Corresponding author: Casey H. Richart (
Academic editor: Adriano Kury
In
The genus
The biogeographic situation in
The goal of this paper is to use morphometrics and molecular phylogenetics to investigate the validity of the four species hypothesized by
Most fieldwork was conducted in the summer of 2008, with additional adult specimens collected from May to September in 2006, 2007, and 2009 (Appendix I - Collection Locality Information).
Genomic DNA was extracted from two legs per specimen using the Qiagen DNeasy kit, per manufacturer’s protocol. Currently, few genes are available for resolving shallow phylogenetic relationships in Opiliones (reviewed in
Bi-directional Sanger reads were assembled into contiguous sequences using Sequencher v4.5 (Gene Codes Corporation, MI). EF-1α haplotypes were reconstructed in PHASE (
Individual gene trees were reconstructed using maximum likelihood and Bayesian inference. Bayesian analyses were implemented using MrBayes v3.1.2 (
All available DNA sequences, with the exception of apparently nuclearized copies of COI (see Results), were concatenated for phylogeny reconstruction. The concatenated matrix was analyzed using both maximum likelihood (RAxML) and Bayesian approaches (MrBayes v3.1.2) using a seven-partition strategy (EF-1α intron + exon, Wnt2, 28S, individual COI codon positions). The Bayesian analysis was run for 1 X 107 generations; tree sampling and burn-in were as above. Outgroup sequences for five of six ischyropsalidoid genera (
Specimen measurements were taken using an Olympus SZX12 dissecting microscope with an ocular micrometer. Individuals with missing data (i.e., no Leg II) were excluded from analyses. Standard measurements (
Principle components analyses (PCA) and discriminant function analyses (DFA) were carried out using SYSTAT 12 (Systat Software, Inc.). Though both PCA and DFA are multivariate analyses, DFA is a validation approach that deals specifically with the problem of separating predetermined groups. In taxonomy, DFAs are almost exclusively used to differentiate between morphologically similar species that are difficult to identify from single characteristics (see
In taxonomy, PCAs are regularly employed for species delimitation. This utilization has ranged from comparing the vector angles in a scatter plot of predefined species groups (
Due to sexual dimorphism, male and female specimens were analyzed separately for all morphometric analyses. Since the data used to conduct PCAs were measured in the same units (mm), analyses were conducted on both correlation and covariance matrices. PCAs using covariance matrices tend to be dominated by characters showing the most variability (
We initiated analyses with seven
North of the Salmon River in Idaho,
A cybertaxonomic approach was undertaken for enhanced dissemination of this work (e.g.,
The data underpinning the analyses reported in this paper are deposited in the Dryad Data Repository at doi:
Fieldwork resulted in the collection of 272
Distribution of
Although the overall geographic distribution of
Mitochondrial COI DNA sequences for a geographical subset of
Sequences showing evidence for nuclearization were discarded, resulting in the removal of all sequences for individuals collected north of the Middle Fork Clearwater River, including all
Phylogenetic taxon sample and GenBank accession numbers.<br/>
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Acfquattuor_OP2230_DeVoto |
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Acfquattuor_OP2275_SplitCkTr |
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Acfquattuor_OP2284_SelwayRvRd |
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Acosmetoides_OP2281_2ShadowsCk |
