Research Article |
Corresponding author: Osvaldo Villarreal ( osvaldovillarreal@gmail.com ) Academic editor: Danilo Harms
© 2024 Osvaldo Villarreal, Daniela Ahumada-C., Gabriel R. Navas-S..
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.
Citation:
Villarreal O, Ahumada-C. D, Navas-S. GR (2024) Exploring the diversity of Eutimesius Roewer, 1913: new species and records from Colombia and Venezuela (Opiliones, Gonyleptoidea, Stygnidae). Zoosystematics and Evolution 100(3): 803-820. https://doi.org/10.3897/zse.100.120207
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This study contributes taxonomic information on the genus Eutimesius Roewer, 1913. Three new species are described: the sympatric species E. aroa sp. nov. and E. guaichia sp. nov. from Yaracuy state and E. canoabo sp. nov. from Carabobo state in Venezuela. The male of E. ephippiatus (Roewer, 1915) is described for the first time. Remarks are made on the distribution of E. ornatus, and a complimentary description of male genitalia is provided for E. simoni Roewer, 1913. New departmental records are added for E. ephippiatus and E. simoni. An updated distribution map and a key to the identification of the males of the species in the genus are presented.
Harvestmen, Heterostygninae, Laniatores, Neotropics
The family Stygnidae is a group of primarily mid-sized harvestmen with 114 species (
The genus Eutimesius Roewer, 1913 belongs to the subfamily Heterostygninae and is characterized by the presence of white, dry spots on the dorsal scutum. These dry spots bear a resemblance to those documented by Kury in
The five species currently included within the genus were originally classified under at least five different genera: Eutimesius; Dichobunistygnus Roewer, 1915; Hoplostygnus Roewer, 1915; Metaphareus Roewer, 1912; and Xanthostygnus Mello-Leitão, 1949.
Most of the species within the genus are restricted to the northern Andes of Colombia and Venezuela (
Only the type species of the genus occurs outside the Andes: Eutimesius simoni Roewer, 1913, which inhabits the Amazonas basin and has been recorded from Brazil, Colombia, Ecuador, and Peru. In Colombia, E. simoni is known to occur in the department of Putumayo (
Despite the relatively low number of described species in this genus, recent unpublished surveys suggest a significantly greater diversity, with at least seven undescribed species (OV unpublished data) known exclusively in the Venezuelan Andean cloud forests exhibiting high levels of endemism.
This paper describes three new species: two sympatric species from Yaracuy State and one from Carabobo State in Venezuela. Additionally, the description of the previously unknown male of E. ephippiatus, remarks on the distribution of E. ornatus, and a complimentary description of male genitalia for E. simoni also report new departamental records of E. ephippiatus and E. simoni in Colombia.
All measurements are in millimeters and were taken with a stereomicroscope, referring to the maximum length and width. Setiferous tubercles (i = small, I = large) on pedipalps are given in proximal to distal order. For color descriptions, we used the standard names of the 267 color centroids of the NBS/IBCC color system as named in
Abbreviations: AL – maximum abdominal scutum length; AW – maximum abdominal scutum width; BaCh – basichelicerite length; br – broken; CL – carapace length; ClPp – pedipalp claw; CoPp – pedipalpal coxa; CW – maximum carapace width; DS – dorsal scutum; DSW – dorsal scutum width; DSL – dorsal scutum length; FeL – femur length; FePp – pedipalpal femur; IOD – interocular distance; LP – lamina parva; MS – macrosetae of penis; PeTr – pedipalpal trochanter; Pp – pedipalps; PaPp – pedipalpal patella; TaPp – pedipalpal tarsus; TiPp – pedipalpal tibia; TiL – tibia length.
Depositories (curators):
Family Stygnidae Simon, 1879
Subfamily Heterostygninae Roewer, 1913
Eutimesius albicinctus (Roewer, 1915); Eutimesius aroa sp. nov.; Eutimesius canoabo sp. nov.; Eutimesius ephippiatus (Roewer, 1915); Eutimesius guaichia sp. nov.; Eutimesius ornatus (Rower, 1943); Eutimesius punctatatus (Roewer, 1913); Eutimesius simoni Roewer, 1913.
Heterostygninae with white, dry spot on the dorsal scutum. Carapace with interocular monticle or spine, and scutal area III with two acute paramedian spines. Genital “bauplan” as Heterostygninae, LP with distal cleft, and long basal “neck.” MS-B ventrally positioned (Fig.
Eutimesius sp., male (
(Fig.
(Fig.
