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RESEARCH ARTICLE

An integrative approach reveals high species diversity in the primitively segmented spider genus Songthela (Mesothelae: Liphistiidae) from Hunan, China

Dengqing Li A , Zhaoyang Chen A , Fengxiang Liu B , Daiqin Li https://orcid.org/0000-0001-8269-7734 C * and Xin Xu https://orcid.org/0000-0001-5632-6622 A *
+ Author Affiliations
- Author Affiliations

A College of Life Sciences, Hunan Normal University, 36 Lushan Road, Changsha, 410081, Hunan Province, PR China.

B State Key Laboratory of Biocatalysis and Enzyme Engineering, and Centre for Behavioural Ecology and Evolution, School of Life Sciences, Hubei University, 368 Youyi Road, Wuhan, 430062, Hubei Province, PR China.

C Department of Biological Sciences, National University of Singapore, 14 Science Drive 4, Singapore 117543, Singapore.


Handling Editor: Mark Harvey

Invertebrate Systematics 36(2) 160-198 https://doi.org/10.1071/IS21058
Submitted: 17 August 2021  Accepted: 13 October 2021   Published: 22 February 2022

© 2022 The Author(s) (or their employer(s)). Published by CSIRO Publishing.

Abstract

Accurate species delimitation is crucial for our understanding of evolution, biodiversity and conservation. However, morphology-based species delimitation alone appears to be prone to taxonomic errors and ineffective for taxa with high interspecific morphological homogeneity or intraspecific morphological variations, as is the case for mesothele and mygalomorph spiders. Combined molecular–morphology species delimitation has shown great potential to delimit species boundaries in such ancient lineages. In the present study, molecular and morphological evidence were integrated to delimit species of the primitively segmented spider genus Songthela Ono, 2000. The cytochrome c oxidase subunit I gene (COI) was sequenced for 192 novel specimens belonging to 12 putative morphospecies. The evolutionary relationships within Songthela and the 12-morphospecies hypothesis were tested in two steps – species discovery and species validation – using four single-locus species delimitation approaches. All species delimitation analyses supported the 12-species hypothesis. Phylogenetic analyses yielded three major clades in Songthela, which are consistent with morphology. Accordingly, we assigned 19 known and 11 new species (S. aokoulong, sp. nov., S. bispina, sp. nov., S. dapo, sp. nov., S. huayanxi, sp. nov., S. lianhe, sp. nov., S. lingshang, sp. nov., S. multidentata, sp. nov., S. tianmen, sp. nov., S. unispina, sp. nov., S. xiujian, sp. nov., S. zizhu, sp. nov.) of Songthela to three species-groups: the bispina-group, the multidentata-group and the unispina-group. Another new species, S. zimugang, sp. nov., is not included in any species groups, but forms a sister lineage to the bispina- and unispina-groups. These results elucidate a high species diversity of Songthela in a small area and demonstrate that integrating morphology with COI-based species delimitation is fast and cost-effective in delimiting species boundaries.

http://zoobank.org/urn:lsid:zoobank.org:pub:AF0F5B31-AFAF-4861-9844-445AE8678B67

Keywords: COI, DNA barcoding, Liphistiidae, Mesothelae, new species, Songthela, species delimitation, taxonomy, trapdoor spiders.


References

Agnarsson, I, LeQuier, SM, Kuntner, M, Cheng, R-C, Coddington, JA, and Binford, G (2016). Phylogeography of a good Caribbean disperser, Argiope argentata (Araneae, Araneidae) and a new ‘cryptic’ species from Cuba. ZooKeys 625, 25–44.
Phylogeography of a good Caribbean disperser, Argiope argentata (Araneae, Araneidae) and a new ‘cryptic’ species from Cuba.Crossref | GoogleScholarGoogle Scholar |

Ahrens, D, Fujisawa, T, Krammer, H, Eberle, J, Fabrizi, S, and Vogler, AP (2016). Rarity and incomplete sampling in DNA-based species delimitation. Systematic Biology 65, 478–494.
Rarity and incomplete sampling in DNA-based species delimitation.Crossref | GoogleScholarGoogle Scholar | 26797695PubMed |

Bao, YH, Yin, CM, and Xu, X (2003). A new species of the genus Heptathela from China (Araneae, Liphistiidae). Acta Zootaxonomica Sinica 28, 459–460.

