Aphonopelma joshua
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Ordo: Araneae
Familia: Theraphosidae
Genus: Aphonopelma
Name
Aphonopelma joshua Prentice, 1997 – Wikispecies link – Pensoft Profile
Diagnosis
Aphonopelma joshua (Fig. 73) is a member of the paloma species group and can be identified by a combination of morphological, molecular, and geographic characteristics. Nuclear and mitochondrial DNA identifies Aphonopelma joshua as a strongly supported monophyletic lineage (Figs 7–8) embedded within the turret-building group (see Hendrixson et al. 2013[1] and Graham et al. 2015[2]) that includes (Aphonopelma atomicum sp. n., Aphonopelma mojave, Aphonopelma icenoglei sp. n., and Aphonopelma prenticei sp. n.). Aphonopelma joshua can easily be differentiated from other turret-building species by possessing stout sternal setae, from Aphonopelma iodius due to its smaller size and reduced extent of scopulation on metatarsus III and IV, and from other members of the paloma species group by their locality. The most significant measurement that distinguishes male Aphonopelma joshua from its closely related phylogenetic and syntopic species is A1. Male Aphonopelma joshua can be distinguished by possessing a smaller Cl/A1 (≤1.57; 1.42–1.57) than Aphonopelma atomicum (≥1.70; 1.70–1.93), Aphonopelma iodius (≥1.64; 1.64–2.23), Aphonopelma icenoglei (≥1.64; 1.64–1.97), Aphonopelma mojave (≥1.58; 1.58–1.94), Aphonopelma prenticei (≥1.68; 1.68–1.92), and Aphonopelma xwalxwal sp. n. (≥1.64; 1.64–1.75). The most significant measurements that distinguish female Aphonopelma joshua from its closely related phylogenetic and syntopic species are F1, A1, and the extent of scopulation on metatarsus IV. Female Aphonopelma joshua can be distinguished by possessing a larger A1/F3 (≥0.57; 0.57–0.64) than Aphonopelma atomicum (≤0.56; 0.53–0.56; by possessing a smaller F1/M4 (≤1.04; 0.98–1.04) than Aphonopelma icenoglei (≥1.07; 1.07–1.23) and Aphonopelma prenticei (≥1.04; 1.04–1.25); by possessing a smaller F1/T3 (≤1.57; 1.47–1.57) than Aphonopelma mojave (≥1.57; 1.57–1.92); and a smaller L4 scopulation extent (34%-40%) than Aphonopelma iodius (59%-83%). Females of Aphonopelma xwalxwal are unknown at this time and cannot be compared.
Description
Male and female originally described by Prentice (1997)[3].
Redescription of male exemplar
(APH_1476; Fig. 74). Specimen preparation and condition: Specimen collected live wandering, preserved in 80% ethanol; deposited in AUMNH; original coloration faded due to preservation. Left legs I, III, IV, and left pedipalp removed for measurements and photographs; stored in vial with specimen. Right legs III & IV removed for DNA and stored at -80°C in the AUMNH (Auburn, AL). General coloration: Black or faded black. Cephalothorax: Carapace 8.48 mm long, 7.26 mm wide; Hirsute; densely clothed with black or faded black pubescence mostly appressed to surface; fringe covered in long setae not closely appressed to surface; foveal groove medium deep and straight; pars cephalica region rises gradually from foveal groove to ocular area; AER procurved, PER slightly recurved; normal sized chelicerae; clypeus extends forward on a curve; LBl 1.153, LBw 1.557; sternum hirsute, clothed with short black or faded black, densely packed setae. Abdomen: Densely clothed in short black pubescence with numerous longer red/orange setae interspersed; possessing a distinct, dense dorsal patch of black Type I urticating bristles (Cooke et al. 1972[4]) - smaller and distinct from large species. Legs: Hirsute; densely clothed in black/faded black pubescence. Metatarsus I straight. F1 8.73; F1w 1.81; P1 2.84; T1 8.05; M1 7.29; A1 5.41; F3 8.11; F3w 1.97; P3 3.20; T3 6.76; M3 8.37; A3 5.42; F4 9.42; F4w 1.34; P4 3.12; T4 8.44; M4 10.39; A4 6.30; femur III is swollen. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 68.3%; leg IV (SC4) = 39.3%. Three ventral and two prolateral spinose setae on metatarsus III, with numerous medium stout setae throughout; eight ventral spinose setae and one prolateral on metatarsus IV, with numerous medium stout setae throughout; three prolateral and three ventral spinose setae on tibia I; one large megaspine is present at the apex on the retrolateral tibia of the mating clasper; one megaspine on the prolateral branch of the mating clasper; two megaspines on either side of the apex of the retrolateral branch of the mating clasper, with numerous thickened setae throughout. Coxa I: Prolateral surface covered by fine, hair-like setae. Pedipalps: Very hirsute, particularly ventrally; densely clothed in the same setal color as the other legs; one spinose seta near the anterior margin of the prolateral palpal femur; one spinose seta on prolateral patella; four spinose setae and two ventral spinose setae on the prolateral palpal tibia, with numerous medium stout setae throughout; PTl 4.846, PTw 1.548. When extended, embolus tapers and gently curves to the retrolateral side; embolus slender, no keels. Variation (6). Cl 7.549–8.48 (7.931±0.14), Cw 6.92–7.62 (7.264±0.11), LBl 1.101–1.339 (1.181±0.04), LBw 1.208–1.709 (1.417±0.08), F1 8.656–9.24 (8.941±0.1), F1w 1.81–2.145 (1.964±0.05), P1 2.84–3.48 (3.158±0.1), T1 7.899–8.96 (8.302±0.17), M1 7.135–7.868 (7.461±0.11), A1 5.13–5.41 (5.266±0.05), L1 length 31.749–34.445 (33.127±0.42), F3 8.028–8.527 (8.23±0.07), F3w 1.97–2.678 (2.5±0.11), P3 2.655–3.20 (2.978±0.1), T3 6.76–7.597 (7.071±0.14), M3 8.213–9.248 (8.707±0.16), A3 5.208–6.21 (5.542±0.14), L3 length 30.918–34.04 (32.527±0.51), F4 9.225–9.93 (9.565±0.11), F4w 1.34–2.084 (1.846±0.12), P4 2.82–3.14 (3.004±0.06), T4 8.172–9.04 (8.676±0.15), M4 10.39–11.45 (11.005±0.19), A4 5.23–6.30 (5.862±0.16), L4 length 36.896–39.52 (38.112±0.41), PTl 4.469–5.021 (4.786±0.08), PTw 1.433–1.58 (1.506±0.03), SC3 ratio 0.532–0.706 (0.628±0.03), SC4 ratio 0.242–0.393 (0.342±0.02), Coxa I setae = very thin tapered, F3 condition = swollen.
Description of female exemplar
(APH_2306; Figs 75–76). Specimen preparation and condition: Specimen collected live from burrow, preserved in 80% ethanol; deposited in AUMNH; original coloration faded due to preservation. Left legs I, III, IV, and pedipalp removed for photographs and measurements; stored in vial with specimen. No tissue for DNA. Genital plate with spermathecae removed and cleared, stored in vial with specimen. General coloration: Black or faded black. Cephalothorax: Carapace 9.15 mm long, 8.02 mm wide; Hirsute, densely clothed with short black or faded black pubescence closely appressed to surface; fringe densely covered in slightly longer setae; foveal groove medium deep and straight; pars cephalica region rises gradually from foveal groove to ocular area; AER slightly procurved, PER recurved; normal chelicerae, clypeus extends forward on a slight curve; LBl 1.36, LBw 1.51; sternum hirsute, clothed with short black or faded black setae. Abdomen: Densely clothed dorsally