Corresponding author: Robert W. Bryson, Jr. (
Academic editor: W. Lourenço
Morphologically conserved taxa such as scorpions represent a challenge to delimit. We recently discovered populations of scorpions in the genus
Bryson Jr RW, Wood DA, Graham MR, Soleglad ME, McCormack JE (2018) Genome-wide SNP data and morphology support the distinction of two new species of
Species delimitation of morphologically conserved taxa has been a historically challenging endeavor for taxonomists. Recent developments in both DNA sequencing and species delimitation modeling have alleviated much of this burden by providing researchers with new ways to systematically classify similar-looking yet evolutionary distinct species. Although many species delimitation methods were originally designed and tested using only a handful of genes (e.g.,
Scorpions represent a well-known group of animals with a relatively conserved morphology. Thought to be derived from amphibious ancestors that lived more than 425 million years ago, their body plan appears to have changed relatively little since their adaptation to land (
California is a global biodiversity hotspot and home to numerous endemic scorpions (i.e.,
We sequenced 36 samples of
Genetic samples of
Sample numbers | Species | Locality |
---|---|---|
sky241, sky263, sky264 |
|
CA: Ventura Co: Yerba Buena Road, Santa Monica Mountains |
sky464, sky465, sky466, sky499 |
|
CA: Los Angeles Co: Kanan-Duma Road, Santa Monica Mountains |
sky266, sky498 |
|
CA: San Bernardino Co: Mountain Home, San Bernardino Mountains |
sky470, sky471, sky472 |
|
CA: Riverside Co: Mountain Center, San Jacinto Mountains |
sky467, sky468, sky469 | San Gabriel Mtns | CA: Los Angeles Co: Eaton Canyon Falls, San Gabriel Mountains |
sky250, sky516, sky517 | Santa Ana Mtns | CA: Orange Co: Trabuco Creek Road, Santa Ana Mountains |
sky518 | Santa Ana Mtns | CA: Orange Co: Silverado Canyon Road, Santa Ana Mountains |
sky248, sky249 |
|
CA: San Diego Co: Palomar Mountain |
sky251 |
|
CA: San Diego Co: San Diego Zoo Safari Park |
sky274, sky276 |
|
CA: San Diego Co: Barrett Flume |
sky277 |
|
CA: San Diego Co: Barrett Lake Road |
sky275, sky519, sky520 |
|
CA: San Diego Co: Escondido |
sky511 |
|
CA: San Diego Co: Indian Valley Road |
sky273, sky504 |
|
CA: San Diego Co: Mission Trails |
sky512, sky513 |
|
CA: San Diego Co: Padre Dam |
sky494, sky514, sky515 |
|
CA: San Diego Co: Santa Ysabel |
We demultiplexed and processed Illumina reads using pyRAD v2.16.1 (
We performed Bayes factor species delimitation using BFD* (
Sampling localities for genetic samples of scorpions in the genus
Bayes factor comparisons of 11 competing models of species delimitation in
Model | Species | Groupings |
|
Rank | 2lnBF |
---|---|---|---|---|---|
M1 | 3 | -25035.20 | 8 | 14165.49 | |
M2 | 3 | -26835.35 | 9 | 15965.64 | |
M3 | 3 | -24961.18 | 7 | 14091.47 | |
M4 | 3 | -28211.23 | 11 | 17341.52 | |
M5 | 5 | -10869.71 | 1 | – | |
M6 | 4 | -14205.08 | 2 | 3335.37 | |
M7 | 4 | -14849.95 | 3 | 3980.24 | |
M8 | 4 | -18460.44 | 4 | 7590.73 | |
M9 | 4 | -19812.85 | 6 | 8943.14 | |
M10 | 4 | -19084.64 | 5 | 8214.93 | |
M11 | 3 | -27947.41 | 10 | 17077.70 |
We generated a final species tree in SNAPP based on the best-ranked species model from BFD*. We ran the analysis for 1,000,000 MCMC generations, sampling every 1,000 steps. We confirmed convergence and high ESS values using Tracer v1.5 (
To examine phylogenetic relationships within
We examined the morphology of 40 preserved specimens of
One sample of
The BFD* model with the best marginal likelihood value strongly supported a five-species model that placed scorpions from the San Gabriel and Santa Ana Mountains into separate lineages (Table
Species tree of scorpions in the genus
Phylogenetic analysis of the concatenated RAD loci produced a well-supported tree (Fig.
