Research Article
Print
Research Article
Taxonomic notes on Barinas: a new generic synonym, a new cave-dwelling species, and new records from Colombia (Arachnida, Opiliones, Agoristenidae)
expand article infoAndrés F. García§, Daniela Ahumada-C.§
‡ Universidade Federal Do Rio de Janeiro, Rio de Janeiro, Brazil
§ Universidad de Cartagena, Cartagena de Indias, Colombia
Open Access

Abstract

Barinas guanenta sp. nov. is described from a cave in Santander, central Andes of Colombia. The new species is recognized by the pedipalps and chelicerae entirely pale yellow and without variegated pattern, the areas I-IV with variegated coloration pattern and the straight stylus of the penis. The monotypic genus Vimina González-Sponga, 1987 is synonymized with Barinas, resulting in Barinas virginis (González-Sponga, 1987) comb. nov. New records of Barinas piragua Ahumada-C. & García, 2020 in La Guajira, Northern Colombia, are given. A key for the species of Barinas is given together with an updated distribution map.

Key Words

biodiversity, harvestmen, Leiosteninae, taxonomy, Vimina

Introduction

The taxonomy of the Neotropical subfamily Leiosteninae Šilhavý, 1973 (12 gen., 59 spp.), widespread in northern South America, has recently aroused the interest of some researchers, resulting in several papers focused on nomenclatural acts (Ahumada-C. et al. 2020; García and Kury 2020; García and Villarreal 2020; García and Pastrana-M. 2021; Villarreal et al. 2021), and/or approaches to the systematics of the group (Villarreal and García 2021).

To continue on this path, the present work brings taxonomical novelties of the genus Barinas González-Sponga, 1987, that currently groups two species: the type species, Barinas flava González-Sponga, 1987 (from the Andean slopes of Venezuela), and Barinas piragua Ahumada-C. & García, 2020 (from the Caribbean region of Colombia), both characterized by the paired armature in the scutal areas I-IV and the stylus of penis dorsally curved (Ahumada-C. et al. 2020).

After the revision of harvestmen material from Colombia and Venezuela, we noted that the monotypic genus Vimina González-Sponga, 1987, whose type species was described from a locality near to that of B. flava, should be synonymized with Barinas. Besides that, a new cave-dwelling species from the Andean region of Colombia was found. So, a new diagnosis for Barinas, a generic synonym with its subsequent new combination, the description of the new Colombian species, and new records for B. piragua are proposed.

Materials and methods

Individuals were photographed using a Leica M205C stereoscope attached to a Leica DFC450 digital camera. The resultant images were posteriorly edited in Photoshop CC 2014 software. Drawings were made using the software Inkscape 0.91. Descriptions of colors use the standard names of the 267 Color Centroids of the NBS/IBCC Color System as named in Centore (2016). The distribution map was made with Quantum GIS 3.12.3 Bucarest software (QGIS Development Team 2020). Colored shapes refer to WWF Terrestrial Eco-regions of the World (Olson et al. 2001).

Patterns of description and AHF (armature height formula, height differences between the armature on the mesotergal areas, where roman numerals represent the corresponding mesotergal area) follow Ahumada-C. et al. (2020). The terminology for dorsal scutum outline types follows Kury and Medrano (2016), with the modifications explained in Villarreal and García (2021), and for chaetotaxy of penis lamina parva and truncus follows Kury and Villarreal (2015).

Morphology abbreviations are AL―maximum abdominal scutum length, AW―maximum dorsal scutum width, ChL―chelicera length, CL―carapace length, CW―maximum carapace width, DH―dorsal hump, DP―dorsal process, DS―dorsal scutum, DSL―dorsal scutum length, Fe―femur, IOD―interocular distance, LP―lamina parva, MS―macrosetae of the penis, Mt―metatarsus, Pa―patella, Ta―tarsus, Ti―tibia, TL―total length, Tr―trochanter. All measurements are in mm unless otherwise noted.

The specimens are deposited in the following institutions: CBUDC―Colección de Ejemplares Biológicos de la Universidad de Cartagena (Cartagena de Indias, Colombia); ICNInstituto de Ciencias Naturales de la Universidad Nacional de Colombia (Bogotá, Colombia); MAGS―Manuel Angel González Sponga collection (donated to MIZA collection); MCNCFundación Museo de Ciencias Naturales de Caracas (Caracas, Venezuela); MIZAMuseo del Instituto de Zoología Agrícola (Maracay, Venezuela); MNRJMuseu Nacional da Universidade Federal do Rio de Janeiro (Rio de Janeiro, Brazil). The material destroyed by the fire in September 2018 is marked with an asterisk (*).

