Monograph |
Academic editor: Gonzalo Giribet
© 2021 Marília Pessoa-Silva, Marcos Ryotaro Hara, Ricardo Pinto-da-Rocha.
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:
Pessoa-Silva M, Hara MR, Pinto-da-Rocha R (2021) Revision of the southern Andean genus Sadocus Sørensen, 1886 (Opiliones, Gonyleptidae, Pachylinae). ZooKeys 1025: 91-137. https://doi.org/10.3897/zookeys.1025.57806
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Species of the genus Sadocus Sørensen, 1886 are conspicuous gonyleptids that occur in Chile and Argentina. Here, the genus is revised for the first time and the cladistic analysis based on morphological characters does not corroborate its monophyly unless a phylogenetically unrelated species is excluded (explained further on). A new classification is proposed for the seven species left in the genus and considered valid, of the 13 nominal species previously recognized. Two out of the seven valid species are considered as species inquirendae: Sadocus allermayeri (Mello-Leitão, 1945) [= Carampangue allermayeri Mello-Leitão, 1945] and Sadocus nigronotatus (Mello-Leitão, 1943) [= Carampangue nigronotatum Mello-Leitão, 1943]. The following synonymies are proposed: Sadocus bicornis (Gervais, 1849) [original combination = Gonyleptes bicornis Gervais, 1849] is a junior synonym of Sadocus asperatus (Gervais, 1847) [= Gonyleptes asperatus Gervais, 1847]; Sadocus conspicillatus Roewer, 1913, Sadocus exceptionalis (Mello-Leitão, 1946) [= Araucanoleptes exceptionalis Mello-Leitão, 1946] and Sadocus guttatus Sørensen, 1902 are junior synonyms of the valid name Sadocus polyacanthus (Gervais, 1847) [= Gonyleptes polyacanthus Gervais, 1847]; and Sadocus calcar (Roewer, 1913) [= Lycomedes calcar Roewer, 1913] is a junior synonym of the valid name Gonyleptes horridus Kirby, 1819. Sadocus brasiliensis Soares & Soares, 1949 is not congeneric with Argentinean/Chilean species of the genus according to the cladistic analysis and is here synonymized with Discocyrtus catharinensis (Mello-Leitão, 1923 [= Sadocus catharinensis Mello-Leitão, 1923]).
Argentina, Chile, harvestmen
Harvestman systematics has advanced greatly in the last few decades, especially in the Neotropical region, with many supraspecific groups being recently revised, such as for example Stygnidae Simon, 1879 (
Pachylinae is the most species-rich subfamily of Gonyleptidae, and is currently considered polyphyletic (
The sister group of Pachylinae sensu stricto is a clade that includes the genus Sadocus Sørensen, 1886, composed of rather large-sized (5.5–13.8 mm of dorsal scutum length) and colorful harvestmen. Although conspicuous and relatively common in Chilean preserved areas, it was never revised in more than 130 years of existence. Historically (see below), the genus has been subjected to many taxonomic acts, resulting in confusing species identities. One has to use poor (for modern standards), hundred-year-old descriptions to identify a given species. In addition, similar species are difficult to distinguish, raising doubts about their identities. Therefore, the revision of Sadocus focuses on determining the identity of the included species, which in turn will allow further understanding of their relationships, and more precise inferences of their distribution and diversity (
The history of Sadocus Sørensen, 1886 can be quite confusing because many of its species were described before the proposition of the genus. Therefore, this historical section mentions many species in different genera and subfamilies that were later transferred to Sadocus (
In 1886, Sørensen proposed the monotypic genus Sadocus, to include the type and new species S. vitellinosulcatus. In 1899, Loman described Discocyrtus calcitrosus and Gonyleptes platei, which are relevant to Sadocus, and transferred G. funestus to Discocyrtus Holmberg, 1878.
In 1902, Sørensen: (i) synonymized S. vitellinosulcatus and G. platei Loman, 1899 with G. polyacanthus; (ii) created the new genus Lycomedes (without indication of a type species), to which he transferred G. asperatus, G. bicornis, D. calcitrosus, D. funestus and Pachylus planiceps; and (iii) described Sadocus guttatus. On that paper, Sørensen placed these species in Gonyleptidae, but without assigning them to any subfamily. Therefore, at the beginning of the 20th century, named species relevant to Sadocus were placed in both Sadocus, comprising two species (S. polyacanthus and S. guttatus) and Lycomedes, with five species (L. asperatus, L. bicornis, L. calcitrosus, L. funestus and L. planiceps).
In 1945, Mello-Leitão described Carampangue allermayeri, and in the next year (1946), he described Araucanoleptes for the new species A. exceptionalis, and placed it in the Gonyleptinae. A few years later,
At the beginning of the 21st century (prior to this study), species relevant to Sadocus were placed in the following genera and subfamilies: the monotypic Araucanoleptes (Gonyleptinae); Carampangue (Pachylinae), with three species (C. allermayeri, C. ingens and C. nigronotatum); Lycomedicus (Pachylinae), with seven species (L. asperatus, L. bicornis, L. brasiliensis, L. calcar, L. dilatatus, L. funestus and L. planiceps); and Sadocus (Gonyleptinae), with three species (S. conspicillatus, S. guttatus, and S. polyacanthus).
Material examined belongs to the following institutions (curators in parentheses) listed below:
URMU Museo Nacional de Historia Natural de Montevideo, Montevideo, Uruguay (M. Simó);
The following abbreviations are used throughout the text, including synonymic listings:
cat catalogue;
cit citation;
coll collected;
desc description;
eco ecology;
rdesc redescription;
syst systematic discussion.
In the examined material:
fe female;
ma male;
juv juvenile;
MS A–E penis ventral plate pairs of macrosetae A–E.
[X(y)] where X is the character number and y, the character state.
The topological nomenclature follows
We chose the outgroups based on available hypotheses including Pachylinae, such as
Acanthopachylus aculeatus (Kirby, 1819) (Gonyleptidae: Pachylinae) (
Acanthoprocta conica Maury, 1991 (Gonyleptidae: Pachylinae) (
Goniosoma varium Perty, 1833 (Gonyleptidae: Goniosomatinae) (
Gonyleptes horridus Kirby, 1819 (Gonyleptidae: Gonyleptinae) (
Metagyndes pulchella (Loman, 1899) (Gonyleptidae: Pachylinae) (
Nanophareus polyhastatus Hara, 2016 (Gonyleptidae: Pachylinae) (
Neogonyleptes docilis (Butler, 1876) (Gonyleptidae: Pachylinae) (
Neogonyleptes karschii (Sørensen, 1902) (Gonyleptidae: Pachylinae) (
Pachylus chilensis (Gray, 1833) (Gonyleptidae: Pachylinae) (
Pachyloides thorellii Holmberg, 1878 (Gonyleptidae: Pachylinae) (
Roeweria bittencourti Mello-Leitão, 1923 (Gonyleptidae: Roeweriinae) (
Stygnus polyacanthus (Mello-Leitão, 1923) (Stygnidae: Stygninae) (
We include records of distribution in maps only for vials with males. We used LibreOffice Calc to edit the character matrix and TNT 1.0 (
To test the monophyly of Sadocus, we used a matrix of morphological characters composed of 18 taxa (13 outgroups and five ingroups) and 64 characters (Table
The single most parsimonious tree retrieved in the cladistic analysis representing Sadocus relationships (182 steps, L = 182; C.I. = 45; R.I. = 53). Number at the nodes are Goodman-Bremer support. Black circles indicate unique transformations, while white circles indicate homoplastic transformations, the arrow indicates the position of the genus Sadocus. The character number is above each circle, and the character state is below. The characters are optimized in ACCTRAN. The character and character states are given in Table
List of character and character states used in the cladistic analysis. All characters of legs and genitalia refer to male.
Character | State | |
---|---|---|
1 | (DS) Ocularium ( |
0. Divided, each eye placed onto different elevations; 1. Single. |
2 | (DS) Ocularium, unpaired armature: | 0. Absent; 1. Present. |
3 | (DS) Ocularium, paired armature: | 0. Absent; 1. Present. |
4 | (DS) Anterior margin of carapace, frontal hump | 0. Inconspicuous; i.e., straight from ocularium to anterior margin of DS in lateral view 1. Conspicuous; i.e., clear elevation from ocularium to anterior margin of DS in lateral view |
5 | (DS) Mesotergum: placement of the maximum width | 0. Maximum width in the middle of the mesotergum; 1. Maximum width placed posteriorly to the middle of mesotergum |
6 | (DS) Dorsal scutum length and width ratio | 0. Wider than long; 1. Longer than wide |
7 | (DS) Posterior margin, shape | 0. Straight; 1. Concave; 2. Convex |
8 | (DS) Area I, state of fusion | 0. Divided in right and left halves by a longitudinal groove between scutal areas I – II (even though the groove of area II slightly invades area I) 1. Divided in right and left halves by invasion of scutal area II into middle of scutal area I. |
9 | (DS) Scutal area I,paramedian armature ( |
0. Absent or with similar sized granules; 1. With a pair of tubercles |
10 | (DS) Scutal area II, paramedian paired armature ( |
0. Absent or with similar sized granules; 1. With a pair of tubercles |
11 | (DS) Scutal area III, paramedian paired armature ( |
0. Absent or with similar sized granules; 1. With a paramedian pair of tubercles |
12 | (DS) Scutal area IV, presence | 0. Absent; 1. Present. |
13 | (DS) Scutal area IV, degree of division | 0. Incompletely divided; i.e., interrupted scutal groove IV; 1. Completely divided |
14 | (DS) Scutal area IV, paramedian paired armature | 0. Absent; 1. Present. |
15 | (DS) Lateral margin, type of integumentary ornamentation | 0. Covered with granules; 1. With tubercles, sometimes clustered |
16 | (DS) Lateral margin, type of armature | 0. Large tubercles or apophyses; 1. similar sized tubercles |
17 | (DS) Posterior margin of the DS, paramedian armature | 0. Absent; 1. Present. |
18 | (DS) Posterior margin of the DS, central unpaired armature | 0. Absent; 1. Present. |
19 | (DS) Free tergites I, paramedian armature | 0. Absent; 1. Present. |
20 | (DS) Free tergites II, paramedian armature | 0. Absent; 1. Present. |
21 | (DS) Free tergites II, unpaired armature | 0. Absent; 1. Present. |
22 | (DS) Free tergites III, paramedian paired armature | 0. Absent; 1. Present. |
23 | (Chelicerae) Chelicerae, sexual dimorphism (#30, |
0. Isomorphic in both sexes 1. Large in male |
24 | (Pedipalp) Tibia, type of retro-lateral apical seta | 0. Single; 1. Bifid |
25 | (Leg) Coxa IV, branch of the prodorsal apophysis | 0. Single; 1. Bifid |
26 | (Leg) Coxa IV, insertion of the prodorsal apical apophysis in relation to the DS main axis ( |
0. Almost transversal; i.e., almost 90 degrees in relation to DS main axis; 1. Oblique; i.e., more than 120 degrees in relation to DS main axis; 2. Parallel to femur IV |
27 | (Leg) Coxa IV, retro-apical apophysis: | 0. Absent; 1. Present. |
28 | (Leg) Leg IV, torsion that begins at the trochanter and ends at the patella | 0. Untwisted; 1. Strongly twisted; i.e., prolateral features becoming dorsal in situ and gradually untwisting towards patella; 2. Strongly twisted from coxa IV towards the patella (Fig. |
29 | (Leg) Trochanter IV, Prolateral basal apophysis (# 30 |
0. Absent; 1. Present. |
30 | (Leg) Trochanter IV, dorso-median subapical apophysis | 0. Absent; 1. Present. |
31 | (Leg) Trochanter IV, prodorsal apical apophysis | 0. Absent; 1. Present. |
32 | (Leg) Trochanter IV, type of prodorsal apical apophysis | 0. As a wart; 1. As a hook-like pointed apophysis of large base, smoothly becoming pointed apically; 2. As a moderate size blunt cone; 3. As a finger shaped, robust apophysis, basal half of ca. uniform diameter |
33 | (Leg) Trochanter IV, retro-dorsal apical apophysis (# 29 |
0. Absent; 1. Present. |
34 | (Leg) Trochanter IV, retro-apical armature ( |
0. Absent; 1. Present. |
35 | (Leg) Trochanter IV, type of retro-apical armature | 0. Pointed tubercle; 1. Moderate apophysis (ca. a quarter of podomere width); 2. Huge apophysis (as long as podomere width). |
36 | (Leg) Trochanter IV, proapical apophysis | 0. Absent; 1. Present. |
37 | (Leg) Trochanter IV, length-width ratio (modified from |
0. As long as wide; 1. Twice longer than wide; 2. Wider than long. |
38 | (Leg) Femur IV, curvature in dorsal view | 0. Straight; 1. Sinuous |
39 | (Leg) Femur IV, size of granules on retro-lateral row | 0. Similar sized granules; 1. Tubercles, twice the size of granules. |
40 | (Leg) Femur IV, spiniform apophyses on basal half of the retro-lateral row of granules | 0. Absent; 1. Present. |
41 | (Leg) Femur IV, the pattern of apophyses distribution at the ⅔ basal region of the retro-lateral row | 0. Just an apophysis in the basal ⅓; 1. Growing from the median region to the basal region; 2. Very high apophyses alternating with low apophyses; 3. Apophyses distributed in the median region; 4. A basal apophysis and one or more in the distal ⅔; 5. An average apophysis; 6. Decreasing from the median region to the basal region. |
42 | (Leg) Patella IV, ventral row of granules | 0. Similar sized granules; 1. Granules becoming tubercles or spines. |
43 | (Leg) Tibia IV, ventro-basal long spine | 0. Absent; 1. Present. |
44 | (Leg) Tibia IV, size of granules on retro-ventral row | 0. Tubercles of similar sizes; 1. Granules increasing in size apically, becoming tubercles. |
45 | (Leg) Tibia IV, proventral row of tubercles size | 0. Tubercles of similar sizes; 1. Larger tubercles, which grows in size apically. |
46 | (Leg) Tibia IV, size of granules on ventral row | 0. Similar sized granules; 1. Increasing in size apically. |
47 | (Penis) Ventral plate, shape of the distal margin | 0. Straight; 1. Slightly concave; 2. Very concave, forming a “U”. |
48 | (Penis) Ventral plate, basal lobes | 0. Inconspicuous; 1. Conspicuous. |
49 | (Penis) Ventral plate, plate format | 0. Rectangular; 1. Hexagonal. |
50 | (Penis) Ventral plate, number of MS C | 0. Three pairs; 1. Four pairs or more. |
51 | (Penis) Glans, dorsal prominence in the distal region of the sac | 0. Absent; 1. Present. |
52 | (Penis) Glans, sac texture | 0. Smooth and turgid; 1. Wrinkled. |
53 | (Penis) Glans, latero-apical region | 0. Without projections; 1. With projections covering part or all of the pedestal in lateral view. |
54 | (Penis) Glans, dorsal process | 0. Absent; 1. Present. |
55 | (Penis) Glans, ventral process | 0. Absent; 1. Present. |
56 | (Penis) Ventral process, presence of stem | 0. Absent; 1. Present. |
57 | (Penis) Ventral process, apex shape | 0. As a flabellum ; 1. Tapered at the apex and rolled; 2. Flattened circular; 3. Flattened quadrangular; 4. Fringed triangular; 5. Large rectangular; 6. Rectangular bifid; 7. Rectangular with pointed projections. |
58 | (Penis) Stylus, ventral process length ratio | 0. Stylus shorter than ventral process; 1. Stylus longer than ventral process. |
59 | (Penis) Stylus, apical lateral projections | 0. Absent; 1. Present. |
60 | (Penis) Stylus, apex shape ( |
0. Rounded; 1. With an apical back beak. |
61 | (Penis) Stylus, trichomes on median apical region | 0. Absent; 1. Present. |
62 | (Penis) Insertion on the glans in lateral view | 0. Ventral; 1. Median |
63 | (Penis) Trunk of the penis, subapical region | 0. Truncated; 1. Projected on the glans |
64 | (Color) Carapace, presence of dry-mark | 0. Absent; 1. Present. |
According to the retrieved tree, Sadocus is not monophyletic, as it excludes Sadocus brasiliensis (Soares & Soares, 1949).
Matrix of character states for the cladistic analysis of the Sadocus (Gonyleptidae: Pachylinae).
Stygnus polyacanthus | 0 | 1 | 0 | 0 | 0 | 1 | 2 | 0 | 1 | 0 | 1 | 0 | - | - | 0 | 1 | 0 | 0 | 1 | 1 | 0 | 1 | 1 | 0 | 0 | 1 | 0 | 0 | 0 | 1 | 0 | - | 0 | 0 | - | 0 | 2 | 0 | 0 | 0 | - | 1 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 1 | 0 | - | - | - | - | 0 | 0 | 1 | 0 | 0 |
Acanthopachylus aculeatus | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 0 | 0 | - | 0 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 4 | 1 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 0 | 4 | 0 | 0 | 0 | 0 | 0 | 1 | 0 |
Acanthoprocta conica | 1 | 0 | 0 | 1 | 0 | 1 | 2 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 2 | 0 | 0 | 1 | 0 | 1 | 0 | 0 | 1 | 2 | 0 | 2 | 1 | 0 | 0 | - | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 0 | 1 | 0 | 4 | 1 | 0 | 0 | 0 | 0 | 0 | 0 |
Goniosoma varium | 1 | 0 | 1 | 1 | 1 | 0 | 1 | 1 | 1 | 0 | 1 | 0 | - | - | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 1 | 0 | 0 | - | 0 | 0 | - | 1 | 0 | 1 | 0 | 0 | - | 0 | 0 | 1 | 1 | 0 | 0 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 | 5 | 0 | 0 | 1 | 0 | 0 | 1 | 1 |
Gonyleptes horridus | 1 | 0 | 1 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 1 | 0 | - | - | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 1 | 0 | 1 | 1 | 0 | - | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 1 | 6 | 0 | 0 | 1 | 1 | 0 | 2 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | 1 | 1 | 0 | 1 | 0 | 0 | 1 | 1 | 1 | 0 |
Metagyndes martensii | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | - | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 3 | 1 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 1 | 0 | 1 | 0 | 0 | 1 | 0 | 4 | 1 | 0 | 0 | 0 | 1 | 0 | 0 |
Nanophareus polyhastatus | 1 | 0 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 1 | 0 | 0 | 1 | 1 | 0 | - | 0 | 1 | 1 | 1 | 0 | 1 | 1 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 1 | 7 | 0 | 0 | 0 | 1 | 1 | 0 | 0 |
Neogonyleptes docilis | 1 | 0 | 0 | 1 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | - | 0 | 0 | 1 | 0 | 1 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 1 | 0 | 1 | 1 | 2 | 1 | 0 | 0 | 0 | 1 | 0 | 0 |
Neogonyleptes karschii | 1 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | - | - | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 1 | 0 | 0 | 0 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | - | 1 | 1 | 0 | 0 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 1 | 2 | 1 | 0 | 0 | 0 | 1 | 0 | 0 |
Pachyloides thorellii | 1 | 0 | 1 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 0 | 0 | - | 0 | 0 | - | ? | 0 | 0 | 0 | 0 | - | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 1 | 1 | 3 | 1 | 0 | 0 | 0 | 0 | 0 | 0 |
Pachylus chilensis | 1 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 1 | 1 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 1 | 0 | 0 | - | 0 | 1 | 1 | 0 | 2 | 1 | 0 | 1 | 5 | 1 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 1 | 0 | 4 | 1 | 0 | 0 | 0 | 0 | 1 | 0 |
Roeweria bittencourti | 1 | 0 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 1 | 1 | 0 | 0 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 1 | 1 | 2 | 0 | 0 | 0 | - | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | - | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 0 | 0 | 1 | 0 | - | - | 1 | 0 | 0 | 0 | 1 | 0 |
Discocyrtus catharinensis | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 1 | 0 | 1 | 1 | 0 | 0 | 1 | 0 | 1 | 0 | 1 | 1 | 0 | 1 | 0 | 0 | 1 | 1 | 1 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 0 | 1 | 1 | 1 | 6 | 1 | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | 1 | 0 | 0 | 1 | 1 | 1 | 0 | 1 | 0 | 0 | 1 | 1 | 0 |
Sadocus asperatus | 1 | 0 | 1 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 0 | 1 | 0 | 0 | 1 | 1 | 1 | 1 | 1 | 0 | 1 | 3 | 1 | 1 | 0 | 0 | 1 | 1 | 0 | 1 | 2 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 1 | 2 | 1 | 0 | 0 | 0 | 0 | 0 | 1 |
Sadocus dilatatus | 1 | 0 | 1 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 0 | 1 | 0 | 0 | 1 | 1 | 1 | 1 | 1 | 0 | 1 | 3 | 0 | 1 | 0 | 0 | 1 | 0 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 1 | 2 | 1 | 0 | 0 | 0 | 0 | 0 | 1 |
Sadocus funestus | 1 | 0 | 1 | 1 | 1 | 0 | 1 | 0 | 1 | 1 | 1 | 1 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 | 1 | 2 | 0 | 1 | 0 | 1 | 0 | 0 | 1 | 0 | - | 1 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 1 | 2 | 1 | 0 | 0 | 0 | 0 | 0 | 1 |
Sadocus ingens | 1 | 0 | 1 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 1 | 0 | 1 | 0 | 0 | 1 | 1 | 1 | 1 | 1 | 0 | 1 | 3 | 0 | 0 | - | 0 | 1 | 1 | 0 | 1 | 1 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 1 | 2 | 1 | 0 | 0 | 0 | 0 | 0 | 0 |
Sadocus polyacanthus | 1 | 0 | 1 | 1 | 1 | 0 | 1 | 0 | 1 | 0 | 1 | 0 | - | - | 1 | 0 | 0 | 0 | 0 | 1 | 0 | 1 | 0 | 0 | 1 | 1 | 1 | 1 | 1 | 0 | 1 | 3 | 0 | 1 | 1 | 0 | 1 | 0 | 1 | 0 | - | 1 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 1 | 2 | 1 | 0 | 0 | 0 | 0 | 0 | 0 |
Once we settled the issue related to S. brasiliensis, we propose a new concept of Sadocus. Under the new definition, Sadocus is monophyletic and supported by seven synapomorphies, four of which are exclusive: leg IV twisted from the trochanter to patella [28(0)]; trochanter IV with a finger shaped, robust prodorsal apical apophysis, its basal half of ca. uniform diameter [32(3)], trochanter IV twice longer than wide [37(1)] (modified from
So far, Sadocus (represented especially by S. polyacanthus, its type species) has often been used in cladistic analysis as outgroups (
The sister taxon closest to Sadocus is also an unsettled issue, mainly because different taxa are employed in those analyses. In the present analysis, Sadocus sister group is the Chilean genus Neogonyleptes, supported by seven synapomophies, two of them exclusive: ventral row of granules increasing in size apically on tibia IV [46(1)]; and apex of glans ventral process flattened circle shaped [57(2)]. This sister group relationship is similar to
The main goals of this study were to revise and to test the monophyly of Sadocus, a hundred-year-old genus, with a convoluted taxonomic history. We believe that we succeeded in those, and the present study is an important step towards the understanding the evolution of the genus. Considering all the evidence (including the taxonomic history), Sadocus seems to be related to the Chilean-Argentinean Pachylinae. We understand that Sadocus relationship within Gonyleptidae is still an unsettled issue that deserves further investigation. As a mean to tackle that, we can suggest the inclusion of more Chilean Pachylinae genera (especially those already used in previous analyses and not belonging to Pachylinae sensu stricto) and Brazilian species as well, such as DRMN (
Pachylinae Sørensen, 1884
Gonyleptes
[part]: Gervais, 1842: 2 [rdesc]; 1844: 105 [rdesc]; 1847: 576–577 [cit]; 1849: 21, 24–26 [desc, rdesc];
Discocyrtus [part]: Loman, 1899: 6 [desc].
Sadocus
Sørensen, 1886: 85 [desc]; 1902: 14–20 [rdesc];
Lycomedes
Sørensen, 1902: 17 [desc];
Lycomedicus
Roewer, 1923: 442 (nom. nov. for Lycomedes Sørensen, 1902, rdesc); 1925: 17 [cit]; 1929: 213 [cat];
Lycomedius
[lapsus]: Kästner, 1937: 389 [rdesc];
Carampangue
Mello-Leitão, 1937: 152 [desc]; 1945: 156 [cit]; 1949: 17 [syst];
Jighas
Roewer, 1943: 28 [desc];
Araucanoleptes
Mello-Leitão, 1946: 4 [desc];
Arauconoleptes [lapsus]: Cekalovic, 1985: 12 [cat].
Sadocus vitellinosulcatus Sørensen, 1886, by monotypy. Synonymized with S. polyacanthus by
S. asperatus (Gervais, 1847), S. dilatatus Roewer, 1913, S. funestus (Butler, 1874), S. ingens (Mello-Leitão, 1937) and S. polyacanthus (Gervais, 1847).
Sadocus are large Pachylinae (dorsal scutum maximum length 5.5–13.8 mm) with paired spines on ocularium and prominent frontal hump on dorsal scutum anterior margin. Dorsal scutum shape types gamma triangular and gamma pyriform, its posterior margin concave. Dorsal scutum mid-bulge placed close to scutal groove IV (scutal groove III in S. funestus) and transversal (S. funestus, S. ingens, S. polyacanthus) or oblique (S. asperatus, S. dilatatus); free tergites II and III each with a pair of spines. Legs IV are twisted retro-laterad from the trochanter and gradually distorted along the femur and patella (except in S. funestus). Coxa IV bearing a long, large prodorsal apical apophysis and a short, retro-apical one (except in S. funestus, in which is lacking). Trochanter IV with a short, blunt prolateral sub-basal apophysis and a long, robust prodorsal apical one. Penis glans turgid and dorsally projected (with antero-lateral projections), with ventral process (half stylus length) and without dorsal process. General color (in living specimens) of the body and most parts of legs and ventral area dark brown, with lighter tones at the tips of podomeres. Yellowish to reddish tone in scutal area, scutal posterior margin, free tergites, part of legs and apophysis. Green on the arthrodial membranes between the free tergites.
Male: Dorsum. Anterior margin of carapace with a prominent median frontal hump (bell shaped in dorsal view). Ocularium with one pair of spines posterior to the eyes. Dorsal scutum type varying from gamma to gamma triangular and gamma pyriform, its posterior margin concave, mid-bulge slightly asymmetrical and displaced posteriorly, widest at the scutal groove IV (scutal groove III in S. funestus). The curvature of mid-bulge can be transversal (S. funestus, S. ingens, and S. polyacanthus) or oblique (S. asperatus and S. dilatatus). Four scutal areas (three in S. polyacanthus); scutal area I divided into right and left halves by a longitudinal median groove. Scutal area III with one pair of paramedian spiniform tubercles or spines. Two pairs of ozopores close to coxa II. Lateral margin of dorsal scutum with an external and internal rows of tubercles (the external row of slightly larger tubercles) (except S. asperatus, with granules covering most of the lateral margin of dorsal scutum and S. ingens, smooth or with only few granules). Posterior margin of dorsal scutum and free tergite I each with one paramedian pair of tubercles (except S. funestus and S. polyacanthus, unarmed). Free tergites II and III each with one paramedian pair of spines. Venter. Coxa I–IV granulate; coxa I with a median longitudinal row of granules increasing in size apically, becoming tubercles. Chelicerae. Isomorphic in males and females. Segment I with well-marked bulla. Segment II fixed finger and segment III toothed. Pedipalps. Trochanter dorsal face inflated; ventral face with one or two setiferous tubercles. Femur bearing sub-apical mesal seta; dorsal face with few granules; ventral face with one basal setiferous tubercle. Tibiae and tarsi dorsal and lateral faces with few minute granules and variable setation. Legs. Coxae I–III each with one prodorsal and one retro-dorsal spiniform tubercles, ventral faces granulate (except S. polyacanthus, coxa I with tubercles and others with setae). Coxa IV dorso-lateral face with sparsely distributed granules, ventral face entirely granulated, with one long, oblique, bifid prodorsal apical apophysis (transversal in S. dilatatus, uniramous in S. funestus), dorsal branch longest and curved ventrad and ventral branch short and blunt; and one ventro-apical retro-lateral spine. Trochanters I–III granulate. Leg IV twisted retro-laterad from the trochanter, gradually untwisting along the femur (except S. funestus, straight). Trochanter IV longer than wide; prolateral face with one short, conical, blunt sub-basal apophysis, and one robust, blunt dorso-apical apophysis. Femora I–IV with granules roughly organized in six longitudinal rows (prodorsal, retro-dorsal, pro- and retro-lateral, proventral and retro-ventral rows); femora I and II unarmed. Femur IV curved, with marked inner curvature on the distal half (S. asperatus and S. ingens) or almost straight (S. dilatatus, S. funestus, and S. polyacanthus). Patellae I–III granulate, unarmed; patella IV dorsal face granulate, ventral face tuberculate. Tibiae I–III granulate, unarmed (except S. dilatatus and S. funestus, tibia III dorsal face granulate with retro-ventral row of tubercles increasing in size apically). Tibia IV dorsal face granulate, ventral face with tubercles sparsely distributed. Metatarsi I–IV minute granulate, unarmed. Tarsus III and IV each with ventral process, tarsal claws smooth. Penis. Ventral plate distal margin with slight (but conspicuous) to moderate concavity, two or three pairs of MS A, one pair of MS B or entirely absent, four or five pairs of MS C, one or two pairs of MS D, and one or two pairs of MS E. Glans sac tall, turgid, dorsally projected with antero-lateral projections, forming a sheath for the stylus. Glans without dorsal process; stylus inserted ventrally and smooth. Glans ventral process is short (half of stylus length), parallel to the stylus, apex curved ventrad, with a short semi-circular antero-lateral projection.
(Fig.
1 | Coxa IV with a bifid prodorsal apical apophysis (Fig. |
2 |
– | Coxa IV with an unbranched prodorsal apical apophysis (Fig. |
S. funestus |
2 | Lateral margin of the dorsal scutum with a posterior large tubercle (Figs |
3 |
– | Lateral margin of the dorsal scutum only with similar sized tubercles (Fig. |
4 |
3 | Dorsal scutum with four areas (Fig. |
S. dilatatus |
– | Dorsal scutum with three areas (Fig. |
S. polyacanthus |
4 | Trochanter IV dorso-apical face only with one prolateral apophysis of similar length as the podomere, strongly curved (in lateral view), pointing frontwards (Fig. |
S. ingens |
– | Trochanter IV dorso-apical face with two apophyses, both shorter than the podomere length, femur IV slightly curved (Fig. |
S. asperatus |
Gonyleptes asperatus
Gervais, 1847: 577 [desc]; 1849: 26, pl. 1, fig 9 [rdesc];
Lycomedes asperatus: Sørensen, 1902: 17 [syst];
Lycomedicus asperatus: Roewer, 1923: 442, fig 556 [rdesc]; 1925: 17 [cit]; 1929: 213 [cat];
Sadocus asperatus: Kury, 2003: 191 [cat];
Sadocus
(?) subsimilis: Sørensen, 1902: 17 [cit];
Gonyleptes polyacanthoides
Gervais, 1847: 577 [desc];
Gonyleptes subsimilis
Gervais, 1849 [desc]: 25, pl. 1, fig 8;
Gonyleptes bicornis
Gervais, 1849: 21, pl. 1, fig 4a–b [desc];
Lycomedes bicornis: Sørensen, 1902: 20, fig 4–4b [rdesc];
Lycomedicus bicornis: Roewer, 1923: 445, fig 561 [cit];
Sadocus bicornis: Kury, 2003: 191 [cat];
Discocyrtus calcitrosus
Loman, 1899: 7, fig 5 [desc];
Lycomedes calcitrosus: Sørensen, 1902: 19 [syst]. Synonymy with L. asperatus established by
Lycomedicus calcitrosus: Moritz, 1971: 193 [cat].
Chile, date and collector unknown, 1 ma (
Sadocus asperatus resembles S. ingens, S. polyacanthus, and S. dilatatus by the bifid prodorsal apical apophysis on coxa IV. S. asperatus can be distinguished from the latter species by the combination of the following characters: lateral margin of dorsal scutum covered by granules; trochanter IV with a blunt retro-dorsal apical apophysis being half of the podomere length, and a rhombus retro-ventral apical tubercle; femur IV curved (in dorsal view), with a retro-lateral row of spiniform apophysis (the middle one longest).
Male (
Coloration. Immersed in ethanol: carapace, trochanter–patella IV brown, tibia light brown, legs I–III, pedipalps, and chelicerae yellowish-brown. Live specimens (Fig.
Variations (n = 56). Scutal area I–IV tubercles (5 minimum, 16 maximum per site), Measurements. Dorsal scutum maximum length 6.0–8.5; dorsal scutum maximum width 6.7–11.6; prosoma maximum length 2.5–3.5; prosoma maximum width 3.2–4.8; leg femora: I 3.2–5.0; II 6.5–10.0; III 5.5–8.5; IV 7.2–11.0.
Female (
Dorsum
(Fig.
Variations (n = 66). Tubercle variation in the scutal areas: I 2–7; II 1–8; III 2–8; IV 3–5. Free tergites I–III each with one pair of blunt or pointed spiniform tubercles. Measurements. Dorsal scutum maximum length 5.9–8.2; dorsal scutum maximum width 8.0–9.6; prosoma maximum length 2.4–3.5; prosoma maximum width 3.7–4.5; leg femora: I 3.5–4.5; II 6.7–8.7; III 5.7–7.0; IV 7.0–8.9.
Of Gonyleptes asperatus and Gonyleptes subsimilis: CHILE. Of Discocyrtus calcitrosus: CHILE, Región de Los Rios, Provincia de Valdivia, Corral.
(Fig.
After examining the original description and the drawing of Gonyleptes bicornis, we concluded that it is of a male of S. asperatus. In the original description, the spines on the free tergite, the two apical apophyses on the trochanter IV and uneven spines in the inner part of the “leg” (referring to the femur IV) are mentioned. Those characters lead us to conclude that it is S. asperatus.
Sadocus dilatatus
Roewer, 1913: 249, fig 102 [desc]; 1923: 493–494, fig 620 [rdesc];
Lycomedicus dilatatus: H. Soares, 1968: 264 [rdesc]; Cekalovic 1985: 18 [cat].
Chile, Región de Biobío, Provincia Concepción, date or, collector unknown, 1 ma (
Sadocus dilatatus resembles S. polyacanthus by the lesser-armed femur IV (compared to other species) and by the posterior large tubercle on the lateral margin of dorsal scutum. Sadocus dilatatus can be distinguished from the other species of the genus by the single retro-ventral central apophysis on femur IV and the very long prodorsal apical apophysis on coxa IV (ca. ⅔ of the scutum width).
Male (
Coloration.
Immersed in ethanol: carapace and leg IV dark brown; legs I–III, pedipalps and chelicerae light brown. Specimen color badly preserved. Live specimens (Fig.
Variations (n = 4). Scutal areas I–IV with 10–13, 14–16, 8–10, 5–6 granules, respectively. In smaller males, the lateral margin of dorsal scutum may bear cluster of tubercles instead of a large tubercle. Measurements. Dorsal scutum maximum length 7.0–8.1; dorsal scutum maximum width 8.5–11.5; prosoma maximum length 3.0–3.4; prosoma maximum width 4.1– 4.4; leg femora: I 5.5–6.5; II 11.4–14.5; III 9.0–11.5; IV 9.7–12.0.
Female (
Chile, Región de Biobío, Provincia Concepción.
(Fig.
The allotype
H.
Gonyleptes funestis
Butler, 1874: 153, figs 5, 5a [desc];
Discocyrtus funestus: Loman, 1899: 6, fig 3 [syst];
Lycomedes funestus: Sørensen, 1902: 20 [syst];
Lycomedicus funestus: Roewer, 1923: 443, fig 557 [rdesc]; 1929: 214 [cat];
Sadocus funestis: Kury, 2003: 191 [cat];
Sadocus funestus:
Chile, Región de Los Ríos, Provincia de Valdivia, Corral, date or collector unknown, 1 ma (
Sadocus funestus can be distinguished from other species of the genus by the following characters: uniramous prodorsal apical apophysis on coxa IV; leg IV straight; and trochanter IV with four apophyses.
Male (
Coloration. Immersed in ethanol: carapace, trochanter, and leg IV brown; legs I–III, pedipalps fading from brown to yellowish. Tubercles and spines of dorsal scutum and free tergites yellowish. Live specimens (Fig.
Variations (n = 21). Granules between lateral margin of carapace and ocularium varying from none to 4–19. Color of granules on carapace range from brown to yellowish. Measurements. Dorsal scutum maximum length 8.3–10.0; dorsal scutum maximum width 8.6–10.5; prosoma maximum length 3.4–4.0; prosoma maximum width 4.7–5.5; leg femora: I 4.5–5.0; II 8.3–9.4; III 7.0–7.9; IV 7.5–8.5.
Female (
Variations (n = 8). Dorsal scutum with 10–24 granules, the armature on the free tergites varying from one paramedian pair of blunt to pointed tubercles. Measurements. Dorsal scutum maximum length 5.5–8.3; dorsal scutum maximum width 8.7–10.0; prosoma maximum length 3.4–3.5; prosoma maximum width 4.5–5.0; leg femora: I 4.8–5.0; II 9.0–9.5; III 7.5–7.8; IV 9.3–9.7.
(Fig.
Carampangue ingens
Mello-Leitão, 1937: 152–153, figs 14–15 [desc]; B.
Sadocus ingens: Kury, 2003; 191 [cat];
Jighas vastus
Roewer, 1943: 28, pl. 3, fig 22 [desc];
Chile, Región de Araucanía, Provincia de Malleco, Contulmo Natural Monument (-38.012778, -73.187500), 12.XII.2010, F. Marques, F. Cadiz & F. Carbayo coll., 1 ma, 1 fe (
Sadocus ingens can be distinguished from the other species of the genus by being the largest among them (and quite large among gonyleptid harvestmen); by the prodorsal apical apophysis on trochanter IV of the same length as the podomere (in other Sadocus species, that apophysis length is up to ½ the podomere length); lateral margin of dorsal scutum smooth posterior to scutal area II.
Male (
Coloration. Immersed in ethanol: carapace, trochanters, femora, patella IV, and tibia IV dark brown. Scutal areas II and III, free tergites, patellae and tibiae I–III orange. Live specimens (Fig.
Variations (n = 6) – Free tergites II and III with one paramedian pair of spines which length varies from similar to slightly longer than the tergite length, its apex varying from blunt to pointed; femur IV with granules in between the retro-lateral spines. Measurements. Dorsal scutum maximum length 12.0–13.8; dorsal scutum maximum width 12.7–16.5; prosoma maximum length 4.5–5.4; prosoma maximum width 7.2–7.8; leg femora: I 8.7–10.0; II 16.8–18.5; III 11.7–15.0; IV 16.0–20.0.
Female (
Variations (n = 4). Tubercle variation in the areas: I–2–8, II–4–6, III–3–6, IV–2–4. Measurements. Dorsal scutum maximum length 11.0–12.0; dorsal scutum maximum width 10.0–11.7; prosoma maximum length 4.5–5.0; prosoma maximum width 6.4–7.2; leg femora: I 7.1–8.4; II 13.8–16.2; III 11.0–12.0; IV 13.0–17.0.
Chile, Región de Araucanía, Provincia de Malleco, Monumento Nacional Contulmo.
(Fig.
Gonyleptes polyacanthus
Gervais, 1847: 576 [desc]; 1849: 24, pl. 1, figs 7–7c [rdesc];
Sadocus polyacanthus: Sørensen, 1902: 14 [syst; rdesc];
Sadocus vitellinosulcatus
Sørensen, 1886: 85, pl. 6, fig 7 [desc]. (Type depository unknown, fe). Synonymy with S. polyacanthus established by
Gonyleptes platei
Loman, 1899: 5, pl. 5, fig 3–3a [desc]. (Holotype
Araucanoleptes exceptionalis
Mello-Leitão, 1946: 5, fig 5 [desc];
Arauconoleptes exceptionalis [spelling mistake]: Cekalovic, 1968: 7 [cat]; 1985: 12 [cat].
Sadocus platei: Moritz, 1971: 207 [cat].
Sadocus conspicillatus
Roewer, 1913: 251 – 253, pl. 1, fig 3 [desc]; 1923: 495 [rdesc];
Sadocus guttatus
Sørensen, 1902: 15 [desc];
Sadocus exceptionalis: Kury, 2003: 191 [cat];
Chile, Región Metropolitana de Santiago, Santiago, date or collector unknown, 4 ma, 2 fe (
Sadocus polyacanthus resembles S. dilatatus by the posterior apophysis on the lateral margin of dorsal scutum, and femur IV almost straight. It is distinguished from the latter by the lack of a long, single retro-central apophysis (present in S. dilatatus) on femur IV, and by the presence of prolateral and retro-lateral rows of similar sized tubercles on femur IV and coxa IV being closer to the body (instead of spread to the sides as S. dilatatus).
Male (
Coloration. Immersed in ethanol: carapace, trochanters I–IV and femur IV dark brown. Legs with a gradient from brown to caramel. Live specimens (Fig.
Variations (n = 58). Even in ethanol there is a great variation of coloration, which ranges from orange to caramel and yellow. The number of granules on the anterior margin of dorsal scutum varying from few sparsely distributed to completely covered. Measurements. Dorsal scutum maximum length 5.6–9.8; dorsal scutum maximum width 6.0–11.9; prosoma maximum length 2.2–4.0; prosoma maximum width 3.0–5.4; leg femora: I 3.5–6.9; II 6.4–13.0; III 5.3–11.4; IV 6.0–11.5.
Female (
Female specimens of Sadocus A, B S. asperatus (
Variations (n = 25). Color in dorsal scutum and femur varying between black and orange. Dorsal scutum with 10–30 granules. Measurements. Dorsal scutum maximum length 7.6–9.3; dorsal scutum maximum width 8.3–10.0; prosoma maximum length 3.0–3.6; prosoma maximum width 4.2–5.0; leg femora: I 4.5–6.0; II 7.0–13.0; III 7.0–11.4; IV 8.4–11.5.
Of G. polyacanthus: Chile. Provincias del Sur. Of S. vitellinosulcatus: either Australasia or South America. Of G. platei: Chile. Región de Los Lagos. Provincia de Valdivia. Corral. Of A. exceptionalis Chile. Región de Los Lagos. Osorno. Barra del Rio Bueno. Of S. conspicillatus: Chile. Región de Biobío. Provincia de Concepción. Concepción. Of Sadocus guttatus: Chile. Región de Biobío. Provincia de Arauco. Lebu.
(Fig.
After examining the holotypes of S. conspicillatus, S. guttatus, and S. exceptionalis, we concluded that they were males within S. polyacanthus size variation. The apophyses size and shape on trochanter IV and the armature of femur IV (especially the retro-dorsal and retro-lateral row of spines size pattern) of all species are the same.
Carampangue allermayeri
Mello-Leitão, 1945: 158 [desc];
Sadocus allermayeri: Kury, 2003: 191 [cat];
Chile. Región de Biobío. Concepción. Concepción.
The type material, belonging to
Carampangue nigronotatum
Mello-Leitão, 1943: 8, fig 7 [desc];
Sadocus nigronotatus: Kury, 2003, 191 [cat];
Chile. Provincia de Llanquihue. Región de Los Lagos. Maullín.
The type material, belonging to the
Sadocus catharinensis
Mello-Leitão, 1923: 152 [desc]; B.
Parasadocus catharinensis: Mello-Leitão, 1927: 8 [syst];
Discocyrtus catharinensis: Soares & Soares, 1987: 458, figs 3–6 [syst, rdesc].
Sadocus aquifugus
Mello-Leitão, 1931a: 136, fig 8 [desc]; 1935: 106 [cit] (Types
Gonyleptes pugilator
Mello-Leitão, 1932: 303, fig 163 [desc]; B.
Gonyleptes acanthopus
Mello-Leitão, 1945: 156 [cit] [nec
Lycomedicus brasiliensis
Soares & Soares, 1949: 52, figs 6–8 [desc]; 1954: 271 [cat] (type material – old collection CGPC, 1 ma, 1 fe –
Sadocus brasiliensis: Kury, 2003: 191 [cat];
Brazil, Paraná, Piraquara, Banhado, IV.1946, Goffergé coll., 1 ma holotype, 1 fe paratype (
S. catharinensis and S. aquifugus: BRAZIL. Santa Catarina. Joinville. Of G. pugilator: BRAZIL. Santa Catarina. Of Lycomedicus brasiliensis: BRAZIL. Paraná. Piraquara: Banhado.
We examined the type material of Sadocus brasiliensis and its external and penial morphology did not match that of other Chilean species of the genus. Based on its type locality, we examined other Brazilian Pachylinae genera and found striking similarities between S. brasiliensis and D. catharinensis. We examined detailed pictures of the type material kindly shared by Rafael N. Carvalho and additional material from the
Gonyleptes horridus
Kirby, 1819: 452, pl. 22, fig 16 [desc];
Lycomedes calcar
Roewer, 1913: 132, fig 59 [desc];
Lycomedicus calcar: Roewer, 1923: 444, fig 558 [rdesc];
Sadocus calcar: Kury, 2003, 191 [cat];
The holotype of Sadocus calcar is in a very bad state of preservation; only part of the carapace, with the ozopores, and leg IV remain. The rest of the prosoma and all of the other legs are absent. Even in this condition, we noted that S. calcar lacks the large tubercles and spines on the free tergites, which are diagnostic of Sadocus. Furthermore, the armature of trochanter IV and the long, bifid, C-shaped dorso-basal apophysis on femur IV are strikingly similar to those of Gonyleptes horridus, a common species in the state of Rio de Janeiro. Therefore, we propose S. calcar as a junior synonym of G. horridus. This synonymy made us conclude that the provenance of S. calcar is mistaken, because G. horridus is endemic to the Brazilian Atlantic rainforest. It is widely known that Roewer, unfortunately, indicated wrong provenance of a few species, and this seems to be the case for this species.
Sadocus planiceps (originally Gonyleptes planiceps Gervais, 1842) has a convoluted taxonomic history, with many previously unknown synonyms detected, which will be published elsewhere (briefly commented in
This study was supported by the project Dimensions US–BIOTA–São Paulo (Fapesp 2013/50297–0, NSF–DOB 1343578 and NASA), and the first author is also grateful for a M.Sc. fellowship (CAPES). MRH thanks the Fundação de Amparo à Pesquisa do Estado de São Paulo (Fapesp 2018/07193-2). We also thank the financial support from CNPq (304933/2014–7). Rafael Carvalho kindly shared pictures of Discocyrtus catharinensis that resulted its synonymy with Sadocus brasiliensis. We are grateful to the editor Gonzalo Giribet and reviewers of an early version, Cristina Rheims, Cibele Bragagnolo, and James Cokendolpher.