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Acosmetoides_OP2296_FS250 |
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Acosmetoides_OP2299_TribOrogrande |
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Acosmetoides_OP2319_MeadowCk |
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Acosmetoides_OP2341_GooseCk |
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Aleonardi_OP2347_IronCk |
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Aleonardi_OP2349_NFGobleCk |
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Aleonardi_OP2712_KjesbuRd |
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Aleonardi_OP2714_UpperIronCk |
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Americkeli_OP2237_FS443 |
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Americkeli_OP2245_FS443 |
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Americkeli_OP2250_RedHorseCk |
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Amakah_OP1699_RubyBeach |
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Amakah_OP2345_CedarCk |
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Amakah_OP2715_HokoFalls |
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Amakah_OP2716_BrownesCk |
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Amakah_OP2719_YahooLkRd |
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Aquattuor_OP2242_GrouseCk |
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Aquattuor_OP2257_SlateCk |
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Aquattuor_OP2270_FS221 |
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Asheari_OP2708_BurgdorfRd |
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Asheari_OP2709_BurgdorfRd |
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Asheari_OP2720_FS592 |
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Ashoshone_OP2316_EmeraldCkRd |
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Ashoshone_OP2323_Hobo |
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Ceratolasma |
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Dendrolasma |
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Hesperonemastoma |
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Ischyropsalis |
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Ortholasma |
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Sabacon |
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Taracus |
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Gblocks removal of ambiguous sites in the 28S matrix resulted in a reduction of alignment length from 1211 to 1173 positions. The aligned matrix included 26 parsimony informative characters for 14 ingroup sequences. For the EF-1α gene, eight of 22
Models of evolution inferred from jModelTest are summarized in
Models of DNA sequence evolution as determined by jModelTest v0.1.1 <br/>
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COI 1st partition | GTR+I+G |
COI 2nd partition | GTR+I+G |
COI 3rd partition | GTR+G |
28S | GTR+I+G |
EF-1α intron | GTR+G |
EF-1α exon | SYM+G |
Wnt2 | SYM+I |
Bayesian phylogram resulting from analysis of concatenated dataset. Numbers at nodes correspond to Bayesian posterior probabilities. The outgroup topology is shown at bottom. The inset picture is a female
Within
All Bayesian individual gene trees had their longest branch separating Washington and Idaho genetic groups (
Bayesian gene trees with posterior probabilities:
DFA and PCA morphometric analyses of male and female data sets most frequently recovered
Since the group frequencies of hypothesized species varied greatly (
Sample sizes for PCA and DFA analyses.<br/>
