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• W2072161119 abstract "Violaceae, subg. Melanium (DC.) Hegi. Native annual with erect to decumbent stems (2–) 3–10 (−20) cm; simple or little branched, more or less grey pubescent all over with short, crisped or deflexed hairs. Stem from single taproot. Lowest leaves sub-orbicular, obtuse; upper to 2 cm, oblong-lanceolate, subacute, crenate-serrate; stipules, at least the lower, with large ovate-lanceolate, crenate-serrate leaf-like mid-lobe and few lateral lobes, similar to those of V. arvensis but much smaller, 0.5–1.0 cm. Single chasmogamous flowers from leaf axils. Peduncles typically exceeding leaves, glabrous, curved downward at apex, 1–2 (−4) cm long, with two minute (1 mm) bracts near base of curve. Sepals 5, subulate to oblong-lanceolate, ±5 mm long, glabrous. Margins of sepals slightly darker green than rest of sepal. Corolla zygomorphic, 5-petalled, very small, c. 5 mm vertically, about half the length of the sepals, somewhat concave, cream-white to yellow or pale blue-violet; lower petal with yellowish base near ‘throat’ of corolla, lateral petals bearded. Spur 1–2 mm, often violet, slightly longer than the calycine appendages. Stamens 5, stigma globose. Stylar flap 0. 70–85% of pollen grains with 5 pores, the remainder with 4 or 6. Capsule globular, truncate. Seed 1.6 × 1.2 mm, ovoid, smooth, yellow (Butcher 1961). Air-dry mass of seeds averages 0.31 mg. Viola is a genus of approximately 400 mainly temperate herbs and rarely undershrubs. Stace (1997) recognizes 15 species present in the wild in Britain. There are many Viola species grown in gardens that temporarily naturalize nearby. Viola kitaibeliana includes V. nana (DC.) Godr. Other commonly used synonyms outside the UK are V. rafinesquii Greene, V. minima Presl. and V. valesiaca Thomas. The dwarf pansy is native in dunes and short turf on sandy soil by the sea. It occurs on the Isles of Scilly (Tresco, Tean and Bryher) and the Channel Islands (Guernsey, including Lihou, Herm (Le Hucquet 1993) and Jersey). On Guernsey it is also found in thin soil on low, granite sea-cliffs in the south (Girard 1964). In north-west Spain and Portugal (Franco 1971) the smaller V. kitaibeliana var. henriquesii (Willk.) Beck and a more robust var. machadiana (Coutinho) Coutinho have been recognized. It is not clear whether this species is native to North America or has been introduced. In Indiana it is regarded as of European origin (http://www.ulib.iupui.edu/wildflowers/plate10.html) whereas in Kansas it is thought probably native. As a garden plant in the USA it goes by the name of ‘johnnyjump violet’ (http://www.missouriplants.com/Bluealt/Viola_kitaibeliana_page.html) with separate variations recognized in Europe and Africa. In Britain Viola kitaibeliana is limited to three islands in the Channel Islands (Jee 1972; Bichard & McClintock 1975; McClintock 1975; Le Sueur 1984) and three in the Isles of Scilly (Lousley 1971; French et al. 1999) where suitable habitats are present. However, G. Fenton (pers. comm.) also found two plants growing on St Martin's in 2003 after an absence of several years. It is currently confined to nine 10-km squares, with a site in a tenth square in the Channel Islands having been recorded pre-1970 (Fig. 1). It has never been recorded on the British mainland. Throughout its British range it is confined to sea level or thereabouts. Viola kitaibeliana occurs along the Atlantic coast of western Europe with its northernmost published locality near Ambleteuse, Pas-de-Calais (Lousley 1956). However, there is a 2003 report of V. kitaibeliana in the Dunes de Ghyvelde, Belgium (R. Slings, pers. comm.). It is frequently found in maritime sands of France in Calvados, Manche, Vendée, Charente-Inférieure and Gironde. It is widespread in south and central Europe and extends eastwards to western Syria, Turkey and the eastern Ukraine. It is also reported from Algeria (Strid 1986) and Palestine (Zohary 1973) (Fig. 2). Greuter (1974) pointed out that although it has been included within the flora of Crete, there is no basis for this. The distribution of Viola kitaibeliana in the British Isles. Each dot represents at least one record in a 10-km