Redefining the Mexican genus Diguetinus Roewer, 1912 (Opiliones, Phalangioidea, Globipedidae) based on morphology and molecular phylogenetic data, with the redescription of Diguetinus spinulatus (Banks, 1898) stat. restit.

Directeur De la publication / Publication director : Gilles Bloch Président du Muséum national d’Histoire naturelle réDactrice en chef / editor-in-chief : Laure Desutter-Grandcolas assistante De réDaction / assistant editor : Anne Mabille ([email protected]) Mise en page / Page layout : Anne Mabille coMité scientifique / scientific board : Nesrine Akkari (Naturhistorisches Museum, Vienne, Autriche) Maria Marta Cigliano (Museo de La Plata, La Plata, Argentine) Serge Gofas (Universidad de Málaga, Málaga, Espagne) Sylvain Hugel (CNRS, Université de Strasbourg, France) Marco Isaia (Università degli Studi di Torino, Turin, Italie) Rafael Marquez (CSIC, Madrid, Espagne) Jose Christopher E. Mendoza (Lee Kong Chian Natural History Museum, Singapour) Annemarie Ohler (MNHN, Paris, France) Jean-Yves Rasplus (INRA, Montferrier-sur-Lez, France) Wanda M. Weiner (Polish Academy of Sciences, Cracovie, Pologne) couverture / cover : Living female of Diguetinus raptator Roewer, 1912. 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Diffusion – Publications scientifiques Muséum national d’Histoire naturelle CP 41 – 57 rue Cuvier F-75231 Paris cedex 05 (France) Tél. : 33 (0)1 40 79 48 05 / Fax : 33 (0)1 40 79 38 40 [email protected] / https://sciencepress.mnhn.fr © Publications scientifiques du Muséum national d’Histoire naturelle, Paris, 2025 ISSN (imprimé / print) : 1280-9551/ ISSN (électronique / electronic) : 1638-9387 731 ZOOSYSTEMA • 2025 • 47 (30) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.zoosystema.com urn:lsid:zoobank.org:pub:203D4929-F31D-443F-94F6-4CCC15B2D8CD Ochoa-Vázquez D., Cruz-López J. A., Rosas-Valdez R. & Martínez-Salazar E. A. 2025. — Redefining the Mexican genus Diguetinus Roewer, 1912 (Opiliones, Phalangioidea, Globipedidae) based on morphology and molecular phylogenetic data, with the redescription of Diguetinus spinulatus (Banks, 1898) stat. restit. Zoosystema 47 (30): 731-749. https:// doi.org/10.5252/zoosystema2025v47a30. http://zoosystema.com/47/30 ABSTRACT The genus Diguetinus Roewer, 1912 (Opiliones, Globipedidae) is a monotypic genus that includes the widespread species Diguetinus raptator Roewer, 1912 from Mexico. Some studies have suggested that additional species could be included within the genus, particularly Metopilio spinulatus (Banks, 1898). This species has a brief but complicated taxonomic history, being originally described in Leptobunus Banks, 1893, considered in Hadrobunus Banks, 1900, later transferred to Diguetinus and currently in Metopilio Roewer, 1911, all changes without a solid justification. The aim of this study was to conduct a morphological revision of D. raptator and M. spinulatus using specimens from their respective type localities. In addition, a phylogenetic hypothesis that includes D. raptator and M. spinulatus, was inferred based on partial sequences of the mitochondrial gene cytochrome c oxidase subunit I (COI) gene. Our results indicate that D. raptator and M. spinulatus are related in a clade that leads to the phylogenetic redefinition of Diguetinus. Accordingly, the following taxonomic changes are proposed: a) a new diagnosis of Diguetinus; and b) the redescription of Diguetinus spinulatus stat. restit. (♂, ♀). Also, comparative detailed illustrations of the male and female genitalia of these species are provided, including taxonomic comments on the use of previous characters for the diagnosis of Diguetinus and Metopilio. Finally, we provide evidence of intrasexual dimorphism in males of D. raptator. Daniel OCHOA-VÁZQUEZ Doctorado en Ciencias Básicas, Unidad Académica de Ciencias Biológicas, Universidad Autónoma de Zacatecas, Av. Preparatoria s/n, Campus Universitario II, Col. Agronómica, CP. 98066. Zacatecas, Zacatecas (Mexico) and Laboratorio de Colecciones Biológicas y Sistemática Molecular, Unidad Académica de Ciencias Biológicas, Universidad Autónoma de Zacatecas, Av. Preparatoria s/n, Campus Universitario II, Col. Agronómica, CP. 98066. Zacatecas, Zacatecas (Mexico) [email protected] Jesús A. CRUZ-LÓPEZ Instituto Nacional de Investigaciones Forestales, Agrícolas y Pecuarias (INIFAP), Centro de Investigación Regional Pacífico Sur (CIRPAS), Campo Experimental Valles Centrales de Oaxaca, Melchor Ocampo 7, CP. 68200, Santo Domingo Barrio Bajo, Villa de Etla, Oaxaca (Mexico) [email protected] Rogelio ROSAS-VALDEZ Elizabeth A. MARTÍNEZ-SALAZAR* Laboratorio de Colecciones Biológicas y Sistemática Molecular, Unidad Académica de Ciencias Biológicas, Universidad Autónoma de Zacatecas, Avenida Preparatoria s/n, Campus Universitario II, Col. Agronómica, CP. 98066. Zacatecas, Zacatecas (Mexico) rogr[email protected]; [email protected] (*corresponding author) Submitted on 21 December 2024 | Accepted on 30 April 2025 | Published on 28 November 2025 Redefining the Mexican genus Diguetinus Roewer, 1912 (Opiliones, Phalangioidea, Globipedidae) based on morphology and molecular phylogenetic data, with the redescription of Diguetinus spinulatus (Banks, 1898) stat. restit. KEY WORDS Emended diagnosis, Harvestmen, alpha male, Diguetinus raptator, COI barcoding. 