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Total evidence phylogeny of the North American harvestman
family Stygnopsidae (Opiliones : Laniatores : Grassatores)
reveals hidden diversity
Jesús A. Cruz-López
A,B,C
and Oscar F. Francke
A
A
Colección Nacional de Arácnidos, Departamento de Zoología, Instituto de Biología,
Universidad Nacional Autónoma de México, Apartado Postal 70-153, Mexico City 04510, Mexico.
B
Posgrado en Ciencias Biológicas, Universidad Nacional Autónoma de México,
Avenida Universidad 3000, 04510 Coyoacán, Mexico City, Mexico.
C
Corresponding author. Email: thelyphonidito@gmail.com
Abstract. Systematic relationships among Laniatores have received considerable attention during the past few years.
Many significant taxonomic changes have been proposed, particularly in the superfamily Gonyleptoidea. As part of
this superfamily, the basalmost Stygnopsidae is the least known family. In order to propose the first total evidence
phylogeny of the family, we produced four datasets: three molecular markers –partial nuclear 28S, mitochondrial ribosomal
16S, mitochondrial protein-encoding cytochrome coxidase subunit I; and 72 morphological characters. With these data,
we performed three different phylogenetic analyses: (1) Bayesian Inference with molecular data, and (2) Bayesian
Inference and (3) Maximum Likelihood using combined data. Our results are congruent: a monophyletic Stygnopsidae
subdivided into two major clades: Stygnopsinae and Karosinae, subfam. nov. The following genera are redefined:
Stygnopsis,Hoplobunus and Serrobunus stat. rev. The following taxa are described: Iztlina venefica, gen. nov., sp. nov.
and Tonalteca, gen. nov. Additionally, the following changes are proposed: Serrobunus queretarius (Šilhavý, 1974), comb.
nov., Stygnopsis apoalensis (Goodnight & Goodnight, 1973), comb. nov., Stygnopsis mexicana (Roewer, 1915), comb.
nov., Stygnopsis oaxacensis (Goodnight & Goodnight, 1973), comb. nov., and Tonalteca spinooculorum (Goodnight
& Goodnight, 1973), comb. nov. We also discuss the status of the genera Isaeus stat. rev. and Mexotroglinus. Finally,
we discuss the evolution of male genitalia and convergence of selected homoplastic diagnostic characters.
Received30July2016,accepted12December2016,publishedonline.BZ
Introduction
The order Opiliones Sundevall, 1833, with ~6500 species, is the
third most diverse group in Arachnida, after Acari and Araneae
(Giribet and Sharma 2015). The group comprises conspicuous
arachnids from humid habitats in both tropical and temperate
zones, but mostly in the Neotropical region (Pinto-da-Rocha
et al.2014). Four major lineages are recognised in the order:
(1) the mite-like harvestmen Cyphophthalmi Simon, 1879; (2) the
daddy-longlegs Eupnoi Hansen and Sørensen, 1904; (3) the
thread-like palpi Dyspnoi Hansen and Sørensen, 1904; and
(4) the armoured harvestmen Laniatores Thorell, 1876 (Wolff
et al.2016). This last lineage is the most diverse, encompassing
more than two-thirds of the diversity in Opiliones (Sharma and
Giribet 2011).
During the last two decades, the systematics of Laniatores
has received special attention (Sharma and Giribet 2014; Giribet
and Sharma 2015). For example, 11 family-group taxa have been
described or re-assigned, many of these previously considered
as part of the ‘waste-basket’Phalangodidae Simon, 1879 under
Roewer’s criterion (revisited in part for selected taxa by Pinto-da-
Rocha et al.(2012), Bragagnolo et al.(2015) and Cruz-López
et al.(2016). These taxa have been erected or re-assigned based
on three general criteria: (1) the re-examination of male genitalia
as in Escadabiidae Kury and Pérez-González, 2003, Icaleptidae
Kury and Pérez-González, 2002 and Kimulidae Pérez-González
et al., 2007 (Kury and Pérez-González 2002; Kury and Pérez-
González in Kury 2003; Pérez-González and Kury 2007);
(2) morphology-based phylogenies as in Cranaidae Roewer,
1013, Cryptogeobiidae Kury, 2014, Globibuninae Kury, 2012,
Manaosbiidae Roewer, 1943 and Nomoclastidae Roewer, 1943
(Kury 2012,2014; Kury and Villarreal 2015); and (3) molecular-
based phylogenies as in Gerdesiidae Bragagnolo et al., 2014,
Metasarcidae Kury, 1994, Petrobunidae Sharma and Giribet,
2011, Pyramidopidae Sharma et al., 2011 and Tithaeidae
Sharma and Giribet, 2011 (Sharma and Giribet 2011; Sharma
et al.2011; Pinto-da-Rocha et al.2014; Bragagnolo et al.2015).
Among Laniatores, the superfamily Gonyleptoidea Sundevall,
1833 is one of the most controversial groups. Phylogenetically,
Journal compilation !CSIRO 2017 www.publish.csiro.au/journals/is
CSIRO PUBLISHING
Invertebrate Systematics, 2017, 31, 317–360
http://dx.doi.org/10.1071/IS16053
relationships have been open to discussion, depending on the data
source and taxon sampling, as in the status of Cranaidae/nae and
Manaosbiidae/nae. Using morphological evidence, these groups
have been considered to form a grade with Gonyleptidae and
preserved their familial status (Kury and Villarreal 2015).
Conversely, based on multi-locus phylogenetic research, these
two families are nested within Gonyleptidae, and are considered
subfamilies (Pinto-da-Rocha et al.2014).
Currently, both data sources, morphology and molecules,
support Stygnopsidae Sørensen, 1932 (although only a few
representatives of the group have been sampled) as sister-
group of the other families of Gonyleptoidea (Giribet et al.
2002,2010; Sharma and Giribet 2011; Kury and Villarreal
2015). However, the monophyly of the family has never been
tested using a strict phylogenetic approach, either with
morphological or molecular data, and the internal relationships
remain unclear. Complementary to the phylogenetic position of
Stygnopsidae, Kury (2003), Pérez-González (2006), Mendes and
Kury (2007) and Sharma et al.(2011) considered that the glans
penis formed by an exposed multifolded follis with small apical
spines could be a synapomorphic feature for the family, although
similar genital structures are present in Epedanidae Sørensen,
1886, Assamiidae Sørensen, 1884 and Pyramidopidae Sharma
et al., 2011.
The taxonomic history of Stygnopsidae is very complex, with
isolated descriptions of genera and species in different families
and many synonymies and transfers. Many of these taxonomic
problems were generated by Roewer (1912,1915) and Goodnight
Table 1. Taxa used in the phylogenetic analysis, including GenBank accession numbers
Taxa Family DNA voucher 28S 16S COI
Zalmoxida sp. Petrobunidae DNA104070 JF786583 JF786462 JF786435
Conomma oedipus Pyramidopidae DNA101051 GQ912801.2 GQ912853 GQ912882
Cynortula granulata Cosmetidae DNA100332 GQ912809.2 JF786464 –
Glysterus sp. Gonyleptidae, Ampycinae DNA101422 GQ912814.2 FJ796472 FJ796493
Zygopachylus sp. Nomoclastidae, Nomoclastinae DNA101425 GQ912818.2 GQ912855 GQ912889
Chapulobunus asper Stygnopsidae, Karosinae CNAN-DNA-0043 KY042066 ––
Chapulobunus poblano Stygnopsidae, Karosinae CNAN-DNA-0072 KY042067 ––
Chapulobunus psilocybe Stygnopsidae, Karosinae CNAN-DNA-0069 KY042068 KY042047 KY097807
Chapulobunus unispinosus Stygnopsidae, Karosinae CNAN-DNA-0042 KY042069 KY042048 KY097808
Chinquipellobunus aff. madlae Stygnopsidae, Stygnopsinae CNAN-DNA-0067 KY042070 ––
Chinquipellobunus aff. russelli Stygnopsidae, Stygnopsinae CNAN-DNA-0065 KY042071 KY042049 KY097809
Chinquipellobunus osorioi Stygnopsidae, Stygnopsinae CNAN-DNA-0068 KY042072 KY042050 KY097810
Crettaros valdezi Stygnopsidae, Karosinae CNAN-DNA-0073 KY042073 KY042051 KY097811
Hoplobunus barretti Stygnopsidae, Stygnopsinae CNAN-DNA-0090 KY042074 KY042052 KY097812
Hoplobunus sp. Stygnopsidae, Stygnopsinae CNAN-DNA-0002 KY042075 KY042053 KY097813
‘Hoplobunus’sp. Stygnopsidae, Stygnopsinae CNAN-DNA-0033 KY042076 KY042054 KY097814
‘Hoplobunus’aff. planus Stygnopsidae, Karosinae CNAN-DNA-0029 KY042082 –KY097820
‘Hoplobunus’zullinii Stygnopsidae, Stygnopsinae CNAN-DNA-0170 KY042065 KY042046 KY097806
Huasteca gratiosa Stygnopsidae, Karosinae CNAN-DNA-0020 KY042077 KY042055 KY097815
Huasteca rugosa Stygnopsidae, Karosinae CNAN-DNA-0004 KY042078 –KY097816
Huasteca sp. Stygnopsidae, Karosinae CNAN-DNA-0060 KY042079 KY042056 KY097817
Iztlina venefica Stygnopsidae, Stygnopsinae CNAN-DNA-0057 KY042080 KY042057 KY097818
Karos barbarikos Stygnopsidae, Karosinae CNAN-DNA-0031 KY042081 –KY097819
Mexotroglinus aff. sbordonii Stygnopsidae, Stygnopsinae CNAN-DNA-0051 KY042083 KY042058 KY097821
Mictlana inops Stygnopsidae, Karosinae CNAN-DNA-0063 KY042084 –KY097822
Paramitraceras aff. granulatum Stygnopsidae, Stygnopsinae CNAN-DNA-0009 KY042085 –KY097823
Paramitraceras aff. hispidulum Stygnopsidae, Stygnopsinae CNAN-DNA-0152 KY042086 –KY097824
Paramitraceras tzotzil Stygnopsidae, Stygnopsinae CNAN-DNA-0035 KY042087 –KY097825
Paramitraceras veracruz Stygnopsidae, Stygnopsinae CNAN-DNA-0007 KY042088 –KY097826
Philora mazateca Stygnopsidae, Stygnopsinae CNAN-DNA-0046 KY042089 ––
Philora tuxtlae Stygnopsidae, Stygnopsinae CNAN-DNA-0045 KY042090 –KY097827
Potosa sp. Stygnopsidae, Karosinae CNAN-DNA-0089 KY042091 KY042059 KY097828
Sbordonia aff. parvula Stygnopsidae, Stygnopsinae CNAN-DNA-0163 KY042092 KY042060 KY097829
Sbordonia sp. Stygnopsidae, Stygnopsinae CNAN-DNA-0055 KY042093 –KY097830
Serrobunus boneti Stygnopsidae, Stygnopsinae CNAN-DNA-0005 KY042094 KY042061 KY097831
Stygnopsis apoalensis Stygnopsidae, Stygnopsinae CNAN-DNA-0064 KY042095 KY042062 KY097832
Stygnopsis mexicana Stygnopsidae, Stygnopsinae CNAN-DNA-0071 KY042096 –KY097833
Stygnopsis oaxacensis Stygnopsidae, Stygnopsinae CNAN-DNA-0083 KY042097 –KY097834
Stygnopsis robusta Stygnopsidae, Stygnopsinae CNAN-DNA-0036 KY042098 KY042063 KY097835
Stygnopsis valida Stygnopsidae, Stygnopsinae CNAN-DNA-0044 KY042099 –KY097836
Tonalteca spinooculorum Stygnopsidae, Stygnopsinae CNAN-DNA-0084 KY042100 KY042064 KY097837
Troglostygnopsis sp. Stygnopsidae, Stygnopsinae CNAN-DNA-0049 KY042101 ––
Troglostygnopsis sp. Stygnopsidae, Stygnopsinae CNAN-DNA-0050 KY042102 –KY097838
318 Invertebrate Systematics J. A. Cruz-López and O. F. Francke
and Goodnight (1953). Sørensen (1932) described the family
Stygnopsidae as those Laniatores with distitarsus I with two
segments and without maxillary lobes on coxae II. Sørensen
(1932) included three genera in the family: Stygnopsis Sørensen,
1902, Tachus Sørensen, 1932 (currently Tibangara Mello-
Leitão, 1940 in Cryptogeobiidae Kury, 2014) and Isaeus
Zygopachylus sp. DNA101425
Mexotroglinus aff. sbordonii 0051
Stygnopsis robusta 0036
Chinquipellobunus aff. madlae 0067
Chinquipellobunus aff. russelli 0065
Zalmoxida sp. DNA104070
Crettaros valdezi 0073
Cynortula granulata DNA100332
Conomma oedipus DNA101051
Serrobunus boneti 0005
Stygnopsis oaxacensis 0083
Paramitraceras veracruz 0007
Paramitraceras tzotzil 0035
Mictlana inops 0063
Glysterus sp. DNA101422
Karos barbarikos 0031
Philora tuxtlae 0045
'Hoplobunus' zullinii 0170
Chapulobunus asper 0043
Huasteca gratiosa 0020
Huasteca rugosa 0004
'Hoplobunus' planus
Philora mazateca 0046
Torreana spinata DNA105839
Stygnopsis valida 0044
Stygnopsis mexicana 0071
Paramitraceras aff. hispidulum 0152
Hoplobunus sp. 0002
Chapulobunus poblano 0072
Paramitraceras aff. granulatum 0009
'Karos' depressus
Sbordonia sp. 0055
Troglostygnopsis sp. 0049
'Hoplobunus' sp. 0033
Tampiconus philippii
Hoplobunus barretti 0090
Potosa sp. 0089
0029
Sbordonia aff. parvula 0163
Troglostygnopsis sp. 0050
Chapulobunus psilocybe 0069
Stygnopsis apoalensis 0064
Huasteca sp. 0060
Chinquipellobunus osorioi 0068
Chapulobunus unispinosus 0042
Tonalteca spinooculorum 0084
Iztlina venefica 0057
0.56
0.63
1
0.95
0.68
0.93
1
0.73
1
0.87 1
0.95
1
0.92
1
0.91
1
0.99
0.87
0.89
1
0.72
1
1
1
0.9
0.99
1
0.82
0.87
0.93
1
0.56
0.86 0.62
0.87
1
1
0.98
1
1
1
Karosinae
Stygnopsinae
.