The altitudinal range occupied by species of this genus is extensive, ranging from low elevations in the Amazon to heights reaching 3,050 meters in the Venezuelan Andes. However, the majority of species are found above 1,000 meters, typically in cloud forest habitats. Only E. simoni has been recorded in lowland areas between 70 and 110 meters in altitude, which is characteristic of the Amazonian region. E. ephippiatus occurs at the foothills of the Andes between 500 and 1,000 meters. All other species occur above 1,000 meters in altitude. The highest altitude record corresponds to a single female from Táchira state in Venezuela, misidentified as E. ornatus (
1 | Paramedian spines of the area III fused (Fig. |
E. simoni |
– | Paramedian spines of the area III separated (Figs |
2 |
2 |
DS yellowish background with conspicuous dark circular spots on the scutal areas ( |
E. punctatus |
– |
DS generally dark brown; rarely yellow, but in no case with that pattern of dark circular spots on the scutal areas (Figs |
3 |
3 | Tibia IV with a conspicuous prodorsal subdistal spine (Fig. |
E. canoabo sp. nov. |
– | Tibia IV without a conspicuous prodorsal subdistal spine (Figs |
4 |
4 | Scutal areas II–III divided (Figs |
5 |
– | Scutal areas II–III entire (Figs |
6 |
5 | Lateral margins of DS and inner margin of eyes without white dry spots (Pinto-da-Rocha, 1997: figs 57, 58); femora III greenish | E. ornatus |
– | Lateral margins of DS and inner margin of eyes with white dry spots, eyes completely encircled by white dry spots (Figs |
E. ephippiatus |
6 | Femur IV slightly curved and distally widened, with long and wide ventrodistal spines; patella IV as wide as long, with two distal very large dorsal tubercles; and tibia IV swollen, about two times as long as wide (Fig. |
E. guaichia sp. nov. |
– | Femur IV almost straight, with small ventrodistal tubercles; patella IV longer than wide, and tibia IV cylindrical and not swollen, between 5.5 and 9.6 times longer than wide | 7 |
7 | Pattern of small, white dry spots forming a lateral line on the margins of the DS and delineating the border of both halves of the scutal area I (Pinto-da-Rocha, 1997: figs 49, 50); tubercles of the pro and retroventral rows of the femora IV confined to the distal portion (Pinto-da-Rocha, 1997: fig. 53) | E. albicinctus |
– | Pattern of dry white spots not forming a lateral line on the DS margins; at most, they are only present in the anterior region of the carapace and do not delineate the border of both halves of the I area; instead, they are mainly confined to the lateral portions of the area I (Figs |
E. aroa sp. nov. |
Venezuela • ♂ holotype; Yaracuy, Road Cocorote – Aroa, sector Las Cumaraguas; (10.3520°N, 68.8298°W); 1,200 m a.s.l.; 09 Mar. 2008; (Villarreal O., Escalona H., Jayaro Y., Viera E. leg.) (
It is distinguishable from all other species in the genus by the pattern of dry white spots, occupying the anterolateral zone of the carapace, lateral areas of the II and III areas, and part of the lateral margins of the dorsal shield (Figs
The species is named in reference to the Sierra de Aroa, the mountain range in which it inhabits. Aroa is a Chibcha word believed to have a meaning associated with the jaguar, or “tigre.” Noun in apposition.
Male. Measurements. DSL 4.0; DSW 3.5; AL 1.9; AW 2.7; IOD 2.0; pedipalp: CoPp 0.8, TrPp 1.1, FePp 3.8, PaPp 1.8, TiPp 2.4, TaPp 2.2, ClPp 1.8; total 13.9; leg IV: FeL 7.2, TiL 3.9. Dorsum (Figs
Female. Measurements. Dorsal scutum length 4.1; dorsal scutum width 3.4; abdominal scutum length 1.8; abdominal scutum width 2.8; interocular distance 1.7; pedipalp: coxa 0.6, trochanter 0.8, femur 3.9, patella 1.6, tibia 2.0, tarsus 2.2, claw 1.7; total 12.8; leg IV: femur 7.3, tibia 3.9. Description (Fig.
Venezuela, Yaracuy. Only known from the type locality (Fig.
Venezuela • ♂ holotype; Carabobo, El Santuario, Posada Ecológica Casa María, near Canoabo, (10.3132°N, 68.2232°W); 1,220 m a.s.l.; 19 Mar. 2008; (Villarreal O., Pereira M.P. leg.); on vegetation abbott 1.5–2 m above the ground (
It is distinguishable from all other species in the genus by the pattern of dry white spots, occupying lateral zones of the scutal areas II and III, the medial zone of the scutal areas I and II, dispersal spots on the medial zone of the carapace, and part of the lateral margins of the dorsal scutum (Figs
Canoabo is an indigenous word of Arawako origin that means “village next to fresh water.” The species name refers to the type locality, a forest near Canoabo, a town and river of the Cordillera de la Costa in Carabobo State, Venezuela.