Time-calibrated phylogeny of scorpions in the genus
Estimated divergence dates among the five major clades of
Morphological assessments revealed several characters that differentiated
Telson of adult females of genus
Our genetic data strongly support the recognition of
Patronym honoring Warren E. Savary for his contributions to vaejovid scorpion taxonomy.
Large sized species for the family, with males up to 50.5 mm and females reaching 57.0 mm; pectinal tooth counts 12–13 for males and 11–13 for females. The species possesses the characteristics of genus
Dorsal and ventral views of
Measurements (in mm) of the type series of
|
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|
|
|
|
|
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Total L | 50 | 50.5 | 49 | 47.5 | 57 | 53 | 47 | 57 |
Cara L | 5.85 | 6.5 | 6.15 | 5.75 | 6.5 | 6.3 | 6.05 | 7.2 |
Meso L | 14.95 | 15.7 | 14.1 | 15.8 | 17.65 | 17.8 | 13.6 | 17.3 |
Met I L | 3 | 2.95 | 2.9 | 2.75 | 3.1 | 2.95 | 2.65 | 3.2 |
Met I W | 2.6 | 2.95 | 2.9 | 2.55 | 2.7 | 2.7 | 2.6 | 2.95 |
Met II L | 3.55 | 3.55 | 3.5 | 3.25 | 3.65 | 3.5 | 3.3 | 3.9 |
Met II W | 2.55 | 2.9 | 2.8 | 2.5 | 2.55 | 2.55 | 4.45 | 2.85 |
Met III L | 3.9 | 3.95 | 3.9 | 3.55 | 3.9 | 3.75 | 3.5 | 4.2 |
Met III W | 2.5 | 2.85 | 2.7 | 2.4 | 2.45 | 2.5 | 2.3 | 2.8 |
Met IV L | 4.8 | 4.95 | 5 | 4.5 | 4.95 | 4.75 | 4.35 | 4.5 |
Met IV W | 2.3 | 2.65 | 2.5 | 2.3 | 2.2 | 2.3 | 2.2 | 2.6 |
Met V L | 7.5 | 7.3 | 7.65 | 6.85 | 7.7 | 7.35 | 6.7 | 8.2 |
Met V W | 2.25 | 2.5 | 2.4 | 2.2 | 2.3 | 2.3 | 2.15 | 2.5 |
Tel L | - | 6.3 | - | 6.2 | 7.2 | 6.9 | 6.2 | 7.3 |
Ves L | 5.2 | 4.75 | 4.1 | 4.5 | 5.4 | 5.1 | 4.6 | 5.55 |
Ves W | 2.6 | 2.6 | 2.8 | 2.35 | 2.6 | 2.5 | 2.5 | 3 |
Ves D | 2.05 | 2.1 | 2.2 | 2 | 2.1 | 2.15 | 1.95 | 2.3 |
Acu L | - | 1.6 | 1.6 | 1.7 | 1.85 | 1.75 | 1.65 | 1.7 |
Fem L | 5.4 | 5.5 | 5.6 | 5.5 | 5.95 | 5.7 | 5.35 | 6.4 |
Fem W | 1.9 | 1.95 | 1.9 | 1.8 | 2.1 | 2 | 1.95 | 2.2 |
Pat L | 5.25 | 5.3 | 5.5 | 5.4 | 5.8 | 5.6 | 5.3 | 6.3 |
Pat W | 2.25 | 2.25 | 2.35 | 2.3 | 2.3 | 2.35 | 2.3 | 2.6 |
Chel L | 10 | 9.95 | 10.4 | 10.1 | 11.5 | 10.8 | 9.85 | 12.35 |
Palm L | 5.55 | 5.8 | 5.9 | 5.55 | 6.4 | 6.05 | 5.5 | 6.8 |
Palm W | 3 | 2.55 | 3.3 | 2 | 3.2 | 3.05 | 2.85 | 3.5 |
Palm D | 4.2 | 4.45 | 4.65 | 2.8 | 4.3 | 3.95 | 4 | 4.75 |
3.85 | 5.2 | 4.1 | 4 | 4.4 | 4.1 | 3.75 | 4.75 | |
5.25 | 5.15 | 5.5 | 5.25 | 6 | 5.55 | 5.05 | 6.45 | |
Pect Teeth | 12/13 | 13/12 | 12/12 | 11/11 | 12/11 | 11/10 | 11/11 | 12/13 |
Slight sexual dimorphism was evident in telson and metasoma morphology for
The specific name is a noun in apposition in reference to Occidental College, commonly referred to as Oxy, which lies at the base of the San Gabriel Mountains near Eaton Canyon, the type locality.