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10 | 14 | 24 |
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6 | 4 | 10 |
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4 | 3 | 7 |
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14 | 17 | 28 |
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19 | 19 | 38 |
22 | 18 | 40 | |
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56 | 58 | 114 |
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131 | 133 | 261 |
The number of individuals from each hypothesized species used in morphometric analyses.
Groupings based on female specimens were not recovered as frequently as male-based groups. Females of
Principle components analyses regularly recovered all hypothesized species of
PCA bi-dimensional plot. PCA results of male data using a covariance matrix; plot of principle components 2 and 3.
Both phylogenetic and morphometric analyses strongly support three new species:
Though they lack molecular support,
The Western Hemlock Zone (WHZ) (
Reconstructed molecular phylogenies show that
The primary defense of
Recent surveys of litter dwelling animals in mesic forests of the Pacific Northwest have led to the discovery of many new species. Taxa include terrestrial gastropods (
The genus
The simple, distally tapering penis (Appendix VIII, Figure I) and short ovipositor (Appendix III,
This dichotomous key should allow users to identify the new species described herein. However, discovery of new morphologies in northern Idaho not described by
1a | Setose mounds dorsally on surface of cheliceral article I; males ( |
2 |
1b | Without raised mounds on chelicerae; females | 8 |
2a (1a) | Paramedian tubercles acute spines on scute area II only ( |
3 |
2b | Paramedian tubercles acute spines on multiple scute areas ( |
6 |
3a (2a) | Ocularium height (ventral edge of eye to tip of ocularium) ≤ 0.60 mm; area II spine height ≤ 0.50 mm from surface of tergite | |
3b | Ocularium and area II spine heights ≥ 0.80 mm | 4 |
4a (3b) | Distal ends of leg femora, patellae, and tibiae distinctly light, contrasting as light joints; palpi white or nearly so; with or without false leg articulations on leg II metatarsi (Appendix IX); with or without distal, dark, prolateral tubercle on palpal patellae; western Washington | 5 |
4b | Distal ends of leg segments not light; palpi light brown or dark; without false leg articulations and palpal tubercles; Idaho | |
5a (4a) | Leg II femur ≤ 3.76 mm; Cascade Mountains | |
5b | Leg II femur ≥ 3.76 mm; Olympic Peninsula | |
6a (2b) | Paramedian tubercles raised into acute spines on scute areas I and II only ( |
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6b | Paramedian tubercles with acute spines on three or more scute areas; scute area III always with such tubercles (Appendix V, |
7 |
7a (6b) | Paramedian tubercles not raised into spines on scute area I, paired spines on areas II, III, and IV | |
7b | Paramedian tubercles raised into acute spines on scute areas I – IV (four pairs of abdominal spines | |
8a (1b) | Paramedian tubercles acute spines on scute area II only ( |
9 |
8b | Tubercles not as previous; if spines on area II only, greatly reduced | 11 |
9a (8a) | As couplet 4a | 10 |
9b | As couplet 4b |
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10a (9a) | Known only from the Cascade Mountains of Washington State; no perceived discriminating features | |
10b | Known only from the Olympic Peninsula of Washington State; no perceived discriminating features | |
11a (8b) | As couplet 6a | |
11b | Paramedian tubercles not as previous | 12 |
12a (11a) | As couplet 7a | |
