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Accepted by J. Rivera & G.J. Svenson: 11 Mar. 2014; published: 21 May 2014
ZOOTAXA
ISSN 1175-5326 (print edition)
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1175-5334
(online edition)
Copyright © 2014 Magnolia Press
Zootaxa 3797 (1): 207
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http://dx.doi.org/10.11646/zootaxa.3797.1.15
http://zoobank.org/urn:lsid:zoobank.org:pub:A03FDF55-8D04-49E6-ADC2-7684BCF7395D
A revision of Miobantia Giglio-Tos, 1917 (Mantodea: Thespidae, Miobantiinae),
with molecular association of dimorphic sexes and immature stages
MARCUS V. SCHERRER
Universidade Federal do Espírito Santo, Departamento de Ciências Biológicas, Av. Fernando Ferrari, 514, Goiabeiras, Vitória, ES,
Brazil, 29075-010. E-mail: marcusscherrer@gmail.com
Table of contents
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 208
Material and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .208
Results and Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .212
Taxonomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .214
Miobantia Giglio-Tos, 1917 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 214
Key to Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .220
Miobantia aptera Giglio-Tos, 1917 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221
Miobantia arctissima Scherrer, n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 231
Miobantia ciliata (Stål, 1860) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 232
Miobantia fuscata (Giglio-Tos, 1915) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 238
Miobantia immanis Scherrer, n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 246
Miobantia nordestina Scherrer, n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 249
Miobantia phryganea (Saussure, 1869) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 250
Miobantia rustica (Fabricius, 1781) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 251
Miobantia sulista Scherrer, n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .253
Concluding Remarks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 255
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 255
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 256
Appendix 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .258
Appendix 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .262
Appendix 3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .262
Appendix 4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .267
Abstract
The Neotropical praying mantis genus Miobantia Giglio-Tos, 1917 currently includes six species with a complex taxonomic
history. Although frequently found in the atlantic forest of Brazil, little is known about these species. Several obstacles make
it difficult identifying these mantids, including high levels of sexual dimorphism and both sexes known only for one of the
six currently known species. The taxonomic boundaries among the species of Miobantia are proposed in this work, through
a cladistic analysis using 99 morphological characters, and the analysis of sequence variation of approximately 700 bp of
the mithocondrial gene COI for association of dimorphic sexes and nymphs. Additionally, an investigation of intraspecific
variation is conducted based on a large number of specimens of M. fuscata (Giglio-Tos) in oder to choose the most relevant
features for the separation of the species to be used in taxonomic descriptions and identification keys. The male of M. aptera
Giglio-Tos, and the females of M. ciliata (Stål) and M. fuscata are described for the first time, and redescription of the males
of these and the female of M. aptera, are provided based on additional data. Miobantia nebulosa (Giglio-Tos) is herein re-
described (male genitalia included) and proposed as a junior synonym of M. rustica (Fabr.). Additionally, four new species
are here described, totaling nine species for the genus: M. immanis n. sp. (Espírito Santo, Brazil), M. arctissima n. sp. (Es-
pírito Santo, Brazil), M. sulista n. sp. (Santa Catarina and Rio Grande do Sul, Brazil; San Bernardino, Paraguay), and M.
nordestina n. sp. (Bahia, Brazil); both sexes of all new species are described, except for M. arctissima, which remain known
by males only. Identification keys and figures of diagnostic features are provided for both males and females of all valid
species. The species distribution is mapped according to literature data and 50 new records.
Key words: praying mantis, Miopteryginae, COI, DNA barcoding
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Introduction
Miobantia is a Neotropical praying mantis genus proposed by the Italian entomologist Ermanno Giglio-Tos in 1917
for two females named by him as Miobantia aptera. The species currently included in this genus, however, were
only recently assembled by Terra (1995), who reported the complicated taxonomic history of Miobantia.
Everything started when Rehn, in April of 1904, chose Miopteryx granadensis Saussure, 1870 as type species of
Miopteryx Saussure, 1969, but Kirby, by October of the same year, chose Miopteryx rustica (Fabricius, 1781) with
the same purpose. Giglio-Tos (1915), proposed Promiopteryx and transferred to it two species of Miopteryx,
including Miopteryx granadensis, which was chosen by him as type species of his new genus. Oblivious to the
procedure of Rehn (1904), Giglio-Tos kept the name Miopteryx for the not transferred species, adopting then
Miopteryx rustica as the type species of Saussure’s genus. From this point, a series of nomenclatural acts were
taken based on this complex scenario, resulting in the organization of the species initially grouped on Miopteryx
sensu Saussure in two different genera: Cnephomantis Rehn, 1920—treated as Antimiopteryx by Giglio-Tos
(1927), and Beier (1935, 1964, 1968)—and Promiopteryx Giglio-Tos, 1915. Terra (1995) deciphered this problem
and declared Cnephomantis as junior synonym of Miobantia, after noting high similarity between the species of
these groups. All this nomenclatural issue was recently summarized by Roger Roy (2013), also resulting in a new
name for the subfamily, the Miobantiinae, formerly known as Miopteryginae, once Miopteryx is no longer used.
The taxonomic treatment herein conforms with Terra’s and Roy’s nomenclatural analyses.
Miobantia is therefore currently composed of six species (Ehrmann, 2002; Otte & Spearman, 2005; Agudelo et
al., 2007): M. aptera Giglio-Tos, 1917, M. ciliata (Stål, 1860), M. fuscata (Giglio-Tos, 1915), M. nebulosa (Giglio-
Tos, 1915), M. phryganea (Saussure, 1869), and M. rustica (Fabricius, 1781). As most Thespidae, they are small
insects, with a generally brown coloration and wingless females. Although very active (often found in leaf litter or
on parts of plants close to the ground), and quite frequently found on collecting expeditions in the atlantic forest of
Brazil (pers. obs.), Miobantia is a poorly known mantid lineage, clearly in need of revision, e.g., as pointed out by
Rivera (2010). Despite the intensive activity that Miobantia has experienced in terms of nomenclature, little
information about its species has been added since their original descriptions. These descriptions date back to the
period between 1781 to 1917, lacking many important details, and are based primarily on variable characters such
as color and size. Similarly, the identification key for Antimiopteryx spp. provided by Giglio-Tos (1927), is based
solely on color patterns of the wings. It keys only males for five of the six currently described species of Miobantia,
but it is hardly useful due to the high intraspecific variation of the employed characters.
Another obstacle for the identification of Miobantia spp. is their high sexual dimorphism. Only two of six of
Miobantia, M. aptera and M. phryganea, are complemented by the description of females (Giglio-Tos, 1927).
However, the male of M. aptera is unknown, and the available information about the female M. phryganea lacks
details. Since the characters used in the literature to separate males of this genus are based on the wings, all of
which exhibit high sexual dimorphism, all efforts for sexual association up to now seem to have been not much
more than vague assumptions. The true range of variation between males and females of the same species, essential
for their association, remain unknown. This led to a situation where most Miobantia females were “traditionally”
classified as M. aptera. There is also no information in the literature about the oothecae and immature stages.
This work aims to contribute to the clarification of the taxonomic boundaries among species of Miobantia by
using cladistic analyses, and a molecular-based approach making use of DNA barcoding techniques to associate
conspecific males, females, and nymphs.
Material and Methods
Material examined. This work is based on 923 specimens and five oothecae of Miobantia, and another three
specimens from genera related to Miobantia, used in the phylogenetic analyses. The specimens analyzed belong to
the following institutions: The Natural History Museum, London, UK (NHM); Museu de Ciências e Tecnologia da
Pontifícia Universidade Católica do Rio Grande do Sul, Porto Alegre, Brazil (MCTP); Museu de Zoologia da
Universidade Estadual de Feira de Santana, Feira de Santana, Brazil (MZUEFS); Museu Nacional do Rio de
Janeiro, Rio de Janeiro, Brazil (QBUM); Universidade Federal do Espírito Santo, Vitória, Brazil (UFES); Museum
für Naturkunde, Berlin, Germany (ZMB).
Cladistics. Phylogenetic relationships among specimens were investigated as an aid for the taxonomic
delimitation of species. All examined Miobantia were separated in groups of specimens according to its
morphological pattern (morphospecies). Because of the high sexual dimorphism, only males were evaluated at this
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A REVISI ON OF MIOBANTIA GIGLIO-TOS
first stage. A representative specimen of each morphospecies as well as the types of Miobantia fuscata and M.
nebulosa were used in the cladistic analysis. Morphological analysis included 99 morphological characters, codified
as listed in Appendix 1. A cotype of M. rustica was codified through digital photographs. The holotype of Miobantia
aptera was not considered in this initial analysis because it is a female. The outgroups included one specimen of each of
the following genera (label data of the specimens in Appendix 2): Anamiopteryx Giglio-Tos (Pseudomiopteriginae),
Chloromiopteryx Giglio-Tos (Miobantiinae), and Eumusonia Giglio-Tos (Thespinae). This outgroup was chosen
aiming to comprise another genus of the same subfamily of Miobantia and other related subfamilies.
In this way, a matrix of 99 rows (characters) and 41 columns (taxa) was constructed. The analyses was made
under parsimony criterion through heuristic searches with TNT 1.1 (Goloboff et al., 2008b), using sectorial
searches and the algorithms Ratchet and Drift, with 1000 iterations for each search, as well as the Tree Fusing
method. Characters were downweighted according to their homoplasy (Goloboff et al., 2008a) through implied
weighting using values of k between 1 and 6 to explore multiple evolutionary scenarios. The generated trees were
visualized with WinClada 1.00.08 (Nixon, 2002). The support of the clades were tested in the program TNT by
symmetric resampling (Goloboff et al., 2003), with 1000 replications.
Preparation of genitalia. The male genitalia were investigated to assist separation of morphological patterns
and its characters included in phylogenetic analysis. The preparations of the phallic complex were conducted
according the classic methodology used in studies of praying mantises (e.g., Cerdá, 1993) which consists in
removing the genitalia with tweezers, after a brief period of submergence of the abdomen tip of the specimen in
water slightly heated. The phallic complex must then be placed in 10% KOH solution for about 30 minutes to
dissolve the associated musculature.
Species delimitation. Male specimens were sorted in species according to the clades supported by the
phylogenetic analysis, based on those that showed conspicuous discontinuities in the morphological patterns
compared to other clades, following the criterion of Darwin (1859). The new taxa were recognized as the species
delimited by this method that (1) were not clustered with any of the included types of Miobantia (i.e., of previously
named spp.), (2) did not match any of the published descriptions, and (3) did not match direct comparisons with the
digital images of the types of M. ciliata and M. phryganea, taxa not included in the cladistic analysis. Since M.
aptera was known only from a female, the identification of males for this species was performed subsequent to the
study aimed to associated males and females (described below). The holotype of M. aptera was then compared
with female specimens to determin the correct clade within the male only phylogenetic analysis that corresponded
to M. aptera.
Association of dimorphic sexes and immature stages. For the association of females and nymphs with their
corresponding conspecific males, a pairwise distance matrix was generated in order to find more genetically related
pairs of individuals. The molecular methodology was conducted by removing one midleg or hindleg from one to
four males of each morphospecies, covering the maximum possible variation, and those of the examined females
and nymphs. Only specimens from UFES were considered for this analysis; the material from other collections
were not included to preserve its integrity, as they are mostly very old and the state of conservation of their genetic
material is unknown. The legs were then subjected to the standard protocol for DNA extraction with NaCl/SDS/
Proteinase K (Bruford et al., 1992). The obtained products were used to amplify approximately 700 bp of the
mitochondrial gene Cytochrome Oxidase I (COI) by Polymerase Chain Reaction (PCR) with the primers LCO
1490 (5'-GGTCAACAAATCATAAAGATATTGG-3') and HCO 2198 (5'-TAAACTTCAGGGTGACCAAAAAAT
CA-3') (Folmer et al., 1994). The protocol used for PCR consists in temperatures of hot start at 94°C (1 min),
denature at 94°C (40 s), anneal at 45°C (40 s), extension at 72°C (1 min), and final extension at 72°C (5 min), with
42 cycles. Sequencing of DNA was performed by using a third party service, and the degree of saturation of its
nucleotide substitutions was observed with Dambe (Xia & Xie, 2001). The generated sequences were submitted to
GenBank (http://www.ncbi.nlm.nih.gov/genbank) with the accession numbers KJ160683–KJ160780.
In a first analysis, intra- and interspecific genetic distances were calculated only for males in MEGA5 (Tamura
et al., 2011) using the evolutionary model Kimura 2-parameter (Kimura, 1980). Then, the pairwise distances were
calculated among all sequenced males, females, and nymphs. The females or nymphs were associated with the
species of the male genetically less distant of them if these distance values of associations were conclusive in
comparison with those genetic distances found in intra- and interspecific analyses.
Although not all the examined males and females have been included in the molecular analyses, the achieved
associations by DNA data were strictly necessary to identify the expected limits of morphological variation among
the dimorphic sexes. From this point, a morphological investigation could be conducted with a reasonable level of
confidence for associating the remaining specimens.
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Intraspecific variability. A total of 867 specimens of Miobantia fuscata were considered in an additional
morphological and molecular investigation. These specimens were obtained from 29 localities in Espírito Santo,
southeastern Brazil, encompassing 19 cities. Morphological variation was discussed, and the genetic distance
calculated, between and within all morphospecies. The more stable characters are identified and selected to
produce the identification keys and the taxonomic comments of the description of each species of the genus. A total
of 60 male genitalia of M. fuscata were dissected and compared.
Measurements, terminology, and descriptions. Specimens were examined and measurements taken under Leica
MZ12.5 stereomicroscope with ocular micrometer. General morphological terminology follows Snodgrass (1935),
except for male genitalia terminology which follows La Greca (1954). The anterior, middle, and posterior leg,
coxa, trochanter, femur, and tibia, are all treated here preceded with the prefixes fore, mid, and hind-, respectively,
as for example in foreleg, and midtibia. Wing venation nomenclature follows Roy (1999), except by tegmina and
metathoracic wing, which are referred respectively as forewing and hindwing, and the addition of the forewing
veins M1 and M2 originated from the bifurcation of vein M, and hindwing veins CuA1 and CuA2 originated from
vein CuA. The main morphological structures for this work are pointed in the figures 4A, 11B, 12A, 14A, 16B,
16J, 17C, and 19A. Descriptions were generated with the aid of the program DELTA 1.04 (Dallwitz et al., 1999),
except for color descriptions. The identification keys were prepared using wing venation features in some couplets
even that intraspecific variation of such structures are known for Mantodea groups. However, the intraspecific
variation analysis herein revealed that abnormal venation patterns were very rarely repeated in the corresponding
wing of the other side of the body for the used features in the keys. Species descriptions, as well as the
measurements and ratios, represent the holotype; DELTA’s characters file is provided in Appendix 3. The head and
the male genitalia of M. fuscata, the wings and vertex of M. rustica, and the forelegs of M. ciliata, could not be
redescribed based on the respective type specimens because these structures were lacking or badly damaged. Also
most characters of the head and abdomen of M. ciliata could not be seen in the available photographs; the male
genitalia of this type could not be examined. In those particular cases, a separate description of such structures is
provided under the item “Complementary description,” based on another specimen. The genicular spines of the
femora and the apical spur of the tibia were not included in the count of total spines of foreleg structures in
descriptions. In the items “Distribution” and “Material Examined”, the countries and states are listed from north to
south, as standardized by Zanella et al. (2000). The entire list of the males, females, and nymphs identified for each
species as results of both molecular and morphological data is provided under the item “Material Examined” of
each species description.
Imaging and mapping. Photographs were taken using the extended-focus system EntoVision (GTVision,
Hagerstown, Maryland) including a Leica M16 APO A macroscope attached to a JVC KY–75U 3-CCD digital
video camera that feeds image data to a desktop computer through Cartograph software. The programs Archimed
5.3.1 and Gimp 2.6 were used to combine the stacks of images and finalize the resulting illustrations. The left
dorsal phallomeres were imaged from the dorsal perspective, unlike the ventral perspective of right dorsal, and
ventral phallomeres, to provide a better view of the anterior process, and of the dorsal face of the area between the
phalloid apophysis and the membranous lobe. Legends of photographic illustrations show information to allow
recognition of the particular specimens from which they were produced, including either the indication of holotype
status or the unique label data.
The geographical coordinates, when not available from label data, were obtained from geoLoc (http://
splink.cria.org.br/geoloc) and from the Global Gazetteer v. 2.2 (http://www.fallingrain.com/world/). Distribution
maps were generated using Quantum GIS 1.8.0 (http://www.qgis.org).
TABLE 1. Comparison of tree costs; tree length, consistency index (Ci), and retention index (Ri) are related to the strict
consensus tree.
KNº of trees fit Score Rearrangements Length Ci Ri
1 5 57.60 33.40 402,342,164 289 47 72
2 4 65.70 25.30 423,828,209 301 45 70
3 6 70.56 20.44 430,989,862 300 46 70
4 5 73.76 17.24 435,016,260 300 46 70
5 5 76.05 14.95 435,386,933 300 46 70
6 5 77.79 13.21 455,750,516 301 45 70
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A REVISI ON OF MIOBANTIA GIGLIO-TOS
FIGURE 1. Strict consensus tree of relationships between specimens of Miobantia resulted in the cladistic analysis using
implied weighting (k = 1). The numbers on branches are the supports of the clades measured with symmetrical resampling
method, showed as frequency differences (%). Clades with support below 50% are collapsed. The results of delimitation and
identification of species is provided in the right side of vertical bars.
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Results and Discussion
Phylogeny and species delimitation for males
Analysis of the data matrix (Appendix 4) using implied weighting with value of concavity of 1–6 produced the
trees with the attributes listed in Table 1. The symmetric resampling support analysis with the most parsimonious
trees of each concavity value presented the same topology, which is provided in Figure 1. The values of branch
support are for analysis with k = 1, that was chosen because of its lower length, and higher Ci and Ri.
The examination of the generated phylogenetic hypothesis combined with investigations of discontinuities of
morphological patterns among the different clades suggests the existence of eight species of Miobantia in the
cladograms, as shown in Figure 1 by the vertical bars on the right. The most useful characters to separate the males
of the different species are: the shape of vertex and frontal sclerite; the ocelli size and relative proximity; pronotal
shape; pattern of distribution of dark spots on wings, primarily on vein R, the forewing vein M1 and hindwing
CuA1 branching pattern; genital features, such as the general form and curvature or extension of the distal and
proximal processes of the ventral phallomere, and the anterior process, the phalloid apophysis, the left margin of
dorsal lamina, and a well sclerotized area between the phalloid apophysis and the membranous lobe of the left
dorsal phallomere. As in most Mantodea genera, the male genitalia was the most distinctive structure to
successfully separate species of Miobantia—even very similar species, such as M. rustica and M. ciliata can be
instantly distinguished with it.
Two different clades are recovered including types of previously named species of Miobantia. The clade 80%
support (Fig. 1) contains most of the codified specimens and includes the holotype of M. fuscata. In addition,
another clade contains both the M. nebulosa holotype and the M. rustica syntype (with 99% of support). The
morphological comparison shows no relevant differences between them, especially in the male genitalia, indicating
that these are the same species. As M. rustica is the oldest name, it is the senior synonym of M. nebulosa according
to the principle of priority, as stated in the article 23 of International Code of Zoological Nomenclature (ICZN,
2000). The sister group of this species in the cladogram comprises three specimens that fit the description of M.
ciliata and share distinctive features with the imaged holotype of M. ciliata, primarily because of the distinctive
forewing vein R that is mostly uniformly dark colored. Therefore, these specimens were identified as M. ciliata.
The well supported monophyly of the group comprising M. ciliata and M. rustica is due to the synapomorphies
which form an easily recognizable group, such as the forewing dark spots distinctly present on the longitudinal and
crossveins, the presence of darkish marks around the branches of vein M of forewing, the straight, short, and
laterally projected anterior process of the left dorsal phallomere, and the left portion of the dorsal lamina of the left
dorsal phallomere not projected, but uniformly curved. Examination of wings and the description of M. phryganea
indicated that this species should be assigned to this group, because it shares at least the wing characteristics cited
above. The other five species supported by the phylogeny did not fit the descriptions and did not present relevant
similarities with the males of previously known species of Miobantia.
However, after the association of males and females, detailed below, the females associated to specimens
MT00087, MT00195 and MT00051, grouped in a clade with support of 98%, proved to be conspecific with the
holotype of M. aptera. Consequently, such males must be considered as M. aptera. The four remaining species are
here interpreted as new to science, and described in the "Taxonomy" section, below. The synapomorphies of each
species are also discussed in that section.
TABLE 2. Intra- and interspecific genetic distance values found between males.
Species n
Minimum –Maximum
Intraspecific Distance
[Average] (%)
Minimum–Maximum Interspecific Distance [Average] (%)
M. aptera M. ciliata M. fuscata
M. aptera 4 0.4–8.9 [6.91] - - -
M. ciliata 4 0.2–1.4 [0.78] 16.1–14.7 [15.3] - -
M. fuscata 44 0.0–17.7 [12.53] 15.6–22.2 [18.6] 13.2–17.9 [15.8] -
M. immanis n.sp. 2 12.5 [12.5] 26.4–22.4 [24.2] 21.9–19.4 [20.6] 20.2–27.2 [23.7]
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Association of dimorphic sexes and immature stages, and species delimitation for females
The intra- and interspecific genetic distances found between males are summarized in Table 2. Analyses of
obtained sequences with DAMBE indicated start of saturation in transition mutation for genetic distances above
15%. The transversion mutations showed no saturation.
The pairwise matrix investigation associated one nymph to Miobantia immanis, four females and one nymph
to males posteriorly identified as M. aptera, two females to M. ciliata, and 29 females and 13 nymphs to M.
fuscata. All genetic distances values between females or nymphs and its respective less distant male were below
7.02%, with average of 1.05%. These association values were far below the maximum intraspecific (17.7%), and
minimum interspecific (13.2%) distances found, and were therefore considered conclusive. Males from MCTP,
MZUEFS, QBUM, and ZMB not included in this molecular analysis, as well as the male of M. arctissima from
UFES not sequenced because of the poor conservation of its genetic material, had no females or nymphs sequenced
since all of these were successfully associated with the males included in molecular analysis.
Non-sequenced females that proved to be conspecific with sequenced females by morphological comparison,
were also associated with the respective males molecularly associated to such females. In this way, another six
females and one nymph were identified as M. fuscata and three females as M. aptera, including the holotype of the
latter. Five oothecae and 28 nymphs of first instar which emerged from them could also be associated to M. fuscata
by including the progenitor females in the molecular analysis.
Morphological investigations of males and females of the species associated by the molecular methods
indicated that the vertex and the pronotum, although noticeably most swollen and robust in females, maintain
forms proportionally similar in both sexes. The same proved to occur with the general form of the frontal sclerite of
the head. Another structure here discovered to be an important taxonomic character is the supranal plate. It is very
short in males, making the visualization of most variation difficult; however, in females it is longer than in males,
and the variation of its shape can be used to successfully separate females of the different species. Additionally,
species with males that show the supranal plate slightly pointed were molecularly associated to females having the
sides of the supranal plate distinctly converging towards a subrounded apex. On the other hand, species with males
that show a supranal plate widely rounded, in most cases were related with females having the sides of the supranal
plate somewhat parallel, and rounded apex.
Morphological analysis focusing on these features conclusively associated one female with males from M.
immanis, three females with males from M. sulista, and one female with males of M. nordestina. In the case of M.
immanis, other features were also important to associate the dimorphic sexes, such as the relative distance from the
internal spine 12 of the forefemora, and the external spine 1 of the foretibiae to the other spines of its respective
sets, and the presence of a small tubercle over which raises the central ocellus. Such features were uniquely found
in specimens of M. immanis.
Although the association of males, females, and immature stages using molecular data was successful, it is
noted that the high values of genetic distances found for the molecular marker used may not be as effective for such
associations when the sample size is small, or when other species are not included for comparison. For example, if
the genetic distance between a female and its less distant male for the present data set showed values between
13.2% (minimum interspecific distance found) and 17.7% (maximum intraspecific distance found), any
conclusions based only on this information would be uncertain. In addition, the saturation level of transition
mutations presented by these DNA sequences reinforces the problem with high distances.
Most studies on DNA barcoding with COI sequences usually find intraspecific divergence values less than 2%,
whereas higher values are considered as indicative of interspecific associations (Hebert et al., 2003a).
Unfortunately, the levels of genetic divergence in Mantodea have not been extensively studied for any particular
marker, therefore there is no data available to compare with the levels found on Miobantia for COI. However, a
further investigation on genetic divergence levels of COI among animal kingdom of Hebert et al. (2003b)
demonstrated that when higher divergences are observed, these variants ordinarily occur as geographical isolates,
probably reflecting their origin in past episodes of gene pool fragmentation, revealing regional lineages. This
information seems to be in agreement with the divergences found in Miobantia. The pairs of males of Miobantia
fuscata most divergent in their COI sequences (15.6–17.7%) represents some of the most distant collect points for
males (Fig. 27C), more specifically the males of Pinheiros and Conceição do Castelo (ES, Brazil) in relation with
males of Itapemirim (ES, Brazil), presenting a average distance of about 310 Km. On other hand, the divergence
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levels between two males collected in Itapemirim was 0.4%. Under this perspective, the divergence of 12.5%
found between the two males of M. immanis can possibly be explained by the geographical distance of their collect
points (about 100 Km from Santa Tereza to Pinheiros, ES, Brazil). However, these biogeographical issues will be
extensively discussed in a ongoing work focusing on the divergences of COI sequences found between Miobantia
specimens along their distribution localities.
Taxono my
Miobantia Giglio-Tos, 1917
Miobantia Giglio-Tos, 1917a: 73–74; 1919: 62; 1927: 259; Beier, 1935: 6; 1964: 944; 1968: 9; Jantsch & Corseuil, 1988: 227;
Terra, 1995: 42–43; Ehrmann, 2002: 225; Dorneles et al., 2005: 224; Agudelo et al., 2007: 120; Ehrmann & Koçak, 2009:
14; Rivera, 2010: 52.