Results

Taxonomy

Order Opiliones Sundevall, 1833

Family Agoristenidae Šilhavý, 1973

Subfamily Leiosteninae Šilhavý, 1973

Barinas González-Sponga, 1987

Barinas González-Sponga 1987: 453; Kury 1997: 344; 2003: 30; Ahumada-C. et al. 2020: 633.

Vimina González-Sponga 1987: 547; Kury 1997: 344; Kury 2003: 34. New synonym

Type species

Barinas flava González-Sponga, 1987, by original designation.

Emended diagnosis

DS Epsilon type 1. Ocularium low, smooth, or with paired high spines (Barinas piragua). Mesotergal areas I–IV with paired paramedian spines (AHF: I < II = III = IV in B. flava, I = II < IV < III in B. piragua, II = I<III = IV in B. virginis comb. nov., and I = IV < II = III in B. guanenta sp. nov.). Posterior margin, free tergites, and anal operculum armed (with tubercles in B. flava, B. virginis comb. nov. and B. guanenta sp. nov.; with spines in B. piragua). Pedipalps concolor with the body (B. flava and B. virginis comb. nov.; lighter than the body in B. piragua and B. guanenta sp. nov.). Coxae I–IV with one anterolateral tubercle dorsally projected. Fe IV length/DSL ratio: less than three in B. virginis comb. nov (1.93) and B. flava (2.58); more than three in B. piragua (3.4) and B. guanenta sp. nov. (3.38). Lamina parva crescent-shaped (in B. flava seems heart-shaped) and with very short corners that do not exceed the insertion point of the MS-A2. Malleus with two pairs of trifid MS-A, one pair of trifid MS-B, two pairs of MS-D, one pair of large and trifid MS-E2, and one pair of MS-E1 (in B. piragua short an conical, in B. guanenta medium-sized and elongated; unknown in B. flava and B. virginis comb. nov.). Stylus elongated, curved dorsally at distal portion (straight in B. guanenta sp. nov.), smooth (in B. flava and B. virginis comb. nov.; with a dorsal hump close to the union of the DP with the stylus in B. piragua and B. guanenta sp. nov.), and with a sharp proximal DP (blunt in B. virginis comb. nov.).

Included species

Barinas flava González-Sponga, 1987, Barinas guanenta sp. nov., Barinas piragua Ahumada-C. & García, 2020, Barinas virginis (González-Sponga, 1987) comb. nov.

Remarks

An emended diagnosis comparing Barinas with the remaining genera of Leiosteninae was offered by Ahumada-C. et al. (2020) and we decided not to repeat such information here. However, we want to emphasize that the presence of paired armature on mesotergal areas plus the basal DP of the stylus and the LP with very short corners not exceeding the insertion point of the MS-A2 are characters exclusive of Barinas.

Key to the species of Barinas (males)

1 Ocularium with paired spines; tarsus and claw of pedipalp of different coloration than the other pedipalpal segments B. piragua
Ocularium smooth; all segments of pedipalp of the same coloration 2
2 Paramedian armature of mesotergal area IV of the same size as that of area III; stylus curved distally 3
Paramedian armature of mesotergal area IV smaller than that of area III; stylus straight B. guanenta sp. nov.
3 Paramedian armature of mesotergal area II smaller than that of area III; dorsal connective tissue between the stylus and the dorsal process present (Fig. 3C, D, F) B. virginis comb. nov.
Paramedian armature of mesotergal area II same size as that of area III; dorsal connective tissue between the stylus and the dorsal process absent (figs 1D-F in Ahumada-C. et al. 2020) B. flava

Barinas guanenta sp. nov.

Figs 1, 2, 4

Type material

Colombia• ♂ holotype: Santander, Cabrera, Cueva del Indio de Doña Joaquina; [6.562261; – 73.237897]; [1100 m]; 14 July 2007; Ca-vita grupo de bioespeleologia leg.; ICN-Ao-742.1 • 1 ♂ 1 ♀ paratypes; same data as the holotype; ICN-Ao-742.