Bond, JE, and Stockman, AK (2008). An integrative method for delimiting cohesion species, finding the population-species interface in a group of Californian trapdoor spiders with extreme genetic divergence and geographic structuring. Systematic Biology 57, 628–646.
An integrative method for delimiting cohesion species, finding the population-species interface in a group of Californian trapdoor spiders with extreme genetic divergence and geographic structuring.Crossref | GoogleScholarGoogle Scholar | 18686196PubMed |

Cabra-García, J, and Hormiga, G (2020). Exploring the impact of morphology, multiple sequence alignment and choice of optimality criteria in phylogenetic inference, a case study with the Neotropical orb-weaving spider genus Wagneriana (Araneae, Araneidae). Zoological Journal of the Linnean Society 188, 976–1151.
Exploring the impact of morphology, multiple sequence alignment and choice of optimality criteria in phylogenetic inference, a case study with the Neotropical orb-weaving spider genus Wagneriana (Araneae, Araneidae).Crossref | GoogleScholarGoogle Scholar |

Camargo A, Sites JJ (2013) Species delimitation, a decade after the renaissance. In ‘The Species Problem – Ongoing Issues’. (Ed. IY Pavlinov) pp. 225–247. (InTech)

Chen, XE, Gao, JC, Zhu, CD, and Luo, ZM (1988). A new species of the genus Heptathela from China (Araneae: Heptathelidae). Sichuan Journal of Agricultural Science 3, 78–81.

Chen, ZF, Zhang, ZH, and Zhu, CD (1981). A new species of genus Heptathela. Journal of Hangzhou University, Natural Science Edition 8, 305–308.

Chen, ZY, Li, DQ, Li, D, and Xu, X (2021). Three new species of the primitively segmented spider genus Songthela (Araneae, Mesothelae) from Guizhou Province, China. ZooKeys 1037, 57–71.
Three new species of the primitively segmented spider genus Songthela (Araneae, Mesothelae) from Guizhou Province, China.Crossref | GoogleScholarGoogle Scholar |

Darriba, D, Taboada, GL, Doallo, R, and Posada, D (2012). jModelTest 2: more models, new heuristics and parallel computing. Nature Methods 9, 772.
jModelTest 2: more models, new heuristics and parallel computing.Crossref | GoogleScholarGoogle Scholar | 22847109PubMed |

Dellicour, S, and Flot, JF (2018). The hitchhiker’s guide to single-locus species delimitation. Molecular Ecology Resources 18, 1234–1246.
The hitchhiker’s guide to single-locus species delimitation.Crossref | GoogleScholarGoogle Scholar | 29847023PubMed |

Eberle, J, Ahrens, D, Mayer, C, Niehuis, O, and Misof, B (2020). A plea for standardized nuclear markers in metazoan DNA taxonomy. Trends in Ecology & Evolution 35, 336–345.
A plea for standardized nuclear markers in metazoan DNA taxonomy.Crossref | GoogleScholarGoogle Scholar |

Folmer, O, Black, M, Hoeh, W, Lutz, R, and Vrijenhoek, R (1994). DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology 3, 294–299.
| 7881515PubMed |

Fouquet, A, Gilles, A, Vences, M, Marty, C, Blanc, M, and Gemmell, NJ (2007). Underestimation of species richness in Neotropical frogs revealed by mtDNA analyses. PLoS One 2, e1109.
Underestimation of species richness in Neotropical frogs revealed by mtDNA analyses.Crossref | GoogleScholarGoogle Scholar | 17971872PubMed |

Gertsch, WJ (1967). A new Liphistiid spider from China (Araneae: Liphistiidae). Journal of the New York Entomological Society 75, 114–118.