.
.
.
.
.
.
.
.
.
.
.
'Hoplobunus' aff. planus
Fig. 1. Phylogenetic relationships of the family Stygnopsidae based on Bayesian Inference (BI) using
combined data. Numbers on nodes correspond to posterior probabilities. Terminals in bold black correspond
to taxa with only morphology data available from females only.
Table 2. Molecular markers, primers sequences and original references
28S
28S D1F 50-GGG ACT ACC CCC TGA ATT TAA GCA T-30Park and Ó Foighil (2000)
28Srd4b 50-CCT TGG TCC GTG TTT CAA GAC-30Edgecombe and Giribet (2006)
28Sa 50-GAC CCG TCT TGA AAC ACG GA-30Whiting et al.(1997)
28Srd5b 50-CCA CAG CGC CAG TTC TGC TTA C-30Schwendinger and Giribet (2005)
28Srd4.8a 50-ACC TAT TCT CAA ACT TTA AAT GG-30Schwendinger and Giribet (2005)
28Srd7b1 50-GAC TTC CCT TAC CTA CAT-30Schwendinger and Giribet (2005)
16S
16Sa 50-CGC CTG TTT ATC AAA AAC AT-30Xiong and Kocher (1991)
16Sb 50-CTC CGG TTT GAA CTC AGA TCA-30Edgecombe et al.(2002)
COI
LCO1490 50-GGT CAA CAA ATC ATA AAG ATA TTG G-30Folmer et al.(1994)
HCOoutout 50-GTA AAT ATA TGR TGD GCT C-30Prendini et al.(2005)
Vf1d_t1 50-TGT AAA ACG ACG GCC AGT TCT CAA CCA ACC ACA ARG AYA TYG G-30Ivanova et al.(2007)
Vr1d_t1 50-CAG GAA ACA GCT ATG ACT AGA CTT CTG GGT GGC CRA ARA AYC A-30Ivanova et al.(2007)
Total evidence phylogeny of Stygnopsidae Invertebrate Systematics 319
Sørensen, 1932, a monotypic genus described in the same work. It
is remarkable that at the same time, other genera currently placed
in Stygnopsidae were allocated to Phalangodidae or Assamiidae
(Banks 1900; Sørensen 1902; Pickard-Cambridge 1905).
Mello-Leitão (1938) revised the familial classification of
Sørensen and adopted proposal of Roewer (1923). Mello-
Leitão (1938) considered Stygnopsinae Sørensen, 1932 as
subfamily within Stygnopsidae with only two genera, Stygnopsis
and Tachus, and erected the subfamily Isaeinae Mello-Leitão,
1938 for Isaeus, this last proposal without any justification.
Goodnight and Goodnight (1942,1944) described four
monotypic genera: Serrobunus Goodnight & Goodnight, 1942,
Chinquipellobunus Goodnight & Goodnight, 1944, Karos
Goodnight & Goodnight, 1944, and Monterella Goodnight &
Goodnight, 1944; all of them in Phalangodinae. Later, Goodnight
and Goodnight (1945,1946) transfered Chinquipellobunus,
Hoplobunus Banks, 1900 and Serrobunus to Stygnopsinae,
and described the monotypic genera Montabunus Goodnight
& Goodnight, 1945 in Phalangodinae and Chapulobunus
Goodnight & Goodnight, 1946 in Stygnopsinae. In subsequent
publications, Goodnight and Goodnight (1947b,1953,1954,
1967,1971,1973,1977) again ignored Stygnopsidae/nae, and
described more monotypic genera, Potosa Goodnight &
Goodnight, 1947 and Philora Goodnight & Goodnight, 1954,
in Phalangodinae. Among the several papers by the Goodnights,
Goodnight and Goodnight (1953) stands out, because the authors
made many unjustified synonymies in laniatorean genera, and
reduced almost all of the monotypic genera previously described
by them to only three genera: Hoplobunus,Karos and
Paramitraceras Pickard-Cambridge, 1905.
Šilhavý(1974) considered Stygnopsidae with two subfamilies:
Stygnopsinae with Hoplobunus and Stygnopsis, and a new
subfamily, Troglostygnopsinae Šilhavý, 1974, for the monotypic
Troglostygnopsis Šilhavý, 1974, a genus described in the same
work. The last action was justified by the high tarsal count
in legs I, with more than two segments in distitarsus I. It is
also remarkable that Šilhavý(1974) considered Karos and
Paramitraceras, placed at the time in Phalangodinae, with
presence of ‘lateral projections’on either side of scutum,
as ‘related’to Troglostygnopsis, which has similar lateral
projections.
On the basis of male genitalic morphology, Kury (1994,
1997,2003) and Kury and Cokendolpher (2000) rediagnosed
and restricted the family to Hoplobunus,Karos,Mexotroglinus
Šilhavý, 1977, Paramitraceras,Sbordonia Šilhavý, 1977,
Stygnopsis,Tampiconus Roewer, 1949 and Troglostygnopsis,
without any subfamilial groupings.
Recently,Cokendolpher(2004) revalidatedChinquipellobunus
from its synonymy under Hoplobunus on the basis of male
genitalia of selected North American stygnopsids. Cruz-López
(A)(B)(C)(D)
(E)(F)
Fig. 2. Dorsal habitus of Stygnopsis.(A)Stygnopsisvalida male. (B)Stygnopsis valida female. (C)Stygnopsis robusta male. (D)Stygnopsis robusta female. (E)
Stygnopsis oaxacensis male. (F)Stygnopsis oaxacensis female. Scale bars: A, B, C, D = 4.0 mm, E, F = 3.0 mm.
320 Invertebrate Systematics J. A. Cruz-López and O. F. Francke
and Francke (2013b) revised Philora and transferred this genus
from incertae sedis to Stygnopsidae. They also described a
unique male genital pattern present only in the Neotropical
stygnopsid genera Paramitraceras and Philora, the type
species of Troglostygnopsis, and presumably in Sbordonia.
Cruz-López and Francke (2015) performed a cladistic
analysis of the genus Karos using morphological data. They
found that the Goodnights’concept of Karos represented a
paraphyletic group, because Troglostygnopsis inops (Goodnight
and Goodnight 1971) is nested within it with high support.
They also proposed the revalidation of all genera currently in
synonymy with Karos, because all of them exhibit sufficient
synapomorphies and diagnostic characters to be considered valid
(Cruz-López and Francke 2015). In addition, Cruz-López and
Francke (2015) proposed two monophyletic genus-groups
within Stygnopsidae: the Paramitraceras-group, diagnosable
according to Cruz-López and Francke (2013b), and the Karos-
group, comprising those stygnopsids with armature on the
meso-apical and mesal surfaces of the pedipalpal femur and
patella, the ocularium located in the middle of the prosoma,
males and females without sexual dimorphism in dentition
and cheliceral size, and penis flattened apically, with lateral
macrosetae forming a longitudinal row.
Currently, Stygnopsidae is composed of 17 genera and 56
species. This family is distributed mainly in the Sierra Madre
Oriental in Mexico, inhabitating pine forest, pine-oak forest,
rainforest and/or caves. However, Chinquipellobunus occurs
in caves in Southern Texas, USA, and Paramitraceras extends
to tropical regions in Guatemala, Belize, El Salvador and
Honduras.
Here, we propose the first total evidence phylogenetic
hypothesis of Stygnopsidae relationships, based on two
mitochondrial markers, COI and 16S, the almost entire nuclear
28S, and 72 morphological characters, and sampling the largest
ever number of representative taxa of the family. Our results
support the recognition of two subfamilies in Stygnopsidae:
Stygnopsinae and Karosinae, subfam. nov. Additionally, we
discuss the evolution of male genitalia, the convergence of
some diagnostic morphological characters, the polyphyly of
the genera Hoplobunus and Paramitraceras, and the taxonomic
status of selected taxa, including the description of the
following taxa: Tonalteca, gen. nov. and Iztlina venefica, gen.
nov., sp. nov.
(A)
(B)
(C)
Fig. 3. Lateral habitus of Stygnopsis.(A)Stygnopsis valida male.
(B)Stygnopsis robusta male. (C)Stygnopsis oaxacensis male. Scale bars:
A, B, C = 2.00 mm.
(A)
(B)
(C)
(D)
(E)
(F)
Fig. 4. Chelicera of Stygnopsis males. (A) Frontal view of Stygnopsis valida.(B) Mesal view of Stygnopsis valida.(C) Frontal view of Stygnopsis robusta.
(D) Mesal view of Stygnopsis robusta.(E) Frontal view of Stygnopsis oaxacensis.(F) Mesal view of Stygnopsis oaxacensis. Arrows indicate the basal blunt
tooth. Scale bars: A = 2.0 mm, B, C = 1.5 mm.
Total evidence phylogeny of Stygnopsidae Invertebrate Systematics 321
Materials and methods
Morphological data and taxon sampling
Diverse material of described and undescribed Stygnopsidae
and additional Laniatores was examined from the following
collections and museums: American Museum of Natural
History, New York, USA (AMNH); the Natural History
Museum, London, UK (NHM); Colección Nacional de
Arácnidos, Mexico City, Mexico (CNAN); Laboratorio de
Acarología ‘Anita Hoffmann’,FacultaddeCiencias,
Universidad Autónoma de México (UNAM), Mexico City,
Mexico (LAAH); Museo Argentino de Ciencias Naturales,
Buenos Aires, Argentina (MACN); Naturmuseum Senckenberg
Sektion Arachnologie, Frankfurt, Germany (SMF); Texas
Memorial Museum, Texas, USA (TMM); and The Museum,
Texas Tech University, Texas, USA (TTU). Types of new taxa
described here and DNA vouchers of ingroup are deposited
in CNAN. For a complete list of material examined, see
Supplementary Materials 1 and 2 (available as Supplementary
material to this paper).
We attempted to include the greatest representation
of stygnopsid taxa in our analyses, but it was not possible to
obtain DNA from ‘Hoplobunus’planus Goodnight & Goodnight,
1973, ‘Karos’depressus Goodnight & Goodnight, 1971 and
Tampiconus philippi Roewer, 1949, species where only the
female types are known. Even so, we included them in the
combined analyses because they are problematic taxa.
The types of P. granulatum Pickard-Cambridge, 1905,
P. hispidulum Pickard-Cambridge, 1905 (both sexes), and
Sbordonia parvula (Goodnight & Goodnight, 1953) (female),
and an additional male of S. parvula (MACN) were examined,
but it was not possible to obtain fresh tissue for DNA extraction
for these species. However, DNA and morphological data of
three closely related species (aff.) were included in the analyses.
We selected six taxa from Sharma and Giribet (2011) as
outgroups (Table 1). The selection was based on the
availability of both morphological and molecular data of some
Gonyleptoidea, and of two more distant groups, Pyramidopidae
and Petrobunidae. Only one male of Conomma oedipus Roewer,
1949 (Pyramidopidae) was examined for morphology. With
respect to the remaining outgroups, the morphological data
were obtained as complete as possible from Goodnight and
Goodnight (1947a), Avram (1977) and Kury and Villarreal
(2015). The ingroup is formed by 41 species (Supplementary
Materials 1 and 2). For the most diverse genus, Hoplobunus, we
included eight of the 10 species known, plus two undescribed
species ‘similar’to the type species Hoplobunus barretti
Banks, 1900. We focused on this genus, because it is
currently the most diverse, although this diversity is an artefact
due to its complex taxonomic history based on some of the
Goodnight’s unjustified synonymies. This problem has been
corroborated for some other Mexican taxa, as reported by
Cruz-López and Francke (2013a,2015) in Paramitraceras
and Karos, respectively.
To collate the morphological data, the specimens were
examined under a Nikon SMZ 625 microscope. Colour
photographs were taken with Nikon Coolpix S10 camera with
adaptor for the microscope. Electronic photographs were
taken using two differents SEMs: Hitachi S-2460N and
Hitachi SU1510, both in the Instituto de Biología, UNAM,
Mexico. All figures were edited using Adobe Illustrator CC
and PhotoShop CS5. We coded 72 morphological characters,
some of which were taken and modified from Kury and
Villarreal (2015) and Cruz-López and Francke (2015). To see
(A)
(B)
(C)
Fig. 5. Mesal view of pedipalps of Stygnopsis males. (A)Stygnopsis robusta.(B)Stygnopsis valida.(C)Stygnopsis oaxacensis. Scale bars: A = 2.5 mm,
B = 2.0 mm, C = 1.5 mm.
322 Invertebrate Systematics J. A. Cruz-López and O. F. Francke
the list and description of all characters and how these were coded,
see Supplementary Materials 3 and 4.
Taxonomic nomenclature
For the description of morphological characters and new taxa, we
adopted different nomenclatural sources, including the scutum
shape from Kury and Medrano (2016), the morphology of the
chelicera, pedipalp and ornamentation of dorsum and legs from
Cruz-López and Francke (2015,2016a), and the pedipalpal
armature from Acosta et al.(2007). To describe the male
genitalia, we adopted the penial setation nomenclature recently
proposed by Kury and Villarreal (2015), with modifications as
treated in Cruz-López and Francke (2016a,2016b), and
additional modifications proposed here. For morphology of the
capsula externa and interna, we follow Pérez-González (2006).
Molecular data
All outgroups sequences were obtained from GenBank
(Table 1). Total DNA of the ingroup was extracted from legs
or complete juveniles using DNeasy tissue kit (Qiagen, Mexico
City, Mexico) following supplier procedure with modifications
proposed by Boyer et al.(2005). Total DNA was used as a
template for amplification of partial nuclear ribosomal 28S
obtained from assembly of three fragments, the mitochondrial
ribosomal 16S and the mitochondrial protein-encoding
cytochrome coxidase subunit I (COI). Primer sequences and
their original references are indicated in Table 2.