=Miopteryx Kirby, 1904: 274; Giglio-Tos, 1915: 137; 1919: 60 (non Saussure, 1869: 55).
=Cnephomantis Rehn, 1920: 225.
=Antimiopteryx Giglio-Tos, 1927: 210; Beier, 1935: 6; Beier, 1964: 944, 1968: 9.
Type species: Miobantia aptera Giglio-Tos, 1917, by monotypy.
Diagnosis. Body small, general coloration darkish brown to pale yellow, ventrally often lighter, with dark spots of
variable distribution; dimorphic sexes, with apterous females and fully winged males, often distinctly smaller than
females. Head with eyes rounded; frontal sclerite laterally wider than long, sides parallel to very slightly
converging towards lower margin; vertex somewhat straight to markedly convex, inconspicuously to distinctly
higher than imaginary line joining apex of eyes; vertex region between parietal suture and eye straight to slightly
convex or markedly convex forming distinct, low bulge (juxtaocular bulge). Pronotum about 3 times as long as
wide; lateral margin without denticulation in males, distinctly present in females; supracoxal dilatation faintly to
markedly pronounced; metazone about 1.6–2.0 times as long as prozone. Forecoxa longer than metazone;
forefemora with claw groove just basad femur middle, 4 discoidal, 4 external, 12 internal spines; foretibiae with 7
external spines, spine 1 distinctly away from spine 2, 9–11 internal spines, dorsal spines absent. General
pigmentation of wings highly variable, even intraspecifically, ranging from grayish to markedly brownish or
blackish; main longitudinal veins with dark spots, crossveins sometimes with dark spots, large or small; forewing
about same length or shorter than hindwing, reaching about half body length, its apex somewhat rounded; vein M
branched, forming veins M1 and M2; vein M1 branched or not; hindwing vein CuA branched, forming veins CuA1
and CuA2; vein CuA1 branched or not. Supranal plate apex pointed, with sides slightly converging towards apex,
or rounded.
Redescription. Body small (length from head to tip of abdomen 16.0–26.4 mm), females as large as males or
distinctly larger (Figs 2–10).
Head (Figs 11–13): Labrum slightly convex, lower margin rounded; clypeus about 2.0 times as wide as long,
sides distinctly converging towards lower margin; frontal sclerite wider than long, longitudinally very to
moderately short, sides parallel to inconspicuously converging towards upper margin, lower margin slightly
convex, upper edge uniformly convex, medially extended between toruli; antenna filiform, markedly longer in
males than females; flagellomeres in males with swollen apex, in females with apex not swollen, entirely thinner
than in males; region between each compound eye and toruli often with tiny tubercle; eyes rounded, protruding
head; ocelli in females vestigial, in males small to large, regularly distant or close to one another, placed over a
slight elevation; central ocellus about as elevated as, or distinctly more elevated than lateral ocellus, placed atop a
small, conical tubercle; lateral ocelli rounded or distinctly elliptical; vertex somewhat straight to markedly convex,
inconspicuously to distinctly higher than imaginary line joining apex of eyes; vertex region between parietal suture
and eye not or more elevated towards parietal suture, straight to slightly convex or markedly convex forming
distinct, low bulge (juxtaocular bulge); parietal suture lower portion markedly curved towards ocelli cuticle
elevation; vertex region between parietal sutures with lateral portion sometimes abruptly or gradually elevated
close to parietal suture.
Pronotum (Figs 14 & 15): About 3.0 times as long as wide, smooth, with dense pilosity mostly distally on
metazone; median keel more evident at metazone, in prozone often anteriorly inconspicuous; lateral margin
without denticulation in males, distinctly present in females, with density of distribution variable across species;
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supracoxal dilatation faintly to markedly pronounced; metazone about 1.6–2.0 times as long as prozone, apical
region with a pair of very low cuticle bulges.
Meso- and Metanotum: Posterior margin of mesonotum with a pair of triangular, densely pilose expansions;
meso- and metanotum with median longitudinal pilosity often higher than general thorax pilosity.
Legs (Fig. 16): Densely pilose in males, pilosity very short and more sparsely distributed in females. Forecoxa
longer than metazone, anterior edge with faint, longitudinal denticulation in females, denticulation absent or
inconspicuous in males; forefemora with anterior edge somewhat straight or very faintly convex, claw groove just
basad femur middle, 4 discoidal spines, 4 external spines, spines 1 and 2 distinctly close to each other, 12 internal
spines, crenulated internally to lateral spine lines; foretibiae with 7 external spines, spine 1 distinctly away from
spine 2, 9–11 internal spines, dorsal spines absent; foretarsus with basal tarsomere longer than remaining segments
combined.
FIGURE 2. Miobantia aptera Giglio-Tos, 1917, dorsal habitus. A, male MT00051; B, female from Bicuíba, Reserva Natural
Vale; C, holotype label (not to scale); D, female holotype.
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FIGURE 3. Miobantia ciliata (Stål, 1860), dorsal habitus. A, male MT00047; B, female collected in 26.XI-02.XII.2006; C,
male holotype (G. Lindberg phot.); D, male MT00033; E, holotype label (not to scale).
Wings (Figs 2–10): Apterous female; fully winged male. Forewing about same length or shorter than hindwing,
about half as long as body length, its apex somewhat rounded; costal area about same width of forewing base, or
slightly thinner; vein Sc somewhat parallel to vein R; vein R apically branched; vein M branched, forming veins
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M1 and M2; vein M1 branched or not; stigma long, very thin. Hindwing basally wide, apically rounded to
distinctly pointed, or apex abruptly interrupted; vein R not branched; vein M basally branched; vein CuA branched,
forming veins CuA1 and CuA2; vein CuA1 branched or not.
FIGURE 4. Miobantia fuscata (Giglio-Tos, 1915), dorsal habitus. A, male MT00030; B, male holotype; C, holotype label (not
to scale); D, female from Reserva Natural Vale, collected in 13.I.2011; E, male MT00013; F, male MT00070.
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FIGURE 5. Miobantia phryganea (Saussure, 1869), dorsal habitus, male syntype (P. Schwendinger phot.).
FIGURE 6. Miobantia rustica (Fabricius, 1781), dorsal habitus. A, male holotype of junior synonym M. nebulosa of Giglio-
Tos; B, label of the holotype of M. nebulosa (not to scale); C, male lectotype (G. Svenson phot., Copyright of The Natural
History Museum, London); D, lectotype label (G. Svenson phot., Copyright of The Natural History Museum, London; not to
scale).
Abdomen: Dorsally slightly compressed in males, cylindrical in females; supranal plate rounded or faintly
pointed, very short in males, longer in females, not overlaying ovipositor, sides somewhat parallel or distinctly
converging towards apex (Fig. 17); cercus regularly sized, circular in cross-section; stylus circular in cross-section.
Male Genitalia (Figs 19–26): Ventral phallomere broad; distal process thin, short to long, its apex rounded or
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pointed; left posterior corner with prominent expansion (proximal process), from slightly to markedly projected,
often long, curved, forming a slit with posterior margin of ventral phallomere. Right dorsal phallomere of regular
shape. Left dorsal phallomere with anterior process short to long; portion between anterior process and phalloid
apophysis triangularly expanded; area between phalloid apophysis and membranous lobe sometimes with well
sclerotized region; phalloid apophysis very short to very long, with basal portion projected to the back or left, then
curved to the right towards apex, apical portion sometimes bent, not or slightly sharp; apical process (titillator)
small, curved; membranous lobe wide to very slender; dorsal lamina posterior margin rounded or bent, sometimes
projected; articular process elongate, slender.
FIGURE 7. Miobantia immanis n. sp., dorsal habitus. A, male holotype; B, male MT00045; C, female allotype.
Color (Figs 2–17): Body generally darkish brown to pale yellow, ventrally often lighter, with many dark spots
of variable distribution. Head with ocelli margin dark in males, often with well pigmented transversal area between
eyes, covering the ocelli cuticle elevation, sometimes pigmentation faint between ocelli. Legs with dark spots often
densely distributed; forefemora sometimes with three, mid- and hinfemur sometimes with two, large transverse
dark marks; tibiae often with three transverse dark marks; forefemora and foretibiae with spines at apex; all
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tarsomeres with dark apex. Wings general pigmentation highly variable, even intraspecifically, from grayish to
markedly brownish, or blackish pigmented; main longitudinal veins with dark spots, crossveins sometimes also
with dark spots, the latter large or small.
Ootheca (Fig. 18): Known for only one species. See M. fuscata redescription.
Distribution. Neotropical (Fig. 27). Recorded from Brazil, Paraguay, and Argentina. The northernmost record
is for a male of M. nordestina, collected in Cachoeira da Fumaça, Feira de Santana, in Bahia state, Brazil
(10°28'43"S 40°12'27"W). The southernmost record corresponds to the types of M. rustica, collected in coastal
Patagonia (exact locality unknown).
FIGURE 8. Miobantia arctissima n. sp., dorsal habitus, male holotype.
Key to Species
Males
1. Forewing dark spots at vein R with marks of distinctly unequal sizes; forewing and hindwing costal margin apically with dis-
tinct bright marks (e.g., Figs 2A, 4A, 7A, 7B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
-. Forewing dark spots at vein R with marks of about same sizes, or vein R entirely or mostly uniformly dark; forewing and hind-
wing costal margin apically without distinct bright marks (e.g., Figs 3A, 5, 6A, 8, 9A, 10A) 4
2(1). Vertex inconspicuously more elevated than eyes (Fig. 12E); forewing vein M1 branched. Dorsal habitus as in Fig. 4 . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Miobantia fuscata (Giglio-Tos, 1915)
-. Vertex distinctly more elevated than eyes (e.g., Fig. 13A); forewing vein M1 not branched . . . . . . . . . . . . . . . . . . . . . . . . . . .3
3(2). Frontal sclerite longitudinally very short, more than 3.0 times as wide as long (Fig. 11A); central ocellus about as elevated as
lateral ocelli (Fig. 11A); forefemora with distance between internal spines 12 and 11 about same distance between spines 10
and 11 or slightly shorter (Fig. 16A); foretibiae external spine 1 away from spine 2 by space equivalent to one spine (Fig. 16J);
hindwing vein CuA1 not branched (Fig. 2); genitalia: apex of distal process of ventral phallomere pointed and lacking denticu-
lations, and proximal process distinctly projected, curved posterad (Fig. 19). Dorsal habitus as in Fig. 2 . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Miobantia aptera Giglio-Tos, 1917
-. Frontal sclerite longitudinally moderately short, less than 3.0 times as wide as long (Fig. 11E); central ocellus distinctly more
elevated than lateral ocellus (Fig. 11E); forefemora with distance between internal spines 12 and 11 distinctly longer than dis-
tance between spines 10 and 11 (Fig. 16D); foretibiae external spine 1 away from spine 2 by space equivalent to two spines
(Fig. 16I); hindwing vein CuA1 branched (Fig. 7); genitalia: apex of distal process of ventral phallomere rounded, exhibiting
well marked denticulations, and proximal process only slightly projected, not curved (Fig. 23) Dorsal habitus as in Fig. 7 . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Miobantia immanis Scherrer, n. sp.
4(1). Vertex somewhat straight or slightly convex (e.g., Figs 12H, 13D, 13F); forewing with crossveins of costal area without dark
spots (e.g., Figs 8–10) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
-. Vertex markedly convex (e.g., Figs 12C, 12G); forewing with crossveins of costal area dark spotted, even if inconspicuously
(e.g., Figs 3, 5, 6) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7
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5(4). Distance between ocelli much narrower than their width (Fig. 12H); forewing slightly shorter than hindwing (Fig. 8). Dorsal
habitus as in Fig. 8. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Miobantia arctissima Scherrer, n. sp.
-. Distance between ocelli as large as their width or larger (e.g., Figs 13D & 13F); forewing about same length than hindwing
(e.g., Figs 9 & 10) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6
6(5). Frontal sclerite longitudinally very short, more than 3.0 times as wide as long (Fig. 11G); ocelli medium-sized, about as wide
as pedicel apex (Fig. 11G); ventral phallomere of genitalia with distal process slightly long, about 2.0 times as long as wide,
basal dilatation present, its tip pointed, proximal process long with apex rounded (Fig. 25). Dorsal habitus as in Fig. 9 . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Miobantia sulista Scherrer, n. sp.
-. Frontal sclerite longitudinally moderately short, less than 3.0 times as wide as long (Fig. 11H); ocelli markedly large, wider
than pedicel apex (Fig. 11H); ventral phallomere of genitalia with distal process short, about as long as wide, basal dilatation
absent, its tip rounded, proximal process short with apex pointed (Fig. 26). Dorsal habitus as in Fig. 10 . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Miobantia nordestina Scherrer, n. sp.
7(4). Supracoxal dilatation very faintly pronounced, prozone middle width/supracoxal maximum greater than 0.80 (Fig. 14C); fore-
wing vein R entirely or mostly uniformly dark. Dorsal habitus as in Fig. 3 . . . . . . . . . . . . . . . . .Miobantia ciliata (Stål, 1860)
-. Supracoxal dilatation slightly pronounced, prozone middle width/supracoxal maximum dilatation between 0.76–0.80 (e.g.,
Fig. 14G); forewing vein R not darkened but entirely spotted (e.g., Figs 5 & 6) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8(7). Distance between ocelli slightly narrower than their width; forewing slightly shorter than hindwing; hindwing apex rounded
(Fig. 6). Dorsal habitus as in Fig. 6 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Miobantia rustica (Fabricius, 1781)
-. Distance between ocelli as large as their width or larger; forewing much shorter than hindwing; hindwing apex slightly pointed
(Fig. 5). Dorsal habitus as in Fig. 5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Miobantia phryganea (Saussure, 1869)
Females (except phryganea, rustica, and arctissima)
1. Supranal plate sides parallel, or nearly so, apex rounded (e.g., Figs 17C, 17D, 17H, 17I) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
-. Supranal plate sides distinctly converging towards subrounded apex (e.g., Figs 17E–17G) . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2(1). Central ocellus on a very small, low tubercle (Fig. 13B); forefemora: distance between internal spines 12 and 11 distinctly lon-
ger than distance between spines 10 and 11 (e.g., Fig. 16D); foretibiae: external spine 1 away from spine 2 by space equivalent
to two spines (e.g., Fig. 16I). Dorsal habitus as in Fig. 7C . . . . . . . . . . . . . . . . . . . . . . . . Miobantia immanis Scherrer, n. sp.
-. Central ocellus site without tubercle (e.g., Figs 12B & 12D); forefemora : distance between internal spines 12 and 11 about
same distance between spines 10 and 11 or slightly shorter (e.g., Fig. 16A); foretibiae: external spine 1 away from spine 2 by
space equivalent to one spine (e.g., Fig. 16J) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3(2). Vertex entirely markedly convex (Fig. 12D); supracoxal dilatation very faintly pronounced, prozone middle width/supracoxal
maximum greater than 0.80 (Fig. 14D); foretrochanter light brown, with few dark marks. Dorsal habitus as in Fig. 3B . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Miobantia ciliata (Stål, 1860)
-. Vertex laterally convex, slightly concave centrally (Fig. 12B); supracoxal dilatation slightly pronounced, prozone middle
width/supracoxal maximum dilatation between 0.76–0.80 (Fig. 14B); foretrochanter entirely dark, with only few evanescent
bright marks laterally. Dorsal habitus as in Fig. 2B & 2D . . . . . . . . . . . . . . . . . . . . . . . . . . Miobantia aptera Giglio-Tos, 1917
4(1). Vertex distinctly convex (Fig. 13E); frontal sclerite markedly short. Dorsal habitus as in Fig. 9B . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Miobantia sulista Scherrer, n. sp.
-. Vertex somewhat straight (e.g., Figs 12F & 13G); frontal sclerite moderately short . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3
4(1). Vertex region between parietal suture and eye inconspicuously convex, nearly straight (Fig. 12F); forefemora with internal
spine 12 distinctly shorter than spine 10 (e.g., Fig. 16C); pronotum with metazone slightly robust (Fig. 14F). Dorsal habitus as
in Fig. 4D . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Miobantia fuscata (Giglio-Tos, 1915)
-. Vertex region between parietal suture and eye slightly but distinctly convex (Fig. 13G); forefemora with internal spine 12
about same size of spine 10 (e.g., Fig. 16F); pronotum with metazone markedly robust (Fig. 15F). Dorsal habitus as in Fig. 10B
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Miobantia nordestina Scherrer, n. sp.
Miobantia aptera Giglio-Tos, 1917
(Figs. 2; 11B; 12A–B; 14A–B; 16A; 17C–D; 19; 27A)
Miobantia aptera Giglio-Tos, 1917a: 74; 1927: 259; Beier, 1935: 6; Jantsch & Corseuil, 1988: 227; Terra, 1995: 43; Ehrmann,
2002: 225; Dorneles et al., 2005: 224; Agudelo et al., 2007: 120; Ehrmann & Koçak, 2009: 14. Holotype ♀ (ZMB),
examined.
Redescription, male from Alfredo Chaves, Brazil. Length of body 17.2 mm from head to tip of abdomen, of
pronotum 4.1 mm, of metazone 2.8 mm, of forecoxa 3.4 mm, of forewing 15.5, of hindwing 17.2; maximum width
of head 3.1 mm, of prozone middle 1.1 mm, of supracoxal dilatation 1.5 mm; metazone minimum width 0.7 mm.
Head (Fig. 12A): Frontal sclerite longitudinally very short, more than 3.0 times as wide as long, upper edge
uniformly convex; small tubercle between each compound eye and toruli not pointed, represented by low cuticle
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elevation; ocelli slightly small, narrower than pedicel apex, distance between ocelli as large as their width or larger,
central ocellus about as elevated as lateral ocelli, lateral ocelli distinctly elliptical; vertex slightly convex, distinctly
higher than imaginary line joining apex of eyes, with region between parietal suture and eye more elevated towards
parietal suture, nearly straight; parietal suture faintly marked.
FIGURE 9. Miobantia sulista n. sp., dorsal habitus. A, male holotype; B, female allotype, paratype of Giglio-Tos’ description
of M. aptera; C, allotype label.
Pronotum (Fig. 14A): With moderately long pilosity, about as long as wing pilosity, supracoxal dilatation
slightly pronounced, prozone middle width/supracoxal maximum dilatation between 0.76–0.80; margin without
denticulation.
Legs (Figs 16A): Forecoxa markedly longer than metazone; forefemora internal spine 6 with shape and size
more similar to spine 5 than spines 2 and 4, distance between internal spines 12 and 11 about same distance
between spines 10 and 11 or slightly shorter, internal spine 12 about same size of spine 10; foretibiae external spine
1 away from spine 2 by space equivalent to one spine, external spine 2 slightly longer than spine 3, with 9 internal
spines or 10 internal spines.
Wings (Fig. 2A): Forewing slightly shorter than hindwing; vein M1 not branched; vein CuA1 branches slightly
sinuous. Hindwing apical angle moderately acute, vein CuA1 not branched, apex slightly short, slightly pointed.
Abdomen: Supranal plate very short, rounded.
Genitalia (Fig. 19): Ventral phallomere distal process short, with most marked sclerotization taking at least
half area of the process, denticulation absent, tip pointed, basal swelling absent; proximal process faintly
sclerotized, markedly projected, long, curved, distinctly turned posterad, with basal portion slightly wide, apex
rounded, slightly slender than base; slit between posterior margin of ventral phallomere and proximal process deep.
Left dorsal phallomere with anterior process slightly long, markedly large, markedly curved at about middle,
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oriented anterad, apex about as wide as middle; expanded portion between anterior process and phalloid apophysis
short, oriented anterad; well sclerotized area between phalloid apophysis and membranous lobe which is marked,
small, reaching anterior area of membranous lobe; phalloid apophysis regularly long, slender, its basal portion
projected to left one, subapically slightly bent, apically almost straight, not sharp; membranous lobe wide; dorsal
lamina posterior margin with left portion projected, expanded, uniformly curved.
FIGURE 10. Miobantia nordestina n. sp., dorsal habitus. A, male holotype; B, female allotype.
Color: Head, thorax except appendices, and abdomen, light brown, dorsally darker. Head: area between
mouthparts and toruli pale yellow, with few, very sparse, irregular dark marks, except by mandible apex, and
maxillary terminal segment dark; scape pale yellow, except by apical frontal dark mark; pedicel dark, with apical
margin pale yellow; flagellum entirely light brown; area between toruli and vertex darkish brown, with several
minute darker spots, mostly on transversal darker band between eyes covering the ocelli cuticle elevation, dark
band inconspicuous between ocelli; head dorsal region with irregular darkish marks, transversal dark band on it
apical area. Pronotum: dorsally entirely with very faint dark spots; margin pale yellow with moderately large dark
spots; ventrally with black median stripe on its posterior 0.30. Legs: light brownish to pale yellow, tarsomeres
apically darkish, tarsomere 1 centrally darkish; forecoxa with many dark spots; forecoxa apical margin medially
dark; foretrochanter entirely dark, with only few evanescent bright marks laterally; forefemora with moderate dark
spots, anteriorly with three large darkish marks, visible at the lateral and medial areas of femur, posteriorly with a
dark longitudinal mark from femur base to claw groove, apical region anteriorly with longitudinal dark mark;
foretibiae with few dark spots of irregular sizes; fore- and midtibia with three large transversal areas of dense
concentration of dark spots, one at each extremity and another at middle, mid- and hindfemur with two on its apical
half, hindfemur with area between dark transversal bands lighter than remaining leg parts. Mid- and hindlegs with
dense dark spots of irregular sizes. Wings: light brownish, regularly translucent; costal margins apically, and some
veins of forewing, with bright large marks, not restricted to vein limits; forewing dark spots present only on main
veins, crossveins without dark spots; vein R spotted, with marks of unequal sizes, spaced from each other by
2.0–3.0 times length of smallest spots; area around vein M branches without darkish marks.
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FIGURE 11. Miobantia, head from the frontal perspective of the frontal sclerite. A, M. aptera Giglio-Tos, 1917, female
holotype; B, M. ciliata (Stål, 1860), male MT00047; C, M. fuscata (Giglio-Tos, 1915), male MT00030; D, M. rustica
(Fabricius, 1781), male holotype of junior synonym M. nebulosa of Giglio-Tos; E, M. immanis n. sp., male holotype; F, M.
arctissima n. sp., male holotype; G, M. sulista n. sp., male holotype; H, M. nordestina n. sp., male holotype.
Female holotype. Length of body 21.4 mm from head to tip of abdomen, of pronotum 5.3 mm, of metazone
3.4 mm, of forecoxa 4.6 mm; maximum width of head 4.2 mm, of prozone middle 1.6 mm, of supracoxal dilatation
2.1 mm; metazone minimum width 1.1 mm. General morphology and coloration similar to male, but larger, stouter
than described male; head with vertex laterally convex, centrally slightly concave (Figs 11A & 12B); ocelli
vestigial; pronotum with distinct denticulation along entire lateral margin, more pronounced posteriorly, metazone
slightly more robust (Fig. 14B); forecoxa anterior edge with median distinct denticulation; foretibiae with 9
internal spines; apterous; supranal plate slightly longer than males, sides somewhat parallel (Fig. 17D); abdomen
with several dark spots, irregular dark marks.
Immature Stages. Nymph, immature male, unknown instar. Length of body11.1 mm; antenna 7.4 mm. General
morphology and coloration as in adults, except by flagellum basally stout, becoming very slender towards apex;
ocelli vestigial; foretibiae with external spines 1 and 3 slightly lower than the others, spine 2 markedly higher than
spines 4–6, spine 7 more than 2.0 times as high as spine 2. Wings vestigial. Cercus basally stout, very slender
towards apex. Subgenital plate with stylus present.
Variability, male. Examinated males very similar to holotype, except by head with parietal suture line
uniformly curved; forewing veins CuA-a and CuA-b1 nearly straight. General body dark marks and spots high
variable on size and intensity; wings with general coloration slightly variable, male from MN with forewing with
evanescent darkish brown marks between veins, male from Sooretama with wings distinctly dark. This specimen
also with general body coloration with pale yellow tint, dark marks distinctly more marked than others specimens.
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FIGURE 12. Miobantia, head from the frontal perspective of the ocelli. M. aptera Giglio-Tos, 1917: A, male MT00087; B,
female holotype. M. ciliata (Stål, 1860): C, male MT00047; D, female collected in 26.XI-02.XII.2006. M. fuscata (Giglio-Tos,
1915): E, male MT00030; F, female from Reserva Natural Vale, collected in 13.I.2011. M. rustica (Fabricius, 1781): G, male
holotype of junior synonym M. nebulosa of Giglio-Tos. M. arctissima n. sp.: H, male holotype.
Female. Other examined females very similar to holotype, except by foretibiae with 10 internal spines. Female
from Sooretama general body coloration with pale yellow tint, dark marks distinctly more marked; female from
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Vitória with general body coloration with dark green tint when observed alive; after drying with same coloration of
holotype, dark marks slightly more marked; females from Guarapari with general body coloration more similar to
holotype, with dark marks more rare and faint than other collected females.
FIGURE 13. Miobantia, head from the frontal perspective of the ocelli. M. immanis n. sp.: A, male holotype; B, female
allotype; M. sulista n. sp.: C, male holotype; D, female allotype; M. nordestina n. sp.: E, male holotype; F, female allotype.
Comments. Miobantia aptera was described by Giglio-Tos (1917a) as single species of the genus based on
two females. However, the holotype differs from the paratype by having the frontal sclerite slightly wider, eyes less
protruding, vertex laterally convex, centrally slightly concave (vs. entirely slightly convex), metazone of pronotum
distinctly longer and more slender, and supranal plate rounded (vs. slightly pointed). Based on such differences, the
types were considered non-conspecific specimens. The sex association procedure detailed above resulted in the
identification of morphological correspondence between the paratype of M. aptera, as well as other two females,
and the males of M. sulista. Therefore, the paratype of M. aptera was taken as paratype of this new species.