square of the National Grid. (○) Pre-1950; (•) 1950 onwards. Mapped by Henry Arnold, Biological Records Centre, Centre for Ecology and Hydrology, mainly from records collected by members of the Botanical Society of the British Isles. The old world distribution of Viola kitaibeliana. Source Floras of various countries: Aeschimann & Burdet (1989), Atl. Br. Fl., Carlström (1987), Davis (1965), Fl. Eur. 2, Fournier (1977), Franco (1971), Hess et al. (1970), Meikle (1977), Palhina (1939), Pignatti (1982), Polunin (1980), Stojanov & Stefanov (1948), Strid (1986), Zohary (1973) and Vergl. Chor. (1978, vol. 2). Fl. Eur. 2 lists V. kitaibeliana as present in Albania, Austria, UK, Bulgaria, Corsica, (Crete), Czechoslovakia, France, Greece, Spain, Italy, Switzerland, Hungary, Jugoslavia, Luxembourg, Portugal, Romania, Russia (west, Crimea and south-east), Sicily and Turkey. Davis (1965) includes Cyprus. Fl. Eur. 2 suggests it was introduced to Germany and possibly Poland. The Isles of Scilly locations have an ‘extreme Atlantic’ climate, equable temperature, adequate and regular supply of precipitation, near freedom from snow, high frequency of fogs and an excellent sunshine record. There is a February mean temperature of 7.3 °C and an August mean temperature of 16.5 °C (with extremes of −3.9 °C and 27.8 °C). There are, on average, per year, 1752 h of sunshine, 810 mm rainfall, 200 rain days and 350 days with a mean air temperature > 5 °C. The climate of the Channel Islands is somewhat similar though slightly sunnier and drier. The February mean temperature is 7.2 °C and the August mean temperature is 16.9 °C (with extremes of 0.3 °C and 23.4 °C). There are on average, per year, 1799 h of sunshine and 724 mm rainfall. In Hungary, its type location, V. kitaibeliana is recorded as a significant pioneer of open Pannonian sand steppe (http://www.botanika.hu/project/habhun/habitats/34A11.html) in the inland dunes of the Pannonic Plain and neighbouring basins north-west to Marchfeld. Here, it is described as part of the ‘therophyte-dominated stage of the first succession phase’. In the Czech Republic, Antonin et al. (http://www.sci.muni.cz/botany/botguide/pavlov.htm) include V. kitaibeliana as one of the sub-Mediterranean or peralpid elements reaching its northern limit in southern Moravia. Meikle (1977) described it, in Cyprus, as a plant of vineyards, gardens and fields or bare, sandy places and grassy hillsides between 625 m and 1250 m. Strid (1980) found it widely on Mount Olympus, Greece, in similar locations to V. arvensis. In other areas of Greece (Strid 1986) it is chiefly a lowland species of dry, open habitats. However, it is also found in rocky pastures and screes at subalpine level, ascending to c. 1900 m where it flowers from April to July according to altitude, escaping low winter temperatures by its annual habit. In Turkey, Davis (1965) also located it on stony slopes, screes and ‘macchie’ from sea level to 1800 m. Polunin (1980) recorded it as widespread in dry, open habitats of Greece and the Balkans except in the Aegean Islands. In Europe it is suggested that the species occurs most commonly on well-drained siliceous, sandy soils (Aeschimann & Burdet 1989). Fitter (1979) described it as a species of infertile soils. The blown sand of Scilly on which it occurs is composed of finely comminuted granite, the dominant constituent being minute grains of quartz and white felspar, the latter imparting the snow-white colour (Barrow 1906). However, much coarser sands are present on Tean. Samples of soil in which V. kitaibeliana was growing were taken for analysis from Bryher, Tean and Tresco. The results are given in Table 1. It is seen that Bryher and Tresco soils appear to be less nutrient poor, slightly more acidic, more organic rich and of a finer grade than those of Tean. McClintock (1975) and Girard (1964) pointed out that, on Guernsey, V. kitaibeliana grows best in the bare soils of the cliff-tops of the south-west, although it also occurs in the sand dunes of the north-west (Jee 1957). Throughout its range Viola kitaibeliana is recorded as occurring with a wide variety of associated species in differing habitats. A precondition of growth seems to be an