PCR reactions were carried out in 12.5 mL volumes, consisting
of 0.2 mM each primer, 0.8 mM deoxynucleotides, 1 !PCR
buffer, 4 mM MgCl
2
, 1.25 U of Taq DNA polymerase
(TaKara LA Taq, CA, USA) and 1–2mL of DNA template for
COI and 16S. Similarly, we carried out PCR reactions using
Green Master Mix Polymerase (WI, USA) for 28S, with the
following mix: 0.5 !of the mix, 0.2 mM each primer and 1–2mL
of DNA template. For all markers, the cycles were according
to Sharma and Giribet (2009), except annealing temperature of
46"C for COI. For the primer pairs Vf1d_t1-Vr1d_t1, the cycles
were: 5 initial cycles of 94"C!2 min, 94"!30 s, 50"!40 s,
72"!1 min; plus 30 cycles of 94"!30 s, 55"!40 s, 72"!1 min,
with a final extention of 72"!5 min. PCR products were
visualised by agarose gel electrophoresis (1.5% agarose).
Unpurified products were sequenced in the Instituto de
Biología (UNAM), Mexico City, and by High Throughput
Sequencing (htSEQ), in the University of Washington, Seattle,
USA. Chromatograms and sequences were viewed and edited
in Geneious ver. 9.1.2 (Kearse et al.2012) and BioEdit ver.
7.2.5 (Hall 1999). The fragments of 28S were assembled in
Geneious. Finally, we obtained 99 sequences of the ingroup
(Table 1). To see the complete collection data of vouchers,
see Supplementary Material 2.
The 16S and COI markers were aligned using MAFFT (Katoh
2013), with default parameters through the interface Mesquite
ver. 3.0.4 (Maddison and Maddison 2015). The 28S marker
was aligned using the L-INS-i algorithms implemented in
(A)
(B)
(C)
(D)
(E)
(F)
Fig. 7. Retrolateral views of legs III and IV of Stygnopsis males. (A) Leg III of Stygnopsis robusta.(B) Leg IV of Stygnopsis robusta.(C) Leg III of
Stygnopsis valida.(D)Leg IV of Stygnopsis valida.(E)Leg IIIof Stygnopsis oaxacensis.(F)Leg IV of Stygnopsis oaxacensis. Scale bars: A = 5.0 mm, B = 4.5 mm,
C, D, F = 3.00 mm, E = 2.5 mm.
(C)
(B)
(A)
Fig. 6. Dorsal view of pedipalp patella of Stygnopsis males. (A)Stygnopsis
valida.(B)Stygnopsis robusta.(C)Stygnopsis oaxacensis. Scale bars: A,
B = 0.5 mm, C = 0.25 mm.
Total evidence phylogeny of Stygnopsidae Invertebrate Systematics 323
(A)(B)(C)
(D)(E)(F)
Fig. 8. Male genitalia of Stygnopsis.(A) Lateral view of Stygnopsis robusta.(B) Ventral view of Stygnopsis robusta.(C) Dorsal view of Stygnopsis
oaxacensis.(D)Dorsal view of Stygnopsis valida.(E)Dorsal view of Stygnopsis robusta.(F)Ventral view of Stygnopsis valida. Small letters A, B, C
and D indicate setal groups. Scale bars: A, B, C, F =250 mm, D, E = 200 mm.
324 Invertebrate Systematics J. A. Cruz-López and O. F. Francke
(A)
(B)
(C)
(D)
Fig. 9. Habitus of Hoplobunus.(A) Dorsal view of Hoplobunus barretti male. (B) Dorsal view of Hoplobunus sp. 0002 male. (C) Dorsal view of
Hoplobunus sp. 0002 female. (D) Lateral view of Hoplobunus barretti male. Scale bars: A = 1.5 mm, B, C = 2.0 mm, D = 0.5 mm.
(A)(B)
(C)
Fig. 10. Pedipalp of Hoplobunus barretti male. (A) Mesal view. (B) Ectal
view. (C) Frontal view. Scale bars: A, B = 1.0 mm, C = 0.7 mm.
(A)
(B)
Fig. 11. Chelicera of Hoplobunus barretti male. (A) Frontal view. (B) Mesal
view. Arrow indicates the basal blunt tooth. Scale bars: A, B =0.7 mm.
Total evidence phylogeny of Stygnopsidae Invertebrate Systematics 325
MAFFT. Aligned sequences were concatenated and file types
modified in Mesquite. Three different matrices were generated:
(1) molecular Nexus file with 44 terminal taxa; (2) total evidence
Nexus file with 47 terminal taxa; and (3) total evidence Phylip
file with 47 terminal taxa (Supplementary Materials 5–7). These
matrices were partitioned for each molecular marker and for
morphological data, respectively.
To select the evolutionary model for each molecular
partition, we used jModelTest ver. 2.1.6 (Darriba et al.2012).
We compared the Akaike Information Criterion (AIC) and the
Bayesian Information Criterion (BIC) to choose the best model
for each molecular marker. The evolutionary models chosen
were: general time reversible (GTR) + proportion of invariable
sites (I) + gamma distribution (G) for both 28S and COI sets,
and Hasegawa–Kishino–Yano (HKY) + G for the 16S set. For the
total evidence matrices, the model applied to the morphological
dataset was MkV (Markov model for variable characters)
(Lewis 2001), with Lewis correction for Maximum Likelihood
analysis (Lewis 2001), as previously used in other studies (e.g.
Monjaraz-Ruedas et al.2016). The models were specified for
the MrBayes block in both molecular and total evidence
matrices, and a partition.txt file and in morphological specific
command line for Maximum Likelihood analysis.
Phylogenetic methods
Phylogenetic trees were reconstructed using Bayesian Inference
(BI) and Maximum Likelihood (ML). BI analyses on molecular
(A)
(B)(C)
(D)
(E)
(F)
(G)(H)
Fig. 12. Legs III and IV of Hoplobunus barretti male. (A) Leg III prolateral view. (B) Leg IV prolateral view. (C) Leg IV retrolateral view. (D) Leg III retrolateral
view. (E) Femur IV prolateral view. (F) Femur IV retrolateral view. (G) Femur III ventral view. (H) Femur IV ventral view. Scale bars: A, D, G =1.0 mm, B,
C = 1.1 mm, E, F, H = 1.5 mm.
326 Invertebrate Systematics J. A. Cruz-López and O. F. Francke
and total evidence matrices were conducted with MrBayes
ver. 3.2.6 on XSEDE (Ronquist and Huelsenbeck 2003)
through the online portal CIPRES gateway (Miller et al.
2010). The ML analysis for total evidence was conducted
using RaxML HPC-AVX ver. 8.2.7 (Stamatakis 2014) on a
Dell XPS 8900 desktop computer. The BI analyses consisted
of two simultaneous runs each, with four chains defaults for
10 000 000 generations sampling every 2000 trees. The initial
25% of sampled trees were discarded as burn-in. Stationarity
parameters were viewed in TRACER ver. 1.6 (Rambaut et al.
2014). The ML analysis consisted of 1000 bootstrap replicates,
with 500 hits to find the tree with the best value of ML.
Results
Phylogenetic analyses
We obtained similar topologies in the three matrices analysed
under BI and ML approaches, with high support values for major
clades (Fig. 1, Figs S1 and S2). In all analyses, Stygnopsidae
was recovered as monophyletic, as sister-group of C. oedipus
(Pyramidopidae). Gonyleptoidea was not recovered as a clade,
which could be due to the use of a different subset of molecular
markers than those used to generate previous laniatorean
phylogenies (see Sharma and Giribet 2011; Cruz-López et al.
2016). However, the support values for Stygnopsidae in both
BI analyses are high, and are considerably higher in the ML
analysis.
Phylogenetic relationships among Stygnopsidae
The BI analysis with the molecular data (Fig. S1) generated
high support values for the family, subfamilies and some
genera. This analysis produced the following major monophyletic
relationships: (Karosinae (Paramitraceras-group (Stygnopsinae +
P. aff. hispidulum))). The BI analysis using combined data
obtained similar general topology to that for molecular data
(A)
(B)
(C)
Fig. 13. Scutum, leg IV and pedipalp of Hoplobunus barretti male. (A) Dorsal view of scutum. (B) Prolateral view of leg IV. (C) Mesal view of pedipalp.
Scale bars: A = 1 cm, B, C = 0.5 cm.
Total evidence phylogeny of Stygnopsidae Invertebrate Systematics 327
(A)(B)(C)
(D)(E)
Fig. 14. Male genitalia of Hoplobunus barretti.(A) Dorsal view. (B) Lateral view. (C) Ventral view. (D) Dorsal view. (E) Lateral view.
Small letters A, B, C, D and E indicate setal groups Scale bars: A, B, C = 200 mm, D, E = 50 mm.
328 Invertebrate Systematics J. A. Cruz-López and O. F. Francke
only, but with the Paramitraceras-group nested within
Stygnopsinae (Fig. 1). Also, the BI analysis of combined data
generated support values slighly lower than the BI analysis
of molecular data, which may be caused by the high number
of homoplastic morphological characters. The ML analysis of
combined data yielded considerable support values for some
clades (Fig. S2).
Both BI and ML analyses using combined data nested
the Paramitraceras-group in Stygnopsinae, with the surprising
inclusion of Serrobunus as sister-group of the remaining
members of the Paramitraceras-group. Another surprising
result is the exclusion of Paramitraceras aff. hispidulum from
Paramitraceras, but forming the clade (P. aff. hispidulum +
‘Hoplobunus’zullinii), with high support values in the three
analyses. Despite the similar external appareance in the known
Paramitraceras species (Fig. 35), the presence of ventral
glandular tubercles on males (Figs 38,39), and the unarmed
pedipalp (Figs 44,45), the analyses did not recover a
monophyletic Paramitraceras. However, the male genitalia of
P. hispidulum and of P. aff. hispidulum (Fig. 41A–C) do not
share the genital pattern present in the remaining Paramitraceras
(A)
(B)
(C)
Fig. 15. Habitus of ‘Hoplobunus’sp. 0033. (A) Dorsal view of male.
(B) Dorsal view of female. (C) Lateral view of male. Scale bars: A, B =
2.0 mm, C = 1.0 mm.
(A)
(B)
Fig. 16. Chelicera of ‘Hoplobunus’sp. 0033 male. (A) Frontal view.
(B) Mesal view. Scale bars: A, B = 1.0mm.
(A)
(B)
Fig. 17. Pedipalp of ‘Hoplobunus’sp. 0033 male. (A) Mesal view. (B) Ectal
view. Scale bars: A, B = 0.9mm.
Total evidence phylogeny of Stygnopsidae Invertebrate Systematics 329
and, unfortunately, the male genitalia of ‘Hoplobunus’zullinii
Šilhavý, 1977 remains unknown.
In all three analyses, we recovered Hoplobunus as a
polyphyletic group. It is remarkable that only the type species,
H. barreti and Hoplobunus sp. 0002 form a clade, sharing
uniform external and male genital morphology (Figs 9–14).
The voucher ‘Hoplobunus’sp. 0033 was recovered as sister-
group of T. philippi (with high support values) in both BI and
ML analyses using combined data. These specimens plus
an undescribed Tampiconus-like taxon are slightly similar to
Hoplobunus, with the following differences: (1) ventral
armature of pedipalpal femur formed by cylindrical, strong
almost fused setiferous tubercles in Hoplobunus, and formed
by scattered spiniform setiferous tubercles in Tampiconus-
like clade (Figs 10,17); (2) mesotergal area III unarmed in
Hoplobunus, armed with two small paramedian spines in
Tampiconus-like clade (Figs 9,15); (3) pedipalp patella with
dorsal crest formed by strong setiferous tubercles ending in a
prominent tubercle in Hoplobunus, absent in Tampiconus-like
clade (Figs 10,17); (4) two ventral rows of rounded tubercles,
increasing in size apically in legs III and IV in Hoplobunus, legs
III and IV without remarkable rows of tubercles in Tampiconus-
like clade (Figs 12,18); (5) presence of E setae on penis in
Hoplobunus and absence of these setae in Tampiconus-like clade
(Figs 14,19); and (6) presence of basal blunt tooth on movable
cheliceral finger in Hoplobunus and absence of this tooth in
Tampiconus-like clade (Figs 11,16). Also, Hoplobunus is found
in pine-forest at relatively high elevations in central and Northern
Mexico, and members of the Tampiconus-like clade are found in
lowland rain-forest in Eastern Mexico. However, without
available males of T. philippii, the type species of this genus,
it is difficult to determine the true identity of this genus and its
related taxa. The‘not-true’Hoplobunus are discussed in the
taxonomic treatments below.
Taxonomy
Family STYGNOPSIDAE Sørensen
Phalangodidae (in part): Roewer, 1923: 69; Mello-Leitão, 1938: 137;
Goodnight & Goodnight, 1942: 1; Goodnight & Goodnight, 1944: 1.
Stygnopsidae Sørensen, 1932: 272; Šilhavý,1974: 176; Šilhavý,1977:
220; Rambla & Juberthie, 1994: 218.
Included subfamilies: Stygnopsinae Sørensen, 1932, and Karosinae,
subfam. nov.
Emended diagnosis
Harvestmen with scutum type zeta (z) (Figs 2,9,15,20,24,28),
with the mid-bulge tending to be almost absent or type gamma
(g) in several taxa (Figs 35A,C,46A). Shape and size of ocularium
varying over a wide range, from small, almost absent, to large,
conical, and armed with long median spine. Chelicera ranging
(A)
(B)(C)
Fig. 18. Legs III and IV of ‘Hoplobunus’sp. 0033 male. (A) Prolateral view of leg III. (B) Ventral view of femur III. (C) Ventral view of femur IV. Scale bars:
A = 1.5 mm, B = 1.0 mm, C = 0.7 mm.
330 Invertebrate Systematics J. A. Cruz-López and O. F. Francke
(A)(B)(C)(C)
(D)(E)
Fig. 19. Male genitalia of ‘Hoplobunus’sp. 0033. (A) Frontal view. (B) Lateral view. (C) Detail of follis in lateral view. (D) Detail of
follis in dorsal view. (E) Detail of setae D and C, lateral view. Small letters A, B, C, and D indicate setal groups. Scale bars: A,
B = 150 mm, C = 50 mm, D, E = 25 mm.
Total evidence phylogeny of Stygnopsidae Invertebrate Systematics 331
(A)
(B)
(C)
Fig. 20. Habitus of Serrobunus boneti.(A) Dorsal view of male. (B) Dorsal view of female. (C) Lateral view of male. Scale bars: A, B = 1.5 mm,
C = 1.0 mm.
