Corresponding author: Alípio Benedetti (
There are over 6,600 described species of
Carl-Friedrich Roewer (1881–1963) published several contributions on the taxonomy of
In the last 20 years, several Neotropical groups of
Twenty years later,
In another publication on Peruvian arachnids,
In his study of the early lineages of
A number of studies have been conducted using
Currently, and because of the historical background outlined above,
The main goal of this contribution is to review the
The material examined is housed in the following depositories (curators in parentheses):
The following abbreviations are used in the synonymic listing:
The following morphological abbreviations are used in the descriptions:
The following abbreviations are used in phylogenetic results and discussion:
The terminology for the armature of the dorsal scutum and legs follows
The drawings were sketched using a Leica stereomicroscope (model MZ APO, Heerbrugg, Switzerland) coupled with a camera lucida. The penises were prepared following
Our choice of outgroups was based on a phylogenetic hypothesis proposed by Pinto-da-Rocha et al. (in prep.) for the
The choice of ingroup for the analyses was based on the availability of biological material from sequences which could be obtained. This includes all species represented by specimens that had been maintained in 98% ethanol at –20ºC in the tissue collection of the Laboratório de Aracnologia (IBUSP).
We extracted DNA from the muscle tissues of the legs (preferably leg IV) (
Five molecular loci were amplified from the extracted DNA: the nuclear ribosomal 28S rRNA gene; the 12S and 16S mitochondrial ribosomal genes; the mitochondrial cytochrome c oxidase subunit I (COI) coding gene; and the nuclear gene encoding histone H3 (H3). We amplified the fragments of 12S, 16S, 28S and COI using the primers as in
List of primer sequences used for amplification and sequencing of H3 gene fragments.
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H3AF | 5’-ATGGCTCGTACCAAGCAGAC(ACG)GC-3’ |
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54 |
H3AR | 5’-ATATCCTT(AG)GGCAT(AG)AT(AG)GTGAC-3’ |
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54 |
H3AF_edit | 5’-GCVMGVAAGTCYACVGGMGG-3’ | This study | 54 |
H3AR_edit | 5’-ATGGTSACTCTCTTGGCGTGR-3’ | This study | 54 |
We assembled contiguous sequences using the package Consed/PhredPhrap (
Additionally, we used sequences of four markers (12S, 16S, 28S and COI) present in
We used Mesquite 2.5 computer software (
The analysis based on molecular and morphological data combined (total evidence) and molecular data alone was conducted under the optimality criteria of maximum parsimony and maximum likelihood (
The search strategy was conducted in two stages, following
During the second step, the topologies selected on
The optimization of the morphological characters present in the analysis of total evidence was visualized through the Winclada-ASADO program (
Node support was evaluated for maximum parsimony analyses using the Goodman-Bremer method (
Our matrix consisted of 33 taxa (eight outgroup taxa and 25 ingroup taxa). There are no missing data in the 12S, 16S, 28S and COI datasets (33 sequences). In contrast, there are missing data from nine exemplars in the H3 gene dataset (24 sequences; see Table
List of species with voucher and GenBank accession numbers for the amplified fragments.
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Our morphological matrix consisted of 68 characters (see “List of morphological characters” below and Table
Morphological data matrix used in the cladistic analysis of
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1000020?04 | 00-2-11113 | 1121?10101 | 0100010001 | 1104325010 | 001000---- | 11110110 |
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0000021?05 | 0112-11010 | 000?010101 | 0000000000 | 1000000000 | 000?-0---- | 11000001 |
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1111300202 | 1010-00000 | 111-200101 | 0100020011 | 1100000000 | 00111100-1 | ??000100 |
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0111300206 | 1010-11?1? | 1021?11000 | 0000000000 | 0000000000 | 001110---- | 01111100 |
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2111300206 | 1000-11?10 | 101-?10000 | 0000010000 | 1003000010 | 001100---- | 01?11100 |
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0000020004 | 0000-11010 | 111-110101 | 1000000000 | 1000000000 | 001000---- | 10111110 |
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2000110001 | 0101111113 | 0021-10100 | 0000110111 | 0000000000 | 001000---- | 20100000 |
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0000021011 | 0111111111 | 111–10101 | 0100120111 | 1023012000 | 101310---- | 12100001 |
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1000220100 | 0010-00000 | 000-000101 | 0100000000 | 1100000000 | 0011121010 | 21111100 |
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1000220100 | 0010-00000 | 000-000101 | 0000000000 | 1100110000 | 0010021000 | 21111100 |
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1000220101 | 0010-00000 | 111-100101 | 0100010001 | 1101221010 | 1010121001 | 21000100 |