Males of M. aptera are most similar to males of M. fuscata and M. immanis based on the presence of forewing
dark spots at vein R, with marks of distinctly unequal sizes, and fore- and hindwings costal margin apically with
distinct bright marks. However, it can be easily separated from these two species by the foretrochanter entirely dark
with only few evanescent bright marks laterally, and the hindwing with vein CuA1 not branched. Although the
examination of male genitalia can successfully separate it from M. immanis based on the distal process of ventral
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phallomere with slender apex and proximal process long and curved, it can also allow the misidentification of it as
M. fuscata. In this case, M. aptera can be isolated by the distal process of ventral phallomere with denticulation
absent, its tip pointed, most marked sclerotization spotted at least the posterior half of the process, and left dorsal
phallomere with well sclerotized area between phalloid apophysis and membranous lobe extended, reaching
anterior area of membranous lobe.
FIGURE 14. Miobantia, dorsal perspective of the pronotum. M. aptera Giglio-Tos, 1917: A, male from Mata Pico do
Eldorado, T3; B, female holotype. M. ciliata (Stål, 1860): C, male MT00043; D, female collected in 26.XI-02.XII.2006. M.
fuscata (Giglio-Tos, 1915): E, male MT00030; F, female from Reserva Natural Vale, collected in 13.I.2011. M. rustica
(Fabricius, 1781): G, M. rustica, male holotype of junior synonym M. nebulosa of Giglio-Tos. M. arctissima n. sp.: H, male
holotype.
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FIGURE 15. Miobantia, dorsal perspective of the pronotum. M. immanis n. sp.: A, female allotype; B, male holotype; M.
sulista n. sp.: C, male holotype; D, female allotype; M. nordestina n. sp.: E, male holotype; F, female allotype.
The females are most likely to be mistaken for M. ciliata due to have the combination of the central ocellus site
without tubercle, the forefemora with distance between internal spines 12 and 11 about same distance between
spines 10 and 11 or slightly shorter, the foretibiae external spine 1 away from spine 2 by space equivalent to one
spine, and the supranal plate with sides parallel and rounded apex. However, females of M. aptera can be readily
separated from this species by having the vertex laterally convex, centrally slightly concave, the supracoxal
dilatation slightly pronounced, prozone middle width/supracoxal maximum dilatation between 0.76–0.80, the
foretrochanter entirely dark with only few evanescent bright marks laterally, and the hindfemur with area between
the two transverse dark bands lighter than remaining leg parts.
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FIGURE 16. Miobantia, structures of the foreleg. A–G, forefemur from medial-posterior perspective: A, M. aptera Giglio-
Tos, 1917, male MT00051; B, M. ciliata (Stål, 1860), male MT00047; C, M. fuscata (Giglio-Tos, 1915), male MT00030; D, M.
immanis n. sp., male holotype; E, M. arctissima n. sp., male holotype; F, M. sulista n. sp., paratype from Porto Alegre; G, M.
nordestina n. sp., male holotype. H, forefemur from medial perspective, M. rustica (Fabricius, 1781), male holotype of junior
synonym M. nebulosa of Giglio-Tos; I–J, foretibia from lateral perspective: I, M. immanis n. sp., male holotype; J, M.
nordestina n. sp., male holotype.
Distribution (Fig. 27A). Brazil. Recorded from Rio de Janeiro (type locality), and eight localities of Espírito
Santo: from Reserva Biológica de Sooretama, Sooretama (19°02'55"S 40°08'48"W); from Fazenda Juliberto Stur,
Pancas (19°12'54.8"S 40°47'52.5"W); from Reserva Natural Vale, Linhares (19°09'06"S 40°04'14"W); from
Fazenda Paulo Seick, Santa Maria de Jetibá (20°02'31.1"S 40°41'51.3"W); from Mata Pico do Eldorado, Domingos
Martins (20°22'17"S 40°39'29"W); from Reserva Biológica de Duas Bocas, Cariacica (20°16'21"S 40°28'40"W);
Universidade Federal do Espírito Santo campus Goiabeiras, Vitória (20°16'29.49"S 40°18'16.87"W); from Reserva
Natural Oiutrem, Matilde, Alfredo Chaves (20°33'S 40°48'W). Agudelo, Lombardo and Jantsch (2007) indicated
the occurrence of this species also in the states of Rio de Janeiro, Paraná and Rio Grande do Sul, in Brazil, based on
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a previous study of Jantsch (1999). However, two females identified by such author as M. aptera and included in
the original distribution record made by him in 1999 (registered for São Borja and Garabi, from Rio Grande do
Sul), was examined and taken as par types of M. sulista. Although these occurrence records were based also on
other specimens, these misidentifications suggest the need for more detailed investigations of such data. The other
specimens examined by Jantsch (1999) were recorded from the locality of Represa Rio Grande, Rio de Janeiro
(around 22°55'S 43°26W); the locality of Morretes, Paraná (around 25°28'S 48°49'W); and Porto Alegre, Rio
Grande do Sul (around 30°02'S 51°12'W). The distribution record for Paraguay (Agudelo et al. 2007, Ehrmann
2002) was based on an original paratype included in Giglio-Tos’ description, now the allotype of M. sulista.
FIGURE 17. Miobantia, supranal plate from the dorsal perspective. A, M. ciliata (Stål, 1860), male MT00043; B, M.
nordestina n. sp., male holotype; C, M. aptera Giglio-Tos, 1917, female from Bicuíba, Reserva Natural Vale; D, M. aptera,
female holotype; E, M. fuscata (Giglio-Tos, 1915), female from Reserva Natural Vale, Trilha atrás do viveiro de mudas, point
1; F, M. sulista n. sp., female allotype; G, M. nordestina n. sp., female allotype; H, M. ciliata (Stål, 1860), female from Alfredo
Chaves; I, M. immanis n. sp., female allotype.
FIGURE 18. Miobantia fuscata (Giglio-Tos, 1915), ootheca laid by female collected at Reserva Biológica Córrego do Veado,
Pinheiros. A, lateral view; B, dorsal view.
Material Examined. 19 ♂♂, 7 ♀♀, 1 nymph. Holotype: ♀ from BRAZIL, Rio de Janeiro, Mus. Berlin,
Miobantia aptera Giglio-Tos, E. Giglio-Tos det., Holotypus Nr. (ZMB). Described male: ♂ from BRAZIL,
Espírito Santo, Alfredo Chaves, Matilde, Reserva Natural Oiutrem, Malaise, COAzevedo et al. leg. [specimen
MT00051 in the cladistic analysis] (UFES). Other specimens: BRAZIL: 1 ♀ from Espírito Santo, Sooretama,
Reserva Biológica de Sooretama, 11-18.XI.2011, Malaise, COAzevedo et al. leg. (UFES); 1 ♂ same data except
Reserva Biológica de Sooretama, Trilha Ana - Malaise 6, 06-14.XII.2011, Malaise, MTTavares et al. leg.
[MT00195] (UFES); 1 ♂ from Espírito Santo, Pancas, Fazenda Juliberto Stur, Área 2, point T7, 24-31.I.2003,
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Malaise, MTTavares, COAzevedo et al. leg. (UFES); 1 nymph from Espírito Santo, Linhares, Reserva Natural
Vale, next to the labs, 13.I.2011, lighting trap, MVScherrer & MCCarreiro leg. (UFES); 1 ♀ same data except
Reserva Natural Vale, Bicuiba, Trilha 1, Pt.02, 29.V-8.VI.2010, Malaise, APAguiar et al. leg.; 1 ♀ from Espírito
Santo, Santa Maria de Jetibá, Fazenda Paulo Seick, Área 1, Bosque 2, 06-13.XII.2002, Malaise, MTTavares et al.
leg. (UFES); 1 ♂ from Espírito Santo, Domingos Martins, Mata Pico do Eldorado, point B2, 03-10.XII.2004,
Malaise, MTTavares et al. leg. (UFES); 3 ♂ ♂ same data except Mata Pico do Eldorado, T2; 1 ♂ same data except
Mata Pico do Eldorado, T3; 1 ♂ same data except Mata Pico do Eldorado, T7; 1 ♂ same data except Mata Pico do
Eldorado, T8; 1 ♂ same data except Mata Pico do Eldorado, B3, 26.XI-03.XII.2004; 1 ♂ same data except Mata
Pico do Eldorado, B4; 1 ♂ same data except Mata Pico do Eldorado, T2 [MT00087]; 1 ♂ same data except Mata
Pico do Eldorado, T3; 1 ♂ same data except Mata Pico do Eldorado, T8; 2 ♀ ♀ from Espírito Santo, Cariacica,
Reserva Biológica de Duas Bocas, Pau Amarelo, 21-27.X.2005, Malaise, APAguiar et al. leg. (UFES); 1 ♂ same
data except Pau Amarelo, point 18, 21-30.X.2005; 1 ♂ same data except Pau Amarelo, point 21, 26.X-05.XI.2005;
1 ♀ from Espírito Santo, Vitória, UFES (Goiabeiras), Prédio da Botânica, 26.IV.2010, 20:10 h, manual, JTCunha
leg. (UFES); 1 ♂ from Espírito Santo, Alfredo Chaves, Matilde, Reserva Natural Oiutrem, Malaise, COAzevedo et
al. leg. (UFES); 1 ♂ from [locality missing] “59.2 = 59.1”, Nº58/Proc.702, Miobantia sp., Agudelo det. 02.XI.2010
(QBUM).
Miobantia arctissima Scherrer, n. sp.
(Figs. 8; 11F; 12H; 14H; 16E; 24; 27F)
Description, male holotype. Length of body 17.1 mm from head to tip of abdomen, of pronotum 4.1 mm, of
metazone 2.6 mm, of forecoxa 3.6 mm; of forewing 18.8; of hindwing 19.6; maximum width of head 3.4 mm,
prozone middle 1.3 mm, supracoxal dilatation 1.7 mm; metazone minimum width 0.8 mm.
Head (Figs 11F & 12H): Frontal sclerite longitudinally very short, more than 3.0 times as wide as long; small
tubercle between each compound eye and toruli not pointed, represented by a low cuticle elevation; ocelli markedly
large, wider than pedicel apex, distance between ocelli much narrower than their width, central ocellus about as
elevated as lateral ocelli, lateral ocelli distinctly elliptical; vertex somewhat straight, distinctly higher than
imaginary line joining apex of eyes, with region between parietal suture and eye more elevated towards parietal
suture, nearly straight; parietal suture faintly marked; vertex region between parietal sutures laterally gradually
elevated close to parietal suture, centrally slightly concave.
Pronotum (Fig. 14H): With moderately long pilosity, about as long as wing pilosity, supracoxal dilatation
regularly sized, prozone middle width/supracoxal maximum dilatation between 0.71–0.75; margin without
denticulation.
Legs (Fig. 16E): Forecoxa markedly longer than metazone; forefemora internal spine 6 with shape and size more
similar to spine 5 than spines 2 and 4, distance between internal spines 12 and 11 about same distance between
spines 10 and 11 or slightly shorter, internal spine 12 about same size of spine 10; foretibiae external spine 1 away
from spine 2 by space equivalent to one spine, external spine 2 slightly longer than spine 3, with 9 internal spines.
Wings (Fig. 8): Forewing slightly shorter than hindwing; vein M1 branched; vein CuA1 branches markedly
sinuous. Hindwing apical angle moderately acute, vein CuA1 branched, apex slightly short, slightly pointed.
Abdomen: Supranal plate very short, rounded.
Genitalia (Fig. 24): Ventral phallomere distal process short, with most marked sclerotization taking entire
process, denticulation markedly present, tip wide, rounded, basal swelling absent; proximal process markedly
sclerotized, markedly projected, long, somewhat straight, turned diagonally posterad lateral portion, with basal
portion slightly wide, apex rounded; slit between posterior margin of ventral phallomere and proximal process
deep. Left dorsal phallomere with anterior process very long, moderately large, markedly curved at about middle,
oriented anterad, apex about as wide as middle; expanded portion between anterior process and phalloid apophysis
short, oriented anterad; well sclerotized area between phalloid apophysis and membranous lobe which is marked,
large, wide; phalloid apophysis regularly long, slender, its basal portion projected to left one, subapically markedly
bent, apically concave, slightly sharp; membranous lobe slender; dorsal lamina posterior margin with left portion
projected, expanded, uniformly curved.
Color. Body entirely pale yellow. Head: area between mouthparts and toruli with few, irregular dark spots,
except by mandible apex, and maxillary terminal segment dark; scape pale yellow, except by apical frontal dark
mark; pedicel dark, with margins pale yellow; flagellum entirely light brown; area between toruli and vertex with
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several minute light brownish spots, mostly on transversal darker band between eyes covering the ocelli cuticle
elevation; head dorsal region with irregular darkish marks, transversal dark band on it apical area. Pronotum: with
few, very sparse, small dark spots, mostly laterally; margin with slightly larger, sparse dark spots; ventrally with
black median stripe on its posterior half. Legs: forecoxa entirely, trochanter mostly medially with regularly sized
dark spots; forefemora anterior edge with 4 faint dark spots along its length, claw groove, area next to internal
spines with irregular, large darkish marks; fore- and midtibia with three large transversal areas of dense
concentration of dark spots, one at each extremity and another at middle. Hindfemur with two faint transversal dark
bands on its apical half; tarsomeres apically darkish, tarsomere 1 centrally darkish. Wings: light brown, forewing
cells, hindwing apical cells with central slightly brownish mark, very translucent; forewing dark spots present only
on main veins, crossveins without dark spots; vein R spotted, with marks distinctly of about same sizes, spaced
from each other by 2.0–3.0 times length of smallest spots; area around vein M branches without darkish marks.
Female. Unknown.
Variability. Unknown.
Comments. This species is most similar to M. nordestina and M. sulista based on the very translucent wings,
with costal margins of both wings without bright marks, costal area of forewing without dark spots, forewing vein
R with dark spots of about same sizes, with vein M1 branched, and hindwing vein CuA1 also branched. Miobantia
arctissima can however be instantly separated from the males of these species by having the characteristic ocelli
markedly large, wider than pedicel apex and the space between ocelli much narrower than their width.
Additionally, the combination of the following features of the male genitalia is also diagnostic for separating M.
arctissima from these species: the ventral phallomere distal process short, with most marked sclerotization taking
entire process and denticulation markedly present, its tip wide and rounded, and with basal swelling absent; and the
proximal process markedly projected and long, somewhat straight but turned diagonally posterad lateral portion,
with basal portion slightly wide, the apex rounded, with the slit between posterior margin of the ventral phallomere
and the proximal process deep.
Distribution (Fig. 27F). Brazil. Recorded only from the type locality: Ribeirão do Meio, in Conceição do
Castelo, Espírito Santo (around 20°21'S 41°14'W).
Material Examined. Holotype: ♂ from BRAZIL, Espírito Santo, Conceição do Castelo, Propriedade Ribeirão
do Meio, trilha ao lado da lagoa, 17-24.III.2007, Malaise, APAguiar et al. leg. [specimen MT00063 in the cladistic
analysis] (UFES).
Etymology. From the Latin word arctissima, meaning “very close”, in reference to the ocelli very close to one
another in this species in comparison with the other species of this genus.
Miobantia ciliata (Stål, 1860)
(Figs. 3; 11B; 12C–D; 14C–D; 16B; 17H; 20; 27B)
Mantis ciliata Stål, 1860: 313. Holotype ♂ (Swedish Museum of Natural History, Stockholm, Sweden - NHRS), examined
through digital images.
Miopteryx ciliata: Saussure, 1871: 111–112; Kirby, 1904: 274; Chopard, 1913: 761; Giglio-Tos, 1917b:154.
Parastagmatoptera ciliata: Kirby, 1904: 298; Rehn, 1911: 10.
Antimiopteryx ciliata: Giglio-Tos, 1927: 211; Sjöstedt, 1930: 7; Beier: 1935: 7.
Miobantia ciliata: Jantsch & Corseuil, 1988: 228; Terra, 1995: 43; Ehrmann, 2002: 225; Agudelo et al., 2007: 120; Ehrmann &
Koçak, 2009: 14.
Redescription, male holotype. Length of body about 22.6 mm from head to tip of abdomen.
Head: Ocelli medium-sized, about as wide as pedicel apex, central ocellus about as elevated as lateral ocelli,
lateral ocelli distinctly elliptical; vertex markedly convex, distinctly higher than imaginary line joining apex of
eyes, with region between parietal suture and eye more elevated towards parietal suture.
Pronotum (Fig. 14C): Supracoxal dilatation very faintly pronounced, prozone middle width/supracoxal
maximum greater than 0.80; margin without denticulation.
Wings (Fig. 3C): Forewing slightly shorter than hindwing; vein M1 branched; vein CuA1 branches slightly
sinuous. Hindwing apical angle moderately acute, vein CuA1 branched, apex slightly short, slightly pointed.
Color: Body entirely light brown, with small dark spots. Legs: light brownish, with several dark spots; Mid-
and hindlegs pale yellow, with several dark spots of irregular sizes, without transversal dark bands. Wings: light
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brown, very translucent; forewing main longitudinal veins with dark spots present, crossveins of discoidal area also
with few spots, of costal area without spots; vein R basal 0.6 entirely dark colored; area around vein M branches
with one larger darkish mark; hindwing main longitudinal veins apically dark spotted.
FIGURE 19. Miobantia aptera Giglio-Tos, 1917, male genitalia, right dorsal and ventral phallomeres from ventral
perspective, and left dorsal phallomere from dorsal perspective. A, MT00051; B, male from Pancas; C, male from Reserva
Biológica de Duas Bocas, Pau Amarelo, point 21; D, MT00195; E, male at MN. Images –.1, right dorsal phallomere; .2, ventral
phallomere; .3, left dorsal phallomere; .4, phalloid apophysis.
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FIGURE 20. Miobantia ciliata (Stål, 1860), male genitalia, right dorsal and ventral phallomeres from ventral perspective, and
left dorsal phallomere from dorsal perspective. A, MT00033; B, MT00047; C, MT00043. Images –.1, ventral phallomere; .2,
left dorsal phallomere.
Complementary description, male from Alfredo Chaves, Brazil. Length of body 19.1 mm from head to tip of
abdomen, of pronotum 3.8 mm, of metazone 2.44 mm, of forecoxa 3.6 mm, of forewing 17.6, of hindwing 19.1;
maximum width of head 3.2 mm, prozone middle 1.2 mm, supracoxal dilatation 1.4 mm; metazone minimum
width 0.9 mm.
Head (Figs 11B & 12C): Frontal sclerite longitudinally very short, more than 3.0 times as wide as long, upper
edge uniformly convex; small tubercle between each compound eye and toruli not pointed, represented by low
cuticle elevation; ocelli medium-sized, about as wide as pedicel apex, distance between ocelli as large as their
width or larger, central ocellus about as elevated as lateral ocelli, lateral ocelli distinctly elliptical; vertex markedly
convex, distinctly higher than imaginary line joining apex of eyes, with region between parietal suture and eye
more elevated towards parietal suture, nearly straight; parietal suture faintly marked.
Legs (Fig. 16B): Forecoxae markedly longer than metazone; forefemora internal spine 6 with shape and size
more similar to spine 5 than spines 2 and 4, distance between internal spines 12 and 11 about same distance
between spines 10 and 11 or slightly shorter, internal spine 12 about same size of spine 10; foretibiae external spine
1 away from spine 2 by space equivalent to one spine, external spine 2 slightly longer than spine 3, with 9 internal
spines.
Abdomen (e.g., Fig. 17A): Supranal plate very short, rounded.
Genitalia (Fig. 20B): Ventral phallomere distal process slightly long, with most marked sclerotization taking at
least half area of the process, denticulation absent, tip pointed, basal swelling absent; proximal process faintly
sclerotized, markedly projected, short, curved, turned diagonally posterad central portion, with basal portion very
wide, apex with slightly long, slender point; slit between posterior margin of ventral phallomere and proximal
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process deep. Left dorsal phallomere with anterior process short, markedly large, straight, oriented to left, apex
about as wide as middle; expanded portion between anterior process and phalloid apophysis short, oriented anterad;
well sclerotized area between phalloid apophysis and membranous lobe which is evanescent; phalloid apophysis
regularly short, moderately stout, its basal portion projected to left one, subapically markedly bent, apically almost
straight, not sharp; membranous lobe wide; dorsal lamina posterior margin with left portion not projected,
uniformly curved.
Color: Body entirely light brown to pale yellow, with several, small dark spots. Head: area between mouthparts
and toruli pale yellow, with few dark spots, except by mandible apex, and maxillary terminal segment, clypeus
apically dark; scape pale yellow, except by apical frontal dark mark; pedicel dark, with margins pale yellow;
flagellum entirely pale yellow; area between eyes covering the ocelli, head posterior area darkish brown, with
several minute darker spots. Pronotum: dorsally entirely, densely marked with small dark spots; ventrally pale
yellow, with dark spots more sparsely distributed, black median stripe. Legs: pale yellow, with only few dark spots;
foretrochanter laterally with larger dark spots; forefemora with dark spots restrict to anterior edge; foretibiae with
inconspicuous, few dark spots. Mid- and hindlegs pale yellow, with several dark spots of irregular sizes, without
transversal dark bands. Wings: light brown, very translucent; forewing main longitudinal and crossveins with dark
spots present, spots of crossveins more evident in costal and discoidal area; vein R entirely dark colored from base
to apical branching; area around vein M branches with two darkish marks; hindwing main longitudinal veins
apically dark spotted.
Female from Guarapari, Brazil. Length of body 23.3 mm from head to tip of abdomen, of pronotum 5.5 mm,
of metazone 3.5 mm, of forecoxa 4.7 mm; maximum width of head 4.3 mm, prozone middle 1.4 mm, supracoxal
dilatation 2.3 mm; metazone minimum width 1.4 mm. General morphology and coloration similar to described
males, except by body larger, more stout than males; ocelli vestigial (Fig. 12D); pronotum with faint denticulation
along lateral margin, more evident posteriorly (Fig. 14D); anterior edge of forecoxa with median, faint
denticulation; apterous; supranal plate longer than in males, but not overlaying ovipositor, sides somewhat parallel
(e.g., Fig. 17H); forefemora medially with three large transversal bands, basal one extends to femur posterior
region, from femur base to claw groove; hind femur with two large transversal areas of dense concentration of dark
spots on its apical half, apical mark darker.
Variability, male. Other examined males very similar to described males, except by foretibiae with 9–10
internal spines, pronotum sometimes with moderately long pilosity. General body dark marks and spots high
variable on size and intensity; wings with general also coloration high variable; non-described males with body
darker, forecoxa entirely, trochanter mostly laterally with larger dark spots; forefemora with dark spots of
moderate, spots larger on anterior edge; foretibiae with few dark spots of irregular sizes; wings distinctly brown,
less translucent than in described male, dark spots on forewing crossveins and hindwing veins sometimes
inconspicuous, area around vein M branches with one to three darkish marks (e.g., 3D).
Female. The other female very similar to described one, except by distinct shorter body length of 20,5 mm;
foretibiae with 9 internal spines; general coloration lighter, darkish marks and spots more rare and faint.
Comments. This species is most similar to M. phryganea and M. rustica, because their wings exhibit a similar
shape and coloration pattern, and all have convex vertex. However, males of M. ciliata can be easily distinguished
from those of M. rustica by having the supracoxal dilatation very faintly pronounced, prozone middle width/
supracoxal maximum greater than 0.80, forewing vein R entirely dark or almost so, ventral phallomere with distal
process slightly long, proximal process with short and robust point, and left dorsal phallomere with phalloid
apophysis very short and basally projected to the back. It can also be separated from males of M. phryganea by
having the supracoxal dilatation very faintly pronounced, prozone middle width/supracoxal maximum greater than
0.80, forewing only slightly shorter than hindwing, and forewing vein R entirely dark or almost so.
The females of M. ciliata are most similar to those of M. aptera because of the combination of the central
ocellus site without tubercle, the forefemora with distance between internal spines 12 and 11 about same distance
between spines 10 and 11 or slightly shorter, the foretibiae external spine 1 away from spine 2 by space equivalent
to one spine, and the supranal plate with sides parallel and rounded apex. However, females of M. ciliata can be
readily separated from this species by having the vertex entirely markedly convex, the supracoxal dilatation very
faintly pronounced, prozone middle width/supracoxal maximum greater than 0.80, the foretrochanter light brown
with few dark marks, and the hindfemur with area between the two transverse dark as pigmented as the remaining
leg parts.
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FIGURE 21. Miobantia fuscata (Giglio-Tos, 1915), male genitalia, right dorsal and ventral phallomeres from ventral
perspective, and left dorsal phallomere from dorsal perspective. A, male from Rserva Biológica de Duas Bocas, Sede, point 4;
B, male from Reserva Natural Vale, Estrada Experimental 1, Farinha Seca; C, MT00030; D, MT00061; E, male from Parque
Estadual de Itaúnas, point 7; F, male from Reserva Biológica de Duas Bocas, Pau Amarelo, point 18; G, MT00069; H, male
from Fazenda Usina Paineiras, Bosque 13. Images –.1, right dorsal phallomere; .2, ventral phallomere; .3, left dorsal
phallomere; .4, anterior process.
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FIGURE 22. Miobantia rustica (Fabricius, 1781), male genitalia, right dorsal and ventral phallomeres from ventral
perspective, and left dorsal phallomere from dorsal perspective. A, holotype of M. nebulosa; B, male at QBUM; C, holotype
(reduced scale). Images –.1, right dorsal phallomere; .2, ventral phallomere; .3, left dorsal phallomere.