Etymology

Guanentá was the leader of the Guanes, an indigenous group that fought bravely against the Spanish conquerors in the region where the species was collected. Noun in apposition.

Diagnosis

Can be distinguished by having pedipalps entirely pale yellow without variegated pattern (in B. piragua distal portion of Tr and Fe black, and tarsi entirely black; in B. virginis comb. nov. with a variegated pattern on Fe-Ti); variegated coloration on areas I-IV (B. flava and B. piragua without variegated coloration); stylus of penis straight (in all other Barinas species the stylus is curved at apical portion) (Figs 1, 2).

Figure 1.

Photographs of Barinas guanenta sp. nov. (ICN-Ao-742.1), male holotype. Habitus in panoramic (A), dorsal (B), ventral (C), lateral (D), and frontal (E) views. Scale bars: 3 mm (A); 1 mm (C, D); 0.5 mm (B, E).

Figure 2.

Drawings of Barinas guanenta sp. nov. Male holotype (ICN-Ao-742.1) habitus in dorsal (A) and lateral (B) views; coxa I, ventral view (C); chelicera, frontal view (D); right pedipalp, mesal view (E). Male paratype (ICN-Ao-742): genitalia in lateral and ventral views (F); detail of stylus (G). The colored letters A-D refers to the chaetotaxy system of Kury and Villarreal (2015). Abbreviations: DH―dorsal hump, DP―dorsal process, LP―lamina parva, Ma―malleus, St―stylus, T―truncus. Scale bars: 0.5 mm (A-E); 0.05 mm (F,G).

Description

(Figs 1, 2). Male holotype (ICN-Ao-742.1) measurements (mm): CL (0.57), AL (1.22), CW (1.20), AW (1.44), DSL (1.79), IOD (0.27), ChL (0.86). Pedipalp: Tr (0.35), Fe (0.48), Pa (0.34), Ti (0.57), Ta (0.44), Claw (0.42). Legs I-IV (Tr/Fe/Pa/Ti/Mt/Ta/TL): I (0.21/2.45/0.43/1.93/3.69/1.24/9.95), II (0.27/6.19/0.65/5.38/7.30/3.57/23.36), III (0.34/4.02/0.75/2.47/4.85/1.29/13.72), IV (0.37/6.11/0.71/3.57/6.80/1.79/19.35). Dorsum: DS outline Epsilon type 2 (Figs 1A, B, 2A). Cheliceral sockets narrow, separated by medial projection (Fig. 2A). Anterior margin of DS smooth. Ocularium high (Figs 1D, E, 2B). Carapace smooth. Mesotergum delimited, divided into four areas (area I divided into two halves), with a transverse row of granules on each (Figs 1B, D, 2A, B). Areas I–IV with two paramedian spines, AHF: I = IV < II = III (Figs 1D, E, 2B); paramedian spines of areas III and IV projected backward (Figs 1D, E, 2B). Posterior margin of DS substraight, with a row of tubercles. Free tergites I–III with a row of tubercles and anal operculum with some tubercles (Figs 1B, 2A, B). Venter: Tegument granular. Coxa I with one trifid tubercle on the anterior margin and a transverse row of tubercles on the medial region (Fig. 2C). Coxa III-IV with some tubercles. Sternites with rows of spaced granules (Fig. 1C, D). Stigmatic area slightly granular. Stigmata large and oval (Fig. 1C). Genital operculum slightly granular. Chelicerae: Chelicera swollen. Segment I rectangular in dorsal view, with three-four tubercles on the proximal region and one-two tubercles on the ectal face (Figs 1B, 2A). Basal region of the movable finger with abundant setiferous tubercles of different sizes (some reaching the medial region of hand) (Fig. 2D). Fixed finger with smooth inner surface (Fig. 2D). Movable finger with one trapezoid sub-basal tooth and distal inner surface dentate (Fig. 2D). Pedipalps: Trochanter with five dorso-basal tubercles and two ventroectal tubercles. Fe with a ventroectal row of four setiferous tubercles (the two basalmost largest and the two distalmost small-sized), and one large ventromesal setiferous tubercle in the distal portion. Pa armed with one large ventromesal setiferous tubercle (Fig. 2E). Tibial setation: ectal iiIi, mesal IIi. Ta setation: ectal I?i, mesal IIi. Legs: Leg I–IV granular. Leg I filiform, the rest getting steadily thicker from leg II-IV (Fig. 1A). Tr I-III slightly granular, Tr IV with two dorsodistal tubercles (Figs 1B, 2A). Tarsal counts: 7(3)-7(3)/17(3)-16(3)/7-7/8-7. Penis: LP small and depressed, crescent-shaped, with anterolateral short sharp corners, apically pointed (Fig. 2F). Malleus with two pairs of trifid MS-A, and one pair of trifid MS-B (Fig. 2F); MS-C absent. Two pairs of MS-D located in a keel between the ventral part of LP and the base of the stylus; MS-E2 large and trifid, MS-E1 medium-sized (half of the size of MS-E2) (Fig. 2F). Stylus straight and elongated, surpassing the LP, with a short DP at the proximal region, and a DH just after the union of the DP with the stylus (Fig. 2G, F). Color (in ethanol) (Fig. 1A–E): Carapace and mesotergum reticulated Dark Grayish Olive (111) on Light Grayish Olive (109). Pedipalps and chelicerae Yellowish White (92). Free tergites Dark Olive (108). Coxae I-IV Light Grayish Olive (109) with lateral margins Dark Olive (108), trochanters I-IV Light Grayish Olive (109). Fe, Pa, and Ti I Dark Olive (108). Femora and tibiae II-IV Light Grayish Olive (109), patellae II-IV Dark Olive (109).