732 ZOOSYSTEMA • 2025 • 47 (30) Ochoa-Vázquez D. et al. RÉSUMÉ Redéfinition du genre mexicain Diguetinus Roewer, 1912 (Opiliones, Phalangioidea, Globipedidae) à partir de la morphologie et de données phylogénétiques moléculaires, avec redéfinition de Diguetinus spinulatus (Banks, 1898) stat. restit. Le genre Diguetinus Roewer, 1912 (Opiliones, Globipedidae) est un genre monotypique qui comprend l’espèce Diguetinus raptator Roewer, 1912, largement répandue au Mexique. Cependant, certaines études ont suggéré que d’autres espèces pourraient être incluses dans le genre, en particulier Metopilio spinulatus (Banks, 1898). Cette espèce a une histoire taxonomique brève mais compliquée: elle a été décrite dans Leptobunus Banks, 1893, puis considérée comme faisant partie de Hadrobunus Banks, 1900, avant d’être transférée dans Diguetinus et actuellement dans Metopilio Roewer, 1911, tous ces changements sans justification solide. L’objectif de cette étude était d’effectuer une révision morphologique de D. raptator et de M. spinulatus à l’aide de spécimens provenant de leurs localités types respectives. En outre, une hypothèse phylogénétique incluant D. raptator et M.spinulatus a été inférée à partir de séquences partielles du gène mitochondrial cytochrome c oxydase subunit I (COI). Nos résultats indiquent que D. raptator et M. spinulatus sont apparentés dans un clade qui conduit à la redéfinition phylogénétique de Diguetinus. En fonction des résultats, les changements taxonomiques suivants sont proposés: a) une nouvelle diagnose du genre Diguetinus ; et b) la redescription de Diguetinus spinulatus stat. restit. (♂, ♀). Des illustrations détaillées et comparatives des genitalia mâles et femelles de ces espèces sont également fournies, accompagnées de commentaires taxonomiques sur l’utilisation des caractères précédemment utilisés pour la diagnose du genre Diguetinus et Metopilio. Enfin, nous apportons la preuve d’un dimorphisme intrasexuel chez les mâles de D. raptator. MOTS CLÉS Diagnose amendée, opilions, mâle alpha, Diguetinus raptator, code-barres COI. INTRODUCTION The order Opiliones Sundevall, 1833 remains understudied in certain geographic regions despite its status as one of the most species-rich groups of arachnids (Kury etal. 2021). Hedin etal. (2012) highlighted the taxonomic uncertainty in certain Mexican groups, due to the limited knowledge of the diversity of Opiliones in Mexico and the scarcity of taxonomic literature, sometimes complicated by inadequate original descriptions and a lack of study of genitalia (Kury& Cokendolpher 2000; Hedin etal. 2012). One of these groups is the recently erected family Globipedidae Kury& Cokendolpher, 2020, previously referred to as the Metopilio-group (Giribet etal. 2002, 2010; Hedin etal. 2012). This family currently comprises 33 species, classified in six genera: Dalquestia Cokendolpher, 1984; Diguetinus Roewer, 1912; Eurybunus Banks, 1893; Globipes Banks, 1893; Lanthanopilio Cokendolpher and Cokendolpher, 1984; and Metopilio Roewer, 1911(Kury etal. 2021). The geographical distribution of globipedids includes Mexico, Central America, and the southwestern United States, extending as far south as Chiriqui in Panama (Kury& Cokendolpher 2020). The taxonomic history of this group is intricate, early authors considered it either as an undescribed subfamily in Sclerosomatidae Simon, 1879, as part of the dissolved Leptobunidae Banks, 1901, or and undescribed subor family assemblage (Gruber 1969; Cokendolpher 1984a; Cokendolpher& Cokendolpher 1984; Crawford 1992). Subsequent phylogenetic research has recovered this group either as monoor polyphyletic group within Phalangioidea Latreille, 1802 (Giribet etal. 2002; 2010; Hedin etal. 2012). Hedin etal. (2012) recovered the Metopilio-group as monophyletic including three species: Dalquestia grasshoffi Cokendolpher, 1984, Eurybunus brunneus Banks, 1893 and Globipes simplex (Schenkel, 1951), being either sister to Neopilionidae Lawrence, 1931 or Phalangiidae Latreille, 1802, both hypotheses with weak support. Recently, Derkarabetian etal. (2023) included Metopiliosp. in a phylogenomic analysis, and retrieved Globipedidae as the sister group of Phalangiidae. So far, there have been no published molecular data for the remaining Globipedidae