae (Sp) oval, ~1.2× longer than wide, relatively large, ~2/3 of epigyne length; two spermathecae close together, separated by ~2/3 of their width. Fertilization ducts (FD) membranous, relatively long, ~2/3 of spermathecae length, located on dorsal-basal surface of spermathecae. Distribution. Known only from the type locality (Fig. 1). Etymology. The specific epithet is derived from the name of the type locality; noun in apposition. Comments. In spite of the stable morphology of the male palp and the consistent coloration of the male habitus, considerable morphological variation is observed among female individuals, primarily related to epigynal structures. These variations involve features such as the presence or absence of a knob-shaped projection (KP), the shape of the copulatory openings (CO), and whether the median septum (MS) is distinct or indistinct. For example, in some females (e.g., YHGY428, as in Fig. 10B, C, E), the KP is distinct, the CO is situated on a comma-shaped window (or within pockets with chitinous posterior margins), and the MS is indistinct. In contrast, in other individuals (e.g., YHGY209, as in Fig. 11A–C, E), the KP is broken off, the CO is positioned on a nearly circular window (or within pockets bordered anteriorly, internally, and posteriorly), and the MS is distinct. In addition, some variation related to the abdominal pattern is also observed: the dorsum of the abdomen bears a median band extending along its entire length in some individuals (e.g., YHGY428), whereas in others (e.g., YHGY209), the median band is restricted to the posterior quarter of the dorsum (cf. Fig. 7A and Fig. 7D). However, the morphological variation was determined to be intraspecific variation based on the molecular species delimitation analysis. Lipocrea fusiformis (Thorell, 1877) Figs 1, 3, 12–14 Meta fusiformis Thorell, 1877: 431 (♀). Lipocrea fusiformis: Thorell 1878: 6. Larinia quadrinotata Simon, 1889: 340 (juv.); Simon 1909: 105 (♀). Larinia lutescens Thorell, 1898: 342 (♂♀). Larinopa fusiformis: Grasshoff 1970: 231, figs 15a, b, 16a–e (♂♀, transfer from Larinia, synonym of Larinia lutescens and L. quadrinotata); Tanikawa 1989: 35, figs 8–14 (♂♀); Chang and Tso 2004: 28, figs 5–8 (♂♀); Tanikawa et al. 2023: 56, figs 15–18. Note. For full list of taxonomic references, see WSC (2025). Material examined. • 1♂, 3♀ (YHGY432, YHGY495, YHGY507, YHGY508), China: Guizhou Prov.: Guiyang City, Huaxi District, University Town, 26.38°N, 106.65°E, c. 1173 m, by beating, 7 VII 2022, Q. Jiang & Q. Du leg; • 1♂, 1♀ (YHGY431, YHGY433), Guiyang City, Huaxi District, Dangwu Town, 26.38°N, 106.60°E, c. 1138 m, by hand, 19 V 2022, H. Yu & Q. Lu leg; Guiyang City, Kaiyang Co., Nanjiang Grand Canyon, 26.94°N, 106.97°E, c. 861 m, by hand, 7 VI 2022, H. Yu & Q. Lu leg. Diagnosis and description. See Tanikawa (1989). Living specimens as in Fig. 3A–C, male habitus as in Fig. 12A–C, male palp as in Fig. 13A–D, female habitus as in Fig. 12D–F, genitalia as in Fig. 14A–F. 229 ZooKeys 1255: 207–237 (2025), DOI: 10.3897/zookeys.1255.158340 Jianshuang Zhang et al.: A survey of Lipocrea from Guiyang Figure 12. Lipocrea fusiformis (Thorell, 1877), male (YHGY431, A–C) and female (YHGY432, D–F), habitus. A, D. Dorsal; B, E. Ventral; C, F. Lateral. Scale bars: 1 mm (A–C), 2 mm (D–F). A B C D E F 230 ZooKeys 1255: 207–237 (2025), DOI: 10.3897/zookeys.1255.158340 Jianshuang Zhang et al.: A survey of Lipocrea from Guiyang Figure 13. Male palp of Lipocrea fusiformis (Thorell, 1877). A. Dorsal; B. Ventral; C. Prolateral; D. Retrolateral. Abbreviations: C = conductor; Cy = cymbium; Em = embolus; HP = hook-shaped process of MA; MA = median apophysis; Pc = paracymbium; R = radix; RA = radix apophysis; St = subtegulum; T = tegulum; TA = terminal apophysis; TAA = terminal apophysis appendix; TE = tegular extension. Scale bar: 0.2 mm. A B C D 231 ZooKeys 1255: 207–237 (2025), DOI: 10.3897/zookeys.1255.158340 Jianshuang Zhang et al.: A survey of Lipocrea from Guiyang Figure 14 Macerated epigynes of females of Lipocrea fusiformis (Thorell, 1877), YHGY432 (A–C) and YHGY507 (D–F). A. Ventral, with scape; B. Lateral, with scape; C. Ventro-posterior, with scape; D. Ventral, without scape (blue dashed line showing the lateral and posterior margins of the epigynal base); E. Dorsal, without scape; F. Ventro-posterior, without scape. Abbreviations: CD = copulatory duct; CO = copulatory opening; FD = fertilization duct; MS = median septum; Sc = scape; Sp = spermatheca. Scale bars: 0.2 mm (A, B, D–F). A B C D E F