open habitat with enough bare ground for seeds to germinate and low enough surrounding vegetation for competition for light to be limited. In its British locations all sites are open, sandy grassland or heathland, or derelict sandy fields, near the coast. Table 2 lists species recorded with V. kitaibeliana on the Isles of Scilly in habitats visited by the author. The commonest associates are Stellaria pallida, Cerastium fontanum, C. diffusum, Sagina procumbens, Aphanes australis, Lotus corniculatus, Trifolium occidentale, Euphorbia portlandica, Geranium molle, Myosotis ramosissima, Veronica arvensis, Galium verum, Hypochaeris radicata, Senecio vulgaris, Festuca rubra and Tortula ruraliformis. These habitats are clearly related to that described by Rodwell (2000) as SD8Festuca rubra–Galium verum fixed dune grassland, Luzula campestris subcommunity. Records of Viola kitaibeliana on the Isles of Scilly have shown that it appears to be temporary where it establishes in sites that are disturbed by man. However, it remains longer term in naturally open locations. At St Aubyn's Bay, St Brelade's Bay & St Ouen's Bay, Jersey, Druce (1907) found V. kitaibeliana growing with Cochlearia danica, Moenchia erecta, Sagina apetala, Arenaria serpyllifolia, Cerastium semidecandrum, Ornithopus perpusillus, Saxifraga tridactylites, Aphanes arvensis, Valerianella locusta, V. carinata, Myosotis ramosissima, Plantago coronopus, Carex arenaria and Bromus hordeaceus ssp. thominei. Less common associates were Capsella bursa-pastoris, Teesdalia nudicaulis, Erophila verna, Stellaria pallida, Erodium cicutarium, Trifolium subterraneum, T. dubium, Vicia sativa ssp. angustifolia, V. lathyroides, Myosotis discolor and Phleum arenarium. More rarely the dwarf pansy is found with Polygala serpyllifolia, Medicago polymorpha, Trifolium micranthum, Romulea columnae and Scilla autumnalis. At Quenvais, Euphobia paralias, E. portlandica. Silene conica and Schoenus nigricans have been recorded with V. kitaibeliana (Druce 1908). The British communities containing V. kitaibeliana do not have a close match with any European alliance but might be placed within the Koelerion albescentis of the Sedo-Scleranthetia or Galio-Koelerion of the Koelerio-Corynephoretea (Westhoff & den Held 1969) of Atlantic coastal Europe. In the open dune swards of Hungary, V. kitaibeliana is found growing with other mostly ephemeral, early flowering annuals: Bromus mollis, B. tectorum, Medicago minima, Cerastium brachypetalum, Erophila verna, Plantago indica, Saxifraga tridactylites, Poa annua, P. bulbosa, Lithospermum arvense, Coryspermum nitidum and Polygonum arenarium. Also present are a few species of longer seasonal visibility, such as Equisetum ramosissimum var. altissimum, Alyssum tortuosum, Sedum acre and Cynodon dactylon (http://www.botanika.hu/project/habhun/habitats/34A11.html). Oberdorfer (Pfl. Exk.) gives V. kitaibeliana as a very rare and inconstant species of waste places and cornfields on summer-dry nutrient- and base-rich but generally non-calcareous sandy loam soils. Viola kitaibeliana responds poorly to competition. It is normally shaded out where encroachment by Pteridium aquilinum occurs but may persist in places, under the shelter of coastal Pteridium in the Isles of Scilly. Where there is a low, closed sward of dune grassland, plants are depauperate but survive to flowering and seed-set. Ideal conditions seem to be in the partial protection and shade of species such as Carex arenaria or Holcus lanatus. Viola kitaibeliana is commonly situated adjacent to paths and in the flatter areas of dune and is consequently subjected to trampling by humans. Disturbance favours the annual by preventing the establishment of perennial species. Trampling may also aid dispersal of seed (see VIII(C)). Even though V. kitaibeliana grows most extensively in heavily rabbit-grazed turf there is no direct evidence of herbivory. Indeed, some of the best-developed plants have been recorded growing in the excavation spoil from rabbit burrows. Rabbits also reduce competition by grazing the more palatable perennials within the community. On Bryher and Tean, where paths have been mown through taller vegetation in the semi-fixed