Distribution (Fig. 27B). Brazil. Recorded from Rio de Janeiro (type locality), and from two localities within
Espírito Santo: Reserva Particular do Patrimônio Natural Oiutrem, Matilde, Alfredo Chaves (20°33'S 40°48'W);
and Parque Estadual Paulo César Vinha, Guarapari (20°36'S 40°25'W). Terra (1995) also recorded this species for
the locality of Campos do Jordão, state of São Paulo, Brazil (around 22°43'S 45°34'W). Agudelo, Lombardo and
Jantsch (2007) indicated the occurrence of this species also in the states of Rio Grande do Sul and Santa Catarina,
in Brazil, based on a previous study of Jantsch (1999). However, one male identified by such author as M. ciliata
and included in the original distribution record made by him in 1999 was examined and taken as the holotype of M.
sulista. Although these occurrence records were based also on other specimens, this misidentification suggests the
need for more detailed investigations of such data. One of these other specimens examined by Jantsch (1999) were
recorded from the locality of Colorado, Rio Grande do Sul (28°31'S 53°00'W); the examined material relating to
the record from Santa Catarina was not listed in the cited work.
Material Examined. 4 ♂♂, 2 ♀♀. Described male: ♂ from BRAZIL, Espírito Santo, Alfredo Chaves,
Matilde, Reserva Natural Oiutrem, 600-800 m, 14-21.X.2009, Malaise, COAzevedo et al. leg. [specimen
MT00047 in the cladistic analysis] (UFES). Described female: ♀ from BRAZIL, Espírito Santo, Guarapari,
Parque Estadual Paulo César Vinha, Restinga, Mata 2, 26.XI-02.XII.2006, Malaise, BAraújo & MSantos leg.
(UFES). Other specimens: BRAZIL: 1 ♀ from Espírito Santo, Alfredo Chaves, Matilde, Reserva Natural Oiutrem,
600-800 m, 14-21.X.2009, Malaise, COAzevedo et al. leg. (UFES); 1 ♂ from Espírito Santo, Guarapari, Parque
Estadual Paulo César Vinha, Restinga, Mata 7, 02-09.XI.2006, Malaise, BAraújo & MSantos leg. (UFES); 1 ♂
same data except Restinga, Mata 9 [MT00043]; 1 ♂ same data except Restinga, Mata 7, 09-16.XI.2006
[MT00033]. Images of the holotype: ♂ from BRAZIL, ciliata Stål, F. Sahlb., Typus, 27-35, NRM-MANT
0000674. Photographs taken by Gunvi Lindberg (NHRS).
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FIGURE 23. Miobantia immanis n. sp., male genitalia, right dorsal and ventral phallomeres from ventral perspective, and left
dorsal phallomere from dorsal perspective. A, holotype; B, MT00090; C, male at MCTP. Images –.1, ventral phallomere; .2,
left dorsal phallomere.
Miobantia fuscata (Giglio-Tos, 1915)
(Figs. 4; 11C; 12E–F; 14E–F; 16C; 17E; 18; 21; 27C)
Miopteryx fuscata Giglio-Tos, 1915: 139–140. Holotype ♂ (ZMB), examined.
Cnephomantis fuscatus: Rehn, 1920: 225.
Antimiopteryx fuscata: Giglio-Tos 1927: 212; Beier, 1935: 7.
Miobantia fuscata: Terra, 1995: 43; Ehrmann, 2002: 225; Agudelo et al., 2007: 120; Ehrmann & Koçak, 2009: 14.
Redescription, male holotype. Length of pronotum 4.3 mm, of metazone 2.8 mm, of forecoxa 3.8 mm, of
forewing 15.5 mm, of hindwing 16.3 mm; maximum width of head 3.2 mm, prozone middle 1.2 mm, supracoxal
dilatation 1.7 mm; metazone minimum width 0.9 mm.
Head (e.g., Figs 11C & 12E): Frontal sclerite longitudinally moderately short, less than 3.0 times as wide as
long, upper edge uniformly convex; small tubercle between each compound eye and toruli not pointed, represented
by low cuticle elevation; ocelli medium-sized, about as wide as pedicel apex, distance between ocelli as large as
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their width or larger, central ocellus about as elevated as lateral ocelli, lateral ocelli distinctly elliptical; vertex
somewhat straight, inconspicuously or not higher than imaginary line joining apex of eyes, with region between
parietal suture and eye not elevated towards parietal suture, nearly straight; parietal suture regularly marked.
Pronotum (e.g., Fig. 14E): With moderately long pilosity, about as long as wing pilosity, supracoxal dilatation
regularly sized, prozone middle width/supracoxal maximum dilatation between 0.71–0.75; margin without
denticulation.
Legs (e.g., Fig. 16C): Forecoxa markedly longer than metazone; forefemora internal spine 6 with shape and
size more similar to spine 5 than spines 2 and 4, distance between internal spines 12 and 11 about same distance
between spines 10 and 11 or slightly shorter, internal spine 12 distinctly shorter than spine 10; foretibiae external
spine 1 away from spine 2 by space equivalent to one spine, external spine 2 slightly longer than spine 3, with 9
internal spines.
Wings (Fig. 4B): Forewing slightly shorter than hindwing; vein M1 branched; vein CuA1 branches markedly
sinuous. Hindwing apical angle moderately acute, vein CuA1 branched, apex slightly short, rounded.
Color: Body dorsally darkish brown, ventrally pale yellow. Head: area between mouthparts and toruli pale yellow,
with few, very sparse, irregular dark marks, except by mandible apex, and maxillary terminal segment dark; scape
pale yellow; pedicel dark, with apical margin pale yellow; flagellum light brown, flagellomeres apically dark
brown; area between toruli and vertex darkish brown with large, marked, irregular dark marks, mostly on
transversal darker band between eyes covering the ocelli cuticle elevation; head dorsal region with irregular
darkish marks, transversal dark band on its apical area. Pronotum: markedly darkly pigmented dorsally, with few
irregular, evanescent bright marks; margin entirely pale yellow with moderately large dark spots; ventrally pale
yellow with few, evanescent dark spots, black median stripe faint, on its entire length. Legs: pale yellow with dark
marks, tarsomeres apically darkish, tarsomere 1 centrally darkish; forecoxa with many dark spots; trochanter
medially darkish; forefemora with dark spots of moderate size, with three inconspicuous, large, transversal dark
marks, posteriorly with dark longitudinal mark from femur base to claw groove, apical region anteriorly with faint,
small longitudinal dark mark; foretibiae with few dark spots of irregular sizes; fore- and midtibia with three large
transversal areas of dense concentration of faint dark spots, one at each extremity and another at middle, mid- and
hindfemur with two on its apical half. Mid- and hindlegs with dense darkish spots of irregular sizes. Wings: brown,
regularly translucent, costal margins apically with few bright marks; forewing dark spots present only on main
longitudinal veins, crossveins without dark spots; vein R spotted, with marks of unequal sizes, spaced from each
other by 2.0–3.0 times length of smallest spots; area around vein M branches without darkish marks.
Complementary description, male from Linhares, Brazil. Length of body 16.8 mm from head to tip of
abdomen, of pronotum 4.3 mm, of metazone 2.7 mm, of forecoxa 4.0 mm; of forewing 15.5; of hindwing 16.1;
maximum width of head 3.2 mm, prozone middle 1.3 mm, supracoxal dilatation 1.7 mm; metazone minimum
width 0.9 mm.
Abdomen: Supranal plate very short, rounded.
FIGURE 24. Miobantia arctissima n. sp., male genitalia, right dorsal and ventral phallomeres from ventral perspective, and
left dorsal phallomere from dorsal perspective, holotype.
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FIGURE 25. Miobantia sulista n. sp., male genitalia, right dorsal and ventral phallomeres from ventral perspective, and left
dorsal phallomere from dorsal perspective. A, male from Porto Alegre; B, holotype. Images –.1, ventral phallomere; .2, left
dorsal phallomere.
Genitalia (Fig. 21C): Ventral phallomere distal process short, with most marked sclerotization taking only
process tip, denticulation faintly present, tip slender but rounded, basal swelling absent; proximal process faintly
sclerotized, markedly projected, long, curved, distinctly turned posterad, with basal portion slightly wide, apex
rounded, distinctly slender than base; slit between posterior margin of ventral phallomere and proximal process
deep. Left dorsal phallomere with anterior process slightly long, moderately large, markedly curved at about
middle, oriented anterad, apex about as wide as middle; expanded portion between anterior process and phalloid
apophysis short, oriented anterad; well sclerotized area between phalloid apophysis and membranous lobe which is
marked, small, not reaching membranous lobe; phalloid apophysis regularly long, slender, its basal portion
projected to left one, subapically markedly bent, apically concave, slightly sharp; membranous lobe wide; dorsal
lamina posterior margin with left portion projected, expanded, uniformly curved.
Female from Linhares, Brazil. Length of body 16.0 mm from head to tip of abdomen, of pronotum 5.0 mm, of
metazone 3.2 mm, of forecoxa 4.2 mm; maximum width of head 3.6 mm, prozone middle 1.2 mm, supracoxal
dilatation 1.7 mm; metazone minimum width 0.9 mm. General morphology and coloration similar to male, except
by body slightly smaller, but stouter than holotype; head with vertex slightly higher than imaginary line joining
apex of eyes, with region between parietal suture and eye slightly more elevated towards parietal suture (Fig. 12F);
ocelli vestigial; pronotum with distinct denticulation along entire lateral margin, more pronounced posteriorly,
metazone slightly more robust (Fig. 14F); forecoxa anterior edge with median distinct denticulation; apterous;
supranal plate longer than in males, but not overlaying ovipositor, with sides distinctly converging towards a
subrounded apex (e.g., 17E); abdomen darkish brown with several dark spots, irregular dark marks.
Immature stages. First instar. Length of body 2.5 mm; antenna 3.3 mm. Head with median carina between
frontal sclerite upper edge middle and central ocellus; ocelli vestigial; vertex straight, not elevated. Foreleg spines
in pilose shape. Forefemora with 4 discoidal spines, the 3º with pilose apex diagonally inserted on a cuticular
elevated base, 4 external spines, 12 internal spines, spines 1–4, and 12 distinctly higher than the others; foretibiae
with 7 external spines, spines 1 and 3 almost indistinct, spines 2 and 7 distinctly higher than the others, spine 7
swollen, sinuous, 8 internal spines. Body general coloration brown with darker marks in similar positions of adults
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dark marks, dark spots absent; antenna with flagellum alternating between segment with apical half whitish and
segment entirely brown, except flagellum apical 0.3 entirely brown. Apterous. Genitalia not modified.
Second instar. Length of body 6.5 mm; antenna 4.0 mm. Head similar to first instar. Foreleg spines shape as in
adult. Forefemora with discoidal spines 3 regularly shaped, internal spines 2, 4, and 12 distinctly higher than the
others; foretibiae with external spines 1 and 3 distinctly lower than the others, spines 2 and 7 distinctly higher than
the others, spine 7 swollen, sinuous, 9–10 internal spines. Body general coloration brown with darker marks in
similar positions of adults dark marks, dark spots present; antenna with flagellum entirely brown. Apterous.
Genitalia not modified.
FIGURE 26. Miobantia nordestina n. sp., male genitalia, right dorsal and ventral phallomeres from ventral perspective, and
left dorsal phallomere from dorsal perspective. A, holotype; B, male from Pindobaçu. Images –.1, right dorsal phallomere; .2,
ventral phallomere; .3, left dorsal phallomere.
Third to last instar. Length of body about 10.0–15.7 mm; antenna 4.7–7.0 mm. General morphology and
coloration as in adults, except by head with median carina between frontal sclerite upper edge middle and central
ocellus progressively indistinct; ocelli vestigial; forefemora internal spine 12 only little higher than spine 10;
foretibiae with external spines 1–6 with about same size, except by spine 2 only little higher than the others, spine
7 distinctly higher than the others. Male wings vestigial in last instar, otherwise apterous. Genitalia progressively
developed, sexes are recognizable.
Ootheca. (Fig. 18) Pale yellow with 10 to 14 eggs positioned in two rows. Measurements: length of 5.94 mm,
height of 4.05 mm, and width of 3.25 mm.
Variability, male. Other examined males similar to holotype, except by foretibiae with 9–10 internal spines;
wings costal margins apically often with more marked bright marks. The analysis of a large set of individuals of
this species showed that male genitalia presented some particularly variable structures. Some different character
states seem to be local varieties. It is observed for males from localities in the northern Espírito Santo, such as
Pinheiros, Conceição da Barra, and some groups from Sooretama and Linhares, which have the ventral phallomere
proximal process expanded, with apex slightly slender than base (e.g., Figs 21C.2, 21E.2, 21G.2) while males from
Fazenda Oriente, in Atílio Vivacqua, and Itapemirim, in southern Espírito Santo, have the ventral phallomere with
proximal process slender, with apex distinctly slender than base (e.g., Fig. 21H.2). Other variable genitalia
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characters were the angle and width of the distal process of the ventral phallomere (e.g., Figs 21–.2), but always
maintaining the overall shape, the apical width of the anterior process of the left dorsal phallomere (e.g., Figs 21–.3
& 21–.4), and the general shape of the membranous lobe (e.g., Figs 21–.3), but always large. These characters also
seem to be related to populations of within the same locality, but apparently not following the geographical north-
south variation shown by the dorsal phallomere proximal process. Although variability in the male genitalia of this
species was frequent, it comprises clearly only slight variations of the same pattern, not overlaying the other
species pattern nor difficulting its identification. Additionally, the number of spines in the forefemora , and external
side of the foretibiae, was consistent in all examined material. The number of internal spines in the foretibiae varies
from 9 to 10, very rarely 11, often with variation in the number of spines between the right and left leg tibia of the
same individual.
The dark spots and general coloration of the body, and wing coloration high variable. Specimens from Aracruz,
Guarapari, Itarana, and Vila Velha have light brown wings, regularly translucent, with very faint, dark spots (e.g.,
Fig. 4E), the specimens from Conceição da Barra, Itapemirim, and some specimens from Santa Leopoldina, and
Santa Tereza have wings dark brown colored, with more evident dark spots, general body coloration darker than
holotype, with dark spots and marks more evident, hindtibia with three large, marked transversal dark bands. In the
other hand, about 60% of individuals from Pinheiros have wings almost white, very translucent, with faint dark
spots, general body coloration lighter than holotype, with dark spots and marks less evident (e.g., Fig. 4F); the
other 40% have similar wings, but less translucent and with slightly grayish tint. The remaining material examined
is more similar to holotype, individuals from Linhares, and Sooretama especially with large darkish evanescent
marks on wings. Though high variability in wing coloration was found, the pattern of forewing vein R dark spots of
distinctly unequal sizes, the absence of dark spots in crossveins, and the presence of bright marks on both wings
costal margins (even if sometimes faintly marked) remained consisted across examined specimens.
It is also remarkable that anomalies of wing venation had an observed frequency of 15,8% among the males of
M. fuscata. The most common abnormalities were the forewing vein M1 not branched or branched twice, the vein
M2 branched, and the hindwing vein CuA1 not branched or branched twice. However, abnormal venation patterns
were very rarely repeated in the corresponding wing of the other side of body.
All variation mentioned above contributed to the separation of 23 morphospecies later associated to M.
fuscata, and in the delimitation of the species. This high variability is also supported by the molecular data, which
indicated the existence of genetic distances of up to 17.7% between individuals of this species. These distances
between individuals of different morphospecies were above 2%, while distances between individuals of the same
morphospecies were rarely above 1% (often 0–0.1%). However, after all analyzes this large number of
morphospecies proved to be only local varieties. Even with high variation of some characteristics, the general
patterns of the distinct species are easily recognizable by the features stated in the "Comments" section for each of
them, specially the comparison of the male genitalia between the examined specimens.
Female. Other examined females very similar to described female, except by mostly slightly larger than
described one, foretibiae with 10–11 internal spines. Body coloration highy variable, with dark marks and spots.
The quantity of dark spots along the body varies from nearly absent with the presence of only a few large marks, to
dark spots densely present along the entire body. Unlike the color variation of males, this variation in females seem
to be little related to the locality. For instance, among females collected in Pinheiros, about 50% have light colored
body with few dark markings, whereas the other half exhibits darker body coloration with more abundant and
evident dark spots.
Comments. Males of M. fuscata are most similar to males of M. aptera and M. immanis based on the dark
spots on vein R of the forewings, with marks of distinctly unequal sizes, and the costal margin of both wings with
distinct bright marks apically. However, M. fuscata can be instantly distinguished from these two species by having
vein M1 of forewings branched. Although the examination of male genitalia can successfully separate it from M.
immanis (based on the distal process of ventral phallomere with slender apex and the proximal process long and
curved), it may also be confused with M. aptera. In this case, M. fuscata can be identified by the distal process of
ventral phallomere with faint denticulation present, its tip rounded, with most marked sclerotization restricted only
to process tip, and the left dorsal phallomere with a well sclerotized area between the phalloid apophysis and the
membranous lobe small, not reaching the membranous lobe.
The females are most likely to be mistaken for M. nordestina due to the combination of having the head with a
distinctly convex vertex, frontal sclerite markedly short, and supranal plate with sides distinctly converging
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towards a subrounded apex. However, it can be isolated from females of M. nordestina by having the vertex region
between parietal suture and eye inconspicuously convex, nearly straight, pronotum with metazone slightly robust,
and forefemora with internal spine 12 distinctly shorter than spine 10.
Distribution. (Fig. 27C) Brazil. The exact type locality is unknown. Recorded from Espírito Santo by Rehn
(1920), here registered for 29 additional localities of this state. The most northern record is for the Parque Estadual
de Itaúnas, Conceição da Barra (18°20'S 39°40'W) and the most southern is for Fazenda Oriente, Atílio Vivacqua
(20°58'38.6"S 41°10'10.6"W). Also registered from the states of Santa Catarina, and of Rio Grande do Sul
(Agudelo et al. 2007), based on a previous study of Jantsch (1999). One male identified by such author as M.
fuscata was examined and matched with the concept of M. fuscata considered in the present work. The exemplar
included in the original distribution record for Santa Catarina is indicated for the locality of Nova Teutônia
(27°11'S 52°23'W); the examined material relating to the record from Rio Grande do Sul was not listed in the cited
work.
Material Examined. 793 ♂♂, 35 ♀♀, 39 nymphs. Holotype: ♂ from BRAZIL, Miopteryx fuscata Giglio-Tos
typus, E. Giglio-Tos det., Holotypus Nr., Mantis, Ca. No. 653. (ZMB). Described male: ♂ from BRAZIL, Espírito
Santo, Linhares, Reserva Natural Vale, estrada próximo à porteira à direita da entrada, point 19, 21-29.V.2010,
Malaise, APAguiar et al. leg. [specimen MT00030 in the cladistic analysis] (UFES). Described female: ♀ from
BRAZIL, Espírito Santo, Linhares, Reserva Natural Vale, ao lado dos Laboratórios, 13.I.2011, lighting trap,
MVScherrer & MCCarreiro leg. (UFES). Other specimens: BRAZIL: 1 ♂ from Espírito Santo, Conceição da
Barra, Parque Estadual de Itaúnas, 23-25.XI.2006, Malaise, MTTavares et.al. leg. (UFES); 3 ♂♂ same data except
23-28.XI.2006; 5 ♂♂ same data except point 7, 23-25.XI.2006; 2 ♂♂ same data except point 14 [one of them
MT00061]; 6 ♂♂ same data except point 3, 31.X-15.XI.2006; 1 ♀ same data except Conceição da Barra, Floresta
Nacional do Rio Preto, 200? [no further data]; 1 ♂ from Espírito Santo, Pinheiros, Reserva Biológica Córrego do
Veado, 08.VI.2011, manual, BFSantos leg. (UFES); 1 ♀ same data except 09.VI.2011, F Dal Vechio leg., 14
nymphs from ootheca laid by the previous female; 1 ♀ same data except lighting trap, BFSantos leg.; 2 ♂♂ same
data except manual, BFSantos leg. [one of them MT00074]; 1 ♂ same data except MVScherrer leg.; 1 ♀ same data
except 10.VI.2011, HYamaguti leg.; 5 ♂♂, 1 ♀, 1 nymph same data except MVScherrer leg.; 1 nymph same data
except 12.VI.2011, BFSantos leg.; 1 ♀ same data except 13.VI.2011; 1 ♂ same data except Reserva Biológica
Córrego do Veado, Estrada Oeste, point 8, 12-14.VI.2011, yellow pan trap, MTTavares et al. leg.; 1 ♂ same data
except Trilha Água Limpa, point 16, 09-11.VI.2011, Malaise, MTTavares et al. leg. [MT00073]; 2 ♂♂ same data
except point 17; 1 ♂ same data except point 18; 3 ♂♂ same data except point 19; 1 ♂ same data except point 20; 1
♂ same data except point 29; 1 ♂ same data except point 16, 09-11.VI.2011, yellow pan trap, MTTavares et al.
leg.; 1 ♂ same data except point 30; 2 ♂ same data except point 30, 11-13.VI.2011, Malaise, MTTavares et al. leg.;
4 ♂♂ same data except point 19, 15-17.VI.2011; 4 ♂♂ same data except Malaise 3, 27.XI-06.XII.2011; 3 ♂♂ same
data except Malaise 5; 2 ♂♂ same data except Malaise 6; 1 ♂ same data except Malaise 7; 1 ♀ same data except
Trilha da Anta, point 5, 09-11.VI.2011, yellow pan trap, MTTavares et al. leg.; 1 ♂ same data except point 8,
Malaise, MTTavares et al. leg.; 1 ♂ same data except point 9; 1 ♂ same data except point 15; 1 ♀ same data except
point 2, 10.VI.2011, manual, MVScherrer leg.; 1 ♂ same data except 10-12.VI.2011, Malaise, MTTavares et al.
leg.; 2 ♂♂ same data except point 1, 11-13.VI.2011; 1 ♂ same data except point 11; 1 ♂ same data except point 14;
1 ♀ same data except point 2, 14-16.VI.2011, yellow pan trap; 2 ♂♂ same data except point 1, 15-17.VI.2011,
Malaise; 1 ♂ same data except point 2; 1 ♂ same data except point 10; 1 ♂ same data except point 11 [MT00070];
1 ♂ same data except point 13; 1 ♂ same data except point 14; 3 ♂♂ same data except point 15; 4 ♂♂, 3 ♀♀,
1nymph same data except Trilha Jaboti, 11.VI.2011, manual, BFSantos leg., 14 nymphs from ootheca laid by a
female of the previous set; 10 ♂♂ same data except Malaise 2, 27.XI-06.XII.2011, Malaise, MTTavares et al. leg.;
3 ♂♂ same data except Malaise 3; 11 ♂♂ same data except Malaise 4; 8 ♂♂ same data except Malaise 5; 11 ♂♂
same data except Malaise 6; 2 ♂♂ same data except Malaise 8; 1 ♂ from Espírito Santo, Sooretama, Reserva
Biológica Sooretama, 14.XII.2012, BCAraújo & TRBrotto leg. (UFES); 1 ♀ same data except Reserva Biológica
Sooretama, Sede, 02.VI.2011, BFSantos leg.; 1 nymph same data except Trilha Sede, 31.V.2011, MTTavares et al.
leg.; 1 ♀ same data except 03.VI.2011, HYamaguti BFSantos leg.; 1 ♂ same data except 08.VI.2011, Malaise,
MTTavares et al. leg.; 15 ♂♂ same data except 11-18.XI.2011, Malaise, COAzevedo et al. leg.; 1 ♂ same data
except Trilha da Sede, point 12, 31.V-2.VI.2011, Malaise, MTTavares et al. leg.; 1 ♂ same data except 06-
08.VI.2011; 3 ♂♂ same data except Trilha Ana, Malaise 3, 06-12.XII.2011; 1 ♂ same data except Malaise 5; 14
♂♂ same data except Malaise 6; 8 ♂♂ same data except Malaise 7; 13 ♂♂ same data except Malaise 8; 5 ♂♂ same
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data except Malaise 1; 9 ♂♂ same data except Malaise 2; 1 ♂ same data except Malaise 3; 11 ♂♂ same data except
Malaise 4; 5 ♂♂ same data except Malaise 5; 4 ♂♂ same data except Malaise 6; 10 ♂♂ same data except Malaise
7; 3 ♂♂ same data except Malaise 8; 1 ♂ same data except Estrada Córrego Cupido, 37 m, 30.X.2011 [MT00058];
1 ♂ same data except Estrada Cupido, point 30, 31.V-2.VI.2011; 1 ♀ from Espírito Santo, Linhares, Reserva
Natural Vale, 08.VII.2010, manual, APAguiar et al. leg. (UFES); 1 ♀ same data except 21.I.2011, MCCarreiro leg.;
7 ♂♂ same data except Reserva Natural Vale, Ao lado do laboratório, 13.I.2011, lighting trap, MVScherrer et al.