genera, such as Lanthanopilio or Diguetinus. The genus Diguetinus currently includes only the species Diguetinus raptator Roewer, 1912, which has a widespread distribution in central Mexico. In the recent revision of the genus, Cokendolpher etal. (2021) mentioned that there is the possibility that members of other genera (e.g., Metopilio) could be included within Diguetinus. One of these species is the enigmatic Metopilio spinulatus (Banks, 1898), which origi - nally was described in Leptobunus Banks, 1893, considered in Hadrobunus Banks, 1900 (Roewer 1910, 1923), transferred to Diguetinus by Goodnight& Goodnight (1942b), and assigned to Metopilio by Cokendolpher (1984b). These consecutive taxonomic changes occurred because M.spinulatus and D.raptator share a large body size and male genitalia pattern, nevertheless some features are different, such as the tuberculation, spination, and the lack of a prominent modification on leg I in M.spinulatus. Additionally, Cokendolpher etal. (2021) proposed that there are still other undescribed species of Diguetinus, and that D.raptator could be a species complex due to the variability of coloration and body proportions. These authors also made a redescription of D.raptator based on topotype specimens, illustrating the male genitalia for the first time and presented information about natural history. 733 The Mexican genus DiguetinusRoewer,1912 ZOOSYSTEMA • 2025 • 47 (30) The aim of the present study is to provide morphological and molecular phylogenetic evidence for the reincorporation of M.spinulatus into the genus Diguetinus. Additionally, a new diagnosis of the genera is proposed differentiating Diguetinus and Metopilio, and male dimorphism in D.raptator is presented with its taxonomic implications. MATERIAL AND METHODS SpecimenS and figureS The specimens examined are deposited in CNAN and CRLSMOp collections. A total of 27 individuals (10 ♀, six ♂ of D.raptator; 10 ♀, one ♂ of D.spinulatus stat. restit.) from their respective type localities, Guadalajara, Jalisco, Mexico (GDL-JAL) and Tepic, Nayarit, Mexico (TEP-NAY) respectively (Fig.1). We examined images of the female holotype of Leptobunus spinulatus (MCZ-IZ-14818) provided by the Museum of Comparative Zoology, Harvard University (MCZ). External morphological characteristics were observed under a LABOMED CZM6 stereo microscope. As a comparative reference, we took dorsal, ventral and lateral habitus photographs of the individuals of both D.raptator and D.spinulatus stat. restit. using a Sony Alpha 6400 digital camera, with a Laowa 100mm f/2.8 2:1 f/2.8 lens, and external illumination. Subsequently, the images were postprocessed in Photoshop© version 22.0.0, and in some cases, photographs were stacked in Helicon Focus version 8.1.0 to obtain images with greater depth of field. For the extraction of genitalia, dissections and preparations were performed following Acosta etal. (2007), photographs of the genitalia were obtained using the same configuration employed for the habitus (+ Raynox DCR-250 lens). Terminology, meaSuremenTS and abbreviaTionS For the description of the morphological structures, the terminology described in Cokendolpher etal. (2021), Kury& Cokendolpher (2020), Rodriguez etal. (2014) and Wijnhoven (2013) were used. To obtain measurements of D.spinulatus stat. restit., one male and ten females were measured, following suggestions of Acosta etal. (2007) and Cokendolpher etal. (2021). All measurements were taken in millimetres, through a HY-900B digital camera adapted to the stereoscope, using Hayear Version ×64 software. Morphology measurements BH body height, taken in lateral view; CL cephalothorax width; DL dorsum length, taken in lateral view; DW dorsum width, taken at the widest portion; GoL genital operculum length; GoWb width of genital operculum at base; GoWn width of genital operculum at neck; OcH ocularium height; OcL ocularium length; OcW ocularium width. Repository CNAN Colección Nacional de Arácnidos, UNAM, Mexico. CRLSM-Op Colección de Referencia del Laboratorio de Sistemática Molecular, UACB, UAZ, Mexico. (Op: Opiliones section); MCZ Museum of Comparative Zoology, Harvard University, Cambridge. molecular meThodS Genomic DNA was extracted from locomotor leg tissues of five and four individuals from TEP-NAY and GDL-JAL respectively, as described in Cruz-López etal. (2016) and Derkarabetian etal. (2019), using a column extraction kit (DNeasy® Blood& Tissue Kit, Qiagen Inc., Valencia, California) according to the manufacturer’s instructions. The mitochondrial proteinencoding cytochrome c oxidase subunit I (COI) was amplified using the primers LCO1490 (5’-GGTCAACAAATCATAAAGATATTG-3’) (Folmer etal. 1994) and HCOoutout (5’-GTAAATATATGRTGDGCTC-3’) (Prendini etal. 2005), following laboratory