Hamilton, CA, Formanowicz, DR, and Bond, JE (2011). Species delimitation and phylogeography of Aphonopelma hentzi (Araneae, Mygalomorphae, Theraphosidae): cryptic diversity in North American tarantulas. PLoS One 6, e26207.
Species delimitation and phylogeography of Aphonopelma hentzi (Araneae, Mygalomorphae, Theraphosidae): cryptic diversity in North American tarantulas.Crossref | GoogleScholarGoogle Scholar | 22022570PubMed |

Hamilton, CA, Hendrixson, BE, Brewer, MS, and Bond, JE (2014). An evaluation of sampling effects on multiple DNA barcoding methods leads to an integrative approach for delimiting species: a case study of the North American tarantula genus Aphonopelma (Araneae, Mygalomorphae, Theraphosidae). Molecular Phylogenetics and Evolution 71, 79–93.
An evaluation of sampling effects on multiple DNA barcoding methods leads to an integrative approach for delimiting species: a case study of the North American tarantula genus Aphonopelma (Araneae, Mygalomorphae, Theraphosidae).Crossref | GoogleScholarGoogle Scholar | 24280211PubMed |

Harrington, RC, and Near, TJ (2012). Phylogenetic and coalescent strategies of species delimitation in snubnose darters (Percidae, Etheostoma). Systematic Biology 61, 63–79.
Phylogenetic and coalescent strategies of species delimitation in snubnose darters (Percidae, Etheostoma).Crossref | GoogleScholarGoogle Scholar | 21828082PubMed |

Harvey, MS (2002). Short-range endemism amongst the Australian fauna: some examples from non-marine environments. Invertebrate Systematics 16, 555–570.
Short-range endemism amongst the Australian fauna: some examples from non-marine environments.Crossref | GoogleScholarGoogle Scholar |

Haupt, J (1983). Vergleichende Morphologie der Genitalorgane und Phylogenie der liphistiomorphen Webspinnen (Araneae: Mesothelae). I. Revision der bisher bekannten Arten. Journal of Zoological Systematics and Evolutionary Research 21, 275–293.
Vergleichende Morphologie der Genitalorgane und Phylogenie der liphistiomorphen Webspinnen (Araneae: Mesothelae). I. Revision der bisher bekannten Arten.Crossref | GoogleScholarGoogle Scholar |

Haupt, J (2003). The Mesothelae – monograph of an exceptional group of spiders (Araneae, Mesothelae): morphology, behaviour, ecology, taxonomy, distribution and phylogeny. Zoologica 154, 1–102.

Hebert, PDN, Cywinska, A, Ball, SL, and deWaard, JR (2003). Biological identifications through DNA barcodes. Proceedings of the Royal Society of London – B. Biological Sciences 270, 313–321.
Biological identifications through DNA barcodes.Crossref | GoogleScholarGoogle Scholar |

Hedin, M, Carlson, D, and Coyle, F (2015). Sky island diversification meets the multispecies coalescent – divergence in the spruce-fir moss spider (Microhexura montivaga, Araneae, Mygalomorphae) on the highest peaks of southern Appalachia. Molecular Ecology 24, 3467–3484.
Sky island diversification meets the multispecies coalescent – divergence in the spruce-fir moss spider (Microhexura montivaga, Araneae, Mygalomorphae) on the highest peaks of southern Appalachia.Crossref | GoogleScholarGoogle Scholar | 26011071PubMed |

Hofmann, EP, Nicholson, KE, Luque-Montes, IR, Köhler, G, Cerrato-Mendoza, CA, Medina-Flores, M, Wilson, LD, and Townsend, JH (2019). Cryptic diversity, but to what extent? Discordance between single-locus species delimitation methods within mainland anoles (Squamata, Dactyloidae) of northern central America. Frontiers in Genetics 10, 11.
Cryptic diversity, but to what extent? Discordance between single-locus species delimitation methods within mainland anoles (Squamata, Dactyloidae) of northern central America.Crossref | GoogleScholarGoogle Scholar | 30804976PubMed |