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1000220100 | 0010-00000 | 000-000101 | 0100000000 | 1100110000 | 0012121000 | 10111100 |
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1000220011 | 0001111010 | 0021010101 | 0000110001 | 0002233020 | 0010021001 | 10?11110 |
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1000220011 | 0001111010 | 0021010101 | 0000110001 | 0002233020 | 0010021001 | 10?11110 |
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1000220101 | 0011100000 | 1220100111 | 0100010001 | 1101221010 | 0011121000 | 10010110 |
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1000220101 | 0011100000 | 111-100101 | 0100010001 | 1101221010 | 1010121001 | 21001100 |
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1000220101 | 0011100000 | 111-100101 | 0100010001 | 1101221010 | 1010121001 | 21001100 |
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0000220100 | 0010-00000 | 000-000101 | 0100000000 | 1100000000 | 0011121010 | 21?11100 |
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1000220101 | 0011110000 | 000-000000 | 0100010001 | 1102002010 | 00121211?0 | 10000100 |
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1000220100 | 0010-00000 | 000-000101 | 0100000000 | 1100110000 | 0010121010 | 21?11110 |
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1000121010 | 0000-00000 | 101-010001 | 0000000001 | 0100334021 | 0010121011 | ?1?11100 |
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1000121010 | 0000-11010 | 1020000000 | 0000000001 | 0102003021 | 0011021011 | 21?11100 |
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1000221010 | 0000-00000 | 000-000000 | 0000000000 | 0100300020 | 0012121010 | 11111100 |
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1000221010 | 0000-00000 | 0020000000 | 0000000000 | 0100300021 | 0010121001 | 21001100 |
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1000121013 | 0011011010 | 1021010000 | 0111000000 | 0100000000 | 0010120011 | 21111100 |
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1000221013 | 0001110012 | 0021010010 | 0010000001 | 0102000000 | 0110021010 | 21121100 |
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0000221012 | 0000-11011 | 1021011000 | 1100101101 | 1000000000 | 0010121011 | 1000?100 |
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1000121013 | 0000-11013 | 0021010000 | 0100100000 | 0010040200 | 0011121000 | 10011100 |
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1000121013 | 0000-11013 | 0021010000 | 0100100000 | 0010040200 | 0011121000 | 10011100 |
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0000121011 | 0000-11011 | 1020010000 | 1100001100 | 0100000000 | 0010020011 | 10011100 |
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0000121012 | 0000-11010 | 001-011000 | 1100100100 | 0000000000 | 0010121011 | 10001110 |
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1000221013 | 0000-11010 | 000-010000 | 0100000000 | 0000000000 | 0012121001 | 10011110 |
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1000221013 | 0001111010 | 1121010000 | 0000000000 | 0002000000 | 0112121000 | 10011110 |
The complete matrix for each of the analyses had the total characters as follows:
We chose
Phylogenetic hypotheses of
The monophyly of
We recognized six genera in
The ocularium armed with a pair of tubercles [15:0] and tarsal claws of legs III–IV pectinated [34:1] are exclusive autapomorphies of
The sister-group of
The monophyly of
The monophyly of
The
The monophyly of
Support for both inner clades is also low: “
The topology of the phylogenetic hypothesis found with the exclusive use of molecular data (
The monophyly of
1 | Leg IV elongate ( |
2 |
1’ | Leg IV short ( |
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2 | Alpha-type |
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2’ | Gamma-type (Fig. |
3 |
3 | Eye mound armed with a pair of tubercles (at least eye length or longer; Fig. |
4 |
3’ | Eye mound unarmed (Fig. |
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4 | Apex of coxa IV reaching area III (Fig. |
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4’ | Apex of coxa IV reaching area IV or posterior border of dorsal scutum (Fig. |
5 |
5 | Posterior margin and free tergites with one or a pair of tubercles larger than the others in the segment (Fig. |
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5’ | Posterior margin and free tergite I unarmed, free tergite II and III unarmed or armed with a pair of large tubercles (Fig. |
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1 | Eye mound unarmed (Fig. |
2 |
1’ | Eye mound with two conspicuous tubercles (Fig. |
7 |
2 | Femur–tibia IV with large tubercles (Figs |
3 |
2’ | Femur–tibia IV minute tuberculate (Figs |
4 |
3 | Areas I–IV with same-sized tubercles (Fig. |
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3’ | Areas I–IV with a median pair of tubercles slightly larger than others (Fig. |
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4 | Penis with 5–6 |
5 |
4’ | Penis with 7–9 |
6 |
5 | Chelicerae strongly inflated (Fig. |
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5’ | Chelicerae moderately inflated (Figs |
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6 | Areas I–IV densely and uniformly tuberculate (Fig. |