Female paratype (ICN-Ao-742). Similar to male, except for the non-hipertelic chelicera and shorter Fe IV. Measurements (mm): CL (0.57), AL (0.93), CW (1.06), AW (1.24), DSL (1.50), IOD (0.23), ChL (0.61). Pedipalp: Tr (0.22), Fe (0.49), Pa (0.32), Ti (0.46), Ta (0.43), Claw (0.33). Legs (Tr/Fe/Pa/Ti/Mt/Ta/TL): I (0.26/2.00/0.42/1.52/2.96/1.05/8.21), II (0.34/4.87/0.50/4.54/6.08/3.69/20.02), III (0.31/3.70/0.64/2.21/4.02/1.16/12.04), IV (0.34/5.52/0.65/?/?/?/?). Tarsal counts: 7-7/17-?/7-?/?-?.

Distribution

Known only from the type locality, a cave in the Magdalena Valley montane forest ecoregion (NT0136) (Fig. 4).

Barinas virginis (González-Sponga, 1987), comb. nov.

Figs 3, 4

Vimina virginis González-Sponga 1987: 547, figs 714–719; Kury 2003: 34.

Type data

Venezuela• ♂ holotype: Trujillo, Trujillo, Cerro de La Virgen, environs of Trujillo; [9.349513, -70.457898]; 1100 m; A.R. Delgado de González, J.A. González Delgado y M.A. González-Sponga leg.; MCNC 981 • 1 ♀ paratype; same data as holotype; MCNC 982 • 1 ♂ paratype; same data as holotype; MAGS.

Other examined material

Venezuela• 3 ex.: Trujillo, Cerro La Virgen; 1100 m; 28 July 2006; O. Villarreal leg.; MNRJ 9324* • 1 ♂; Trujillo, Cerro de La Virgen; 1300 m; MAGS 852.

Complementary description

DS Epsilon type 2 (Fig. 3A). Anterior and lateral margins of DS smooth, posterior margin with some granules. Ocularium low, with median concavity, unarmed (Fig. 3B). Mesotergum delimited, divided into four areas, granulate; area I divided longitudinally, areas II–IV undivided; AHF: II = I<III = IV (Figs 3B–E). Posterior margin of DS substraight. Free tergites I–III with granules (Fig. 3E). Cheliceral hand swollen. Legs increasing in thickness from leg I to leg IV, unarmed, leg I filiform. Fe IV two times DS length (Fig. 3B). Penis with small LP (height less than width), apex with anterolateral sharp corners. Malleus with two pairs of branched MS-A; one pair of branched MS-B; MS-C absent; MS-D absent (see remarks); MS-E2 large and trifid, MS-E1 absent. Stylus elongated, curved at distal third, surpassing the lamina parva, with dorsal process and dorsal connective tissue (Fig. 3F).

Figure 3.