332 Invertebrate Systematics J. A. Cruz-López and O. F. Francke
from small-sized in Karosinae and Mexotroglinus, to slightly
swollen in Paramitraceras, and large in Hoplobunus and
Stygnopsis. Cheliceral dentition formed by many similar-sized
teeth in non-sexually-dimorphic species (Fig. 32); prominent
basal and/or medial teeth on movable finger in sexually
dimorphic species (Figs 4,43). Pedipalps armed with
setiferous tubercles, unarmed or with only spiniform setae
in Paramitraceras and Sbordonia (Figs 44,45). Sexual
dimorphism on legs occurs in many different ways within the
family: presence of glandular opening in males (Potosa); males
with legs longest (Karos); modified articles on males, possibly
with glandular pores (Huasteca Cruz-López & Francke, 2015,
Mexotroglinus, Fig. 46C,G); male legs strongly armed
(Chapulobunus,Chinquipellobunus,Hoplobunus,Sbordonia,
Stygnopsis); or without sexual dimorphism (Crettaros Cruz-
López & Francke, 2015, Mictlana Cruz-López & Francke,
2015, Montabunus,Monterella,Paramitraceras,Philora
and Troglostygnopsis). Calcaneous covering apical portion of
metatarsus. Tarsi without scopulae or pseudonychium, claws
non-pectinate. Penis without well defined ventral plate, truncus
generally cylindrical, ventral portion elongated, forming a
flimsy lamina (Figs 8,14,19,23,27,31,34,40,41,47).
Three major penial macrosetal patterns are found in the
family: (1) Paramitraceras-pattern: with several macrosetae
C forming a row along lateral margins; macrosetal complex
A+B latero-basally to follis, sometimes extending to ventral
portion; two ventral pairs of setae E, the inner micro and the
external macro; 1–3 pairs of microsetae D near lateral margins
(Fig. 40); (2) Karos-pattern: setal complex A+B+C forming
longitudinal rows along the lateral margins, sometimes setae
B can be recognised by small size; ventral microsetae E in
two pairs or forming two longitudinal rows of several setae;
2–4 pairs of macrosetae D near to base of follis (Fig. 34); and
(3) Stygnopsis-pattern: macrosetae (micro in Stygnopsis) setal
groups A, B and C recognisable and separated, each formed by
two or three pairs; one pair of small setae E, absent or only
vestigial socket; two pairs of small or microsetae D (Figs 8,14,
19,31). Capsula externa formed by an exposed and rigid multi-
folded follis, apical portion covered by small spines (except in
Stygnopsis). Capsula interna formed by internal stylus, slightly
evertible, cylindrical, with small apical bristles (Figs 14E,23,
27,34,40).
Subfamily STYGNOPSINAE Sørensen
Phalangodinae (in part): Roewer, 1912: 108; Roewer, 1923: 69;
Goodnight & Goodnight, 1942: 1; Goodnight & Goodnight, 1944: 1.
Isaeinae (in part): Mello-Leitão, 1938: 137.
Troglostygnopsinae (in part): Šilhavý,1974: 182.
Stygnopsinae: Mello-Leitão, 1938: 137; Šilhavý,1974: 176; Šilhavý,
1977: 220; Rambla & Juberthie, 1994: 218.
Included taxa. Hoplobunus Banks, 1900, Stygnopsis Sørensen, 1902,
Paramitraceras Pickard-Cambridge, 1905, Serrobunus Goodnight &
Goodnight, 1942 stat. rev., Chinquipellobunus Goodnight &
(A)
(B)
Fig. 21. Chelicera of Serrobunus boneti male. (A) Frontal view. (B) Mesal
view. Arrow indicates the basal blunt tooth. Scale bars: A, B = 1.5 mm.
(A)(B)
(C)(D)(E)(F)
Fig. 22. Leg IV and pedipalp of Serrobunus boneti male. (A) Retrolateral
view of leg IV. (B) Mesal view of pedipalp. (C) Retrolateral view of femur
IV. (D) Dorsal view of femur IV. (E) Prolateral view of femur IV. (F) Ventral
view of femur IV. Arrows indicate tubercles of ventral retrolateral row. Scale
bars: A, C, D, E, F = 3.0mm, B =1.5 mm.
Total evidence phylogeny of Stygnopsidae Invertebrate Systematics 333
(A)(B)(C)
(D)(E)(F)
Fig. 23. Male genitalia of Serrobunus.(A) Dorsal view of Serrobunus boneti.(B) Lateral view of Serrobunus boneti.(C) Ventral view of
Serrobunus boneti.(D)Dorsal view of Serrobunus queretarius.(E)Lateral view of Serrobunus queretarius.(F)Ventralview of Serrobunus queretarius.
Small letters A+B, C, and D indicate setal groups. A, B, C, D, F= 100 mm, E = 150 mm.
334 Invertebrate Systematics J. A. Cruz-López and O. F. Francke
(A)(B)
(C)
Fig. 24. Habitus of Iztlina venefica.(A)Dorsal view of male holotype. (B) Dorsal view of female paratype. (C) Lateral view of male holotype.
Scale bars: A, B, C = 2.0mm.
Total evidence phylogeny of Stygnopsidae Invertebrate Systematics 335
Goodnight, 1944, Tampiconus Roewer, 1949, Philora Goodnight &
Goodnight, 1954, Troglostygnopsis Šilhavý, 1974, Mexotroglinus
Šilhavý, 1977, Sbordonia Šilhavý, 1977, Iztlina, gen. nov.,
Tonalteca, gen. nov., and the uncertain ‘Hoplobunus’zullinii
Šilhavý, 1977.
Type genus:Stygnopsis Sørensen, 1902.
Emended diagnosis
Large and medium-sized harvestmen (scutum length ~6 or
>8 mm), except the small Mexotroglinus,Philora and some
Sbordonia (<3 mm). Ocularium at frontal margin of prosoma
(Figs 3,9,15,20,35D–H,46A), medium-sized and rounded
in Chinquipellobunus and Mexotroglinus, very large, usually
with an apical spine in the remaining members. Mesotergal
sulci straight (Figs 13A,35D–H,46A,E). Chelicera usually
large, movable finger with prominent basal teeth (Figs 4,11,
21); usually with sexual dimorphism in cheliceral size and
dentition. Mexotroglinus exhibits Karosinae-type chelicera
(Fig. 46D). Pedipalpal femur and patella without mesal
setiferous tubercles, except in Mexotroglinus. Penis with setal
patterns Stygnopsis,Paramitraceras and an unrecognisable
pattern (Figs 8,14,31,40). Pars distalis forming an angle of
40"or more with respect to flimsy ventral lamina (Figs 14E,
31B). Mexotroglinus is the only member of the subfamily that
exhibits many convergences with Karosinae in male genitalia,
such as: apical projection of pars distalis contiguous with
truncus; macrosetae D near to follis and similar size of
macrosetae A+B+C complex; and setal complex A+B+C forming
a longitudinal row (Fig. 47). Iztlina, gen. nov., exhibits a unique
and unrecognisable setal pattern within the subfamily, which
is described below.
Remarks
Because the polyphyletic Hoplobunus has created much
confusion in some taxonomic determinations, we put special
emphasis on the taxonomy of selected stygnopsine taxa
previously considered as Hoplobunus according to Goodnight
and Goodnight (1953): the type Stygnopsis,Hoplobunus,
Serrobunus stat. rev. and the new genera Tonalteca, gen. nov.
and Iztlina, gen. nov.
Genus Stygnopsis Sørensen
Stygnus (in part): Sørensen, 1884: 644.
Stygnopsis Sørensen, 1902: 4; Roewer, 1912: 153; Roewer, 1923: 116;
Mello-Leitão, 1926: 329; Roewer, 1927: 272; Sørensen, 1932: 273
(=Haehnelia Roewer 1915); Šilhavý,1974: 176; Šilhavý,1977: 220.
Included taxa. Stygnopsis valida (Sørensen, 1884), Stygnopsis mexicana
(Roewer, 1915), comb. nov., Stygnopsis robusta (Goodnight &
Goodnight, 1971), Stygnopsis apoalensis (Goodnight & Goodnight,
1973), comb. nov., Stygnopsis oaxacensis (Goodnight & Goodnight,
1973), comb. nov.
Type species:Stygnus validus Sørensen, 1884, by subsequent designation.
Emended diagnosis
Large harvestmen (scutum length >6 mm). Ocularium high,
usually with strong median spine. Chelicera large and sexually
dimorphic in amales, but there are bmales with chelicera
similar to those of females. Cheliceral dentition heterogeneous,
basal tooth on movable finger blunt (Fig. 4). Pedipalpal patella
smooth, roundly swollen apically, with small mesal apophysis
(Figs 5,6). Femora to tibiae III and IV armed with two ventral
rows of spiniform tubercles, increasing in size apically (Fig. 7).
Penis setal Stygnopsis-pattern, with all setae of penis are
microsetae. Three to four pairs of setae C, two pairs of setae
A and B, and two or three pairs of setae D, the last on dorsal
margin (Fig. 8). Pars distalis with apical depression, follis
inserted inside it. Latero-apical apices of pars distalis rolled
ventrally, pointed, ventral margin concave (Fig. 8). Follis
without small apical spines.
Remarks
Sørensen(1932) synonymised the monotypic Haehnelia Roewer,
1915 under Stygnopsis, forming the new combination Stygnopsis
mexicana (Roewer, 1915). However, Goodnight and Goodnight
(1953) ignored this synonymy, and considered Haehnelia a
synonym of Hoplobunus.
Šilhavý(1974) was the first to rediagnose the genus and
illustrated for the first time the male genitalia of Stygnopsis
valida (Sørensen, 1884) and Stygnopsis robusta (formerly
Hoplobunus robustus Goodnight & Goodnight, 1971). Šilhavý
mentioned that the main character to differentiate between
Hoplobunus and Stygnopsis is the presence of one pair of
paramedian long spines in mesotergal area III in the latter
(Fig. 3A,B). These spines are present in S. valida,S. mexicana
and S. robusta. It is remarkable that Šilhavýdid not mention
anything about Haehnelia mexicana, a species redescribed as
Hoplobunus mexicanus by Goodnight and Goodnight (1971),
with well developed dorsal spines (Goodnight and Goodnight
1971:fig. 20). Additionally, Goodnight and Goodnight (1971)
reported several additional records for H. mexicanus; however,
the record labelled as: ‘Cueva Arriba del Presidente, 1½km N
(A)
(B)
Fig. 25. Chelicera of Iztlina venefica male paratype. (A) Frontal view.
(B) Mesal view. Arrow indicates the basal blunt tooth. Scale bars: A,
B = 1.0 mm.
336 Invertebrate Systematics J. A. Cruz-López and O. F. Francke
(A)(B)
(C)(D)(E)
Fig. 26. Ocularium, pedipalp, legs III and IV, and posterior habitus of Iztlina venefica.(A) Ocularium frontal view of male paratype. (B) Posterior
view of habitus of holotype. (C) Ventral view of pedipalp tibia and tarsus. (D) Ventral view of femur III. (E) Ventral view of femur IV. Scale bars:
A = 1.5 mm, B = 2.0 mm, C, D = 1.0 mm, E = 0.5 mm.
Total evidence phylogeny of Stygnopsidae Invertebrate Systematics 337
of Huautla, Oaxaca, August 12, 1967 (one female), collected
by J. Reddell and J. Fish’corresponds to an undescribed species.
Like Sørensen (1932), we consider that S. mexicana could
be a junior synonym of S. valida, but with incomplete locality
data for the types, and the great similarities among male genitalia
in the five known species of the genus, it is impossible to
corroborate the proposed synonymy at this time. Maybe further
molecular studies from specimens from a range of localities
would help clarify this situation.
Genus Hoplobunus Banks
Hoplobunus Banks, 1900: 200; Pickard-Cambridge, 1905: 585; Roewer,
1912: 149; Roewer, 1923: 112; Roewer, 1927: 272; Goodnight &
Goodnight, 1942: 1; Goodnight & Goodnight, 1945: 3; Goodnight &
Goodnight, 1953: 19; Goodnight & Goodnight, 1967: 1, Goodnight
& Goodnight, 1971: 38; Goodnight & Goodnight, 1973: 86; Šilhavý,
1974: 176; Šilhavý,1977: 220; Edgar, 1990: 548; Rambla & Juberthie,
1994: 218.
Included taxa:Hoplobunus barretti Banks, 1900, by monotypy.
Emended diagnosis
Medium-size harvestmen (scutum length ~5 mm). Ocularium
very high, conical, with small apical spine (Fig. 9). Posterior
margin of scutum wider than mid-bulge width (Fig. 9). Chelicera
large and sexually dimorphic. Cheliceral dentition heterogeneous,
basal tooth on movable finger blunt (Fig. 11). Pedipalpal femur
compressed laterally, with two dorsal longitudinal rows of
spiniform tubercles, ending in a spiniform apical apophysis
(Figs 10,13C). Pedipalpal patella with dorsal crest of
contiguous, procurved spiniform tubercles (Figs 10,13C).
Sockets of pedipalpal tibia, tarsus and ventral femur
cylindrical, very wide and contiguous (Figs 10,13C). Femora
to tibiae III and IV armed with two ventral rows of spiniform
tubercles, increasing in size apically and ending in spiniform
apophysis (Figs 12,13B). Trochanter III globose, basal-most
portion of femur III dorsally curved abruptly (Fig. 12E,F). Penis
setal patterns Stygnopsis-pattern, two pairs of small setae D
on dorsal projection of truncus, latero-dorsal to follis base
(Fig. 14D), one pair of microsetae E, over setal group C
(Fig. 14B,E), setal groups A, B and C formed by two or three
macrosetae (Fig. 14A–C). Pars distalis compressed laterally in
almost one-half the length of penis, slightly curved dorsally
(Fig. 14A–C). Surface of follis rugose (Fig. 14E).