Hundsdoerfer, AK, Lee, KM, Kitching, IJ, and Mutanen, M (2019). Genome-wide SNP data reveal an overestimation of species diversity in a group of hawkmoths. Genome Biology and Evolution 11, 2136–2150.
Genome-wide SNP data reveal an overestimation of species diversity in a group of hawkmoths.Crossref | GoogleScholarGoogle Scholar | 31143925PubMed |

Kapli, P, Lutteropp, S, Zhang, J, Kobert, K, Pavlidis, P, Stamatakis, A, and Flouri, T (2017). Multi-rate poisson tree processes for single-locus species delimitation under maximum likelihood and Markov Chain Monte Carlo. Bioinformatics 33, 1630–1638.
Multi-rate poisson tree processes for single-locus species delimitation under maximum likelihood and Markov Chain Monte Carlo.Crossref | GoogleScholarGoogle Scholar | 28108445PubMed |

Kishida, K (1923). Heptathela, a new genus of liphistiid spiders. Annotationes Zoologicae Japonenses 10, 235–242.

Klimov, PB, Skoracki, M, and Bochkov, AV (2019). Cox1 barcoding versus multilocus species delimitation, validation of two mite species with contrasting effective population sizes. Parasites & Vectors 12, 8.
Cox1 barcoding versus multilocus species delimitation, validation of two mite species with contrasting effective population sizes.Crossref | GoogleScholarGoogle Scholar |

Kumar, S, Stecher, G, Li, M, Knyaz, C, and Tamura, K (2018). MEGA X: Molecular evolutionary genetics analysis across computing platforms. Molecular Biology and Evolution 35, 1547–1549.
MEGA X: Molecular evolutionary genetics analysis across computing platforms.Crossref | GoogleScholarGoogle Scholar | 29722887PubMed |

Leavitt, DH, Starrett, J, Westphal, MF, and Hedin, M (2015). Multilocus sequence data reveal dozens of putative cryptic species in a radiation of endemic Californian mygalomorph spiders (Araneae, Mygalomorphae, Nemesiidae). Molecular Phylogenetics and Evolution 91, 56–67.
Multilocus sequence data reveal dozens of putative cryptic species in a radiation of endemic Californian mygalomorph spiders (Araneae, Mygalomorphae, Nemesiidae).Crossref | GoogleScholarGoogle Scholar | 26025426PubMed |

Li, DQ, Liu, H, and Xu, X (2019). Three new species of the primitively segmented spiders from Mt Yuelu, Hunan, China (Mesothelae, Liphistiidae). Zootaxa 4691, 139–152.
Three new species of the primitively segmented spiders from Mt Yuelu, Hunan, China (Mesothelae, Liphistiidae).Crossref | GoogleScholarGoogle Scholar |

Li, DQ, Liu, FX, Li, D, and Xu, X (2020). Two new species of the primitively segmented spider genus Songthela from Hunan Province, China (Mesothelae, Liphistiidae). ZooKeys 937, 1–19.
Two new species of the primitively segmented spider genus Songthela from Hunan Province, China (Mesothelae, Liphistiidae).Crossref | GoogleScholarGoogle Scholar |

Luo, A, Ling, C, Ho, SYW, and Zhu, CD (2018). Comparison of methods for molecular species delimitation across a range of speciation scenarios. Systematic Biology 67, 830–846.
Comparison of methods for molecular species delimitation across a range of speciation scenarios.Crossref | GoogleScholarGoogle Scholar | 29462495PubMed |

Masters, BC, Fan, V, and Ross, HA (2011). Species delimitation – a GENEIOUS plugin for the exploration of species boundaries. Molecular Ecology Resources 11, 154–157.
Species delimitation – a GENEIOUS plugin for the exploration of species boundaries.Crossref | GoogleScholarGoogle Scholar | 21429114PubMed |