Large sized species for the family, with males up to 51.0 mm and females reaching 52.0 mm; pectinal tooth counts 12 for males and 11–13 for females. The species possesses the characteristics of genus
Dorsal and ventral views of
Measurements (in mm) of the type series of
Male holotype (DMNS ZA.38178) | Male paratype 1 (DMNS ZA.38179) | Male paratype 2 (DMNS ZA.38180) | Female paratype 1 (DMNS ZA.38181) | Female paratype 2 (DMNS ZA.38182) | Female paratype 3 (DMNS ZA.38183) | Female paratype 4 (DMNS ZA.38184) | |
---|---|---|---|---|---|---|---|
Total L | 45 | 51 | 47 | 41 | 42.5 | 52 | 43 |
Cara L | 5.35 | 6.2 | 5.8 | 5.75 | 6 | 6.1 | 6.1 |
Meso L | 14.7 | 15.2 | 13.2 | 10.8 | 10.5 | 17.5 | 10.5 |
Met I L | 2.5 | 3 | 2.7 | 2.4 | 2.5 | 2.65 | 2.6 |
Met I W | 2.45 | 2.95 | 2.55 | 2.45 | 2.5 | 2.7 | 2.6 |
Met II L | 3 | 3.5 | 3.3 | 2.9 | 2.95 | 3.2 | 3.1 |
Met II W | 2.3 | 2.9 | 2.55 | 2.35 | 2.35 | 2.5 | 2.5 |
Met III L | 3.55 | 3.95 | 3.7 | 3.2 | 3.35 | 3.6 | 3.4 |
Met III W | 2.3 | 2.8 | 2.4 | 2.2 | 2.3 | 2.45 | 2.35 |
Met IV L | 4.45 | 4.85 | 4.7 | 4 | 4.15 | 4.4 | 4.3 |
Met IV W | 2.15 | 2.7 | 2.25 | 2.1 | 2.1 | 2.3 | 2.2 |
Met V L | 6.6 | 7.6 | 7.1 | 6.2 | 6.3 | 6.8 | 6.6 |
Met V W | 2.1 | 2.5 | 2.2 | 2.05 | 2.15 | 2.2 | 2.15 |
Tel L | 5.75 | 6.55 | 6.25 | 5.8 | 6 | 6.25 | na |
Ves L | 4.25 | 4.9 | 4.6 | 4 | 4.25 | 4.5 | 4.35 |
Ves W | 2.4 | 2.7 | 2.6 | 2.25 | 2.45 | 2.5 | 2.55 |
Ves D | 1.85 | 2.2 | 2 | 1.75 | 2 | 2.05 | 2 |
Acu L | 1.6 | 1.7 | 1.65 | - | 1.75 | 1.8 | |
Fem L | 5 | 5.75 | 5.5 | 5.1 | 5.35 | 5.55 | 5.5 |
Fem W | 1.65 | 1.95 | 1.7 | 1.7 | 1.9 | 2 | 1.85 |
Pat L | 4.9 | 5.65 | 5.2 | 5 | 5.35 | 5.55 | 5.4 |
Pat W | 1.95 | 2.25 | 2.05 | 2.05 | 2.2 | 2.25 | 2.2 |
Chel L | 9.1 | 10.4 | 10.05 | 9.4 | 9.85 | 10.35 | 10.1 |
Palm L | 5 | 6.05 | 5.6 | 5 | 5.45 | 5.8 | 5.55 |
Palm W | 2.7 | 3.1 | 3.15 | 2.8 | 2.85 | 3.05 | 2.85 |
Palm D | 3.85 | 4.5 | 4.3 | 3.75 | 3.95 | 4.1 | 4.1 |
3.6 | 4.15 | 4.05 | 3.8 | 3.9 | 4.25 | 4.2 | |
4.9 | 5.55 | 5.45 | 5.05 | 5.3 | 5.6 | 5.45 | |
Pect Teeth | 12/12 | -/12 | 12/12 | 11/11 | 11/11 | 13/12 | 11/11 |
Sexual dimorphism was evident in several morphological characters for
1 | Chelal fingers are relatively long when compared to the telson vesicle width: vesicle width / movable finger 0.35–0.41 (0.382) in the female and vesicle width / fixed finger 0.46–0.57 (0.510) |
|
– | Chelal fingers are relatively short when compared to the telson vesicle width: vesicle width / movable finger length 0.43–0.50 (0.463) for the female and 0.43–0.51 (0.480) for the male, and vesicle width / fixed finger length 0.59–0.67 (0.619) for the female and 0.58–0.68 (0.642) for the male |
|
