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Taxonomy of the Thelyphonid Genus Typopeltis Pocock, 1894, Including Homology Proposals for the Male Gonopod Structures (Arachnida, Thelyphonida, Typopeltinae)

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Taxonomy of the Thelyphonid Genus Typopeltis Pocock, 1894, Including Homology Proposals for the Male Gonopod Structures (Arachnida, Thelyphonida, Typopeltinae) A peer-reviewed open-access journal ZooKeys 848:Taxonomy 21–39 (2019) of the thelyphonid genus Typopeltis Pocock, 1894, including homology... 21 doi: 10.3897/zookeys.848.32263 RESEARCH ARTICLE http://zookeys.pensoft.net Launched to accelerate biodiversity research Taxonomy of the thelyphonid genus Typopeltis Pocock, 1894, including homology proposals for the male gonopod structures (Arachnida, Thelyphonida, Typopeltinae) Gabriel Seraphim1, Alessandro Ponce de Leão Giupponi1, Gustavo Silva de Miranda2 1 Laboratório de Referência Nacional em Vetores das Riquetsioses, LIRN-IOC-FIOCRUZ. Manguinhos, 21040-360, Rio de Janeiro, RJ, Brazil 2 Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington D.C., USA Corresponding author: Alessandro Ponce de Leão Giupponi ([email protected]) Academic editor: C. Haddad | Received 8 December 2018 | Accepted 12 March 2019 | Published 20 May 2019 http://zoobank.org/BC8282C2-C971-453F-B521-2A1BA06D77DA Citation: Seraphim G, Giupponi APL, Miranda GS (2019) Taxonomy of the thelyphonid genus Typopeltis Pocock, 1894, including homology proposals for the male gonopod structures (Arachnida, Thelyphonida, Typopeltinae). ZooKeys 848: 21–39. https://doi.org/10.3897/zookeys.848.32263 Abstract The genus Typopeltis Pocock, 1894 is poorly known regarding its systematics, natural history, and distribu- tion, despite important taxonomic advances during the 1990s. Currently, only 13 species are known from East Asia, including areas in south China, Japan, Vietnam, Laos, Thailand, and Taiwan. In this work, we describe and illustrate a new species of Typopeltis from Vietnam and provide a new description for the male of T. guangxiensis Haupt & Song, 1996. Additionally, we describe and illustrate the female gonopod of T. guangxiensis for the first time and propose a new homology hypothesis for the male gonopod parts. The male of T. laurentianus sp. n. is characterized by the unique patellar apophysis that presents a smooth texture and no spines. Typopeltis laurentianus sp. n. is the third species of this genus to be described from Vietnam. Keywords Asian fauna, taxonomy, Uropygi, Vinegaroons, whip scorpion Copyright Gabriel Seraphim et al. 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. 22 Gabriel Seraphim et al. / ZooKeys 848: 21–39 (2019) Introduction The order Thelyphonida Latreille, 1804 (also known as whip scorpions) is a conspicuous, yet small arachnid order with only 124 living species in 15 genera described so far (Zhang 2013, Barrales-Alcalá et al. 2018). Despite not being extremely diverse, the group is quite old and is estimated to have originated around 333 mya in tropical Pangea (Clouse et al. 2017). The oldest Thelyphonida fossil is from 318 mya and currently only seven fossils are known (Dunlop et al. 2008, Wolfe et al. 2016). Recent estimates of time divergence indicate an increase in diversification rates during the Cretaceous (Clouse et al. 2017). Thelyphonida is currently composed of one family (Thelyphonidae) and four sub- families: Hypoctoninae Pocock, 1899, Mastigoproctinae Speijer, 1933, Thelyphoninae Lucas, 1973 and Typopeltinae Rowland & Cooke, 1973. The classification history of the order goes back to Pocock (1899), who divided the family Thelyphonidae into two subfamilies, Thelyphoninae and Hypoctoninae (Pocock 1899). The two subfamilies were defined by the presence or absence of a keel between the medial and lateral eyes, respectively. Gravely (1916) set up an organization scheme that would become the cur- rent classification of the order, although some of the names were given only afterwards (such as Mastigoproctinae Speijer, 1933 and Typopeltinae Rowland & Cooke, 1973). Gravely (1916) divided Thelyphoninae into three groups, Mastigoproctinae and Ty- popeltinae, and a new one characterized by a strongly modified patellar process. This last group became the currently defined subfamily Typopeltinae, which includes the single genus Typopeltis Pocock, 1894 (Rowland and Cooke 1973). The genus Typopeltis is endemic to Asia, with 13 valid species (including the no- men dubium T. amurensis (Tarnani, 1889) from Russia). Six of these species are known from Southeast Asia: two from Vietnam (T. harmandi Kraepelin, 1900; T. soidaoensis Haupt, 1996), three from southern China (T. vanoorti (Speijer, 1936); T. sinensis (But- ler, 1872); T. guangxiensis Haupt & Song, 1996) and one from Laos (T. magnificus Haupt, 2004) (Haupt 1996, 2004a, Haupt and Song 1996, Harvey 2003). Several areas