Remarks
Since the description of the genus and type species by
Banks (1900), the diagnosis of the genus was revised only
once by Goodnight and Goodnight (1953). They proposed a
brief rediagnosis based on ambiguous characters, the reason
they synonymised Haehnelia,Isaeus,Serrobunus and
Chinquipellobunus under Hoplobunus. Recently, Cokendolpher
(2004) revalidated Chinquipellobunus from its synonymy
under Hoplobunus on the basis of the male genitalia. He also
transferred Hoplobunus madlae Goodnight & Goodnight, 1967
and Hoplobunus russelli Goodnight & Goodnight, 1967 to
Chinquipellobunus. Before this study, Hoplobunus included
10 species under the diagnosis of Goodnight and Goodnight
(1953). It is remarkable that after the phylogenetic analyses
based on combined data, the genus now becomes monotypic
(but some undescribed species are known). As mentioned earlier,
three species were transferred to Stygnopsis. Hoplobunus
spinooculorum Goodnight & Goodnight, 1973 is here
transferred to Tonalteca, gen. nov., H. boneti is restored as a
valid species of Serrobunus, and Hoplobunus queretarius
Šilhavý, 1974 is transferred to Serrobunus. Finally, ‘H.’
(A)(B)(C)
Fig. 27. Male genitalia of Iztlina venefica male paratype. (A) Dorsal view. (B) Lateral view. (C) Ventral view. Small letters A, C, and E indicate setal groups.
Scale bars: A, B and C = 50mm.
338 Invertebrate Systematics J. A. Cruz-López and O. F. Francke
(A)(B)
(C)
Fig. 28. Habitus of Tonalteca spinooculorum.(A) Dorsal view of male. (B) Ventral view of female. (C) Lateral view of male. Scale bars: A, B = 1.0 mm,
C = 1.5 mm.
Total evidence phylogeny of Stygnopsidae Invertebrate Systematics 339
planus and ‘H.’zullinii remain as incertae sedis in Karosinae and
Stygnopsinae, respectively. These new taxonomic proposals are
described below.
Genus Serrobunus Goodnight & Goodnight, stat. rev.
Serrobunus Goodnight & Goodnight, 1942: 2.
Hoplobunus (in part): Goodnight & Goodnight, 1953: 19.
Included taxa:Serrobunus boneti Goodnight & Goodnight, 1942 stat.
rev., Serrobunus queretarius (Šilhavý, 1974), comb. nov.
Emended diagnosis
Large harvestmen (scutum length ~6 mm), troglomorphic.
Ocularium very high, conical, with long apical spine (Fig. 20).
Chelicera large and sexually dimorphic. Cheliceral dentition
heterogeneous, basal tooth on movable finger blunt (Fig. 21).
Legs very long, femur IV longer than scutum, straight. Femur
IV ornate with longitudinal rows of spiniform tubercles, all
tubercles of the rows increasing in size apically, except on the
ventral retrolateral row, which decrease in size (Fig. 22C–F).
Trochanter III slightly globular. Trochanter IV with both dorsal
and ventral apophyses (Fig. 22C–F). Penis setal Paramitraceras-
pattern, with the following modifications: setae C formed by three
orfour macrosetae, the apical-most small, three or four setae A+B,
laterally on truncus, slightly below of follis base, one or two pairs
of setae D, slightly shorter than other macrosetae, lateral to follis,
without setae E (Fig. 23). Apical margin of pars distalis rounded.
Follis with bilobular dorsal projection and apically covered by
acute spines (Fig. 23A,D).
Remarks
Kury and Villarreal (2015) examined two males of S. boneti
labelled ‘Mexico, SL Potosí, Cueva de los Sabinos, near Valles,
underground waterway to Devil’s Hole, 26.iii.1946, EJ Dontzin
& E Ruda leg.’. These authors mentioned the presence of two
median pairs of microsetae E forming a rectangle (Kury and
Villarreal 2015:fig. 1B, C). Examining material from different
localities of S. boneti (Supplementary Material 3), we did not find
evidence of the presence of these setae. Therefore, it is possible
that Kury and Villarreal (2015) examined a different taxon.
Despite not including H. queretarius in the molecular
analyses, we have transferred it to Serrobunus because it
shares external and male genital morphology with S. boneti
(Fig. 23D–F).
Genus Iztlina, gen. nov.
(Figs 24–27)
http://zoobank.org/NomenclaturalActs/urn:lsid:zoobank.org:act:
A6450531-D49F-46F9-BD91-0202703BA758
Type species:Iztlina venefica, sp. nov.
Diagnosis
Medium-size harvestmen (scutum length <5 mm), troglomorphic.
Ocularium very high, conical, apex divided into two divergent
tips (Fig. 26A). Mesotergal area IV with two long paramedian
spines (Fig. 26B). Chelicera large and sexually dimorphic.
Cheliceral dentition heterogeneous, basal tooth on movable
finger blunt (Fig. 25). Legs very long, femur IV longer than
scutum, straight. Femora III and IV almost smooth, femur III with
one ventral longitudinal row of spiniform tubercles increasing in
size distally (Fig. 26D), femur IV with two ventral rows apically
(Fig. 26E). Trochanter III slightly globose. Setation of penis
unique for subfamily: five or six setae C laterally to follis, two
pairs of setae A below setae C, two ventral longitudinal rows of
three pairs of setae E, setae D absent (Fig. 27). All penial setae are
uniform in size and shape. Truncus cylindrical, almost contiguous
with ventral projection, ventro-apical margin with U-shaped
(A)(B)
Fig. 30. Pedipalp and femur IV of Tonalteca spinooculorum male.
(A) Mesal view of pedipalp. (B) Ventral view of femur IV. Scale bars:
A = 1.0 mm, B = 1.5 mm.
(A)
(B)
Fig. 29. Chelicera of Tonalteca spinooculorum male. (A) Frontal view.
(B) Mesal view. Black arrow indicates the basal tooth, white arrow indicates
the median tooth. Scale bars: A, B = 1.0mm.
340 Invertebrate Systematics J. A. Cruz-López and O. F. Francke
concavity (Fig. 27C). Follis globose, with dorsal lobular
projection (Fig. 27A,B).
Remarks
This genus can be easily distinguished from other genera of
Stygnopsinae by the unique presence of two long paramedian
spines on mesotergal area IV and apex of ocularium divided in
two tips and general shape of male genitalia.
Etymology
‘Iztli’is the name of a god of sacrifice and stone knives in the
Mexica culture. This god is associated with the deities of death
and darkness. The name was modified to feminine ending.
Iztlina venefica, sp. nov.
(Figs 24–27)
http://zoobank.org/NomenclaturalActs/urn:lsid:zoobank.org:
act:0E780459-A572-41C2-B7A3-F0DB9A279862
Material examined
Holotype. MEXICO: Chiapas: Municipio de Cintalapa: male, Cueva
del Arco, 16"50051.600N, 93"43004.500 W, 19.xi.2011, coll. K. Zárate, male
(CNAN-T1092).
Paratypes. MEXICO: Chiapas: Municipio de Cintalapa: 2 males,
collected with holotype (CNAN-T1093); 1 male, 1 female, Cueva Ejidal,
EjidoLópezMateos,16"51054.100 N 93"42045.300 W, 23.iv.2013, coll. K.Zárate
(CNAN-T1101).
Diagnosis
As for the genus.
Description
Male (holotype)
Scutum length 4.5 mm, scutum width 3.7, femur II 9.5 mm,
femur IV 11.7 mm.
Dorsum. Scutum type z, mid-bulge gently rounded, both
constrictions 1 and 2 shallow (Fig. 24A,B). Ocularium at frontal
margin of prosoma, very high and conical shape, with the
(A)(B)(C)
Fig. 31. Male genitalia of Tonalteca spinooculorum.(A) Dorsal view. (B) Lateral view. (C) Ventral view. Small letters A, B, and C indicate setal groups,
Er indicates vestigial insertion bases of E setae. Scale bars: A, B and C = 100mm.
Total evidence phylogeny of Stygnopsidae Invertebrate Systematics 341
apex divided in two diverging tips (Fig. 26A). Eyes well
marked but small-sized, at the base of ocularium. Dorsum
smooth, mesotergal sulci wide and shallow. Mesotergal
areas I–III with transverse row of minute and scattered
setiferous tubercles, area IV with one pair of long paramedian
spines (Fig. 26B). Lateral pegs forming a continuous and
distintive row.
Venter. Coxae I and II ornate with long spiniform
setiferous tubercles, coxae III and IV with small ones and
setae. Length of coxa IV similar to length of coxa III. Lateral
margins posterior to genital operculum parallel.
Chelicera. Basichelicerite elongated, bulla softly marked.
Cheliceral hand swollen, dorsal portion elevated, frontal face
covered by small tubercles and spiniform setae (Fig. 25B). Fixed
finger with a row of six teeth, from the base to almost the end,
increasing in size slightly apically. Movable finger with basal
blunt tooth and four flat teeth medially (Fig. 25A).
Pedipalp. All segments almost rectangulgar in cross-
section. Trochanter and basal femur with one long spiniform
setiferous tubercle ventrally (Fig. 24C). Ventrally femur with
longitudinal row of five scattered tubercles. Patella unarmed.
Tibia and tarsus armed with long spiniform setiferous tubercles
as follows: both margins of tibia IiIi (3 >1>4>2), ectal margin of
tarsus IIII (1 = 2 = 3 = 4), mesal margin of tarsus III (1 = 2 = 3)
(Fig. 26C). Claw elongated.
Legs. All segments long and slenders, anterior legs
covered with few small spiniform setae. Posterior legs covered
with small tubercles. Femur III with two ventral rows of scattered
spiniform tubercles. Femur IV with two ventral rows of tubercles
confined only to apical portion (Fig. 26D,E). Tarsal count:
8(3):20(4):8:9/10.
Penis. Truncus cylindrical, flimsy lamina slightly projected,
contiguous with the truncus, with apical concavity U-shaped
(Fig. 27C). All macrosetae of penis of the same size and shape,
with longitudinal sulcus and rounded apices (Fig. 27). Setation
of penis: five or six setae C laterally to follis, two pairs of setae
A basal to setae C, two ventral longitudinal rows of three pairs
of setae E, setae D absent (Figs 27). Follis with dorsal lobe
projection (Fig. 27A).
Female (paratype)
Similar to male, but with chelicera slightly smaller
(Fig. 24A,B).
(A)(B)(C)(D)
(E)(F)
Fig. 32. Chelicera of Karosinae males. (A) Frontal view of Karos barbarikos.(B) Mesal view of Karos barbarikos.(C) Frontal view of Chapulobunus
unispinosus.(D) Mesal view of Chapulobunus unispinosus.(E) Frontal view of Huasteca gratiosa.(F) Mesal view of Huasteca gratiosa. Scale bars: A, B, E,
F = 0.2 mm, C, D = 0.3 mm.
342 Invertebrate Systematics J. A. Cruz-López and O. F. Francke
Etymology
From Latin ‘venefica’, which means ‘a female who poisons’
in reference to a sorceress who used poisons and potions for
various reasons.
Genus Tonalteca, gen. nov.
(Figs 28–31)
http://zoobank.org/NomenclaturalActs/urn:lsid:zoobank.org:
act:7DECBCBC-5A22-4E6D-89A9-286AFF240825
Hoplobunus (in part): Goodnight & Goodnight, 1973: 86.
Type species:Hoplobunus spinooculorum Goodnight & Goodnight,
1973.
Diagnosis
Medium-size harvestmen (scutum length ~5mm), troglomorphic.
Ocularium very high, conical, with long apical spine (Fig. 28C).
Chelicera large and sexually dimorphic. Basal tooth on movable
finger blunt (Fig. 29A). Legs long, femur IV slightly longer
than scutum, straight. Ornamentation of all legs formed by
longitudinal rows of small, scattered spiniform setiferous
tubercles (Fig. 30B). Penis setal Stygnopsis-pattern, two
longitudinal pairs of setae C, two transversal pairs of setae A,
one basalmost pair of setae B, and two pairs of small setae D,
near to follis base (Fig. 31). With one ventral pair of vestigial
insertion bases of setae E (Fig. 31C). Pars distalis forming
an angle of 90"at the base of ventral projection. Pars distalis
slightly compressed laterally (Fig. 31A). Follis with dorsal lobe
rounded (Fig. 31A).
Remarks
Tonalteca can be distinguished from the following epigean
stygnopsin genera (Hoplobunus,Paramitraceras,Philora,
Tampiconus and Sbordonia) in having femur IV straight and
longer than the scutum. It can be separated from the troglomorphic
Chinquipellobunus,Mexotroglinus and Troglostygnopsis by
the ocularium with a long apical spine. Tonalteca can be
distinguished from the troglomorphic Iztlina by the absence of
two paramedian spines on mesotergal area IV. It can be separated
from Serrobunus by legs III and IV without remarkable rows of
spiniform tubercles, and finally, from Stygnopsis by pedipalpal
patella covered with few small tubercles and the absence of a
mesal apophysis.
Etymology
The ‘Tonalteca’is the initial waltz of a folkloric dance named
‘Flor de Piña’from Oaxaca, Mexico. This dance is an emblematic
cultural reference to Oaxaca, specially the Northern region of
Papaloapan, where this genus occurs. The name is feminine.
Subfamily KAROSINAE, subfam. nov.
http://zoobank.org/NomenclaturalActs/urn:lsid:zoobank.org:act:
B6392819-5994-4F15-9EEB-409F3BB5C4B1
Phalangodinae (in part): Roewer, 1912: 108; Roewer, 1923: 69;
Goodnight & Goodnight, 1944: 1; Goodnight & Goodnight, 1947b:
3; Goodnight & Goodnight, 1953: 13; Goodnight & Goodnight, 1971:
33; Goodnight & Goodnight, 1973: 83; Šilhavý,1974: 185; Šilhavý,
1977: 227; Rambla & Juberthie, 1994: 218.
(A)(B)(C)
Fig. 33. Dorsal views of pedipalp femur and patella of Karosinae males. (A)Karos barbarikos.(B)Chapulobunus unispinosus.(C)Huasteca gratiosa. Scale
bars: A, B, C = 0.3mm.
Total evidence phylogeny of Stygnopsidae Invertebrate Systematics 343
(A)(B)(C)
(D)(E)(F)
Fig. 34. Male genitalia of Karosinae. (A) Dorsal view of Karos barbarikos.(B) Lateral view of Karos barbarikos.(C) Ventral view of Karos barbarikos.