12b | Paramedian tubercles not enlarged into spines, but paired, low, rounded tubercles on all scutal areas ( |
13 |
13a (11b) | Palpal femur ≤ 0.88 mm; leg II tarsus ≤ 3.92 mm; leg II femur ≤ 2.75 mm; currently known south of Salmon River | |
13b | Palpal femur ≥ 0.90 mm; leg II tarsus ≥ 3.95 mm; leg II femur ≥ 2.72 mm, north of Middle Fork Clearwater River |
Male
The specific epithet is a patronym in honor of the naturalist and careful observer William P. Leonard for his work on litter-dwelling organisms in the poor-person’s rainforest of the Pacific Northwest.
Distinguished from all
Eye tubercle at anterior edge of carapace prolonged into an acute spine lacking hemispherical warts on distal half; standing 1.25 mm above the surface of the carapace (
All scutal tergites with pairs of median tubercles, these prolonged into large spines on area II only; lateral tubercles distinct. Tergite I with paired median tubercles as raised mounds adorned with warts standing 0.09 mm (
Abdominal sternite warty sculpturing strongest laterally and on posterior margins; sternites brown. Sclerotized areas of genital operculum relatively setose. Prosomal sternum (
Chelicerae brown or light brown (
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Palpus | trochanter | 0.55 | 0.52–0.58 | 3 | 0.61 | 0.58–0.65 | 3 |
femur | 0.94 | 0.84–1.00 | 7 | 0.94 | 0.90–0.98 | 5 | |
patella | 0.67 | 0.66–0.68 | 3 | 0.71 | 0.70–0.73 | 3 | |
tibia | 0.74 | 0.73–0.74 | 3 | 0.73 | 0.67–0.80 | 3 | |
tarsus | 0.69 | 0.67–0.72 | 3 | 0.66 | 0.64–0.68 | 3 | |
Leg I | trochanter | 0.49 | 0.48–0.52 | 3 | 0.50 | 0.45–0.56 | 3 |
femur | 2.37 | 2.28–2.44 | 3 | 2.25 | 1.96–2.60 | 3 | |
patella | 0.88 | 0.88 | 3 | 0.87 | 0.84–0.92 | 3 | |
tibia | 1.60 | 1.48–1.72 | 3 | 1.46 | 1.44–1.50 | 3 | |
metatarsus | 2.17 | 1.84–2.36 | 3 | 2.19 | 2.04–2.28 | 3 | |
tarsus | 2.89 | 2.72–3.08 | 3 | 2.80 | 2.68–2.88 | 3 | |
Leg II | trochanter | 0.57 | 0.52–0.60 | 7 | 0.55 | 0.52–0.56 | 5 |
femur | 3.53 | 3.32–3.76 | 7 | 3.46 | 2.88–3.92 | 5 | |
patella | 1.02 | 0.96–1.08 | 7 | 1.00 | 0.92–1.08 | 5 | |
tibia | 2.38 | 2.16–2.60 | 7 | 2.36 | 2.12–2.60 | 5 | |
metatarsus | 3.70 | 3.44–4.05 | 7 | 3.69 | 3.28–3.96 | 5 | |
tarsus | 4.93 | 4.74–5.19 | 7 | 4.58 | 4.40–4.70 | 5 | |
Leg III | trochanter | 0.48 | 0.48 | 3 | 0.54 | 0.52–0.60 | 3 |
femur | 2.23 | 2.12–2.28 | 3 | 2.10 | 1.88–2.25 | 3 | |
patella | 0.85 | 0.80–0.88 | 3 | 0.84 | 0.80–0.88 | 3 | |
tibia | 1.57 | 1.56–1.60 | 3 | 1.50 | 1.48–1.52 | 3 | |
metatarsus | 2.49 | 2.40–2.56 | 3 | 2.38 | 2.24–2.45 | 3 | |
tarsus | 3.13 | 3.08–3.20 | 3 | 2.95 | 2.80–3.05 | 3 | |
Leg IV | trochanter | 0.57 | 0.52–0.64 | 3 | 0.58 | 0.55–0.64 | 3 |
femur | 3.23 | 3.08–3.32 | 3 | 3.09 | 2.76–3.35 | 3 | |
patella | 1.03 | 0.96–1.08 | 3 | 0.99 | 0.96–1.00 | 3 | |
tibia | 2.01 | 1.96–2.08 | 3 | 1.99 | 1.96–2.05 | 3 | |
metatarsus | 4.03 | 3.88–4.12 | 3 | 3.85 | 3.64–4.00 | 3 | |
tarsus | 4.09 | 4.08–4.12 | 3 | 3.87 | 3.76–4.99 | 3 |
All measurements in millimeters;
Leg measurements given in
Penis length 2.48 mm (
Eye tubercle height above surface of carapace 1.18 mm (
Paramedian tubercles or tergite I height 0.08 mm (
Sternites brown. Transverse furrow and membranous lateral sutures of genital operculum less distinct than in other species. Prosomal sternum (
Horn-shaped process of epistome decurved, projecting 0.42 mm from sulcus. Chelicerae brown or light brown; article I length 1.18 mm (
Leg measurements given in
Ovipositor length 0.80 mm, width 0.44 mm; corona of setae at furcal base surrounding lobes, apical setae on lobes; furca without dorsoventral differentiation.
Male