2 | Telson vesicular ridges are well developed and protrude beyond the aculeus juncture; telson vesicle is relatively wide, chelal palm depth / vesicle width 1.52–1.89 (1.680) for the female and 1.73–1.86 (1.803) for the male |
|
– | Telson vesicular ridges are of medium development and do not protrude beyond the aculeus juncture; telson vesicle is relatively thin, chelal palm depth / vesicle width 1.96–2.19 (2.052) for the female |
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3 | Pectinal tooth counts of male 12–13 and female 11–13 |
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– | Pectinal tooth counts of male 9–11 and female 10–11 |
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4 | Telson vesicle is relatively long when compared to the chelal fingers, movable finger length / vesicle length 1.09–1.16 (1.115) for the female and 1.01–1.08 (1.047) for the male, and fixed finger length / vesicle length 0.80–0.86 (0.822) for the female and 0.74 (0.740) for the male |
|
– | Telson vesicle is relatively short when compared to the chelal fingers, movable finger length / vesicle length 1.24–1.26 (1.252) for the female and 1.13–1.18 (1.157) for the male, and fixed finger / vesicle length 0.92–0.97 (0.944) for the female and 0.85–0.88 (0.858) for the male |
|
* excluding male |
We thank P. Cushing, C. Grinter, and J. Stephenson of the Denver Museum of Nature and Science (DMNS); D. Stokes, R. Wood, S. Wood, C. Rochester and E. Zarza for assistance in the field; A. Leaché for help with BFD*; R. Hansen for help editing drafts of this manuscript; and M. Adams for assistance with images. Research was conducted under scientific permits issued by CDFW. This work used the Vincent J. Coates Genomics Sequencing Laboratory at UC Berkeley, supported by NIH S10 OD018174 Instrumentation Grant. The use of trade, product, or firm names in this publication does not imply endorsement by the U.S. Government.
Morphological variation among female
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(N = 3) | (N = 4) | (N = 6) | (N = 4) | (N = 5) | |
Total L | 44.67±5.80 (38.8–50.0) | 49.38±1.89 (47.5–51.0) | 48.92±1.59 (47.0–50.5) | 44.63±4.99 (41.0–43.0) | 52.30±4.89 (47.0–57.0) |
Cara L | 5.83±0.67 (5.10–6.40) | 6.66±0.13 (6.50–6.80) | 6.96±0.33 (6.40–7.30) | 2.99±0.14 (5.75–6.10) | 6.36±0.55 (5.75–7.20) |
Met I L | 2.65±0.25 (2.40–2.90) | 2.70±0.22 (2.45–2.95) | 2.88±0.13 (2.70–3.00) | 2.54±0.10 (2.40–2.65) | 2.93±0.23 (2.65–3.20) |
Met I W | 2.55±0.28 (2.30–2.85) | 2.75±0.04 (2.70–2.80) | 2.85±0.11 (2.70–3.05) | 2.56±0.10 (2.45–2.70) | 2.70±0.15 (2.55–2.95) |
Met II L | 3.13±0.25 (2.90–3.40) | 3.29±0.11 (3.15–3.40) | 3.41±0.20 (3.15–3.60) | 3.04±0.12 (2.90–3.20) | 3.52±0.27 (3.25–3.90) |