Mateos, E, Winkler, D, Riutort, M, and Álvarez-Presas, M (2021). New morphological and molecular data reveal an important underestimation of species diversity and indicate evolutionary patterns in European Lepidocyrtus (Collembola: Entomobryidae). Invertebrate Systematics 35, 471–492.
New morphological and molecular data reveal an important underestimation of species diversity and indicate evolutionary patterns in European Lepidocyrtus (Collembola: Entomobryidae).Crossref | GoogleScholarGoogle Scholar |

McGuire, JA, Linkem, CW, Koo, MS, Hutchison, DW, Lappin, AK, Orange, DI, Lemos-Espinal, J, Riddle, BR, and Jaeger, JR (2007). Mitochondrial introgression and incomplete lineage sorting through space and time: Phylogenetics of crotaphytid lizards. Evolution 61, 2879–2897.
Mitochondrial introgression and incomplete lineage sorting through space and time: Phylogenetics of crotaphytid lizards.Crossref | GoogleScholarGoogle Scholar | 17941840PubMed |

Minh, BQ, Schmidt, HA, Chernomor, O, Schrempf, D, Woodhams, MD, von Haeseler, A, and Lanfear, R (2020). IQ-TREE 2: new models and efficient methods for phylogenetic inference in the genomic era. Molecular Biology and Evolution 37, 1530–1534.
IQ-TREE 2: new models and efficient methods for phylogenetic inference in the genomic era.Crossref | GoogleScholarGoogle Scholar | 32011700PubMed |

Moritz, C, and Cicero, C (2004). DNA barcoding, promise and pitfalls. PLoS Biology 2, e354.
DNA barcoding, promise and pitfalls.Crossref | GoogleScholarGoogle Scholar | 15486587PubMed |

Ono, H (1999). Spiders of the genus Heptathela (Araneae, Liphistiidae) from Vietnam, with notes on their natural history. Journal of Arachnology 27, 37–43.

Ono, H (2000). Zoogeographic and taxonomic notes on spiders of the subfamily Heptathelinae (Araneae, Mesothelae, Liphistiidae). Memoirs of the National Science Museum Tokyo – A. Zoology 33, 145–151.

Ono, H (2010). Four new spiders (Arachnida, Araneae) of the families Liphistiidae, Ctenizidae, Araneidae and Ctenidae from Vietnam. Memoirs of the National Museum of Nature and Science Tokyo 46, 1–12.

Pocock, RI (1892). XXXVIII. – Liphistius and its bearing upon the classification of spiders. The Annals and Magazine of Natural History Series 6, 306–314.
XXXVIII. – Liphistius and its bearing upon the classification of spiders.Crossref | GoogleScholarGoogle Scholar |

Puillandre, N, Lambert, A, Brouillet, S, and Achaz, G (2012). ABGD, automatic barcode gap discovery for primary species delimitation. Molecular Ecology 21, 1864–1877.
ABGD, automatic barcode gap discovery for primary species delimitation.Crossref | GoogleScholarGoogle Scholar | 21883587PubMed |

Rambaut, A, Drummond, AJ, Xie, D, Baele, G, and Suchard, MA (2018). Posterior summarization in bayesian phylogenetics using Tracer 1.7. Systematic Biology 67, 901–904.
Posterior summarization in bayesian phylogenetics using Tracer 1.7.Crossref | GoogleScholarGoogle Scholar | 29718447PubMed |

Renner, MAM, Heslewood, MM, Patzak, SDF, Schäfer-Verwimp, A, and Heinrichs, J (2017). By how much do we underestimate species diversity of liverworts using morphological evidence? An example from Australasian Plagiochila (Plagiochilaceae: Jungermanniopsida). Molecular Phylogenetics and Evolution 107, 576–593.
By how much do we underestimate species diversity of liverworts using morphological evidence? An example from Australasian Plagiochila (Plagiochilaceae: Jungermanniopsida).Crossref | GoogleScholarGoogle Scholar | 28007566PubMed |

Ronquist, F, Teslenko, M, van der Mark, P, Ayres, DL, Darling, A, Höhna, S, Larget, B, Liu, L, Suchard, MA, and Huelsenbeck, JP (2012). MrBayes 3.2, efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61, 539–542.
MrBayes 3.2, efficient Bayesian phylogenetic inference and model choice across a large model space.Crossref | GoogleScholarGoogle Scholar | 22357727PubMed |

Satler, JD, Carstens, BC, and Hedin, M (2013). Multilocus species delimitation in a complex of morphologically conserved trapdoor spiders (Mygalomorphae, Antrodiaetidae, Aliatypus). Systematic Biology 62, 805–823.
Multilocus species delimitation in a complex of morphologically conserved trapdoor spiders (Mygalomorphae, Antrodiaetidae, Aliatypus).Crossref | GoogleScholarGoogle Scholar | 23771888PubMed |

Song, DX, and Haupt, J (1984). Comparative morphology and phylogeny of liphistiomorph spiders (Araneae: Mesothelae). 2. Revision of new Chinese heptathelid species. Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg (NF) 27, 443–451.

Song, DX, and Wu, KY (1997). On a new species of the genus Heptathela (Araneae: Liphistiidae) from Hong Kong, China. Chinese Journal of Zoology 32, 1–3.

Thorell, T (1869). On European spiders. Part I. Review of the European genera of spiders, preceded by some observations on zoological nomenclature. Nova Acta Regiae Societatis Scientiarum Upsaliensis 7, 1–108.

Wang, HZ, and Jiao, YC (1995). A new species of the family Heptathelidae in China. Journal of Yunnan Normal University 15, 80–81.

Wheeler WC, Giribet G, Edgecombe GD (2004) Arthropod systematics, the comparative study of genomic, anatomical, and paleontological information. In ‘Assembling the Tree of Life’. (Eds J Cracraft, MJ Donoghue) pp. 281–295. (Oxford University Press: New York, NY, USA)

Wilson, JD, Hughes, JM, Raven, RJ, Rix, MG, and Schmidt, DJ (2018). Spiny trapdoor spiders (Euoplos) of eastern Australia: Broadly sympatric clades are differentiated by burrow architecture and male morphology. Molecular Phylogenetics and Evolution 122, 157–165.
Spiny trapdoor spiders (Euoplos) of eastern Australia: Broadly sympatric clades are differentiated by burrow architecture and male morphology.Crossref | GoogleScholarGoogle Scholar | 29428510PubMed |

Xu, X, and Yin, CM (2001). A new species of the genus Heptathela from China (Araneae, Liphistiidae). Acta Arachnologica Sinica 10, 8–10.

Xu, X, Liu, F, Chen, J, Li, D, and Kuntner, M (2015a). Integrative taxonomy of the primitively segmented spider genus Ganthela (Araneae: Mesothelae: Liphistiidae): DNA barcoding gap agrees with morphology. Zoological Journal of the Linnean Society 175, 288–306.
Integrative taxonomy of the primitively segmented spider genus Ganthela (Araneae: Mesothelae: Liphistiidae): DNA barcoding gap agrees with morphology.Crossref | GoogleScholarGoogle Scholar |

Xu, X, Liu, F, Chen, J, Ono, H, Li, D, and Kuntner, M (2015b). A genus-level taxonomic review of primitively segmented spiders (Mesothelae, Liphistiidae). ZooKeys 488, 121–151.
A genus-level taxonomic review of primitively segmented spiders (Mesothelae, Liphistiidae).Crossref | GoogleScholarGoogle Scholar |

Xu, X, Liu, F, Cheng, R-C, Chen, J, Xu, X, Zhang, ZS, Ono, H, Pham, DS, Norma-Rashid, Y, Arnedo, MA, Kuntner, M, and Li, D (2015c). Extant primitively segmented spiders have recently diversified from an ancient lineage. Proceedings of the Royal Society of London – B. Biological Sciences 282, 20142486.
Extant primitively segmented spiders have recently diversified from an ancient lineage.Crossref | GoogleScholarGoogle Scholar |