leg.; 1 ♂ same data except Bicuíba, trilha 2, point 4, V.2010, Malaise, APAguiar et al. leg.; 1 ♂ same data except
point 3 [MT00029]; 2 ♂♂ same data except trilha 1, point 1, 29.V-08.VII.2010; 3 ♂♂ same data except point 2; 1
♂ same data except point 4; 7 ♂♂ same data except Descampado Macanaiba Pele de Sapo, point 10; 1 ♀ same data
except Estrada Bicuíba, point 28, 24-26.I.2011, yellow pan trap; 1 ♂ same data except Estrada Bicuíba, trilha 2,
point 15, 18-25.I.2011, Malaise; 7 ♂♂ same data except Estrada da Gávea, borda do rio, point 16, 19-25.I.2011; 3
♂♂ same data except Estrada da Gávea, point 6, 21-26.I.2011; 6 ♂♂ same data except Estrada da Oiticica próximo
a Ponte, point 18, 29.V-08.VII.2010; 4 ♂♂ same data except Estrada do Campo; 1 ♀ same data except Estrada do
Campo Próx. a Administração, point 20, 24-26.I.2011, yellow pan trap; 1 ♂ same data except Estrada do Campo,
point 05, 23-29.V.2010, Malaise; 5 ♂♂ same data except point 06, 17-23.V.2010; 5 ♂♂ same data except 23-
29.V.2010; 13 ♂♂ same data except 29.V-08.VII.2010; 1 nymph same data except Estrada entre Roxinho e
Bicuiba, point 7, 22-24.V.2010, yellow pan trap; 1 ♂ same data except 23-29.V.2010, Malaise; 2 ♂♂ same data
except 29.V-08.VII.2010; 1 ♂ same data except point 8; 1 ♂ same data except point 15, 16-18.I.2011, yellow pan
trap; 1 ♀ same data except 20-22.I.2011; 1 ♂ same data except Estrada Experimental 1, Farinha Seca, point 15, 19-
29.V.2010, Malaise; 15 ♂♂ same data except 29.V-08.VII.2010; 12 ♂♂ same data except Estrada Experimental 2,
Farinha Seca, point 16, 19-29.V.2010 [one of them MT00027]; 2 ♂♂ same data except Estrada Farinha Seca, point
12, 21-26.I.2011; 1 ♂ same data except Estrada Flamengo após a Estrada da Oiticica e Guaribu Sabão, point 4; 3
♂♂ same data except 14-21.I.2011; 7 ♂♂ same data except Estrada Flamengo próximo a porteira, point 5; 1 ♂
same data except 21-26.I.2011; 1 nymph same data except Estrada Flamengo, point 53, 24-26.V.2010, yellow pan
trap; 1 ♀ same data except Estrada Ipê Amarelo, point 27, 17-19.I.2011; 1 ♂ same data except Estrada Macanaiba
Pele de Sapo, point 7, 15-21.I.2011, Malaise; 1 ♂ same data except 21-26.I.2011; 5 ♂♂ same data except Estrada
Oiticica, point 10, 15-21.I.2011; 4 ♂♂, 1 nymph same data except 21-26.I.2011; 1 ♀ same data except point 17, 16-
18.I.2011, yellow pan trap; 4 ♂♂ same data except Estrada Orelha da Onça, point 9, 21-26.I.2011, Malaise; 4 ♂♂
same data except 15-21.I.2011; 7 ♂♂ same data except Estrada Paraju, point 14, 17-25.I.2011; 4 ♂♂ same data
except Estrada próximo a porteira, point 13, 16-21.I.2011; 2 ♂♂ same data except Estrada próximo à porteira à
direita da entrada, point 19, 21-29.V.2010; 5 ♂♂ same data except 29-08.V.2010; 2 ♀♀ same data except Estrada
Roxinho, point 13, 16-18.I.2011, yellow pan trap; 7 ♂♂ same data except Estrada X, point 6, 29.V-08.VII.2010,
Malaise; 1 ♂ same data except Flamengo, baixada após a torre, point 12, 23-30.V.2010; 4 ♂♂, 1 ♀ same data
except Macanaiba Pele de Sapo, point 10, 29.V-08.VII.2010; 1 ♀ same data except Margem da Farinha Seca,
19.V.2010, manual, MCCarreiro & FFGomes leg.; 1 nymph same data except Margem do Rio, alagado, point 22,
17-19.I.2011, yellow pan trap, APAguiar et al. leg.; 19 ♂♂ same data except Mussununga Flamengo, point 09,
29.V-08.VII.2010, Malaise; 6 ♂♂ same data except 23-30.V.2010; 1 nymph same data except Trilha atrás do
viveiro de mudas, 14.I.2011, manual, MVScherrer leg.; 1 ♀ same data except Trilha atrás do viveiro de mudas,
point 1, 24-26.V.2011, yellow pan trap, APAguiar et al. leg.; 1 ♂ same data except Trilha atrás do viveiro de mudas,
point 3, 14-21.I.2011, Malaise; 3 ♂♂ same data except Trilha do campo aberto próximo a administração, point 2,
13-20.I.2011; 1 ♀ same data except point 19, 16-18.I.2011, yellow pan trap; 1 nymph same data except Trilha
Estrada Flamengo, point 39, 24-26.V.2011; 14 ♂♂ same data except Trilha próximo ao rio, Gávea, point 17, 19-
29.V.2010, Malaise; 1 ♀ same data except Trilha próximo ao viveiro, 27-29.V.2010, yellow pan trap; 20 ♂♂, 1 ♀
same data except Trilha próximo ao viveiro, point 14, 29.V-08.VII.2010, Malaise; 5 ♂♂ same data except 19-
29.V.2010; 1 ♂ same data except Linhares, Desengano - Salvador Lagoa Azul, point mata 1, Malaise 1, 03-
10.XI.2007, Malaise, FGRampinelli et al. leg.; 1 ♂ same data except point mata 1, Malaise 2; 1 ♂ same data except
point mata 1, Malaise 3; 1 ♂ from Espírito Santo, Pancas, Faz. Juliberto Stur, [no further data] (UFES); 2 ♂♂ same
data except Fazenda Juliberto Stur, área 1, point B1, 31.I-07.II.2003, Malaise, MTTavares & COAzevedo leg.; 6
♂♂ same data except área 1, B2, 24-31.I.2003, Malaise; 7 ♂♂ same data except T2; 3 ♂♂ same data except área 2,
T7; 1 ♂ from Espírito Santo, Aracruz, Fazenda São José, Malaise 1, 06-13.X.2007, Malaise, FGRampinelli et al.
leg. (UFES); 3 ♂ same data except point 2 [one of them MT00013]; 3 ♂♂ same data except point 3; 2 ♂♂ same
data except point 4; 1 ♂ from Espírito Santo, Santa Teresa, Malaise 4, 19-26.X.2007, Malaise, FGRampinelli et al.
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leg. (UFES); 1 ♂ same data except Santa Teresa, Estação Biológica Santa Lúcia, B2, 19-23.X.2007, Malaise,
MTTavares et al. leg.; 1 ♂ same data except Estação Biológica Santa Lúcia, Trilha Tapinoã, 16-22.X.2010
[]MT00049]; 1 ♂ same data except Estação Biológica Santa Lúcia, Trilha Itapinuã, 17-23.X.2011; 1 ♂ same data
except Estação Biológica Santa Lúcia, Trilha Indaiaçu, área de pedra, B2, 06-09.XI.2009; 2 ♂♂ same data except
Estação Biológica Santa Lúcia, point P2, Trilha, 15-19.X.2010; 1 ♂ same data except Santa Teresa, Museu de
Biologia Mello Leitão, 23.II.2001, manual, RKawada leg.; 3 ♂♂ from Espírito Santo, Itarana, Bom Destino, point
P2, 22-25.II.2004, Malaise, BCAraújo leg. (UFES) [one of them MT00069]; 1 ♂ from Espírito Santo, Santa Maria
de Jetibá, Fazenda Clarindo Kruger, point T7, 06-13.XII.2002, Malaise, MTTavares et al. leg. (UFES); 3 ♂♂ same
data except B5, 29.XI-06.XII.2002; 1 ♂ same data except B8; 1 ♂ same data except T5; 1 ♂ same data except
Santa Maria de Jetibá, Fazenda Paulo Seick; 1 ♂ same data except Fazenda Paulo Seick, Área 1, point B2, 06-
13.XII.2002; 1 ♂ same data except B4; 2 ♂♂ same data except 29.XI-06.XII.2002; 4 ♂♂ same data except T2, 06-
13.XII.2002; 2 ♂♂ same data except 29.XI-06.XII.2002; 4 ♂♂ same data except B1; 1 ♂ from Espírito Santo,
Santa Leopoldina, Suiça, Mata, 361 m, 05-12.XI.2007, Malaise, COAzevedo et al. leg. (UFES); 1 ♂ same data
except Mata, 361 m, Malaise 6; 1 ♂ same data except Malaise 7 [MT00085]; 2 ♂♂ same data except Malaise 8; 2
♂♂ same data except Santa Leopoldina, Alto Rio das Farinhas, 14-24.V.2008, Malaise, CWaichert & Furieri leg.
[one of them MT00021]; 3 ♂♂ same data except Santa Leopoldina, Meia Légua, Mata próximo a trilha, 31.III-
06.IV.2007, Malaise, FGRampinelli et al. leg.; 5 ♂♂ same data except Meia Légua, Point próximo a cerca de
divisão; 5 ♂♂ same data except Meia Légua, Trilha 1; 1 ♂ same data except Trilha 2; 1 ♂ same data except Trilha
3; 12 ♂♂, 1 nymph same data except Trilha 5 [one ♂ MT00084]; 1 ♂ from Espírito Santo, Serra, Mestre Álvaro,
16.IV.1987, manual, APAguiar leg (UFES).; 1 ♀ same data except Serra, Cidade Continental, Setor África,
16.XI.2009, MVScherrer leg.; 3 ♂♂ from Espírito Santo, Domingos Martins, Mata Pico do Eldorado, point B1, 03-
10.XII.2004, Malaise, MTTavares et al. leg. (UFES); 6 ♂♂ same data except B2; 5 ♂♂ same data except B4; 2 ♂♂
same data except B5; 2 ♂♂ same data except B8; 10 ♂♂ same data except T1; 3 ♂♂ same data except T2; 1 ♂
same data except T3; 4 ♂♂ same data except T4; 3 ♂♂ same data except T5; 4 ♂♂ same data except T6; 11 ♂♂
same data except T7; 3 ♂♂ same data except T8; 3 ♂♂ same data except B2, 26.XI-03.XII.2004, Malaise,
MTTavares et al. leg.; 2 ♂♂ same data except B3; 2 ♂♂ same data except B4; 1 ♂ same data except B6; 4 ♂♂
same data except T2 [one of them MT00088]; 2 ♂♂ same data except T6; 8 ♂♂ same data except T7; 1 ♂ from
Espírito Santo, Cariacica, Reserva Biológica de Duas Bocas, 08.II.2003, MTTavares et al. leg. (UFES); 1 ♀, 1
nymph same data except Reserva Biológica de Duas Bocas, Pau Amarelo, 21-27.X.2005, Malaise, APAguiar et al.
leg.; 2 ♂♂ same data except 20-28.X.2005; 1 ♂ same data except 21-30.X.2005; 3 ♂♂ same data except Pau
Amarelo, point 1, 20-28.X.2005; 2 ♂♂ same data except point 5; 2 ♂♂ same data except 28.X-05.XI.2005; 3 ♂♂
same data except point 7, 20-28.X.2005; 1 ♂ same data except point 8; 9 ♂♂ same data except point 10, 21-
30.X.2005; 1 ♂ same data except point 13; 7 ♂♂ same data except point 14; 2 ♂♂ same data except point 17; 1 ♂
same data except point 18; 1 ♂ same data except point 9, 28.X-05.XI.2005; 3 ♂♂ same data except point 21; 2 ♂♂
same data except point 22; 1 ♂ same data except Reserva Biológica de Duas Bocas, Sede, point 2, 09-13.III.2009,
Malaise, BFSantos et al. leg.; 4 ♂♂ same data except Sede, point 4; 2 ♂♂ same data except Sede, point 5 [one of
them MT00077]; 2 ♂♂ from Espírito Santo, Vila Velha, Reserva Ecológica Jacarenema, point 4, 30.X-06.XI.2002,
Malaise, COAzevedo et al. leg. (UFES); 4 ♂♂ same data except point 7 [one of them MT00064]; 4 ♂♂ same data
except point 9 [one of them MT00016]; 1 ♂ from Espírito Santo, Marechal Floriano, 09.IV.2004, CLMPereira leg.
(UFES); 8 ♂♂ from Espírito Santo, Alfredo Chaves, Matilde, Reserva Natural Oiutrem, 600-800 m, 14-21.X.2009,
Malaise, COAzevedo et al. leg. [one of them MT00052] (UFES); 1 ♀ same data except 14.X.2009, manual,
COAzevedo leg.; 4 ♂♂ same data except Alfredo Chaves, Picadão, Mata, Malaise 4, 714 m, 8-15.X.2007, Malaise,
COAzevedo et al. leg. [one of them MT00053]; 1 ♂ same data except Malaise 8; 1 ♂ from Espírito Santo,
Guarapari, Parque Estadual Paulo Cesar Vinha, Restinga, Mata 3, 02-10.XII.2006, Malaise, BAraújo & MSantos
leg. (UFES); 1 ♂ same data except Mata 12, 15-26.V.2006; 1 ♂ same data except Mata 8, 2-10.XII.2006; 2 ♂♂
same data except Mata 2, 26.XI-2.XII.2006; 1 ♂ same data except Mata 4, 08-15.V.2006 [MT00034]; 1 ♂ same
data except 02-09.XII.2006; 1 ♂ same data except Mata 10, 16-26.XI.2006; 1 ♂ same data except Mata 5; 1 ♂
same data except Mata 7; 2 ♂♂ same data except Mata 8; 1 ♂ same data except 26.XI-02.XII.2006; 1 ♀ from
Espírito Santo, Itapemirim, Mata da Usina Paineiras, point 14, 01-02.III.2008, yellow pan trap, APAguiar et al. leg.
(UFES); 1 ♂ same data except Itapemirim, Fazenda Usina Paineiras, 19-26.XI.2010, Malaise, MTTavares et al.
leg. [MT00031]; 3 ♂♂ same data except Fazenda Usina Paineiras, point B1/Pt.1; 4 ♂♂ same data except B2/Pt.1;
2 ♂♂ same data except B3/Pt.1; 4 ♂♂ same data except B4/Pt.1; 3 ♂♂ same data except B5/Pt.1; 2 ♂♂ same data
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except B5/Pt.2; 4 ♂♂ same data except B6/Pt.1; 5 ♂♂ same data except B7/Pt.1; 3 ♂♂ same data except B9/Pt.2;
3 ♂♂ same data except B10/Pt.2; 2 ♂♂ same data except B11/Pt.2; 3 ♂♂ same data except B13/Pt.2; 1 ♂ same
data except B15/Pt.2; 1 ♂ from Espírito Santo, Atílio Vivacqua, Fazenda José Carlos Lutoza (UFES); 1 ♂ same
data except Fazenda José Carlos Lutoza, Área 1, point B3, 13-20.II.2003, Malaise, MTTavares & COAzevedo leg.;
1 ♂ same data except points T2+T4+T6+B2, 20-27.II.2003; 1 ♂ same data except points T3; 4 ♂♂ same data
except Atílio Vivacqua, Fazenda Oriente, Área 2 [one of them MT00066]; 3 ♂m3, 1 nymph same data except Área
2, points T8+B4+B6+B8, 13-20.II.2003; 1 ♂ same data except 20-27.II.2003; 11 ♂♂ same data except point T5,
20-27.II.2003; 1 ♂ same data except 20-27.II.2003; 1 ♂ from Espírito Santo, exact locality unknown, point trilha 2,
31.III-6.IV.2007, Malaise, FGRampinelli leg. (UFES).
Miobantia immanis Scherrer, n. sp.
(Figs. 7; 11E; 13A–B; 15A–B; 16D, I; 17C ,I; 23; 27F)
Description, male holotype. Length of body 21.4 mm from head to tip of abdomen, of pronotum 5.3mm, of
metazone 3.5 mm, of forecoxa 4.1 mm; of forewing 16.0; of hindwing 18.3; maximum width of head 3.4 mm,
prozone middle 1.3 mm, supracoxal dilatation 2.0 mm; metazone minimum width 0.9 mm.
Head (Figs 11E & 13A): Frontal sclerite longitudinally moderately short, less than 3.0 times as wide as long,
upper edge uniformly convex; small tubercle between each compound eye and toruli in form of slightly pointed
crest; ocelli medium-sized, about as wide as pedicel apex, distance between ocelli as large as their width or larger,
central ocellus distinctly more elevated than lateral ocelli by low tubercle, lateral ocelli rounded; vertex somewhat
straight, distinctly higher than imaginary line joining apex of eyes, with region between parietal suture and eye
more elevated towards parietal suture, markedly convex, forming distinct, low bulge (juxtaocular bulge); parietal
suture regularly marked; vertex region between parietal sutures laterally abruptly elevated close to parietal suture,
centrally slightly concave.
Pronotum (Fig. 15B): With very long pilosity, longer than wing pilosity, supracoxal dilatation markedly
pronounced, prozone middle width/supracoxal maximum dilatation less than 0.70; margin without denticulation.
Legs (Figs 16D & 16I): Forecoxa slightly longer than metazone; forefemora internal spine 6 with shape and size
more similar to spines 2 and 4 than the spine 5, distance between internal spines 12 and 11 distinctly longer than
distance between spines 10 and 11, internal spine 12 distinctly shorter than spine 10; foretibiae external spine 1
away from spine 2 by space equivalent to two spines, external spine 2 about same size of spine 3, with 10 internal
spines.
Wings (Fig. 7A): Forewing much shorter than hindwing; vein M1 not branched; vein CuA1 branches nearly
straight. Hindwing apical angle slightly acute, vein CuA1 branched, apex abruptly interrupted.
Abdomen: Supranal plate very short, rounded.
Genitalia (Fig. 23A): Ventral phallomere distal process short, with most marked sclerotization taking entire
process, denticulation markedly present, tip wide, rounded, basal swelling present; proximal process markedly
sclerotized, only slightly projected, with basal portion slightly wide, apex rounded. Left dorsal phallomere with
anterior process slightly long, markedly large, slightly curved at base, oriented anterad, apex about as wide as
middle; expanded portion between anterior process and phalloid apophysis long; well sclerotized area between
phalloid apophysis and membranous lobe which is marked, large, wide; phalloid apophysis regularly long, slender,
its basal portion projected to left one, somewhat contiguously curved with entire phalloid apophysis, apically
concave, slightly sharp; membranous lobe wide; dorsal lamina posterior margin with left portion projected,
distinctly bent.
Color: Head, thorax except appendices, dorsally dark, ventrally pale yellow, abdomen entirely dark. Head: area
between mouthparts and toruli pale yellow, with few, very sparse, small dark spots, except by mandible apex, and
maxillary terminal segment dark; scape pale yellow, except by apical frontal dark mark; pedicel dark, with apical
margin pale yellow; flagellum mostly dark, alternating between segment with apical half pale yellow and segment
entirely dark from segment 10 to apex; area between toruli and vertex dark pigmented, with large bright marks of
irregular forms and intensities; pale yellow median line starting at foramen magnum, bifurcating just before the
ocelli cuticle expansion, going along with parietal suture to eye; ocelli cuticle expansion centrally with another pale
yellow median line. Pronotum: dorsally entirely dark colored, with pale yellow evanescent marks of irregular
forms; margin pale yellow with moderately large dark spots; ventrally pale yellow, with few, sparse dark spots,
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black median stripe on its posterior half. Legs: pale yellow, with dark marks, tarsomeres apically darkish,
tarsomere 1 centrally darkish; forecoxa with large dark spots; trochanter laterally with few large dark spots,
medially with large longitudinal dark mark; forefemora with dark spots of moderate, anteriorly with two distinct
areas of spots concentrations, visible at the lateral and medial areas of femur, posteriorly with dark longitudinal
mark from femur base to claw groove, apical region anteriorly with longitudinal dark mark; foretibiae with few
dark spots of irregular sizes; fore- and midtibia with three large transversal areas of dense concentration of dark
spots, one at each extremity and another at middle, hindfemur with two on its apical half. Mid- and hindlegs with
dense dark spots of irregular sizes. Wings: mostly dark, slightly translucent; costal margins apically, some veins of
forewing, hindwing apical part with bright large marks, not restricted to vein limits; forewing dark spots present
only on main veins, crossveins without dark spots; vein R spotted, with marks of unequal sizes, spaced from each
other by 2.0–3.0 times length of smallest spots; area around vein M branches without darkish marks.
Female allotype. Length of body 26.4 mm from head to tip of abdomen, of pronotum 8.1 mm, of metazone 5.3
mm, of forecoxa 6.4 mm; maximum width of head 4.7 mm, prozone middle 1.7 mm, supracoxal dilatation 3.0 mm;
metazone minimum width 1.4 mm. General morphology and coloration very similar to male, except by body
markedly larger, more stout than holotype; head with vertex centrally straight (Fig. 13B); ocelli vestigial; pronotum
with distinct denticulation along entire lateral margin, more pronounced anteriorly, metazone slightly more robust
(Fig. 15A); forecoxa anterior edge with median distinct denticulation; apterous; supranal plate longer than in
males, but not overlaying ovipositor, side parallel (Fig. 17I). General coloration lighter, dark marks less marked;
head with dark marks on parietal suture, area between parietal sutures with densely distributed evanescent dark
spots.
Immature Stages. Nymph, unknown instar. Length of body 8.7 mm; antenna 4.7 mm. General morphology and
coloration as in adults, except by ocelli vestigial; forefemora internal spine 11 vestigial; foretibiae with external
spines 1 and 3 lower than the others, spine 2 only slightly higher than spines 4–6, spine 7 distinctly higher than the
others. Apterous. Cercus basally stout, very slender towards apex.
Variability, male. Paratypes very similar to holotype, except by the foretibiae with 11 internal spines;
hindwing large bright marks restricted to costal area apical parts; genitalia ventral phallomere of paratypes form
Domingos Martins and Santa Teresa with posterior right quadrant centrally with three distinct, thick hairs, left
dorsal phallomere of paratype from Linhares with anterior process apex slightly wider than its middle. General
body dark marks and spots high variable on size and intensity; wings with general coloration high variable,
paratypes from Domingos Martins and Linhares with wings distinctly brown, paratype from Santa Teresa with
wings light brown, very translucent, bright marks inconspicuous (Fig. 7B). Flagellum with about flagellomere 25
to apex alternating between segment with apical half pale yellow and three segments entirely dark, apparently
because flagellomeres are duplicated from this point.
Comments. It is the most distinctive species examined, both male and female can be instantly separated from
all other species of Miobantia by having the small juxtaocular tubercle between each compound eye and toruli
prominent, in the form of a low and lightly pointed crest, the central ocellus on small and low tubercle, the
flagellum with bright transverse marks, the forecoxa only slightly longer than metazone, the forefemora with
distance between internal spines 12 and 11 distinctly longer than distance between spines 10 and 11, and the
foretibiae external spine 1 away from spine 2 by space equivalent to two spines. Males also have the forewing vein
M1 not branched, while the hindwing vein CuA is branched, a unique venation pattern among species of the genus,
as well as the hindwing apex abruptly interrupted. In addition, the male genitalia have the ventral phallomere with
proximal process only slightly projected and the dorsal lamina of the left dorsal phallomere with the left portion
distinctly bent.
Distribution (Fig. 27F). Brazil. Recorded from five localities of Espírito Santo: Reserva Biológica Córrego do
Veado, Pinheiros (type locality) (18°22'04"S 40°8'29"W); Reserva Natural Vale, Linhares (19°09'06"S
40°04'14"W); Estação Biológica Santa Lúcia, Santa Teresa (19º57'55"S 40º32'25"W); Mata Pico do Eldorado,
Domingos Martins (20°22'17"S 40°39'29"W); and Vitória (around 20°19'S 40°21'W).
Material Examined. 4 ♂♂, 1 ♀, 1 nymph. Holotype: ♂ from BRAZIL, Espírito Santo, Pinheiros, Reserva
Biológica Córrego do Veado, Trilha da Anta, point 1, 09-11.VI.2011, Malaise, MTTavares et al. leg. [specimen
MT00046 in the cladistic analysis] (UFES). Allotype: ♀ from BRAZIL, Espírito Santo, Vitória, 18.VI.1981
(UFES). Paratypes: BRAZIL: 1 nymph from Espírito Santo, Linhares, Reserva Natural Vale, Estrada Baba de Boi,
point 18, 16-18.I.2011, yellow pan trap, APAguiar et al. leg. (UFES); 1 ♂ same data except Linhares, 25.VI.1990,
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JSSantos leg., Miobantia rustica, L.Jantsch det., 28.VII.1993 (MCTP); 1 ♂ from Espírito Santo, Santa Teresa,
Estação Biológica Santa Lúcia, T1, 19-23.X.2007, Malaise, MTTavares et al. leg. [MT00045] (UFES); 1 ♂ from
Espírito Santo, Domingos Martins, Mata Pico do Eldorado, point T3, 03-10.XII.2004, Malaise, MTTavares et al.
leg. [MT00090] (UFES).
Etymology. From the Latin word immanis, meaning “enormous, huge,” in reference to the large body of this
species in comparison with the other species of this genus.
FIGURE 27. Distribution maps for species of Miobantia with new records provided in the text and localities found on
literature.
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A REVISI ON OF MIOBANTIA GIGLIO-TOS
Miobantia nordestina Scherrer, n. sp.
(Figs. 10; 11H; 13F–G; 15E–F; 16G, J; 17B, G; 26; 27F)
Description, male holotype. Length of body 19.2 mm from head to tip of abdomen, of pronotum 4 mm, of
metazone 2.5 mm, of forecoxa 3.9 mm; of forewing 16.9; of hindwing 17.1; maximum width of head 3.2 mm,
prozone middle 1.2 mm, supracoxal dilatation 1.7 mm; metazone minimum width 0.9 mm.
Head (Figs 11H & 13F): Frontal sclerite longitudinally moderately short, less than 3.0 times as wide as long,
upper edge uniformly convex; small tubercle between each compound eye and toruli not pointed, represented by
low cuticle elevation; ocelli markedly large, wider than pedicel apex, distance between ocelli as large as their width
or larger, central ocellus about as elevated as lateral ocelli, lateral ocelli distinctly elliptical; vertex somewhat
straight, inconspicuously or not higher than imaginary line joining apex of eyes, with region between parietal
suture and eye not elevated towards parietal suture, slightly convex; parietal suture regularly marked.
Pronotum (Fig. 15E): With moderately long pilosity, about as long as wing pilosity, supracoxal dilatation
regularly sized, prozone middle width/supracoxal maximum dilatation between 0.71–0.75; margin without
denticulation.