in short black or faded black setae with longer, lighter setae (generally red or orange in situ) focused near the urticating patch; small but dense dorsal patch of black Type I urticating bristles (Cooke et al. 1972[4]) - smaller and distinct from large species. Spermathecae: Paired and separate, very simple, with capitate bulbs and wide bases that are not fused. Legs: Hirsute; densely clothed in short black or faded black pubescence; F1 7.23; F1w 2.05; P1 2.72; T1 6.30; M1 4.48; A1 3.74; F3 6.54; F3w 1.90; P3 2.82; T3 4.88; M3 5.62; A3 4.13; F4 8.11; F4w 2.07; P4 2.76; T4 6.58; M4 7.34; A4 4.28. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 66.9%; leg IV (SC4) = 39.9%. Three ventral spinose setae on metatarsus III; six ventral spinose setae and one prolateral on metatarsus IV, with numerous thicker setae throughout. Coxa I: Prolateral surface covered by fine, hair-like and very thin tapered setae. Pedipalps: Densely clothed in the same setal color as the other legs; one spinose seta on the apical, prolateral femur, six prolateral (two at the apical, prolateral border with the tarsus) and two ventral spinose setae (one at the apical, ventral border with the tarsus) on the tibia. Variation (6). Cl 6.05–9.82 (8.238±0.59), Cw 5.13–8.02 (7.057±0.48), LBl 0.901–1.36 (1.188±0.07), LBw 1.133–1.52 (1.383±0.07), F1 5.048–8.45 (6.805±0.51), F1w 1.436–2.256 (1.869±0.14), P1 2.07–3.322 (2.712±0.22), T1 4.231–6.91 (5.813±0.4), M1 2.853–5.35 (4.235±0.4), A1 2.597–4.16 (3.541±0.22), L1 length 16.909–28.16 (23.106±1.72), F3 4.163–7.21 (5.864±0.45), F3w 1.319–2.154 (1.732±0.12), P3 1.839–2.92 (2.453±0.2), T3 3.241–5.49 (4.486±0.32), M3 3.484–5.82 (4.887±0.4), A3 2.98–4.147 (3.718±0.2), L3 length 15.707–25.52 (21.409±1.55), F4 5.668–8.55 (7.329±0.47), F4w 1.265–2.182 (1.818±0.14), P4 2.175–3.54 (2.745±0.22), T4 4.788–7.34 (6.271±0.38), M4 5.019–8.15 (6.732±0.48), A4 3.488–4.73 (4.195±0.22), L4 length 21.138–32.31 (27.766±1.99), SC3 ratio 0.627–0.696 (0.657±0.01), SC4 ratio 0.345–0.399 (0.368±0.01), Coxa I setae = very thin tapered. Spermathecae variation can be seen in Figure 76.
Material examined
United States: California: Riverside: Joshua Tree National Park, 34.028757 -116.315665 6, 4880ft., [APH_0883, 2007, 1♀, Josh Richards, AUMNH]; Joshua Tree National Park, 1 mile off Keys View Road, 33.87529 -116.038777 5, 3589ft., [APH_2306, 3/5/1989, 1♀, T.R. Prentice, AMNH]; Joshua Tree National Park, Cottonwood Springs area, 33.743197 -115.811558 1, 3116ft., [APH_1498-1499, 5/9/2012, 1♂, 1 juv, Brent E. Hendrixson, AUMNH]; [APH_2292, 23/8/1989, 1♂, T.R. Prentice, AMNH]; [APH_2300, 23/8/1993, 1♂, T.R. Prentice, AMNH]; [APH_2315, 23/8/1989, 1♂, T.R. Prentice, AMNH]; Joshua Tree National Park, Covington Flats area, 34.00862 -116.304733 1, 4831ft., [APH_1475-1477, 29/7/2012, 1♂, 1♀, 1 juv, Brent E. Hendrixson, Brendon Barnes, Austin Deskewies, AUMNH]; [APH_2293, 27/7/1992, 1♂, T.R. Prentice, AMNH]; [APH_2301, 2/8/1962, 1♂, Jim Ortez, AMNH]; [AUMS_2567, 2/8/1962, 1♂, unknown, AUMNH]; Joshua Tree National Park, Hidden Valley Campground, 34.01651 -116.16179 1, 4200ft., [APH_0335-0337, 8/5/2008, 3 juv, Brent E. Hendrixson, Zach Valois, AUMNH]; [APH_1200, 29/7/2010, 1♂, Brent E. Hendrixson, Brendon Barnes, Nate Davis, AUMNH]; Joshua Tree National Park, off El Dorado Mine Rd, Belle Campground, 33.99305 -116.0213 1, 3872ft., [APH_1007, 10/5/2010, 1♀, Chris A. Hamilton, AUMNH]; Joshua Tree National Park, picnic area, Covington Flats