dune, V. kitaibeliana seems to grow particularly well at the boundary between mown and unmown vegetation. Viola kitaibeliana usually forms small, patchy populations, many of which persist over a number of decades (Lousley 1971). Perennial plants within the same habitat probably limit the size of patches and the density of V. kitaibeliana within them (see IV(A)). Owing to the short seed dispersal distance of the species (see VIII(C)), seedlings are often found aggregated around the sites of plants from the previous year. The largest population in Scilly extends over c. 200 m2 on Bryher. At other sites a few plants may occur within 1 or 2 m2. The population at Pellistry Bay, St Mary's was in a sandy field near the shore. A similar location for the species was in St Martin's below the school. However, the plant does not seem to persist in such locations. In natural habitats on sand dune systems it is usually found in disturbed places such as around rabbit holes or on the edge of pathways but it can also occur in closed dune turf. In such locations it rarely grows to more than 2 cm. Where it is present in arable ground or between tussocks of coarse grasses, plants may be up to +10 cm and flower later into the summer. Frost is rare at coastal locations where Viola kitaibeliana grows in the UK. An Iversen plot (Fig. 3) shows that V. kitaibeliana has a tri-modal distribution pattern. Sites in Britain are at the limits of the plant's temperature tolerance for Atlantic locations. It would appear to be best assigned to the European Southern-temperate floristic element of Preston & Hill (1997), thriving best where there is minimal frost in the growing season and adequate moisture. In its continental interior sites, the species seems to grow as a summer annual rather than overwintering as a seedling. The occurrence of Viola kitaibeliana relative to the mean temperature of the warmest and coldest months. Open circles (○) represent the presence in the British Isles and filled circles (•) presence elsewhere in the world. The broken lines give the inferred thermal limits for the species in its main distribution ranges. Records of occurrence were taken from Floras (see caption of Fig. 2); temperature data were derived from climatological Atlases for sites adjacent to those where the plant occurs. Drought is also infrequent in the Channel Islands and Scilly but years with low spring precipitation have coincided with poor flowering in the Rushy Bay, Bryher colony during the 1990s. The coastal sands of the Isles of Scilly and the Channel Islands become dry enough in summer for this species to select for summer dormancy as seed. The Rushy Bay population experienced severe sea flooding and sand burial after the ‘hurricane’ of 1987. After this event the local English Nature warden brushed away the sand covering to expose the surface of the underlying turf. The following year the best flowering of V. kitaibeliana ever seen by the author occurred! Growth of seedlings under glasshouse conditions showed that the first true leaves emerged as an opposite pair with somewhat different sizes when mature (length 45 days after germination = 2.5 and 3.0 mm). The next two true leaves came as a more even pair, but thereafter leaves emerged singly, larger (length 45 days after germination = 3.5 mm) and became noticeably alternate. Leaves of more decumbent specimens are more or less all basal and smaller. Stomatal counts from 5 leaves of different specimens from Bryher gave a mean of 135 mm−2 for the adaxial surface and 210 mm−2 for the abaxial surface. The root system of V. kitaibeliana growing in its natural habitat in Scilly is shallow, to a maximum of 3 cm, with the white primary root extending further than any laterals (Fig. 4). Root of Viola kitaibeliana after 200 days’ growth. Harley & Harley (1987) report arbuscular mycorrhiza but note that specimens are rarely mycorrhizal. Viola kitaibeliana is a winter-annual therophyte and reproduction is entirely by seed. Chromosome number 2n = 48 (Fl. Br. Is.), 2n = 16, 48 (Fl. Eur. 2). According to Pettet (1964), however, the plant of Scilly appears to be identical with that of Jersey and material from Tresco and Bryher has been found to have the same