protocols outlined by Boyer etal. (2005) and Sharma& Giribet (2009). The following amplification conditions were used: an initial denaturation step at 95°C for 5minutes, followed by 40 cycles of 95°C for 30 seconds, annealing at 46°C for 40 seconds, and extension at 72°C for 45 seconds; followed by a final extension at 72°C for 7minutes. The Taq DNA Polymerase Amplification Kit (Vivantis Technologies PL1202) was used, following the manufacturer’s recommended volumes and concentrations of reagents. The final volume of each reaction was 25 μL, consisting of 1μL DNA (30ng/μL), 0.2μL Taq DNA Polymerase (5u/μL), 0.5dNTPs (10mM), 1 μL of each oligonucleotide (10 μM), 2.5 μL 10× buffer, 1 μL MgCl2 (25mM), and 17.9μL H2O. The amplified products were sent to Macrogen Inc. (South Korea) for Sanger sequencing. phylogeneTic reconSTrucTion and geneTic diSTanceS Forward and reverse chromatograms were automatically ensembled with CodonCode Aligner version 7.1.2 and AliView programs. Sequence identity was confirmed by searches in BLAST (Basic Local Alignment Search Tool) from NCBI (National Center for Biotechnology Information). The final COI alignment included four D.raptator sequences and five D.spinulatus stat. restit. sequences. Additionally, we incorporated a segment of a sequence of Metopiliosp. from Oaxaca, Mexico (SAMN35540793, MCZ-IZ-95161), based on sequence capture of ultraconserved elements (UCEs) from Derkarabetian etal. (2023). We included a sequence of Dalquestia formosa (Banks, 1910) generated by Giribet etal. (2001). As proposed by Hedin etal. (2012) and Derkarabetian etal. (2023), the most closely related family to Globipedidae is Phalangiidae. Therefore, available sequences of Phalangiidae were used as outgroups in the phylogenetic analysis, including Phalangium opilio Linnaeus, 1758 (Masta& Boore 2008), Odiellus lendii (Sørensen, 1894), Opilio canestrinii (Thorell, 1876), Mitopus morio (Fabricius, 1779) (Astrin etal. 2016) and Lophopilio palpinalis (Herbst, 1799). Additionally, we include sequences 734 ZOOSYSTEMA • 2025 • 47 (30) Ochoa-Vázquez D. et al. of Sclerosomatidae from Dewaard etal. (2019) as outgroups (GenBank accession numbers in Fig.8). The COI matrix was subjected to a maximum likelihood (ML) phylogenetic inference analysis. The most appropriate evolutionary model was estimated from the jModelTest software version 2.1.10 (Darriba etal. 2012) based on the Akaike information criteria (AICc) and Bayesian approaches (BIC) according to Posada& Buckley (2004). The General Time Reversible (GTR) model with Gamma variation rate and parameters for invariant sites (GTR+G+I) was identified as the most appropriate evolutionary model. ML analyses were conducted using the MEGA X program (Kumar etal. 2018). To evaluate the reliability of the branches, a bootstrap analysis was conducted with 10 000 replicates. The resulting topologies were exported from the Dendroscope version 3 software (Huson etal. 2007), and figures were subsequently processed in the Illustrator© version 22.0.0 program. Average genetic distances were calculated between all nucleotide sequences obtained, as well as between the clades identified in the phylogenetic analysis. These values were estimated using the MEGA X program (Kumar etal. 2018). RESULTS Order OPILIONES Sundevall, 1833 Suborder EUPNOI Hansen& Sørensen, 1904 Superfamily phalangioidea Latreille, 1802 Family globipedidae Kury& Cokendolpher, 2020 Genus Diguetinus Roewer, 1912 Diguetinus Roewer, 1912: 271, pl.1 fig.25; 1923: 863, fig.1030; 1956: 252.— Bronn 1932: 7, fig.9b.— Di Caporiacco 1938: 280.— Goodnight& Goodnight 1942a: 15.— Weidner 1959: 121.— Gruber 1969: 273.— Cokendolpher 1984a: 27-28; 1984b: 377.— Cokendolpher& Cokendolpher 1984: 168.— Cokendolpher& Lee 1993: 16.— Crawford 1992: 17.— Kury& Cokendolpher 2000: 150; 2020: 52.— Cokendolpher etal. 2021: 119, fig.1-9. Type SpecieS.— Diguetinus raptator Roewer, 1912. i ncluded SpecieS .— Diguetinus raptator Roewer, 1912 and Diguetinus spinulatus (Banks, 1898) stat. restit. emended diagnoSiS.— Diguetinus can be recognized from others members of Globipedidae by the following combination of characters: a) large bodied (adult individuals dorsum length from 10 to 12.5mm); b) dorsum with transverse rows of spines running across each opisthosomal tergite, from a single row configuration to a randomly ordered arrangement of at least two rows of spines; c) tergite spines yellowish-white and black tipped, usually with at least one chaetal sensilla present near the base of each spine, none spine extended to a height greater than the ocularium; d) stylus of the penis coiled in a complete circle. Diguetinus raptator Roewer, 1912 (Figs1A, B, E; 2A, B; 3A, B; 6A-F; 7A-C; 8-9) Diguetinus raptator Roewer, 1912: 271, pl.1 fig.25; 1923: 863, fig.1030; 1956: 252.