Xu, X, Liu, F, Chen, J, Ono, H, Agnarsson, I, Li, D, and Kuntner, M (2016). Pre-Pleistocene geological events shaping diversification and distribution of primitively segmented spiders on East Asian Margins. Journal of Biogeography 43, 1004–1019.
Pre-Pleistocene geological events shaping diversification and distribution of primitively segmented spiders on East Asian Margins.Crossref | GoogleScholarGoogle Scholar |

Xu, X, Liu, F, Ono, H, Chen, J, Kuntner, M, and Li, D (2017). Targeted sampling in Ryukyus facilitates species delimitation of the primitively segmented spider genus Ryuthela (Araneae: Mesothelae: Liphistiidae). Zoological Journal of the Linnean Society 181, 867–909.
Targeted sampling in Ryukyus facilitates species delimitation of the primitively segmented spider genus Ryuthela (Araneae: Mesothelae: Liphistiidae).Crossref | GoogleScholarGoogle Scholar |

Xu, X, Kuntner, M, Liu, F, Chen, J, and Li, D (2018). Formation of rivers and mountains drives diversification of primitively segmented spiders in continental East Asia. Journal of Biogeography 45, 2080–2091.
Formation of rivers and mountains drives diversification of primitively segmented spiders in continental East Asia.Crossref | GoogleScholarGoogle Scholar |

Xu, X, Ono, H, Kuntner, M, Liu, FX, and Li, D (2019). A taxonomic monograph of the Liphistiid spider genus Heptathela, endemic to Japanese islands. ZooKeys 888, 1–50.
A taxonomic monograph of the Liphistiid spider genus Heptathela, endemic to Japanese islands.Crossref | GoogleScholarGoogle Scholar | 31754317PubMed |

Xu, X, Kuntner, M, Bond, JE, Ono, H, Ho, SYW, Liu, F, Yu, L, and Li, D (2020). Molecular species delimitation in the primitively segmented spider genus Heptathela endemic to Japanese islands. Molecular Phylogenetics and Evolution 151, 106900.
Molecular species delimitation in the primitively segmented spider genus Heptathela endemic to Japanese islands.Crossref | GoogleScholarGoogle Scholar | 32599078PubMed |

Xu, X, Su, YC, Ho, SYW, Kuntner, M, Ono, H, Liu, FX, Chang, CC, Warrit, N, Sivayyapram, V, Aung, KP, Pham, DS, Norma-Rashid, Y, and Li, D (2021). Phylogenomic analysis of ultraconserved elements resolves the evolutionary and biogeographic history of segmented trapdoor spiders. Systematic Biology 70, 1110–1122.
Phylogenomic analysis of ultraconserved elements resolves the evolutionary and biogeographic history of segmented trapdoor spiders.Crossref | GoogleScholarGoogle Scholar | 33367903PubMed |

Yin, CM (2001). A new species of the genus Heptathela and its variant type from China (Araneae: Liphistiidae). Dong Wu Fen Lei Xue Bao 26, 297–300.

Yin, CM, Tang, G, Zhao, JZ, and Chen, J (2002). Two new species of the genus Heptathela from China (Araneae: Liphistiidae). Acta Arachnologica Sinica 11, 18–21.

Yin, CM, Tang, G, and Xu, X (2003). Two new species of the genus Heptathela from China (Araneae: Liphistiidae). Acta Arachnologica Sinica 12, 1–5.

Yu, H, Li, F, and Zhang, JS (2018). A new species of the genus Songthela from Guizhou Province China (Araneae: Mesothelae: Liphistiidae). Turkish Journal of Zoology 42, 715–721.
A new species of the genus Songthela from Guizhou Province China (Araneae: Mesothelae: Liphistiidae).Crossref | GoogleScholarGoogle Scholar |