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Description of two new species of Progonyleptoidellus (Opiliones: Gonyleptidae), with a cladistic analysis of the genus, an overview of relationships in the K92 group, and taxonomic notes on Deltaspidium

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As part of an ongoing revision and cladistic analysis of the "K92 clade" (Gonyleptidae), the Brazilian genus Progonyleptoidellus Piza, 1940 is revised and two new species from São Paulo State are described: P. bocaina sp. nov. and P. picinguaba sp. nov. A cladistic analysis of the genus was performed using these two new species plus the three previously described species of the genus [P. fuscopictus (B. Soares, 1942); P. orguensis (Soares & Soares, 1954); and P. striatus (Roewer, 1913)], and 25 more additional gonyleptoid outgroup species, most being representatives of the K92 clade. The data matrix is composed of 109 characters: three from the ocularium, 24 from the dorsal scutum, six from the free tergites, nine from the pedipalp, 41 from the legs and 26 from male genitalia. The genus Progonyleptoidellus was recovered as monophyletic only with the exclusion of P. orguensis and was supported on the basis of only one exclusive synapomorphy: presence of dry-marks on sulci of dorsal scutum. Based on the cladistic analysis, P. orguensis was reallocated in Deltaspidium Roewer, 1927, herein considered a senior synonym of Adhynastes Roewer, 1930, and two new combinations are proposed: Deltaspidium orguense (Soares & Soares, 1954) and Deltaspidium tenue (Roewer, 1930). Diagnoses are given for Progonyleptoidellus, the previously described species, (P. fuscopictus and P. striatus) and the two new species.
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ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Accepted by L. E. Acosta: 2 Oct. 2019; published: 4 Nov. 2019 461
Zootaxa 4691 (5): 461–490
https://www.mapress.com/j/zt/
Copyright © 2019 Magnolia Press Article
https://doi.org/10.11646/zootaxa.4691.5.3
http://zoobank.org/urn:lsid:zoobank.org:pub:B2156154-5C5B-4BC8-815D-8C3BC26E5C2A
Description of two new species of Progonyleptoidellus (Opiliones: Gonyleptidae),
with a cladistic analysis of the genus, an overview of relationships in the
K92 group, and taxonomic notes on Deltaspidium
ALÍPIO R. BENEDETTI & RICARDO PINTO-DA-ROCHA*
Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, Caixa Postal: 11.461, 05422-970, São Paulo, SP,
Brazil.
*Corresponding author. E-mail: ricrocha@usp.br
Abstract
As part of an ongoing revision and cladistic analysis of the “K92 clade” (Gonyleptidae), the Brazilian genus
Progonyleptoidellus Piza, 1940 is revised and two new species from São Paulo State are described: P. bocaina sp. nov.
and P. picinguaba sp. nov. A cladistic analysis of the genus was performed using these two new species plus the three
previously described species of the genus [P. fuscopictus (B. Soares, 1942); P. orguensis (Soares & Soares, 1954); and P.
striatus (Roewer, 1913)], and 25 more additional gonyleptoid outgroup species, most being representatives of the K92
clade. The data matrix is composed of 109 characters: three from the ocularium, 24 from the dorsal scutum, six from
the free tergites, nine from the pedipalp, 41 from the legs and 26 from male genitalia. The genus Progonyleptoidellus
was recovered as monophyletic only with the exclusion of P. orguensis and was supported on the basis of only one
exclusive synapomorphy: presence of dry-marks on sulci of dorsal scutum. Based on the cladistic analysis, P. orguensis
was reallocated in Deltaspidium Roewer, 1927, herein considered a senior synonym of Adhynastes Roewer, 1930, and
two new combinations are proposed: Deltaspidium orguense (Soares & Soares, 1954) and Deltaspidium tenue (Roewer,
1930). Diagnoses are given for Progonyleptoidellus, the previously described species, (P. fuscopictus and P. striatus) and
the two new species.
Key words: taxonomy, systematics, Neotropical fauna, Progonyleptoidellinae, Gonyleptinae, K92
Introduction
The Roewerian system of harvestmen classification emphasized differences instead of shared features (Pinto-da-
Rocha 2002; Hara & Pinto-da-Rocha, 2010). For genera in the suborder Laniatores it consisted of a combination
of number of tarsi on legs and type of armature on dorsal scutum and free tergites, what ultimately rendered an
impressive number of unrelated taxa placed together. It also introduced subjectivity into the classification resulting
in some species being placed in different genera. The gonyleptid genus Progonyleptoidellus Piza 1940 is one good
example of the fragility of the Roewerian system (see Hara & Pinto-da-Rocha, 2010; Pinto-da-Rocha et al., 2012).
In fact, we can also say that the subfamily Progonyleptoidellinae suffers the same taxonomic problem as its type
genus. This taxon belongs to a large monophyletic clade within Gonyleptidae, formed of five subfamilies (including
also Caelopyginae, Gonyleptinae, Hernandariinae, and Sodreaninae), mostly endemic to the Atlantic Rain Forest,
called “K92” by Caetano & Machado (2013).
In this paper, we revise Progonyleptoidellus, describe two new species and perform a cladistic analysis of the
genus. To accomplish the latter goal, all treated species are analyzed together with an ample outgroup, including
selected members of the mentioned K92 group. Our objective is to test the monophyly of Progonyleptoidellus, and
to accurately determine the position of P. orguensis, which was suggested to be external to this genus in a previous
work (Pinto-da-Rocha & Bragagnolo 2010).
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462 · Zootaxa 4691 (5) © 2019 Magnolia Press
Material and methods
List and acronyms of repositories: The material examined is deposited in the following repositories (curators
within parentheses): IBSP—Instituto Butantan, São Paulo, São Paulo, Brazil (Antônio D. Brescovit); MNRJ—Mu-
seu Nacional do Rio de Janeiro, Rio de Janeiro, Brazil (Adriano B. Kury); MZLQ—Museu de Zoologia Luiz de
Queiroz, now housed at the IBSP; MZSP—Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil
(Ricardo Pinto-da-Rocha); SMF—Naturmuseum Senckenberg, Frankfurt am Main, Germany (Peter Jäger).
Abbreviations: The following abbreviations are used in the annotated synonymic listing: cat = catalogue; cit =
citation; desc = description; diag = diagnosis; dist = distribution; mat = character matrix; rdesc = redescription; syst
= systematic discussion. The following abbreviations are used in the descriptions: CL = carapace maximum length;
CW = carapace maximum width; DS = dorsal scutum; DSL = dorsal scutum maximum length;; DSW = dorsal scu-
tum maximum width; FeP = femur of the pedipalp length; FI–FIV = femur of the legs I-IV length, FT I–III = free
tergites I–III; MS A–E = penis ventral plate pairs of macrosetae A–E; VP, ventral plate of the penis.
Cladistic analysis.
Outgroup: The selection of outgroups was based on a phylogenetic hypothesis of Gonyleptoidea proposed by
Pinto-da-Rocha et al. (2014), using Hutamaia caramaschii Soares & Soares, 1977 (Gonyleptidae, Ampycinae) to
root the tree. Terminals were chosen to account for the morphological and systematical diversity of the K92 clade.
The outgroup includes the following 25 terminals (listed with voucher specimens): Acanthogonyleptes fulvigra-
nulatus (Mello-Leitão, 1922) (Gonyleptidae: Gonyleptinae) (MZSP 59844, Brazil, São Paulo, Santo Antonio do
Pinhal, Estação Eugenio Lefreve); Acanthogonyleptes singularis (Mello-Leitão, 1935a) (Gonyleptidae: Gonylepti-
nae) (MZSP 73017, Brazil, Rio de Janeiro, Itatiaia, Maromba); Ampheres luteus (Giltay, 1928) (Gonyleptidae: Ca-
elopyginae) (MZSP 59804, Brazil, São Paulo, Santo Antonio do Pinhal); Cadeadoius niger (Mello-Leitão, 1935b)
(Gonyleptidae: Progonyleptoidellinae) (MZSP 73121, Brazil, São Paulo, Cajati, Parque Estadual do Rio Turvo);
Caelopygus elegans (Perty, 1833) (Gonyleptidae: Caelopyginae) (MZSP 73773, Brazil, Terezópolis, Parque Nacio-
nal da Serra dos Órgãos); Cearinus corniger Roewer, 1929 (Gonyleptidae: Gonyleptinae) (MZSP 72883, Brazil,
Pará, Monte Alegre); Geraeocormobius sylvarum Holmberg, 1887 (Gonyleptidae: Gonyleptinae) (MZSP 72953,
Brazil, Paraná, Floresta Nacional Irati); Goniosoma macracanthum (Mello-Leitão, 1922) (Gonyleptidae: Gonio-
somatinae) (MZSP 73133, Brazil, Rio de Janeiro, Itatiaia, Cachoeira Poranga); Gonyleptes fragilis Mello-Leitão,
1923 (Gonyleptidae: Gonyleptinae) (MZSP 72861, Brazil, São Paulo, Cubatão, Copebras); Gonyleptes horridus
Kirby, 1819 (Gonyleptidae: Gonyleptinae) (MZSP 59820, Brazil, Terezópolis, Parque Nacional da Serra dos Ór-
gãos); Heliella singularis B. Soares, 1945b (Gonyleptidae: Progonyleptoidellinae) (MZSP 73113, Brazil, Paraná,
Fazenda Rio Grande, Fazenda Gralha Azul); Hernandaria armatifrons (Roewer, 1917) (Gonyleptidae: Hernandarii-
nae) (MZSP 59801, Brazil, Rio Grande do Sul, São Miguel das Missões); Hutamaia caramaschii Soares & Soares,
1977 (Gonyleptidae: Ampycinae) (MZSP 59968, Brazil, Rondônia, Porto Velho, Caiçara); Iguapeia melanocephala
Mello-Leitão, 1935b (Gonyleptidae: Progonyleptoidellinae) (MZSP 59935, Brazil, São Paulo, Iporanga Parque
Estadual Turístico Alto Ribeira); Iporangaia pustulosa Mello-Leitão, 1935b (Gonyleptidae: Progonyleptoidellinae)
(MZSP 59875, Brazil, São Paulo, Guapiara, Parque Estadual Intervales); Liogonyleptoides inermis (Mello-Leitão,
1922) (Gonyleptidae: Gonyleptinae) (MZSP 59827, Brazil, Minas Gerais, Barroção); Mischonyx cuspidatus (Ro-
ewer, 1913) (Gonyleptidae: Gonyleptinae) (MZSP 59809, Brazil, São Paulo, São Paulo, Parque Estadual da Canta-
reira); Mischonyx processigerus (Soares & Soares, 1970) (Gonyleptidae: Gonyleptinae) (MZSP 72922, Brazil, São
Paulo, Pindamonhangaba, Parque Municipal Trabijú); Mitopernoides variabilis B. Soares, 1945a (Gonyleptidae:
Progonyleptoidellinae) (MZSP 59975, Brazil, São Paulo, Ubatuba, Fazenda Angelim); Multumbo terrenus Roewer,
1927 (Gonyleptidae: Hernandariinae) (MZSP 73139, Brazil, Terezópolis, Parque Nacional da Serra dos Órgãos);
Neosadocus maximus (Giltay, 1928) (Gonyleptidae: Gonyleptinae) (MZSP 59953, Brazil, São Paulo, Santo An-
dré, Estação Biológica de Paranapiacaba); Promitobates ornatus (Mello-Letião, 1922) (Gonyleptidae: Mitobatinae)
(MZSP 59812, Brazil, São Paulo, Cubatão, Copebras); Sodreana barbiellinii (Mello-Leitão, 1927) (Gonyleptidae:
Sodreaninae) (MZSP 73043, Brazil, São Paulo, Ubatuba, Fazenda Angelim); Sodreana sodreana Mello-Leitão,
1922 (Gonyleptidae: Sodreaninae) (MZSP 59813, Brazil, São Paulo, Cubatão, Copebras)
Matrix: We used Mesquite 2.5 computer software (Maddison & Maddison 2017) to edit the character matrix.
The characters were encoded, preferably, in a reductive form (Maddison, 1994) following, for instance, Strong &
Lipscomb (1999). The multistate characters were not polarized a priori. Our matrix consisted of 30 taxa (25 out-
CLADISTICS AND TWO NEW PROGONYLEPTOIDELLUS Zootaxa 4691 (5) © 2019 Magnolia Press · 463
groups and five ingroup taxa) and 109 characters (Tables 1 and 2). The 109 characters were distributed as follows:
three from the ocularium, 24 from the dorsal scutum, six from the free tergites, nine from the pedipalps, 41 from
the legs and 26 from male genitalia.
Analysis and settings: We used TNT (Goloboff et al. 2003, 2008) to perform a heuristic search under parsi-
mony, using 100 rounds of 1,000 replications of Wagner trees built using random sequence addition followed by
TBR branch swapping, keeping 100 trees. The command “collapse branches after search” was used to eliminate
unsupported nodes. Goodman-Bremer support (see Grant & Kluge, 2008) was used on each node (Goodman et al.,
1982; Bremer, 1988), calculated using the Bremer Support Script for TNT 1.0 written by Pablo Goloboff. We have
used equal weights because it provides the most severe test of a phylogenetic hypothesis (Kluge, 1997, 1998) and
maximizes character congruence over all data (Grant and Kluge, 2005).
TABLE 1. List of characters and characters states used in the cladistic analysis.
1 Pedipalp: coxa length (Pinto-da-Rocha & Bragagnolo, 2010): 0. Short; 1. Elongate, reaching the distal margin of trochanter
I.
2 Pedipalp: femur thickness. (DaSilva & Pinto-da-Rocha, 2010): 0. Thick (diameter similar to tibia and tarsus); 1. Thin (di-
ameter much narrower than tibia and tarsus).
3 Pedipalp: femur length (Pinto-da-Rocha & Bragagnolo, 2010): 0. Shorter than prosoma; 1. Longer than prosoma, shorter
than dorsal scutum; 2. Longer than dorsal scutum.
4 Pedipalp: femur, subapical prolateral seta (modified from Pinto-da-Rocha & Bragagnolo, 2010): 0. A small seta (twice
longer than wide); 1. Absent; 2. A large seta (five or more times longer than wide).
5 Pedipalp: patella length (modified from Pinto-da-Rocha & Bragagnolo, 2010): 0. Short (twice as long as wide); 1. Elongate
(five times longer than wide); 2. Very elongate (fifteen times longer than wide).