— Bronn 1932: 7, fig.9b.— Di Caporiacco 1938: 280.— Goodnight& Goodnight 1942a: 15.— Weidner 1959: 121.— Cokendolpher& Lee 1993: 16.— Crawford 1992: 17.— Kury& Cokendolpher 2000: 150; 2020: 52.— Cokendolpher etal. 2021: 119, fig.1-9. Type daTa.— See Cokendolpher etal. (2021). maTerial examined.— Mexico • 6 ♂, 10 ♀; Jalisco: Guadalajara: Bosque El Centinela; 20°45’44.4”N, 103°22’51.1”W; 1584m a.s.l.; 26.X.2023; D. Ochoa-Vázquez, O. I. Ochoa-Vázquez leg.; GenBank accession numbers: PV430733-36; CRLSM-OP_01; CNAN-OP2021. emended diagnoSiS.— Diguetinus raptator can be distinguished from Diguetinus spinulatus stat. restit. by the combination of the following characters: a) dorsum with transverse rows of spines running across each tergite of the opisthosoma, not in a single row (as in Diguetinus spinulatus stat. restit.) but rather with random to at least two rows of spines per tergite; b) leg femur predominantly dark brown; c) glans length less than length of alae section, about 1/2 the length of alae; d) alae section proportion approximately 1/6 of the total length of the penis; e) alae section internal texture with a reticulate pattern, dark in colour, present over the entire alae surface. Diguetinus spinulatus (Banks, 1898) stat. restit. (Figs1C-E; 2C-D; 3C-D; 4-5; 6G-L; 7D-F) Leptobunus spinulatus Banks, 1898: 182. Hadrobunus spinulatus – Roewer 1910: 254, 257; 1923: 920. Diguetinus spinulatus – Goodnight& Goodnight 1942b: 11. Metopilio spinulatus – Cokendolpher 1984b: 376. Type daTa.— Holotype (examined by photographs). Mexico • ♀; Nayarit: Tepic; 21°30’34.2”N, 104°53’44.5”W (error radius: 8832m); Banks leg.; MCZ:IZ:14818. maTerial examined.— Mexico • 1 ♂, 10 ♀ ; Nayarit: Tepic: Cerro de la Cruz; 21°32’4.5“N, 104°53’5.3”W; 1091m a.s.l.; 28.X.2023; D. Ochoa-Vázquez, O. I. Ochoa-Vázquez leg.; GenBank accession numbers: PV430728-32; CRLSM-OP_02; CNAN-OP2022. diagnoSiS.— Diguetinus spinulatus stat. restit. can be distinguished from D.raptator by the combination of the following characters: a) leg femur bicolored, first 1/4 of all femurs light yellow-brown, more evident in femur II; b) transverse rows of spines across each tergite of the opisthosoma, with one linear row configuration per tergite; c) alae and glans penis sections similar in length (c.1mm), each proportion 1/8 of the total length of the penis; d) alae section internal texture with a reticulate pattern, dark in colour, present only on apical margin. redeScripTion Body leathery in texture, with a tuberculate-microgranulate cuticle morphology (with rounded or pointed protuberances surrounded by small obtuse to acute granules). Dorsum length 10.33mm (male) and an average length of 11.59mm ±0.61 (females: ranging from 10.68 to 12.3mm). Male dorsum light yellow-brown tone (alcohol preserved), dorsally with black patches on the cephalothorax across each opisthosomal tergite (Fig.2C), female dorsum coloration dark-brown tone (Fig.1C; 3C). Leg coloration with a gradual transition from light yellow-brown at the base of femurs to brown (male) and dark-brown (female) at patella and tibia, with tarsi and meta- 735 The Mexican genus DiguetinusRoewer,1912 ZOOSYSTEMA • 2025 • 47 (30) A CD E 105° O 105° O 21° N 21° N 104° O 104° O SIMBOLOGY Type localities Political state division Pacific coast Volcanic Belt Western Sierra Madre 40 km B Fig. 1. — Living females and type locality habitats of Diguetinus raptator Roewer, 1912 (A, B) and Diguetinus spinulatus (Banks, 1898) stat. restit. (C, D); E, geographical location of the type localities of D. spinulatus stat. restit. (Tepic, Nayarit: TEP-NAY; green hexagon) and D. raptator (Guadalajara, Jalisco: GDL-JAL; purple hexagon). Biogeographic regionalization following Arriaga et al. (1997). Photos: A-D, DOV. 736 ZOOSYSTEMA • 2025 • 47 (30) Ochoa-Vázquez D. et al. AB CD Fig. 2 . — Comparison of males of Diguetinus raptator Roewer, 1912 and Diguetinus spinulatus (Banks, 1898) stat. restit. from Guadalajara, Jalisco, Mexico and Tepic, Nayarit Mexico, respectively: A, B, D. raptator dorsal (A) and ventral (B) habitus; C, D, D. spinulatus stat. restit. dorsal (C) and ventral (D) habitus. Scale bars: 1 mm. 737 The Mexican genus DiguetinusRoewer,1912 ZOOSYSTEMA • 2025 • 47 (30) AB CD Fig. 3 . — Comparison of females of Diguetinus raptator Roewer, 1912 and Diguetinus spinulatus (Banks, 1898) stat. restit. from Guadalajara, Jalisco, Mexico and Tepic, Nayarit Mexico, respectively: A, B, D. raptator dorsal (A) and ventral (B) habitus; C, D, D. spinulatus stat. restit. dorsal (C) and ventral (D) habitus. Scale bars: 1 mm. 744 ZOOSYSTEMA • 2025 • 47 (30) Ochoa-Vázquez D. et al. The sequence of Dalquestia formosa was recovered paraphyletic with respect to the Globipedidae (only Diguetinus + Metopilio) and Phalangiidae clades. A total of