Legs (Figs 16G & 16J): Forecoxa markedly longer than metazone; forefemora internal spine 6 with shape and
size more similar to spine 5 than spines 2 and 4, distance between internal spines 12 and 11 about same distance
between spines 10 and 11 or slightly shorter, internal spine 12 about same size of spine 10; foretibiae external spine
1 away from spine 2 by space equivalent to one spine, external spine 2 slightly longer than spine 3, with 10 internal
spines.
Wings: Forewing about same length than hindwing; vein M1 branched; vein CuA1 branches slightly sinuous.
Hindwing apical angle moderately acute, vein CuA1 branched, apex slightly short, rounded.
Abdomen (Fig. 17B): Supranal plate very short, subrounded, faintly pointed.
Genitalia (Fig. 26A): Ventral phallomere distal process short, with most marked sclerotization taking only
process tip, denticulation faintly present, tip slender but rounded, basal swelling absent; proximal process faintly
sclerotized, markedly projected, short, curved, distinctly turned posterad, with basal portion slender, apex with
slightly long, slender point; slit between posterior margin of ventral phallomere and proximal process deep. Left
dorsal phallomere with anterior process very long, very slender, slightly curved at base, oriented anterad, apex
about as wide as middle; expanded portion between anterior process and phalloid apophysis short, oriented anterad;
well sclerotized area between phalloid apophysis and membranous lobe which is marked, small, not reaching
membranous lobe; phalloid apophysis very long, slender, its basal portion projected to left one, subapically
markedly bent, apically almost straight, not sharp; membranous lobe wide; dorsal lamina posterior margin with left
portion projected, narrow, uniformly curved.
Color: Body entirely pale yellow, dorsally slightly darker than ventrally. Head: area between mouthparts and
toruli with few, irregular darkish marks, except by mandible apex, and maxillary terminal segment dark; scape pale
yellow, except by apical frontal dark mark; pedicel dark, with margins pale yellow; flagellum entirely light brown;
area between toruli and vertex with few, large, irregular darkish marks, mostly on transversal darker band between
eyes covering the ocelli cuticle elevation; head dorsal region with irregular darkish marks, transversal dark band on
it apical area. Pronotum: with large, irregular darkish marks; margin with few, inconspicuous dark spots restricted
to prozone; ventrally with black median stripe on its posterior half. Legs: entirely pale yellow, except by tarsomeres
apically dark, femora apically with two, tibiae with three large, inconspicuous transversal darkish bands. Wings:
grayish, except by forewing cells, hindwing apical cells with central inconspicuously brownish mark, very
translucent; forewing dark spots present only on main veins, crossveins without dark spots; vein R spotted, with
marks distinctly of about same sizes, spaced from each other by 2.0–3.0 times length of smallest spots; area around
vein M branches without darkish marks.
Female allotype. Length of body 19.2 mm from head to tip of abdomen, of pronotum 4.8 mm, of metazone 2.9
mm, of forecoxa 4.4 mm; maximum width of head 4.0 mm, prozone middle 1.5 mm, supracoxal dilatation 1.9 mm;
metazone minimum width 1.1 mm. General morphology and coloration similar to male, except by body larger,
more stout than male; vertex inconspicuously more elevated (Fig. 13G); ocelli vestigial; pronotum with distinct
denticulation along lateral margin, metazone markedly more robust (Fig. 15F); forecoxa anterior edge with median
distinct denticulation; apterous; supranal plate longer than in males, but not overlaying ovipositor, with sides
distinctly converging towards a subrounded apex (Fig. 17G). General body with slightly, inconspicuous green tint,
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more evident on tibiae and tarsi; head with darkish marks smaller, more densely distributed, darker band between
eyes less evident than in male; legs medially with few darkish spots, forefemora dorsally with two transversal,
large, faint darkish bands; abdomen with several dark spots, irregular dark marks.
Variability, male. Paratypes very similar to holotype, except by foretibiae with 9–10 internal spines; hindwing
with apex slightly but distinctly pointed; paratype from Feira de Santana with right forewing vein M-a1 and
hindwing vein CuA-a1 branched. General body dark marks and spots high variable on size and intensity; paratype
from Pindobaçu with forewing with evanescent darkish brown marks between veins.
Comments. Males of this species are most similar to those of M. arctissima and M. sulista, based on the very
translucent wings, with costal margins of both wings without bright marks, the forewing costal area without dark
spots, the forewing vein R with dark spots of about same sizes, with vein M1 branched, and the hindwing vein
CuA1 also branched. It can however be securely distinguished from males of those two species by having the head
with frontal sclerite longitudinally moderately short, less than 3.0 times as wide as long, the vertex general
elevation inconspicuously higher than imaginary line joining apex of eyes, the hindwing apex rounded, the genitalia
with ventral phallomere distal process most marked sclerotization taking only process tip, its tip slender but
rounded, the proximal process projection short, and the left dorsal phallomere with phalloid apophysis very long.
Females of M. nordestina are most similar to those of M. fuscata due to have the combination of head with
vertex distinctly convex, frontal sclerite markedly short, and supranal plate with sides distinctly converging
towards a subrounded apex. It can however be separated from females of M. fuscata by having the vertex region
between parietal suture and eye distinctly, but slightly convex, pronotum with metazone markedly robust; and
forefemora with internal spine 12 about same size of spine 10.
Distribution (Fig. 27F). Brazil. Recorded from three localities of Bahia: Cachoeira da Fumaça, Pindobaçu,
Feira de Santana (10°28'43"S 40°12'27"W); Serra de São José, district of Maria Quitéria, Feira de Santana
(12°09'19"S 38°59'48"W); and Serra da Jibóia, Pedra Branca, Santa Terezinha (type locality) (12°51'S 39°28'W).
Material Examined. 3 ♂♂, 1 ♀. Holotype: ♂ from BRAZIL, 1 ♂ from Santa Terezinha, Pedra Branca, Serra
da Jibóia, 18.IX.2009, manual noturna, MMenezes leg. [specimen MZUEFS46390 in the cladistic analysis]
(MZUEFS). Allotype: ♀ from BRAZIL, Santa Terezinha, Pedra Branca, Serra da Jibóia, 24.VII.2009, manual,
MMiranda & MMenezes leg. (MZUEFS). Paratypes: BRAZIL: 1 ♂ from Feira de Santana, Pindobaçu, Cachoeira
da Fumaça, 18.XII.2009, lighting trap, EMenezes et al. leg. (MZUEFS); Bahia, Serra de São José, 10.V.2010,
manual, MMenezes leg. (MZUEFS).
Etymology. From the Potuguese word nordestina, meaning “northeastern” in reference to localities where the
examined specimens were collected, in Bahia state, northeastern Brazil.
Miobantia phryganea (Saussure, 1869)
(Figs. 5; 6A; 27D)
Miopteryx phryganea Saussure, 1869: 64; Kirby, 1904: 274; Giglio-Tos, 1917b:154. Two syntypes ♂♂ (Natural History
Museum of Geneva, Switzerland - MHNG), one of them examined through digital images.
= Miopteryx pryganea Saussure, 1870: 237 (misspelled).
Antimiopteryx phryganea: Giglio-Tos 1927: 211; Beier, 1935: 7
Miobantia phryganea: Jantsch & Corseuil, 1988: 228; Terra, 1995: 43; Ehrmann, 2002: 225; Agudelo et al., 2007: 120;
Ehrmann & Koçak, 2009: 14
Comments. Saussure (1869) proposed Miopteryx phryganea for two males syntypes examined by him. The
examined pictures showed one of the syntypes is in good condition, whereas the other one lacks both forelegs (Peter
Schwendinger pers. comm.). No lectotype is here designed. This species was considered by Kirby (1904) as junior
synonym of M. ciliata. Giglio-Tos (1917b) however disagreed and kept these two species separated. The results of
the present investigation supports Giglio-Tos’ opinion, with males of M. phryganea differing from those of M.
ciliata by having the supracoxal dilatation slightly pronounced, prozone middle width/supracoxal maximum
dilatation between 0.76–0.80, forewing much shorter than hindwing, and forewing vein R entirely dark or almost so
(Fig. 5). Males of Miobantia phryganea are also similar to M. rustica because they have similar shape and
coloration pattern of the wings, but it can be easily separated from this species by having the distance between ocelli
as large as their width or larger, the forewing much shorter than hindwing, and the apex of hindwing
inconspicuously pointed.
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Distribution (Fig. 27D). Brazil. The exact type locality is unknown. Terra (1995) recorded this species for the
locality of Santo Amaro (23°38'60"S 46°42'00"W), district of São Paulo, São Paulo state, Brazil. Agudelo,
Lombardo and Jantsch (2007) indicated the occurrence of this species also in the states of Rio de Janeiro, São
Paulo, Paraná, Santa Catarina, and Rio Grande do Sul, Brazil, based on a previous study of Jantsch (1999).
However, one male identified by such author as M. phryganea and included in the original distribution record made
by him in 1999, from Porto Alegre (Rio Grande do Sul) was examined and taken as paratype of M. sulista.
Although these records were based also on other specimens, this misidentification suggests the need for more
detailed investigations of such data. The other specimens examined by Jantsch (1999) were recorded from the
locality of E. P. Rio Grande, Rio de Janeiro (around 22°55'S 43°26W); three localities of Paraná: Morretes (around
25°28'S 48°49'W), Fênix (around 23°53'S 51°57'W), and Pinhão (25°43'S 51°37'W); the locality of Florianópolis,
Santa Catarina (around 27°35'S 48°34'W); and six localities of Rio Grande do Sul: Novo Hamburgo (around
29°41'S 51°08'W); a second specimen from Porto Alegre (around 30°02'S 51°12'W); Viamão (around 30°05'S
51°02'W); Guaíba (around 30°07'S 51°19'W); Gravataí (around 29°57'S 50°59'W); and Sapiranga (around 29°38'S
51°00'W).
Material Examined. Images of the syntype: ♂ from BRAZIL, Miopteryx phryganea Saussure, Syntypus.
Photographs taken by Peter Schwendinger (MHNG).
Miobantia rustica (Fabricius, 1781)
(Figs. 6; 11D; 12G; 14G; 16H; 17C; 22; 27E)
Mantis rustica Fabricius, 1781: 350; 1787: 229; 1793: 23; Gmelin-Linné, 1790: 2051; Olivier, 1792: 630. Lectotype ♂ (Natural
History Museum, London, England - BM), here designed, examined through digital images.
Miopteryx rustica: Saussure, 1871: 260; 1872, 277; Chopard, 1913: 76; Kirby, 1904: 274.
Antimiopteryx rustica: Giglio-Tos 1927: 211–212; Beier, 1935: 7.
Miobantia rustica: Jantsch & Corseuil, 1988: 228; Terra, 1995: 43; Ehrmann, 2002: 225–226; Agudelo et al., 2007: 120;
Ehrmann & Koçak, 2009: 14.
= Miopteryx perloides Saussure, 1869: 64; 1870: 237.
= Miopteryx nebulosa Giglio-Tos, 1915: 139 (n. syn.)
Antimiopteryx nebulosa: Giglio-Tos, 1927: 212; Beier: 1935: 7.
Miobantia nebulosa: Jantsch & Corseuil, 1988: 228; Terra, 1995: 43; Ehrmann, 2002: 225; Agudelo et al., 2007: 120; Ehrmann
& Koçak, 2009: 14.
Redescription, male lectotype. Length of body about 20.3 mm.
Head (e.g., Figs 11D & 12G): Frontal sclerite longitudinally very short, more than 3.0 times as wide as long,
upper edge uniformly convex; small tubercle between each compound eye and toruli not pointed, represented by
low cuticle elevation; ocelli medium-sized, about as wide as pedicel apex, distance between ocelli slightly
narrower than their width, central ocellus about as elevated as lateral ocelli, lateral ocelli distinctly elliptical; vertex
markedly convex, distinctly higher than imaginary line joining apex of eyes, with region between parietal suture
and eye more elevated towards parietal suture, nearly straight; parietal suture faintly marked.
Pronotum (e.g., Fig. 14G): With moderately long pilosity, about as long as wing pilosity, supracoxal dilatation
slightly pronounced, prozone middle width/supracoxal maximum dilatation between 0.76–0.80; margin without
denticulation.
Legs (Fig. 16H): Forecoxa markedly longer than metazone; forefemora internal spine 6 with shape and size
more similar to spine 5 than spines 2 and 4, distance between internal spines 12 and 11 about same distance
between spines 10 and 11 or slightly shorter, internal spine 12 about same size of spine 10; foretibiae external spine
1 away from spine 2 by space equivalent to one spine, external spine 2 slightly longer than spine 3, with 9 internal
spines.
Wings (Fig. 6C): Vein M1 branched. Hindwing apical angle moderately acute.
Genitalia (Fig. 22C): Ventral phallomere distal process very long, with most marked sclerotization taking at
least half area of the process, denticulation absent, tip pointed, basal swelling absent; proximal process markedly
sclerotized, markedly projected, short, curved, turned diagonally posterad central portion, with basal portion very
wide, apex with short, robust point; slit between posterior margin of ventral phallomere and proximal process
shallow. Left dorsal phallomere with anterior process short, markedly large, straight, oriented to left, apex about as
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wide as middle; expanded portion between anterior process and phalloid apophysis short, oriented to left; well
sclerotized area between phalloid apophysis and membranous lobe which is indistinct; phalloid apophysis very
short, moderately stout, its basal portion projected to the back, subapically markedly bent, apically almost straight,
not sharp; membranous lobe wide; dorsal lamina posterior margin with left portion not projected, uniformly
curved.
Color: Body entirely light brown to pale yellow. Head: area between mouthparts and toruli pale yellow; area
between toruli and vertex with small, few, irregular dark marks, transversal darker band between eyes covering the
ocelli cuticle elevation markedly evanescent, between ocelli absent, except around ocelli. Pronotum: dorsally with
irregular dark marks, mostly laterally; margin with moderately large dark spots more evident on prozone. Legs:
generally with dark spots. Wings: light brown, very translucent; forewing dark spots present, on main and
crossveins, spots very marked, crossveins spots most evident on costal area; vein R spotted, with marks distinctly
of about the same size, spaced from each other by 2.0–3.0 times length of smallest spot.
Complementary description, male holotype of Miopteryx nebulosa n. syn. Length of body 22.1 mm from
head to tip of abdomen, of pronotum 4.9 mm, of metazone 3.1 mm, of forecoxa 4.6 mm; of forewing 18.0; of
hindwing 19.3; maximum width of head 3.7 mm, prozone middle 1.3 mm, supracoxal dilatation 1.8 mm; metazone
minimum width 1.0 mm.
Wings (Fig. 6A): Forewing slightly shorter than hindwing; vein M1 branched; vein CuA1 branches markedly
sinuous. Hindwing apical angle moderately acute, vein CuA1 branched, apex slightly short, rounded.
Abdomen: Supranal plate very short, subrounded, faintly pointed.
Color: Body entirely light brown to pale yellow. Head: area between mouthparts and toruli pale yellow, with
few, small dark spots, except by mandible apex, and maxillary terminal segment, dark, frontal sclerite with upper
corners dark; scape pale yellow, except by apical frontal dark mark; pedicel dark, with margins pale yellow; area
between around toruli and vertex, head posterior area darkish brown, with small, few, irregular dark marks,
transversal darker band between eyes covering the ocelli cuticle elevation markedly present, except between ocelli,
but present around it. Pronotum: dorsally with irregular dark marks, mostly laterally; margin with moderately large
dark spots; ventrally with dark spots more sparsely distributed, black median stripe on its posterior 0.10. Legs: with
several dark spots, tarsomeres apically darkish, tarsomere 1 centrally darkish; forecoxa entirely, trochanter mostly
medially with larger dark spots; forefemora with dark spots of moderate, with three large, transversal dark marks
more evident medially; fore- and midtibia with three large transversal of dense concentration of faint dark spots,
one at each extremity and another at middle, mid- and hindfemur with two on its apical half. Mid- and hindlegs
with dense darkish spots of irregular sizes. Wings: light brown, very translucent; forewing dark spots present on
main and crossveins, crossveins spots most evident on costal area; vein R spotted, with marks distinctly of about
same sizes, spaced from each other by 2.0–3.0 times length of smallest spots; area around vein M branches with
two darkish marks on right forewing, one on left forewing. Abdomen: with dark spots densely distributed; dorsally
with median, longitudinal large dark band.
Female. Unknown.
Variability, male. Specimens from QBUM and coloration very similar to described specimens, except by dark
marks and spots high variable on size and intensity, in individual collected at Estrada Rio-São Paulo in 1947 spots
rare, inconspicuous on pronotum and forelegs. Transversal darker band between eyes covering the ocelli cuticle
elevation faint to restricted only to ocelli margin.
Comments. Fabricius’ (1781) description of Mantis rustica was based on two syntypes. Martin Stiewe
(Natural History Museum, London) dissected the male genitalia of both types and found no differences between
them (George Beccaloni pers. comm.). The imaged syntype is proposed as lectotype because it is in better
condition; the other is lacking the head (George Beccaloni pers. comm.).
Males of this species are most similar to M. ciliata and M. phryganea because they have similar shape and
coloration pattern of the wings, and the vertex is convex. However, M. rustica can be easily separated from males
of M. ciliata by having the supracoxal dilatation slightly pronounced, prozone middle width/supracoxal maximum
dilatation between 0.76–0.80, the forewing vein R spotted, the ventral phallomere with distal process very long,
proximal process with short and robust point, and the left dorsal phallomere with phalloid apophysis very short and
basally projected to the back. It can also be isolated from males of M. phryganea by having the distance between
ocelli slightly narrower than their width, forewing only slightly shorter than hindwing, and apex of hindwing more
rounded.
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The holotype of Miopteryx nebulosa was examined and it shares with the lectotype of Miobantia rustica all the
same character states mentioned above. In fact, Giglio-Tos (1915) originally described it as similar to Miopteryx
rustica, except by having the central ocellus not pronounced, vertex testaceous, transversal darker band between
eyes passing over ocelli cuticle elevation markedly present, forewing more slender and longer, with crossveins of
costal area darker, the discoidal area with two dark larger marks, hindwings with costal area apically opaque-
ferruginous. However, the material here in examined pointed such characters as being highly variable within the
other species of the genus, especially the dark band between the eyes, which varies even among the additional
material examined for this species. Additionally, the male genitalia of the types of both species was examined and
showed no relevant differences (Figs 22A & 22C). Therefore, Miopteryx nebulosa is here designed as junior
synonym of Miobantia rustica and is taken for complementing the redescription of this species.
Distribution (Fig. 27E). The type locality is the coastal Patagonia, Argentina. Recorded also from Brazil, on
two localities of Rio de Janeiro: Fazenda Peuedo, Itatiaia (around 22°30'S 44°34'W); and Km 47 of Rio-São Paulo
road (around 22°46'S 43°41'W). Terra (1995) recorded this species for three localities of São Paulo, Brazil:
Fazenda Pau d’Alho, Itu (23°08'38"S 47°40'45"W); Floresta Estadual Edmundo Navarro de Andrade, Rio Claro
(22°24'43"S 47°33'5"W); and São Sebastião (around 23°48'S 45°25'W). Though not mentioned in the given
distribution of the species, Terra (1995) also listed examined material for M. rustica from Viçosa, on Minas Gerais
(around 20°45'S 42°53'W), and Angra dos Reis, on Rio de Janeiro (around 23°00'S 44°18'W). Agudelo et al.
(2007) also indicated the occurrence of this species in the state of Espírito Santo, based on a previous study of
Jantsch (1999). However, the particular specimen identified by such author as M. rustica and included in the
distribution record made by him in 1999, from Linhares, in Espírito Santo, was examined and taken as paratype of
M. immanis herein described.
Miopteryx nebulosa is recorded from Brazil (exact type locality unknown), but Terra (1995) added the record
for Santo Amaro, São Paulo district, on São Paulo (23°38'60"S 46°42'00"W).
Material Examined. 4 ♂♂. Described male: ♂ from BRAZIL, Miopteryx nebulosa Giglio-Tos typus, E.
Giglio-Tos det., Holotypus Nr., Mantis, Ca. No. 653. (ZMB). Other specimens: BRAZIL: 1 ♂ from Rio de Janeiro,
Itatiaia, Fazenda Peuedo, 15.XI.1942, Wygod. (QBUM); 1 ♂ from Rio de Janeiro, Estrada Rio-São Paulo, Km 47,
II.1945, Wygod.; 1 ♂ same data except 3.I.1947. Images of the lectotype: ♂ from ARGENTINA, Patagonia, 63 47,
BMNH(E)#877364, BMNH(E)#669455. Type specimen habitus photographed by Gavin Svenson (Cleveland
Museum of Natural History), and their genitalia dissected and photographed by Martin Stiewe (Natural History
Museum, London).
Miobantia sulista Scherrer, n. sp.
(Figs. 9; 11G; 13D–E; 15C–D; 16F; 17F; 25; 27F)
Description, male holotype. Length of body 17.5 mm from head to tip of abdomen, of pronotum 4.0 mm, of
metazone 2.4 mm, of forecoxa 4 mm; of forewing 16.9; of hindwing 17.3; maximum width of head 3.2 mm,
prozone middle 1.2 mm, supracoxal dilatation 1.7 mm; metazone minimum width 1.0 mm.
Head (Figs 11G & 13D): Frontal sclerite longitudinally very short, more than 3.0 times as wide as long, upper
edge uniformly convex; small tubercle between each compound eye and toruli not pointed, represented by low
cuticle elevation; ocelli medium-sized, about as wide as pedicel apex, distance between ocelli as large as their
width or larger, central ocellus about as elevated as lateral ocelli, lateral ocelli distinctly elliptical; vertex slightly
convex, distinctly higher than imaginary line joining apex of eyes, with region between parietal suture and eye
more elevated towards parietal suture, slightly convex; parietal suture regularly marked; vertex region between
parietal sutures laterally abruptly elevated close to parietal suture, centrally following lateral part overall outline.
Pronotum (Fig. 15C): With moderately long pilosity, about as long as wing pilosity, supracoxal dilatation
markedly pronounced, prozone middle width/supracoxal maximum dilatation less than 0.70; margin without
denticulation.
Legs (e.g., Fig. 16F): Forecoxa markedly longer than metazone; forefemora internal spine 6 with shape and
size more similar to spine 5 than spines 2 and 4, distance between internal spines 12 and 11 about same distance
between spines 10 and 11 or slightly shorter, internal spine 12 about same size of spine 10; foretibiae external spine
1 away from spine 2 by space equivalent to one spine, external spine 2 slightly longer than spine 3, with 9 internal
spines.
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Wings (Fig. 9A): Forewing about same length than hindwing; vein M1 branched; vein CuA1 branches
markedly sinuous. Hindwing apical angle moderately acute, vein CuA1 branched, apex slightly long, markedly
pointed.
Abdomen: Supranal plate very short, subrounded, faintly pointed.
Genitalia (Fig. 25B): Ventral phallomere distal process slightly long, with most marked sclerotization taking
entire process, denticulation markedly present, tip pointed, basal swelling present; proximal process markedly
sclerotized, markedly projected, long, somewhat straight, turned diagonally posterad lateral portion, with basal
portion slightly wide, apex rounded; slit between posterior margin of ventral phallomere and proximal process
deep. Left dorsal phallomere with anterior process very long, moderately large, markedly curved at about middle,
oriented anterad, apex distinctly wider than middle width; expanded portion between anterior process and phalloid
apophysis short, oriented anterad; well sclerotized area between phalloid apophysis and membranous lobe which is
marked, large, slender; phalloid apophysis regularly long, slender, its basal portion projected to left one, somewhat
contiguously curved with entire phalloid apophysis, apically concave, not sharp; membranous lobe slender; dorsal
lamina posterior margin with left portion projected, expanded, uniformly curved.
Color: Body dorsally darkish brown, ventrally pale yellow. Head: area between mouthparts and toruli entirely
pale yellow, except by mandible apex, and maxillary terminal segment dark; scape pale yellow; pedicel dark, with
margins pale yellow; flagellum entirely light brown; area between toruli and vertex with several minute light
brownish spots, mostly on transversal darker band between eyes covering the ocelli cuticle elevation; head dorsal
region with few, faint irregular darkish marks. Pronotum: dorsally with irregular darker marks; margin few, faint,
small dark spots; ventrally with black median stripe on its posterior half. Legs: pale yellow, with darkish marks,
tarsomeres apically darkish; forecoxa medially with few, faint dark spots; forefemora medially with three areas of
dense concentration of evanescent spots, restricted to anterior area, posteriorly with basal area, claw groove
brownish. Mid- and hindleg femur, tibia without dark spots or marks. Wings: faintly light brownish, with few,
evanescent, irregularly sized darkish marks mostly on forewing; forewing dark spots present only on main veins,
crossveins without dark spots; vein R spotted, with marks distinctly of about same sizes, very away from each other
by more than 3 times length of smallest spots; area around vein M branches without darkish marks.
Female allotype. Length of body 18.5 mm from head to tip of abdomen, of pronotum 4.7 mm, of metazone 2.8
mm, of forecoxa 4.6 mm; maximum width of head 3.8 mm, prozone middle 1.6 mm, supracoxal dilatation 2.1 mm;
metazone minimum width 1.4 mm. General morphology and coloration similar to male, except by body larger,
more stout than allotype; head with vertex distinctly more elevated (Fig. 13E); ocelli vestigial; pronotum with
distinct denticulation along lateral margin, more evident posteriorly, metazone markedly more robust (Fig. 15D);
forecoxa anterior edge with median distinct denticulation; apterous; supranal plate longer than in males, but not
overlaying ovipositor, with sides distinctly converging towards a subrounded apex (Fig. 17F); area between
mouthparts and toruli entirely with few dark spots, mostly on labrum; pronotum dorsally light brown to pale
yellow, with small dark spots, mostly laterally; foreleg with tarsomere 1 centrally darkish, coxa, femur, tibia with
more darkish spots, darkish marks more evident than in males; fore- and midtibia with three large transversal areas
of dense concentration of dark spots, one at each extremity and another at middle, mid- and hindfemur with two
faint transversal dark bands on its apical half; abdomen with several darkish spots, irregular dark marks.