area, 34.027454 -116.30194 1, 4656ft., [APH_0492-0494, 16/5/2009, 3 juv, Brent E. Hendrixson, Bernadette DeRussy, Sloan Click, Jason Bond, AUMNH]; [APH_1199, 29/7/2010, 1 juv, Brent E. Hendrixson, Brendon Barnes, Nate Davis, AUMNH]; Joshua Tree National Park, Upper Covington Flats Area, near back country hiking trail parking area, 34.0085 -116.30694 1, 4800ft., [APH_0329-0331, 8/5/2008, 3 juv, Brent E. Hendrixson, Zach Valois, AUMNH]; Lower Covington Flats, Joshua Tree National Monument, 34.004968 -116.307553 5, 4861ft., [AUMS_2636, 31/7/1972, 1♂, Marqua, AUMNH]; [AUMS_2639, 31/7/1972, 1♂, D.G. Marqua, AUMNH]; Pleasant Valley, Joshua Tree National Park, 33.903782 -116.043557 5, 4180ft., [APH_2291, 27/4/1965, 1♀, E.L. Sleeper and S.L. Jenkins, AMNH]; [APH_2316, 23/9/1967, 1♂, E.L. Sleeper and S.L. Jenkins, AMNH]; Smoke Tree Wash, In Joshua Tree National Park, 33.796978 -115.787031 5, 2812ft., [APH_2296, 31/8/1989, 1♂, T.R. Prentice, AMNH]; Squaw Tank, Joshua Tree National Park, 34.087561 -116.153877 5, 3442ft., [APH_2317, 9/9/1966, 1♂, E.L. Sleeper and S.L. Jenkins, AMNH]; San Bernardino: Burns Canyon, 8.3 miles west of Hwy 247 , 34.206942 -116.596597 4, 5400ft., [APH_2290, 2/11/1991, 1♀, T.R. Prentice, AMNH]; [APH_2295, 2/11/1991, 1♀, T.R. Prentice, AMNH]; Burns Canyon, 8.3 miles west of Pipes Canyon Rd. , 34.227029 -116.668521 4, 6096ft., [APH_2289, 2/11/1989, 1♀, T.R. Prentice, AMNH]; Covington Flats, 34.077473 -116.357061 5, 4587ft., [APH_2302, 10/8/1989, 1♂, T.R. Prentice, AMNH]; [APH_2305, 10/8/1989, 1♂, T.R. Prentice, AMNH]; [APH_2308, 27/7/1990, 1♂, T.R. Prentice, AMNH]; [APH_2311, 27/12/1990, 1♂, T.R. Prentice, AMNH]; [APH_2313, 24/7/1989, 1♂, T.R. Prentice, AMNH]; [APH_2314, 3/8/1989, 1♂, T.R. Prentice, AMNH]; [AUMS_2479, 26/7/1990, 1♂, T.R. Prentice, AUMNH]; Covington Flats, below Joshua Tree National Park entrance, 34.051039 -116.342447 5, 4577ft., [APH_2307, 30/7/1992, 1♀, T.R. Prentice, AMNH]; [APH_2312, 27/7/1992, 1♂, T.R. Prentice, AMNH]; [APH_2304, 28/7/1992, 1♂, T.R. Prentice, AMNH]; Covington Flats, area before the entrance, 34.077473 -116.357061 5, 4587ft., [APH_2303, 28/7/1992, 1♂, T.R. Prentice, AMNH]; Covington Flats, in Joshua Tree National Monument, .5 miles north of split in the road, 34.071626 -116.367566 5, 4206ft., [APH_2310, 12/8/1992, 1♂, T.R. Prentice, AMNH]; Joshua Tree National Monument, 4.5 miles SE of entrance on Quail Springs Road., 34.05728 -116.224505 4, 3878ft., [APH_2286, 28/3/1989, 1♂, T.R. Prentice, AMNH]; Joshua Tree National Monument, below the entrance, 34.081762 -116.254111 5, 3996ft., [APH_2309, 27/7/1992, 1♂, T.R. Prentice, AMNH]; Joshua Tree National Monument, Lost Horse Valley, 1.1 miles south of Quail Springs Rd on Keys View Rd, then 1 mile west, 33.968748 -116.184906 4, 4469ft., [APH_2287, 3/5/1989, 1♂, T.R. Prentice, AMNH]; New Dixie Mine Rd, 6.3 miles west of Hwy 247, then .5 miles south, 34.264951 -116.5958 4, 5331ft., [APH_2288, 11/11/1992, 1♀, T.R. Prentice, AMNH]; Pipes Canyon Road, 34.199004 -116.492984 5, 4065ft., [APH_2294, 4/11/1989, 1♂, T.R. Prentice, AMNH]; Pipes Canyon, 2.5 miles west of Hwy 247 towards Burns Canyon, 34.194486 -116.479814 4, 4377ft., [APH_2298, 18/4/1990, 1♂, T.R. Prentice, AMNH]; Pipes Canyon, 4.3 miles north of Hwy 247, 34.246525 -116.493804 4, 4042ft., [APH_2297, 1/8/1992, 1♂, T.R. Prentice, AMNH]; Pipes Canyon, 5.9 miles west of Hwy 247 , 34.190832 -116.536894 4, 4288ft., [APH_2299, 13/11/1992, 1♀, T.R. Prentice, AMNH]; Upper Morongo Valley, 5.1 miles north of the Post Office off Hwy 62., 34.093965 -116.512293 4, 2799ft., [APH_2285, 6/9/1993, 1♀, T.R. Prentice, AMNH]; San Bernardino Mtns., Pipes Canyon road, 3.2 miles W of jct with Hwy 247, 34.198916 -116.486393 5, 4000ft., [AUMS_3323, 6/9/1978, 1♂, W. Icenogle, AUMNH].