two main subclades were recovered within the Diguetinus s.l. clade, both exhibited a bootstrap value of 100%. The purple subclade grouped the sequences from GDL-JAL (the type locality of D.raptator), while the green clade includes sequences from TEP-NAY (the type locality of D.spinulatus) (Fig.8). The genetic differences expressed as pairwise distance between the clades identified in the phylogenetic hypothesis described above (Table2) showed that the highest divergence values were observed between the outgroup sequences (Sclerosomatidae and Phalangiidae) and the remaining groups (ranging from mean 31.9 to 34.6% and 21.6 to 26.8%, respectively), as well as Dalquestia formosa sequence and the remaining groups (from 26.8 to 32.7), then the Metopiliosp. lineage and the remaining groups (from 20.2 to 34.6% of mean distance). The mean divergence between subclades within the Diguetinus s.l. clade was 6.3% (between the D.spinulatus and D.raptator clades, with percentages ranging from 6.1 to 6.9%). Intraspecific divergence observed in D.raptator were from identical sequences to 0.2%, and within D.spinulatus from identical sequences to 0.04%. DISCUSSION Diguetinus vS Metopilio Some authors have proposed that the genus Diguetinus is similar to Metopilio, implying that they may eventually be considered synonyms, the main distinction being in the number of tubercles found in the transverse rows of the dorsum tergites and in the curved morphology of the stylus (Gruber 1969; Cokendolpher 1984a; Cokendolpher& Cokendolpher 1984; Cokendolpher& Stockwell 1986; Cokendolpher& Sissom 2000; Cokendolpher etal. 2021). However, in the taxonomic work by Roewer (1911, 1912, 1923), Metopilio is distinguished from Diguetinus by the presence of two median spines on each tergite of the dorsum, described as follows: “the last cephalothorax and abdominal segments with more or less clear transverse rows of pointed tubercles, the two medians of which are reinforced like thorns, so that two parallel longitudinal rows of such arise” (translated from the German from Roewer 1912: 262-263). These original definitions do not establish the number of tubercle rows per tergite as a diagnostic character, as subsequently suggested by several authors (Cokendolpher 1984a; Cokendolpher& Cokendolpher 1984; Cokendolpher& Stockwell 1986; Cokendolpher& Sissom 2000; Kury& Cokendolpher 2020; Cokendolpher etal. 2021). Nevertheless, Roewer (1911, 1912, 1923) stated the features that define Diguetinus, i.e., rough tuberculated dorsal segments, but without large spines, coarsely granulated coxae; and sexual dimorphism, in males, legs I and III strongly clubbed, tibia I ventrally with a strong tuberculate tooth and articulated against the S-shape metatarsus I, ventrally toothed, thus forming a grasping hook. Furthermore, Cokendolpher etal. (2021) pointed out that, in Diguetinus, the opisthosomal spines are not arranged in a single transverse row per tergite, but rather randomly with at least two rows of tubercles per tergite. The specimens from D.raptator and D.spinulatus stat. restit analysed in the present study (Figs1-4; 6-7) exhibited the diagnostic characters described by Cokendolpher etal. (2021) (in part) and Roewer (1912), with a notableabsence of a pair of long spines on each dorsal tergite, with only one row of spines per tergite (D.spinulatus stat. restit.) and notably more than two rows of spines per tergite (D.raptator).We suggest that the dichotomy for the separation of these genera by one row of spines or irregular rows of spines on each tergite, is a misinterpretation of the original descriptions. Otherwise, the situation in Metopilio, the most specious genus in the family with 18 described species, is a result of the absence of data, because the penial morphology has not been fully documented, and the genus in general requires taxonomic revision to clarify the definitions of the species (Cokendolpher& Cokendolpher 1984). Currently, only the penis morphologies of Metopilio australis (Banks, 1909) (Gruber 1969), Metopilio ornatipes (Banks, 1909), and Metopilio niger Goodnight& Goodnight, 1942 (Rodriguez etal. 2014) are known. Furthermore, an examination of various species within the genus revealed a diverse range of penis morphologies, including examples with a markedly pronounced alate region, as observed in M.australis (Gruber, 1969, figs1, 4, 5), and those lacking an alate region (undescribed Metopilio species from Table 2. — Pairwise genetic distance (expressed as percentage) between clades identified in the phylogenetic hypothesis (Fig. 8). Mean values in bold, followed by minimum-maximum ranges, OG, outgroup (see Fig. 8). D. raptator D. spinulatus Metopilio sp. Dalquestia formosa OG: Phalangiidae D. raptator ––––– D. spinulatus 6.3 6.1-6.9 – – – – Metopilio