Male. Measurements. DSL 3.4; DSW 3.1; AL 1.8; AW 3.1; IOD 1.9; pedipalp: CoPp 0.5, TrPp 0.9, FePp 3.3, PaPp 1.5, TiPp 1.2, TaPp 1.1, ClPp 1.1; total 9.6; leg IV: FeL 7.4, TiL 3.6. Dorsum (Figs
Female. Measurements. Dorsal scutum length 3.9; dorsal scutum width 3.5; abdominal scutum length 1.9; abdominal scutum width 2.9; interocular distance 1.5; pedipalp: coxa 0.8, trochanter 0.7, femur 2.8, patella 1.2, tibia 1.8, tarsus 1.6, claw 1.3; total 10.2; leg IV: femur 8.0, tibia 3.4. Description (Fig.
Venezuela, Yaracuy. Only known from the type locality (Fig.
Dichobunistygnus ephippiatus Roewer C-F, (1915): 105, figs 57a–b.
Eutimesius ephippiatus:
Colombia • 1 ♂, 1 ♀; Boyacá (New department record), Santa María, sendero Hyca Quye, 5.5 km NW from Santa María, forest edge roadside; (4.89811°N, 73.29344°W); Ago. 2016; (Rodríguez, C. leg.) (MPUJ_ENT 004 8876). • 1 ♂, Boyacá, Santa María, Quebrada La Argentina, forest edge, 0.5 km SSW from Santa María; (4.85673°N, 73.26375°W); manual capture; 850 m a.s.l.; Ago-Sep. 2015; (Guzmán, K. leg.) (MPUJ_ENT 0039466). • 1 ♀; Casanare (new department record), Agua Azul; [5.1866°N, 72.5558°W]; 14 Oct. 1978; (La Rotta, C. leg.) (
It is distinguishable from all other species in the genus by the pattern of dry white spots that encircled completely the eyes (Figs
Male. Measurements. DSL: 1.4; DSW: 2.8; AL: 1.7; AW: 3.0; IOD: 1.6; pedipalp: CoPp 0.7, TrPp 0.7, FePp 2.9, PaPp 1.3, TiPp 1.5, TaPp 1.3, ClPp 0.8; total 9.2; Leg IV: FeL 7.6, TiL 2.6. Dorsum (Figs
Female. Measurements. DSL: 1.2–1.5; DSW: 2.5–3.2; AL: 1.5–2.2; AW: 2.9–3.5; IOD: 1.3–1.7; pedipalp: CoPp 0.4–0.8, TrPp 0.5–0.6, FePp 1.9–2.6, PaPp 1.0–1.4, TiPp 0.8–1.4, TaPp 0.8–1.3, ClPp 0.7–0-9, total 4.3–8.0; Leg IV: FeL 6.2–8.2, TiL 3.4–4.1. Description of female (Fig.
Colombia, Casanare, Cundinamarca, Quindio (Fig.
Holotype. Venezuela • ♂, Yaracuy, road Cocorote - Aroa, sector Las Cumaraguas; (10.3520°N, 68.8298°W); 1,200 m a.s.l.; 09 Mar. 2008; (Villarreal O., Escalona H., Jayaro Y., Viera E. leg.) (
It can be distinguished from other congeneric species by the pattern of dry white spots occupying the anterolateral zone of the carapace, the posterior zone of the eyes, and the lateroposterior corner of the DS (Figs
The species is named after Guaichía, another name given to María Lionza, a female deity belonging to Venezuelan spiritism, which originates from the state of Yaracuy, where the species inhabits. The myth of María Lionza has been interpreted as a symbol of the biological and cultural mestizaje, or intermixing, that characterizes Venezuela. It expresses the Indigenous, European, and African roots that comprise the cultural diversity of Venezuela. This is a noun in apposition.
Male. Measurements. DSL: 4.4; DSW: 3.7; AL: 1.8; AW: 3.0; IOD: 2.4; pedipalp: CoPp 0.7, TrPp 0.9, FePp 3.3, PaPp 1.3, TiPp 1.8, TaPp 1.7, ClPp 1.3, total 11.0; Leg IV: FeL 4.2, TiL 2.7. Dorsum (Figs
Female. Measurements. DSL: 4.3; DSW: 3.8; AL: 2.1; AW: 3.1; IOD: 1.8; pedipalp: CoPp 0.7, TrPp 0.5, FePp 2.9, PaPp 1.4, TiPp 1.7, TaPp 1.8, ClPp 1.2, total 10.2; Leg IV: FeL 4.7, TiL 3.0. Description (Fig.