Met II W | 2.40±0.20 (2.20–2.60) | 2.65±0.06 (2.60–2.70) | 2.68±0.15 (2.50–2.95) | 2.43±0.08 (2.35–2.50) | 2.95±0.83 (2.50–4.45) |
Met III L | 3.32±0.18 (3.15–3.50) | 3.65±0.13 (3.50–3.80) | 3.73±0.23 (3.45–4.00) | 3.39±0.14 (3.20–3.60) | 3.78±0.28 (3.50–4.20) |
Met III W | 2.30±0.20 (2.10–2.50) | 2.55±0.09 (2.45–2.65) | 2.56±0.14 (2.40–2.80) | 2.33±0.09 (2.20–2.45) | 2.49±0.19 (2.30–2.80) |
Met IV L | 4.22±0.23 (4.00–4.45) | 4.58±0.14 (4.40–4.75) | 4.50±0.36 (4.00–4.95) | 4.21±0.15 (4.00–4.40) | 4.61±0.24 (4.35–4.95) |
Met IV W | 2.15±0.15 (2.00–2.30) | 2.31±0.05 (2.25–2.35) | 2.39±0.12 (2.25–2.60) | 2.18±0.08 (2.10–2.30) | 2.32±0.16 (2.20–2.60) |
Met V L | 6.67±0.50 (6.20–7.20) | 6.84±0.23 (6.65–7.15) | 7.27±0.44 (6.80–7.80) | 4.28±0.18 (4.00–4.50) | 7.36±0.62 (6.70–8.20) |
Met V W | 2.15±0.23 (1.90–2.35) | 2.24±0.05 (2.20–2.30) | 2.36±0.14 (2.20–2.55) | 2.14±0.05 (2.05–2.20) | 2.29±0.13 (2.15–2.50) |
Ves L | 4.57±0.18 (4.40–4.75) | 4.40±0.15 (4.25–4.60) | 4.79±0.54 (4.00–5.45) | 4.28±0.18 (4.00–4.50) | 5.03±0.47 (4.50–5.55) |
Ves W | 2.47±0.15 (2.30–2.60) | 2.21±0.08 (2.15–2.30) | 2.50±0.21 (2.20–2.80) | 2.44±0.11 (2.25–2.55) | 2.59±0.25 (2.35–3.00) |
Ves D | 2.03±0.16 (1.85–2.15) | 1.94±0.05 (1.90–2.00) | 2.11±0.22 (1.80–2.40) | 1.95±0.12 (1.75–2.05) | 2.10±0.14 (1.95–2.30) |
Fem L | 5.17±0.60 (4.50–5.65) | 6.18±0.25 (6.00–6.55) | 6.38±0.31 (5.90–6.70) | 5.38±0.18 (5.10–5.55) | 5.78±0.41 (5.35–6.40) |
Fem W | 2.18±0.63 (1.60–2.85) | 2.19±0.03 (2.15–2.22) | 2.24±0.09 (2.15–2.35) | 1.86±0.11 (1.70–2.00) | 2.01±0.15 (1.80–2.20) |
Pat L | 5.03±0.57 (4.40–5.50) | 5.71±0.13 (5.60–5.90) | 6.04±0.28 (5.60–6.40) | 5.33±0.20 (5.10–5.55) | 5.68±0.40 (5.30–6.30) |
Pat W | 2.18±0.28 (1.90–2.45) | 2.53±0.12 (2.40–2.65) | 2.60±0.14 (2.40–2.75) | 2.18±0.08 (2.05–2.25) | 2.37±0.13 (2.30–2.60) |
Palm L | 5.50±0.56 (4.90–6.00) | 6.11±0.17 (6.00–6.35) | 6.52±0.29 (6.10–6.80) | 5.45±0.29 (5.00–5.80) | 6.06±0.56 (5.50–6.80) |
Palm W | 3.50±0.88 (2.85–4.50) | 3.09±0.10 (3.00–3.20) | 3.29±0.28 (2.90–3.65) | 2.89±0.10 (2.80–3.05) | 2.37±0.13 (2.00–3.50) |
Palm D | 3.83±0.35 (3.50–4.20) | 4.54±0.16 (4.40–4.70) | 4.56±0.28 (4.15–4.90) | 3.98±0.14 (3.75–4.10) | 3.96±0.72 (2.80–4.75) |
3.90±0.28 (3.60–4.15) | 4.58±0.15 (4.40–4.70) | 4.74±0.31 (4.30–5.10) | 4.04±0.19 (3.80–4.25) | 4.20±0.39 (3.75–4.75) | |
5.20±0.52 (4.60–5.50) | 6.03±0.10 (5.95–6.15) | 6.38±0.38 (5.80–6.75) | 5.35±0.20 (5.05–5.60) | 5.66±0.57 (5.05–6.45) | |
Pect Teeth | 10.20±0.84 (9–11) | 11.50±0.53 (11–12) | 12.00±0.47 (11–13) | 11.38±0.74 (11–13) | 11.30±0.82 (10–13) |
Morphological variation among male