Variability, male. Male paratypes very similar to holotype, except by foretibiae with 9–10 internal spines. General
body dorsally pale yellow in paratypes, dark marks and spots high variable on size and intensity; head with area
between toruli and vertex with transversal darker band between eyes sometimes absent, only ocelli margin marked
dark.
Female. Female paratypes very similar to allotype, except by female from Garabi with foretibiae with 9
internal spines; body dark marks distinctly less marked; both female paratypes with limit between supracoxal
dilatation and metazone distinctly evident, margin with denticulation inconspicuous on prozone. Female from
Garabi with dark marks distinctly less marked.
Comments. Males of this species are most similar to those of M. arctissima and M. nordestina, based on the
very translucent wings, with costal margins of both wings without bright marks, the forewing costal area without
dark spots, the forewing vein R with dark spots of about same sizes, with the vein M1 branched, and the hindwing
vein CuA1 also branched. However, it can be easily isolated from males of these two species by having the ocelli
medium-sized, about as wide as pedicel apex, vertex slightly convex, with region between parietal sutures laterally
abruptly elevated next to the parietal suture, supracoxal dilatation markedly pronounced, prozone middle width/
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supracoxal maximum dilatation less than 0.70, hindwing apex distinctly pointed, and ventral phallomere with distal
process slightly long, pointed, with basal swelling present.
The female of M. sulista is somewhat similar to those of M. fuscata and M. nordestina due to the presence of a
supranal plate with sides distinctly converging towards a subrounded apex, but it can be readily distinguished from
those species by having the vertex distinctly convex and the frontal sclerite markedly short (vs. vertex somewhat
straight and frontal sclerite moderately short).
Distribution (Fig. 27E). Recorded from San Bernardino, Paraguay (around 25°16'S 57°19'W), and Brazil, for
Nova Teutônia, Santa Catarina (27°11'S 52°23'W), and on three localities of Rio Grande do Sul: Garabi; São Borja
(28°39'S 56°00'W); and Porto Alegre (type locality) (30°02'S 51°12'W).
Material Examined. 4 ♂♂, 3 ♀♀. Holotype: ♂ from BRAZIL, Rio Grande do Sul, Porto Alegre, 03.II.1985,
Jantsch leg., Miobantia ciliata, LJantsch det., 0776 [specimen MCTP0776 in the cladistic analysis] (MCTP).
Allotype: ♀ from PARAG U AY, 3952, KFiebrig, 27.VII.1905, F. Nº 803/05, [paratype of] Miobantia aptera Giglio-
Tos, E. Giglio-Tos det. (ZMB). Paratypes: BRAZIL: 2 ♂♂ from Santa Catarina, Nova Teutônia, X.1995, Fritz
Plaumann leg., G1115 (QBUM); 1 ♀ from Rio Grande do Sul, Garabi, 3-12.I.1989, ELerner leg., Miobantia aptera
L.Jantsch det., 1505 (MCTP); 1 ♀ from Rio Grande do Sul, São Borja, 08.VII.1980, AALise leg., Miobantia
aptera L.Jantsch det., 0469 (MCTP); 1 ♂ from Porto Alegre, 25.XI.1988, ALAzevedo leg., Miobantia phryganea
L.Jantsch det., 1504 (MCTP).
Etymology. From the Portuguese word sulista, meaning “southern,” in reference to localities where the
examined specimens were collected, in Paraguay and southern Brazil.
Concluding Remarks
The use of both morphological and molecular analyses allowed the successful separation of species of Miobantia,
overcoming the challenges of the high sexual dimorphism, a recurrent issue for taxonomic studies in Thespidae.
Thus, four new species have been described: M. immanis n. sp., M. arctissima n. sp., M. sulista n. sp., and M.
nordestina n. sp.. Miobantia nebulosa has been designated new synonym of M. rustica. The male of M. aptera, and
the females of M. ciliata and M. fuscata were described for the first time. Immature stages are also first described
for M. aptera, M. fuscata, and M. immanis.
Additionally, 50 new records for the species of the genus were herein presented and, along with a extensive
literature review, the distribution of all Miobantia were mapped. The northernmost record for this genus is now
northern Bahia, Brazil, having been previously recorded only as far north as Espírito Santo, also in Brazil. It is
important to mention that the misidentifications of M. aptera and M. ciliata by Jantsch, commented in the
descriptions above, showed that the distribution records of his study from 1999, as well as the distributions
reported by Agudelo, Lombardo, and Jantsch (2007) accordingly with those of Jantsch (1999), need to be
considered critically.
The DNA barcoding for association of males, females, and immature stages proved to be effective regardless
the high values and saturation level of transition mutations present in the COI sequences of Miobantia. However, it
may be less useful with small sets of samples, or without other species to compare. Tests with sequencing of more
conservative DNA regions of these mantises can possibly lead to more optimal choices of molecular markers that
may be effective with smaller taxon sample numbers. It is also remarkable that the genetic distances of such COI
sequences were very high even between intraspecific groups. The lack of such investigations in other Mantodea
groups makes it impossible to compare the results obtained in this work with other related data. New studies on
DNA barcoding for other mantid populations may help to understand what is happening with the evolution of this
gene in this group.
Acknowledgments
Graduate scholarships were provided by the Conselho Nacional de Desenvolvimento Científico e Tecnológico /
CNPq, Brazil. This work benefitted directly from research funding provided by Fundação de Amparo à Pesquisa
do Espírito Santo/FAPES (grant 45.440.611/2009), and CNPq (grants 480507/2010-4 and 620068/2008-6), to my
advisor, Alexandre P. Aguiar (UFES/Brazil). The curators Gervásio S. Carvalho (MCTP), Freddy Bravo
(MZUEFS), Miguel A. Monné (QBUM), Marcelo T. Tavares (UFES), and Michael Ohl (ZMB) helped with
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valuable loans of Miobantia specimens. Photographs of the type of M. ciliata are a courtesy of the Naturhistoriska
Riksmuseets (Stockholm, Sweden), and were kindly sent by Gunvi Lindberg and Kjell A. Johanson. Peter
Schwendinger (Muséum d'Histoire Naturelle, Switzerland), taked the photographs of M. phryganea. The
photographs of the type of M. rustica were provided by George Beccaloni and are copyrighted to the Natural
History Museum, London (England). Gavin Svenson (Cleveland Museum of Natural History) kindly took the
images of the general morphology and labels of this type during his visit to the Natural History Museum, and
Martin Stiewe dissected and imaged the male genitalia. Eliomar Menezes promptly helped with the process of
obtention of the loan from MZUEFS. The molecular methodology was supported by Valéria Fagundes (UFES),
who helpfully provided the technical equipments and lab workspace, as well as by the help of many colleagues of
the Laboratório de Genética Animal and of the Núcleo de Genética Aplicada à Conservação da Biodiversidade
(UFES) especially Mariana S. Azevedo, Lucas A. Vianna, Arturo B. Martinelli, Lorena L. Dinelli, Victor H.
Collombi, Rosana R. A. Nunes, Cristina D. A. Nogueira, and Juliana F. Justino. Alexandre P. Aguiar (UFES)
contributed with many important suggestions to the manuscript and fully supported this research throughout its
development; without his comments and assistance the present work would be quite limited. Mariana, and the lab
colleagues Adriana C. B. Ramos, Anazélia M. Tedesco, Bernardo F. Santos, Fabiana G. Rampinelli, and Maria C.
Carreiro, also helped with valuable discussions, in addition to contributions with collecting and sorting of many
specimens of Miobantia. Gavin Svenson and Celso O. Azevedo (UFES) contributed with many enriching
comments on the manuscript.
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APPENDIX 1
Character list used for the codification of the cladistic matrix.
Head
1. Length of frontal sclerite: [0] very short, ratio width/length greater than 3; [1] moderately short, width/
length less than 3
2. Alignment of lateral margins of frontal sclerite: [0] distinctly converging anteriorly; [1] somewhat
parallel or inconspicuously converging anteriorly
3. Development of small tubercle between each compound eye and toruli: [0] prominent, in form of low
crest moderately pointed; [1] very low, represented by low cuticle elevation
4. Ocelli size: [0] large, larger than pedicel apex; [1] medium-sized, about as wide as pedicel apex; [2]
slightly small, narrower than pedicel apex; [3] markedly small, vestigial
5. Ocelli proximity: [0] regularly distant to one another, the distance between ocelli as large as their width or
larger; [1] slightly close to one another, the distance between ocelli slightly narrower than their width; [2]
very close to one another, the distance between much narrower than the width of a ocellus
6. Relative height of central ocellus: [0] about as elevated as lateral ocelli; [1] distinctly more elevated than
lateral ocellus
7. Shape of cuticle projection of central ocellus: [0] somewhat conical; [1] almost rounded, slightly medially
pointed; [2] distinctly rounded
8. Form of central ocellus: [0] distinctly transversally elliptical; [1] somewhat rounded
9. Form of lateral ocelli: [0] rounded; [1] distinctly elliptical
10. Depth of parietal suture: [0] shallow; [1] deep
11. Overall shape of vertex: [0] markedly convex (e.g., Fig. 12C); [1] slightly convex (e.g., Fig. 12A); [2]
somewhat straight (e.g., Fig. 13E)
12. General elevation of vertex: [0] inconspicuously or not higher than imaginary line joining apex of eyes;
[1] distinctly higher than imaginary line joining apex of eyes
13. Form of region of vertex between parietal suture and eyes: [0] straight or slightly convex; [1] slightly
high, forming smooth bulge; [2] very high, forming marked juxtaocular tubercle
14. Angle of region between parietal suture and eyes: [0] not elevated towards parietal suture; [1] more
elevated towards parietal suture
15. Form of lateral part of vertex region between parietal suture: [0] abruptly elevated close to parietal suture;
[1] gradually elevated close to parietal suture; [2] continuously lined with remaining regions of vertex
16. Form of central part of vertex region between parietal suture: [0] continuously lined with lateral part
overall outline; [1] concave
Pronotum
17. Density of pronotum pilosity: [0] very sparse, the distance between each pilosity about as wide as its
height or wider; [1] moderately dense, the distance between each pilosity narrower than its height
18. Length of pronotum pilosity: [0] slightly long, about as long as wing pilosity or longer; [1] short, shorter
than wing pilosity
19. Presence of margin denticulation: [0] absent; [1] present
20. Development of median keel of prozone: [0] faint; [1] moderately marked, with anterior part very low to
indistinctly marked
21. Supracoxal dilatation: [0] markedly pronounced, prozone middle width/supracoxal maximum dilatation
less than 0.70; [1] regularly sized, prozone middle width/supracoxal maximum dilatation between
0.71–0.75; [2] slightly pronounced, prozone middle width/supracoxal maximum dilatation between
0.76–0.80; [3] very faintly pronounced, prozone middle width/supracoxal maximum greater than 0.80
22. Development of median keel of metazone: [0] faint; [1] marked
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23. Size of pair of tubercles at posterior area of metazone: [0] moderately low or high, a pair of bulges or
crests; [1] very low, cuticle inconspicuously elevated
Meso- and Metanotum
24. General shape of triangular expansion of posterior margin of mesonotum: [0] small; [1] slightly large; [2]
very large
25. Tip length of triangular expansion of posterior margin of mesonotum: [0] short; [1] slightly long; [2] very
long
26. Pilosity length of triangular expansion of jugal field of mesonotum: [0] short; [1] high
27. Size of meso- and metanotum median longitudinal pilosity: [0] same pattern of other areas of thorax; [1]
higher than pilosity of other areas of thorax
Legs
28. Forecoxa relative length: [0] about same length of metazone or slightly longer; [1] much longer than
metazone
29. Relative distance of external spines 1 and 2 of forefemora : [0] markedly shorter than distance of spines 2
and 3; [1] slightly shorter than distance of spines 2 and 3; [2] same distance of spines 2 and 3
30. Relative size of internal spine 6 of forefemora : [0] more similar to spines 2 and 4 than spine 5; [1] more
similar to spine 5 than spines 2 and 4
31. Relative distance of internal spines 12 and 11 of forefemora : [0] distinctly longer than distance of spines
10 and 11; [1] about same distance of spines 10 and 11 or slightly shorter
32. Relative size of internal spine 12 of forefemora : [0] distinctly shorter than spine 10; [1] about same size
of spine 10
33. Number of external spines of foretibia: [0] 5; [1] 6; [2] 7
34. Distance of base to external spine 1 of foretibia: [0] about 0.2 of tibia length; [1] about 0.4 of tibia length
35. Distance between external spine 1 and 2 of foretibia: [0] distinctly larger than distance between spine 2
and 3; [1] as large as distance between spine 2 and 3
36. Length of distance of external spine 1 of foretibiae from spine 2, if markedly distant: [0] equivalent to two
spines; [1] equivalent to one spine
37. Size of external spine 2 of foretibia: [0] slightly shorter than spine 3; [1] same size of spine 3; [2] slightly
longer than spine 3
38. Number of internal spines of foretibia: [0] 9; [1] 10; [2] 11; [3] 13
39. Presence of transversal dark marks on hind femur: [0] indistinct or absent; [1] distinct
Wings
40. Wings general color: [0] light brownish; [1] dark brownish; [2] blackish
41. Uniformity of coloration: [0] uniform; [1] distinctly tarnished
42. Pattern of stains: [0] large, irregularly distributed; [1] medium, somewhat sparsely distributed; [2] small,
densely distributed
43. Translucency of wings: [0] very translucent, wings hyaline or almost so; [1] regularly translucent; [2]
faintly translucent, wing pigmentation very marked
44. Relative length of forewing: [0] longer than hindwing; [1] about same length than hindwing; [2] shorter
than hindwing
45. Presence of dark spots at forewing: [0] absent; [1] present
46. Locality of forewing dark spots: [0] only on main veins, crossveins without dark spots; [1] on main and
crossveins
47. Presence of dark spots at vein R of forewing: [0] indistinct, vein R entirely dark; [1] present
48. Form of dark spots at vein R of forewing: [0] some spots distinctly larger than other; [1] spots of about
equal size
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49. Presence of darkish marks around vein M branches of forewing: [0] absent; [1] present
50. Number of darkish marks around vein M branches of forewing: [0] 3; [1] 2; [2] 1
51. Branching of vein M1 of forewing: [0] vein M1 not branched; [1] vein M1 branched
52. Form of branches of vein CuA1 of forewing: [0] nearly straight; [1] sinuous, even if slightly
53. Ratio between AA1 length and hindwing maximum length: [0] very short, less than 0.82; [1] slightly
short, between 0.83-0.86; [2] slightly long, between 0.87-0.94; [3] very long, greater than 0.96
54. Hindwing angle between imaginary line tangent to costal border and imaginary line tangent to apical
border: [0] very acute, less than 30 degrees; [1] regularly acute, between 31–60 degrees; [2] slightly
acute, greater than 61 degrees
55. Presence of branches of vein CuA1 of hindwing: [0] branched; [1] not branched
Abdomen
56. Shape of supranal plate: [0] markedly pointed; [1] subrounded, slightly pointed; [2] rounded
57. Length of supranal shield: [0] short, distinctly less than 2.0 times as long as wide; [1] slightly long, about
2.0 times as long as wide; [2] very long, distinctly more than 2.0 times as long as wide
Male Genitalia, Ventral Phallomere
58. Most marked sclerotization of distal process of ventral phallomere: [0] taking entire process; [1] taking at
least half area of process; [2] taking only process tip
59. Presence of denticulation of distal process of ventral phallomere: [0] present, even if inconspicuously; [1]
absent
60. Intensity of denticulation of distal process of ventral phallomere: [0] denticulation evident; [1]
inconspicuous
61. Width of apex of distal process of ventral phallomere: [0] wide; [1] slender
62. Form of tip of distal process of ventral phallomere: [0] rounded; [1] pointed
63. Presence of basal swelling of distal process of ventral phallomere: [0] absent; [1] present
64. Basal width of distal process of ventral phallomere: [0] large, basal width distinctly greater than length;
[1] slender, process as wide as long or longer than wide
65. General length of distal process of ventral phallomere: [0] short, about as long as maximum width or
slightly longer; [1] slightly long, about 2.0–2.5 times as long as wide ; [2] very long, about 3.0 times as
longer as wide or larger
66. Sclerotization intensity of proximal process of ventral phallomere: [0] faintly sclerotized; [1] slightly
marked; [2] very marked
67. Insertion of proximal process of ventral phallomere: [0] continuous with posterior margin of ventral
phallomere; [1] with slit between it and posterior margin of ventral phallomere, even if very shallow
68. Projection of proximal process of ventral phallomere: [0] only slightly projected; [1] markedly projected
69. Length of projected proximal process of ventral phallomere: [0] short (e.g., Figs 22A, 26A); [1] long
(e.g., Fig. 19A)
70. Overall curvature of projected proximal process of ventral phallomere: [0] somewhat straight; [1] curved
71. Overall width of projected proximal process of ventral phallomere: [0] regular; [1] slender; [2] expanded
72. Apical width of projected proximal process of ventral phallomere: [0] slightly slender than base; [1]
distinctly slender than base
73. Direction of projected proximal process of ventral phallomere: [0] turned diagonally posterad lateral
portion; [1] distinctly turned posterad ; [2] turned diagonally posterad central portion
74. Slit between posterior margin of ventral phallomere and proximal process: [0] deep; [1] shallow
75. Basal width of projected proximal process of ventral phallomere: [0] regularly wide, basal width less than
3 times the length of process; [1] very wide, basal width greater than 4 times the length of process
76. Form of tip of projected proximal process of ventral phallomere: [0] rounded; [1] pointed
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Male Genitalia, Left Dorsal Phallomere
77. Presence of anterior process of left dorsal phallomere: [0] indistinct; [1] distinct
78. Length of anterior process of left dorsal phallomere: [0] very long, distinctly longer than ventral
phallomere half length; [1] slightly long, about as long as ventral phallomere half length or slightly
longer; [2] short, slightly shorter than ventral phallomere half length
79. Shape of anterior process of left dorsal phallomere: [0] curved, even if slightly; [1] straight
80. Curvature of anterior process of left dorsal phallomere: [0] markedly curved; [1] slightly curved
81. Width of anterior process of left dorsal phallomere, if large: [0] moderately large, process distinctly
longer than 2.5 times its width; [1] markedly large, about 2.5 times as long as wide or less
82. Direction of anterior process of left dorsal phallomere: [0] laterad; [1] anterad
83. Apical width of anterior process of left dorsal phallomere: [0] about same width than middle or
inconspicuously wider; [1] distinctly wider than middle width
84. Shape of expanded portion between anterior process and phalloid apophysis of left dorsal phallomere: [0]
not triangular; [1] triangular
85. Length of expanded portion between anterior process and phalloid apophysis of left dorsal phallomere:
[0] long; [1] short
86. Direction of expanded portion between anterior process and phalloid apophysis of left dorsal phallomere:
[0] forward; [1] lateral
87. Presence of well sclerotized area between phalloid apophysis and membranous lobe of left dorsal
phallomere: [0] indistinct; [1] evanescent; [2] marked
88. Extension of well sclerotized area between phalloid apophysis and membranous lobe of left dorsal
phallomere, if medium or small: [0] medium, expanded posterad ; [1] small
89. Width of small well sclerotized area between phalloid apophysis and membranous lobe of left dorsal
phallomere: [0] extended, reaching anterior area of membranous lobe; [1] small, not reaching
membranous lobe
90. Width of medium sized well sclerotized area between phalloid apophysis base and membranous lobe of
left dorsal phallomere: [0] wide (e.g., Fig. 23A); [1] slender (e.g., Fig. 25A)
91. Length of phalloid apophysis of left dorsal phallomere: [0] vestigial, about as long as wide; [1] very short,
about as long as half left dorsal phallomere width or less; [2] slightly short, slightly but distinctly longer
than half left dorsal phallomere width; [3] moderately long, its length between 0.75–1.00 the left dorsal
phallomere width; [4] very long, distinctly longer than left dorsal phallomere width
92. Width of phalloid apophysis of left dorsal phallomere: [0] moderately stout; [1] slender
93. Basal direction of phalloid apophysis of left dorsal phallomere: [0] right; [1] back; [2] left
94. Form of apical curvature of phalloid apophysis of left dorsal phallomere, if bent posterad : [0] not or
inconspicuously bent; [1] distinctly bent
95. Apical shape of phalloid apophysis of left dorsal phallomere: [0] concave, even if slightly; [1] almost
straight or convex
96. Apical width of phalloid apophysis of left dorsal phallomere: [0] contiguously wide with entire phalloid
apophysis; [1] slightly sharpen
97. General width of membranous lobe of left dorsal phallomere: [0] wide (e.g., Fig. 19A); [1] slender (e.g.,
Fig. 25A)
98. Form of left portion of posterior margin of dorsal lamina of left dorsal phallomere: [0] projected,
expanded, uniformly curved; [1] projected, distinctly bent; [2] projected, narrow, uniformly curved; [3]
not projected, uniformly curved
99. Size of apical process (titillator) of left dorsal phallomere: [0] large; [1] small
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APPENDIX 2. Label data of the specimens of the outgroup in cladistic analyses.
Anamiopteryx sp.: BRAZIL, Espírito Santo, Pinheiros, Reserva Biológica Córrego do Veado, Água Limpa, 27.XI-06.XII.2011,
Malaise, MTTavares, et al. leg.
Chloromiopteryx sp.: BRAZIL, Espírito Santo, Santa Teresa, Estação Biológica Santa Lúcia, 17-23.IV.2011, DBarbosa leg.
Eumusonia sp.: BRAZIL, Espírito Santo, Pinheiros, Reserva Biológica Córrego do Veado, 09.VI.2011, LAVianna leg.
APPENDIX 3. DELTA characters file used to generate the morphological descriptions.