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6’ | Areas I–IV with central region less tuberculate than laterals (Fig. |
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7 | Coxa IV with large dorsoapical tubercle (e.g. Fig. |
8 |
7’ | Coxa IV without large dorsoapical tubercle (Fig. |
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8 | Area III with two long and acute tubercles (Fig. |
9 |
8’ | Area III with same-sized tubercles or a pair slightly larger on median region (Fig. |
10 |
9 | Areas I–II and IV with a pair of tubercles larger than others in same area (Fig. |
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9’ | Areas I–II and IV with same-size tubercles (Fig. |
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10 | Femur IV with rows of tubercles of varied morphology (apex can be blunt, acuminate or lanceolate; Figs |
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10’ | Femur IV with rows of acuminate tubercles (e.g. Figs |
11 |
11 | Femur IV with dorsal row of large tubercles (Fig. |
12 |
11’ | Femur IV without dorsal row of large tubercles (Fig. |
14 |
12 | Larger tubercles on dorsal femur IV concentrated on distal half (Fig. |
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12’ | Larger tubercles on dorsal femur IV concentrated on basal half (Figs |
13 |
13 | Eye mound with divergent large tubercles (Fig. |
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13’ | Eye mound with large parallel tubercles (Fig. |
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14 | Femur IV with two ventral rows of tubercles (Fig. |
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14’ | Femur IV with one ventral row of tubercles (Fig. |
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1 | Area III unarmed (Fig. |
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1’ | Area III armed with a pair of spines (e.g. Figs |
2 |
2 | White or silver patches on dorsal scutum | 3 |
2’ | Without patches on dorsal scutum | 4 |
3 | White patches on prosoma behind ocularium and on area I (Fig. |
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3’ | Silver patches on prosoma, area I and lateral margin; femur IV with ventral row of conspicuous tubercles; patella IV with two long and acute dorsoapical tubercles (Figs |
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4 | Dorsal scutum with sparse granules (Fig. |
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4’. | Dorsal scutum densely granulate (Fig. |
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1 | Free tergite III with conspicuous apophysis (Figs |
2 |
1’ | Free tergite III unarmed or with a pair of acute tubercles (Figs |
5 |
2 | Area I divided (Figs |
3 |
2’ | Area I undivided (Figs |
4 |
3 | Coxa IV with long retrolateral apophyses; free tergite with short and simple apophysis (Fig. |
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3’ | Coxa IV without retrolateral apophysis; free tergite with bifid apophysis (Fig. |
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4 | Dorsal scutum grooves virtually inconspicuous; free tergite III with short, wide and trifid apophysis (Fig. |
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4’ | Dorsal scutum grooves conspicuous; free tergite III with long and simple apophysis (Fig. |
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5 | Area III with one pair of spines (Figs |
6 |
5’ | Area III unarmed |
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6 | Trochanter IV with a retrolateral apophysis; Femur IV robust, with a retrolateral basal apophysis and two dorsal rows of tubercles (Figs |
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6’ | Trochanter IV without apophysis | 7 |
7 | Gamma-P type |
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7’ | Kappa type |
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1 | Free tergite III armed with a pair of tall tubercles (longer than its tergite length; Figs |
2 |
1’ | Free tergite III unarmed, tubercles absent or small (Figs |
3 |
2 | Ventral distal half of femur IV and patella IV with tubercles longer than segment width (Fig. |
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2’ | Leg IV with small tubercles (Figs |
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3 | Femur IV with tall tubercles (most similar-sized or longer than segment width); Figs |
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3’ | Femur IV with small tubercles (Figs |
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The family
Most species (28 spp.) are known only from their type-locality, and those known from a few records of distribution (6 spp.) are endemic to small areas, where the maximum distance between two records is 150km. A few localities possess sympatric species, such as: Parque Nacional Yanachaga-Chemillén/Peru (
All Bolivian species of
The monotypic genus
PERU.
Alpha-type
Habitus, dorsal of
Habitus, dorsal of males of
Habitus, dorsal of males of
Similar to
Considering that
(Fig.
Habitus, dorsal of
Similar to
Habitus, dorsal of males of
The specific epithet of masculine gender, in the genitive form, dedicated to the Brazilian filmmaker, actor and writer Glauber de Andrade Rocha (1939–1981).
(Fig.
Male trochanter–tibia IV of
Resembles
(Fig.
It differs from females of
This species is known only by the female holotype, which ends up being a taxonomic problem, since it is not uncommon for females of
(Fig.
Male trochanter–tibia IV of leg IV of
It differs from other species of the genus by the set of following characteristics: dorsal scutal area III with a pair of long spines (Fig.
Male trochanter–patella/tibia IV of
The specific epithet of masculine gender, in the genitive form, dedicated to the Italian writer and filmmaker Pier Paolo Pasolini (1922–1975).
(Fig.
Male trochanter–patella/tibia IV of
Similar to
The specific epithet, a noun in apposition, in reference to the type locality, Laguna Pomacocha (Junín, Peru), a beautiful pond surrounded by grass and large rocks that harbor this species.
(Fig.
Male trochanter–patella/tibia IV of
Similar to
The specific epithet, a noun in apposition, in reference to the type locality, Laguna Querococha, a blue waters lagoon, from glacier of Parque Nacional Huascaran, Department of Ancash, type locality of the species.
(Fig.
Penis of
It differs from other species of the genus with known females by having a high median spine in the free tergites I–II (Fig.
Considering that
Penis of
The specific epithet of masculine gender, in the genitive form, dedicated to the German arachnologist Carl Friedrich Roewer (1881–1963), the author of original name in secondary homonymy.
(Fig.
Penis of
Similar to
The specific epithet of feminine gender, in the genitive form, dedicated to Dr. Diana Silva D. (
(Fig.
It differs from other species of the genus by the combination of the following characteristics: Gamma-type
The specific epithet of masculine gender, in the genitive form, dedicated to the American filmmaker, producer and screenwriter Steven Allen Spielberg.
(Fig.
Penis of
Similar to
(Fig.
Penis of
Similar to
Roewer described
The specific epithet, a noun in apposition, in reference to the type locality, Tapacocha, Ancash, Peru, as well as in relation to the name of the genus where the species was originally described by
(Fig.
Penis of
Similar to
The male holotype and paratypes examined by Roewer (
(Fig.
Penis of
Similar to
Roewer described
The specific epithet, an adjective in nominative singular, formed by Latin prefix
(Fig.
It differs from other species of the genus (with males known) by the set of the following characteristics: ocularium with a pair of spines; scutal areas unarmed; free tergites I–III with a pair of tubercles (Fig.
Regarding the type of
(Fig.
Penis of
Similar to
The specific epithet of masculine gender, in the genitive form, dedicated to the Peruvian writer, politician, journalist, essayist, filmmaker, college professor and Nobel Prize winner Jorge Mario Pedro Vargas Llosa (born 1936), more commonly known as Mario Vargas Llosa.
(Fig.
Penis of
Similar to
(Fig.
Penis of
Kappa-type
The genus name, a noun in the nominative singular, from Quechua,
(Fig.
As for the genus.
The specific epithet of masculine gender, in the genitive form, dedicated to the American director, producer and screenwriter Stanley Kubrick (1928–1999).
(Fig.
Alpha-type
(Fig.
It differs from other species of the genus by silver-white patches on carapace, area I and lateral margins of dorsal scutum; femur IV with two rows of acuminate large tubercles, a retroventral one with 35–37 tubercles and a proventral one with 31–32 tubercles (Fig.
(Fig.
Live specimens of
It differs from other species of the genus by the set of following characters:
(Fig.
It differs from other species of the genus by the set of following characters:
(Fig.
It differs from other species of the genus by the set of following characters:
(Fig.
Live specimens of
It differs from other species of the genus by the set of following characters:
(Fig.
Kappa-type
The genus name, a noun in the nominative singular, is derived from Auguste Marie Louis Nicholas Lumière (1862–1954) and Louis Jean Lumière (1864–1948), the Lumière brothers, who were the inventors of cinematograph, being frequently referred like the parents of the “Cinema”. Gender feminine.
(Fig.
Live specimens of
Similar to
The specific epithet of masculine gender, in the genitive form, dedicated to the Italian filmmaker, editor, screenwriter, painter and writer Michelangelo Antonioni (1912–2007).
(Fig.
Live specimens of
Similar to
The specific epithet of masculine gender, in the genitive form, dedicated to the American filmmaker, actor, musician and writer Heywood Allen (born Allan Stewart Königsberg in 1935), known as Woody Allen.
(Fig.
Gamma-type
(Fig.
Live specimens of
Similar to
The specific epithet, a noun in apposition, in reference to Beni Department (“El Beni”), Bolivia, Department of the type locality of the species.
(Fig.
It differs from other species of the genus by having a large retrolateral apophysis on coxa IV, curved toward externally, and a short spiniform apophysis in free tergite III (Fig.
The specific epithet of masculine gender, in the genitive form, dedicated to the Swedish filmmaker, director, producer and writer Ernst Ingmar Bergman (1918–2007).
(Fig.
Very similar to females of
It is intriguing how Roewer described the same species from the Bolivian Chaco not only in different species and genera, but in different subfamilies (
BOLIVIA. Chaco.
Live specimens of
Differs from other species of the genus by having femur IV swollen and with a long retrobasal apophysis; trochanter IV with long retroapical spiniform apophysis (Fig.
See the remarks on section 3.39 for an observation on the recent taxonomic history of the species. Considering that the type material is formed from syntypes, we chose the
(Fig.
Distribution of
It differs from other species of the genus because males present an extremely long apophysis in free tergite III with forked apex (Fig.
The specific epithet of masculine gender, in the genitive form, dedicated to the Italian filmmaker and screenwriter Federico Fellini (1920–1993).
(Fig.
Similar to
The specific epithet of masculine gender, in the genitive form, dedicated to the Japanese filmmaker, producer and screenwriter Akira Kurosawa (黒澤明; 1910–1998).
(Fig.
It differs from other species of the genus because by having area I undivided; long spiniform apophysis on free tergite III (Fig.
The specific epithet of masculine gender, in the genitive form, dedicated Miguel Limachi, from Coleccíon Boliviana de Fauna (
(Fig.
It differs from other species of the genus by a trifurcated and acuminate apophysis in free tergite III (Fig.
The specific epithet, an adjective in nominative singular, formed by Latin prefix
(Fig.
Distribution of
It differs from other species of the genus by the following set of characteristics: alpha type
The specific epithet of feminine gender, in the genitive form, in honor to Maria René Vacaflores, from Coleccíon Boliviana de Fauna (
(Fig.
Kappa-type, with straight posterior margin of
(Fig.
Similar to
The specific epithet of masculine gender, in the genitive form, dedicated to the English actor, composer, director and producer Sir Charles Spencer Chaplin (1889–1977), a worldwide icon in the era of silent film through his screen persona “The Tramp”.
(Fig.
Distribution of
It differs from other species of the genus by the following set of characteristics: gamma-P-type
The specific epithet of masculine gender, in the genitive form, dedicated Dr. José Ochoa C. (
(Fig.
Distribution of
Similar to
The specific epithet of masculine gender, in the genitive form, dedicated to the American director, producer, screenwriter and cinema historian Martin Charles Scorsese (1942–), an exponent of New Hollywood (American New Wave).
(Fig.
It differs from other species of the genus by the following set of characteristics: kappa-type
(Fig.
We chose parsimony analysis to obtain a working phylogenetic hypothesis of
Besides using
The family
Our phylogenetic analyses based on molecular and morphological markers resulted in a new classification with several new combinations proposed for
Since the choice of ingroup taxa for the total evidence analysis was based on the availability (see section 2.5) of specimens, it was necessary to adopt a strategy that would combine the results of the phylogenetic hypothesis with the results of the classical taxonomy (or α-taxonomy) and achieve a new classification of the
Analysis of the type-material of all described species ensured that we would be able to reliably recognize the new species from Bolivia and Peru. Considering the large number of new species and the large number of monotypic genera of
Of the six genera of
Before this study,
Of the three new species not included in the phylogenetic analyses,
The internal relationships of
We were not able to test the monophyly of
Prior to this revision,
The internal relationships of
All unambiguous synapomorphies and those recovered only with ACCTRAN are listed in Section 3.3. The slightly flattened, sub cylindrical pedipalpus femur, [8:1] is found in all representatives of
It is important to note that of
Finally, it is worth mentioning that the “
We present a strongly supported, comprehensive phylogenetic hypothesis of
The following generic synonymies are proposed:
The following new genera are described from Peru:
The following specific synonymies are proposed:
The following new combinations are proposed:
The following new species are described from Bolivia:
The following secondary homonym species names are replaced:
A.R.B. and R.P.R. designed the study and contributed to collecting materials. A.R.B. conducted the taxonomic revision and phylogenetic analyses with assistance from R.P.R. A.R.B and R.P.R. discussed the results and drafted and approved the final version of the manuscript.
We thank the curators Adriano Kury (
Supplementary list
Figures S1–S5