chromosome number, 2n = 24. Van Loon (1980) also found 2n = 24 in plants from Konitsa, Greece, as did Franzen & Gustavsson (1983) in Yiorla. Hess et al. (1970) recorded 2n = 14, 16, 36, 48. Pettet (1964) considered chromosome number to be an important criterion for the separation of Viola arvensis (2n = 34) from the Viola kitaibeliana complex, a pair of taxa difficult to delimit satisfactorily on purely morphological criteria (Schmidt 1964). In field locations in Britain, water availability does not seem to limit growth. However, salinity appears to be significant with no plants being recorded within 30 m distance of High Water Mark High Spring Tides, even though the vast majority of plants occur in extreme sub-maritime locations. None reported. Lousley (1971) recorded that in Scilly V. kitaibeliana commences to flower at the end of March and is in full flower in April and May. In its dune and close-cropped coastal turf habitats it dries up and disappears early, but in arable ground or damper sites straggling, late-flowering plants persist into June or early July. McClintock (1975) also recorded it as being in flower as late as 5 July in 1902, although it is usually over by early summer. Knight (1996) shows it as ‘possibly in flower’ as early as February. Up to seven single, chasmogamous flowers arise from leaf axils as day-length increases in mid-February/March. Cleistogamous flowers absent. Individual plants can flower for 4–5 weeks. Flower size may vary from 4 mm to 6 mm across, and colour may range from predominantly cream/yellow to an almost total suffusion of violet. Small insects, particularly flies and small bees, visit the flowers freely on calm, sunny days but the flowers are usually automatically self-pollinated. Knuth (Poll.2) describes the mechanism of insect pollination in the genus Viola. The following larger insect species have been noted visiting Viola kitaibeliana in the field: HYMENOPTERA: Apidae: Bombus terrestris (L.), B. lucorum (L.), Apis mellifera L. and LEPIDOPTERA: Pieridae: Pieris rapae L. There is a complex of forms from south and east Europe that require further study. Davis (1965) suggested that in Turkey intermediates occur between V. kitaibeliana and V. arvensis Murray and V. modesta Fenzl. No hybrids have been recognized in Britain. However, at Pellistry Bay, St Mary's during the 1950s the Dwarf Pansy persisted in decreasing numbers with the flowers and small fruits of V. kitaibeliana but with a growth habit and stipule form nearer V. arvensis. Trampling and seed becoming lodged in shoe treads may sometimes disperse seed. In 1980 plants were recorded for a year at a dump on Tresco and in 1986 plants were recorded at Great Porth, Bryher, away from the main population but along a frequently used pathway. Viola kitaibeliana is a diplochore (Beattie & Lyons 1975), explosively ejecting its seeds prior to ant transportation. The small seeds may be dispersed a metre or more from the parent plant. They are held tightly in each valve under pressure until dehiscence of the capsule occurs. The capsule is carried above the leaves and is held clear of the ground but still low, because of the minute size of the plant. Ants of the genus Formica have been recorded taking smaller Viola seeds below ground in less than 1 h after they were shed (Beattie & Lyons 1975). Seeds from the central row in each valve are ejected first and fall near the parent whereas others under greater pressure from the valve walls are ejected further. Seeds germinate in autumn and continue to grow through the mild winter so that plants in the more sheltered locations can flower by mid-March. Colonies have been known to be rejuvenated (see section V(C)) or reappear on former sites following disturbance (Preston et al. 2002), suggesting the presence of a persistent seed bank. Seed freshly collected from Bryher in late spring 2002 did not germinate within 3 months when placed on filter paper in an unheated glasshouse and watered with distilled water as required. However, air-dried seed showed 20% germination (emergence of radicle) after 30 days when sown on 21 September 2002. Seed collected in 2002 and sown on 1 October 2003 also resulted in 20% germination by 31 