The specific epithet refers to the Makah Nation, which historically occupied much of the known distribution of the species. The name Makah was given to these people by their neighbors; it means “generous with food”. These people have shared with many people access to their beautiful land, next to the rocks and gulls. For more information on the Makah Nation see:
Distinguished from all
Eye tubercle at anterior edge of carapace, prolonged anteriad into a sharp conical spine 1.27 mm (
All scutal areas with pairs of paramedian tubercles. Area I paramedian tubercles cluster of cuticular warts standing 0.04 mm above the surrounding scute (
Abdominal sternites with infrequent setae; warty sculpturing strongest on posterior and lateral margins; brown to dark brown. Prosomal sternum length 0.14 mm (
Acuclavella makah palpus and leg measurements.<br/>
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Palpus | trochanter | 0.58 | 0.54-0.62 | 3 | 0.57 | 0.54-0.60 | 3 |
femur | 0.95 | 0.90-0.98 | 10 | 0.98 | 0.88-1.05 | 14 | |
patella | 0.67 | 0.62-0.72 | 3 | 0.70 | 0.67-0.74 | 3 | |
tibia | 0.79 | 0.73-0.84 | 3 | 0.72 | 0.68-0.76 | 3 | |
tarsus | 0.73 | 0.71-0.76 | 3 | 0.68 | 0.67-0.68 | 3 | |
Leg I | trochanter | 0.57 | 0.56-0.60 | 3 | 0.53 | 0.50-0.56 | 3 |
femur | 2.67 | 2.60-2.72 | 3 | 2.46 | 2.36-2.56 | 3 | |
patella | 0.97 | 0.92-1.00 | 3 | 0.90 | 0.88-0.92 | 3 | |
tibia | 1.75 | 1.72-1.76 | 3 | 1.57 | 1.55-1.60 | 3 | |
metatarsus | 2.36 | 2.24-2.44 | 3 | 2.13 | 2.08-2.20 | 3 | |
tarsus | 2.93 | 2.76-3.04 | 3 | 2.72 | 2.56-2.80 | 3 | |
Leg II | trochanter | 0.60 | 0.56-0.64 | 10 | 0.61 | 0.56-0.64 | 14 |
femur | 4.01 | 3.76-4.35 | 10 | 3.85 | 3.60-4.20 | 14 | |
patella | 1.12 | 1.00-1.20 | 10 | 1.14 | 1.04-1.28 | 14 | |
tibia | 2.65 | 2.40-2.88 | 10 | 2.64 | 2.40-2.92 | 14 | |
metatarsus | 3.91 | 3.60-4.25 | 10 | 3.80 | 3.52-4.75 | 14 | |
tarsus | 5.11 | 4.55-5.50 | 10 | 4.62 | 4.25-4.94 | 14 | |
Leg III | trochanter | 0.55 | 0.52-0.56 | 3 | 0.57 | 0.55-0.60 | 3 |
femur | 2.47 | 2.32-2.56 | 3 | 2.21 | 2.12-2.32 | 3 | |
patella | 0.93 | 0.92-0.96 | 3 | 0.96 | 0.88-1.05 | 3 | |
tibia | 1.64 | 1.60-1.68 | 3 | 1.60 | 1.52-1.65 | 3 | |
metatarsus | 2.64 | 2.56-2.76 | 3 | 2.48 | 2.40-2.64 | 3 | |
tarsus | 3.16 | 3.00-3.32 | 3 | 2.95 | 2.85-3.12 | 3 | |
Leg IV | trochanter | 0.67 | 0.60-0.72 | 3 | 0.65 | 0.60-0.72 | 3 |
femur | 3.64 | 3.52-3.80 | 3 | 3.34 | 3.16-3.60 | 3 | |
patella | 1.05 | 0.96-1.12 | 3 | 1.11 | 1.04-1.20 | 3 | |
tibia | 2.32 | 2.32 | 3 | 2.30 | 2.16-2.44 | 3 | |
metatarsus | 4.16 | 4.05-4.30 | 3 | 3.99 | 3.76-4.25 | 3 | |
tarsus | 4.17 | 3.85-4.35 | 3 | 3.86 | 3.68-3.95 | 3 |
All measurements in millimeters;
Leg measurements given in
Penis 2.39 mm in length (
Eye tubercle height above surface of carapace 1.37 mm (
Paramedian tubercles of tergite I height 0.06 mm (
Sternites brown. Prosomal sternum length 0.17 mm (
Chelicerae article I length 1.35 mm (
Leg measurements given in
Ovipositor length 0.74 mm (
Known from the northwest areas of the Olympic Peninsula in Clallam and Jefferson Counties, Washington State (Appendix I). Found in coniferous or riparian forests along small, perennial water-features such as headwater streams, springs, and seeps; underneath woody debris and moss.
Male
The specific epithet honors Dr. William A. Shear, eminent milliped and opilionid taxonomist. His influence has been important to the authors’ aspirations to be systematic biologists, and we thank him sincerely, and with pleasure.
Generally reduced dimensions overall. Males with one pair of scutal spines on area II only; scutal spines ≤ 0.50 mm, distance from ventral edge of eye to tip of ocularium ≤ 0.60 mm distinguishes it from other males with single pair of spines. Females lack scutal spines. Diagnosed from spine-less