6 Pedipalp: tibia, socket size (Pinto-da-Rocha & Bragagnolo, 2010): 0. Short; 1. Elongate.
7 Pedipalp: patella–tibia articulation placement (Hara et al. 2012): 0. Posteriorly articulated (distal patella entirely connected
to the base of tibia); 1. Dorsally articulated (distal ventral part of patella connected with dorso-basal part of tibia).
8 Pedipalp: tarsus shape (Pinto-da-Rocha & Bragagnolo, 2010): 0. Straight; 1. Biconvex
9 Pedipalp: tarsus, two ventral rows of setae: (DaSilva & Pinto-da-Rocha, 2010; Pinto-da-Rocha & Bragagnolo, 2010): 0.
Absent; 1. Present.
10 Ocularium: armature type: 0. A pair of small blunt tubercles; 1. A pair of spines.
11 Ocularium: shape, median depression. 0. Absent; 1. Present.
12 Ocularium: width (Pinto-da-Rocha, 2002). 0. Narrow; 1. Wide.
13 Dorsal scutum: number of ozopores (modified from DaSilva & Pinto-da-Rocha, 2010): 0. Two openings; 1. One large open-
ing.
14 Dorsal scutum: carapace, frontal hump height: 0. Low (less than half the height of the ocularium); 1. High (more than half
the height of the ocularium).
15 Dorsal scutum: dry-marks on DS sulci I–IV: 0. Absent; 1. Present.
16 Dorsal scutum: mid-bulge on males, shape in dorsal view: 0. Slightly asymmetrical; 1. Strongly asymmetrical, oblique
posteriorly; 2. Symmetrical, long and round; 3. Subrectangular.
17 Dorsal scutum: coda on males, shape in dorsal view: 0. Long, well separated from mid-bulge; 1. Engulfed by mid-bulge; 2.
Totally absent.
18 Dorsal scutum: mid-bulge on females, shape in dorsal view: 0. Slightly asymmetrical; 1. Strongly asymmetrical, oblique
posteriorly; 2. Symmetrical, long and round.
19 Dorsal scutum: coda on females, shape in dorsal view: 0. Long, well separated from mid-bulge; 1. Long, well separated
from mid-bulge, with flaring sides (alpha-keyhole type); 2. Totally absent; 3. Engulfed by mid-bulge.
20 Dorsal scutum: shape in lateral view: 0. Elevated on areas I–III; 1. Flattened on areas I–III.
21 Dorsal scutum: camouflage with debris (DaSilva & Pinto-da-Rocha, 2010): 0. Absent; 1. Present.
22 Dorsal scutum: pair of tubercles behind ocularium (Pinto-da-Rocha & Bragagnolo, 2010): 0. Absent; 1. Present.
23 Dorsal scutum: sulci II–IV: 0. Conspicuous; 1. Inconspicuous.
24 Dorsal scutum: posterior margin, shape: 0. Straight; 1. Concave; 2. Convex; 3. Sinuous.
25 Dorsal scutum: Area I, state of fusion (modified from Pinto-da-Rocha et al. in prep.): 0. Divided by a longitudinal groove
between scutal grooves I–II; 1. Divided by a scutal groove II; 2. Divided by projections both scutal groove I and II towards
each other.
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464 · Zootaxa 4691 (5) © 2019 Magnolia Press
26 Dorsal scutum: area III, number of armature elements: 0. A pair, separated; 1. A pair, fused; 2. Single one.
27 Dorsal scutum: area III, relative size of armature: 0. similar to those of area II; 1. Larger than those of area II.
28 Dorsal scutum: areas III–IV, state of fusion: 0. Completely fused; 1. As separate scutal areas; 2. Partially fused.
29 Dorsal scutum: area III, sexual dimorphism (Modified from DaSilva & Pinto-da-Rocha, 2010): 0. Absent (unarmed in both
or similarly armed); 1. Present (stronger armature on females); 2. Present (stronger armature on males).
30 Dorsal scutum: areas I–IV, granulation density. 0. Lower than 30 tubercle in total; 1. Higher than 50 tubercles in total.
31 Dorsal scutum: lateral margins, tubercles size (DaSilva & Pinto-da-Rocha, 2010): 0. Tubercles of similar size; 1. Some
tubercles larger than the others.
32 Dorsal scutum: lateral margin on posterior half of opisthosoma, whitish rounded tubercles: 0. Absent; 1. Present.
33 Dorsal scutum: carapace lateral margin, whitish tubercles: 0. Absent; 1. Present.
34 Dorsal scutum: anal operculum, white patches (Pinto-da-Rocha, 2002): 0. Absent; 1. Present.
35 Dorsal scutum: black transversal bands on areas I–III (Pinto-da-Rocha & Bragagnolo, 2010): 0. Absent; 1. Present.
36 Dorsal scutum: maximal width: 0. At area II; 1. Posterior to area II.
37 Free tergites I–III, sexual dimorphism on armature. 0. Absent; 1. Present.
38 Free tergite I: armature in males: 0. Unarmed; 1. Long armature (spine or apophysis); 2. Low armature (tubercle).
39 Free tergite II: armature in males: 0. Unarmed; 1. Long armature (spine or apophysis); 2. Low armature (tubercle).
40 Free tergite III: armature in males: 0. Unarmed; 1. Long armature (spine or apophysis); 2. Low armature (tubercle).
41 Free tergite II: armature in females: 0. Unarmed; 1. Long spine; 2. Median tubercle.
42 Free tergite III: armature in females: 0. Unarmed; 1. Long spine; 2. Median tubercle.
43 Leg I: male basitarsus: 0. Similar to other tarsal articles; 1. Swollen.
44 Leg I: basitarsus I, number of segments (Modified from Pinto-da-Rocha & Bragagnolo, 2010): 0. Three; 1. Four or more;
2. Two.
45 Leg II: distitarsus segmentation: 0. 3-segmented; 1. 4/5-segmented.
46 Leg IV: Coxa hidden by dorsal scutum in males: 0. Absent; 1. Present.
47 Leg IV: Coxa hidden by dorsal scutum in females: 0. Absent; 1. Present.
48 Leg IV: apical width of coxa IV on males in ventral view (Pinto-da-Rocha et al. in prep.): 0. As wide as coxa III apex; 1.
1.5 to twice wider than coxa III apex.
49 Leg IV: coxa, prolateral apical apophysis on males: 0. Present; 1. Absent.
50 Leg IV: coxa, prolateral apical apophysis length: 0. Very short (shorter than trochanter IV); 1. Medium size (similar to the
length of the trochanter IV); 2. Very Large (longer than trochanter IV).
51 Leg IV: coxa, prolateral apical apophysis, secondary lobe subdistal: 0. Absent; 1. Present.
52 Leg IV: coxa, prolateral apical apophysis, direction in dorsal view (in relation to the axis of the base of coxa IV): 0. Slightly
inclined; 1. Transversal; 2. Oblique.
53 Leg IV: coxa, prolateral apical apophysis, apex width: 0. Acuminate, base more than 4 times wider than apex, tapering api-
cally; 1. Acuminate, base twice as wide as apex; 2. Slightly blunt, base as wide as apex.
54 Leg IV: coxa, prolateral apical apophysis, apex sinuous: 0. Absent; 1. Present.
55 Leg IV: coxa, prolateral apical apophysis, thickness: 0. Strong; 1. Thin.
56 Leg IV: coxa, basal process on prolateral apical apophysis: 0. Absent; 1. Present.
57 Leg IV: coxa, prolateral apical apophysis in females: 0. Absent or strongly reduced (a tiny tubercle); 1. Smaller than males;
2. Similar to apophysis on males.
58 Leg IV: coxa, retrolateral apical apophysis in males: 0. Absent; 1. Present.
59 Leg IV: coxa, retrolateral apical apophysis, size: 0. as a small tubercle; 1. As a large apophysis.
60 Leg IV: trochanter, retrolateral apical armature: 0. Absent; 1. Present, a tubercle; 2. Present, strong apophysis.
61 Leg IV: trochanter, basal prolateral tubercle: 0. Absent; 1. Present.
62 Leg IV: femur, curvature inward: 0. Absent; 1. Weakly curved; 2. Strongly curved.
63 Leg IV: femur, curvature outward: 0. Absent; 1. Present.
64 Leg IV: femur, dorso-basal apophysis in males (DBA): 0. Absent; 1. Present.
65 Leg IV: femur, dorso-basal apophysis in males, smaller apophysis, more basal than DBA: 0. Absent; 1. Present.
66 Leg IV: femur, dorso-basal apophysis shape: 0. Bifid, geniculate; 1. Bifid, anvil-shaped; 2. Single, acuminate; 3. Single,
quadrangular apex.
67 Leg IV: femur, dorso-basal apophysis size: 0. Small; 1. Very large (more than twice longer than wide); 2. Large (two times
longer than wide).
CLADISTICS AND TWO NEW PROGONYLEPTOIDELLUS Zootaxa 4691 (5) © 2019 Magnolia Press · 465
68 Leg IV: femur, dorsal armature, row of tubercles: 0. Absent; 1. Present.
69 Leg IV: femur, retrodorsal strong armature in the middle third: 0. Absent; 1. Present.
70 Leg IV: femur, retrodorsal strong armature in the distal third: 0. Absent; 1. Present.
71 Leg IV: femur, retroventral strong armature in the basal third: 0. Absent; 1. Present.
72 Leg IV: femur, retroventral armature, acessory small spines in the basal third: 0. Absent; 1. Present.
73 Leg IV: femur, retroventral strong armature in the distal third: 0. Absent; 1. Present.
74 Leg IV: femur, retroventral acessory small spines in the distal third: 0. Absent; 1. Present.
75 Leg IV: femur, prodorsal spines on basal half: 0. Absent; 1. Present.
76 Leg IV: femur, prodorsal spines on distal half: 0. Absent; 1. Present.
77 Leg IV: femur, proventral spines on basal half: 0. Absent; 1. Present.
78 Leg IV: femur, proventral spines on distal half: 0. Absent; 1. Present.
79 Leg IV: femur, sexual dimorphism of armature: 0. Absent; 1. Present, females unarmed; 2. Present, females with different
armature than males.
80 Leg IV: number of tarsomeres: 0. Less than 11; 1. More than 16;
81 Leg IV: metatarsus, astragalus on male: 0. As wide as calcaneus; 1. Swollen.
82 Legs III–IV: shape of tarsomeres: 0. Cylindrical; 1. Globular.
83 Legs III–IV: tarsus, claw: 0. Smooth; 1. Pectinate.
84 Penis: glans frontal base heel-shape: 0. Absent; 1. Present.
85 Penis:. glans, insertion of the pedestal in the glans (in lateral view): 0. Medial; 1. Ventral.
86 Penis VP: shape in dorsal view: 0. Trapezoid; 1. Rectangular; 2. Quadrate.
87 Penis VP: shape in lateral view: 0. Globous at base, thin at apex; 1. Without globous base, straight; 2. Without globous base,
rounded.
88 Penis VP: cleft on distal margin. (Kury, 1992): 0. Absent; 1. Present.
89 Penis VP: cleft shape: 0. U-shaped; 1. Parabolic.
90 Penis VP: cleft depth: 0. Deep; 1. Shallow; 2. Very deep (approximately half of VP).
91 Penis: VP: base of cleft: 0. Curved; 1. Straight.
92 Penis: VP: lateral lobes (Kury, 1992): 0. Absent; 1. Present.
93 Penis: VP: lateral lobes, shape in lateral view: 0. Rounded; 1. Rectangular.
94 Penis VP: lateral lobes, placement: 0. Basal; 1. Sub basal.
95 Penis: apical part of divided VP: 0. Unfolded; 1. Folded.
96 Penis VP: macrosetae A + B, shape: 0. Conical with apex slightly rounded; 1. Lanceolate; 2. Spatulate.
97 Penis VP: macrosetae A + B, arrangement in lateral view: 0. Arranged forming an arch; 1. Arranged forming a row; 2. Ar-
ranged forming a triangle; 3. Arranged forming a U.
98 Penis VP: macroseta B, size: 0. Extremely reduced or absent; 1. As same size than macrosetae A 0; 2. Smaller than macro-
setae A.
99 Penis VP: macrosetae C, shape. 0. Straight. 1. Helicoidal; 2. Curved.
100 Penis VP: macrosetae C, placement: 0. Distal; 1. Sub distal.
101 Penis VP: microsetae (ventral view): 0. Absent; 1. Present.
102 Penis: microsetae on lateral lobes: 0. Absent; 1. Present.
103 Penis: microsetae on truncus (lateral view): 0. Absent; 1. Present.
104 Penis: ventral process, length: 0. Short; 1. Long.
105 Penis ventral process: flabellum apex: 0. Serrated; 1. With one terminal longer than others; 2. Without projections.
106 Penis: stylus, ventral microspines oriented in a longitudinal row: 0. Absent; 1. Present.
107 Penis: stylus, microspines at subapical region: 0. Absent; 1. Present.
108 Penis: stylus, apex shape: 0. Inclined to the axis of the stylus, without ventral projection; 1. Inclined to the axis of the stylus,
with ventral projection; 2. Not inclined, short.
109 Penis: glans, stylus insertion: 0. Stylus as a continuation of glans; 1.Candelabrum. 2. V-Shaped; 3. Close together with
ventral process.
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TABLE 2. Data matrix used in the cladistic analysis of Progonyleptoidellus spp. (continued on next pages).