#1. Frontal sclerite longitudinally/
1. very short, more than 3.0 times as wide as long/
2. moderately short, less than 3.0 times as wide as long/
#2. <Frontal sclerite> upper edge/
1. uniformly convex/
2. <bent>/
#3. Small tubercle between each compound eye and toruli <prominence>/
1. in form of slightly pointed crest/
2. not pointed, represented by low cuticle elevation/
3. indistinct/
#4. Ocelli <size>/
1. markedly large, wider than pedicel apex/
2. medium-sized, about as wide as pedicel apex/
3. slightly small, narrower than pedicel apex/
4. vestigial/
#5. <Ocelli proximity>/
1. distance between ocelli as large as their width or larger/
2. distance between ocelli slightly narrower than their width/
3. distance between ocelli much narrower than their width /
#6. Central ocellus <height>/
1. about as elevated as lateral ocelli/
2. distinctly more elevated than lateral ocelli by low tubercle/
#7. Lateral ocelli <form>/
1. rounded/
2. distinctly elliptical/
#8. Vertex <overall shape>/
1. markedly convex/
2. slightly convex/
3. somewhat straight/
4. laterally convex, centrally slightly concave/
#9. <Vertex general elevation>/
1. inconspicuously or not higher than imaginary line joining apex of eyes/
2. distinctly higher than imaginary line joining apex of eyes/
#10. <Vertex> with region between parietal suture and eye <angle>/
1. not elevated towards parietal suture/
2. more elevated towards parietal suture/
#11. <Vertex region between parietal suture and eye, dilatation>/
1. nearly straight/
2. slightly convex/
3. markedly convex, forming distinct, low bulge (juxtaocular bulge)/
#12. Parietal suture <depth>/
1. faintly marked/
2. regularly marked/
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#13. Vertex region between parietal sutures laterally/
1. abruptly elevated close to parietal suture/
2. gradually elevated close to parietal suture/
#14. <Vertex region between parietal sutures> centrally/
1. following lateral part overall outline/
2. slightly concave/
#15. <pilosity length> with/
1. very long pilosity, longer than wing pilosity/
2. moderately long pilosity, about as long as wing pilosity/
#16. Supracoxal dilatation/
1. markedly pronounced, prozone middle width/supracoxal maximum dilatation less than 0.70/
2. regularly sized, prozone middle width/supracoxal maximum dilatation between 0.71–0.75/
3. slightly pronounced, prozone middle width/supracoxal maximum dilatation between 0.76–0.80/
4. very faintly pronounced, prozone middle width/supracoxal maximum greater than 0.80/
#17. <Pronotum> margin/
1. without denticulation/
2. with distinct denticulation along entire lateral margin, more pronounced posteriorly/
#18. Forecoxa <relative length>/
1. slightly longer than metazone/
2. markedly longer than metazone/
#19. <Forecoxa> anterior edge/
1. with denticulation inconspicuous/
2. with median very faint denticulation/
#20. Forefemora internal spine 6 with shape and size <relative size>/
1. more similar to spines 2 and 4 than the spine 5/
2. more similar to spine 5 than spines 2 and 4/
#21. <Forefemora > distance between internal spines 12 and 11/
1. distinctly longer than distance between spines 10 and 11/
2. about same distance between spines 10 and 11 or slightly shorter/
#22. <Forefemora > internal spine 12 <relative size>/
1. distinctly shorter than spine 10/
2. about same size of spine 10/
#23. Foretibiae external spine 1 away from spine 2 by space equivalent to/
1. two spines/
2. one spine/
#24. <Foretibiae> external spine 2 <size>/
1. about same size of spine 3/
2. slightly longer than spine 3/
#25. <Foretibiae number of internal spines> with/
1. 0 internal spines/
2. 6 internal spines/
3. 9 internal spines/
4. 10 internal spines/
5. 11 internal spines/
6. 13 internal spines/
#26. Forewing <relative length>/
1. about same length than hindwing/
2. slightly shorter than hindwing/
3. much shorter than hindwing/
#27. <Forewing> vein M1 <number of branches>/
1. not branched/
2. branched/
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#28. <Forewing> vein CuA1 branches <form>/
1. nearly straight/
2. slightly sinuous/
3. markedly sinuous/
#29. Hindwing apical angle <between imaginary line tangent to costal border and imaginary line tangent to apical
border>/
1. very acute/
2. moderately acute/
3. slightly acute/
#30. <Hindwing> vein CuA1 <branches>/
1. branched/
2. not branched/
#31. <Hindwing> apex <length>/
1. abruptly interrupted/
2. slightly short/
3. slightly long/
#32. <Hindwing apex shape>/
1. rounded/
2. slightly pointed/
3. markedly pointed/
#33. Supranal plate <length>/
1. very short/
2. sligthly elongate, not overlaying ovipositor/
#34. <Supranal plate shape>/
1. subrounded, faintly pointed/
2. rounded/
#35. Ventral phallomere distal process <size>/
1. short/
2. slightly long/
3. very long/
#36. <Ventral phallomere distal process> with most marked sclerotization taking/
1. entire process/
2. at least half area of the process/
3. only process tip/
#37. <Ventral phallomere distal process> denticulation/
1. markedly present/
2. faintly present/
3. absent/
#38. <Ventral phallomere distal process> tip/
1. wide, rounded/
2. slender but rounded/
3. pointed/
#39. <Ventral phallomere distal process> basal swelling/
1. absent/
2. present/
#40. <Ventral phallomere> proximal process <sclerotization intensity>/
1. faintly sclerotized/
2. markedly sclerotized/
#41. <Ventral phallomere proximal process length>/
1. only slightly projected/
2. markedly projected/
#42. <Ventral phallomere proximal process projection>/
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1. short/
2. long/
#43. <Ventral phallomere proximal process overall curvature>/
1. somewhat straight/
2. curved/
#44. <Ventral phallomere proximal process overall direction>/
1. turned diagonally posterad lateral portion/
2. distinctly turned posterad/
3. turned diagonally posterad central portion/
#45. <Ventral phallomere proximal process> with basal portion/
1. slender/
2. slightly wide/
3. very wide/
#46. <Ventral phallomere proximal process> apex/
1. rounded/
2. with slightly long, slender point/
3. with short, robust point/
#47. <Ventral phallomere proximal process apex width>/
1. slightly slender than base/
2. distinctly slender than base/
#48. Slit between posterior margin of ventral phallomere and proximal process/
1. deep/
2. shallow/
#49. Left dorsal phallomere with anterior process <length>/
1. very long/
2. slightly long/
3. short/
#50. <Left dorsal phallomere anterior process width>/
1. markedly large/
2. moderately large/
3. very slender/
#51. <Left dorsal phallomere anterior process shape>/
1. markedly curved at about middle/
2. slightly curved at base/
3. straight/
#52. <Left dorsal phallomere anterior process> oriented/
1. to left/
2. anterad/
#53. <Left dorsal phallomere anterior process> apex <width>/
1. about as wide as middle/
2. distinctly wider than middle width/
#54. <Left dorsal phallomere> expanded portion between anterior process and phalloid apophysis <length>/
1. long/
2. short/
#55. <Left dorsal phallomere expanded portion between anterior process and phalloid apophysis> oriented/
1. anterad/
2. to left/
#56. <Left dorsal phallomere> well sclerotized area between phalloid apophysis and membranous lobe which is
<presence>/
1. indistinct/
2. evanescent/
3. marked/
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#57. <Left dorsal phallomere well sclerotized area between phalloid apophysis and membranous lobe size>/
1. large, slender/
2. large, wide/
3. small, not reaching membranous lobe/
4. small, reaching anterior area of membranous lobe/
#58. <Left dorsal phallomere> phalloid apophysis <length>/
1. very short/
2. regularly short/
3. regularly long/
4. very long/
#59. <Left dorsal phallomere phalloid apophysis width>/
1. moderately stout/
2. slender/
#60. <Left dorsal phallomere phalloid apophysis> its basal portion projected to/
1. back/
2. left one/
#61. <Left dorsal phallomere phalloid apophysis apical portion>/
1. somewhat contiguously curved with entire phalloid apophysis/
2. subapically slightly bent/
3. subapically markedly bent/
#62. <Left dorsal phallomere phalloid apophysis apical portion shape> apically/
1. concave/
2. almost straight/
#63. <Left dorsal phallomere phalloid apophysis apical portion sharpness>/
1. not sharp/
2. slightly sharp/
#64. <Left dorsal phallomere> membranous lobe <general width>/
1. wide/
2. slender/
#65. <Left dorsal phallomere> dorsal lamina posterior margin with left portion/
1. projected, expanded, uniformly curved/
2. projected, distinctly bent/
3. projected, narrow, uniformly curved/
4. not projected, uniformly curved/
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Appendix 4. Data matrix for the cladistic analysis. External morphology.
......continued on the next page
Specimen 1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 2 2 3 3 3 3 3 3 3 3 3 3 4 4 4 4 4 4 4 4 4 4 5
1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0
Eumusonia sp. 0 0 1 1 0 0 2 0 1 1 0 1 0 1 2 0 0 1 1 1 3 1 1 0 2 - 0 0 2 0 - - 0 1 1 - - 3 0 0 0 - 0 0 0 - - - 0 -
Chloromiopteryx sp. 1 1 1 3 0 0 1 0 1 1 1 1 0 1 0 0 0 1 1 0 0 0 1 0 1 0 0 0 1 0 1 0 2 0 1 - - 0 0 - - - 0 1 0 - - - 0 -
Anamiopteryx sp. 0 0 0 1 1 0 1 1 1 0 2 1 2 1 1 1 0 1 1 0 0 0 0 2 1 1 0 1 0 0 1 0 1 1 1 - - 1 1 2 1 0 0 1 1 0 1 0 0 -
M. fuscata Holotype 1 1 1 1 0 0 2 1 1 1 2 0 0 0 2 0 1 0 0 0 1 1 0 ? ? ? ? 1 0 1 1 0 2 0 0 1 2 0 1 1 0 - 1 2 1 0 1 0 0 -
M. nebulosa Holotype 0 1 1 1 1 0 2 1 1 0 0 1 0 1 2 0 1 0 0 0 2 1 0 1 1 1 1 1 0 1 1 1 2 0 0 1 2 1 1 0 0 - 0 2 1 1 1 1 1 2
M. rustica Syntype 0 1 1 1 1 0 2 1 1 0 0 1 0 1 ? ? 1 0 0 0 2 1 0 ? ? ? ? 1 0 1 1 1 2 0 0 1 2 0 ? 0 0 - 0 ? 1 1 1 1 ? ?
MT00013 1 1 1 1 0 0 2 1 ? 1 2 0 0 0 2 0 1 0 0 0 ? 1 0 2 1 1 1 1 0 1 1 0 2 0 0 1 2 1 1 0 1 0 1 2 1 0 1 0 0 -
MT00016 1 1 1 1 0 0 2 1 1 1 2 0 0 0 2 0 1 0 0 0 1 1 0 1 1 1 1 1 0 1 1 0 2 0 0 1 2 [01] 1 0 0 - 1 2 1 0 1 0 0 -
MT00021 1 1 1 1 0 0 2 1 1 1 2 0 0 0 2 0 1 ? 0 ? 1 1 0 1 2 1 1 1 0 1 1 0 2 0 0 1 2 1 1 1 0 - 1 2 1 0 1 0 0 -
MT00027 1 1 1 1 0 0 2 1 1 1 2 0 0 0 2 0 1 0 0 0 1 1 0 1 0 1 1 1 0 1 1 0 2 0 0 1 2 1 1 1 0 - 1 2 1 0 1 0 0 -
MT00029 1 1 1 1 0 0 2 1 1 1 2 0 0 0 2 0 1 0 0 0 1 1 0 1 0 1 1 1 0 1 1 0 2 0 0 1 2 1 1 0 1 0 1 2 1 0 1 0 0 -
MT00030 1 1 1 1 0 0 2 1 1 1 2 0 0 0 2 0 1 1 0 0 1 1 0 1 0 1 1 1 0 1 1 0 2 0 0 1 2 0 1 1 0 - 1 2 1 0 1 0 0 -
MT00031 1 1 1 1 0 0 2 1 1 1 2 0 0 0 2 0 1 0 0 0 1 1 0 1 0 1 1 1 0 1 1 0 2 0 0 1 2 1 1 1 0 - 1 2 1 0 1 0 0 -
MT00033 0 1 1 1 0 0 2 1 1 0 0 1 0 1 2 0 1 1 0 0 3 1 0 0 2 1 1 1 0 1 1 1 2 0 0 1 2 0 0 0 0 - 1 2 1 1 0 - 1 0
MT00034 1 1 1 1 0 0 2 1 1 1 2 0 0 0 2 0 1 0 0 0 1 1 0 1 0 1 1 1 0 1 1 0 2 0 0 1 2 1 1 0 0 - 1 2 1 0 1 0 0 -
MT00043 0 1 1 1 0 0 2 1 1 0 0 1 0 1 2 0 1 1 0 0 3 1 0 0 2 1 1 1 0 1 1 1 2 0 0 1 2 0 0 0 0 - 1 2 1 1 0 - 1 1
MT00045 1 1 0 1 0 1 2 1 0 1 2 1 1 1 0 1 1 0 0 1 0 1 0 2 2 1 1 0 0 0 0 0 2 0 0 0 1 [12] 1 1 0 - 0 2 1 0 1 0 0 -
MT00046 1 1 0 1 0 1 0 1 0 1 2 1 1 1 0 1 1 0 0 0 0 1 0 2 2 1 ? 0 0 0 0 0 2 0 0 0 0 1 1 2 1 0 2 2 1 0 1 0 0 -
MT00047 0 1 1 1 0 0 2 1 1 0 0 1 0 1 2 0 1 0 0 1 3 1 0 0 2 1 1 1 0 1 1 1 2 0 0 1 2 0 0 0 0 - 0 2 1 1 0 - 1 1
MT00049 1 1 1 1 0 0 2 1 1 1 2 0 0 0 2 0 1 0 0 1 1 1 0 1 1 1 1 1 0 1 1 0 2 0 0 1 2 1 1 1 0 - 1 2 1 0 1 0 0 -
MT00051 0 1 1 2 0 0 2 1 1 0 1 1 0 1 2 0 1 0 0 0 2 1 0 1 0 1 1 1 0 1 1 1 2 0 0 1 2 [01] 1 0 0 - 1 2 1 0 1 0 0 -
MT00052 1 1 1 1 0 0 2 1 1 1 2 0 0 0 2 0 1 0 0 0 1 1 0 1 1 1 1 1 0 1 1 0 2 0 0 1 2 1 1 1 1 0 1 2 1 0 1 0 0 -
MT00053 1 1 1 1 0 0 2 1 1 1 2 0 0 0 2 0 1 0 0 0 1 1 0 0 0 1 1 1 0 1 1 0 2 0 0 1 2 1 1 1 0 - 1 2 1 0 1 0 0 -
MT00058 1 1 1 1 0 0 2 1 1 1 2 0 0 0 2 0 1 0 0 0 1 1 0 1 0 1 1 1 ? ? ? ? 2 0 0 1 2 1 ? 1 1 0 1 2 1 0 1 0 0 -
MT00061 1 1 1 1 0 0 2 1 1 1 2 0 0 0 2 0 1 0 0 0 1 1 0 1 1 1 1 1 0 1 1 0 2 0 0 1 2 1 1 1 0 - 1 2 1 0 1 0 0 -
MT00063 0 1 1 0 2 0 2 1 1 0 2 1 0 1 1 1 1 0 0 0 2 1 0 0 1 1 1 1 0 1 1 1 2 0 0 1 2 0 1 0 1 1 0 2 1 0 1 1 0 -
MT00064 1 1 1 1 0 0 2 1 1 1 2 0 0 0 2 0 1 0 0 0 1 1 0 1 1 1 1 1 0 1 1 0 2 0 0 1 2 1 1 0 0 - 1 2 1 0 1 0 0 -
MT00066 1 1 1 1 0 0 2 1 1 1 2 0 0 0 2 0 1 0 0 0 1 1 0 1 1 1 1 1 0 1 1 0 2 0 0 1 2 1 ? 0 0 - 1 2 1 0 1 0 0 -
MT00069 1 1 1 1 0 0 2 1 1 1 2 0 0 0 2 0 1 0 0 0 1 1 0 1 0 1 1 1 0 1 1 0 2 0 0 1 2 1 1 0 0 - 1 2 1 0 1 0 0 -
MT00070 1 1 1 1 0 0 2 1 1 1 2 0 0 0 2 0 1 0 0 0 1 1 0 1 1 1 1 1 0 1 1 0 2 0 0 1 2 1 1 1 0 - 0 2 1 0 1 0 0 -
MT00073 1 1 1 1 0 0 2 1 1 1 2 0 0 0 2 0 1 0 0 0 1 1 0 1 1 1 1 1 0 ? ? ? 2 0 ? ? ? [01] 1 1 ? ? ? 2 1 ? ? ? 0 -
MT00074 1 1 1 1 0 0 2 1 1 1 2 0 0 0 2 0 1 0 0 0 1 1 0 1 0 1 1 1 0 1 1 0 2 0 0 1 2 [12] 1 1 1 0 0 2 1 0 1 0 0 -
MT00077 1 1 1 1 0 0 2 1 1 1 2 0 0 0 2 0 1 0 0 0 1 1 0 1 2 1 1 1 0 1 1 0 2 0 0 1 2 1 1 1 0 - 1 2 1 0 1 0 0 -
MT00084 1 1 1 1 0 0 2 1 1 1 2 0 0 0 2 0 1 1 0 0 1 1 0 1 0 1 1 1 0 1 1 0 2 0 0 1 2 1 1 0 1 0 1 2 1 0 1 0 0 -
MT00085 1 1 1 1 0 0 2 1 1 1 2 0 0 0 2 0 1 0 0 0 1 1 0 1 2 1 1 1 0 1 1 0 2 0 0 1 2 1 1 1 0 - 1 2 1 0 1 0 0 -
MT00087 0 1 1 2 0 0 2 1 1 0 1 1 0 1 2 0 1 0 0 0 2 1 0 1 0 1 1 1 0 1 1 1 2 0 0 1 2 [01] 1 0 0 - 1 2 1 0 1 0 0 -
MT00088 1 1 1 1 0 0 2 1 1 1 2 0 0 0 2 0 1 1 0 1 1 1 0 1 0 1 1 1 0 1 1 0 2 0 0 1 2 1 1 0 1 0 1 2 1 0 1 0 0 -
MT00090 1 1 0 1 0 1 1 1 0 1 2 1 1 1 0 1 1 0 0 0 0 1 0 2 2 1 1 0 0 0 0 0 2 0 0 0 0 [12] ? 1 0 - 2 2 1 0 1 0 0 -
MT00195 0 1 1 2 0 0 2 1 1 0 1 1 0 1 2 0 1 0 0 0 2 1 0 1 0 1 1 1 0 1 1 1 2 0 0 1 2 [12] 1 2 1 0 1 2 1 0 1 0 0 -
MZUEFS46390 1 1 1 0 0 0 2 1 1 1 2 0 0 0 2 0 1 0 0 0 1 1 0 0 1 1 0 1 0 1 1 1 2 0 0 1 2 1 0 0 1 2 0 1 1 0 1 1 0 -
PUCRS0776 0 1 1 1 0 0 2 1 1 1 1 1 0 1 0 0 1 0 0 0 0 1 ? ? ? ? 1 1 0 1 1 1 2 0 0 1 2 0 0 0 0 - 0 1 1 0 1 1 0 -
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Zootaxa 3797 (1) © 2014 Magnolia Press
Appendix 4. (Continued)
Specimen 5 5 5 5 5 5 5 5 5 6 6 6 6 6 6 6 6 6 6 7 7 7 7 7 7 7 7 7 7 8 8 8 8 8 8 8 8 8 8 9 9 9 9 9 9 9 9 9 9
1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9
Eumusonia sp. 1 0 0 0 0 0 2 1 0 1 0 0 0 0 0 1 0 - - - - - - - - - 0 - - - - - - 0 - - 0 - - - 0 0 0 0 1 0 1 - 0
Chloromiopteryx sp. 1 1 0 0 0 1 1 0 0 0 0 0 0 0 0 0 1 0 - - - - - - 1 0 ? ? ? ? ? ? ? 0 - - - - - - - 0 2 ? ? ? ? ? 0
Anamiopteryx sp. 0 1 0 0 0 1 0 2 0 0 1 0 0 1 1 1 0 - - - - - - - - - 1 0 0 0 0 1 0 1 0 - 2 1 1 - 3 0 2 1 0 0 0 0 0
M. fuscata Holotype 1 1 2 1 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?
M. nebulosa Holotype 1 1 1 1 0 1 0 2 1 - 1 1 0 1 2 2 1 1 0 1 - - 2 1 1 1 1 2 1 - 1 0 0 1 1 1 0 - - - 1 0 1 1 1 0 0 3 1
M. rustica Syntype 1 ? ? 1 ? ? 0 2 1 - 1 1 0 1 2 2 1 1 0 1 - - 2 1 1 1 1 2 1 - 1 0 0 1 1 1 0 - - - 1 0 1 1 1 0 0 3 1
MT00013 1 1 2 1 0 2 0 2 0 1 0 0 0 1 0 1 1 1 1 1 0 1 1 0 0 0 1 1 0 0 0 1 0 1 1 0 2 1 1 - 3 1 2 1 0 1 0 0 1
MT00016 1 1 2 1 0 2 0 2 0 1 0 0 0 1 0 1 1 1 1 1 0 0 1 0 0 0 1 1 0 0 0 1 ? 1 1 0 2 1 1 - 3 1 2 1 0 1 0 0 1
MT00021 1 1 2 1 0 2 0 2 0 1 0 0 0 1 0 1 1 1 1 1 0 0 1 0 0 0 1 1 0 0 0 1 1 1 1 0 2 1 1 - 3 1 2 1 0 1 0 0 1
MT00027 1 1 2 1 0 2 0 2 0 1 0 0 0 1 0 1 1 1 1 1 2 0 1 0 0 0 1 1 0 0 0 1 1 1 1 0 2 1 1 - 3 1 2 1 0 1 0 0 1
MT00029 1 1 2 1 0 2 0 2 0 1 1 0 0 1 0 1 1 1 1 1 0 1 1 0 0 0 1 1 0 0 0 1 0 1 1 0 2 1 1 - 3 1 2 1 0 1 0 0 1
MT00030 1 1 2 1 0 2 0 2 0 1 1 0 0 1 0 1 1 1 1 1 2 0 1 0 0 0 1 1 0 0 0 1 1 1 1 0 2 1 1 - 3 1 2 1 0 1 0 0 1
MT00031 1 1 2 1 0 2 0 2 0 1 0 0 0 1 0 1 1 1 1 1 1 1 1 0 0 0 1 1 0 0 0 1 1 1 1 0 2 1 1 - 3 1 2 1 0 1 0 0 1
MT00033 1 1 1 1 0 2 0 1 1 - 1 1 0 1 1 1 1 1 0 1 - - 2 0 1 1 1 2 1 - 1 0 0 1 1 0 1 - - - 2 0 2 1 1 0 0 3 1
MT00034 1 1 2 1 0 2 0 2 0 1 0 0 0 1 0 1 1 1 1 1 0 0 1 0 0 0 1 1 0 0 0 1 0 1 1 0 2 1 1 - 3 1 2 1 0 1 0 0 1
MT00043 1 1 1 1 0 2 0 1 1 - 1 1 0 1 1 1 1 1 0 1 - - 2 0 1 1 1 2 1 - 1 0 0 1 1 0 1 - - - 2 0 2 1 1 0 0 3 1
MT00045 0 0 3 2 0 2 0 0 0 0 0 0 1 1 0 2 1 0 - - - - - - 0 0 1 1 0 1 1 1 0 1 0 - 2 0 - 0 3 1 2 0 0 1 0 1 1
MT00046 0 0 3 2 0 2 0 0 0 0 0 0 1 1 0 2 1 0 - - - - - - 0 0 1 1 0 1 1 1 0 1 0 - 2 0 - 0 3 1 2 0 0 1 0 1 1
MT00047 1 1 1 1 0 2 0 1 1 - 1 1 0 1 1 1 1 1 0 1 - - 2 0 1 1 1 2 1 - 1 0 0 1 1 0 1 - - - 2 0 2 1 1 0 0 3 1
MT00049 1 1 2 1 0 2 0 2 0 1 1 0 0 1 0 1 1 1 1 1 0 0 1 0 0 0 1 1 0 0 0 1 1 1 1 0 2 1 1 - 3 1 2 1 0 1 0 0 1
MT00051 0 1 2 1 1 2 0 1 1 - 1 1 0 1 0 1 1 1 1 1 0 0 1 0 0 0 1 1 0 0 1 1 0 1 1 0 2 1 0 - 3 1 2 1 1 0 0 0 1
MT00052 1 1 2 1 0 2 0 2 0 1 0 0 0 1 0 1 1 1 1 1 0 0 1 0 0 0 1 1 0 0 0 1 1 1 1 0 2 1 1 - 3 1 2 1 0 1 0 0 1
MT00053 1 1 2 1 0 2 0 2 0 1 0 0 0 1 0 1 1 1 1 1 0 1 1 0 0 0 1 1 0 0 0 1 1 1 1 0 2 1 1 - 3 1 2 1 0 1 0 0 1
MT00058 1 1 2 1 0 2 0 2 0 1 0 0 0 1 0 1 1 1 1 1 2 0 1 0 0 0 1 1 0 0 0 1 0 1 1 0 2 1 1 - 3 1 2 1 0 1 0 0 1
MT00061 1 1 2 1 0 2 0 2 0 1 0 0 0 1 0 ? 1 1 1 ? ? ? ? ? 0 0 1 1 0 0 0 1 1 1 1 0 2 1 1 - 3 1 2 1 0 1 0 0 1
MT00063 1 1 1 1 0 2 0 0 0 0 0 0 0 1 0 2 1 1 1 0 - - 0 0 0 0 1 0 0 0 0 1 0 1 1 0 2 0 - 0 3 1 2 1 0 1 1 0 1
MT00064 1 1 2 1 0 2 0 2 0 1 0 0 0 1 0 1 1 1 1 1 0 1 1 0 0 0 1 1 0 0 0 1 1 1 1 0 2 1 1 - 3 1 2 1 0 1 0 0 1
MT00066 1 1 2 1 0 2 0 2 0 1 0 0 0 1 0 1 1 1 1 1 1 1 1 0 0 0 1 1 0 0 0 1 1 1 1 0 2 1 1 - 3 1 2 1 0 1 0 0 1
MT00069 1 1 2 1 0 2 0 2 0 1 1 0 0 1 0 1 1 1 1 1 2 1 1 0 0 0 1 1 0 0 0 1 0 1 1 0 2 1 1 - 3 1 2 1 0 1 0 0 1
MT00070 1 1 2 1 0 2 0 2 0 1 1 0 0 1 0 1 1 1 1 1 2 0 1 0 0 0 1 1 0 0 0 1 0 1 1 0 2 1 1 - 3 1 2 1 0 1 0 0 1
MT00073 - - 2 1 0 2 0 2 0 1 0 0 0 1 0 1 1 1 1 1 0 1 1 0 0 0 1 1 0 0 0 1 0 1 1 0 2 1 1 - 3 1 2 1 0 1 0 0 1
MT00074 1 1 2 1 0 2 0 2 0 1 0 0 0 1 0 1 1 1 1 1 2 0 1 0 0 0 1 1 0 0 0 1 0 1 1 0 2 1 1 - 3 1 2 1 0 1 0 0 1
MT00077 1 1 2 1 0 2 0 2 0 1 0 0 0 1 0 1 1 1 1 1 0 1 1 0 0 0 1 1 0 0 0 1 1 1 1 0 2 1 1 - 3 1 2 1 0 1 0 0 1
MT00084 1 1 2 1 0 2 0 2 0 1 0 0 0 1 0 1 1 1 1 1 0 0 1 0 0 0 1 1 0 0 0 1 1 1 1 0 2 1 1 - 3 1 2 1 0 1 0 0 1
MT00085 1 1 2 1 0 2 0 2 0 1 0 0 0 1 0 1 1 1 1 1 0 1 1 0 0 0 1 1 0 0 0 1 1 1 1 0 2 1 1 - 3 1 2 ? ? ? 0 0 1
MT00087 0 1 2 1 1 2 0 1 1 - 1 1 0 1 0 1 1 1 1 1 0 0 1 0 0 0 1 1 0 0 1 1 0 1 1 0 2 1 0 - 3 1 2 1 1 0 0 0 1
MT00088 1 1 2 1 0 2 0 2 0 1 0 0 0 1 0 1 1 1 1 1 0 0 1 0 0 0 ? ? ? ? ? ? ? 1 1 0 ? ? ? ? 3 1 ? 1 0 1 ? 0 1
MT00090 0 0 3 2 0 2 0 0 0 0 0 0 1 1 0 2 1 0 - - - - - - 0 0 1 1 0 1 1 1 0 1 0 - 2 0 - 0 3 1 2 0 0 1 0 1 1
MT00195 0 0 2 1 1 2 0 1 1 - 1 1 0 1 0 1 1 1 1 1 0 0 1 0 0 0 1 1 0 0 1 1 0 1 1 0 2 1 0 - 3 1 2 1 1 0 0 0 1
MZUEFS46390 1 1 1 1 0 1 0 2 0 1 1 0 0 1 0 1 1 1 0 1 - - 1 0 - 1 1 0 0 1 - 1 0 1 1 0 2 1 1 - 4 1 2 1 1 0 0 2 1
PUCRS0776 1 1 1 1 0 1 0 0 0 0 0 1 1 1 1 2 1 1 1 0 - - 0 0 0 0 1 0 0 0 0 1 1 1 1 0 2 0 - 1 3 1 2 0 0 0 1 0 1