Venezuela, Yaracuy. Only known for the type locality (Fig.
Bunistygnellus ornatus Roewer, C.-F. (1943): 37, figs 37–37a.
Eutimesius ornatus:
Eutimesius ornatus (Roewer, 1943) was described from Bogotá, Colombia (
Eutimesius simoni Roewer, C.-F. (1913): 453, figs 178.
Xanthostygnus fractus Mello-Leitão, C.F. (1949): 32.
Xanthostygnus fractus
Mello-Leitão, 1949 is a junior subjective synonym of Eutimesius simoni Roewer, 1913 in
Eutimesius miles Henriksen, K.L. (1932): 293, fig. 10.
Eutimesius miles
Sørensen, 1932, is a junior subjective synonym of Eutimesius simoni Roewer, 1913, in
Colombia • 1 ♂; Amazonas (New department record), Leticia, Km 11 vía a Tarapacá; [4.1209°S, 69.9508°W]; (Animal systematics [students leg.]) (
Can be distinguished from all other species of the genus by the fusion of the paired spines of scutal area III (Fig.
Male. Measurements. Males (
Subtle variations were observed in the shape of the LP, which presents less pronounced and rounded lateral projections (instead of acute), a wider neck, and the MS-A, which are separated from each other, with one pair positioned almost ventrally and the other almost dorsally (Fig.
Eutimesius cf. simoni: A. From Puerto Montúfar, Sucumbíos, Ecuador; B–D. From Iquitos, Maynas, Peru; E. From Leticia, Amazonas, Colombia; F. From Parque Nacional Yasuní Tipu Tini, Napo, Ecuador. Photographs by: Mark Silverstein (A), Jiri Hodecek (B), Martin Cejka (C), Phill Kahler (D), Traveler Dan (E), and Cinthya Villegas (F).
Brazil, Amazonas; Colombia, Amazonas; Ecuador, Napo, and Los Ríos; Peru, Loreto (Fig.
Within Heterostygninae, at least four genera share a similar genital morphology, namely Eutimesius and Innoxius Pinto-da-Rocha, 1997, from the Andean region; Minax Pinto-da-Rocha, 1997; and Yapacana Pinto-da-Rocha, 1997, from the Amazon. The latter two can be distinguished from Eutimesius by the reduction and shape of the LP and the widening of the malleus in the former, and by the increase in the number of MS-A and the loss of the dorsal process of the stylus in Minax. Meanwhile, Innoxius is separated from Eutimesius solely by somatic characters, including the DS shape, the ornamentation of scutal areas, and the presence of white dry spots (
The monophyly of Eutimesius was demonstrated by
Despite raising questions about intergeneric relationships in our work, we have chosen not to conduct a phylogenetic analysis in this study due to several key factors. Recent taxonomic changes and the reinterpretation of genital traits in the Stygnidae family have occurred independently of a phylogenetic analysis (e.g.,
Although the need for a complete phylogenetic analysis of the family is increasingly evident, this goes beyond the main objective of our present work and is part of future projects that involve a broader group of authors. In addition, due to the nature of the group studied, the lack of this analysis does not limit or affect the results obtained in this work.
All authors of this work contributed significantly to the project. OV designed and led the project, collecting Venezuelan species, diagnosing new species from Venezuela and Colombia, and working on the description, photography, and illustration of the new species. DAC visited Colombian collections, analyzed the material from this country, and generated the corresponding figures. DAC also contributed to the description of the new species and created the map. GRN provided the facilities for the project to be carried out, including funding for the visits to collections and laboratory facilities. All authors worked together on the writing of the article and agreed on the final version of the manuscript presented.
Claudia Medina, scientific manager of the collection at the Instituto Alexander von Humboldt, coordinated the loan of material to Adriano B. Kury. We would like to thank Adriano B. Kury for his invaluable assistance during our research in his laboratory (MNRJ), as well as for his outstanding coordination in arranging the loan of materials from the Instituto Alexander von Humboldt. To Eduardo Flórez, Daniela Martínez, and Sebastian Galvis for their collaboration during the visit of DAC and OV to the arachnological collection of ICN. To Julian Clavijo-Bustos for his collaboration during the visit of the DAC to the