Distribution and natural history
Aphonopelma joshua has a rather limited distribution within Joshua Tree National Park and surrounding areas in south-central San Bernardino and north-central Riverside Counties, California (Fig. 77). These spiders have been collected from elevations between 850 and 1500 meters in habitats characteristic of the Mojave and Sonoran deserts including the Eastern Mojave Low Ranges and Arid Footslopes Level III Ecoregion. The species distribution model suggests that suitable habitat may be present well beyond the boundaries of Joshua Tree National Park (Fig. 77B), but many of these regions are either unrealistically discontinuous (e.g., Arizona and northern Baja California) or already occupied by close relatives (e.g., Aphonopelma xwalxwal near the San Jacinto and Santa Rosa Mountains and Aphonopelma prenticei in Mojave National Preserve). Consequently, we feel that our sampling of Aphonopelma joshua provides a reasonable estimate of the extent of its distribution. This species is syntopic with Aphonopelma iodius (burrows of both species have been located within several meters of each other) and probably shares portions of its range with Aphonopelma icenoglei in the vicinity of Yucca Valley. Burrow entrances are generally surrounded by a distinct mound or turret made of excavated soil and silk (Fig. 2D–E). Mating is nocturnal and occurs during the summer (July-September, earlier in the northwestern portion of its range and later in the southeastern portion). A more extensive account of the natural history of Aphonopelma joshua is provided by Prentice (1997)[3].
Conservation status
Despite its narrow distribution, Aphonopelma joshua is largely protected by Joshua Tree National Park. Recreational activities may pose some concern but this species is common throughout the park and is likely secure.
Remarks
Aphonopelma joshua is one of the larger species of miniature Aphonopelma and is morphologically very similar to the new species Aphonopelma xwalxwal. Other important ratios that distinguish males: Aphonopelma joshua possess a smaller F1/M4 (≤0.84; 0.78–0.84) than Aphonopelma atomicum (≥0.89; 0.89–0.91), Aphonopelma iodius (≥0.89; 0.89–1.04), Aphonopelma icenoglei (≥0.90; 0.90–1.02), Aphonopelma mojave (≥0.87; 0.87–0.94), Aphonopelma prenticei (≥0.91; 0.91–0.95); by possessing a smaller L1/L3 (≤1.03; 0.99–1.03) than Aphonopelma iodius (≥1.05; 1.05–1.20), Aphonopelma mojave (≥1.06; 1.06–1.11), Aphonopelma prenticei (≥1.05; 1.05–1.11), Aphonopelma xwalxwal (≥1.07; 1.07–1.12); by possessing a smaller L3 scopulation extent (53%-71%) than Aphonopelma iodius (78%-96%); by possessing a smaller L4 scopulation extent (24%-39%) than Aphonopelma iodius (62%-88%); by possessing a smaller M1/F3 (<0.92; 0.88–0.92) than Aphonopelma xwalxwal (>0.98; 0.98–1.02). Other important ratios that distinguish females: Aphonopelma joshua possess a larger Cl/P1 (≥2.77; 2.77–3.53) than Aphonopelma atomicum (≤2.50; 2.43–2.50); by possessing a smaller L1/Cl (≤2.87; 2.67–2.87) than Aphonopelma icenoglei (≥2.92; 2.92–3.13); by possessing a smaller Cl/M3 (≤1.74; 1.62–1.74) than Aphonopelma prenticei (≥1.77; 