Barinas virginis (González-Sponga, 1987) comb. nov. (MAGS 852), male. Habitus in dorsal (A), panoramic (B), frontal (C), lateral (D), and posterior (E) views. Penis drawings (F) of holotype (MCNC 981) (González-Sponga, 1987: 550, figs 718–719), lateral and ventral views (modified to show the chaetotaxy system of Kury and Villarreal (2015)). Scale bars: 1 mm. Photos courtesy of Osvaldo Villarreal.

Figure 4.

Map showing the distribution of Barinas in Northern South America. Colored shapes refer to WWF Terrestrial Eco-regions of the World (Olson et al. 2001).

Remarks

It seems that MS-D and MS-E1 went unnoticed or ignored in some works of González-Sponga (1981, 1987, 1998). However, Barinas species has these MS, as explained by Ahumada-C. et al. (2020). Therefore, we suspect that they are present in B. virginis comb. nov. as well, but only a revision of the male genitalia could settle this issue.

Barinas piragua Ahumada-C. & García, 2020

Barinas piragua: Ahumada-C. et al. 2020: 635.

Previous records

Colombia: Bolívar and Magdalena departments.

New records

Colombia• 2 ♀: La Guajira, Hatonuevo, Cerro Bañaderos, 11°7’33.3"N, 72°47’06.9"W [11.125899, –72.785241]; 785 m; 19 May 2018; Miguel Gutiérrez leg.; CBUDC-ARA 346.

Discussion

The penis of the Barinas species shows remarkable characters, as the short size of the corners of the LP and the shape of the stylus. For the first case, the genus has a LP with very short corners that do not exceed the insertion point of the MS-A2 (just known in Paravima Caporiacco, 1951 as far as we know). On the other hand, the distal half of the stylus exhibits a 45-degree curvature at the dorsal region (except B. guanenta sp. nov) not present in the other Leiosteninae genera. Nevertheless, the presence of paired armature in all the mesotergal areas plus the basal DP on the stylus is a combination of characters exclusive of Barinas, so, we do no doubt about the inclusion of B. guanenta sp. nov. in it.

Additionally, we noted that B. flava lacks additional ornamentations above the insertion point of the DP (see Ahumada-C. et al. 2020, fig. 1F), while the other species of Barinas exhibits both a DH (see Ahumada-C. et al. 2020, fig. 4D for B. piragua and Fig. 2G for B. guanenta sp. nov.) or connective tissue in B. virginis comb. nov. (Fig. 3F). Nonetheless, it is difficult to know if the DH is the remnant of membranous tissue or corresponds to another structure. We think that SEM images of the genital of B. flava could help to better understand such an issue and that a phylogenetic analysis including all the species of Barinas and more representatives of Leiosteninae could help in the understanding of the internal relationships of the group.

Finally, we would like to reinforce the importance to do scientific expeditions in the cave systems in Colombia, mainly because those have perfect conditions of humidity and darkness for the harvestmen. Besides B. guanenta, there are three oficial records of the order Opiliones in caves: Andrescava sturmi Roewer, 1963 (Agoristenidae) in National Natural Park Cueva de los Gúacharos (Huila); Phareus antrophilus Villarreal & Rodríguez, 2006 (Stygnidae), in Gámbita (Santander); and Phalangodus briareos Villarreal & García, 2016 (Cranaidae), from Zapatoca (Santander) (Barriga et al. 2019). However, there are many undescribed species and genera, as evidenced by some recent works on the cave systems of Santander (Casallas-Pabón et al. 2013; Barriga et al. 2019).

Acknowledgments

Thanks to Eduardo Flórez for the loan of material from the ICN collection. To Adriano Kury (MNRJ), for all the facilities in the laboratory work and for maintaining the OmniPaper Project, a very helpful tool that provided access to harvestmen literature. To Miguel Gutiérrez for sharing with us the material of B. piragua from La Guajira collected by him. To Osvaldo Villarreal for providing the photographs of Vimina virginis from the MIZA collection used in Fig. 3. The photographs of Fig. 1 were taken with the stereomicroscope (CNPq Universal 14/2013) at the Invertebrate department (MNRJ). The suggestions of Miguel Medrano and one anonymous referee greatly improved the present work. This work was supported by a scholarship from the Coordination for the Improvement of Higher Education Personnel (CAPES) to AFG, and financed in part by the PROEX Program [CAPES-Finance Code 001, assistance number 0721/2018] to PPGZoo and by international mobility scholarship from Universidad de Cartagena (Resolution 01787 of 2020) to DAC.