sp. 20.2 20.1-20.3 21.3 21.3-21.5 – – – Dalquestia formosa 30.1 27.1 26.927.4 28.1 – – OG: Phalangiidae 23.7 22.4-27 21.6 20.3-24.7 24.4 24.2-25.6 26.8 26.4-29.2 – OG: Sclerosomatidae 34.8 34.4-35 31.9 31.5-32.3 34.6 34.4-34.8 32.7 32.6-32.7 32.4 31.2-36 745 The Mexican genus DiguetinusRoewer,1912 ZOOSYSTEMA • 2025 • 47 (30) Mexico; Cruz-López, personal observations). Moreover, it is notablethat none of the above-mentioned specimens exhibit the distinctive stylus morphology described in Diguetinus (Kury& Cokendolpher 2020; Cokendolpher etal. 2021; this study), which is characterized by a coiled structure that forms a complete ring (Fig.6). This observation indicates that the stylus morphology is an important taxonomic character for delineating the boundaries of Diguetinus and Metopilio. Diguetinus SySTemaTicS and geographical diSTribuTion Our phylogenetic results, morphological description, and geographic distribution provide evidence supporting the separation of D.spinulatus stat. restit. and D.raptator as distinct species. The average genetic distance between the two clades was 6.3%, with percentages ranging from 6.1 to 6.9%. To date, no information on genetic variation within the family Globipedidae has been reported. Hedin& Thomas (2010) reported COI genetic distances between species within the family Phalangodidae to range from 12 to 30%. As an example, within Opiliones species, intraspecific variation within Fumontana deprehendor Shear, 1977 (Laniatores, Triaenonychoidea, Buemarinoidae) was observed to range from identical sequences to 6%. In a study conducted as part of the Barcode of Life project in Germany, Astrin etal. (2016) analysed the sequences of approximately 200Opiliones species from the region, showing that the range of percent divergence between closely related species is 7 to 21.8%. Considering all evidence A B C D Leg III Leg III Leg I Leg I Fig. 9. — Intrasexual dimorphism of Diguetinus raptator Roewer, 1912 from Guadalajara, Jalisco, Mexico: A, C, alpha male; B, D, beta male. Lateral view (A, B), prolateral view of legs I and III (C, D). Scale bars: 2 mm. 746 ZOOSYSTEMA • 2025 • 47 (30) Ochoa-Vázquez D. et al. presented here, the comparable genetic divergence among other harvestman and between D.spinulatus stat. restit. and D.raptator, the phylogenetic hypothesis, morphology of the stylus and other features, and the disjunct geographic distribution, we conclude that they represent distinct evolutionary lineages. Cokendolpher etal. (2021) presented a distribution map of the genus Diguetinus based on historical records in literature, original records, and records of citizen science observations (iNaturalist platform). They noted that the distribution of the “species” of the genus aligns with the boundaries of two biogeographic provinces defined by Arriaga etal. (1997): the southern Mexican Plateau and the western and central Transmexican Volcanic Belt. The type localities of D.spinulatus stat. restit. and D.raptator are found in the Pacific Coast and Volcanic Belt provinces, respectively (Fig.1E). This information demonstrates that the genus Diguetinus is distributed along the Mexican Pacific coast and, probably, even more widespread than previously documented. Consequently, further research under an integrative taxonomic approach will facilitate a comprehensive understanding of the phylogenetic relationships, evolutionary history, and biogeography of the potential species present in the central Mexican territory (Cokendolpher etal. 2021). Finally, our phylogenetic results included a sequence of Dalquestia formosa (AF370833) (Giribet etal. 2001), which resulted in a paraphyletic relationship with respect to the Phalangiidae and Globipedidae clades. This could be due to the lack of taxa terminals sequenced in Globipedidae. It is crucial that future efforts to resolve the phylogenetic relationships of the family take this into account. In addition, there are two globipedid COI sequences, Dalquestia grasshoffti (JQ437182) and Eurybunus brunneus (JQ437181), generated by Hedin etal. (2012). However, these were excluded in the present study since they correspond to a different COI fragment analysed here and are therefore not alignable. Sexual dimorphiSm in Diguetinus Roewer (1910, 1912, 1923) used the sexually dimorphic character of legs I and III to recognize Diguetinus among relatives. However, in the analysed material of D.raptator from GDL-JAL, three adult males were observed with the described morphology and three without this morphology (legs I and III not modified) (Fig.9). In Opiliones, several authors have employed the terms “alpha/beta” or “major/minor” to describe the dimorphic condition observed in males. This condition presents larger and more strongly armed males (classified as alpha or major) and smaller males similar to females with reduced or absent armor (beta or minor) (Kury 2008; Ferreira& Kury 2010; Zatz etal. 2011; Kury& Ferreira 2012; Buzatto& Machado 2014; Cruz-López etal. 2016; Townsend Jr.