(D) Dorsal view of Chapulobunus asper.(E) Lateral view of Chapulobunus asper.(F) Ventral view of Chapulobunus asper. Small letters D and E indicate
setal groups, B? indicates possibly B setae. Scale bars: A, B, C, D, E, F= 100 mm.
344 Invertebrate Systematics J. A. Cruz-López and O. F. Francke
(A)(B)(C)
(D)(E)(F)
(G)(H)
Fig. 35. Dorsal habitus of Karosinae and Paramitraceras-group. (A)Karos barbarikos.(B)Huasteca gratiosa.(C)Chapulobunus
unispinosus.(D)Philora tuxtlae.(E)Paramitraceras veracruz.(F)Paramitraceras aff. hispidulum.(G)Troglostygnopsis sp. 0049.
(H)Paramitraceras aff. granulatum. Scale bars: A, B, C, D = 0.5 cm, E, F, G, H= 1 cm.
Total evidence phylogeny of Stygnopsidae Invertebrate Systematics 345
(A)(B)
(C)(D)
(E)(F)
Fig. 36. Lateral projections of scutum and mesotergal sulci of Karosinae and Paramitraceras-group. (A) Lateral projection of Paramitraceras aff.
granulatum.(B) Lateral projection of Paramitraceras veracruz.(C) Lateral projection of Huasteca gratiosa.(D) Mesotergal sulci III and IV of Huasteca
gratiosa.(E) Detail of mesotergal sulci II and III of Karos barbarikos.(F) Lateral projection of Karos barbarikos. MsII to MsIV indicate mesotergal sulcus II,
III and IV respectively. Scale bars: A = 0.5 mm, B = 250 mm, C, E, F = 100 mm, D = 200 mm.
346 Invertebrate Systematics J. A. Cruz-López and O. F. Francke
(A)(B)
(C)(D)
(E)(F)
(G)(H)
Fig. 37. Details of lateral projections and associated pores in Karosinae and Paramitraceras-group. (A) Pores at level of mesotergal
area I in Karos barbarikos.(B) Projection of corner on mesotergal area V in Karos barbarikos.(C) Lateral projection of Huasteca
gratiosa.(D) Pores on lateral projection in Huasteca gratiosa.(E) Lateral projection of Paramitraceras veracruz.(F) Pores on lateral
projection in Paramitraceras veracruz.(G) Lateral projection in Troglostygnopsis sp. 0049. (H) Detail of lateral projection in
Troglostygnopsis sp. 0049. Arrows indicate some pores. Scale bars: A, D, H = 25 mm, B, C, F, G =50= mm, E = 100 mm.
Total evidence phylogeny of Stygnopsidae Invertebrate Systematics 347
Stygnopsinae (in part): Goodnight & Goodnight, 1945: 1; Goodnight &
Goodnight, 1946: 1.
Karos-group (in part): Cruz-López & Francke, 2015: 838.
Type genus:Karos Goodnight & Goodnight, 1944.
Included taxa:Karos Goodnight & Goodnight, 1944, Monterella Goodnight
&Goodnight,1944,Montabunus Goodnight & Goodnight, 1945,
Chapulobunus Goodnight & Goodnight, 1946, Potosa Goodnight &
Goodnight, 1947, Crettaros Cruz-López & Francke, 2015, Huasteca
Cruz-López & Francke, 2015, Mictlana Cruz-López & Francke, 2015,
and the uncertain ‘Karos’depressus Goodnight & Goodnight, 1971 and
‘Hoplobunus’planus Goodnight & Goodnight, 1973.
Diagnosis
Modified from Cruz-López and Francke (2015). Small to medium
harvestmen (<to ~6 mm). Ocularium in the middle of prosoma,
(A)(B)
(C)(D)
(E)(F)
Fig. 38. Colour detail of ventral glandular tubercles on stigmatic area in Paramitraceras-group males. (A)Paramitraceras aff. hispidulum.(B)Paramitraceras
tzotzil.(C)Paramitracerasaff. hispidulum.(D)Sbordonia aff. parvula.(E)Paramitracerasveracruz.(F)Philora tuxtlae. Scale bars: A = 1.3 mm, B, E = 0.8 mm,
C, D = 0.7 mm, F = 0.4 mm.
348 Invertebrate Systematics J. A. Cruz-López and O. F. Francke
small to medium size, generally unarmed (Fig. 35A–C). Usually
with spiniform bulge anterior to ocularium (Fig. 35A–C).
Mesotergal sulci generally sinuous (Figs 35A–C,36D,E).
Lateral channels at level of mesotergal areas I, II, V and usually
in free tergites forming rounded projections, these projections have
the surface covered by small pores (Figs 35A–C,36C,F,37A–D).
Chelicera small, both fingers with similar, small and uniform
dentition (Fig. 32). Meso-apical surface of pedipalp femur and
patela armed with spiniform setiferous tubercles (Fig. 33). Penis
setation Karos-pattern. Pars distalis contiguous with ventral
projection, not forming an angle (Fig. 34).
Remarks
Karosinae exhibits more uniform diagnostic characters than
Stygnopsinae. The ocularium in the middle of prosoma and
mesal armature of pedipalpal femur and patella are consistent
also in ‘H.’planus and ‘K.’depressus, species known from
females only. The spiniform bulge anterior to the ocularium is
an inconspicuous character present in almost all Karosinae
examined (Fig. 35A–C). Mictlana inops and ‘H.’planus
exhibit two dorsal bulges on the prosoma. Cruz-López and
Francke (2015) considered the anterior bulge in M. inops as
the ocularium; a similar structure is present in ‘H.’planus. In
this work, we are not sure which of the two bulges is the
ocularium because without evidence of eyes, retina or eye-spots,
we are uncertain, as occurs in Jarmilana pecki (Goodnight and
Goodnight 1977), as indicated by Cruz-López et al.(2016).
Conflictive taxa, the case of Mexotroglinus and Isaeus
The genus Mexotroglinus is the most controversial taxon within
the family. This genus has small chelicerae without sexual
(A)(B)(C)
(D)(E)(F)
(G)(H)
Fig. 39. Detail of ventral glandular tubercles on stigmatic area in Paramitraceras-group males. (A) Stigmatic area of Paramitraceras aff. hispidulum.
(B) Glandular tubercles of Paramitraceras aff. hispidulum.(C) Detail of one glandular tubercle of Paramitraceras aff. hispidulum.(D) Stigmatic area of
Philora tuxtlae.(E) The four glandular tubercles of Philora tuxtlae.(F) Detail of one glandular tubercle of Philora tuxtlae.(G) Detail of one glandular tubercle
of Paramitraceras aff. granulatum.(H) Detail of one glandular tubercle of Paramitraceras veracruz. Scale bars: A = 1 mm, B = 100 mm, C, G = 25 mm,
D = 200 mm, E = 50 mm, F = 10 mm, H = 20 mm.
Total evidence phylogeny of Stygnopsidae Invertebrate Systematics 349
(A)(B)(C)
(D)(E)(F)
Fig. 40. Male genitalia of Paramitraceras-group members. (A) Dorsal view of Paramitraceras aff. granulatum.(B) Lateral view of Paramitraceras aff.
granulatum.(C) Ventral view of Paramitraceras aff. granulatum.(D) Dorsal view of Paramitraceras tzotzil.(E) Lateral view of Paramitraceras tzotzil.(F)
Ventral view of Paramitraceras tzotzil. Small letters C, D, E and A+B indicate setal groups. Scale bars: A, B, D, E, F = 100mm, C = 200 mm.
350 Invertebrate Systematics J. A. Cruz-López and O. F. Francke
(A)(B)(C)
(D)(E)(F)
Fig. 41. Male genitalia of Paramitraceras-group members. (A) Dorsal view of Paramitraceras aff. hispidulum.(B) Lateral view of Paramitraceras aff.
hispidulum.(C) Ventral view of Paramitraceras aff.hispidulum.(D) Dorsal view of Sbordonia aff.parvula.(E) Lateral view ofSbordonia aff.parvula.(F) Ventral
view of Sbordonia aff. parvula. Small letters C, D, E and A+B indicate setal groups. Scale bars: A, B, D, E, F = 50mm, C = 100 mm.
Total evidence phylogeny of Stygnopsidae Invertebrate Systematics 351
dimorphism (Fig. 46D), mesal armature on the pedipalpal femur
and patella (Fig. 46H,I), and the male genitalia is similar to the
Karos-pattern (Fig. 47). This combination of character states
places Mexotroglinus in Karosinae. However, the ocularium
located on the frontal margin of the prosoma (Fig. 46A,B), the
straight mesotergal sulci (Fig. 46A,B,E), the lateral channels
not projected laterally and the absence of associated pores
(Fig. 46A,B,E,F) suggest its position within Stygnopsinae.
The three analyses are consistent in the inclusion of
Mexotroglinus in Stygnopsinae. In our results, both BI and
ML analyses of total evidence showed a sister-group
relationship between Mexotroglinus and Chinquipellobunus,
with high support value in the BI analysis, but without
significant support in ML. Based on the phylogenetic evidence,
(A)(B)(C)
(D)(E)(F)
Fig. 42. Male genitalia of Paramitraceras-group members. (A) Dorsal view of Sbordonia sp. 0055. (B) Lateral view of Sbordonia sp. 0055. (C) Ventral view of
Sbordonia sp. 0055. (D) Dorsal view of Troglostygnopsis sp. 0049. (E) Lateral view of Troglostygnopsis sp. 0049. (F) Ventral view of Troglostygnopsis sp. 0049.
Small letters C, D, E and A+B indicate setal groups. Scale bars: A, B, C, D, F = 100mm, E = 50 mm.
352 Invertebrate Systematics J. A. Cruz-López and O. F. Francke
Mexotroglinus is the most extreme case of morphological
convergence, allocated to Stygnopsinae, but also showing some
important diagnostic morphological characters of Karosinae, as
mentioned above.
Together with the description of Stygnopsidae, Sørensen
(1932) described the monotypic genus Isaeus from Mexico.
He considered that the ectal and mesal armature on the
pedipalpal patella were significant characters to differentiate
between Stygnopsis and Isaeus, respectively. Also, Sørensen
(1932) considered Isaeus related to the monotypic Mexican
genus Metaconomma Pickard-Cambridge, 1905, a taxon
which currently is incertae sedis in Grassatores (Laniatores)
(Kury 2003). Later, Goodnight and Goodnight (1953)
considered Isaeus a junior synonym of Hoplobunus. We here
(A)(B)(C)(D)
(E)(F)(G)(H)
(I)(J)(K)(L)
Fig. 43. Chelicera of Paramitraceras-group males. (A) Frontal view of Paramitraceras aff. granulatum.(B) Mesal view of Paramitraceras aff.
granulatum.(C) Frontal view of Paramitraceras aff. hispidulum.(D) Mesal view of Paramitraceras aff. hispidulum.(E) Frontal view of Paramitraceras
tzotzil.(F) Mesal view of Paramitraceras tzotzil.(G) Frontal view of Paramitraceras veracruz.(H) Mesal view of Paramitraceras veracruz.(I) Frontal
view of Sbordonia sp. 0055. (J) Mesal view of Sbordonia sp. 0055. (K) Frontal view of Sbordonia aff. parvula.(L) Mesal view of Sbordonia aff. parvula.
Black arrows indicate the basal blunt tooth on movable finger, green arrows indicate the basal tooth on fixed finger, and white row indicates the median
tooth un movable finger. Scale bars: A, B = 1.6mm, C, D = 0,4 mm, E, F= 1.0 mm, G, H = 0.5 mm, I, J = 0.8 mm, K, L = 0.7 mm.
Total evidence phylogeny of Stygnopsidae Invertebrate Systematics 353
demonstrate the polyphyly of Hoplobunus, revalidating all of the
synonymised genera. In the case of Isaeus, we revalidate it below,
hoping that in the future this genus can be studied properly, a
nomenclatural act similar to those generic revalidations in
Cosmetidae proposed by Kury (2003).
Genus Isaeus Sørensen, stat. rev.
Isaeus Sørensen, 1932: 276.
Hoplobunus (in part): Goodnight & Goodnight, 1953: 19.
Included taxa:Isaeus mexicanus Sørensen, 1932, by monotypy.
Remarks
We were unable to examine any specimens of I. mexicanus, and
therefore cannot assign Isaeus to a subfamily.
Discussion
Genital evolution and homoplastic characters
The capsula externa forming a multifolded follis has been
considered a plesiomorphic condition by some authors under
different approaches (Kury 1997; Mendes and Kury 2007; Kury
and Villarreal 2015). This multifolded follis has been considered
a synapomorphy or a convergence among some Epedanoidea,
Assamioidea and Stygnopsidae, depending on different authors
(Kury 1997; Sharma et al.2011). The detailed examination
of penes in Assamiidae, Epedanidae, Pyramidopidae and
Tithaeidae, corroborated that the genital capsula externa and
glans are different structures (not homologous), and that they
exhibit morpho-mechanical differences when the capsula
externa is expanded using chemical sustances such as lactic
acid (Martens 1986; Zhang and Zhang 2010; Lian et al.2011;
Sharma et al.2011; Cruz-López et al.2016).
Stygnopsidae exhibits a great diversity in genital morphology,
especially in the shape of the pars distalis and in penial setation.
However, within each genus, the general pattern of the genital
morphology is stable. The only known genital convergence
occurs between Karos and Crettaros, according to the
morphological cladistics analysis presented by Cruz-López
and Francke (2015). This convergence is corroborated here in
the BI and ML analyses using combined data. In the present
contribution, we observed another genital convergence between
Hoplobunus and the Tampiconus-like clade (Figs 1,14,19),
and, unlike Karos and Crettaros, these taxa are similar externally,
as discussed previously.
All members of Paramitraceras, plus Sbordonia parvula
and S. aff. parvula exhibit very uniform external morphology
(A)(B)(C)
(D)
(E)
(F)
(G)
Fig. 44. Pedipalps of members of Paramitraceras-group males. (A) Mesal view of Paramitraceras aff. granulatum.(B) Mesal view of Paramitraceras aff.
hispidulum.(C) Mesal view of Paramitraceras tzotzil.(D) Mesal view of Paramitraceras veracruz.(E) Ectal view of Sbordonia sp. 0055. (F) Mesal view of
Sbordonia aff. parvula.(G) Mesal view of Sbordonia armigera. Scale bars: A = 3.0 mm, B = 0.9 mm, C = 1.9 mm, D = 1.3 mm, E = 1.5 mm, F, G = 1.0 mm.