in East and Southeast Asia remain unsampled or undersampled, such as Cambo- dia and Thailand (with no records of Typopeltis at all), and Laos (with one record). Not only is there little information regarding the group’s distribution, but the systematics of Typopeltis is also still in its infancy. The greatest contributor to the understanding of the genus was Joachim Haupt (Haupt and Song 1996, Haupt 2004a, 2009); however, several details of the morphology of the species were not addressed and continue to be unknown, such as the form of the male gonopod. The genus Typopeltis can be easily recognized by the presence of a marked keel between the lateral and median eyes and by the absence of a suture dividing the ab- dominal tergites (Rowland and Cooke 1973). The males have a well-developed patel- lar apophysis and no projection on sternite III (Rowland and Cooke 1973). Females have modifications of the tarsomeres of leg I (antenniform), but according to Gravely (1916) this character can vary depending on the age and reproductive period of the specimen. In addition, the females have clear modifications of sternite II (genital plate) compared to males. The trochanter spines, despite being conspicuous, are not used in Taxonomy of the thelyphonid genus Typopeltis Pocock, 1894, including homology... 23 the taxonomy of the group because they vary considerably, with differences between the right and left pedipalps of a single individual (Gravely 1916). Not much is known regarding the phylogenetic relationships of Typopeltis species. In a molecular phylogeny of Thelyphonida, Clouse et al. (2017) included only one named species of Typopeltis, T. crucifer, which was recovered as sister to what was most likely an unnamed species in the same genus from Vietnam. Interestingly, Typopelti- nae was recovered as being more closely related to Thelyphoninae and Mastigoprocti- nae than to Hypoctoninae. In this work, we aim to contribute to the understanding of the morphological characters of Typopeltis by describing and illustrating a new species from Vietnam. In addition, we provide the first description of the male of T. guangxiensis Haupt & Song, 1996, provide detailed images of the female of that species, and present a hypothesis of homology of the male gonopod structures based on Giupponi and Kury (2013). Our homology hypothesis is made based on the consistent sister group relationship between Amblypygi and Thelyphonida (e.g. Ballesteros and Sharma 2019). With this, we intend to set the basis for the evaluation of new characters in future morphological phylogenetic studies. Material and methods Specimens were identified based on Rowland and Cooke (1973) and Haupt (1996). The description was adapted from Haupt (1996), Víquez and Armas (2007), Giupponi and Vasconcelos (2008), Villarreal and Giupponi (2009). The descriptions were made with NIKON SMZ745 and LEICA MZ15 stereomicroscopes. Photographs were made with a Leica M205C and Leica Application Suite V. 4.7 software. Scanning electron microscope (SEM) images were produced in a JEOL JSM-6390LV. The map was made with ArcGIS 10.3. All images have been edited with Adobe Photoshop CS6 and Adobe InDesing CS6. Acronyms: Fi = fistula;GO = genital operculum; LaM = lamina medialis; LoD = lobus dorsalis; LoL1 = lobus lateralis primus; LoL2 = lobus lateralis secundus; PI = processus inter- nus; Me = Mensa (new name); Fu = Fulcrum (new name); RS = receptaculum semi- nis; CCh = circulus chitinosus (new name); ACh = arcus chitinosus. MNRJ Museu Nacional, Rio de Janeiro, Brazil (the thelyphonid specimens were on loan from the collection when the Museum burned in 2018, so the material survided the incident; Dr. Adriano B. Kury); CAVAISC Coleção de Artrópodes Vetores Ápteros de Importância em Saúde das Co- munidades FIOCRUZ, Rio de Janeiro, Brazil (Dr. Marinete Amorim); MNHN Muséum national d’Histoire naturelle, Paris, France (Dr. Mark Judson); CAS California Academy of Sciences, San Francisco, USA (Dr. Darell Ubick). 24 Gabriel Seraphim et al. / ZooKeys 848: 21–39 (2019) Results Taxonomy Thelyphonidae Lucas, 1835 Typopeltinae Rowland & Cook, 1973 Typopeltis Pocock, 1894 Type species: Typopeltis: Typopeltis crucifer Pocock, 1894, by original designation. Gipopeltis: Typopeltis harmandi Kraepelin, 1900, by original designation. Teltus: Teltus vanoorti Speijer, 1936, by monotypy. List of Typopeltis species: 1 – T. amurensis (Tarnani, 1889) (Russia), nomen dubium; 2 – T. cantonensis Speijer, 1936 (China); 3 – T. crucifer Pocock, 1894 (Japan, Taiwan); 4 – T. dalyi Pocock, 1900 (Thailand); 5 – T. guangxiensis Haupt & Song, 1996 (China); 6 – T. harmandi Kraepelin, 1900 (Vietnam); 7 – T. kasnakowi Tarnani, 1900 (Thailand); 8 – T. mag- nificus Haupt, 2004 (Laos); 9 – T. sinensis (Butler, 1872) (China); 10 – T. soidaoensis Haupt, 1996 (Thailand, Vietnam); 11 –T. stimpsonii
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