Taxa 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
Hutamaia caramaschii 0 0 0 ? 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0
Promitobates ornatus 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 3 0 0 0
Goniosoma macracanthum 0 0 0 2 0 1 0 0 0 1 0 1 0 0 0 1 1 1 3 1 0 0
Ampheres luteus 0 1 1 1 1 1 1 1 1 0 1 1 0 0 0 1 2 1 2 0 0 1
Caelopygus elegans 0 1 1 0 0 1 1 1 1 0 1 1 0 0 0 1 2 1 2 1 0 0
Hernandaria armatifrons 0 0 0 1 0 0 0 0 1 1 0 0 1 0 0 0 1 0 3 1 1 0
Multumbo terrenus 0 0 0 1 0 0 0 1 1 1 0 0 1 0 0 0 1 0 3 1 1 1
Sodreana barbiellinii 1 1 2 1 2 1 1 1 1 1 0 0 0 1 0 0 1 0 3 1 0 1
Sodreana sodreana 1 1 2 0 2 1 1 1 1 1 0 0 0 0 0 0 2 0 2 1 0 0
Cadeadoius niger 0 1 0 1 0 0 0 1 1 1 0 0 0 0 0 1 2 1 3 1 0 1
Heliella singularis 0 0 0 1 0 0 0 1 1 1 0 0 0 0 0 1 2 1 2 1 0 0
Mitopernoides variabilis 0 1 2 1 1 1 1 1 1 1 0 0 0 0 0 0 1 0 3 1 0 1
Iporangaia pustulosa 0 1 1 1 0 0 1 1 1 1 0 0 0 0 0 0 1 0 3 0 0 1
Deltaspidium asperum 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 1 2 1 2 1 0 1
Deltaspidium tenue comb.nov. 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 1 2 1 2 1 0 1
Deltaspidium orguense comb. nov. 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 1 2 1 2 1 0 1
Neosadocus maximus 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 1 1 3 0 0 1
Liogonyleptoides inermis 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 2 0 2 1 0 0 0
Cearinus corniger 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 2 0 2 1 0 0 0
Geraeocormobius sylvarum 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 2 0 2 0 1 0 0
Acanthogonyleptes singularis 0 1 1 0 0 0 1 1 1 0 0 0 0 0 0 0 2 0 3 1 0 1
Acanthogonyleptes fulvigranulatus 0 1 1 0 0 0 1 1 1 0 0 0 0 1 0 0 2 0 3 1 0 0
Gonyleptes horridus 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 3 1 0 0
Gonyleptes fragilis 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 1 0 3 1 0 0
Mischonyx cuspidatus 0 0 0 1 0 0 0 0 1 1 0 0 0 0 0 0 1 0 0 0 1 1
Mischonyx processigerus 0 0 0 1 0 0 0 0 1 1 0 0 0 0 0 0 1 0 0 0 1 1
Progonyleptoidellus bocaina sp. nov. 0 1 1 1 1 1 1 1 1 1 0 0 0 1 1 1 2 0 3 1 0 1
Progonyleptoidellus fuscopictus 0 1 1 1 1 1 1 1 1 1 0 0 0 1 1 1 2 1 2 1 0 1
Progonyleptoidellus picinguaba sp.
nov.
1 1 1 0 1 1 1 1 1 1 0 0 0 1 1 1 2 1 2 1 0 1
Progonyleptoidellus striatus 0 1 1 1 1 1 1 1 1 1 0 0 0 1 1 1 2 1 2 1 0 1
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CLADISTICS AND TWO NEW PROGONYLEPTOIDELLUS Zootaxa 4691 (5) © 2019 Magnolia Press · 467
TABLE 2. (Continued)
Taxa 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44
Hutamaia caramaschii 0 0 0 0 - 1 0 1 1 0 0 0 0 0 0 0 0 1 0 1 0 0
Promitobates ornatus 0 0 2 0 - 1 0 1 1 0 0 0 0 0 0 0 0 1 2 2 0 1
Goniosoma macracanthum 1 2 1 0 - 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1
Ampheres luteus 1 2 0 0 1 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 1 1
Caelopygus elegans 0 2 0 0 1 0 0 1 0 0 0 1 0 1 0 0 0 0 0 0 0 1
Hernandaria armatifrons 1 0 0 0 0 2 0 1 1 0 0 0 0 0 1 2 2 2 1 1 0 2
Multumbo terrenus 0 0 0 0 1 0 1 1 1 0 0 0 0 0 1 2 2 2 2 2 1 0
Sodreana barbiellinii 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1
Sodreana sodreana 0 3 0 2 1 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Cadeadoius niger 0 2 0 0 1 0 1 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0
Heliella singularis 1 2 0 0 1 0 1 0 1 0 1 0 0 1 1 0 1 0 1 0 1 0
Mitopernoides variabilis 0 0 2 0 1 0 0 0 1 0 0 0 0 0 0 2 1 2 2 2 1 0
Iporangaia pustulosa 0 0 0 0 1 0 0 0 1 1 1 0 0 0 0 0 2 1 0 1 0 0
Deltaspidium asperum 0 2 0 1 1 0 1 1 1 0 0 0 0 1 0 0 0 0 0 0 0 1
Deltaspidium tenue comb.nov. 0 2 0 1 1 0 1 1 1 0 0 0 0 1 0 0 0 0 0 0 0 1
Deltaspidium orguense comb. nov. 0 2 0 0 1 0 1 1 1 0 0 0 0 1 0 0 0 0 0 0 0 1
Neosadocus maximus 0 0 0 0 1 0 1 1 0 0 1 0 0 1 0 0 0 0 0 0 0 0
Liogonyleptoides inermis 0 0 0 - - 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Cearinus corniger 0 0 0 0 1 1 2 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0
Geraeocormobius sylvarum 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0
Acanthogonyleptes singularis 0 3 0 2 1 0 0 1 1 0 0 0 0 1 1 0 0 0 1 1 0 1
Acanthogonyleptes fulvigranulatus 0 2 0 0 1 0 0 1 1 0 0 0 0 1 1 0 2 2 1 1 1 0
Gonyleptes horridus 0 0 0 0 - 0 0 0 1 1 0 0 0 0 1 0 0 0 2 2 0 0
Gonyleptes fragilis 0 0 0 0 - 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0
Mischonyx cuspidatus 0 0 0 0 1 0 2 0 1 1 0 0 0 0 0 1 1 1 1 1 1 0
Mischonyx processigerus 0 0 0 0 1 0 2 0 1 1 0 0 0 0 0 2 1 1 1 1 0 0
Progonyleptoidellus bocaina sp.
nov.
0 3 0 0 - 0 0 0 1 0 0 0 0 1 0 2 2 1 0 1 0 1
Progonyleptoidellus fuscopictus 0 0 0 0 1 0 0 0 1 0 0 0 0 1 0 0 0 2 1 1 0 1
Progonyleptoidellus picinguaba sp.
nov.
1 3 0 0 1 0 0 0 1 0 0 0 0 1 0 2 1 1 1 1 0 1
Progonyleptoidellus striatus 0 2 0 - - 0 1 0 0 0 0 0 1 1 0 0 0 0 0 0 0 1
......continued on the next page
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TABLE 2. (Continued)
Taxa 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66
Hutamaia caramaschii 0 1 1 0 0 1 0 2 1 0 1 0 0 0 - 2 0 0 1 0 0 -
Promitobates ornatus 0 0 1 0 0 1 1 2 1 1 0 1 0 0 0 1 1 0 0 0 0 -
Goniosoma macracanthum 0 0 0 1 0 0 0 2 1 1 0 0 1 1 1 1 1 2 0 0 0 -
Ampheres luteus 1 1 1 0 0 1 0 0 1 0 0 0 1 0 - 1 0 0 0 0 0 -
Caelopygus elegans 1 0 1 1 0 2 0 0 0 1 0 0 0 1 0 0 1 0 0 0 0 -
Hernandaria armatifrons 0 0 1 1 0 2 0 0 0 0 0 0 0 0 - 0 1 0 0 1 0 2
Multumbo terrenus 0 1 1 0 1 - - - - - - - - 0 - 0 0 0 0 0 0 -
Sodreana barbiellinii 1 1 1 0 0 1 0 2 1 0 1 0 0 0 - 1 1 0 0 1 0 2
Sodreana sodreana 0 0 1 0 0 2 1 0 0 1 0 1 0 1 0 1 1 0 0 1 1 2
Cadeadoius niger 1 1 0 0 0 1 0 2 1 0 0 1 0 0 - 2 1 2 0 0 0 -
Heliella singularis 1 1 1 0 0 1 0 2 1 0 0 1 0 1 1 2 1 1 0 0 0 -
Mitopernoides variabilis 1 1 1 0 0 0 0 2 1 0 1 0 0 1 0 1 0 0 0 0 0 -
Iporangaia pustulosa 0 1 1 0 1 - - - - - - - 0 1 0 0 0 0 0 0 0 -
Deltaspidium asperum 1 0 1 1 0 2 0 2 1 1 0 1 0 1 0 1 1 0 0 1 1 2
Deltaspidium tenue comb.nov. 1 0 1 1 0 2 0 2 1 1 0 1 0 1 0 1 1 0 0 1 1 2
Deltaspidium orguense comb. nov. 1 0 1 1 0 2 0 2 1 1 0 1 0 1 0 1 1 0 0 1 1 2
Neosadocus maximus 1 0 1 1 0 2 1 0 1 1 0 1 0 1 0 1 1 1 1 1 0 2
Liogonyleptoides inermis 0 0 1 1 0 1 1 0 2 0 0 1 0 0 - 0 1 1 0 1 1 3
Cearinus corniger 0 0 1 1 0 2 1 0 0 1 0 1 0 0 - 1 1 1 0 1 1 2
Geraeocormobius sylvarum 0 0 1 1 0 2 1 0 0 1 0 1 0 0 - 0 1 0 0 1 1 2
Acanthogonyleptes singularis 0 0 1 1 0 2 1 0 0 1 0 1 0 1 0 1 1 0 0 1 1 1
Acanthogonyleptes fulvigranulatus 0 0 1 1 0 2 1 0 0 1 0 1 0 1 0 1 1 0 0 1 1 1
Gonyleptes horridus 0 0 1 1 0 2 1 0 0 1 0 1 0 1 1 1 1 0 0 1 1 0
Gonyleptes fragilis 0 0 1 1 0 2 1 0 0 1 0 1 0 1 0 1 0 0 0 1 1 0
Mischonyx cuspidatus 0 0 1 1 0 1 1 0 0 1 0 1 0 1 1 1 1 1 0 1 1 0
Mischonyx processigerus 0 0 1 1 0 1 1 2 0 1 0 1 0 1 0 1 1 0 1 1 1 0
Progonyleptoidellus bocaina sp.
nov. 1 1 1 1 0 2 0 0 1 1 0 0 1 1 0 1 1 0 0 0 0 -
Progonyleptoidellus fuscopictus 1 0 1 1 0 2 0 0 1 1 0 0 1 1 0 1 1 0 0 1 0 2
Progonyleptoidellus picinguaba sp.
nov. 0 0 1 0 0 1 0 0 1 0 0 0 2 1 0 1 1 0 0 0 0 -
Progonyleptoidellus striatus 1 0 1 1 0 2 1 0 1 1 0 1 1 1 0 1 1 0 0 1 1 3
......continued on the next page
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TABLE 2. (Continued)
Taxa 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88
Hutamaia caramaschii - 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 - 0 2 1
Promitobates ornatus - 0 0 0 0 0 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 2 0
Goniosoma macracanthum - 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0
Ampheres luteus - 1 0 0 0 1 0 0 0 0 0 0 ? 1 0 1 1 1 1 2 0 1
Caelopygus elegans - 1 0 0 0 0 0 1 0 0 0 0 2 1 0 1 1 1 1 2 1 1
Hernandaria armatifrons 0 1 0 1 0 0 0 0 0 0 1 1 2 0 0 0 0 1 1 1 2 1
Multumbo terrenus - 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 2 1
Sodreana barbiellinii 0 1 0 0 0 0 0 0 0 0 0 0 2 0 0 0 0 1 1 1 1 1
Sodreana sodreana 2 1 1 0 1 1 0 1 1 1 0 0 1 0 1 0 0 1 1 1 0 1
Cadeadoius niger - 1 1 1 0 0 0 0 0 0 0 0 1 0 0 0 0 1 1 2 0 1
Heliella singularis - 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 1 1 1 0 1
Mitopernoides variabilis - 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 1 1 1 2 1
Iporangaia pustulosa - 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 1 1 2 ? 1
Deltaspidium asperum 2 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 2 1
Deltaspidium tenue comb.nov. 2 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 2 1
Deltaspidium orguense comb. nov. 2 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 2 1
Neosadocus maximus 2 1 1 0 0 0 0 0 1 0 0 0 1 0 1 0 0 1 1 1 0 1
Liogonyleptoides inermis 0 0 1 0 0 0 1 0 0 0 0 0 1 0 0 0 0 1 1 2 0 1
Cearinus corniger 2 1 0 1 0 0 0 1 0 1 0 0 1 0 0 0 0 1 1 2 0 1
Geraeocormobius sylvarum 2 0 0 0 1 0 1 0 0 1 0 1 1 0 0 0 0 1 1 1 2 1
Acanthogonyleptes singularis 2 1 1 0 0 0 0 1 0 1 0 1 2 0 0 0 0 ? ? ? ? ?
Acanthogonyleptes fulvigranulatus 2 1 1 0 0 0 0 1 0 0 0 1 2 0 0 0 0 1 1 0 1 1
Gonyleptes horridus 1 1 1 1 0 0 0 1 1 1 0 1 1 0 0 0 0 1 1 1 0 1
Gonyleptes fragilis 2 1 1 0 1 1 0 1 1 1 1 1 1 0 0 0 0 1 1 2 0 1
Mischonyx cuspidatus 2 1 1 0 0 0 0 0 0 1 0 1 1 0 0 0 0 1 1 1 2 1
Mischonyx processigerus 1 1 1 0 0 0 0 0 0 0 0 1 1 0 0 0 0 1 1 1 2 1
Progonyleptoidellus bocaina sp.
nov.
- 1 0 0 0 0 0 1 0 0 0 1 2 1 0 1 0 1 1 1 2 1
Progonyleptoidellus fuscopictus 2 1 1 0 0 1 0 1 1 1 1 1 2 1 0 1 0 1 1 1 2 1
Progonyleptoidellus picinguaba
sp. nov.