354 Invertebrate Systematics J. A. Cruz-López and O. F. Francke
and unarmed pedipalps, ornate only with spiniform setae on
longitudinal keels on tibia and tarsus, and sometimes with few
spiniform setiferous tubercles (Figs 44,45). Surprisingly, males
of these taxa have glandular tubercles on the stigmatic area
(Figs 38,39), characters reported here for the first time in
Laniatores. According to Cruz-López and Francke (2013a),
the robust body, a dorsally convex opisthosoma and unarmed
pedipalps are diagnostic characters for Paramitraceras.
Subsequently, Cruz-López and Francke (2013b) described the
male genitalia as Paramitraceras-pattern, which is present in
(A)(B)(C)
(D)
(E)(F)(G)
Fig. 45. Ventral views of pedipalp tibia and tarsus of Paramitraceras-group males. (A)Paramitraceras aff. granulatum.(B)Paramitraceras aff. hispidulum.
(C)Paramitraceras tzotzil.(D)Paramitraceras veracruz.(E)Sbordonia sp. 0055. (F)Sbordonia aff. parvula.(G)Sbordonia armigera. Scale bars: A = 1.7 mm,
B = 0.4 mm, C = 0.9 mm, D = 0.6 mm, E = 0.7 mm, F, G = 0.5 mm.
Total evidence phylogeny of Stygnopsidae Invertebrate Systematics 355
(A)(B)(C)(D)
(E)(F)(G)
(H)(I)
Fig. 46. External morphology of Mexotroglinus aff. sbordonii male. (A) Dorsal habitus. (B) Lateral habitus. (C) Mesal view of tibia II. (D) Frontal view of
chelicera. (E) Detail of mesotergal sulci II–IV. (F) Lateral channels at level of mesotergal area I. (G) Ventro-mesal detail of apical swollen in tibia II. (H) Dorsal
view of pedipalp femur and tibia. (I) Ventral view of pedipalp femur and tibia. Scale bars: A, B, C =250 mm, D = 150 mm, E = 200 mm, F, G = 50 mm.
356 Invertebrate Systematics J. A. Cruz-López and O. F. Francke
Paramitraceras,Philora and Troglostygnopsis. Surprisingly, the
male genitalia of Paramitraceras tzotzil Cruz-López & Francke,
2013, exhibits penial modifications, as dorso-ventral constriction
of pars distalis and nonrecognition of setal groups A, B and
C (Fig. 40D–F). The Paramitraceras genital pattern in the
clade (Serrobunus (Sbordonia aff. parvula (P. veracruz
((P. aff. granulatum (Troglostygnopsis +Sbordonia sp. 0055))
(Philora +P. tzotzil)))), according to the BI analyses of
combined data (Fig. 1), has apparently evolved as follows. In
Serrobunus, the setae D are present, with similar lengths of the
other macrosetae, whereas setae E are absent. In S. parvula and
S. aff. parvula, the setal pattern is unrecognisable, and these taxa
also have a dorsal expandable lobe, which is broken through by
the one pair of long setae D (Fig. 41D–F). In the remaining taxa,
the Paramitraceras genital pattern is expressed as Cruz-
López and Francke (2013b) described, except in P. tzotzil, in
which the pars distalis and the ventral lamina are contiguos,
without apical depression, and the lateral macrosetae groups
are unrecognisable. Also, this modified genital pattern is
present in an undescribed Paramitraceras species similar to
P. tzotzil.
According to the BI analysis of combined data, the unarmed
pedipalps could be a homoplastic character, present in the
Paramitraceras-group in one clade, and in P. hispidulum +P.
aff. hispidulum in a totally different clade. In the Paramitraceras-
group, the unarmed pedipalps could have appeared only
once in the clade, with subsequent reversal in Philora and
Troglostygnopsis +Sbordonia sp. 0055 (Figs 44,45). However,
spiniform setiferous tubercles in Philora and Sbordonia sp. 0055
are reduced in size compared with Troglostygnopsis. This may be
due to the troglobitic habits in Troglostygnopsis, since the only
other known cave-dwelling species in the clade, Philora nympha
Cruz-López & Francke, 2016, also exhibits very long pedipalpal
armature (Cruz-López and Francke 2016b). The glandular
tubercles present ventrally on males of Paramitraceras,
S. parvula,S. aff. parvula and Philora are another remarkable
convergence, which may have evolved in a similar way as the
unarmed pedipalps, as in the Paramitraceras-group clade, with
subsequent loss in Troglostygnopsis, and S. parvula +S. aff.
parvula group.
Pores associated with median projections on the lateral
channels are convergently found in Karosinae and Paramitraceras,
Philora and Sbordonia aff. parvula (Fig. 37A–F). Detailed
examination of Troglostygnopsis did not reveal the presence of
these pores, although this genus has median lateral projections
similar to those on Paramitraceras (Fig. 37F,G). The function
of these pores and their relationship to the lateral median
projections are unclear.
Acknowledgements
Thanks are extended to the Graduate Program in Biological Sciences of the
National Autonomous University of Mexico (UNAM). This paper constitutes
a partial fulfilment of the Graduate Program in Biological Science of
the UNAM. We thank the scholarship and financial support provided by
the National Council of Science and Technology (CONACYT) (scholarship
number 249637) and the Institute of Biology (UNAM) for the infrastructure
(A)(B)(C)
Fig. 47. Male genitalia of Mexotroglinus aff. sbordonii.(A) Dorsal view. (B) Lateral view. (C) Ventral view. Small letters D indicate setal groups. Scale bars:
A, B, C = 50 mm.
Total evidence phylogeny of Stygnopsidae Invertebrate Systematics 357
provided. Also we thank CONACYT project #271108 ‘Red temática
Código de Barras de la Vida’(continuidad de redes temáticas) for
providing financial support to field and molecular work. The authors are
grateful to many people who have contributed to this study. Field work
was possible thanks to all members of Colección Nacional de Arácnidos
(CNAN) and Colección Nacional de Ácaros (CNAC) from UNAM, especially
toD.Barrales,G.Contreras,J. Mendoza, R. Monjaraz, G. Montiel, R. Paredes,
C. Santibáñez and A. Valdez. Additionally, people who collected several
specimens examined from different institutions: J. Bokma, B. Damron,
E. Goyer, S. Longhorn, E. Miranda, A. Pérez, P. Sprouse, K. Zárate. We
thank the following speleological groups: Grupo Espeleológico La
Venta, Grupo Espeleológico Jaguar and Proyecto Espeleológico Sistema
Huautla (PESH). We thank the curators, researchers and colleages from
museums and collections whichs loaned types and additionall material:
L. Prendini (AMNH), J. Cokendolpher (TTU), J. Reddell (TMM),
I. Vázquez (CAAH), J. Beccaloni and S. Longhorn (BMNH), M. Ramírez,
A. Pérez and C. Grismado (MACN), P. Jaeger (SMF), B. Huber (ZFMK),
A. Valdez (IBUNAM), and J. Mendonza, and D. Candia (CNAN). Molecular
work was possible with the help of A. Jiménez, L. Márquez and
P. Rosas from Biología Molecular laboratory of Laboratorio Nacional de
Biodiversidad (LANABIO) at UNAM. Scanning electronic photographs
were possible with the help of B. Mendoza from Microscopia Electrónica
(LANABIO), with recommendations of A. Pérez and D. Proud (MACN).
Important comments and suggestions are thanked to J. Morrone and
A. Zaldívar (UNAM), C. Santibáñez (IBT), M. Harvey (Western
Australian Museum) and two anonymous reviewers. Finally, JACL thanks
his wife Catalina Juárez, for her patience and eternal unconditional support.
References
Acosta, L. E., Pérez-González, A., and Tourinho, A. L. (2007). Methods
for taxonomic study. In ‘Harvestmen: The Biology of Opiliones’. (Eds
R. Pinto-da-Rocha, G. Machado and G. Giribet.) pp. 494–505. (Harvard
University Press: Cambridge, MA.)
Avram, S. (1977). Recherches sur les Opilionides de Cuba IV. Genres et
espèces nouveaux d’Agoristeninae (Agoristenidae, Gonyleptomorphi).
Resultados expédititions biospéologiques cubano-rumana Cuba 2,
137–143.
Banks, N. (1900). New genera and species of American Phalangida. Journal
of the New York Entomological Society 8, 199–201.
Boyer, S. L., Karaman, I., and Giribet, G. (2005). The genus Cyphophthalmus
(Arachnida, Opiliones, Cyphophthalmi) in Europe: a phylogenetic
approach to Balkan Peninsula biogeography. Molecular Phylogenetics
and Evolution 36, 554–567. doi:10.1016/j.ympev.2005.04.004
Bragagnolo, C., Hara, M. R., and Pinto-da-Rocha, R. (2015). A new family of
Gonyleptoidea from South America (Opiliones, Laniatores). Zoological
Journal of the Linnean Society 173, 296–319. doi:10.1111/zoj.12207
Cokendolpher, J. C. (2004). Revalidation of the genus Chinquipellobunus
(Opiliones: Stygnopsidae). Texas Memorial Museum, Speleological
Monographs 6, 143–152.
Cruz-López, J. A., and Francke, O. F. (2013a). Two new species of the
genus Paramitraceras Pickard-Cambridge, 1905 (Opiliones: Laniatores:
Stygnopsidae) from Chiapas, Mexico. Zootaxa 3641, 481–490.
doi:10.11646/zootaxa.3641.4.13
Cruz-López, J. A., and Francke, O. F. (2013b). On the enigmatic genus
Philora: familial assignment and taxonomic revision (Opiliones:
Laniatores: Stygnopsidae). The Journal of Arachnology 41, 291–305.
doi:10.1636/Ha13-13.1
Cruz-López, J. A., and Francke, O. F. (2015). Cladistic analysis and
taxonomic revision of the genus Karos Goodnight & Goodnight, 1944
(Opiliones, Laniatores, Stygnopsidae). Zoological Journal of the
Linnean Society 175, 827–891. doi:10.1111/zoj.12299
Cruz-López, J. A., and Francke, O. F. (2016a). Three new species of the
Mexican harvestman genus Chapulobunus (Opiliones: Stygnopsidae).
The Journal of Arachnology 44, 65–75. doi:10.1636/Ha15-44.1
Cruz-López, J. A., and Francke, O. F. (2016b). Three new harvestman
species of the genus Philora (Opiliones: Gonyleptoidea, Stygnopsidae)
with comments on troglomorphisms. Revista Mexicana de Biodiversidad
87, 328–336. doi:10.1016/j.rmb.2016.02.004
Cruz-López, J. A., Proud, D., and Pérez-González, A. (2016). When
troglomorphism dupes taxonomists: morphology and molecules reveal
the first pyramidopid harvestman (Arachnida, Opiliones, Pyramidopidae)
from the New World. Zoological Journal of the Linnean Society 177,
602–620. doi:10.1111/zoj.12382
Darriba, D., Taboada, G. L., Doallo, R., and Posada, D. (2012). jModelTest 2:
more models, new heuristics and parallel computing. Nature Methods 9,
772. doi:10.1038/nmeth.2109
Edgar, A. L. (1990). Opiliones (Phalangida). In ‘Soil Biology Guide’. (Ed.
D. L. Dindal.) pp. 529–581. (John Wiley & Sons: New York.)
Edgecombe,G.D.,andGiribet,G. (2006). A century later –a total evidence re-
evaluation of scutigeromorph centipedes (Myriapoda: Chilopoda).
Invertebrate Systematics 20, 503–525. doi:10.1071/IS05044
Edgecombe, G. D., Giribet, G., and Wheeler, W. C. (2002). Phylogeny of
Henicopidae (Chilopoda: Lithobiomorpha): a combined analysis of
morphology and five molecular loci. Systematic Entomology 27,
31–64. doi:10.1046/j.0307-6970.2001.00163.x
Folmer,O.,Black,M.,Hoeh, W., Lutz, R., and Vrijenhoek, R.C.(1994).DNA
primers for amplification of mitochondrial cytochrome coxidase subunit I
from diverse metazoan invertebrates. Molecular Marine Biology and
Biotechnology 3, 294–299.
Giribet, G., and Sharma, P. P. (2015). Evolutionary biology of harvestmen
(Arachnida, Opiliones). Annual Review of Entomology 60, 157–175.
doi:10.1146/annurev-ento-010814-021028
Giribet, G., Edgecombe, G. D., Wheeler, W. C., and Babbitt, C. (2002).
Phylogeny and systematic position of Opiliones: a combined analysis of
chelicerate relationships using morphological and molecular data.
Cladistics 18,5–70.
Giribet, G., Vogt, L., Pérez-González, A., Sharma, P. P., and Kury, A. B.
(2010). A multilocus approach to harvestman (Arachnida: Opiliones)
phylogeny with emphasis on biogeography and the systematics of
Laniatores. Cladistics 26, 408–437. doi:10.1111/j.1096-0031.2009.
00296.x
Goodnight, C. J., and Goodnight, M. L. (1942). Phalangida from Mexico.
American Museum Novitates 1211,1–18.
Goodnight, C. J., and Goodnight, M. L. (1944). More Phalangida from
Mexico. American Museum Novitates 1249,1–13.
Goodnight, C. J., and Goodnight, M. L. (1945). Additional Phalangida from
Mexico. American Museum Novitates 1281,1–17.
Goodnight, C. J., and Goodnight, M. L. (1946). Additional studies of the
phalangid fauna of Mexico. American Museum Novitates 1310,1–17.
Goodnight, C. J., and Goodnight, M. L. (1947a). Studies of the phalangid
fauna of Trinidad. American Museum Novitates 1351,1–13.
Goodnight, C. J., and Goodnight, M. L. (1947b). Phalangida from Tropical
America. Fieldiana Zoology 32,1–58.
Goodnight, C. J., and Goodnight, M. L. (1953). The opilionid fauna
of Chiapas, Mexico, and adjacent areas (Arachnoidea, Opiliones).
American Museum Novitates 1610,1–81.