1.77–2.20); by possessing a smaller A1/T4 (≤0.59; 0.54–0.59) than Aphonopelma mojave (≥0.58; 0.58–0.67; small overlap). For both males and females, certain morphometrics have potential to be useful, though due to the amounts of variation, small number of specimens, and the small differences between species, no others are claimed to be significant at this time (see Suppl. material 2). During evaluation of traditional PCA morphospace, males of Aphonopelma joshua separate in PCA morphological space from Aphonopelma atomicum, Aphonopelma iodius, Aphonopelma icenoglei, Aphonopelma mojave, and Aphonopelma prenticei but do not separate from Aphonopelma xwalxwal. Female Aphonopelma joshua separate from Aphonopelma iodius and Aphonopelma prenticei in morphological space, but do not separate from Aphonopelma atomicum, Aphonopelma icenoglei, and Aphonopelma mojave. Interestingly, Aphonopelma joshua males separate from Aphonopelma iodius and Aphonopelma xwalxwal, as well as Aphonopelma atomicum, Aphonopelma icenoglei, Aphonopelma mojave, and Aphonopelma prenticei in three-dimensional PCA morphospace (PC1~PC2~PC3). Aphonopelma joshua females separate from Aphonopelma iodius and Aphonopelma prenticei but do not separate from Aphonopelma atomicum, Aphonopelma icenoglei, or Aphonopelma mojave. PC1, PC2, and PC3 explain ≥97% of the variation in all analyses.
Taxon Treatment
- Hamilton, C; Hendrixson, B; Bond, J; 2016: Taxonomic revision of the tarantula genus Aphonopelma Pocock, 1901 (Araneae, Mygalomorphae, Theraphosidae) within the United States ZooKeys, (560): 1-340. doi
Images
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![Figure 7. Maximum Likelihood inferred CO1 gene tree phylogeny for the 1,032 Aphonopelma specimen dataset. Species delimitations followed the integrative methodological approach outlined in Hamilton et al. (2014)[5]. Black circles denote bootstrap support ≥ 80%; white squares denote bootstrap support ≤ 80%. White triangles indicate species clades supported with ≥ 80% bootstrap support; grey triangles indicate lineages with putative mitochondrial introgression events. Asterisks at the tips of branches indicate undescribed diversity.](/wiki/File:Zookeys-560-e6264-g007.jpg)
Other References
- ↑ Hendrixson B, DeRussy B, Hamilton C, Bond J (2013) An exploration of species boundaries in turret-building tarantulas of the Mojave Desert (Araneae, Mygalomorphae, Theraphosidae, Aphonopelma). Molecular Phylogenetics and Evolution 66: 327–340. doi: 10.1016/j.ympev.2012.10.004
- ↑ Graham M, Hendrixson B, Hamilton C, Bond J (2015) Miocene extensional tectonics explain ancient patterns of diversification among turret-building tarantulas (Aphonopelma mojave group) in the Mojave and Sonoran deserts. Journal of Biogeography 42(6): 1052–1065. doi: 10.1111/jbi.12494
- ↑ 3.0 3.1 Prentice T (1997) Theraphosidae of the Mojave Desert west and north of the Colorado River (Araneae, Mygalomorphae, Theraphosidae). Journal of Arachnology 25: 137–176.
- ↑ 4.0 4.1 Cooke J, Roth V, Miller F (1972) The urticating hairs of theraphosid spiders. American Museum Novitates, 1–43.
- ↑ Hamilton C, Hendrixson B, Brewer M, Bond J (2014) An evaluation of sampling effects on multiple DNA barcoding methods leads to an integrative approach for delimiting species: A case study of the North American tarantula genus Aphonopelma (Araneae, Mygalomorphae, Theraphosidae). Molecular Phylogenetics and Evolution 71: 79–93. doi: 10.1016/j.ympev.2013.11.007