Eye tubercle at anterior edge of carapace, hemispherical warts cover entirety of ocular spine, prolonged into an acute spine standing 0.67 mm above the surface of the carapace (
All scutal tergites with pairs of paramedian tubercles; relatively short, pointed spines on area II only. Fused tergite I with paramedian tubercles as raised mounds, relatively tall but not spike-like, standing 0.125 mm above the surrounding scute (
Abdominal sternites with infrequent setae; warty sculpturing strongest laterally and on posterior margin; sternites brown, dark brown, or black. Sclerotized areas of genital operculum relatively setose. Prosomal sternum (
Chelicerae light brown, dark brown, or black; lighter individuals with article I darker dorsally, article II with darker, more sclerotized striations on prolateral and retrolateral surfaces (
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Palpus | trochanter | 0.51 | 0.49–0.54 | 3 | 0.50 | 0.43–0.54 | 3 |
Femur | 0.85 | 0.78–0.88 | 4 | 0.85 | 0.83–0.88 | 4 | |
Patella | 0.63 | 0.62–0.65 | 3 | 0.66 | 0.60–0.70 | 3 | |
Tibia | 0.74 | 0.72–0.75 | 3 | 0.73 | 0.65–0.79 | 3 | |
Tarsus | 0.70 | 0.67–0.71 | 3 | 0.65 | 0.59–0.68 | 3 | |
Leg I | trochanter | 0.53 | 0.48–0.56 | 3 | 0.45 | 0.40–0.50 | 3 |
Femur | 1.80 | 1.76–1.84 | 3 | 1.73 | 1.48–1.94 | 3 | |
Patella | 0.87 | 0.84–0.92 | 3 | 0.81 | 0.68–0.91 | 3 | |
Tibia | 1.33 | 1.28–1.44 | 3 | 1.30 | 1.16–1.40 | 3 | |
metatarsus | 1.97 | 1.96–2.00 | 3 | 1.88 | 1.68–2.09 | 3 | |
Tarsus | 2.29 | 2.24–2.32 | 3 | 2.22 | 1.88–2.41 | 3 | |
Leg II | trochanter | 0.57 | 0.56–0.60 | 4 | 0.52 | 0.44–0.63 | 4 |
Femur | 2.53 | 2.48–2.64 | 4 | 2.52 | 2.16–2.75 | 3 | |
Patella | 0.98 | 0.88–1.12 | 4 | 0.93 | 0.76–1.03 | 3 | |
Tibia | 1.96 | 1.88–2.08 | 4 | 1.90 | 1.64–2.09 | 3 | |
metatarsus | 2.99 | 2.88–3.12 | 4 | 3.00 | 2.56–3.24 | 3 | |
Tarsus | 3.67 | 3.64–3.72 | 4 | 3.65 | 3.20–3.92 | 3 | |
Leg III | trochanter | 0.53 | 0.52–0.56 | 3 | 0.52 | 0.44–0.59 | 3 |
Femur | 1.72 | 1.64–1.80 | 3 | 1.71 | 1.56–1.84 | 3 | |
Patella | 0.85 | 0.84–0.88 | 3 | 0.79 | 0.48–0.97 | 3 | |
Tibia | 1.33 | 1.20–1.48 | 3 | 1.43 | 1.32–1.56 | 3 | |
metatarsus | 2.21 | 2.16–2.28 | 3 | 2.18 | 1.84–2.38 | 3 | |
Tarsus | 2.56 | 2.52–2.64 | 3 | 2.53 | 2.20–2.84 | 3 | |
Leg IV | trochanter | 0.59 | 0.52–0.68 | 3 | 0.59 | 0.52–0.69 | 3 |
Femur | 2.41 | 2.32–2.56 | 3 | 2.38 | 2.12–2.63 | 3 | |
Patella | 0.96 | 0.88–1.04 | 3 | 0.89 | 0.80–0.94 | 3 | |
Tibia | 1.84 | 1.76–1.88 | 3 | 1.77 | 1.56–2.03 | 3 | |
metatarsus | 3.31 | 3.28–3.32 | 3 | 3.24 | 2.80–3.48 | 3 | |
Tarsus | 3.23 | 3.08–3.32 | 3 | 3.29 | 2.80–3.68 | 3 |
All measurements in millimeters;
Leg measurements given in
Penis length 2.94 mm (
Eye tubercle height above surface of carapace 0.67 mm (
Dorsal armature lacking. Dorsal adornment generally with median tubercles as raised mounds adorned with warts, decreasing in size away from midline and posteriorly across tergites. Two lateral pairs of warts on areas III and IV, area V with median pair of tubercles barely discernable, two pairs of lateral tubercles, or shallowly raised mounds adorned with warts. Free tergites with tubercles in the form of clusters of warts, or without discernable tubercles. Paramedian tubercles heights above the surface of the tergite (
Sternites brown, black, or light brown. Prosomal sternum (
Chelicerae article I length 0.98 mm (
Leg measurements given in
Ovipositor length 0.82 mm, width 0.46 mm (
Known from north-facing, horizontal band of
Males (AMNH; CAS, CASENT9039220; UWBM, ID0013/5661) from the type locality: Slate Creek Road 10.2 miles east of US 95, Nez Perce National Forest, Idaho County, Idaho.