& Enzmann III 2018; Powell etal. 2020; Machado& Burns 2024). This morphological differentiation has been documented from males in the suborders Eupnoi and Laniatores, particularly in the families Neopilionidae Lawrence, 1931 (Taylor& Hunt 2009; Powell etal. 2020), Protolophidae Banks, 1893 (Tsurusaki& Cokendolpher 1990), Cosmetidae Koch, 1839 (Pérez-González& Vasconcelos 2003; Kury& Ferreira 2012), Triaenonychidae Sørensen, 1886 (Forster 1954), and Gonyleptidae Sundevall, 1833 (Pinto-Da-Rocha& Bragagnolo 2010), among others [see Buzatto& Machado (2014) table1]. Buzatto& Machado (2014) described the Alternative Reproductive Tactics (ARTs) in Opiliones as a strategy to avoid complete exclusion from the mating pool. These tactics manifest as morphological dimorphisms in males, which promote sexual interactions such as competition and monopolization of territories, sperm competition, exclusive access to gametes, access to non-fertilized eggs, and competition for unfertilized eggs (see Buzatto& Machado (2014)). The observations made in this study represent the first documentation of intrasexual dimorphism within the family Globipedidae. We show that males of D.raptator exhibit dimorphism manifested in the presence or absence of armature and swelling of the legs I and III (Fig.9) and may indicate the potential existence of ARTs in Diguetinus. However, currently, there is a lack of comprehensive data on sexual behavior within the genus. Field observations of Diguetinussp. revealed a scarcity of males, with only one male observed among at least 50females, or in extreme cases, the absence of males for more than 70females (Ochoa-Vázquez, obs. pers.). In our sampled locality of D.raptator, more than 20female individuals were observed, along with three alpha males and three beta males. In contrast, the male specimen of D.spinulatus stat. restit. exhibits a beta male morphology. Finding a single male individual provides the possibility that there are alpha forms in nature, as well as the close phylogenetic relationship with D.raptator and the observation by citizen scientists in the type locality, wich possibly represents the alpha form of the species, as evidenced by the armor of legs I and III (Fig.5C). In light of these observations, future studies aimed at elucidating the role of alpha/beta males in populations and the overall reproductive behaviour of these species are needed, focusing on behaviour in general and particularly sexual behaviour within the genus Diguetinus. Finally, considering the intrasexual dimorphism reported here, we suggest that sexual dimorphism as diagnostic taxonomic character is insufficient to define the genus Diguetinus, so its use should be taken with caution and not lead to misidentifications. Acknowledgements DOV thanks the Secretaría de Ciencia, Humanidades, Tecnología e Innovación (SCIHTI), for the financial support in form of scholarship during his graduate studies. We thank Marshal Hedin, anonymous referee and editors for their useful and invaluable comments and suggestions that significantly improved the manuscript. Thanks to Edmundo GonzálezSantillán (IB-UNAM) for providing the CNAN catalogue numbers and to Ligia Benavides (Curatorial Assistant of Arachnids and Myriapods, MCZ, Harvard University) for providing images of the holotype of Leptobunus spinulatus. Special thanks to Shahan Derkarabetian (Curator of Entomology, San Diego Natural History Museum) for kindly providing the UCE sequences of Metopiliosp. Thanks to Jesus Guzmán-Moreno for his assistance in the management 747 The Mexican genus DiguetinusRoewer,1912 ZOOSYSTEMA • 2025 • 47 (30) of the UCE sequences. Special thanks to the Laboratorio de Genómica Evolutiva (UACB-UAZ) (Melina del Real Monroy, Lenin Sánchez Calderón and Luciano A. Juárez Díaz) for allowing us to access the Nanodrop. We would like to thank the following people for their help in obtaining the material analyzed in this paper: Emmeth Rodríguez, Carlos Vázquez and especially Gillermo Alatorre (D.raptator locality) and the invaluable technical help of Octavio I. Ochoa Vázquez. Thanks to the Laboratorio de Entomología Agrícola team at INIFAP-OAX for their hospitality and support during DOV short research stay in August 2024. These results were part the project UAZ-2023-38899 (E.A.M.S is the principal investigator). REFERENCES acoSTa l. e., gonzález a. p. & lúciaTourinho a. 2007.— Methods for taxonomic study, in pinTo-da-rocha r., machado g. & giribeT g. (eds), Harvestmen: The Biology of Opiliones. 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