October 2003. Each capsule contains 36–48 smooth, yellow, ovoid seeds 1.6 × 1.2 mm (Fig. 5a). Germination is epigeal. After imbibition, the radicle emerges from the proximal part of the seed. Radicle emergence takes a minimum of 5 days and cotyledons appear 3–4 days later (Fig. 5b,c). Lateral root growth commences by 21 days and the first true leaves appear around the same time (Fig. 5d). Stages in germination of Viola kitaibeliana (grown in air on moist filter paper). There are no published records of animals or birds feeding on V. kitaibeliana. However, Traba et al. (2003) have found the seed in cattle dung near Madrid, suggesting that the species is grazed. A. J. Beattie (Beattie & Lyons 1975) found heavy pre-dispersal damage to Viola seed crops as a result of slug and caterpillar predation of unripe ovaries. The rust Puccinia violae is common in many species of Viola (Moore 1958) but has been seen on V. kitaibeliana only in specimens that grew temporarily in the 1980s in a garden rubbish dump on Tresco. It tends to attack all green parts of the plant. According to N. Robertson in Balme (1954) this material may be P. depauperans Sydow. Peronospora violae DeBary, Ramularia lactea (Desm.) Sacc. and R. agrestis Sacc. have been recorded on closely related species of Viola. The first known record of V. kitaibeliana in the British Isles was that of Trimen who saw it at St Aubyn's Bay on his spring visit to Jersey in the Channel Islands in 1871 (Trimen 1871). In this account, he referred to the species as Viola nemausensis Jord. Over the next few years it was recorded from St Brelade's Bay, Jersey, in 1885 by Rev’d A. Ley and at Grand Havre, Guernsey, in 1890 by Miss Dawber (Bennett 1901). The earliest published record for Scilly was that of Curnow (1876) when he recorded it as V. curtisii. The species was discovered on Tresco by J. Ralfs in July 1873 and named by him as V. curtisii var. mackaii in a letter to F. Townsend in 1876 (Ralfs 1876). According to Bennett (1901), Ralfs later found it in St Martin's in 1877. It was recognized as V. nana Corb. by Baker (1901) who mentioned a recent record by C.P. Andrews from Rousse Towers, Guernsey, but Babington (1881) had placed it in the correct group when he described a ‘small form from Scilly [as] very like V. parvula Tin. The status of Viola kitaibeliana in Britain is vulnerable, but not threatened in Europe (Wigginton 1999). The present overall distribution of the species (Preston et al. 2002) is similar to that shown in the 1962 Atlas (Atl. Br. Fl.) although one Guernsey site has not recurred since before 1970. Sympathetic habitat management practices, in particular light grazing and/or mowing regimes, are vital for this species. All V. kitaibeliana locations in the Isles of Scilly are within SSSIs and are part of a RAMSAR site. On Bryher and Tean mowing of footpaths through Pteridium and Calluna encourages open, sandy grassland in which the dwarf pansy can thrive. On Tresco, coarse vegetation and timber storage has shaded out part of the extant colony and in the past building development has wiped out previously recorded sites. G. Fenton (pers. comm.) recently has persuaded the Tresco Estate to move the timber to help conserve the pansy. At L’Ancresse common, Guernsey, cattle grazing has maintained an open pasture (Rees 1931) suitable for this species. With rising sea-level, coastal erosion is an increasing threat everywhere. Traba et al. (2003) show V. kitaibeliana to be a species that can be introduced into abandoned grasslands by experimental endozoochory via sown cattle dung, despite low frequency presence in grazed areas. I am indebted to Professor Arthur Willis for his meticulous editing, to Tony Davy, Chris Page, Michael Proctor, David Streeter and the late Franklyn Perring for critical advice, and to Geoffrey Fenton, William Wagstaff and Rosemary Parslow for unpublished data. I thank H. R. Arnold (Biological Records Centre, Centre for Ecology and Hydrology, Monks Wood) for the preparation of the British distribution map and Ian Agnew (Cartography Office, Department of Geography, Cambridge University) for other maps, graphs and drawings." @default.
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