Goodnight, C. J., and Goodnight, M. L. (1954). The opilionid fauna of
an isolated volcano in southeastern Veracruz. Transactions of the
American Microscopical Society 73, 344–350. doi:10.2307/3223579
Goodnight, C. J., and Goodnight, M. L. (1967). Opilionids from Texas
caves (Opiliones, Phalangodidae). American Museum Novitates 2301,
1–8.
Goodnight, C. J., and Goodnight, M. L. (1971). Opilionids (Phalangida) of
the family Phalangodidae from Mexican caves. Bulletin of the Association
for Mexican Cave Studies 4, 33–45.
358 Invertebrate Systematics J. A. Cruz-López and O. F. Francke
Goodnight, C. J., and Goodnight, M. L. (1973). Opilionids (Phalangida) from
Mexican caves. Bulletin of the Association for Mexican Cave Studies 5,
83–96.
Goodnight, C. J., and Goodnight, M. L. (1977). Laniatores (Opiliones) of
the Yucatan Peninsula and Belize (British Honduras). Bulletin of the
Association for Mexican Cave Studies 6, 139–166.
Hall, T. A. (1999). BioEdit: a user-friendly biological sequence alignment
editor and analysis program for Windows 95/98/NT. Nucleic Acids
Symposium Series 41,95–98.
Ivanova, N. V., Zemlak, T. S., Hanner, R. H., and Hebert, P. D. N. (2007).
Universal primer cocktails for fish DNA barcoding. Molecular Ecology
Notes 7, 544–548. doi:10.1111/j.1471-8286.2007.01748.x
Katoh, S. (2013). MAFFT multiple sequence alignment software version 7:
improvements in performance and usability. Molecular Biology and
Evolution 30, 772–780. doi:10.1093/molbev/mst010
Kearse, M., Moir, R., Wilson, A., Stones-Havas, S., Cheung, M., Sturrock, S.,
Buxton, S., Cooper, S., Markowitz, S., Duran, C., Thierer, T., Ashton, T.,
Mentjies, P., and Drummond, A. (2012). Geneious Basic: an integrated
and extendable desktop software platform for the organization and
analysis of sequence data. Bioinformatics 28, 1647–1649. doi:10.1093/
bioinformatics/bts199
Kury, A. B. (1994). Rela¸cões filogenéticas entre as famílias de Laniatores
(Arachnida, Opiliones). In ‘Resumos do XX Congresso Brasileiro de
Zoologia, Rio de Janeiro, Brazil’. p. 72.
Kury, A. B. (1997). Os Stygnopsidae na filogenia de Gonyleptoidea, com
comentários sobre a biogeografia da superfamília. In ‘Resumos do Primer
encuentro de Aracnólogos del Cono Sur Montevideo, Uruguay’.
Abstracts: 30.
Kury, A. B. (2003). ‘Annotated catalogue of the Laniatores of the New
World’. Revista Ibérica de Aracnología, Volumen Especial Monográfico
1. (Grupo Ibérico de Aracnología (GIA) and Sociedad Entomológica
Aragonesa (SEA): Zaragoza, España.)
Kury, A. B. (2012). First report of the male of Zamora granulata
Roewer, 1928, with implications on the higher taxonomy of the
Zamorinae Kury, 1997 (Opiliones, Laniatores, Cranaidae). Zootaxa
3546,29–42.
Kury, A. B. (2014). Why does the Tricommatinae position bounce so much
within Laniatores? A cladistic analysis with description of a new family
of Gonyleptoidea (Opiliones, Laniatores). Zoological Journal of the
Linnean Society 172,1
–48. doi:10.1111/zoj.12165
Kury, A. B., and Cokendolpher, J. C. (2000). Opiliones. In ‘Biodiversidad,
taxonomía y biogeografía de artrópodos de México: hacía una síntesis
de su conocimiento, Volumen II’. (Eds J. Llorente-Bousquets,
J. E. González-Soriano and N. Papavero.) pp. 137–157. (Facultad de
Ciencias, UNAM, CONABIO and Bayer: Mexico City.)
Kury, A. B., and Medrano, M. (2016). Review of terminology for the outline
of dorsal scutum in Laniatores (Arachnida, Opiliones). Zootaxa 4097,
130–134. doi:10.11646/zootaxa.4097.1.9
Kury, A. B., and Pérez-González, A. (2002). A new family of Laniatores
from Northwestern South America (Arachnida, Opiliones). Revista
Ibérica de Aracnología 6,3–11.
Kury, A. B., and Villarreal, O. (2015). The prickly blade mapped: establishing
homologies and a chaetotaxy for macrosetae of penis ventral plate in
Gonyleptoidea (Arachnida, Opiliones, Laniatores). Zoological Journal
of the Linnean Society 174,1–46. doi:10.1111/zoj.12225
Lewis, P. O. (2001). A likelihood approach to estimating phylogeny from
discrete morphological character data. Systematic Biology 50, 913–925.
doi:10.1080/106351501753462876
Lian, W. G., Zhang, C., and Zhang, F. (2011). Review of the genus
Plistobunus Pocock, 1903, with description of a new species from
Hainan Island, China (Opiliones, Laniatores, Epedanidae). ZooKeys
112,39–52. doi:10.3897/zookeys.112.1110
Maddison,W.P.,andMaddison, D. R. (2015). Mesquite: a modularsystemfor
evolutionary analysis. Ver. 3.0.4. Available at http://mesquiteproject.org.
Martens, J. (1986). Die Grossgliederung der Opiliones und die evolution
der ordnung (Arachnida). In ‘Actas del X Congreso Internacional de
Aracnología’. (Ed. J. A. Barrientos.) pp. 289–310.
Mello-Leitão, C. F. (1926). Notas sobre Opiliones Laniatores sul-americanos.
Revista do Museu Paulista 14, 327–383.
Mello-Leitão, C. F. (1938). Considera¸cões sobre os Phalangodoidea Soer.
com descri¸cão de novas formas. Annaes da Academia Brasileira de
Sciencias 10, 135–145.
Mendes, A. C., and Kury, A. B. (2007). Stygnopsidae Sørensen, 1932.
In ‘Harvestmen: the Biology of Opiliones’. (Eds R. Pinto-da-Rocha,
G. Machado and G. Giribet.) pp. 232–233. (Harvard University Press:
Cambridge, MA.)
Miller, M. A., Pfeiffer, W., and Schwartz, T. (2010). Creating the CIPRES
Science Gateway for inference of large phylogenetic trees. In
‘Proceedings of the Gateway Computing Environments Workshop,
New Orleands, Louisiana, USA’. pp. 1–8.
Monjaraz-Ruedas, R., Francke, O. F., Cruz-López, J. A., and Santibáñez-
López, C. E. (2016). Annuli and setal patterns in the flagellum of female
micro-whipscorpions (Arachnida: Schizomida): hypotheses of homology
across an order. Zoologischer Anzeiger 263,118–134. doi:10.1016/
j.jcz.2016.05.003
Park, J.-K., and Ó Foighil, D. (2000). Sphaeriid and corbiculid clams
represent separate heterodont bivalve radiations into freshwater
environments. Molecular Phylogenetics and Evolution 14,75–88.
doi:10.1006/mpev.1999.0691
Pérez-González, A. (2006). Revisão Sistemática e Análise Filogenética
de Stygnommatidae (Arachnida: Opiliones: Laniatores). Ph.D. Thesis,
Museo Nacional, Universidad Federal do Rio de Janeiro, Brazil.
Pérez-González, A., and Kury, A. B. (2007). Kimulidae Pérez-González,
Kury and Alonso-Zarazaga, new name. In ‘Harvestmen: the Biology of
Opiliones’. (Eds R. Pinto-da-Rocha, G. Machado and G. Giribet.)
pp. 207–209. (Harvard University Press: Cambridge, MA.)
Pickard-Cambridge, F. O. (1905). Order Opiliones. In ‘Biologia centrali-
Americana, Vol. 2 Arachnida, Araneidea and Opiliones’. (Eds
F. D. Godman and O. Salvin.) pp. 545–560. (Natural History Museum:
London, UK.)
Pinto-da-Rocha, R., Benedetti, A. R., Vasconcelos, E. G., and Hara, M. R.
(2012). New systematic assignment in Gonyleptoidea (Arachnida,
Opiliones, Laniatores). ZooKeys 198,25–68. doi:10.3897/zookeys.198.
2337
Pinto-da-Rocha, R., Bragagnolo, C., Marques, F. P. L., and Junior, M. A.
(2014). Phylogeny of harvestmen family Gonyleptidae inferred from a
multilocus approach (Arachnida: Opiliones). Cladistics 30, 519–539.
doi:10.1111/cla.12065
Prendini, L., Weygoldt, P., and Wheeler, W. C. (2005). Systematic of
the Damon variegatus group of African whip spiders (Chelicerata:
Amblypygi): evidence from behaviour, morphology and DNA.
Organisms, Diversity & Evolution 5, 203–236. doi:10.1016/j.ode.2004.
12.004
Rambaut,A.,Suchard,M. A., Xie, D., and Drummond, A. J.(2014).TRACER
v. 1.6. Available at at http://beast.bio.ed.ac.uk/Tracer.
Rambla, M., and Juberthie, C. (1994). Opiliones. In ‘Encyclopaedia
Biospeologica’. (Eds C. Juberthie and V. Decu.) pp. 215–230. (Société
de Biospéologie: Netherlands.)
Roewer, C. F. (1912). Die familien der Assamiden und Phalangodiden
der Opiliones-Laniatores. Archiv für Naturgeschichte 78,1–242.
Roewer, C. F. (1915). 106 neue Opilioniden. Archiv für Naturgeschichte 81,
1–152.
Roewer, C. F. (1923). ‘Die Weberknechte der Erde. Systematische
Bearbeitung der bisher bekannten Opiliones.’(Gustav Fischer: Jena,
Germany)
Roewer, C. F. (1927). Weitere Weberknechte I. (1. Ergänzung der:
Weberknechte der Erde, 1923). Abhandlungen der Naturwissenschaftlichen
Verein zu Bremen 26,261–402.
Total evidence phylogeny of Stygnopsidae Invertebrate Systematics 359
Ronquist, F., and Huelsenbeck, J. P. (2003). MrBayes 3: Bayesian
phylogenetic inference under mixed models. Bioinformatics 19,
1572–1574. doi:10.1093/bioinformatics/btg180
Schwendinger, P. J., and Giribet, G. (2005). The systematics of the
Southeast Asian genus Fangensis Rambla (Opiliones: Cyphophthalmi:
Stylocellidae). Invertebrate Systematics 19, 297–323. doi:10.1071/
IS05023
Sharma, P. P., and Giribet, G. (2009). Sandokanid phylogeny based on eight
molecular markers –the evolution of a southeast Asian endemic family of
Laniatores (Arachnida, Opiliones). Molecular Phylogenetics and
Evolution 52, 432–447. doi:10.1016/j.ympev.2009.03.013
Sharma, P. P., and Giribet, G. (2011). The evolutionary and biogeographic
history of the armoured harvestmen –Laniatores phylogeny based on
ten molecular markers, with the description of two new families of
Opiliones (Arachnida). Invertebrate Systematics 25,106–142. doi:10.1071/
IS11002
Sharma, P. P., Prieto, C., and Giribet, G. (2011). A new family of Laniatores
(Arachnida: Opiliones) from the Afrotropics. Invertebrate Systematics
25, 143–154. doi:10.1071/IS11003
Sharma, P. P., and Giribet, G. (2014). A revised dated phylogeny of the
arachnid order Opiliones. Frontires in Genetics 5, 255. doi:10.3389/
fgene.2014.00255
Šilhavý, V. (1974). Cavernicolous Opilionids from Mexico. Subterranean
fauna of Mexico. Part. II. Quaderno della Accademia Nazionale dei
Lincei, Problemi Attuali di Scienza e Cultura 171, 175–194.
Šilhavý, V. (1977). Further cavernicolous opilionids from Mexico.
Subterranean fauna of Mexico. Part. III. Quaderno della Accademia
Nazionale dei Lincei, Problemi Attuali di Scienza e Cultura 171,
219–233.
Sørensen, W. E. (1884). Opiliones Laniatores (Gonyleptides W. S. Olim)
Musei Hauniensis. Naturhistorisk Tidsskrift 14, 555–646.
Sørensen, W. E. (1902). Gonyleptiden (Opiliones, Laniatores). Ergebnisse
der Hamburger Magalhaensischen Sammelreise 6,1–36.
Sørensen, W. E. (1932). Descriptiones Laniatorum (Arachnidorum
Opilionum Subordinis). (Opus posthumum recognovit et edidit Kai
L. Henriksen). Det Kongelige Danske Videnskabernes Selskabs skrifter –
Mémoires de l’Académie Royale des Sciences et des Lettres de Danemark.
Ser. 9 3, 197–422.
Stamatakis, A. (2014). RaxML version 8: a tool for phylogenetic analysis
and post-analysis of large phylogenies. Bionformatics 30, 1312–1313.
doi:10.1093/bioinformatics/btu033
Whiting, M. F., Carpenter, J. M., Wheeler, Q. D., and Wheeler, W. C. (1997).
The Strepsiptera problem: phylogeny of the holometabolous insect
orders inferred from 18S and 28S ribosomal DNA sequences and
morphology. Systematic Biology 46,1–68.
Wolff, J. O., Schönhofer, A. L., Martens, J., Wijnhoven, H., Taylor,C. K., and
Gorb, S. N. (2016). The evolution of pedipalps and glandular hairs as
predatory devices in harvestmen (Arachnida, Opiliones). Zoological
Journal of the Linnean Society 177, 558–601. doi:10.1111/zoj.12375
Xiong, B., and Kocher, T. D. (1991). Comparison of mitochondrial DNA
sequences of seven morphospecies of black flies (Diptera: Simuliidae).
Genome 34, 306–311. doi:10.1139/g91-050
Zhang, C., and Zhang, F. (2010). A new Tithaeus species from Hainan
Island, China (Arachnida, Opiliones, Laniatores, Epedanidae), with
a key to the Chinese species. ZooKeys 67,65–72. doi:10.3897/
zookeys.67.705
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360 Invertebrate Systematics J. A. Cruz-López and O. F. Francke
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