Diagnosed from all
Eye tubercle at anterior edge of carapace, erect, spine-like, standing 0.98 mm above the surface of the carapace (
Scutum of opisthosoma rounded anteriorly, squared of posteriorly. All fused tergites with paramedian pair of tubercles, these in the form of erect spines on areas I and II. Spines of tergites I and II curve posteriad, stand 0.82 mm (
Abdominal sternite warty sculpturing strongest laterally and on posterior margin; sternites brown or dark brown. Genital operculum tongue-shaped, clearly delineated by transverse furrow, lateral ends of furrow membranous suture, distal margin rebordered, glossy. Prosomal sternum wider than long; length 0.18 mm (
Chelicerae light brown or brown; darker dorsally; article II with darker, more sclerotized striations on prolateral and retrolateral surfaces (
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Trochanter | Mean | 0.53 | 0.61 | 0.61 | 0.51 | 0.64 |
Range | 0.52–0.54 | 0.60–0.62 | 0.56–0.68 | 0.48–0.56 | 0.56–0.75 | |
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3 | 3 | 14 | 3 | 3 | |
Femur | Mean | 1.01 | 2.31 | 3.14 | 2.00 | 2.88 |
Range | 0.95–1.10 | 2.20–2.44 | 2.72–3.44 | 1.96–2.04 | 2.76–3.00 | |
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14 | 3 | 14 | 3 | 3 | |
Patella | Mean | 0.67 | 0.97 | 1.08 | 0.95 | 1.10 |
Range | 0.64–0.73 | 0.92–1.03 | 1.00–1.20 | 0.92–1.00 | 0.96–1.20 | |
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3 | 3 | 14 | 3 | 3 | |
Tibia | Mean | 0.83 | 1.54 | 2.21 | 1.54 | 2.11 |
Range | 0.80–0.88 | 1.48–1.58 | 2.00–2.41 | 1.48–1.64 | 2.00–2.20 | |
|
3 | 3 | 14 | 3 | 3 | |
Metatarsus | Mean | - | 2.36 | 3.55 | 2.61 | 3.98 |
Range | - | 2.16–2.63 | 2.96–4.08 | 2.48–2.80 | 3.68–4.25 | |
|
- | 3 | 14 | 3 | 3 | |
Tarsus | Mean | 0.77 | 2.74 | 4.71 | 3.10 | 4.04 |
Range | 0.73–0.85 | 2.48–3.03 | 3.90–5.25 | 2.76–3.28 | 3.56–4.31 | |
|
3 | 3 | 14 | 3 | 3 |
All measurements in millimeters,
Leg measurements given in
Penis length 2.49 mm (
For female description see
For species descriptions see
The American Arachnological Society (Vincent Roth Fund for Systematic Research) and the American Museum of Natural History (Theodore Roosevelt Memorial Grant) provided funding to CHR. Several institutional scholarships (Harry E. Hamber Memorial Scholarship, Frank Alverson Memorial Scholarship, and Jordan Dale Covin Memorial Scholarship) to CHR also helped contribute to this work. Specimens were collected with help from Daniel Richart, William Leonard, Shahan Derkarabetian, Jeffery Underwood, Adrienne Richart, Paul Marek, and Dean Levitt. Hours of morphological measurements were expedited with help from Kristen Emata, David Carlson, Nicasia O’Neal, Adrienne Richart, and Kyle Wilson. Nicasia O’Neal, Roy Larimer, and Joseph Warfel helped with images. Maureen McCormack, Shahan Derkarabetian, and Robin Keith Hedin helped with laboratory training. Stas Vidyakin helped with accessioning MorphBank images. This research was enriched from discussions of concepts and methodologies with Ricardo Carretero, Rod Crawford, Shahan Derkarabetian, Andrew Gottscho, William Leonard, Dean Levitt, Maureen McCormack, Tod Reeder, Jordan Satler, Axel Schönhofer, Peter Scott, William Shear, Jack Sullivan, and others. The Wnt2
Collection Locality Information (doi:
PCR Primer Information. (doi:
Male and Female Morphometric Data and Measurement Scheme. (doi:
PCA Methods and Results. (doi:
Morphologies of Acuclavella cf. cosmetoides and Geography of Morphotypes.
Acuclavella distribution. KMZ file of Acuclavella sampling effort, species distributions, and distribution of individuals used in concatenated phylogenetic reconstruction. (doi:
Phylogenetic Trees.
Acuclavella genitalia.
K. Leg II Morphologies. With and without femoral false leg articulations. (doi: