Research Article |
Corresponding author: Alípio Rezende Benedetti ( alipiobenedetti@gmail.com ) Academic editor: Lorenzo Prendini
© 2022 Alípio Rezende Benedetti, Ricardo Pinto-da-Rocha.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Benedetti AR, Pinto-da-Rocha R (2022) Systematic revision and total evidence phylogenetic analysis of the Andean family Metasarcidae Kury, 1994 (Opiliones: Laniatores), with description of two new genera and twenty new species. Arthropod Systematics & Phylogeny 80: 309-388. https://doi.org/10.3897/asp.80.e73829
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Metasarcidae Kury, 1994 (Gonyleptoidea) is exclusively distributed in the Andean region of South America, from northern Peru to southern Bolivia. This contribution reviews the family using traditional taxonomy and cladistics. The cladistic analysis is based on total evidence (TE), maximum likelihood (ML) and direct optimization of molecular and morphological characters. The data matrix is composed of DNA sequences from three mitochondrial loci (12S rRNA, 16S rRNA and COI), from two nuclear loci (28S rRNA and H3) and 68 phenotypic characters. The dataset consists of 25 ingroup terminals (representing 20 species of the family) and eight outgroup species. The parsimony analysis resulted in one most parsimonious tree (L=4910) that recovered a monophyletic Metasarcidae, sister-group of Cosmetidae. As a result of the taxonomic revision and cladistic hypothesis, a new classification is proposed with six genera and 38 valid species of Metasarcidae, of which two genera and 20 species are described for the first time. Additionally, the following are established: six generic name synonymies; four species synonymies; seven new species combinations; and three secondary homonym species names replaced.
Direct Optimization, Gonyleptoidea, Neotropical fauna, parsimony, systematics, taxonomy
There are over 6,600 described species of Opiliones, This order is subdivided into five suborders: Tetrophthalmi
Carl-Friedrich Roewer (1881–1963) published several contributions on the taxonomy of Opiliones during the 1920s, the most important of which is “Die Weberknechte der Erde” in 1923. Roewer pioneered the use of standardized descriptions and illustrations and advanced the systematics of Opiliones. However, the so-called “Roewerian system” of classification attributed subjective weights to a limited set of characters. This created a rigid system that did not take into account new evidence (characters) (
In the last 20 years, several Neotropical groups of Opiliones have been subjected to systematic revisions that include analysis of type material, examination of large numbers of specimens, and cladistic analyses using characters from the external morphology and the male genitalia (
Twenty years later,
In another publication on Peruvian arachnids,
In his study of the early lineages of Gonyleptidae,
A number of studies have been conducted using Metasarcinae species as outgroups in analyses involving Gonyleptidae (e.g.
Currently, and because of the historical background outlined above, Metasarcidae is composed of nine genera and 22 species.
The main goal of this contribution is to review the Metasarcidae family based on combined analyses of morphology and molecular data.
The material examined is housed in the following depositories (curators in parentheses):
The following abbreviations are used in the synonymic listing: cat = catalogue; cit = citation; desc = description; diag = diagnosis; dist = distribution; key = taxonomic key; mat = character matrix; morp = morphology; rdesc = redescription; syst = systematic discussion.
The following morphological abbreviations are used in the descriptions: ChL = Chelicera length; CL = carapace maximum length; DS = dorsal scutum; DSL = dorsal scutum maximum length; DSS = dorsal scutum shape; DSW = dorsal scutum maximum width; FIVL = femur IV length; FT I–III = Free Tergites I–III; MS A–E = penis ventral plate pairs of macrosetae A–E; VP= ventral plate of the penis.
The following abbreviations are used in phylogenetic results and discussion: DO = Direct optimization; GB = Goodman-Bremer support; IP = Iterative pass; ML01 = Total evidence hypothesis under Maximum Likelihood; ML02 = Molecular only hypothesis under Maximum Likelihood; MP01 = Total evidence hypothesis under Maximum Parsimony; MP02 = Molecular only hypothesis under Maximum Parsimony.
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 Gonyleptoidea and the tree was rooted with Stygnus multispinosus (Piza, 1938) (Stygnidae). We complemented our taxon sampling by adding representatives of the families Cryptogeobiidae, Nomoclastidae, Gonyleptidae and Cosmetidae, to account for the morphological and systematic diversity of Metasarcidae and related clades. The specimens and their collection numbers, collecting data and depository institutions are in the Supplementary list.
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.
Primer Name | Primer sequence | Source | Annealing (°C) |
H3AF | 5’-ATGGCTCGTACCAAGCAGAC(ACG)GC-3’ |
|
54 |
H3AR | 5’-ATATCCTT(AG)GGCAT(AG)AT(AG)GTGAC-3’ |
|
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 (ML).
Parsimony analyses: The analyses of total evidence with molecular and phenotypic data (MP01) and with only molecular data (MP02) were implemented in the POY 5.1.1 program (
The search strategy was conducted in two stages, following
During the second step, the topologies selected on DO were re-diagnosed using the iterative pass algorithm (IP;
Maximum Likelihood Analysis: The sequences aligned in MAFFT were used in the maximum likelihood analysis. The analyses of total evidence with molecular and phenotypic data (ML01) and with only molecular data (ML02) were implemented in the parallel version of the IQ-TREE version 1.5.4 program (
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.
Family | Species | Voucher | 12s | 16s | 28s | COI | H3 |
Cosmetidae | Cynorta coxaepunctata |
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MG797727 | MG798345 | MG798032 | MG769081 | MG769400 |
Cosmetidae | Metalibitia paraguayensis |
|
KF726468 | KF726580 | KF726692 | KF726804 | — |
Cosmetidae | Taito insperatus |
|
MG797960 | MG798601 | MG798288 | MG769317 | MG769680 |
Cranaidae | Phareicranaus hermosa |
|
KF767675 | KF767679 | KF767683 | KF767687 | — |
Cryptogeobiidae | Pseudopachylus alticola |
|
KF726498 | KF726604 | KF726716 | KF726828 | MG769520 |
Nomoclastidae | Napostygnus bispinosus |
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KF726454 | KF726566 | KF726678 | KF726790 | — |
Gonyleptidae | Gonyleptes fragilis |
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MG797707 | MG798323 | MG798010 | MG769061 | MG769359 |
Stygnidae | Stygnus multispinosus |
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MG797776 | MG798407 | MG798092 | MG769131 | MG769502 |
Metasarcidae | Ayacucho glauberrochai sp. nov. |
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MG797827 | MG798460 | MG798146 | MG769179 | MG769560 |
Metasarcidae | Ayacucho pomacocha sp. nov. |
|
MG797828 | MG798461 | MG798147 | MG769180 | MG769561 |
Metasarcidae | Ayacucho querococha sp. nov. |
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MG797808 | MG798440 | MG798126 | MG769161 | MG769534 |
Metasarcidae | Ayacucho silvae sp. nov. |
|
MG797825 | MG798458 | MG798144 | MG769177 | MG769558 |
Metasarcidae | Ayacucho spielbergi sp. nov. |
|
MG799804 | MG798436 | MG798122 | MG769158 | — |
Metasarcidae | Ayacucho spielbergi sp. nov. |
|
MG799806 | MG798438 | MG798124 | MG769160 | MG769532 |
Metasarcidae | Ayacucho spiniger comb. nov. |
|
MG797826 | MG798459 | MG798145 | MG769178 | MG769559 |
Metasarcidae | Ayacucho tapacocha nom.nov. |
|
KF726439 | KF726551 | KF726663 | KF726775 | MG769427 |
Metasarcidae | Ayacucho tapacocha nom.nov. |
|
MG797811 | MG798443 | MG798129 | MG769164 | MG769537 |
Metasarcidae | Ayacucho titschacki |
|
MG797823 | MG798456 | MG798142 | MG769175 | MG769555 |
Metasarcidae | Ayacucho uniseriatus comb. nov. |
|
KF726497 | KF726609 | KF726721 | KF726833 | MG769528 |
Metasarcidae | Ayacucho vargasllosai sp. nov. |
|
MG799824 | MG798457 | MG798143 | MG769176 | MG769556 |
Metasarcidae | Incasarcus argenteus |
|
MG799801 | MG798433 | MG798119 | MG769155 | — |
Metasarcidae | Incasarcus dianae |
|
KF726444 | KF726556 | KF726668 | KF726780 | — |
Metasarcidae | Incasarcus ochoai |
|
MG797786 | MG798416 | MG798102 | MG769141 | MG769511 |
Metasarcidae | Incasarcus viracocha |
|
MG797787 | MG798417 | MG798103 | MG769142 | MG769512 |
Metasarcidae | Huancabamba kubricki gen. nov. et sp. nov. |
|
MG797802 | MG798434 | MG798120 | MG769156 | MG769529 |
Metasarcidae | Lumieria antonionii gen. nov. et sp. nov. |
|
MG797822 | MG798455 | MG798141 | MG769174 | MG769554 |
Metasarcidae | Metasarcus bergmani sp. nov. |
|
MG797783 | MG798413 | MG798099 | MG769138 | MG769507 |
Metasarcidae | Metasarcus clavifemur comb. nov. |
|
MG797781 | MG798411 | MG798097 | MG769136 | MG769506 |
Metasarcidae | Metasarcus clavifemur comb. nov. |
|
MG797782 | MG798412 | MG798098 | MG769137 | — |
Metasarcidae | Metasarcus fellinii sp. nov. |
|
KF726489 | KF726601 | KF726713 | KF726825 | MG769508 |
Metasarcidae | Metasarcus trispinosus sp. nov. |
|
MG797856 | MG798493 | MG798179 | MG769210 | — |
Metasarcidae | Metasarcus vacafloresae sp. nov. |
|
MG797780 | MG798410 | MG798096 | MG769135 | — |
Metasarcidae | Tschaidicancha chaplini sp. nov. |
|
MG797819 | MG798452 | MG798138 | MG769171 | MG769553 |
Our morphological matrix consisted of 68 characters (see “List of morphological characters” below and Table
Morphological data matrix used in the cladistic analysis of Metasarcidae.
Taxa | Characters | ||||||
0000000001 | 1111111112 | 2222222223 | 3333333334 | 4444444445 | 5555555556 | 66666666 | |
1234567890 | 1234567890 | 1234567890 | 1234567890 | 1234567890 | 1234567890 | 12345678 | |
Stygnus multispinosus | 1000020?04 | 00-2-11113 | 1121?10101 | 0100010001 | 1104325010 | 001000---- | 11110110 |
Pseudopachylus alticola | 0000021?05 | 0112-11010 | 000?010101 | 0000000000 | 1000000000 | 000?-0---- | 11000001 |
Metalibitia paraguayensis | 1111300202 | 1010-00000 | 111-200101 | 0100020011 | 1100000000 | 00111100-1 | ??000100 |
Cynorta coxaepunctata | 0111300206 | 1010-11?1? | 1021?11000 | 0000000000 | 0000000000 | 001110---- | 01111100 |
Taito insperatus | 2111300206 | 1000-11?10 | 101-?10000 | 0000010000 | 1003000010 | 001100---- | 01?11100 |
Napostygnus bispinosus | 0000020004 | 0000-11010 | 111-110101 | 1000000000 | 1000000000 | 001000---- | 10111110 |
Phareicranaus hermosa | 2000110001 | 0101111113 | 0021-10100 | 0000110111 | 0000000000 | 001000---- | 20100000 |
Gonyleptes fragilis | 0000021011 | 0111111111 | 111–10101 | 0100120111 | 1023012000 | 101310---- | 12100001 |
Ayacucho glauberrochai sp. nov. | 1000220100 | 0010-00000 | 000-000101 | 0100000000 | 1100000000 | 0011121010 | 21111100 |
Ayacucho pomacocha sp. nov. | 1000220100 | 0010-00000 | 000-000101 | 0000000000 | 1100110000 | 0010021000 | 21111100 |
Ayacucho querococha sp. nov. | 1000220101 | 0010-00000 | 111-100101 | 0100010001 | 1101221010 | 1010121001 | 21000100 |
Ayacucho silvae sp. nov. | 1000220100 | 0010-00000 | 000-000101 | 0100000000 | 1100110000 | 0012121000 | 10111100 |
Ayacucho spielbergi sp. nov. | 1000220011 | 0001111010 | 0021010101 | 0000110001 | 0002233020 | 0010021001 | 10?11110 |
Ayacucho spielbergi sp. nov. | 1000220011 | 0001111010 | 0021010101 | 0000110001 | 0002233020 | 0010021001 | 10?11110 |
Ayacucho spiniger comb. nov. | 1000220101 | 0011100000 | 1220100111 | 0100010001 | 1101221010 | 0011121000 | 10010110 |
Ayacucho tapacocha nom.nov. | 1000220101 | 0011100000 | 111-100101 | 0100010001 | 1101221010 | 1010121001 | 21001100 |
Ayacucho tapacocha nom.nov. | 1000220101 | 0011100000 | 111-100101 | 0100010001 | 1101221010 | 1010121001 | 21001100 |
Ayacucho titschacki | 0000220100 | 0010-00000 | 000-000101 | 0100000000 | 1100000000 | 0011121010 | 21?11100 |
Ayacucho uniseriatus comb. nov. | 1000220101 | 0011110000 | 000-000000 | 0100010001 | 1102002010 | 00121211?0 | 10000100 |
Ayacucho vargasllosai sp. nov. | 1000220100 | 0010-00000 | 000-000101 | 0100000000 | 1100110000 | 0010121010 | 21?11110 |
Incasarcus argenteus | 1000121010 | 0000-00000 | 101-010001 | 0000000001 | 0100334021 | 0010121011 | ?1?11100 |
Incasarcus dianae | 1000121010 | 0000-11010 | 1020000000 | 0000000001 | 0102003021 | 0011021011 | 21?11100 |
Incasarcus ochoai | 1000221010 | 0000-00000 | 000-000000 | 0000000000 | 0100300020 | 0012121010 | 11111100 |
Incasarcus viracocha | 1000221010 | 0000-00000 | 0020000000 | 0000000000 | 0100300021 | 0010121001 | 21001100 |
Huancabamba kubricki gen. et.sp. nov. | 1000121013 | 0011011010 | 1021010000 | 0111000000 | 0100000000 | 0010120011 | 21111100 |
Lumieria antonionii gen. et. sp. nov. | 1000221013 | 0001110012 | 0021010010 | 0010000001 | 0102000000 | 0110021010 | 21121100 |
Metasarcus bergmani sp. nov. | 0000221012 | 0000-11011 | 1021011000 | 1100101101 | 1000000000 | 0010121011 | 1000?100 |
Metasarcus clavifemur comb. nov. | 1000121013 | 0000-11013 | 0021010000 | 0100100000 | 0010040200 | 0011121000 | 10011100 |
Metasarcus clavifemur comb. nov. | 1000121013 | 0000-11013 | 0021010000 | 0100100000 | 0010040200 | 0011121000 | 10011100 |
Metasarcus fellinii sp. nov. | 0000121011 | 0000-11011 | 1020010000 | 1100001100 | 0100000000 | 0010020011 | 10011100 |
Metasarcus trispinosus sp. nov. | 0000121012 | 0000-11010 | 001-011000 | 1100100100 | 0000000000 | 0010121011 | 10001110 |
Metasarcus vacafloresae sp. nov. | 1000221013 | 0000-11010 | 000-010000 | 0100000000 | 0000000000 | 0012121001 | 10011110 |
Tschaidicancha chaplini sp. nov. | 1000221013 | 0001111010 | 1121010000 | 0000000000 | 0002000000 | 0112121000 | 10011110 |
The complete matrix for each of the analyses had the total characters as follows: MP01 = 2,981; MP02 = 2,921; ML01 = 2,756; ML02 = 2,688.
1) Chelicerae. Segment II. Shape: (0) monomorphic (Figs
2) Pedipalpus. Trochanter. Shape: (0) distally enlarged, with a short basal neck; (1) distally enlarged, with a long basal neck (longer than wide).
3) Pedipalpus. Tibia. Dorsal-ventral shape: (0) not flat; (1) flat, spoon-shaped.
4) Pedipalpus. Tarsus. Setae length: (0) long, greater than or equal to half length of the tarsus; (1) short, less than half the length of the tarsus.
5) Pedipalpus. Femur. Ventral armature: (0) smooth or with sparse granules; (1) with large spines; (2) with conical short tubercles; (3) with short tubercles, laterally depressed
6) Pedipalpus. Femur. Dorsal armature: (0) with blunt tubercles; (1) with pointed, enlarged tubercles; (2) unarmed.
7) Pedipalpus. Femur. Proapical spine in males: (0) absent; (1) present.
8) Pedipalpus Femur. Shape. (0) subcylindrical, not flattened; (1) subcylindrical, slightly flattened; (2) dorsally curved, ventrally straight, considerably flattened laterally.
9) Pedipalpus. Patella. Proapical spine: (0) absent; (1) present.
10) Dorsal scutum. Shape: (0) type alpha (Fig.
11) Dorsal scutum. Areas: (0) Delimited areas (Fig.
12) Dorsal scutum. Number of ozopore openings: (0) One opening; (1) Two openings.
13) Dorsal scutum. Ocularium shape: (0) saddle-shaped (Fig.
14) Dorsal scutum. Ocularium. Armature: (0) unarmed (Fig.
15) Dorsal scutum. Ocularium. Type of paired armature: (0) tubercles (Fig.
16) Dorsal scutum. Carapace. Granulation: (0) densely granulate (Fig.
17) Dorsal scutum. Lateral margin. Granulation: (0) densely granulate (Fig.
18) Dorsal scutum. Scutal areas III–IV: (0) as divided scutal areas (Fig.
19) Dorsal scutum. Scutal areas granulation: (0) densely granulate (Fig.
20) Dorsal scutum. Scutal area I. State of fusion: (0) undivided (Fig.
21) Dorsal scutum. Scutal area I. A pair of median tubercles: (0) absent (Fig.
22) Dorsal scutum. Scutal area II. Armature: (0) unarmed (Fig.
23) Dorsal scutum. Scutal area III. Armature: (0) unarmed (Fig.
24) Dorsal scutum. Scutal area III. Armature. Spine insertion: (0) direct on integument (Fig.
25) Dorsal scutum. Scutal area IV. Armature: (0) unarmed (Fig.
26) Dorsal scutum. Posterior margin. Granulation: (0) densely granulate (Fig.
27) Dorsal scutum. Posterior margin. Shape: (0) straight (Fig.
28) Dorsal scutum. Posterior margin. A row of tubercles: (0) absent (Fig.
29) Dorsal scutum. Posterior margin. A pair of spines: (0) absent (Fig.
30) Free tergite I–IV. A row of tubercles: (0) absent (Fig.
31) Free tergite III. Apophysis: (0) absent (Fig.
32) Leg III. Coxa III. Number of apophysis: (0) one (Fig.
33) Leg II. Distitarsus: (0) 3-segmented; (1) 4/5-segmented.
34) Legs III–IV. Tarsal claws: (0) smooth; (1) pectinate.
35) Leg IV. Coxa IV. Coxae dorsally reaching the posterior margin of DS: (0) Not reaching (Fig.
36) Leg IV. Coxa IV. Prolateral apical armature: (0) unarmed (Fig.
37) Leg IV. Coxa IV. Retrolateral apical armature: (0) unarmed (Fig.
38) Leg IV. Coxa IV. Apical width of coxa IV: (0) as wide as coxa III apex (Fig.
39) Leg IV. Trochanter IV. Prolateral armature: (0) absent (Figs
40) Leg IV. Trochanter IV. Retrolateral armature: (0) absent (Figs J–L); (1) with a small distal tubercle (Figs
41) Leg IV. Femur IV. Length of femur IV: (0) long (FIV/DSL > 1,5; Fig.
42) Leg IV. Femur IV. Granulation: (0) smooth or slightly granulate (Figs
43) Leg IV. Femur IV. Retrodorsal row of tubercles: (0) absent or with small granules (Figs
44) Leg IV. Femur IV. Retrolateral row of tubercles: (0) absent or with small granules (Figs
45) Leg IV. Femur IV. Retroventral armature: (0) absent or with small granules (Figs
46) Leg IV. Femur IV. Proventral armature: (0) absent or with small granules (Figs
47) Leg IV. Femur IV. Prolateral row of tubercles: (0) absent or with small granules (Figs
48) Leg IV. Femur IV. Prodorsal row of tubercles: (0) absent or with small granules (Figs
49) Leg IV. Patella. Apical armature: (0) absent (Figs
50) Leg IV. Patella. Ventral armature: (0) absent (Fig.
51) Leg IV. Tibia. Proventral armature: (0) absent (Figs
52) Leg IV. Tibia. Retrolateral armature: (0) absent (Figs
53) Penis. Shape: (0) malleus plus Lamina Parva; (1) with VP well defined (Figs
54) Penis. Ventral plate. Distal margin. Shape: (0) straight (Figs
55) Penis. Apex of truncus invading VP: (0) absent (Fig.
56) Penis. Ventral plate. Lateral sacs: (0) absent; (1) present, bump-like; (2) present, finger-like;
57) Penis. Ventral plate. Lateral sacs. Length: (0) very short (base width equal to the length of the lateral sac; Figs
58) Penis. Ventral plate. Lateral sacs. Microsetae: (0) present (Figs
59) Penis. Ventral plate. Lateral sacs. Microsetae shape: (0) long (Figs
60) Penis. Ventral plate. Lateral sacs. Apex: (0) acuminate (Figs
61) Penis. Ventral plate. MS C: (0) two pairs; (1) three pairs (Figs 19DD–F); (2) four or more pairs (Figs
62) Penis. Ventral plate. MS C. Shape: (0) straight (Figs
63) Penis. Ventral plate. Microsetae on ventral face: (0) absent (Fig.
64) Penis. Stylus. Shape: (0) cylindrical (Figs
65) Penis. Stylus. Caruncle on apex: (0) absent (Figs
66) Penis. Insertion of the pedestal in the glans: (0) ventral; (1) medial.
67) Penis. Dorsal process: (0) absent (Figs
68) Penis. Ventral process: (0) absent (Figs
We chose MP01 analysis to obtain a working phylogenetic hypothesis of Metasarcidae (see Discussion section). Therefore, the results of this hypothesis will be presented in more detail (the main differences between the topologies will be indicated). Iterative pass optimization resulted in a single topology with 4,626 steps (Fig.
Phylogenetic hypotheses of Metasarcidae species. Total evidence hypothesis under Maximum Parsimony (IP) of Metasarcidae species, based on five molecular markers (12S, 16S, 28S, COI and H3) and 68 morphological characters. Goodman-Bremer support is given near each node. Asterisks represent clades that were not found in the Maximum Likelihood analysis.
The monophyly of Metasarcidae (sensu
We recognized six genera in Metasarcidae, of which four had been previously described (Ayacucho, Incasarcus, Metasarcus and Tschaidicancha) and two new genera (Huancabamba gen. nov. and Lumieria gen. nov.). Three genera are represented only by one terminal taxon in the present analysis, one of which is monotypic (Huancabamba gen. nov.). The other genera are Lumieria gen. nov. (two species recognized in this study) and Tschaidicancha (four species recognized in this study). Huancabamba kubricki gen. et sp. nov. is recovered as the sister-group of the Lumieria antonionii gen. et sp. nov. (GB=12). There is an exclusive unambiguous morphological synapomorphy: distitarsus II 4/5-segmented [33:1]; and a homoplastic unambiguous synapomorphy: four or more pairs of MS C [61:2], also present in several other Metasarcidae species. This clade is not recovered in ML02, each of the genera as independent basal lineages of Metasarcidae.
The ocularium armed with a pair of tubercles [15:0] and tarsal claws of legs III–IV pectinated [34:1] are exclusive autapomorphies of H. kubricki gen. et sp. nov. Two exclusive unambiguous autapomorphies were recovered for Lumieria antonionii gen. et sp. nov.: DS scutal area I divided by scutal groove II [20:2]; and penial stylus broad and sturdy, laterally flattened [64:2]. L. antonionii gen. et sp. nov. is also supported by a homoplastic unambiguous autapomorphy: presence of a retrolateral row of acuminate tubercles on tibia IV [65:1], also present in Tschaidicancha chaplini sp. nov.
The sister-group of Huancabamba gen. nov. + Lumieria gen. nov. is the clade Incasarcus + Metasarcus + Tschaidicancha + Ayacucho (GB=15), supported by one unambiguous yet homoplastic morphological synapomorphy: VP of penis without microsetae on ventral face [63:0]. Additionally, under ACCTRAN, MS C straight [62:0]. This group contains two well-supported clades: Incasarcus + Metasarcus (GB=22) and Tschaidicancha + Ayacucho (GB=31). The clade Incasarcus + Metasarcus is supported by one homoplastic morphological synapomorphy under ACCTRAN: ocularium unarmed [14:0]. The sister-group relationship between Incasarcus and Metasarcus corroborates the hypothesis of
The monophyly of Incasarcus (represented by four of the five described species) was recovered (GB=55), supported by four unambiguous, yet homoplastic morphological synapomorphies: DSS alpha-type [10:0]; spine insertion direct on integument on scutal area III [24:0]; posterior margin of DS with densely granulation [26:0]; patella IV apically with large tubercles [49:2]. Additionally, there is an exclusive synapomorphy under ACCTRAN: patella IV with two ventral rows of tubercles [50:1], absent in I. ochoai; and two more homoplastic synapomorphies [61:2; 62:1].
The monophyly of Metasarcus (sensu hoc) was recovered (GB=38), supported by one homoplastic unambiguous morphological synapomorphy: coxa III with two apophyses [32:1]. Also supporting this clade, there is one additional homoplastic morphological synapomorphy under ACCTRAN: femur IV densely granulated [42:1]. Internally, there are two clades, one grouping two species from the Bolivian department of La Paz (Metasarcus clavifemur and Metasarcus vacafloresae sp. nov.; GB=13) and the other with species from the Bolivian departments of Cochabamba (Metasarcus trispinosus sp. nov. and Metasarcus bergmani sp. nov.) and Tarija (Metasarcus fellinii sp. nov.; GB=10).
The Tschaidicancha + Ayacucho clade is supported by two homoplastic unambiguous synapomorphies: long microsetae on penial lateral sacs [59:0] and lateral sacs with acuminate apex [60:0]. The internal relationships of this clade are the most inconstant when comparing all the hypotheses of the different analyses. When morphological data are used, Tschaidicancha chaplini sp. nov. is recovered at the base of the clade, as a sister-group of Ayacucho (ML01; MP01). In the analyses built only with molecular data (ML02; MP02), the monophyly of Ayacucho is not obtained, because T. chaplini sp. nov. is recovered nested in the genus (see Discussion section). Tschaidicancha was previously monotypic. Tschaidicancha chaplini sp. nov. is supported by five homoplastic unambiguous autapomorphies [22:1; 44:2; 52:1; 54:2; 67:1] (see Discussion section).
The monophyly of Ayacucho (sensu hoc) was recovered in the MP01 analysis with low support (GB=7). The clade has an unambiguous exclusive synapomorphy: femur of pedipalpus sub cylindrical, slightly flattened [8:1] and 10 homoplastic unambiguous morphological synapomorphies: femur and patella of pedipalpus in males without proapical spine [7:0; 9:0]; DS with carapace, lateral margins, scutal areas and posterior margin densely granulated [16:0; 17:0; 19:0; 26:0]; posterior margin of DS with a row of tubercles [28:1]; free tergites I–III with a row of tubercles [30:1]; coxa III with two apophyses [32:1]; and femur IV short (FIV/DSL < 1,2) [41:1]. Additionally, the genus is supported by one exclusive synapomorphy under ACCTRAN; femur IV with a distal retroventral row of acuminate tubercles [45:1]. Furthermore, under ACCTRAN, three additional homoplastic morphological synapomorphies support the clade [10:0; 13:1; 46:1; see discussion section].
Ayacucho, as recovered by the MP01 analysis, is split into two clades: 1) A. silvae sp. nov. + A. titschacki + A. vargasllosai sp. nov. + A. pomacocha sp. nov. + A. glauberrochai sp. nov. (henceforth “silvae clade”); 2) A. spiniger comb. nov. + A. uniseriatus comb. nov. + A. spielbergi sp. nov. + A. querococha sp. nov. + A. tapacocha nom. nov. (henceforth “spiniger clade”).
Support for both inner clades is also low: “silva clade” (GB=9); “spiniger clade” (GB=5). The “silvae clade” is supported by three homoplastic unambiguous synapomorphies: DS area I [21:0] and III [23:0] unarmed; VP with microseta on ventral face [63:1]. Under ACCTRAN, ocularium without armature [14:0] is a homoplastic synapomorphy for the clade. The “spiniger clade” is supported by two exclusive unambiguous morphological synapomorphies: femur IV with a retrolateral [44:1] and a prolateral [47:1] rows of laminate tubercles, both absent on A. uniseriatus comb. nov. and A. spielbergi sp. nov.; and three homoplastic synapomorphies: coxa IV with a proapical short apophysis [36:1]; trochanter IV with a small distal tubercle [40:1]; and patella IV with small apical tubercles [49:1]. Additionally, under ACCTRAN, the clade is supported by one exclusive synapomorphy, femur IV with a distal retroventral row of acuminate tubercles [45:2]; and four homoplastic characters [10:1; 25:1; 46:2; 65:0].
The topology of the phylogenetic hypothesis found with the exclusive use of molecular data (MP02) is identical to that of MP01, with exception of the relationships within clade Tschaidicancha + Ayacucho (Supplementary Fig.
ML01: The best replacement model performances for the six partitions (five molecular markers + morphology) were evaluated, based on the AICc – corrected Akaike information criterion (
ML02: The best replacement model performances for the five partitions (five molecular markers) were evaluated based on the AICc. The data matrix consisted of 2,688 characters, of which IQ-TREE recognized 1,067 character state distribution patterns. AICc favored the choice of the following replacement models: TIM3+F+G4 for non-coding 12S; GTR+F+I+G4 for non-coding 16S; TIM+F+R4 for non-coding 28S; TIM2+F+I+G4 for coding COI; and TIM+F+I+G4 for coding H3. The analysis resulted in a topology with lnL = –20724.827.
The monophyly of Metasarcidae was recovered in both analyses. The topology in ML01 (Supplementary fig. 4) is very similar to the one recovered by MP01, with the exception that the “spiniger clade” is paraphyletic to the “silvae clade”. A. spielbergi sp. nov. is recovered as the sister group of A. uniseriatus comb. nov., which is the sister-group of the other Ayacucho. The ML02 hypothesis (Supplementary fig. 5) recovered T. chaplini sp. nov. nested within Ayacucho, as a sister-group of “silvae clade”, as in MP02. The “spiniger clade” is paraphyletic to the clade T. chaplini sp. nov. + “silvae clade”. Additionally, H. kubricki gen. et sp. nov. is a sister-group to the other Metasarcidae, with Lumieria antonionii gen. et sp. nov. as the most basal lineage of this clade. This result is different from the other phylogenetic hypotheses (MP01, MP02, ML01), in which Huancabamba gen. nov. and Lumieria gen. nov. are sister-groups.
1 | Leg IV elongate (FIVL/DSL > 1.6; Figs |
2 |
1’ | Leg IV short (FIVL/DSL < 1.5; Figs |
Ayacucho |
2 | Alpha-type DSS (Fig. |
Incasarcus |
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. |
Metasarcus |
4 | Apex of coxa IV reaching area III (Fig. |
Huancabamba gen. nov. |
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. |
Lumieria gen. nov. |
5’ | Posterior margin and free tergite I unarmed, free tergite II and III unarmed or armed with a pair of large tubercles (Fig. |
Tschaidicancha |
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. |
A. silvae sp. nov. |
3’ | Areas I–IV with a median pair of tubercles slightly larger than others (Fig. |
A. querococha sp. nov. |
4 | Penis with 5–6 MS C (Figs |
5 |
4’ | Penis with 7–9 MS C (Figs |
6 |
5 | Chelicerae strongly inflated (Fig. |
A. titschacki |
5’ | Chelicerae moderately inflated (Figs |
A. glauberrochai sp. nov. |
6 | Areas I–IV densely and uniformly tuberculate (Fig. |
A. pomacocha sp. nov. |
6’ | Areas I–IV with central region less tuberculate than laterals (Fig. |
A. vargasllosai sp. nov. |
7 | Coxa IV with large dorsoapical tubercle (e.g. Fig. |
8 |
7’ | Coxa IV without large dorsoapical tubercle (Fig. |
A. pasolinii sp. nov. |
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. |
A. spiniger comb. nov. |
9’ | Areas I–II and IV with same-size tubercles (Fig. |
A. spielbergi sp. nov. |
10 | Femur IV with rows of tubercles of varied morphology (apex can be blunt, acuminate or lanceolate; Figs |
A. tapacocha nom. nov. |
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. |
A. weyrauchi comb. nov. |
12’ | Larger tubercles on dorsal femur IV concentrated on basal half (Figs |
13 |
13 | Eye mound with divergent large tubercles (Fig. |
A. triarmatus nom. nov. |
13’ | Eye mound with large parallel tubercles (Fig. |
A. bambamarca comb. nov. |
14 | Femur IV with two ventral rows of tubercles (Fig. |
A. inermis comb. nov. |
14’ | Femur IV with one ventral row of tubercles (Fig. |
A. uniseriatus comb. nov. |
1 | Area III unarmed (Fig. |
I. ochoai |
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. |
I. pictus |
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 |
I. argenteus |
4 | Dorsal scutum with sparse granules (Fig. |
I. dianae |
4’. | Dorsal scutum densely granulate (Fig. |
I. viracocha |
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. |
M. bergmani sp. nov. |
3’ | Coxa IV without retrolateral apophysis; free tergite with bifid apophysis (Fig. |
M. fellinii sp. nov. |
4 | Dorsal scutum grooves virtually inconspicuous; free tergite III with short, wide and trifid apophysis (Fig. |
M. trispinosus sp. nov. |
4’ | Dorsal scutum grooves conspicuous; free tergite III with long and simple apophysis (Fig. |
M. limachii sp. nov. |
5 | Area III with one pair of spines (Figs |
6 |
5’ | Area III unarmed | M. vacafloresae sp. nov. |
6 | Trochanter IV with a retrolateral apophysis; Femur IV robust, with a retrolateral basal apophysis and two dorsal rows of tubercles (Figs |
M. clavifemur |
6’ | Trochanter IV without apophysis | 7 |
7 | Gamma-P type DSS; Coxa III without apophysis (Fig. |
M. beni sp. nov. |
7’ | Kappa type DSS; Coxa III with two dorsal apophyses (Fig. |
M. kurosawai sp. nov. |
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. |
T. weyrauchi |
2’ | Leg IV with small tubercles (Figs |
T. chaplini sp. nov. |
3 | Femur IV with tall tubercles (most similar-sized or longer than segment width); Figs |
T. joseochoai sp. nov. |
3’ | Femur IV with small tubercles (Figs |
T. scorsesei sp. nov. |
Phalangodidae
Tricommatinae
[part]:
Prostygninae
[part.]:
Mitobatinae
[part.]:
Metasarcinae
Kury, 1994: 349 (desc);
Metasarcidae:
Metasarcus Roewer, 1913.
Ayacucho Roewer, 1949; Huancabamba gen. nov.; Incasarcus Kury and Maury, 1998; Lumieria gen. nov.; Metasarcus Roewer, 1913; and Tschaidicancha Roewer, 1957.
Metasarcidae can be easily diagnosed by other Gonyleptoidea by only one feature, the penis with lateral finger-like sacs. Only one genus (Metalibitia, Cosmetidae) of Gonyleptoidea possess lateral sacs on ventral plate but its shape and position is different from it and not homologous. It differs from Stygnidae by having ocularium undivided; by Gonyleptidae by pedipalpal femur with long spines; by Cosmetidae by pedipalpus somewhat cylindrical and with spines; by Agoristenidae by having tarsal process; by Cranaidae by pedipalpal femur smooth or small-tuberculate.
Gonyleptoidea with eye mound tall and rounded (Ayacucho) or low, medially depressed (the other genera); ocularium with a pair of low tubercles, a pair of high spines or unarmed. Chelicerae swollen in males of some species (also in some females of A. titschacki). Pedipalpus long and robustly armed; femur sub cylindrical, not flattened (slightly flattened in Ayacucho); femur and patella in males with a proapical spine (except Ayacucho). Alpha-type DSS (Incasarcus and majority of Ayacucho), gamma-type DSS (Metasarcus fellinii sp. nov. and Ayacucho spielbergi sp. nov.), gamma-P-type DSS (some Metasarcus and Tschaidicancha joseochoai sp. nov.) and kappa-type DSS (Huancabamba gen. nov., Lumieria gen. nov., Tschaidicancha and some Metasarcus). DS moderate to densely granulate. Scutal area I undivided or divided (Lumieria gen. nov. and some Metasarcus and Tschaidicancha); area III generally armed with a pair of high spines, a pair of low spines (I. argenteus) or tubercles (most Ayacucho, Metasarcus trispinosus sp. nov.) or unarmed (some Ayacucho, I. ochoai, Metasarcus vacafloresae sp. nov.). Male coxa IV generally unarmed; armed with an acute long prolateral tubercle in most Ayacucho or with a retrolateral armature in Metasarcus bergmani sp. nov. and M. limachii sp. nov. Femur IV shorter than DSL in most Ayacucho, about same size in Huancabamba gen. nov. and much longer in the other genera. Tarsal process present. VP of penis well defined, generally subrectangular, without cleft, with three to many (more than 13) pairs of MS C, and lateral finger-like sacs. Stylus long and generally laterally flattened, dorsoventrally widened (broad and sturdy Lumieria gen. nov. and cylindrical in some species); generally with swollen apex and with a caruncle. Dorsal process of glans absent or present.
The family Metasarcidae occurs in Andean Mountains of Bolivia and Peru, the southern limit being the border with Argentina and the northern limit the Huancabamba depression, situated in northern Peru (Figs
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 (T. chaplini sp. nov., T. joseochoai sp. nov., A. pasolinii sp. nov., T. scorsesei sp. nov.), Centro Turistico Ilpa/Peru (L. woodyalleni gen. et sp. nov., L. antonionii gen. et sp. nov.), Zongo/Bolivia (M. kurosawai sp. nov., M. vacafloresae sp. nov.), Cutervo/Peru (A. uniseriatus comb. nov., H. kubricki gen. et sp. nov., A. spielbergi sp. nov.). Ayacucho titschacki, which occurs in the Peruvian Central Andes, (near to Ocollo, Virgem de Cacharras de Cocha) and Ayacucho tapacocha nom. nov., which occurs in northern Peru, are the species with the largest distributions recorded in the Ayacucho and Ancash areas, respectively. This high level of endemism is comparable to the eastern coast of South America, where most species occupy small areas of endemism (see Da-Silva et al. 2017). However, the harvestmen fauna from Peru and Bolivia is poorly sampled, which prevents a more detailed comparison with other regions.
All Bolivian species of Metasarcidae belong to the type genus, Metasarcus, and occur in the eastern Andes from La Paz to Tarija Province (M. fellinii sp. nov. is the southernmost species of the family), the Altiplano being the northern distribution limit. Its sister genus, Incasarcus is present only in the Peruvian Cusco Department, in Montane tall grass vegetation (Puna) and scrub and montane Rain Forest. Both genera are separated by Puna Seca and Titicaca lake, which means the Altiplano.
Ayacucho is widespread in most of the Peruvian Andean region, from Cajamarca to Ayacucho departments, the Rio Apurimac being the southern limit of its distribution. The only metasarcid species recorded from the western Andean foothills is A. roeweri nom. nov., from Rio Fortaleza (2700 m above sea level, Ancash, Cajacay, Peru), where the riparian forest El Bosque de Fortaleza is found. Most species can be found in two types of vegetation, the Mountain short grass and Andean wastes (Quechua) and Mountain tall grass and scrub (Puna). One species, A. pasolinii sp. nov. was recorded from the Mountain Rain Forest (Parque Nacional Yanachaga-Chemillén, Oxapampa, Peru).
The monotypic genus Huancabamba gen. nov. is recorded only in Cutervo (Cajamarca Department), in Mountain Rain Forest.
Lumieria gen. nov. has only two species, sympatrically distributed at Centro Turistico Ilpa (Junin Department – Bolivia). This locality is covered by Mountain tall grass and scrub.
Tschaidicancha has four species recorded in only three regions, two of which are very close to each other. T. scorsesei sp. nov., T. joseochoai sp. nov., and T. chaplini sp. nov. occur in Mountain Rain Forest, and T. weyrauchi in areas with scrubs of Mountain Tall Grass and Scrub.
Ayacucho
Roewer, 1949: 57 (desc);
Ayachuco
[lapsus calami]:
Cajamarca
Roewer, 1952: 41 (desc);
Cargaruaya
Roewer, 1956: 439 (desc);
Palcares
Roewer, 1957: 72 (desc);
Cajacaybia
Roewer, 1957: 73 (desc);
Pinocchio [in part] Roewer, 1957: 70 (desc).
Tapacochana
Roewer, 1957: 73 (desc);
PERU. Ancash, Ayacucho, Cajamarca, Huancavellica, Junín, La Libertad and Pasco (Fig.
Ayacucho can be differentiated from all other Metasarcidae genera by its short leg IV (femur IV length/DS length < 1.5). Most of its species can be distinguished from other genera by having, alpha-type DSS; a femur of pedipalpus slightly flattened and males without a proapical spine (present in all other genera), an ocularium high and rounded and a DS densely granulated.
Alpha-type DSS (Figs
Habitus, dorsal of Ayacucho. A A. bambamarca (Roewer, 1957) comb. nov., male; B A. glauberrochai sp. nov., male; C A. inermis (Roewer, 1957) comb. nov., male; D A. insignitus (Roewer, 1956) comb. nov., female; E A. pasolinii sp. nov., male; F A. pomacocha sp. nov., male; G A. querococha sp. nov., male; H A. roeweri nom. nov., female; I A. silvae sp. nov., male; Legend bars = 1 mm.
Habitus, dorsal of males of Ayacucho. A A. spielbergi sp. nov.; B A. spiniger (Roewer, 1957) comb. nov.; C A. tapacocha nom. nov.; D A. titschacki Roewer, 1949; E Ayacucho triarmatus nom. nov.; F A. weyrauchi (Roewer, 1952) comb. nov.; G Ayacucho vargasllosai sp. nov.; H A. uniseriatus (Roewer, 1959) comb. nov.; Legend bars = 1 mm.
Ayacucho bambamarca (Roewer, 1957) comb. nov.; Ayacucho inermis (Roewer, 1957) comb. nov.; Ayacucho glauberrochai sp. nov.; Ayacucho insignitus (Roewer, 1956) comb. nov.; Ayacucho pasolinii sp. nov.; Ayacucho pomacocha sp. nov.; Ayacucho querococha sp. nov.; Ayacucho roeweri nom. nov.; Ayacucho silvae sp. nov.; Ayacucho spielbergi sp. nov.; Ayacucho spiniger (Roewer, 1957) comb. nov.; Ayacucho tapacocha nom. nov.; Ayacucho titschacki Roewer, 1949; Ayacucho triarmatus nom. nov.; Ayacucho uniseriatus (Roewer, 1959) comb. nov.; Ayacucho vargasllosai sp. nov.; Ayacucho weyrauchi (Roewer, 1952) comb. nov.
Cajamarca bambamarca
Roewer, 1957: 76 (desc.), fig. 33 (male femur IV);
Cajamarca triseriata
Roewer, 1957: 75 (desc.), fig. 32 (male femur IV);
MALE: Measurements (n=7) DSW: 3.6–5.1 (5.1); DSL: 3.8–5.2 (5.2); CL: 1.5–1.7 (1.7). FIVL: 4.4–5.2 (4.6). ChL: 1.5–3.7 (3.6). Coloration (in ethanol): Predominantly yellowish. Dark spots on carapace. Areas I–IV, lateral and posterior margin of dorsal scutum and free tergites I–III more brownish. Dorsum: (Fig.
Similar to Ayacucho triarmatus nom. nov. and A. weyrauchi comb. nov. by possessing three rows of spiniform tubercles in male femur IV (Fig.
Considering that Cajamarca bambamarca and C. triseriata were described in the same work (
(Fig.
Type material
: Of C. bambamarca: Holotype ♂, ‘PERU, Cajamarca, Bambamarca | 2,800m | 29/VI/1956, Weyrauch leg. (
MALE: Measurements (n=9) DSW: 3.3–3.4 (3.3); DSL: 4.6–5.2 (4.6); CL: 1.5–2.0 (1.5). FIVL: 5.0–5.3 (5.0). ChL: 1.7–3.0 (2.1). Coloration: (Fig.
Habitus, dorsal of Metasarcus. A M. beni sp. nov., male; B M. bergmani sp. nov., male; C M. bolivianus Roewer, 1913, female; D M. clavifemur (Roewer, 1929), male; E M. fellinii sp. nov., male; F M. kurosawai sp. nov., male; G M. limachii sp. nov., male; H M. trispinosus sp. nov., male; I M. vacafloresae sp. nov., male; Legend bars = 1 mm.
Similar to Ayacucho pomacocha sp. nov., A. silvae sp. nov., A. titschacki and A. vargasllosai sp. nov. in the following combination of characteristics: dorsal scutum densely granulate; ocularium and areas I–IV of DS unarmed or armed with tiny tubercles, slightly greater than granules; posterior margin of DS and free tergites I–III with median rows of acuminate tubercles (Fig.
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.
Type material: Holotype ♂, ‘PERU, Huancavellica, Huancavellica, Quebrada Potreros | 12°46′10.7″S 75°01′02.5″W | 28/IV/2011, R. Pinto-da-Rocha, A. Benedetti, J. Ochoa & D. Silva leg. (MUBI) – Paratypes 1 ♂, 5 ♀, ‘ditto’ (MUBI); Paratypes 5 ♂, 11 ♀, ‘ditto’ (
Pinocchio inermis Roewer, 1957: 70 (desc.), fig. 3 (male coxa–patella IV).
Palcares inermis:
MALE: Measurements (n=1) DSW: 2.8; DSL: 3.2; CL: 1.0. FIVL: 2.0. ChL: 1.2. Coloration (in ethanol): Uniformly yellowish. In the original description: rusty-yellow body, dorsally darker than ventrally; legs rusty-yellow, slightly blackish. Dorsum: (Fig.
Male trochanter–tibia IV of Ayacucho, dorsal and ventral view, respectively. A–B A. bambamarca (Roewer, 1957) comb. nov.; C–D A. glauberrochai sp. nov.; E–F A. inermis (Roewer, 1957) comb. nov.; G–H A. pasolinii sp. nov.; I–J A. pomacocha sp. nov.; K–L A. querococha sp. nov.; M–N A. silvae sp. nov.; O–P A. spielbergi sp. nov.; Q–R A. spiniger (Roewer, 1957) comb. nov.; S–T: A. tapacocha nom. nov.; U–V A. titschacki Roewer, 1949; W–X: A. triarmatus nom. nov.; Legend bars = 1 mm.
Resembles Ayacucho spiniger comb. nov. because two ventral rows of tubercles in femur IV (Fig.
(Fig.
Type material: Holotype ♂, ‘PERU, La Libertad, Huamachuco, 3,200m a.s.l. without date, Weyrauch leg. (
Cargaruaya insignita
Roewer, 1956: 439 (desc.), figs. 11–12 (ocularium), 13 (female pedipalp).
FEMALE: Measurements (n=1) DSW: 4.0; DWL: 4.5; CL: 1.5. FIVL: 4.2. ChL: 1.5. Coloration (in ethanol): Predominantly yellowish with dark spots at carapace, areas, lateral and posterior margin of the dorsal scutum and free tergites. Dorsum: (Fig.
It differs from females of Ayacucho uniseriatus comb. nov. and A. weyrauchi comb. nov. because it has one pair of median tubercles in areas I–IV; from A. inermis comb. nov., A. spiniger comb. nov. and A. querococha sp. nov. because it has sparser granulation on carapace, and it is larger than A. inermis comb. nov.; A. bambamarca comb. nov. and A. tapacocha nom. nov. because it has smaller tubercles present in the posterior margin of dorsal scutum and free tergite I–II; and A. roeweri nom. nov. because it does not present a huge median spine in free tergites I–II. (Fig.
This species is known only by the female holotype, which ends up being a taxonomic problem, since it is not uncommon for females of Ayacucho, whose males are clearly morphologically distinct, to be very similar to each other. This similarity between the females justifies the not so much informative diagnosis (see above). Geographically, A. insignitus comb. nov. occurs near to three other species of the genus: A. inermis comb. nov., A. triarmatus nom. nov. and A. weyrauchi comb. nov. (Fig.
(Fig.
Type material: Holotype ♀, ‘PERU, La Libertad. Hacienda Llaguén, Rejo Cargaruay forest, 2,650m a.s.l., 14/XII/1952, Koepcke leg. (
MALE: Measurements (n=1) DSW: 3.2; DSL: 3.6; CL: 1.2. FIVL: 5.4. ChL: 1.0. Coloration: (Fig.
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 Incasarcus. A–B Trochanter–tibia IV of I. argenteus Kury & Maury, 1998, dorsal and ventral view, respectively; C–D Trochanter–tibia IV of I. dianae Kury & Maury, 1998, dorsal and ventral view, respectively; E–F Trochanter–tibia IV of I. pictus Kury & Maury, 1998, dorsal and ventral view, respectively; G–H Trochanter–tibia IV of I. viracocha Kury & Maury, 1998, dorsal and ventral view, respectively; I Trochanter–patella of I. viracocha, lateral view; J Trochanter–patella of I. ochoai Kury & Maury, 1998, large male, dorsal view; K–L Trochanter–tibia of I. ochoai, small male, dorsal and ventral view, respectively; M Trochanter–patella of I. ochoai, large male, lateral view; N Trochanter–patella of I. ochoai, small male, lateral view; Legend bars = 1 mm.
The specific epithet of masculine gender, in the genitive form, dedicated to the Italian writer and filmmaker Pier Paolo Pasolini (1922–1975).
(Fig.
Type material: Holotype ♂, ‘ PERU, Pasco, Oxapampa, Parque Nacional Yanachaga-Chemillén, 10°32′42.1″S 75°21′24.4″W, 22/IV/2011, R. Pinto-da-Rocha, A. Benedetti, J. Ochoa & D. Silva leg. (MUBI) – Paratypes 2 ♀, ‘ditto’ (
MALE: Measurements (n=12). DSW: 3.2–4.0 (3.2); DSL: 4.7–5.3 (4.7); CL: 1.7–1.0 (1.7); FIVL: 4.5–5.0 (4.5); ChL: 1.2–3.5 (2.2). Coloration: (Fig.
Male trochanter–patella/tibia IV of Metasarcus, dorsal and ventral view, respectively. A–B M. beni sp. nov.; C–D M. bergmani sp. nov.; E–F M. clavifemur (Roewer, 1929); G–H M. fellinii sp. nov.; I–J M. kurosawai sp. nov.; K–L M. limachii sp. nov.; M–N M. trispinosus sp. nov.; O–P M. vacafloresae sp. nov.; Legend bars = 1 mm.
Similar to Ayacucho glauberrochai sp. nov., A. silvae sp. nov., A. titschacki and A. vargasllosai sp. nov. in the following combination of characteristics: dorsal scutum densely granulate; ocularium and areas I–IV of DS unarmed or armed with tiny tubercles, slightly greater than granules; posterior margin of DS and free tergites I–III with median rows of acuminate tubercles (Fig.
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.
Type material: Holotype ♂, ‘ PERU, Junín, Laguna Pomacocha, 4,500m a.s.l., 11°46′36″S, 75°14′07″W , 27/IV/2011, R. Pinto-da-Rocha, A. Benedetti, J. Ochoa & D. Silva leg. (MUBI) – Paratypes 2 ♂, 3 ♀, ‘ditto’ (MUBI); Paratypes 3 ♂, 3 ♀, ‘ditto’ (
MALE: Measurements (n=3) DSW: 3.4–4.1 (4.0); DSL: 4.2–5.0 (4.8); CL: 1.4–1.7 (1.7). FIVL: 3.3–4.0 (3.5). ChL: 1.7–2.8 (2.4) Coloration (alive): Yellowish with black spots on carapace, lateral part of the scutal areas I–IV, lateral margins of dorsal scutum and legs. Dorsum: (Fig.
Male trochanter–patella/tibia IV of Huancabamba gen. nov., Lumieria gen. nov. and Tschaidicancha, dorsal and ventral view, respectively. A–B H. kubricki gen. et sp. nov.; C–D L. antonionii gen. et sp. nov.; E–F L. woodyalleni gen. et sp. nov.; G–H T. chaplini sp. nov.; I–J T. joseochoai sp. nov.; K–L T. scorsesei sp. nov.; M–N T. weyrauchi Roewer, 1957; Legend bars = 1 mm.
Similar to Ayacucho tapacocha nom. nov. because the tibia IV is armed (Fig.
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.
Type material: Holotype ♂, ‘PERU, Ancash, Parque Nacional Huascarán, Laguna Querococha, 4,024 m a.s.l., 09°43′38.8″S 77°19′47.9″W , 17/V/2010, R. Pinto-da-Rocha & D. Silva leg. (
Cajacaybia spinigera Roewer, 1957: 74 (desc.), figs. 27 (dorsal habitus), 28 (pedipalpus); Kury, 2003: 144 (cat.).
FEMALE: Measurements (n=1) DWS: 3.6; DSL: 4.0; CL: 1.4. FIVL: 3.5. ChL: 1.3. Coloration (in ethanol): Uniformly yellowish (reddish-brown, in the original description), with darker spots on the carapace, especially behind ocularium and legs. Dorsum: (Fig.
Penis of Metasarcidae, dorsal and lateral views, respectively. A–B A. bambamarca (Roewer, 1957) comb. nov.; C–D Ayacucho inermis (Roewer, 1957) comb. nov.; E–F Ayacucho pasolinii sp. nov.; G–H Ayacucho spielbergi sp. nov.; I–J Ayacucho titschacki Roewer, 1949; K–L Ayacucho triarmatus nom. nov.; Legend bars = 0.1 mm.
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 Cajacaybia spinigera and Palcares spiniger were described in the same work (
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.
Type material: Hototype ♀, ‘PERU. Ancash. Cajacay, Rio Fortaleza, 2,700 m a.s.l., 5/III/1956, Weyrauch leg. (
MALE: Measurements (n=10) DSW: 1.7–3.0 (2.8); DSL: 4.0–4.3 (4.2); CL: 1.3–1.5 (1.5). FIVL: 2.6–2.8 (2.6). ChL: 1.2–2.5 (2.2). Coloration: (Fig.
Penis of Metasarcidae, dorsal and lateral views, respectively. A–B Lumieria woodyalleni gen. et sp. nov.; C–D Metasarcus beni sp. nov.; E–F Metasarcus kurosawai sp. nov.; G–H Metasarcus limachii sp. nov.; I–J Tschaidicancha joseochoai sp. nov.; K–L Tschaidicancha weyrauchi Roewer, 1957; Legend bars = 0.1 mm.
Similar to Ayacucho glauberrochai sp. nov., A. pomacocha sp. nov., A. titschacki and A. vargasllosai sp. nov. in the following combination of characteristics: dorsal scutum densely granulate; ocularium and areas I–IV of DS unarmed or armed with tiny tubercles, slightly greater than granules; posterior margin of DS and free tergites I–III with median rows of acuminate tubercles (Fig.
The specific epithet of feminine gender, in the genitive form, dedicated to Dr. Diana Silva D. (
(Fig.
Type material: Holotype ♂, ‘ PERU, Pasco, Near to Cerro de Pasco, 10°41′39″S 76°13′08″W, 23/IV/2011 , R. Pinto-da-Rocha, A. Benedetti, J. Ochoa & D. Silva leg. (MUBI) – Paratypes 2 ♂, 5 ♀, ‘ditto’ (MUBI); Paratypes 2 ♂, 5 ♀, ‘ditto’ (
MALE: Measurements (n=3) DSW: 4.0–4.9 (4.9); DSL: 4.6–5.0 (5.0); CL: 1.6–2.0 (2.0). FIVL: 7.6–8.6 (8.6). ChL: 2.1–3.9 (3.9). Coloration: (Fig.
It differs from other species of the genus by the combination of the following characteristics: Gamma-type DSS (Fig.
The specific epithet of masculine gender, in the genitive form, dedicated to the American filmmaker, producer and screenwriter Steven Allen Spielberg.
(Fig.
Type material: Holotype ♂, ‘PERU, Cajamarca, Parque Nacional Cutervo, Puente Suro, 6°12′10″S 78°44′22″W, 22/V/2010, R. Pinto-da-Rocha & D. Silva leg. (
Palcares spiniger
Roewer, 1957: 72 (desc.), fig. 4 (dorsal habitus, femur IV), 5 (details of femur IV armature), 6 (pedipalpus);
Palcares serrifemur
Roewer, 1959: 70 (desc.) fig. 2A (dorsal habitus, femur IV), 2b (details of femur IV armature);
MALE: Measurements (n=4) DSW: 3.4–3.8 (3.7); DSL: 4.2–4.6 (4.6); CL: 1.4–1.5 (1.4). FIVL: 4.9–5.9 (5.9). ChL: 1.3–2.7 (2.7). Coloration: (Fig.
Similar to Ayacucho inermis comb. nov. because of two rows of tubercles on the femur IV (pro and retrolateral; Fig.
(Fig.
Type material
: Of P. spiniger: Holotype ♂, ‘PERU, Junín, Campañillaya, near to Palca in Rio Tarma, 2,600 m a.s.l., 06/I/1955, Weyrauch leg. (
Tapacochana insignita
Roewer, 1957: 73 (desc.), figs. 25 (dorsal habitus), 26 (pedipalpus);
MALE: Measurements (n=15) DSW: 3.3–5.1 (3.3); DSL: 4.0–5.6 (4.0); CL: 1.3–2.2 (1.3). FIVL: 6,2–6,5. ChL: 1.3–4.0 (1.3). Coloration: (Fig.
Similar to Ayacucho querococha sp. nov. because armed tibia IV (Fig.
Roewer described Cargaruaya insignita in 1956 and Tapacochana insignita in 1957. Since they are both species in the genus Ayacucho, Tapacochana insignita Roewer, 1957 is a secondary homonym of Cargaruaya insignita Roewer, 1957 and must be replaced. Consequently, we create Ayacucho tapacocha nom. nov. as a replacement name for Roewer’s specific epithet.
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.
Type material: Holotype ♂, ‘PERU, Ancash, Tapacocha | Rio Fortaleza, 3,200 m a.s.l., 04/III/1956, Weyrauch leg. (
Ayacucho titschacki
Roewer, 1949: 57 (desc.), figs. 114a (dorsal habitus), 114b (ventral habitus), 114c (chelicera, pedipalpus and carapace in lateral view); 114d (detail of coxa IV), 114e (sternum), 114f (chelicera in lateral view);
MALE: Measurements (n=2) DSW: 3.0–4.1; DSL: 4.5–5.4; CL: 1.5–2.2. FIVL: 4.8–5.4. ChL: 2.5–4.0. Coloration: (Fig.
Similar to Ayacucho glauberrochai sp. nov., A. pomacocha sp. nov., A. silvae sp. nov. and A. vargasllosai sp. nov. in the following combination of characteristics: dorsal scutum densely granulate; ocularium and areas I–IV of DS unarmed or armed with tiny tubercles, slightly greater than granules; posterior margin of DS and free tergites I–III with median rows of acuminate tubercles (Fig.
The male holotype and paratypes examined by Roewer (
(Fig.
Type material. Holotype ♀, ‘PERU, Ayacucho, Ayacucho, without date and leg. (
Tapacochana triseriata
Roewer, 1959: 69 (desc.), fig. 1 (dorsal habitus and trochanter–femur IV);
MALE: Measurements (n=1) DSW: 4.0; DSL: 4.4; CL: 1.4. FIVL: 4.2. ChL: 1.2. Coloration (in ethanol): Predominantly yellow with dark spots on the scutal areas I–IV, lateral and posterior margin of the dorsal scutum and free tergites. Dorsum: (Fig.
Similar to Ayacucho bambamarca comb. nov. and A. weyrauchi comb. nov. because three rows of spiniform tubercles in femur IV (Fig.
Roewer described Cajamarca triseriata in 1957 and Tapacochana triseriata in 1959. Since they are considered here as belonging to the genus Ayacucho, Tapacochana triseriata Roewer, 1959 is a secondary homonym of Cajamarca triseriata Roewer, 1957 and must be replaced. C. triseriata Roewer, 1957 is considered here synonymy of Cajamarca bambamarca Roewer, 1957, but junior synonyms are also combinations with the genus where they are included and consequently compete for homonymy. Therefore, we create Ayacucho triarmatus nom. nov. as a replacement name for Roewer’s previous name.
The specific epithet, an adjective in nominative singular, formed by Latin prefix tri- + Latin armātus, ta, tum (armed), in reference to three rows of tubercles on male femur IV.
(Fig.
Type material: Holotype ♂, ‘PERU, Cajamarca, near San Juan, between Chiclayo and Cajamarca, 1,900 m a.s.l., 06/VII/1956, Weyrauch leg. (
Cajamarca uniseriata
Roewer, 1957: 76 (desc.), fig. 34 (femur IV);
MALE: Measurements (n=5) DSW: 3.6–4.6 (4.4); DSL: 4.1–4.8 (4.3); CL: 1.3–1.7 (1.7). FIVL: 3.5–4.0 (4.0). ChL: 1.8–2.7 (2.6). Coloration: (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 C. uniseriata: Roewer designated one male as holotype and two males as paratypes, but the type material is preserved without any distinction in the same vial. Therefore, it is not possible to recognize with absolute certainty which of the males is the holotype. Because of this, one of the males, whose femur IV most closely resembles the drawing in the original description, was separated as the holotype (although it is important to point out that the drawing does not faithfully represent any of the specimens).
(Fig.
Type material: Holotype ♂, ‘PERU, Cajamarca, Cutervo, 15/VI/1956, Weyrauch leg. (
MALE: Measurements (n=1) DSW: 3.3; DSL: 4.6; CL: 1.7. FIVL: 4.7. ChL: 2.5. Coloration: (Fig.
Similar to Ayacucho glauberrochai sp. nov. A. pomacocha sp. nov., A. silvae sp. nov., and A. titschacki in the following combination of characteristics: dorsal scutum densely granulate; ocularium and areas I–IV of DS unarmed or armed with tiny tubercles, slightly greater than granules; posterior margin of DS and free tergites I–III with median rows of acuminate tubercles (Fig.
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.
Type material: Holotype ♂, ‘PERU, Junín, Cruce Mina, Cemento Andino, 11°22′45.4″S 75°52′43.5″W,| 22/IV/2011, R. Pinto-da-Rocha, A. Benedetti, J. Ochoa & D. Silva leg. (MUBI) – Paratype 1 ♀, ‘ditto’ (MUBI); Paratype 1 ♀, ‘ditto’ (
Cajamarca weyrauchi
Roewer, 1952: 41 (desc.);
Cajamarca affinis
Roewer, 1957: 75 (desc.), fig. 31 (male femur IV);
MALE: Measurements (n=2) DSW: 4.5; DSL: 4.6–5.0 (5.0); CL: 1.5–1.7 (1.5). FIVL: 4.2–4.5 (4.5). ChL: 1.5–3.0 (3.0). Coloration (in ethanol): Predominantly yellowish. Areas I–IV, posterior margin of DS and free tergites I–IV brown. Dorsum: (Fig.
Similar to Ayacucho bambamarca comb. nov. and A. triarmatus nom. nov. for presenting three rows of acuminate tubercles in the male femur IV (Fig.
(Fig.
Type material
: Of C. weyrauchi: Holotype ♂, ‘PERU, Cajamarca, Cajamarca,| 2,750 m a.s.l., without date, Weyrauch leg. (
Huancabamba kubricki gen. et. sp. nov. by present designation.
Huancabamba gen. nov. can be distinguished from all other Metasarcidae genera by the combination of following: Kappa-type DSS; male femur IV at least 1.6 longer than DS length and with low tubercles; ocularium with two low tubercles; area III with spines; coxa IV apex reaching area III; penis with more than 13 MS C; a dry-mark in depression of ocularium, carapace, and lateral region to ocularium.
Kappa-type DSS, with carapace very wide, constriction I weakly marked and coda undefined, coalescing with mid-bulge. Ocularium low, medially depressed. Ocularium with two low tubercles. Areas of dorsal scutum moderately tuberculate. Area I undivided. Area III armed with two high spines. Posterior margin of DS armed with a pair of high tubercles. Coda short, without constriction. Coxa IV reaching area III. Coxa IV unarmed (Fig.
The genus name, a noun in the nominative singular, from Quechua, huanca (stone) + bamba (plain). It refers to Huancabamba depression, an interruption of the Andean Mountains, located between southern Ecuador and northern Peru. This depression constitutes a biogeographic barrier between the northern Andes and the central Andes. The name is a reference to this Metasarcidae genus occurred fartherst north. Gender: feminine.
(Fig.
Huancabamba kubricki gen. et. sp. nov.
MALE: Measurements (n=3) DSW: 4.0–4.2 (4.2); DSL: 3.6–4.3 (4.1); CL: 1.5–2.0 (2.0). FIVL: 6.6–7.3 (7.3). ChL: 1.2–3.6 (3.2). Coloration: (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.
Type material: Holotype ♂, ‘PERU, Cajamarca, near Cutervo, 06°20′42″S 78°49′19″W, 20/V/2011, R. Pinto-da-Rocha & D. Silva leg. (
Incasarcus
Kury & Maury, 1998: 145 (desc);
Incasarcus can be differentiated from other Metasarcidae genera by the combination of following: alpha-type DSS; males with a proapical spine on the pedipalpus femur; area I undivided; male femur IV at least 1.6 longer than dorsal scutum; ocularium with two low tubercles or spines; area III with spines; coxa IV apex reaching area IV or posterior margins of DS; penis with less than 10 MS C and stylus thin thickness.
Alpha-type DSS. Ocularium low, medially depressed. Ocularium with two low tubercles or tall spines. Areas of dorsal scutum moderately to densely tuberculate. Area I undivided. Area III armed with two tall spines (most species), a pair of short spines (I. argenteus), or unarmed (I. ochoai). Posterior margin unarmed. Coxa IV reaching area IV or posterior margin. Coxa IV unarmed (Fig.
(Fig.
Incasarcus argenteus Kury & Maury, 1998; Incasarcus dianae Kury & Maury, 1998; Incasarcus ochoai Kury & Maury, 1998; Incasarcus pictus Kury & Maury, 1998; Incasarcus viracocha Kury & Maury, 1998;
Incasarcus argenteus
Kury & Maury, 1998: 155 (desc.), 159 (key), figs. 32 (male dorsal habitus, chelicerae, pedipalpus), 33 (penis dorsal view), 34 (penis lateral view), 35 (male lateral habitus, chelicera, pedipalpus), 36 (female dorsal habitus, chelicerae, pedipalpus), 37 (male trochanter–tibia IV);
MALE: Measurements (n=3) DSW: 5.5–6.1 (6.1); DSL: 6.1–7.4 (7.4); CL: 2.5–3.1 (3.1). FIVL: 9.7–10.8 (10.8). ChL: 3.4–5.2 (5.2). Coloration (in ethanol): Carapace (more accentuated behind and next to ocularium), area I and lateral margins of dorsal scutum, free tergites I–II, coxa IV (dorsal and ventral surfaces) and free sternites white-silver. Remaining mesotergum, pedipalpus and legs dark brown. Chelicerae yellowish brown. Dorsum: (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.
Type material: Holotype ♂, ‘PERU, Cusco, Urubamba province, Ollantaytambo district, Abra de Málaga, Cancayoc, 3,000 m a.s.l., 13°16′S 72°16′W, 27/VIII/1995, J. Ochoa leg. (
Incasarcus dianae
Kury & Maury, 1998: 146 (desc.), 159 (key), figs. 1 (male dorsal habitus, chelicerae, pedipalpus, trochanter–patella IV), 2 (penis dorsal view), 3 (penis lateral view), 4 (male lateral habitus), 5 (female dorsal habitus, chelicerae, pedipalpus), 6 (male ventral habitus); 7–10 (tarsi I–IV);
MALE: Measurements (n=3) DSW: 4.0–4.3 (4.0); DSL: 4.5–5.1 (4.5); CL: 1.1–1.5 (1.1). FIVL: 11.3–11.4 (11.4). ChL: 3.3–3.4 (3.4). Coloration (in ethanol): Yellow with black spots covering practically the entire dorsal scutum; area III brown. Pedipalpus, chelicerae and legs I–IV brown. Dorsum: (Fig.
It differs from other species of the genus by the set of following characters: DS without silver-white coloration; DS slightly granulate; area I with a pair of lateral tubercles; area III with a pair of spines (Fig.
(Fig.
Type material: Holotype ♂, ‘PERU, Cusco, Paucartambo province, Manu National Park, road to Paucartambo-Pilcopata, 2,900 m a.s.l., 13°01′40″S 71°16′40″W | 19/II/1990, A. Cano & D. Silva leg. (
Incasarcus ochoai
Kury & Maury, 1998: 152 (desc.), 160 (key), figs. 21 (male dorsal habitus, chelicerae, pedipalpus, trochanter–patella IV), 22 (penis dorsal view); 23 (penis lateral view), 24 (male lateral habitus, chelicera, pedipalpus), 25 (female dorsal habitus), 26 (male sternum, coxae I–IV); 27 (male trochanter–patella IV), 28–31 (tarsi I–IV);
MALE: Measurements (n=3) DSW: 5.1–5.2 (5.2); DSL: 6.3–6.5 (6.5); CL: 1.6–1.7 (2.6). FIVL: 11.0–11.2 (11.2). ChL: 3.5–4.7 (3.5). Coloration (in ethanol): Yellow with black spots covering the areas of the dorsal scutum; pedipalpus and chelicerae brown. Legs I–IV yellow. Dorsum: (Fig.
It differs from other species of the genus by the set of following characters: DS without silver-white coloration; DS granulate; ocularium, areas I–IV and free tergites I–III unarmed (Fig.
(Fig.
Type material: Holotype ♂, ‘PERU, Cusco, Urubamba Province, Huayllabamba district, Yanacocha, Huayocari, Huayoccare, 3,000–4,000 m a.s.l., 13°20′S 72°02′W, 14/XI/1992, J.C. Chaparro leg. (
Incasarcus pictus
Kury & Maury, 1998: 149 (desc.), 160 (key), figs. 11 (male dorsal habitus, chelicerae, pedipalpus, trochanter–patella IV), 12 (penis dorsal view), 13 (penis lateral view), 14 (male lateral habitus), 15 (female dorsal habitus), 16 (male sternum and coxae I–IV), 17–20 (tarsi I–IV);
MALE: Measurements (n=1) DSW: 4.9; DSL: 5.8; CL: 1.6. FIVL: 11.5. ChL: 3.2. Coloration (in ethanol): DS with conspicuous white spots, a pair of small rounded spots on carapace, behind ocularium and a large spot covering all area I. The remaining DS yellow with small black spots; chelicerae, pedipalpus, area III and legs I–III brown; leg IV black. Dorsum: (Fig.
It differs from other species of the genus by the set of following characters: DS with white coloration on carapace (behind ocularium) and area I; DS granulate; ocularium and areas II and IV unarmed; area I with a pair of tubercles; area III with a pair of spines; free tergites I–III with a row of tubercles (Fig.
(Fig.
Type material: Holotype ♂, ‘PERU, Cusco, Wiñayhuaina, Inca trail, 2,700–3,100 m a.s.l., 13°09′S 72°31′W, 10/II/1990, D. Silva leg. (
Incasarcus viracocha
Kury & Maury, 1998: 157 (desc.), 160 (key), figs. 38 (male dorsal habitus, chelicerae, pedipalpus, trochanter–patella IV), 39 (penis dorsal view), 40 (penis lateral view), 41 (male lateral habitus), 42 (female dorsal habitus), 43 (male femur IV);
MALE: Measurements (n=3) DSW: 4.9–5.0 (4.9); DSL: 5.9–6.6 (5.9); CL: 2.4–3.0 (2.4). FIVL: 11.7–11.8 (11.8). ChL: 3.5–4.2 (4.2). Coloration (in ethanol): Brownish yellow body with reticular spots covering practically the entire carapace and chelicerae. Dorsum: (Fig.
It differs from other species of the genus by the set of following characters: DS without white spots on DS; DS granular; ocularium unarmed and with few granules or smooth; areas I, II and IV unarmed; area III with a pair of spines; free tergites I–III with a row of tubercles, unarmed (Fig.
(Fig.
Type material: Holotype ♂, ‘PERU, Cusco, Urubamba province, Machu Picchu, 2,200–2,500m a.s.l., 13°07′S 72°34′W, 25/II/1994, J. Ochoa & J. Achicahuala leg. (
Lumieria antonionii gen. et. sp. nov., by present designation.
Lumieria gen. nov. can be distinguished from all other Metasarcidae genera by the combination of following: Kappa-type DSS; area I divided; ocularium medially depressed, with two high tubercles, area III with two spines; posterior margin and free tergites with one or two higher tubercles; femur IV much longer than dorsal scutum length; penis VP and stylus robust and thick in lateral view; conspicuous dry-marks on the carapace, grooves of DS and free tergites.
Kappa-type DSS, with constriction I well marked and constriction II absent. Coda undefined, coalescing with mid-bulge. Ocularium low, medially depressed. Ocularium unarmed, small tuberculate or smooth. Areas of dorsal scutum small to moderately tuberculate. Area I divided in two halves. Area III armed with two high spines. Posterior margin armed with one or a pair of high tubercles. Coda short, without constriction. Coxa IV reaching area IV or posterior margin. Coxa IV unarmed (Figs
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.
Lumieria antonionii gen. et sp. nov. and Lumieria woodyalleni gen. et sp. nov.
MALE: Measurements (n=10) DSW: 4.6–5.5 (5.5); DSL: 5.0–5.8 (5.6); CL: 1.1–1.5 (2.2). FIVL: 10.8–12.4 (12.4). ChL: 1.7–5.2. Coloration: (Fig.
Similar to Lumieria woodyalleni gen. et sp. nov. because ocularium is armed with a pair of spines, area I divided, area III with a great pair of spines, free tergites with large tubercles (Fig.
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.
Type material: Holotype ♂, ‘PERU, Junín, Centro Turístico Ilpa, near Comas, 11°42′37.1″S 75°04′20.2″W, 27/IV/2011, R. Pinto-da-Rocha, A. Benedetti, J. Ochoa & D. Silva leg. (MUBI) – Paratypes 5♂, 5 ♀, ‘ditto’ (
MALE: Measurements (n=1) DSW: 5.4; DSL: 5.8; CL: 1.3. FIVL: 11,8. ChL: 3.5. Coloration: (Fig.
Similar to Lumieria antonionii gen. et. sp. nov. by combination of following characters: ocularium with a pair of spines, area I divided, a long pair of spines in area III, free tergites with large tubercles (Fig.
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.
Type material: Holotype ♂, ‘PERU, Junín, Centro Turístico Ilpa, near Comas, 1°42′37.1″S 75°04′20.2″W, 27/IV/2011, R. Pinto-da-Rocha, A. Benedetti, J. Ochoa & D. Silva leg. (MUBI) – Paratype ♀, ‘ditto’ (
Metasarcus
Roewer, 1913: 304 (desc);
Metarascus
[lapsus calami]:
Chaconatus
Roewer, 1929: 275 (key), 276 (desc);
Chacoikeontus
Roewer, 1929: 275 (key), 278 (desc);
Metasarcus can be diagnosed from other Metasarcidae genera with long legs (except Ayacucho) by the combination of the following features: gamma-type; gamma-P-type or kappa-type DSS; ocularium unarmed; area III with two spines. Half of Metasarcus species present a spine, long projection or trifid projection in free tergite III, present in no other Metasarcidae species; coxa III with two apophyses; coxa IV reaching posterior margin of dorsal scutum or surpassing it; femur IV smooth or slightly granulate; penis stylus and VP thin thickness, with less than 10 MS C
Gamma-type DSS (M. bolivianus and M. fellinii sp. nov.; Fig.
(Fig.
Metasarcus beni sp. nov.; Metasarcus bergmani sp. nov.; Metasarcus bolivianus Roewer, 1913; Metasarcus clavifemur (Roewer, 1929); Metasarcus fellinii sp. nov.; Metasarcus kurosawai sp. nov.; Metasarcus limachii sp. nov.; Metasarcus trispinosus sp. nov. and Metasarcus vacafloresae sp. nov.
MALE: Measurements (n=2) DSW: 5.0–5.1 (5.1); DSL: 5.0–5.2 (5.2); CL: 1.2–1.3 (1.3). FIVL: 16.0–17.0 (17). ChL: 3.3–3.4 (3.4). Coloration (in ethanol): Yellow. Dorsum: (Fig.
Similar to Metasarcus kurosawai sp. nov. because ocularium unarmed, area I divided, area III with a pair of long spines (Fig.
The specific epithet, a noun in apposition, in reference to Beni Department (“El Beni”), Bolivia, Department of the type locality of the species.
(Fig.
Type material: Holotype ♂, ‘BOLIVIA, Beni, southwest of Yucumo | 15°23′S 66°59′W | 15–19/XI/1989, Coddington, Griswold, Silva, Larcher & Penaranda leg. (
MALE: Measurements (n=5) DSW: 5.2–6.0 (5.2); DSL: 5.5–6.8 (5.5); CL: 2.1–2.5 (2.1). FIVL: 12.5–13.0 (12.9). ChL: 2.0–2.1 (2.1). Coloration (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.
Type material: Holotype ♂, ‘BOLIVIA, Cochabamba, road next to Corani,| 17°11′18.9″S 65°53′49.2″W, 01/XII/2010, R. Pinto-da-Rocha, A. Benedetti, J. Ochoa & A. Saravia leg. (
Metasarcus bolivianus
Roewer, 1913: 305 (desc.), fig.123 (female dorsal habitus, pedipalpus);
Chaconatus armatipalpus
Roewer, 1929: 275 (key), 276 (desc.), fig. 43 (female lateral habitus, pedipalpus), fig. 43a (ocularium);
Metasarcus armatipalpus:
FEMALE: Measurements (n=2) DSW: 5.4–5.6 (5.4); DSL: 5.2–5.5 (5.5); CL: 2.2 (2.2). FIVL: 11.2–12.7 (12.7). ChL: 2.0–2.3 (2.0). Coloration (according to Roewer, 1913): Body, chelicerae and pedipalpus yellow-brown, in dark tone. Black legs. Granules of areas I–II yellow. Dorsum. (Fig.
Very similar to females of Metasarcus fellinii sp. nov. because a long spiniform apophysis in tergite III (Fig.
It is intriguing how Roewer described the same species from the Bolivian Chaco not only in different species and genera, but in different subfamilies (M. bolivianus was placed in Mitobatinae and Chaconatus armatipalpus in Prostygninae). As
BOLIVIA. Chaco.
Type material
: Of M. bolivianus: Holotype ♀, ‘BOLIVIA, Gran Chaco, whitout date and leg. (
Chacoikeontus clavifemur
Roewer, 1929: 279 (desc.), fig. 45;
Metasarcus clavifemur:
MALE: Measurements (n=5) DSW: 4.6–5.1 (5.0); DSL: 4.3–5.1 (5.1); CL: 1.7–2.0 (1.9). FIVL: 6.6–10.6 (10.6). ChL: 3.1–5.3. Coloration: (Fig.
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.
Type material: Lectotype ♂, ‘BOLIVIA, La Paz. Without date and collector data.’ (
MALE: Measurements (n=5) DSW: 4.0–4.7 (4.7); DSL: 3.8–4.3 (4.3); CL: 1.5–1.7 (1.7). FIVL: 7.4–9.0 (9.0). ChL: 1.5–1.6 (1.6). Coloration: (Fig.
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.
Type material: Holotype ♂, ‘BOLIVIA, Tarija, near Entre Ríos, Paraíso del Tordo, 21°35′54.7″S 64°08′58.4″W, 05/XII/2010, R. Pinto-da-Rocha, A. Benedetti, J. Ochoa & A. Saravia leg. (
MALE: Measurements (n=2) DSW: 4.0; DSL: 4.0–4.5 (4.5); CL: 1.5 FIVL: 9.2–10.0 (9.2). ChL: 3.3–3.8 (3.8). Coloration: (Fig.
Similar to Metasarcus beni sp. nov. because the ocularium is unarmed, area I divided, a pair of spines in the area III (Fig.
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.
Type material: Holotype ♂, ‘BOLIVIA, La Paz, Zongo, 16°10′32.4″S 68°08′11.9″W, 10/XII/2010, R. Pinto-da-Rocha, A. Benedetti & A. Saravia leg. (
MALE: Measurements (n=1) DSW: 6.1; DSL: 6.8; CL: 1.4. FIVL: 14.2. ChL: 2.4. Coloration: (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.
Type material: Holotype ♂, ‘BOLIVIA, La Paz, Yanacachi, 16°23′54.5″S 67°44′11.6″W, 12/XII/2010, R. Pinto-da-Rocha, A. Benedetti, J. Ochoa & A. Saravia leg. (
MALE: Measurements (n=5) DSW: 6.5–7.7 (7.6); DSL: 6.4–7.0 (7.0); CL: 2.3–2.8 (2.6). FIVL: 10–12 (11.5). ChL: 2.5–3.0 (3.0). Coloration: (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 tri- + Latin spīnōsus, a, um (thorny), in reference to the trifurcated apophysis of free tergite III.
(Fig.
Type material: Holotype ♂, ‘BOLIVIA, Cochabamba, road to Corani, 17°11′18.9″S 65°53′49.2″W, 01/XII/2010, R. Pinto-da-Rocha, A. Benedetti, J. Ochoa & A. Saravia leg. (
MALE: Measurements (n=4) DSW: 4.2–4.9 (4.7); DSL: 5.1–5.4 (5.4); CL: 2.1–2.5 (2.5). FIVL: 13.0–14.2 (14.2). ChL: 2.9–4.2 (3.6). Coloration: (Fig.
It differs from other species of the genus by the following set of characteristics: alpha type DSS; scutal area I undivided; areas I–IV unarmed (Fig.
The specific epithet of feminine gender, in the genitive form, in honor to Maria René Vacaflores, from Coleccíon Boliviana de Fauna (
(Fig.
Type material: Holotype ♂, ‘BOLIVIA, La Paz, Zongo, 16°10′32.4″S 68°08′11.9″W, 10/XII/2010, R. Pinto-da-Rocha, A. Benedetti, J. Ochoa & A. Saravia leg. (
Tschaidicancha
Roewer, 1957: 80 (desc);
Tschaidicancha can be distinguished from all other Metasarcidae genera by the combination of following: Kappa-type or gamma-P-type DSS; ocularium low, with two pairs of high spines; area III with two pairs of high spines; long leg IV (femur IV length/DS length > 1.6); male coxa IV apex reaching posterior margin; penis VP subrectangular, with distal-lateral projections (in most species), thin thickness; with less than five MS C; stylus laterally flattened, with apex inflated; dorsal process present.
Kappa-type, with straight posterior margin of DS; carapace long and wide; constriction I well marked, constriction II absent, mid-bulge slightly larger than carapace and coda undefined, coalescing with mid-bulge (T. chaplini sp. nov., T. scorsesei sp. nov. and T. weyrauchi; Fig.
(Fig.
Tschaidicancha chaplini sp. nov.; Tschaidicancha joseochoai sp. nov.; Tschaidicancha scorsesei sp. nov.; Tschaidicancha weyrauchi Roewer, 1957.
MALE: Measurements (n=5) DSW: 4.4–4.7 (4.6); DSL: 4.6–5.5 (5.5); CL: 1.9–2.4 (2.2). FIVL: 14.1–14.7 (14.7). ChL: 2.9–4.5 (4.5). Coloration: (Figs
Similar to Tschaidicancha scorsesei sp. nov. by the set of the following characteristics: reddish-brown coloration (Fig.
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.
Type material: Holotype ♂, ‘PERU, Pasco, Oxapampa, Parque Nacional Yanachaga-Chemillén, 10°32′42.1″S 75°21′24.4″W, 22/IV/2011, R. Pinto-da-Rocha, A. Benedetti, J. Ochoa & D. Silva leg. (MUBI) – Paratype 1 ♀, ‘ditto’ (MUBI); Paratypes 1 ♂, 1 ♀, ‘ditto’ (
MALE: Measurements (n=1) DSW: 4,4; DSL: 4,2; CL: 1,7; FIVL: 7,8; ChL: 3,0. Coloration: (Fig.
It differs from other species of the genus by the following set of characteristics: gamma-P-type DSS; ocularium with a pair of acuminate tubercles; DS and scutal areas smooth; scutal area I undivided; free tergites I–III unarmed (Fig.
The specific epithet of masculine gender, in the genitive form, dedicated Dr. José Ochoa C. (MUBI), for his great help in fieldwork in Bolivia and Peru.
(Fig.
Holotype ♂, ‘PERU, Pasco, Oxapampa, Parque Nacional Yanachaga-Chemillén 10°32′42.1″S 75°21′24.4″W, 22/IV/2011, R. Pinto-da-Rocha, A. Benedetti, J. Ochoa & D. Silva leg. (MUBI) — Paratype 1 ♀, ‘ditto’ (
MALE: Measurements (n=10) DSW: 4.1–5.5 (5.5); DSL: 4.6–5.6 (5.6); CL: 1.6–2.2 (2.2). FIVL: 11.2–15.0 (12.4). ChL: 2.8–5.2 (5.2). Coloration: Brownish red DS. Chelicerae, pedipalpus and legs brown. Dorsum: (Fig.
Similar to Tschaidicancha chaplini sp. nov. by the set of the following characteristics: reddish brown coloration; femur IV with a retrolateral row with more than 30 small tubercles; tibia IV with a retrolateral row with more than ten small tubercles (Fig.
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.
Type material: Holotype ♂, ‘PERU, Pasco, Oxapampa, Parque Nacional Yanachaga-Chemillén, 10°32′42.1″S 75°21′24.4″W, 22/IV/2011, R. Pinto-da-Rocha, A. Benedetti, J. Ochoa & D. Silva leg. (MUBI) – Paratypes 1 ♂, 1 ♀, ‘ditto’ (
Tschaidicancha weyrauchi
Roewer, 1957: 81 (desc.), fig. 11 (female dorsal habitus), 12 (female pedipalpus), 13 (female apical femur–patella IV);
MALE: Measurements (n=2) DSW: 3.7–4.0; DSL: 3.7; CL: 1.5–1.6. FIVL: 6.8–7.7. ChL: 2.7. Coloration: (Fig.
It differs from other species of the genus by the following set of characteristics: kappa-type DSS; ocularium with a pair of long spines; scutal area I undivided; free tergite III with a pair of large spiniform tubercles (Fig.
(Fig.
Type material: Holotype ♂, ‘PERU, Huánuco, Tschaidicancha, near Huánuco, 2,800 m a.s.l., 2/VIII/1955, Weyrauch leg. (
We chose parsimony analysis to obtain a working phylogenetic hypothesis of Metasarcidae because it minimizes the number of needed ad hoc hypotheses of character transformations, thereby maximizing the explanatory power of phylogenetic hypotheses (
Besides using TE under parsimony, we decided to test how the molecular data (most of the data in the total evidence matrix) behaved in an analysis of phylogenetic inference under Maximum Likelihood, a statistical optimality criterion, as well as parsimony. The objective was to observe whether a highly conflicting hypothesis would be recovered, in relation to the DO analysis. Furthermore, a TE dataset was also submitted to the maximum likelihood criterion. We are aware that, since both analyses differ in terms of optimality criterion (and consequently different inference mode), data set (TE versus molecular data only) and treatment of molecular data (static and dynamic homology), it is necessary to interpret the comparisons with caution (the differences in topologies resulting from all analyses are reported in section 3.3 and below in the discussion; additionaly see
Metasarcidae was recovered as a monophyletic group in previous analyses and as the sister group of Cosmetidae (see
The family Metasarcidae includes six genera and 38 species, of which two new genera and 20 new species were described here. The high proportion of newly described taxa reflects the fact that South American harvestmen are poorly known and suggests that we need more diversity studies to uncover the true species richness of this family. Arachnologists have been working at Argentinean and Brazilian museums since last century and have been responsible for most of what we know about the diversity and distribution of the Opiliones in the Brazilian Atlantic Rain Forest (Pinto-da-Rocha et al. 1995), Argentina (
Our phylogenetic analyses based on molecular and morphological markers resulted in a new classification with several new combinations proposed for Metasarcidae. The only genus that remained unchanged was Incasarcus Kury and Maury, 1998. The drastic reorganization within Metasarcidae is consistent with what has happened to other Neotropical harvestmen that were revised more recently, continuing to prove that the Roewerian system is flawed (e.g.
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 Metasarcidae that reflects its evolutionary history. The role of α-taxonomy, in this case, is fundamental to expand the knowledge of the biodiversity of the target group (see Martens and Segers 2005;
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 Metasarcidae established within the context of the Roewerian system (see Introduction), the strategy mentioned in the previous paragraph was very useful for the new supraspecific classification of Metasarcidae (see section 2.5).
Of the six genera of Metasarcidae recognized in this study, Huancabamba gen. nov. (monotypic) and Lumieria gen. nov. (two species) are described for the first time. The establishment of these two genera resulted from the phylogenetic analysis, since both were recovered in a clade at the base of the tree chosen to represent the phylogeny of the family (MP01; and also in MP02 and ML01). Only in ML02 the two genera do not form a clade, being the two most basal lineages of Metasarcidae. The choice to describe two distinct genera, even if they are sister groups in the working hypothesis, is due to the evident morphological differences (see below), especially those related to the stylus’ format and the number of MS C of the penis. As Huancabamba gen. nov. has only one species known, the autopomorphies of H. kubricki gen. et sp. nov. mentioned in section 3.3 are the putative morphological synapomorphies of the genus. Furthermore, an important diagnostic feature, penial VP with more than 13 MS C, may be another putative synapomorphy of this monotypic genus, since no other Metasarcidae has that many MS C. Additionally, very short VP lateral sacs is another important diagnostic feature, even though it is plesiomorphic and homoplastic (it occurs in a few other species of the family, e.g. Metasarcus fellinii sp. nov., and also in the cosmetid Metalibitia paraguayensis).
Lumieria gen. nov. was represented in the analysis by one of the two described species. The autapomorphies of L. antonionii gen. et sp. nov. were listed in section 3.3. The first autapomorphy listed [20:2] is not found in the other species of the genus (Lumieria woodyalleni gen. et sp. nov.), but the second [64:2], related to the penial stylus is shared by both. The broad and sturdy penis stylus shape of Lumieria gen. nov. species are completely different from all other Metasarcidae. Therefore, the character state of the penis stylus can be understood as an important morphological putative synapomorphy of Lumieria gen. nov. Furthermore, the homoplastic autapomorphy of L. antonionii gen. et sp. nov., retrolateral row of acuminate tubercles on tibia IV [52:1], also present in L. woodyalleni gen. et sp. nov., is not included in the analysis due a lack of fresh samples and may be another putative synapomorphy of the genus. These characteristics mentioned allow the classification of L. woodyalleni gen. et sp. nov. in this genus.
Incasarcus was well represented in the analysis with four of its five known species (I. pictus is not represented). This is the first time that the monophyly of Incasarcus was tested since
Before this study, Metasarcus included three species: M. bolivianus Roewer, 1913 and M. armatipalpus (Roewer, 1929) from Bolivian Chaco, which were synonymized here (see section 3.42) and M. clavifemur from department of La Paz, Bolivia. The type-species of Metasarcus is known only from a female holotype (1913) (plus the female holotype of M. armatipalpus, here established as its junior synonym) and consequently the species was not included in our phylogenetic analysis. Both holotypes are extremely similar, which justified the synonymization of both species. In addition, due to the presence of a spiniform apophysis on free tergite III, these females are very similar to females of M. fellinii sp. nov., another species from the Bolivian Chaco, collected near Entre Ríos, Department of Tarija. The similarity of the females indicates that they are closely related species, which supports the allocation of M. fellinii sp. nov. within this genus. Despite their similarity, they are clearly different species, due to the large difference in size and in the density of the granulation throughout the dorsal scutum. Metasarcus clavifemur was included in the genus without any justification and without a discussion of Metasarcus after the inclusion of the species (
Of the three new species not included in the phylogenetic analyses, M. limachii sp. nov. is most similar to the species of the clade (M. trispinosus sp. nov. (M. bergmani sp. nov. + M. fellinii sp. nov.)) due to the presence of the spiniform apophysis in the free tergite III of male and female. Metasarcus kurosawai sp. nov. resembles M. clavifemur in their DSS and scutal area I shape.
The internal relationships of Incasarcus and Metasarcus are identical in all analyses, as well as the relationship of the two genera as sister groups.
We were not able to test the monophyly of Tschaidicancha. The genus was monotypic prior to this work and the type-species, Tschaidicancha weyrauchi, is known only from a few specimens. It was not possible to extract its DNA due to tissue old age. Additionally, we were not able to include two of the three new species in our analysis. Among the autapomorphies of T. chaplini sp. nov. listed in section 3.3, only one is shared by the other species of the genus: scutal area II with a pair of tubercles. This characteristic is also present in Ayacucho querococha sp. nov. + Ayacucho tapacocha nom. nov. clade among the Metasarcidae. Amidst the four species of the genus (sensu hoc), T. chaplini sp. nov. and T. scorsesei sp. nov. are very similar morphologically, especially in the DSS and armature pattern of femur IV (see diagnosis in sections 3.50 and 3.52). Tschaidicancha joseochoai sp. nov. resembles T. weyrauchi sp. nov. in having a more robust armature on femur IV of male (although the armature pattern is totally different), but it has a series of characteristics that are distinct from the other species of the genus (see diagnosis in section 3.51). The four Tschaidicancha species are quite distinct from most Ayacucho species (granulation; legs/DS ratio; presence of a proapical spine on pedipalpus femur in males, etc). The Tschaidicancha species superficially resembles the Incasarcus, Metasarcus, Lumieria species more than the Ayacucho species. When using morphological data in the analyses (MP01; ML01), T. chaplini sp. nov. is recovered as a sister group of the Ayacucho, which does not occur in analyses with only molecular data, in which it is retrieved nested in Ayacucho (MP02; ML02). This demonstrates that, morphologically, there is a distinction between this species and the Ayacucho (even taking into account that A. spielbergi sp. nov. also differs morphologically from the other Ayacucho, see below). Even considering that, the monophyly of the genus has not been tested and none of the diagnostic characteristics of the genus (see section 3.49) are exclusive, we have chosen to tentatively allocate the three new species in Tschaidicancha. We made this decision while keeping in mind that we wanted to avoid describing new monotypic genera that did not result from a phylogenetic analysis.
Prior to this revision, Ayacucho contained only its type-species from Ayacucho, south-central Peru. We expanded the delimitation of the genus and included the northern Peruvian genera Cajamarca (4 spp.), Cargaruaya (1 spp.), Cajacaybia (1 spp.) and Tapacochana (1 spp.); Central Peruvian genus Palcares (2 spp.); and seven new species. The resulting clade consists of A. titschacki (type-species), A. uniseriatus comb. nov. (previously in Cajamarca), A. spiniger comb. nov. (previously in Palcares), A. tapacocha nom. nov. (previously in Tapacochana), and six new species (12 terminals representing 10 species; 17 species recognized in this study). The type-species of the monotypic genera Cargaruaya and Cajacaybia are known only from the female holotypes described in 1956 and 1957 (respectively) and were not included in the analysis. Tapacochana and Palcares are represented in the ingroup by their type-species, and the type-species of Cajamarca is known only from the male holotype (1952) and is not present in the analysis.
The internal relationships of Ayacucho differ between the different analyses. The two inner clades. “silva clade” and “spinigera clade”, are recovered as sister groups only in MP01. The “silva clade” is monophyletic in all four analyses, but the “spinigera clade” is not (only in MP01).
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 Ayacucho used in the analysis, except in Ayacucho spielbergi sp. nov., which has a subcylindrical and not flattened pedipalpus femur [8:0], similar to all remaining metasarcids. The absence of the proapical spine on pedipalpus femur in males [7:0] is an important homoplastic synapomorphy. This character state is also found in representatives of the outgroup (e.g. Napostygnus, Phareicranaus [Cranaidae], Cosmetidae genera); all the remaining metasarcids have a proapical spine on pedipalpus femur in males [7:1]. The alpha-type DSS [10:0] is another important homoplastic synapomorphy (also found in Incasarcus). Only Ayacucho spielbergi sp. nov. has a different shape (gamma-type). The presence of a row of tubercles on free tergites II–III [30:1] only occurs in I. argenteus and in some representatives of the external group. An exclusive synapomorphy under ACCTRAN optimization, femur IV with a distal row of acuminate tubercles [45:1:] is present only in Ayacucho silvae sp. nov., Ayacucho vargasllosai sp. nov. and Ayacucho pomacocha sp. nov. Despite the low support (GB = 7), and keeping in mind the morphological differences listed above (e.g. presence of proapical spine on pedipalpus femur in males, dense granulation on DS, length of the femur IV; see also diagnosis in section 3.10) that separate Ayacucho from other metasarcids genera, we opted for a broader definition of the genus. Ayacucho bambamarca comb. nov., A. triarmatus nom. nov. and A. weyrauchi comb. nov., absent from the phylogenetic analysis, were described together with A. uniseriatus comb. nov. within the genus Cajamarca (
It is important to note that of Ayacucho pasolinii sp. nov., which was not included in the phylogenetic analysis, and the previously discussed A. spielbergi sp. nov., share a series of morphological features that distinguish them from the other species of the genus: ocularium saddle-shaped (although depression is shallow), low granulation on DS and femur IV long (FIV/DSL = 1.5). The length of the legs (represented by the FIVL/DSL ratio in this study) is especially important to differentiate Ayacucho from other genera and for this reason the condition found in A. spielbergi sp. nov. and A. pasolinii sp. nov. is especially striking. The DSS of A. pasolinii sp. nov. (with not so rounded mid-bulge) is similar to the condition found in the “Ayacucho silvae sp. nov. clade”. Although this can indicate proximity, there are important differences such as the presence of a pair of strong spines on area III, ocularium with large spines and male femur IV with a more robust armature. Additionally, the yellowish spots on the lateral margins of the DS are unique among Ayacucho species. The working phylogenetic hypothesis of this study supports the inclusion of A. spielbergi sp. nov. in Ayacucho, despite the morphological differences, but the same does not occur with A. pasolinii sp. nov., which is not included in the analysis due to lack of fresh tissues. Considering that A. spielbergi sp. nov. has a high number of autapomorphies, we have hypothesized that the same could occur with another species. In addition, there is no indication, based on morphology, that the species belong in the other metasarcid genera. Therefore, we opted to include A. pasolinii sp. nov. in Ayacucho because we consider that the morphological characteristics of the species fit this genus
Finally, it is worth mentioning that the “silvae clade” is especially noteworthy for presenting an extremely conservative external morphology. Males are very similar (with subtle differences) and females are virtually indistinguishable (see diagnoses of A. glauberrochai sp. nov., A. pomacocha sp. nov., A. silvae sp. nov., A. titschacki and A. vargasllosai sp. nov.). The differentiation among these species strongly relies on the penial morphology.
We present a strongly supported, comprehensive phylogenetic hypothesis of Metasarcidae based on morphological and molecular datasets. The family is monophyletic and was recovered as the sister-group of Cosmetidae. Additionally, this contribution increases our knowledge of the taxonomy of the group. The taxonomic acts that derived from this systematic review of the family are compiled below.
The following generic synonymies are proposed: Ayacucho Roewer, 1949 = Cajamarca Roewer, 1952, Cargaruaya Roewer, 1956, Palcares Roewer, 1957, Cajacaybia Roewer, 1957 and Tapacochana Roewer, 1957.
The following new genera are described from Peru: Huancabamba gen. nov. (type species Huancabamba kubricki gen. et sp. nov.) and Lumieria gen. nov. (type species Lumieria antonionii gen. et sp. nov.).
The following specific synonymies are proposed: Cajamarca weyrauchi Roewer, 1952 = Cajamarca affinis Roewer, 1957; Cajamarca bambamarca Roewer, 1957 = Cajamarca triseriata Roewer, 1957; Metasarcus bolivianus Roewer, 1913 = Chaconatus armatipalpus Roewer, 1929; Palcares spiniger Roewer, 1957 = Palcares serrifemur Roewer, 1959.
The following new combinations are proposed: Ayacucho bambamarca (Roewer, 1957) comb. nov.; Ayacucho inermis (Roewer, 1957) comb. nov.; Ayacucho insignitus (Roewer, 1956) comb. nov.; Ayacucho spiniger (Roewer, 1957) comb. nov.; Ayacucho uniseriatus (Roewer, 1959) comb. nov. and Ayacucho weyrauchi (Roewer, 1952) comb. nov.
The following new species are described from Bolivia: Metasarcus beni sp. nov., Metasarcus bergmani sp. nov., Metasarcus fellinii sp. nov., Metasarcus kurosawai sp. nov., Metasarcus limachii sp. nov., Metasarcus trispinosus sp. nov. and Metasarcus vacafloresae sp. nov. The following new species are described from Peru: Ayacucho glauberrochai sp. nov.; Ayacucho pasolinii sp. nov.; Ayacucho pomacocha sp. nov., Ayacucho querococha sp. nov., Ayacucho silvae sp. nov., Ayacucho spielbergi sp. nov., Ayacucho vargasllosai sp. nov.; Huancabamba kubricki gen. et sp. nov., Lumieria antonionii gen. et sp. nov., Lumieria woodyalleni gen. et sp. nov., Tschaidicancha chaplini sp. nov., Tschaidicancha joseochoai sp. nov. and Tschaidicancha scorsesei sp. nov.
The following secondary homonym species names are replaced: Cajacaybia spinigera Roewer, 1957 replaced by Ayacucho roeweri nom. nov.; Tapacochana insignita Roewer, 1957 replaced by Ayacucho tapacocha nom. nov. and Tapacochana triseriata Roewer, 1959 replaced by Ayacucho triarmatus nom. nov.
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 (