References

  • Ahumada-CD, García AF, Navas-S GR (2020) The spiny agoristenid genus Barinas (Arachnida: Opiliones), with the description of a new species from the Colombian Caribbean. Arachnology 18(6): 632–641. https://doi.org/10.13156/arac.2020.18.6.632
  • Barriga JC, Martínez-Torres D, López-Orozco CM, Villarreal O, Murcia MA (2019) Capítulo 3. Artrópodos terrestres de las cuevas y cavernas de El Peñón (Andes), Santander, Colombia. In: Lasso CA, Barriga JC, Fernández-Auderset J (Eds) Volumen VII. Biodiversidad subterránea y epigea de los sistemas cársticos de El Peñón (Andes), Santander, Colombia. Serie Editorial Fauna Silvestre Neotropical. Instituto de Investigación de Recursos Biológicos Alexander von Humboldt. Bogotá, D.C., Colombia, 99–156.
  • Casallas-Pabón D, Medellín-Ruiz MC, Martínez-Torres SD, Murcia-López MA (2013) Biota. Artrópodos. Filo Arthropoda. In: Muñoz-Saba Y, González-Sánchez I, Calvo-Roa N (Eds) Cavernas de Santander, Colombia: Guía de campo. Serie de Guías de Campo del Instituto de Ciencias Naturales No. 13 Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogotá D.C., Colombia, 157–240.
  • Centore P (2016) sRGB Centroids for the ISCC-NBS Colour System. Self Published, 21 pp.
  • García AF, Kury AB (2020) The neotropical harvestman genus Vima Hirst, with description of a new species from Colombia (Arachnida: Opiliones: Agoristenidae). Journal of Arachnology 48: 67–76. https://doi.org/10.1636/0161-8202-48.1.67
  • García AF, Villarreal O (2020) Description of a new species of Leptostygnus Mello-Leitão, 1940 and notes on the male genitalia in the subfamily Leiosteninae (Opiliones: Agoristenidae). Studies on Neotropical Fauna and Environment 1–15. https://doi.org/10.1080/01650521.2020.1724496.
  • González-Sponga MA (1981) Tres nuevas especies del genero Vima del sistema montañoso de la costa en Venezuela (Arachnida: Opiliones: Laniatores). Acta Macarao; Nueva Serie, Instituto Universitário Pedagógico de Caracas 2: 33–50.
  • González-Sponga MA (1987) Arácnidos de Venezuela. Opiliones Laniatores I. Familias Phalangodidae y Agoristenidae. Academia de Ciencias Físicas, Matemáticas y Naturales, Caracas, 562 pp.
  • González-Sponga MA (1998) Arácnidos de Venezuela. Cuatro nuevas especies de la familia Agoristenidae (Opiliones, Laniatores). Acta Biologica Venezuelica 18(3): 21–33.
  • Kury AB, Villarreal MO (2015) The prickly blade mapped: establishing homologies and a chaetotaxy for macrosetae of penis ventral plate in Gonyleptoidea (Arachnida, Opiliones, Laniatores). Zoological Journal of the Linnean Society 174(1): 1–46. https://doi.org/10.1111/zoj.12225
  • Olson DM, Dinerstein E, Wikramanayake ED, Burgess ND, Powell GVN, Underwood EC, D’amico JA, Itoua I, Strand HE, Morrison JC, Loucks CJ, Allnutt TF, Ricketts TH, Kura Y, Lamoreux JF, Wettengel WW, Hedao P, Kassem KR (2001) Terrestrial ecoregions of the world: a new map of life on earth. BioScience 51: 933–938. https://doi.org/10.1641/0006-3568(2001)051[0933:TEOTWA]2.0.CO;2
  • QGIS DEVELOPMENT TEAM (2020) QGIS geographic information system. Open Source Geospatial Foundation Project. http://qgis.osgeo.org
  • Villarreal O, García AF (2021) On the phylogenetic relationships of Muscopilio, a new Andean genus of basibiont harvestmen (Opiliones: Agoristenidae). Zoologischer Anzeiger 292: 150–162. https://doi.org/10.1016/j.jcz.2021.03.006