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|
|
|
|
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(N = 4) | (N = 3) | (N = 3) | (N = 3) | |
Total L | 43.45±1.94(41.30–46.00) | 50.67±2.08(49.0–53.0) | 47.67±3.06(45.0–51.0) | 49.83±0.76(49.0–50.5) |
Cara L | 5.25±3.33(4.80–5.50) | 7.05±0.25(6.80–7.30) | 5.78±0.43(5.32–6.20) | 6.18±0.33(5.85–6.50) |
Met I L | 5.12±5.07(2.42–2.70) | 3.40±0.48(3.10–3.95) | 2.73±0.25(2.50–3.00) | 2.95±0.05(2.90–3.00) |
Met I W | 2.48±0.20(2.22–2.70) | 3.05±0.13(2.95–3.20) | 2.65±0.27(2.45–2.95) | 2.82±0.19(2.60–2.95) |
Met II L | 3.08±0.21(2.80–3.30) | 3.77±0.06(3.70–3.80) | 3.27±0.25(3.00–3.50) | 3.53±0.03(3.50–3.55) |
Met II W | 2.40±0.16(2.18–2.55) | 2.95±0.18(2.80–3.15) | 2.58±0.30(2.30–2.90) | 2.75±0.18(2.55–2.90) |
Met III L | 3.30±0.22(3.00–3.50) | 4.13±0.08(4.05–4.20) | 3.73±0.20(3.55–3.95) | 3.92±0.03(3.90–3.95) |
Met III W | 2.30±0.19(2.03–2.45) | 2.87±0.18(2.70–3.05) | 2.50±0.26(2.30–2.80) | 2.68±0.18(2.50–2.85) |
Met IV L | 4.19±0.24(3.90–4.45) | 4.90±0.62(4.20–5.40) | 4.67±0.20(4.45–4.85) | 4.92±0.10(4.80–5.00) |
Met IV W | 2.09±0.13(1.90–2.20) | 2.67±0.18(2.50–2.85) | 2.37±0.29(2.15–2.70) | 2.48±0.18(2.30–2.65) |
Met V L | 6.41±0.41(5.95–6.80) | 8.00±0.22(7.85–8.25) | 7.10±0.50(6.60–7.60) | 7.48±0.18(7.30–7.65) |
Met V W | 2.04±0.13(1.90–2.15) | 2.65±0.13(2.50–2.75) | 2.27±0.21(2.10–2.50) | 2.38±0.13(2.25–2.50) |
Ves L | 4.18±0.32(3.90–4.50) | 5.52±0.32(5.15–5.75) | 4.58±0.33(2.40–2.70) | 4.68±0.55(4.10–5.20) |
Ves W | 2.26±0.17(2.10–2.40) | 2.87±0.15(2.70–3.00) | 2.57±0.15(2.40–2.70) | 2.67±0.12(2.60–2.80) |
Ves D | 1.81±0.24(1.55–2.10) | 2.40±0.18(2.20–2.55) | 2.02±0.18(1.85–2.20) | 2.12±0.08(2.05–2.20) |
Fem L | 4.88±0.26(4.50–5.10) | 6.70±0.33(6.40–7.05) | 5.42±0.38(5.00–5.75) | 5.50±0.10(5.40–5.60) |
Fem W | 1.69±0.10(1.55–1.80) | 2.25±0.15(2.10–2.40) | 1.78±0.16(1.65–1.95) | 1.92±0.03(1.90–1.95) |
Pat L | 4.79±0.22(4.50–5.00) | 6.22±0.30(5.90–6.50) | 5.25±0.38(4.90–5.65) | 5.35±0.13(5.25–5.50) |
Pat W | 1.98±0.14(1.82–2.15) | 2.63±0.15(2.50–2.80) | 2.08±0.15(1.95–2.25) | 2.28±0.06(2.25–2.35) |
Palm L | 4.80±0.56(4.20–5.50) | 6.93±0.40(6.50–7.30) | 5.55±0.53(5.00–6.05) | 5.75±0.18(5.55–5.90) |
Palm W | 2.66±0.22(2.45–2.90) | 3.60±0.26(3.40–3.90) | 2.98±0.25(2.70–3.15) | 2.95±0.38(2.55–3.30) |
Palm D | 3.70±0.27(3.35–4.00) | 5.17±0.25(4.90–5.40) | 4.22±0.33(3.85–4.50) | 4.43±0.23(4.20–4.65) |
3.70±0.18(3.50–3.90) | 5.02±0.38(4.65–5.40) | 3.93±0.29(3.60–4.15) | 4.38±0.72(3.85–5.20) | |
4.69±0.41(4.15–5.10) | 6.67±0.60(6.10±7.30) | 5.30±0.35(4.90–5.55) | 5.30±0.18(5.15–5.50) | |
Pect Teeth | 10.88±0.35(10–11) | 12.83±0.41(12–13) | 12.00±0.00(12) | 12.33±0.52(12–13) |