- 1 0 0 0 1 0 1 0 0 1 1 0 1 0 1 0 1 1 1 2 1
Progonyleptoidellus striatus 2 1 1 0 0 1 0 1 0 0 1 1 2 1 0 1 0 1 1 1 2 1
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TABLE 2. (Continued)
Taxa 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 105 106 107 108 109
Hutamaia caramaschii 1 0 0 0 - - 0 0 2 1 0 0 0 - 0 - - 0 0 0 0
Promitobates ornatus - - - 0 - - 0 1 2 0 0 0 1 - 0 0 0 0 0 2 2
Goniosoma macracanthum - - - 0 - - 0 0 1 0 0 1 0 - 0 0 - 0 0 0 2
Ampheres luteus 0 1 0 1 1 0 0 1 1 0 0 0 1 ? ? 1 1 0 1 0 1
Caelopygus elegans 0 1 1 1 0 0 1 1 0 1 1 0 1 1 ? 1 1 0 1 0 1
Hernandaria armatifrons 0 0 0 1 1 0 0 1 1 0 1 1 1 0 0 1 1 1 1 0 1
Multumbo terrenus 1 0 0 1 1 0 0 1 1 0 0 1 1 1 1 1 1 0 0 0 1
Sodreana barbiellinii 1 0 0 1 0 1 0 2 3 1 0 1 0 0 0 1 1 0 0 0 1
Sodreana sodreana 1 2 0 1 0 1 0 0 1 0 0 0 1 0 0 1 1 0 1 0 1
Cadeadoius niger 1 0 0 1 0 0 0 1 0 0 1 0 1 1 0 1 0 0 1 0 1
Heliella singularis 1 0 0 1 0 0 0 1 0 0 1 0 ? ? ? 1 0 0 1 0 1
Mitopernoides variabilis 1 0 0 1 0 0 1 1 1 0 1 0 1 0 0 1 0 0 1 0 1
Iporangaia pustulosa 1 0 0 1 1 0 0 1 0 0 1 0 1 1 ? 1 0 0 0 0 1
Deltaspidium asperum 1 0 0 1 0 0 0 1 0 2 0 1 1 0 0 1 1 0 1 0 3
Deltaspidium tenue comb.nov. 1 0 0 1 0 0 0 1 0 2 0 1 1 0 0 1 1 0 1 0 3
Deltaspidium orguense comb. nov. 1 2 0 1 0 0 1 1 0 1 0 1 1 1 0 1 1 0 1 0 3
Neosadocus maximus 1 0 0 1 1 0 1 1 3 1 1 0 1 1 0 0 0 0 1 0 1
Liogonyleptoides inermis 0 0 0 1 0 0 1 1 0 1 1 1 1 1 0 1 0 0 0 0 1
Cearinus corniger 0 0 0 1 0 0 1 1 1 1 0 1 1 1 1 1 1 0 0 0 1
Geraeocormobius sylvarum 1 0 0 1 0 0 1 1 2 2 0 1 1 0 0 1 0 0 1 1 1
Acanthogonyleptes singularis ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?
Acanthogonyleptes fulvigranulatus 0 0 1 1 1 1 1 1 1 1 0 1 1 0 0 1 0 0 0 0 3
Gonyleptes horridus 0 0 0 1 0 0 1 1 0 1 2 1 1 1 1 1 0 1 0 0 1
Gonyleptes fragilis 0 0 0 1 0 0 0 1 0 1 2 1 1 1 1 1 0 0 0 2 1
Mischonyx cuspidatus 1 0 0 1 0 1 0 1 0 2 1 1 1 0 0 1 1 1 0 0 1
Mischonyx processigerus 1 0 0 1 0 1 0 1 0 1 2 0 1 ? ? 1 0 1 0 0 1
Progonyleptoidellus bocaina sp.
nov.
1 0 0 1 0 0 0 1 0 0 1 0 1 0 0 1 0 0 1 0 1
Progonyleptoidellus fuscopictus 1 2 0 1 0 0 1 1 0 0 1 0 1 1 0 1 0 0 1 0 1
Progonyleptoidellus picinguaba
sp. nov.
1 0 0 1 1 1 1 1 0 0 1 0 1 0 0 1 0 0 1 1 1
Progonyleptoidellus striatus 1 2 0 1 0 0 0 1 0 1 1 1 1 0 0 1 0 0 1 0 1
CLADISTICS AND TWO NEW PROGONYLEPTOIDELLUS Zootaxa 4691 (5) © 2019 Magnolia Press · 471
Descriptive section.
Records information: The localities of the examined material are organized as follows “COUNTRY. State. Lo-
cality”. On the map, we depicted only the localities from the samples that we analyzed. Localities from the literature
are only mentioned in the text.
Description and figures: Terminology for the armature of the dorsal scutum and legs follows DaSilva & Gnaspi-
ni (2010) with some modifications to best fit the characteristics of the group. Granules (always in plural) are tiny
elevations (height = width) concentrated at an area or article. Tubercles are small elevations (height at most twice
the width) that stand out from the granules and may be blunt or acuminate, the latter (acuminate tubercles) gener-
ally taller than the former. Spines are medium-sized to high elevations (higher than tubercles; height always more
than twice the width), approximately conical, always acuminate, and that can be found on the anterior margin and
scutal areas, the ocularium, pedipalps, free tergites I–III and legs. Apophyses are projections of the integument, ir-
regularly shaped, found on coxae IV, free tergites I–III, anterior margin of dorsal scutum and the basal portion of
femur IV. The area sometimes referred to as scutal “area V” is here referred to as the posterior margin of the dorsal
scutum. An area or article are considered unarmed when there is no large structure that stands out (i.e., larger than
mere granules), and smooth when there is no structure, not even granules. The topological terms for the append-
ages follow Acosta et al. (2007b). The code for the relative length of setae of pedipalp tibia and tarsus follows the
definition of Pinto-da-Rocha (2002). Tarsal counts display the numbers of tarsomeres in tarsus I to IV, with numbers
in parentheses indicating the number of tarsomeres in distitarsi I–II. The nomenclature for the shape of the dorsal
scutum in dorsal view follows Kury & Medrano (2016). A ”dry-mark” is a patch in the more external serose layer of
the cuticle that forms white drawings in the animal when it dries up when removed from liquid (DaSilva & Gnaspini
2010). The penial macrosetae and microsetae nomenclature follows Kury & Villarreal M. (2015) and Kury (2016),
respectively. Coloration is described on specimens preserved in 70% ethanol. All measurements are in millimeters
unless otherwise stated. The descriptions were based on all the examined material. The description of the male holo-
type is presented following the variation in other males. Female characteristics are described only if they differ from
the male. Drawings were made using a Leica stereomicroscope (model MZ APO, Heerbrugg, Switzerland) with a
camera lucida. The penes were prepared for electron microscopy following Pinto-da-Rocha (2002); structures were
observed and photographed with a Leica LEO 440 scanning electron microscope, at the Museu de Zoologia, Uni-
versidade de São Paulo.
Results
Cladistic analysis. The cladistic analysis resulted in one most parsimonious tree (L = 500; C.I. = 0.26; R.I. = 0.52;
Fig. 1) and Progonyleptoidellus was recovered as monophyletic except for the exclusion of P. orguensis, which
should therefore be removed from this genus. In our hypothesis (Fig. 1), based on more terminals (and more specifi-
cally selected) than Pinto-da-Rocha & Bragagnolo (2010), Progonyleptoidellus orguensis confirms its position as
not related to Progonyleptoidellus and forms a clade with Deltaspidium asperum and Adhynastes tenuis; we then
recognize this clade (Goodman-Bremer = 4) to represent the genus Deltaspidium (subfamily Gonyleptinae). The
mentioned clade is supported by six unambiguous synapomorphies [79(0); 86(0); 87(2); 99(0); 105(1); 109(3)],
all of which are homoplastic. The remaining species of the genus (P. fuscopictus and P. striatus) plus the two new
species (P. bocaina sp. nov. and P. picinguaba sp. nov.) form a clade, here recognized as the actual extent of Progo-
nyleptoidellus (Fig. 1), supported by four synapomorphies, three of which are unambiguous [14(1); 78(1); 87(2)]
and one is exclusive: presence of dry-marks on the sulci of the dorsal scutum (Goodman-Bremer support = 4).
Progonyleptoidellus was recovered as the sister group of Caelopyginae, supported by seven synapomorphies [3(1);
6(1); 7(1); 29(0); 56(0); 79(2); 80(1)], all homoplastic. Progonyleptoidellus was divided into two clades: one with
Progonyleptoidellus bocaina sp. nov. as the sister group of Progonyleptoidellus picinguaba sp. nov. and another
containing P. striatus and P. fuscopictus.
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FIGURE 1. Most parsimonious tree representing the cladistic relationships of Progonyleptoidellus and Deltaspidium with other
members of the K92 clade (L=500; C.I.=0.26; R.I.=0.52). Black circles indicate unique transformations, while white circles
mean homoplastic transformations. The character number is above each circle, the character state is below. Only unambigu-
ous characters were represented. Characters and character states are detailed in Table 1, the data matrix is in Table 2. Absolute
Bremer support is given near each node.
Taxonomic account
Gonyleptidae Sundevall, 1833
Progonyleptoidellinae Soares & Soares, 1985
Progonyleptoidellus Piza, 1940
Ampheres (part): Roewer, 1913: 334 (desc)
Progonyleptoidellus Piza, 1940:63 (desc); Soares & Bauab-Vianna, 1972: 205 (syst) [revalidated from the synonymy of Ger-
aeocormobius]; Soares & Soares, 1985: 181 (rdesc, syst); Kury & Pinto-da-Rocha, 1997: 109 (diag, syst); Kury, 2003 (cat);
Kury & Alonso-Zarazaga, 2011: 58 (cat).
Laneius B. Soares, 1942: 6 (desc); Soares & Soares, 1985: 184 (rdesc, syst) [revalidated from the synonymy of Gonazula
Roewer, 1930] (type species Laneius fuscopictus B. Soares, 1942, by original designation). Synonymy established by Kury
& Pinto-da-Rocha, 1997.
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Gonazula (part): Soares & Soares, 1949: 173 (diag, syst).
Piraquara Piza, 1943: 258 (desc); Soares & Soares, 1949: 167 (cit) (type species Piraquara schubarti Piza, 1943, by original
designation). Synonymy established by Soares & Bauab-Vianna, 1972.
Cadeadoius (part): Roewer, 1943: 45 (desc).
Gonyleptoides (part): B. Soares, 1944a: 257 (syst).
Geraecormobius (part): Soares & Soares, 1949: 167 (syst).
Type species. Progonyleptoidellus androgynus Piza, 1940, by original designation.
Etymology. The genus name derives from the pre-existing genus Progonyleptoides + masculine form of Latin suffix
ellus. Gender masculine.
Included species. Progonyleptoidellus bocaina sp. nov., Progonyleptoidellus fuscopictus (Soares, 1942), Prog-
onyleptoidellus picinguaba sp. nov. and Progonyleptoidellus striatus (Roewer, 1913).
Diagnosis. Progonyleptoidellus can be distinguished from the other genera of Progonyleptoidellinae by the
following combination of characters: Dorsal scutum outline (Figs. 2–7) gamma pyriform (γP); mesotergum divided
into 3 scutal areas; posterior margin of DS convex or sub straight; scutal areas I–II unarmed (Figs. 2–7); area III
armed with two paramedian large backwards-pointing spines (larger in some females), except in P. striatus; coxa
IV with large proapical apophysis in males (Figs. 2–7) and females, generally larger and with curved apex in males
(except P. picinguaba sp. nov., straight and equal-sized in both sexes); femur IV with (P. fuscopictus and P. striatus)
or without DBA (P. bocaina sp. nov. and P. picinguaba sp. nov.); femur IV of females armed (usually with an ar-
mature different than males except P. picinguaba sp. nov.); tarsus IV with more than 16 tarsomeres; carapace dark
colored and DS yellowish, with black spots or bands on scutal areas. Penis (Fig. 6) with deep cleft on VP, lateral
lobes placed sub-basally; VP rounded, without globular base in lateral view; MS A + B placed forming an arch; MS
B smaller than MS A (except in P. striatus); MS C with helicoidal apex; flabellum fan-like; stylus sub sigmoid, with
apex inclined to the axis of stylus.
Nomenclatural history. Roewer (1913) described Ampheres striatus in Caelopyginae. Later, Piza (1940) created
the new genus Progonyleptoidellus, which was placed in Gonyleptinae, with a single species Progonyleptoidellus
androgynus, based on a female from Alto da Serra, São Paulo. Two years later, B. Soares (1942) described the
monotypic genus Laneius, for Laneius fuscopictus B. Soares, 1942, based on a female holotype and a male allotype
from Boracéia, Salesópolis, São Paulo state, placing it in Gonyleptinae too. In the following year, Piza (1943)
created in Gonyleptinae the new monotypic genus Piraquara, with the single species Piraquara schubarti Piza,
1943 from Serra da Piraquara, Itanhaém, São Paulo. In the same year, Roewer (1943) described Cadeadoius
atroluteus Roewer, 1943 in Goniosomatinae. One year later, B. Soares (1944a) synonymized Progonyleptoidellus
and Piraquara under Gonyleptoides Roewer, 1913, a formerly monotypic genus from Rio de Janeiro. Soares &
Soares (1949) synonymized Laneius with Gonazula Roewer, 1930, a formerly monotypic genus from Santa Catarina,
producing the new combination Gonazula fuscopicta. In the same paper, Soares & Soares (1949: 167) synonymized
Gonyleptoides with Geraeocormobius Holmberg, 1888, making the new combinations Geraeocormobius androgynus
and G. schubarti; as pointed out by Acosta et al. (2007a), the correct (original) spelling of the generic name is
Geraeocormobius, while the majority of subsequent authors employed the incorrect spelling “Geraecormobius”.
Still in the same contribution, Soares & Soares (1949) transferred Cadeadoius atroluteus to Geraeocormobius.
Five years later, Soares & Soares (1954) described Geraeocormobius orguensis based on a female from Serra dos
Órgãos, Teresópolis, Rio de Janeiro state. Later, Soares & Bauab-Vianna (1972) revalidated Progonyleptoidellus and
considered Piraquara as its junior subjective synonym. Thirteen years later, Soares & Soares (1985) synonymized
Piraquara schubarti and Cadeadoius atroluteus with Progonyleptoidellus androgynus, and revalidated Laneius.
Furthermore, Soares & Soares (1985) also erected the new subfamily Progonyleptoidellinae, with Progonyleptoidellus
as the type genus, and included the genera Iguapeia Mello-Leitão, 1935b, Laneius, Heliella B. Soares, 1945b,
Cadeadoius Mello-Leitão, 1936, Moreiranula Roewer, 1930, Gonyleptoides, Opisthoplites Sørensen, 1884 and
Stylopisthos Roewer, 1930.
Several years later, Kury & Pinto-da-Rocha (1997) considered the monotypic genus Laneius as a junior subjective
synonym of Progonyleptoidellus, resulting in a new combination, Progonyleptoidellus fuscopictus. Moreover, the
authors also considered Progonyleptoidellus androgynus as a junior subjective synonym of Ampheres striatus, so
they removed this species from Ampheres C.L. Koch, 1839 (Caelopyginae) and assigned it to Progonyleptoidellus
(Progonyleptoidellinae), resulting in the new combination Progonyleptoidellus striatus. Five years later, Kury (2003),
in his annotated catalogue, transferred Geraeocormobius orguensis to Progonyleptoidellus, without justification.
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Key to the males of Progonyleptoidellus
1. Femur IV with dorso-basal apophysis (DBA, see Kury & Pinto-da-Rocha, 1997: figs. 1, 2) . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- Femur IV without DBA (Figs. 2B, 4B) ....................................................................3
2. DBA (=anvil-shaped), merged into the base with the sub DBA; with two small tubercles on scutal area III ...............
............................................................................. Progonyleptoidellus striatus
- DBA acuminate, simple, retrolaterally directed; with a pair of spines on scutal area III (see Kury & Pinto-da-Rocha, 1997: fig.
1) ......................................................................... Progonyleptoidellus fuscopictus
3. Pedipalp coxa long (reaching the apical margin of trochanter IV); basitarsus I not swollen (like the distitarsus); femur III un-
armed; tibia IV unarmed (Fig. 4) ..........................................Progonyleptoidellus picinguaba sp. nov.
- Pedipalp coxa short (not reaching apical margin of trochanter IV); basitarsus I swollen; femur III with ventroapical acuminate
tubercles; tibia IV ventrally armed ...........................................Progonyleptoidellus bocaina sp. nov.
Progonyleptoidellus bocaina sp. nov.
(Figs. 2, 3, 6, 9)
Type material. BRAZIL. São Paulo. São José do Barreiro. Parque Nacional da Serra da Bocaina, Mambucaba river,
1400m, 21–24.iii.1997, Pinto-da-Rocha, Campaner & Vanin leg., ♂ holotype (MZSP 73602); idem, 1 ♂ paratype
(MZSP 18939); same loc., 5–11.i.2008, F. Carbayo et al. leg., 1 ♀ paratype (MZSP 73776); São José do Barreiro, E.
Bocaina road, 44°36’S 22°44’W, 1500m, 8.ii.2008, H.S. de Freitas leg., 1 ♀ paratype (MZSP 30155).
Etymology. The name, a noun in apposition, refers to the type locality, Parque Nacional Serra da Bocaina, a
national park situated between the states of Rio de Janeiro and São Paulo, on a segment of the slopes of the Planalto
Atlântico facing the sea, known as Serra do Mar.
Type locality. BRAZIL, São Paulo, São José do Barreiro: Parque Nacional da Serra da Bocaina.
Geographical distribution (Fig. 9). Known only from the type locality.
Diagnosis. Males. Progonyleptoidellus bocaina sp. nov. resembles P. fuscopictus and P. striatus in having pro-
apical apophysis of coxa IV strong, large and with curved apex, and P. picinguaba sp. nov. in having free tergite III
armed with long spine and absence of DBA on femur IV. It can be distinguished from P. fuscopictus and P. striatus
by the absence of black bands on scutal areas of DS, absence of DBA on femur IV, and free tergite III armed with
a long spine; from P. picinguaba sp. nov. by coxa IV proapical apophysis with curved apex, and femur III with a
retrodorsal and a proventral rows of conical tubercles.
Females. Progonyleptoidellus bocaina sp. nov. resembles P. fuscopictus in coloration, apical apophysis of coxa
IV and armature of femur IV. It can be distinguished from P. fuscopictus by the large DS; coxa IV most prominent
in dorsal view, not totally hidden by DS; free tergite III with a larger spine.
Description. Male (holotype, MZSP 73602)
Measurements: CL 2.5; CW 3.4; DSL 5.6; DSW 6.3; FeP 3.5; FI 4.1; FII 9.8; FIII 7.4; FIV 8.9.
Dorsum (Figs. 2A, 3A, F): DS shape type gamma-pyriform with two median acuminate tubercles on the cor-
ners of its anterior margin, projected obliquely. Frontal hump low, slightly projected anteriorly, with two median
acuminate tubercles upwards. Ocularium narrow, low, with two median and parallel acuminate tubercles directed
upwards, and one pair of tiny tubercles on the posterior face. Carapace with a pair of rounded tubercles posterior
to ocularium and with an oblique row of 3–4 tubercles on each side, in the posterior half. Scutal area I with scat-
tered small tubercles on the posterior half, the paramedian ones slightly larger; scutal area II with scattered small
tubercles, paramedian ones slightly larger; scutal area III with scattered small tubercles, and with a paramedian pair
of large spines directed posteriorly. Lateral margins of DS with 5–6 small acuminate tubercles on posterior half, the
median ones larger. Two ozopores each side. Posterior margin of DS slightly convex, with a row of small tubercles
of similar size. Free tergites I–II each with scattered small tubercles (one median tubercle larger on FT II); free ter-
gite III with a median long spine. Anal operculum unarmed, with some granules.
Venter (Fig. 3C): Coxa I with a median row of enlarged setiferous tubercles; coxa II with two median rows of
small rounded tubercles; coxae III–IV with scattered granules. Coxae II–IV connected by a row of tubercles. Stig-
matic area: posterior margin convex, most of lateral parts touching coxa IV. Stigmata elliptical and slanted.
Chelicerae: Segment I smooth, bulla weakly marked; segment II with scattered granules, fixed finger with 4
teeth; movable finger with 2 teeth.
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FIGURE 2. Progonyleptoidellus bocaina sp. nov., male paratype (MZSP 18939): A, habitus, dorsal view; B, right trochanter–
tibia IV, dorsal view; C, coxa, trochanter and basal part of femur IV, retrodorsal view. Scale bars: A: 3 mm, B: 4 mm, C: 2 mm.
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FIGURE 3. Progonyleptoidellus bocaina sp. nov., male holotype (MZSP 73602) and female paratype (MZSP 73776): A, male
habitus, dorsal view; B, female habitus, dorsal view; C, male habitus, ventral view; D, female trochanter–femur IV, dorsal view;
E, male trochanter–tibia IV, dorsal view; F, male habitus, left lateral view. Scale bars: 1 mm.
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Pedipalps (Fig. 3F): Coxa smooth and enlarged, reaching the cheliceral bulla. Trochanter with a dorsoapical
small elevation, with one apical ventral setiferous tubercle. Femur and patella elongated, slender and smooth. Tibial
setation: mesal IiIi; ectal IiIi. Tarsus with 2 ventral median rows of setae; tarsal setation: mesal IiI; ectal iIiI.
Legs: Coxae I–II with one prodorsal and one retrodorsal small apophyses, the prodorsal ones larger; coxa
III with a prolateral short apophysis fused at apex with the retrolateral apophysis of coxa II. Coxa IV (Figs. 2B,
3A,C) with scattered granules and tubercles; with a strong, large and obliquely directed proapical apophysis, with
an acuminate apex, inclined to the axis of the apophysis, and with a small process on the basal half of apophysis
(basal process); a retroapical small acuminate apophysis. Trochanters I–III unarmed, with some apical tubercles on
retro and prolateral faces; trochanter IV (Figs. 2B, 3A,C,E) with some granules on dorsal face; prolaterally with a
median dorsal large, conical and pointed tubercle and two apical small tubercles; retrolaterally with an apical large,
conical and pointed tubercle and with some small tubercles on the apical half; ventral face with scattered tubercles.
Femora I-II unarmed, with scattered granules; femur III with a retrodorsal and a proventral rows of conical tu-
bercles, increasing in size apically; femur IV (Figs. 2B, 3E) long and slightly sinuous at both the base and the apex;
dorsobasally with a row of spines, decreasing in size apically, and with an adjacent row (retrolateral one) of small
tubercles; retrolaterally with a row of spines, the basal ones larger (the largest armature of the article), decreasing
in size apically; ventrally with two rows of tubercles increasing in size apically, the apical ones spine-like. Patellae
I–IV unarmed, with some scattered tubercles. Tibiae I–II unarmed; tibiae III–IV ventrally with two rows of apical
tubercles, increasing in size apically. Metatarsi I–IV unarmed; astragalus swollen. Claws smooth. Tarsal process
large. Tarsal counts: 7(3), 13–14(4), 9, 9–10.
Penis (MZSP 18939, Figs. 6AB): VP subrectangular, ventral face totally covered with microsetae type 1, distal
margin with a U- shaped cleft, curved in lateral view, with a rounded base. Lateral lobe sub-basal, subquadrangular,
dorsally directed, with microsetae type 3 on its distal part. Macrosetae (MS) A1–A3 well developed, lanceolate,
forming a slanted row on lateral lobe; MS B1 short, conical, separated from MS A; MS C1–C3 long, well developed,
basally straight and distally helicoidal; MS D1 short and conical, placed between MS C and MS A; MS E1–E2 short
and conical, placed near MS C1–C3. Glans stylus subcylindrical with inclined apex, with subapical microspines.
Glans ventral process with stem thicker than stylus, with flabelliform apex. Flabellum fan-like, with multi-serrated
apex. Glans sac short, multi-folded, heel-shaped, projected as a dorsal process. Stylus stem arising from the ventral-
most part of the glans sac, with a candelabrum-like insertion.
Coloration (in ethanol; Fig. 3): Light yellow with small black spots scattered on anterior, posterior and lateral
margins of DS, scutal areas, free tergites I–II, pedipalps, chelicerae and legs I–III. Dark yellow on leg IV (coxa–tib-
ia). Predominantly black on the carapace around and behind the ocularium, in the spines of the scutal area III and
in dorsoapical apophyses of coxae IV. Dry-marks as a strip on ocularium (between the eyes), continuing anteriorly
and posteriorly on the carapace; and weakly in scutal areas sulci.
paratype: Measurements: CL 2.8; CW 3.8; DSL 6.4; DSW 6.9; FeP 3.6; FI 4.2; FII 10.5; FIII 7.6; FIV 9.2.
Anterior margin of dorsal scutum with 3 medium-sized acuminate tubercles on the corners. Lateral margins with
8-9 small acuminate tubercles on posterior half. Free tergite I unarmed; free tergite II with large tubercle on the left
half. Tarsal counts: 7(3), 13–14(4), 8–9, 10.
Female (n=1; Figs. 3B, D)
Measurements: CL 2.7; CW 3.8; DSL 5.8; DSW 6.8; FeP 3.8; FI 4.2; FII 11.2; FIII 7.4; FIV 9.6.
Dorsum: Spines on scutal area III larger and stronger than in males. Free tergites I–II unarmed; FT III with a
huge median spine, slightly stronger than in males. Dry-marks on ocularium and posterior half of carapace (behind
ocularium), and weakly in the middle part of scutal areas I–II.
Legs: Coxa IV shorter than in male, hidden under the dorsal scutum, except its apex and apophysis; with a
proapical large, strong and acuminate apophysis, straight and obliquely directed, much smaller than that of males.
Trochanter IV inserted prolaterally as in male, but with apical tubercle smaller; retrolaterally with small tubercles.
Femur IV with armature as in male, but spines are smaller. Tarsal counts: 7(3), 13(4), 9, 11.
Progonyleptoidellus fuscopictus (B. Soares, 1942)
(Figs. 7C, 9)
Laneius fuscopictus B. Soares, 1942: 6 (desc), fig. 3 (dorsal habitus); 1944a: 258 (cit); 1946: 503 (cit); Soares & Soares, 1985:
184 (cit), fig. 29 (male dorsal habitus), 30 (male pedipalp), 31 (male femur IV), 32 (female dorsal habitus), 33 (female
femur IV), 34 (penis dorsal view), 35 (penis lateral view).).
Gonazula fuscopicta: Soares & Soares, 1949: 173 (cit).
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Progonyleptoidellus fuscopictus: Kury & Pinto-da-Rocha, 1997: 110 (diag, syst), figs. 1 (male dorsal view), 5 (penis dorsal
view), 6 (penis lateral view); Kury, 2003: 198 (cat);
Type series. ♀ holotype, ♂ paratype: BRAZIL, São Paulo, Salesópolis, Boracéia (MZSP 96), examined.
Other material examined. BRAZIL. Rio de Janeiro. Paraty (Paraty–Cunha road, 23º12’65’’S 44º49’19’’W),
C. Bragagnolo et al. leg., 20.iv.2012, 1 ♀ (MZSP 73775). São Paulo. Caraguatatuba, Petrobras Road (km 45), Sítio
Mariscas, D.S. Amorim leg., 8.i.1983, 1 ♂ (MZSP 18940); Cunha (Parque Estadual da Serra do Mar), R. Pinto-
da-Rocha leg., 15.i.2016, 1 ♂ (MZSP 73774); Salesópolis (Estação Biológica da Boracéia, tunnel, 23°38’44.5’’S
45°56’36.2’’W), A. Nogueira et al leg., 5.ii.2014, 3 ♂ 2 ♀ (MZSP 73108); idem, Boracéia, 29.xi.1957, H. Camargo
leg., 1 ♂ (MZSP 14837); same loc., 19.xi.1991, F.A.G. Mello leg., 1 ♀ (MZSP 30176); São Luiz do Paraitinga,
Parque Estadual Serra do Mar, Núcleo Santa Virgínia, Trilha Poço do Pito, G. Machado leg., 18.xii.2004, 1
(MZSP 30161); idem, Parque Estadual Serra do Mar, Núcleo Santa Virgínia, Trilha do Ipiranga, without collector
name , 20.i.2005, 1 ♀ (MZSP 30170); Ubatuba (Oswaldo Cruz road, km 80), C. Bragagnolo et al leg., 19.iv.2012,
1 ♀ (MZSP 73045).
Type locality. BRAZIL. São Paulo. Salesópolis: Boracéia.
Geographical distribution (Fig. 9). BRAZIL. Southeastern portion of São Paulo State.
Diagnosis. Males (fig 7C): Progonyleptoidellus fuscopictus can be distinguished from congeners by the fol-
lowing combination of characters: Scutal area III with pair of median spines; free tergites I–II unarmed; free tergite
III usually unarmed (with median spines in some specimens); proapical apophysis of coxa IV long, strong, with
curved apex; femur IV with DBA (sinple, acuminate, prolaterally directed), with dorsal row of spines on basal half,
retrolaterally directed, decreasing in size apically; retrodorsally with row of strong and large spines, decreasing in
size apically; ventrally with two rows of small spines; carapace dark black; lateral margin of DS yellowish; scutal
areas I–III with black bands, but not continuous as in P. striatus; dry-mark on anterior part of carapace (in front of
ocularium) and on DS sulci.
Females: Progonyleptoidellus fuscopictus can be distinguished from the other species of the genus by the fol-
lowing combination of characters: Proapical apophysis of coxa IV strong, long, straight and acuminate; femur IV
without DBA, dorsally armed like in males; retrodorsally with spines smaller than in males; proventrally with me-
dian spines (larger than males).
Progonyleptoidellus picinguaba sp. nov.
(Figs. 4, 5, 6, 9)
Progonyleptoidellus sp.: Almeida-Neto et al, 2006: 367 (dist).
Type material. BRAZIL. São Paulo. Ubatuba, Picinguaba, Morro do Cuscuzeiro, 21.xii.1995, G. Machado leg., ♂
holotype (MZSP 73601), 2 ♀ paratypes (MZSP 16257); same loc., 20.xii.1995, G. Machado leg., 3 ♂ 4 ♀ (MZSP
16258), 3 ♂ 7 ♀ (MZSP 16259); same loc., 25–29.iv.1991, P. Gnaspini leg., 1 ♂ (MZSP 16009); Ubatuba, Parque
Estadual Serra do Mar, núcleo Picinguaba, 600m, 14.iii.2008, F. Esteves & R. Feitosa leg., 1 ♀ paratype (MZSP
30176).
Etymology. The name, a noun in apposition, refers to the type locality, a region in the municipality of Ubatuba,
São Paulo, Brazil.
Type locality. BRAZIL. São Paulo. Ubatuba: Picinguaba, Morro do Cuscuzeiro
Geographical distribution (Fig. 9). Known only from the type locality.
Diagnosis. Males. Progonyleptoidellus picinguaba sp. nov. can be distinguished from the other species of the
genus by the following combination of characters: Free tergite III armed with a long spine; proapical apophysis on
coxa IV straight, without curved apex; femur III unarmed; femur IV without DBA. Females. This species differs
from the other species of the genus in the absence of sexual dimorphism on armature of coxa IV and femur IV.
Description. Male holotype (MZSP 73601)
Measurements: CL 2.2; CW 3.2; DSL 4.3; DSW 5.1; FeP 3.7; FI 4.5; FII 12; FIII 8; FIV 10.6.
Dorsum (Figs. 4A, 5A, F): DS shape type gamma-P, with two median acuminate tubercles on the corner of
its anterior margin. Frontal hump low, slightly projected anteriorly, with two upwards-pointing median acuminate
tubercles. Ocularium narrow, low, with two parallel acuminate tubercles upwards and one pair of small tubercles
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posteriorly. Carapace with a pair of rounded tubercles posterior to the ocularium and a few scattered granules. Scutal
areas I–II each with a paramedian pair of tiny tubercles; scutal area III with a paramedian pair of large spines di-
rected posteriorly, and scattered granules. Lateral margins of DS with 2–3 small tubercles and one median spine on
posterior 1/3. Two ozopores. Posterior margin of DS straight with prominent corners, with a row of small tubercles
(two median ones larger). Free tergites I–III each with a row of varied-sized tubercles, predominantly small; free
tergites II–III each with a large median spine. Anal operculum unarmed.
FIGURE 4. Progonyleptoidellus picinguaba sp. nov., male holotype (MZSP 73602): A, habitus, dorsal view; B, right trochant-
er–tibia IV, dorsal view; C, coxa, trochanter and basal part of femur IV, retrodorsal view. Scale bars: A, C: 2 mm, B: 3 mm.
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FIGURE 5. Progonyleptoidellus picinguaba sp. nov., male paratype (MZSP 16009) and female paratype (MZSP 16257): A,
male habitus, dorsal view; B, female habitus, dorsal view; C, male habitus, ventral view; D, male trochanter–tibia IV, dorsal
view; E, female trochanter–patella IV, dorsal view; F, male habitus, left lateral view. Scale bar: 1 mm.
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FIGURE 6. Scanning electron micrographs showing the distal part of Progonyleptoidellus spp. penis: A–B, P. bocaina sp. nov.
(MZSP 18939), A. dorsal, B. lateral; C–D, P. picinguaba sp. nov. (MZSP 16259) , A. dorsal, B. lateral. Scale bars: 20 µm.
Venter (Fig. 5C): Coxa I with a median row of enlarged setiferous tubercles; coxa II with a median row of small
tubercles; coxae III–IV with scattered tubercles. Coxae II–IV connected by a row of tubercles. Posterior margin of
the stigmatic area convex, most lateral parts touching coxa IV. Stigmata elliptical and slanted.
Chelicerae: Segment I with scattered granules; bulla weakly marked; segment II with scattered granules; fixed
finger with 3–4 teeth; movable finger with 3–4 teeth.
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Pedipalps (Fig. 5F): Coxa smooth and enlarged, reaching the cheliceral bulla and trochanter I. Trochanter with
a dorsoapical small elevation, with one apical and one basal ventro-median small setiferous tubercle. Femur and
patella elongated, slender and smooth. Tibial setation: mesal IiIi; ectal IiIi. Tarsus with 2 ventral median row of
setae; tarsal setation: mesal Ii; ectal II.
Legs: Coxae I–II with one prodorsal apophysis; coxa III with two retrolateral apophyses. Coxa IV with scat-
tered granules; a strong, large and straight proapical apophysis, with an acuminate apex, obliquely directed, and a
retroapical small, acuminate apophysis. Trochanters I–III unarmed, with some apical small tubercles; trochanter IV
(Figs. 4B, 5A,D) prolaterally with one sub-basal and one apical tubercle, these short and acuminate, the sub-basal
one the largest; retrolaterally with one median and one apical tubercles, these short and acuminate, the apical larg-
est; ventral face with scattered granules. Femora I–III unarmed, with scattered granules; femur IV (Figs. 4B, 5D)
straight and long; dorsobasally with a row of 4–6 median spines (those located in the middle of the row larger);
retrodorsally with a row of intercalated tubercles and spines of irregular size (those located in the middle of the
row larger); proventrally with a row of very short conical tubercles. Patellae and tibiae I–IV unarmed, with ventral
scattered granules. Metatarsi I–IV smooth, astragalus swollen. Claws smooth. Tarsal process large. Tarsal counts:
.7(3), ?, 12, 15.
Penis (MZSP 16259, see Figs. 6C–D): VP subrectangular, ventral face totally covered with microsetae of type
1; distal margin with a U shaped cleft, slightly curved in lateral view. Lateral lobe sub-basal, sub-rectangular shaped,
dorsally directed, with microsetae type 3 on its distal part. Macrosetae (MS) A1–A3 well developed, lanceolate,
forming a slanted row on the lateral lobe; MS B1 short, conical, separated from MS A; MS C1–C3 long, well devel-
oped, basally straight and distally helicoidal; MS D1 short and conical, placed between MS C and MS A; MS E1–E2
short and conical, placed near MS C1–C3. Glans stylus subcylindrical with inclined apex, ornate with microspines
subapically. Glans ventral process with stem thicker than stylus, and with flabelliform apex. Flabellum fan-like,
with multi-serrated apex. Glans sac short, multi-folded, heel-shaped, projected as a dorsal process. Stylus stem aris-
ing from the ventralmost part of glans sac, inserted as a candelabrum-like.
Coloration (in ethanol; see Fig. 5): Light yellow with small black spots scattered on anterior, posterior and
lateral margins, as well as on scutal areas of DS, ocularium, free tergites I–III, pedipalps, chelicerae and legs. Pre-
dominantly black on the carapace around ocularium, on the scutal area III spines and on dorsoapical apophyses of
coxae IV. Dry-marks on scutal areas sulci, anterior margin of DS, anterior half of carapace and sparsely on posterior
half of carapace.
Variation in males (n=7): Measurements: CL 2.1–2.4; CW 3–3.2; DSL 4.4–5; DSW 4.6–5.5; FeP 3.2–3.6; FI
4.2–4.7; FII 11.1–12.4; FIII 7.5–8.2; FIV 9.9–11.5. Anterior margin of DS with 3–4 median acuminate tubercles on
the corners (if more than three, the fourth much smaller than the others). Posterior margin of DS slightly sinuous
in some males; armed with one or more acuminate tubercles. Free tergite I–II armed with one or more spines; free
tergite III with more than one large spine. The armature of posterior margin of DS and free tergites I–III extremely
variable in number, size, position and could be absent in some specimens. Pedipalps: tibia setation: mesal, IiIi, IIiIi,
IiII; ectal IiIi, II; tarsal setation: mesal Ii, II, IIII; ectal Ii, II; proapical apophysis with apex slightly slanted. Armature
of femur IV weak, with spines like tubercles. Dry-mark weak (inconspicuous in some regions); in some specimens
totally or partially absent (in the latter case, notably absent in scutal areas sulci). Tarsal counts: 7(3), 13–14(3–4),
11–13, 15–18.
Female (n=9; Figs. 5B, E)
Measurements: CL 2.2–2.5; CW 3–3.4; DSL 5–5.4; DSW 5.1–5.6; FeP 3–3.6; FI 3.9–4.4; FII 11.2–12.3; FIII
7.5–8.7; FIV 10.3–12.6.
Dorsum: Spines on scutal area III greater than in males in some specimens. Posterior margin of dorsal scutum
sub straight (sinuous appearance); posterior margin of DS unarmed or armed with acuminate tubercles. Free tergites
I–III armed. Like in males, the armature of posterior margin of DS and free tergites I–III extremely variable in
number, size, position, usually with more than one spine; free tergites I–III armature generally greater than those on
males (although some specimens have less robust armature).
Pedipalps: tibial setation: mesal, IiIi, IiII; ectal IiIi; tarsal setation: mesal Ii, II; ectal Ii, II.
Legs: Proapical apophysis of coxa IV like in males in size; vary in stoutness (some specimens with apophysis
stouter than in males). Tarsal counts: 6–8(3), 13–14(4), 12–13, 15–16.
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FIGURE 7. Photographs of living specimens of Progonyleptoidellus: A, P. striatus (male from Brazil, São Paulo, Santo André,
Estação Biológica Alto da Serra de Paranapiacaba); B, P. striatus (female from same locality as male); C, P. fuscopictus (male
from Brazil, São Paulo, Cunha, Parque Estadual da Serra do Mar). Photos by R. Pinto-da-Rocha.
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Progonyleptoidellus striatus (Roewer, 1913)
(Figs. 7A, 7B, 9)
Ampheres striatus Roewer, 1913: 337 (desc), pl. 1, fig. 5 (dorsal habitus); 1923: 531 (rdesc), fig. 664 (dorsal habitus); Mello-
Leitão, 1923: 175 (cit); 1932: 372 (rdesc), fig. 236 (dorsal habitus); Soares & Soares, 1948: 570 (cit).
Progonyleptoidellus striatus: Kury & Pinto-da-Rocha, 1997: 110 (diag, syst), figs. 2 (male dorsal habitus), 3 (female dorsal
habitus), 7 (penis dorsal view) and 8 (penis lateral view); Pinto-da-Rocha, 2002: 361 (cit), 371 (cit), 387 (cit), 379 (syst);
Kury, 2003: 198 (cat); DaSilva & Pinto-da-Rocha, 2010: 579 (cit), 581 (syst), 585–586 (syst), 641–642 (mat); DaSilva &
Gnaspini, 2010: 535 (cit), 539–541 (mat), 600 (syst), 621 (syst); Pinto-da-Rocha & Bragagnolo, 2010: 510 (cit); 511 (syst),
512 (mat); DaSilva & Pinto-da-Rocha, 2012: 27 (mat), 37 (syst); Caetano & Machado, 2013: 593 (cit), 595 (syst), 596
(cit), 609 (mat); Mendes & Barros, 2013: 322 (syst); Pinto-da-Rocha et al, 2014: 6 (cit), 12 (syst); Mori & Pinto-da-Rocha,
2015: 543 (mat); 549 (syst).
Metarthrodes melanacanthus Mello-Leitão, 1923: 174 (cit) [non Roewer 1913: misidentification]
Progonyleptoidellus androgynus Piza, 1940: 63 (desc), fig 11 (dorsal habitus); Soares & Soares, 1985: 182 (rdesc), figs. 17
(male dorsal habitus), 18 (male ocularium), 19 (dorsobasal apophysis of femur IV), 20 (female dorsal habitus), 21 (female
ocularium), 22 (female scutal area III armature), 23 (female femur IV), 24 (male tarsus I), 25 (male chelicera), 26 (male
pedipalp), 27 (penis dorsal view) and 28 (penis lateral view) [=Cadeadoius atroluteus Roewer, 1943] [=Piraquara schu-
barti Piza, 1943]; Kury & Pinto-da-Rocha, 1997: 109–110 (cit, syst). Synonymy established by Kury & Pinto-da-Rocha,
1997.
Gonyleptoides androgynus: B. Soares, 1944a: 258 (syst); 1944b: 281 (cit); 1946: 501 (cit).
Geraecormobius androgynus: Soares & Soares, 1949: 167 (cit).
Piraquara schubarti Piza, 1943: 258 (desc), fig. 3 (dorsal habitus); Paschoal & Barros, 1983: 78 (cat); Coronato et al., 2013:
508 (cat), fig 8I (dorsal habitus).
Gonyleptoides schubarti: B. Soares, 1944a: 258 (syst); 1946: 502 (cit).
Geraeocormobius schubarti: Soares & Soares, 1949: 172 (cit).
Cadeadoius atroluteus Roewer, 1943: 45 (desc), pl. 6, figs 51 (dorsal habitus), 51a (dorsobasal apophysis of femur IV), 51b
(pedipalp tarsus).
Geraecormobius atroluteus: Soares & Soares, 1949: 168 (cit).
Type series. BRAZIL, Bahia. 2 ♂ syntypes (SMF RI). Type of Progonyleptoidellus androgynus: BRAZIL, São
Paulo, Alto da Serra, ♀ holotype (MZSP 1579, examined). Type of Piraquara schubarti: BRAZIL, São Paulo, Ser-
ra Piraquara, Itanhaém, ♂ holotype (MZLQ A0046, now IBSP 10084. Types of Cadeadoius atroluteus: BRAZIL,
Santa Catarina, Seara, Nova Teutônia, 5 ♀ 2 ♂ syntypes (SMF RII 6428/61).
Other material examined. BRAZIL. São Paulo. Cotia, Reserva Morro Grande, without collector’s name,
28.iii.2003, 1 ♂ (MZSP 25922); Miracatu, Fazenda Iterei (24º03’S 47º13’W), Equipe Biota leg., 20–26.ix.2001,
3 ♂ (MZSP 30837), 3 ♂ 1 juvenile (MZSP 30832); Santo André, Paranapiacaba, Estação Biológica Alto da Serra,
R. Pinto-da-Rocha & S.A.Casari leg., 25–26.i.1997, 1 1 ♀ (MZSP 17650); same loc., R. Pinto-da-Rocha leg.,
27.viii.1999, 2 ♂ 2 ♀ (MZSP 18892); Salesópolis, Estação Biológica Boracéia, S.A. Casor & G.J.M. Santos leg.,
23–28.i.2002, 1 ♀ (MZSP 30180); Santos, trilha Caetê, C. Magenta leg., x.2005, 1 ♂ 1 ♀ (MZSP 30168); São Paulo,
Parelheiros Clube Campestre, M. Vicente leg., 23.ix.2001, 1 ♂ (MZSP 30166); São Vicente, Japuí, trilha Itaquitan-
duva, DaSilva et al. leg., 7.iii.2004, 2 ♀ (MZSP 30169).
Type localities. BRAZIL. Of Cadeadoius atroluteus: Santa Catarina, Seara: Nova Teutônia. Of Piraquara
schubarti: São Paulo, Itanhaém: Serra Piraquara. Of Ampheres striatus: Bahia. Of Progonyleptoidellus androgynus:
São Paulo, Santo André: Alto da Serra. The type localities in the Brazilian states of Bahia and Santa Catarina are
dubious and probably mislabeled considering that there are no additional records of the whole genus from these
states.
Geographical distribution (Fig. 9). BRAZIL: Southeastern portion of the state of São Paulo.
Diagnosis. Males (see Fig. 7A): Progonyleptoidellus striatus can be distinguished from the other species of the
genus by the following combination of characters: pedipalp coxa small; all scutal areas unarmed; scutal area III with
two paramedian short tubercles; free tergites I–III unarmed. Proapical apophysis of coxa IV strong and often with
curved and bifurcated apex, ventral branch rounded and small. Trochanter IV retrolaterally with short apical spine;
prolaterally with large, triangular sub basal spine. Femur IV: DBA anvil-shaped, fused basally with the sub DBA
(triangular shaped); with seven rows of armature — dorsally with two rows, one more prolateral (with a few small
spines), and one retrolateral of spines along the entire article, decreasing in size apically; prodorsally with a row of
short spines, all article long (more concentrated in the apical half); retrodorsally with strong spines along the article
(the largest in the medial part; these are the strongest spines on femur IV); an extra row in the apical part, between
CLADISTICS AND TWO NEW PROGONYLEPTOIDELLUS Zootaxa 4691 (5) © 2019 Magnolia Press · 485
retrodorsal and second dorsal rows; and ventrally with two rows of spines throughout article, increasing in size
apically; carapace black with lateralmost parts yellowish. DS areas with dark black bands; dry-mark on ocularium
(between the eyes), carapace (behind the ocularium), weakly on transversal DS sulci. Penis with macroseta B of the
same size as macrosetae A.
Female (see Fig. 7B): This species can be distinguished from the others in the genus by the following combina-
tion of characters: Scutal area III with two small acuminate tubercles; proapical apophysis of coxa IV long (smaller
than males), straight and acuminate (apex more rounded than males); femur IV without DBA, armed with seven
rows of spines (in a different pattern than males); coloration like in males.
Deltaspidium Roewer, 1927
Deltaspidium Roewer, 1927: 348 (desc); 1930: 416 (diag); Mello-Leitão, 1932: 288 (diag); 1935c: 104 (cit); Soares & Soares,
1949: 164 (diag); 1986: 91 (diag, syst); Kury, 2003: 123 (cat); Bragagnolo & Pinto-da-Rocha, 2003: 5 (cit), 6 (cit); Kury
& Alonso-Zarazaga, 2011: 54 (cat).
Adhynastes Roewer, 1930: 438 (desc); Mello-Leitão, 1930: 214 (cit); 1932: 273 (diag); 1935: 103 (cit); Soares & Soares, 1949:
158 (diag); 1985: 194 (cit); Kury, 2003: 121 (cat). . Syn. nov.
Geraecormobius (part): Soares & Soares, 1954 (desc); Bragagnolo & Pinto-da-Rocha, 2003: 5 (cit).
Progonyleptoidellus (part): Kury, 2003 (cat, syst); Pinto-da-Rocha & Bragagnolo, 2010: 510 (cit), 511 (syst), 512 (mat), 513
(cit).
Type species. Deltaspidium bresslaui Roewer, 1927, by monotypy (junior synonym of Gonyleptes asper Perty, 1833). Type
species of Adhynastes: Adhynastes tenuis Roewer, 1930, by monotypy.
Included species. Deltaspidium asperum (Perty, 1833); Deltaspidium tenue comb. nov. (Roewer, 1930) and Del-
taspidium orguense comb. nov. (Soares & Soares, 1954).
Etymology. From the Greek noun “delta” (a letter of alphabet) and “hispidium”, small shield, in reference to
the triangular shape of dorsal scutum. Gender neutrum.
Distribution. Deltapisdium spp. were recorded in Serra dos Órgãos mountain range, and Tijuca, both in central
Rio de Janeiro State, Brazil. The type-species of the genus has no Brazilian locality explicitly assigned. However,
the collectors of the type material, J.B. Spix and C.F von Martius collected in Rio de Janeiro State.
Diagnosis. Deltaspidium can be distinguished from the other genera of Progonyleptoidellinae by the following
combination of characters: dorsal scutum outline (Fig. 8) gamma triangular (γT); mesotergum divided into three
scutal areas; posterior margin of DS convex or sub straight; scutal areas I–II unarmed; area III armed with two para-
median spines directed posteriorly (greater and closer to each other in some females); coxa IV with large proapical
apophysis with curved apex in males, reduced and straight females; femur IV without DBA, similar in both sexes;
tarsus IV with more than 16 tarsomeres; penis with deep cleft on VP, lateral lobes placed sub basally; VP trapezoid,
without globous base in lateral view; MS A + B placed in line; MS B smaller than MS A; MS C straight; flabellum
with one apical long projection; stylus sub sigmoid, parallel to ventral process shaft.
Taxonomical note. In addition to the evidence of the present phylogenetic analysis, a comparison among the
three species revealed several similarities including robust pedipalps, outline of dorsal scutum gamma triangular
(γT), huge spines in area III, usually fused at base, proapical apophysis of coxa IV of males directed proximally to
trochanter IV and mottled coloration. These features and the hypothesis of species relationships led us to propose a
new delimitation of the genus Deltaspidium.
Deltaspidium asperum (Perty, 1833)
Gonyleptes asper Perty, 1833: 202 (desc) (Type locality and repository unknown; probably formerly in Zoologische Staatssam-
mlung München, now lost).
Ampheres asper: Koch, 1839a: 17 (desc); 1839b: 71 (rdesc), pl 235, fig 570 (dorsal habitus); Roewer, 1913: 338 (rdesc); 1923:
532 (rdesc); Mello-Leitão, 1923: 175 (cit); Roewer, 1931: 136 (rdesc); Mello-Leitão, 1932: 374 (rdesc); B. Soares, 1945c:
348 (cit); Soares & Soares, 1948: 567 (cit).
Deltaspidium asper: Soares & Soares, 1986: 91 (syst), figs. 7 (penis dorsal view), 8 (penis lateral view); Kury, 2003: 123;
Bragagnolo & Pinto-da-Rocha, 2003: 5 (cit), 6 (cit). Nomenclatural note: previous spelling of the specific name is incorrect,
since when combined to Deltaspidium (grammatical gender neuter), it is to be corrected to asperum.
BENEDETTI & PINTO-DA-ROCHA
486 · Zootaxa 4691 (5) © 2019 Magnolia Press
Deltaspidium bresslaui Roewer, 1927: 348 (desc), fig. 16 (male dorsal habitus); 1930: 416 (rdesc), pl 6, fig. 3 (male dorsal ha-
bitus); Mello-Leitão, 1932: 288 (rdesc); Soares & Soares, 1949: 164 (cit) (BRAZIL. Rio de Janeiro. Teresópolis: Serra dos
Órgãos. 1 ♂ 1 ♀ syntypes (SMF) examined from digital photos). Synonymy established by Soares & Soares, 1986.
Deltaspidium tenue (Roewer, 1930) comb. nov.
Adhynastes tenuis Roewer, 1930: 439 (desc), fig. 41 (male dorsal habitus); Mello-Leitão, 1932: 274 (rdesc); Soares & Soares,
1949: 159 (cit); Kury, 2003: 121 (cat). (BRAZIL. Rio de Janeiro. Rio de Janeiro, Tijuca, 1 ♂ 1 ♀ syntypes (SMF RII
1340/28) examined from digital photos).
Deltaspidium orguense (Soares & Soares, 1954) comb. nov.
(Fig. 8)
Geraecormobius orguensis Soares & Soares, 1954: 492 (desc), fig. 4 (female dorsal habitus); Bragagnolo & Pinto-da-Rocha,
2003: 5 (cit). Holotype ♀ (MZSP 1153) examined): BRAZIL. Rio de Janeiro. Teresópolis, Parque Nacional Serra dos
Órgãos.
Progonyleptoidellus orguensis: Kury, 2003: 198 (cat, syst); Pinto-da-Rocha & Bragagnolo, 2010: 510 (cit), 511 (syst), 512
(mat), 513 (cit).
FIGURE 8. Photographs of living specimens of Deltaspidium orguense comb. nov. (Brazil, Rio de Janeiro, Parque
Nacional da Serra dos Órgãos); A, male; B, female. Photos by Arthur Anker.
Discussion
This is the eleventh cladistic analysis in which representatives of Progonyleptoidellus were included (Pinto-da-
Rocha, 2002; Vasconcelos, 2009; DaSilva & Gnaspini, 2010; DaSilva & Pinto-da-Rocha, 2010; Pinto-da-Rocha &
Bragagnolo, 2010; DaSilva & Pinto-da-Rocha, 2012; Caetano & Machado, 2013; Mendes & Barros, 2013; Pinto-da-
Rocha et al., 2014; Mori & Pinto-da-Rocha, 2015). However, in most of them only P. striatus was included either
as an outgroup or as a representative of Progonyleptoidellinae. The monophyly of the genus has been previously
CLADISTICS AND TWO NEW PROGONYLEPTOIDELLUS Zootaxa 4691 (5) © 2019 Magnolia Press · 487
tested only partially by Pinto-da-Rocha & Bragagnolo (2010), as a by-product of the analysis of Sodreaninae. These
authors included P. orguensis and P. striatus in their analysis as outgroups, resulting in the recovery of P. orguensis
as the sister group of a clade containing (Sodreaninae (Caelopyginae + Progonyleptoidellinae), the latter including
P. striatus (hence, suggesting the non-monophyletic condition of the genus in that extension). However, Pinto-da-
Rocha & Bragagnolo (2010) did not make any taxonomical decision regarding the placement of P. orguensis.
As in most previous analyses, the monophyly of the subfamily Progonyleptoidellinae has not been corrobo-
rated here. One exception can be found in Pinto-da-Rocha (2002), who recovered the monophyly of the subfamily
with a smaller taxa representation (only P. striatus and Heliella singularis). Pinto-da-Rocha & Bragagnolo (2010)
demonstrated that Progonyleptoidellinae was monophyletic (and sister group to Caelopyginae) only if P. orguensis
is removed from the subfamily. Vasconcelos (2009) and Caetano & Machado (2003) found that Progonyleptoidel-
linae was paraphyletic with respect to Caelopyginae. DaSilva & Pinto-da-Rocha (2010) also recovered a close
relationship between Progonyleptoidellinae and Caelopyginae, but with Sodreaninae also inserted in the clade.
Alternatively, Pinto-da-Rocha et al. (2014) demonstrated that Progonyleptoidellinae was paraphyletic with respect
to Neosadocus (Gonyleptinae). The proximity of Neosadocus with Progonyleptoidellinae was also demonstrated by
Vasconcelos (2009) and Caetano & Machado (2013).
In the present analysis, Progonyleptoidellinae was recovered as polyphyletic, with three independent lineages:
1—Iporangaia pustulosa plus Mitopernoides variabilis; 2—Heliella singularis plus Cadeadoius niger; 3—Prog-
onyleptoidellus, with exclusion of P. orguensis. This result is not supported by other analyses in the literature.
Furthermore, Caelopyginae was here recovered nested within a clade with some alleged Progonyleptoidellinae
(Heliella, Cadeadoius and Progonyleptoidellus), a result in line with some previous hypotheses (Pinto-da-Rocha
2002, DaSilva & Pinto da Rocha 2012). The proximity of Neosadocus with Progonyleptoidellinae was recovered
only with the inclusion of Deltaspidium and Caelopyginae. However, it is not the purpose of this paper to clarify the
relations of Progonyleptoidellinae and the K92 clade, since a more comprehensive analysis, including molecular
evidence, conducted by both authors is in progress.
FIGURE 9. Distribution of Progonyleptoidellus spp. in Coastal Atlantic Rain Forest of São Paulo and Rio de Janeiro States,
Brazil. White symbols represent literature records.
BENEDETTI & PINTO-DA-ROCHA
488 · Zootaxa 4691 (5) © 2019 Magnolia Press
Acknowledgments
We are grateful to Adriano Kury (MNRJ) for his advices on nomenclatural issues, and also for suggestions made by
him and Daniel Proud on an early draft of the manuscript. We thank the curators Adriano Kury (MNRJ), Antônio
Brescovit (IBSP) and Peter Jäger (SMF) for access to the deposited material. Arthur Anker (UFGO) gently sent
us the photos displayed on Fig. 8. Victor Townsend kindly reviewed the English of last version of the manuscript.
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 also thanks CNPq for a PhD fellowship (142170/2013-5).
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... Gonyleptidae is one of the largest families of Grassatores, and it has been the focus of many phylogenetic analyses (Pinto-da- Rocha et al. 2014;Benavides et al. 2021). One of the most used species in such analyses is Gonyleptes horridus Kirby, 1819 (for instance, Pinto-da- Rocha et al. 2014;Benedetti and Pinto-da-Rocha 2019;Ázara et al. 2020), because it is the type-species of Gonyleptes Kirby, 1819, that names both the polyphyletic subfamily Gonyleptinae (second largest one; Pinto-da- Rocha et al. 2014) and the family. However, despite its pivotal taxonomic importance, the defensive secretion of G. horridus is still unknown. ...
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... The inclusion of more molecular data and a morphological cladistic analysis may assist the elucidation of the relationships among the species of Neosadocus. [58], also based on molecular data. Caetano & Machado (2013) [76] also suggested that Neosadocus and Progonyleptoidellinae are closely related, based on behavioral, chemical and ecological data. ...
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... The results of the multilocus Bayesian analysis suggests that N. 964 maximus and N. robustus are more closely related to each other than each is to N. bufo; 965 however, this result needs to be looked at with caution, since the posterior probability 966 of this clade is relatively low (0.82), as in all other phylogenetic trees (Fig 12, S3 Fig, S4 967 Fig, S5 Fig). The inclusion of more molecular data and a morphological cladistic (2014) [42] and Benedetti & Pinto-da-Rocha (2019) [54], also based on molecular data. 973 Caetano & Machado (2013) [75] also suggested that Neosadocus and 974 Progonyleptoidellinae are closely related, based on behavioral, chemical and ecological 975 data. ...
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Neosadocus harvestmen are endemic to the Southern Brazilian Atlantic Forest. Although they are conspicuous and display great morphological variation, their evolutionary history and the biogeographical events underlying their diversification and distribution are still unknown. This contribution about Neosadocus includes the following: a taxonomic revision; a molecular phylogenetic analysis using mitochondrial and nuclear markers; an investigation of the genetic structure and species' diversity in a phylogeographical framework. Our results show that Neosadocus is a monophyletic group and comprises four species: N. bufo, N. maximus, N. robustus and N. misandrus (which we did not find on fieldwork and only studied the female holotype). There is astonishing male polymorphism in N. robustus, mostly related to reproductive strategies. The following synonymies have resulted from this work: Bunoweyhia variabilis Mello-Leitão, 1935 = Neosadocus bufo (Mello-Leitão, 1926); and Bunoweyhia minor Mello-Leitão, 1935 = Neosadocus maximus (Giltay, 1928). Most divergences occurred during the Miocene, a geological epoch marked by intense orogenic and climatic events in the Brazilian Atlantic Forest. Intraspecific analyses indicate strong population structure, a pattern congruent with the general behavior and physiological constraints of Neotropical harvestmen.
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The subfamily Gonyleptinae is the second largest in Gonyleptidae, harboring over 100 species. Gonyleptinae is polyphyletic, nestled in the clade K92, and despite its richness, several species of that subfamily have not had their chemicals of the defensive secretions analyzed. Among these are Gonyleptes curticornis (Mello-Leitão, 1940) and G. horridus Kirby, 1819, the latter being particularly important because it is the type species of the genus, which in turn names the subfamily. Gonyleptes horridus is also used in many phylogenetic analyses, be it using morphological or molecular data. The chemical study of the secretions of these two species by GC-MS and NMR ¹ H showed the presence of 1-(6-isopropyl-3,4-dihydro-2H-pyran-2-yl)-methylbutanone, 1-(6-isopropyl-3,4-dihydro-2H-pyran-2-yl) isobutanone and 4- methyl-1-hepten-3-one in both species. On the other hand, 4-methyl-1 hexen-3-one, benzaldehyde and 3-octanone were observed only in G. curticornis . Both species are Gonyleptinae and chemical mapping of the group corroborates that vinyl ketones are synapomorphy of K92 and that the compound 1-(6-isopropyl-3,4-dihydro-2H-pyran-2-yl) isobutanone is synapomorphic to G. curticornis and G. horridus , but homoplastic to the genus Sodreana Mello-Leitão, 1922. 1-(6-(1-methyl-propyl)3,4-dihydro-2H-pyran-2yl)2-methylbutanone and 4-methyl-1-hepten3-one is also synapomorphic to G. curticornis and G. horridus , but homoplastic in Moreiranula saprophila .
Article
Background K92 is a clade of Gonyleptidae harvestmen of which the greatest diversity lies within the large non-monophyletic subfamily Gonyleptinae. K92 formally includes five subfamilies: Caelopyginae Sørensen, 1884, Gonyleptinae Sundevall, 1833, Hernandariinae Sørensen, 1884, Progonyleptoidellinae Soares & Soares, 1985 and Sodreaninae Soares & Soares, 1985. The basic taxonomy of K92 presently finds itself in an unsatisfactory state, with many species poorly known and supraspecific taxa defined by a combination of few trivial and/or misconstrued features. Questions Three lines of investigation regarding K92 are pursued in this project: (1) some formal gonyleptine genera, such as Liogonyleptoides Mello-Leitão and de, 1925 and Parapachyloides Roewer, 1913a were suspected to be especially related to each other as members of an undiscovered non-gonyleptine taxon. Do those species, distributed along the semi-arid diagonal in South America plus the Atlantic Forest in eastern Brazil really form a clade? (2) Has the acquisition of the caelopygine facies by species of Progonyleptoidellinae and Sodreaninae been gradual and independent, associated with diurnal and arboreal habits? (3) Does the attenuation of sexual dimorphism present in K92 have a phylogenetic signal? Methods A cladistic analysis of some critical taxa in K92 is performed by means of maximum parsimony and Bayesian inference. A matrix with 112 morphological characters and 61 terminals has been analyzed for this project, focusing on (a) potential relatives of Liogonyleptoides, and (b) species of K92 with varied degrees of sexual dimorphism. Main results For question (1), a phylogenetic hypothesis is proposed herein for the larger clades of K92 as follows: (a) a branch in the phylogeny of K92 is here recognized and formally described as the new subfamily Cearininae. The Cearininae subfam. nov. includes the following genera: Cearinus Roewer, 1929 (type genus), Inhuma Piza, 1938, Liogonyleptoides Mello-Leitão and de, 1925 (incl. Pachyleptes Mello-Leitão and de, 1932), Pajeuia gen. nov., Parapachyloides Roewer, 1913b and Ypsilonurus Mello-Leitão and de, 1933a. (b) Gonyleptinae did not resolve here as a clade. (c) A monophylum never reported before is recovered here (clade I, here nicknamed CAPS), consisting of the nominal subfamilies Caelopyginae, Progonyleptoidellinae and Sodreaninae. Progonyleptoidellinae is polyphyletic. All of these subfamilies have their internal composition altered. For question (2), we found that the caelopygine facies (diurnal, arboreal, triangular bodied, brightly colored animals with long legs, high tarsal counts and pectinated claws) probably arose independently in (a) the classic Caelopyginae (including pectinated tarsal claws, the full caelopygine facies), (b) partial caelopygine facies of extended Sodreaninae, as represented by Heliella, and (c) partial caelopygine facies of extended Caelopyginae, as represented by Progonyleptoidellus. For question (3), we found that attenuated sexual dimorphism may evolve via two different processes: females acquiring armature and males losing armature. The three different forms of sexual dimorphism provided insight to the phylogeny of K92: (a) strong sexual dimorphism, where females are plump and unarmed at leg IV in contrast with lean and strongly armed males is typical for most members of K92 and other Gonyleptidae and is the primitive condition; (b) a process of masculinization, where the lean, polygonal females resemble males, only with the armature of leg IV somewhat attenuated, occurs in some Caelopyginae and Progonyleptoidellinae in the traditional sense and reaches the most extreme degree in the members of the genus Moreiranula Roewer, 1930; (c) a process of feminization, where males have weakly developed coxa IV, no differential armature on leg IV, and are almost indistinguishable from the females. This occurs in some Hernandariinae such as Piassagera Roewer, 1929, one species of Moreiranula and some Progonyleptoidellinae, such as Iporangaia Mello-Leitão and de, 1935a and Mitopernoides B. Soares, 1945, and one Caelopyginae strictu sensu, Thereza Roewer, 1923. Other taxonomic results As byproducts of this project, not direct consequences of the phylogenetic analysis, taxonomic changes are proposed for some genera and species in K92. For conciseness, the equality sign (x = y) is here used instead of the expression “x is here newly considered a junior subjective synonym of y”: (1) Cearinus was hitherto monotypic. A second species is described here – Cearinus torulosus sp. nov. – from Mato Grosso State, Central Brazil. (2) Liogonyleptoides is reviewed. There are currently 5 species included, of which one (Liogonyleptoides heliae Kury 2003) is a species inquirenda, another is a synonym (Liogonyleptoides capichaba = Liogonyleptoides tetracanthus) and a third (Liogonyleptoides minensis) is here transferred to the new monotypic genus Pajeuia. A new diagnosis of the genus Liogonyleptoides is proposed and emended diagnoses are given to Liogonyleptoides inermis and L. tetracanthus. Two new species are described in Liogonyleptoides – Liogonyleptoides mirificus sp. nov. and Liogonyleptoides venatus sp. nov., from northern Espírito Santo. (3) Inhuma Piza, 1938, hitherto monotypic, is expanded to include another three species, each of which originally constituted a monotypic genus. Accordingly, the genera Schubartesia B. Soares, 1944a, Hugoesia H. Soares 1968 and Stefanesia Soares and Soares, 1988 all = Inhuma. (4) Metagonyleptes Roewer, 1913a is dismantled, the genus is newly considered a junior subjective synonym of Mischonyx Bertkau, 1880 (because the synonymy of the type species), while its species are transferred to varied genera, notably to Guascaleptes (herein revalidated from the synonymy with Geraeocormobius Holmberg, 1887) and to the undescribed genera SOD-01 and SOD-02. (5) Gonyleptes carinatus Sørensen, 1884 (type species of Metagonyleptes) = Gonyleptes scaber Kirby 1819 (currently Mischonyx scaber (Kirby 1819)). (6) Geogonys Mello-Leitão 1937, currently a synonym of Metagonyleptes, is newly transferred to the synonymy of Guascaleptes Mello-Leitão and de, 1933b to include some species formerly in Metagonyleptes. (7) Geogonys pallidipalpis Mello-Leitão and de, 1937, type species of Geogonys and currently a valid species of Metagonyleptes is newly synonymized with Metagonyleptes grandis Roewer, 1913b, both newly combined under Guascaleptes. (8) Gonyleptes saprophilus Mello-Leitão and de, 1922 and Sodreana curupira Pinto-da-Rocha and Bragagnolo, 2011 are newly transferred to Moreiranula Roewer, 1930, forming the new combinations Moreiranula saprophila and Moreiranula curupira. (9) Gonyleptes espiritosantensis B. Soares, 1944a = Proctobunoides tuberosus Mello-Leitão and de, 1944.
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The type species of Mischonyx Bertkau 1880, Mischonyx squalidus, was described based on a juvenile. The holotype is lost. Based on a revision of publications, the genus includes 12 species, all in Brazil. The objectives of this research are: to propose a phylogenetic hypothesis for Mischonyx based on Total Evidence (TE); propose taxonomic changes based on the phylogeny; and analyze the phylogenetic hypothesis biogeographically. Using the exemplar approach to taxon selection, we studied 54 specimens, 15 outgroups and 39 ingroup taxa using seven molecular markers (28S, 12S and 16S ribosomal genes, citochrome oxidase subunit I gene, carbamoyl-phosphate synthetase gene, internal transcribed spacer subunit 2 and histone H3 gene), totaling 3,742 bp, and 128 morphological characters. We analyzed the dataset under three optimality criteria: Maximum likelihood (ML), Maximum parsimony (MP) and Bayesian. We discuss the transformation of character states throughout the phylogeny, the different phylogenetic hypotheses using different datasets and the congruence of evidence between the clades obtained by the phylogenetic analysis and the biogeographical hypothesis for the Atlantic Forest areas of endemism. We estimate that Mischonyx clade diverged 50.53 Mya, and inside the genus there are two major clades. One of them cointains species from Paraná, Santa Catarina, South of São Paulo and Serra do Mar Areas of Endemism and the other has species from Espinhaço, Bocaina, South coast of Rio de Janeiro and Serra dos Órgãos Areas of Endemism. The first split inside these two clades occurred at 48.94 and 44.80 Mya, respectively. We describe three new species from Brazil: Mischonyx minimus sp. nov. (type locality: Petrópolis, Rio de Janeiro), Mischonyx intervalensis sp. nov. (type locality: Ribeirão Grande, São Paulo) and Mischonyx tinguaensis sp. nov (type locality: Nova Iguaçu, Rio de Janeiro). The genus Urodiabunus Mello-Leitão, 1935 is considered a junior synonym of Mischonyx. Weyhia spinifrons Mello-Leitão, 1923; Weyhia clavifemur Mello-Leitão, 1927 and Geraeocormobius reitzi Vasconcelos, 2005 were transferred to Mischonyx. Mischonyx cuspidatus (Roewer, 1913) is a junior synonym of M. squalidus Bertkau, 1880. In the results of the phylogenetic analyses, Gonyleptes antiquus Mello-Leitão, 1934 (former Mischonyx antiquus) does not belong in Mischonyx and its original combination is re-established. As it is now defined, Mischonyx comprises 17 species, with seven new combinations.
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No volume anterior destes Arquivos [Cf. Arquivos de Zoologia do Estado de São Paulo, 1948, 5 (9) : 553-635] demos inicio a publicação da Monografia dos gêneros de opiliões neotrópicos." Conforme ai justificamos, vimo-nos na contingencia de publica-la por partes, de acordo com o espago disponível na revista deste Departamento. Por mera conveniência pratica, resolvemos inicia-la com a família Gonyleptidae, a que contem os opiliões mais representativos da nossa fauna e, pela mesma razão, as subfamílias e os gêneros foram colocados em ordem alfabética.
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The macrosetae of the ventral plate of Laniatores penis have been occasionally described and used for systematic purposes in the literature, to the point of having a whole system created for them in the recent study Of Kury & Villarreal (2015), in a paper where a phylogenetic analysis of the subfamily Gonyleptoidea was performed. Contrastingly, the microsetae of ventral plate of Gonyleptoidea are left undescribed in descriptive works, where the penis is often illustrated without a ventral view of the ventral plate. In some works with SEM images microsetae do appear, although they remain undescribed and/or unacknowledged ( e.g. , Townsend et al. , 2010 figs 8–10; Coronato-Ribeiro & Pinto-da-Rocha 2015, where microsetae are visible in photos, but only barely mentioned). In the case of line drawings, exceptionally a few microsetae are shown mainly in the lateral view of the penis ventral plate (Acosta 2001, figs 17–21; Weber 1988, fig 6b). Exceptions to this treatment are Kury (2012: 41, figs 14–15) in which besides microsetae are illustrated their distribution pattern is described and Kury (2014), where even a subgroup of Gonyleptoidea was created and named—Microsetata—for species possessing a mat of microsetae on the ventral surface of ventral plate. Furthermore, among the characters used for the analysis in Kury & Villarreal (2015), the authors listed one containing five types of microsetae, which were not explained. Here, these types and their topology are described to make up for that omission.
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