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W1976945654 abstract "A perennial hemicryptophyte with a basal rosette of three to eight overwintering leaves arising from a rootstock. Erect leafy stem up to c. 1 m high, terminating in a much branched panicle with many capitula. Older rootstocks occasionally producing more than one rosette of leaves and erect stem per plant. Leaves light green in summer and dark green in winter in shaded locations, with or without red–crimson undersides. In more open habitats undersides of leaves usually crimson, but entire leaves and other above-ground parts may also be crimson. Leaves lobed, 40–220 mm long, 20–135 mm wide, glabrous. Rosette leaves and leaves on lower portion of erect stem lyrate-pinnatifid with a flange of the leaf blade of variable width connecting individual lobes. Capitula 9–15 mm across (Mejías 1994; Stace 1997), usually with five ligulate florets; capitula borne in a large open panicle. Involucre 7–10 mm, narrowly cylindrical, inner bracts linear, outer bracts very small and spreading (Fl. Eur. 4; Clapham et al. 1987). Florets yellow and glabrous. Achene black or sometimes brown, 3–4 mm long including pale short, slender beak, 0.9 mm wide, with pappus of white simple hairs in two unequal rows, c. 5.5 mm in length; achene weight 0.34 mg (Fl. Eur. 4; Clapham et al. 1987; Grime et al. 1988). Plants self-fertile (Mejías 1994). Although evidence for population differentiation has not been published, screening of plants from open limestone pavement in the Burren, Co. Clare, Ireland, and from woodland near Durrow, Co. Laois, Ireland, using random amplified polymorphic (RAPD) primers (C. Kavanagh, T. Gallagher & B.A. Osborne, unpublished data) indicates genetic differences between these two populations. The two populations could also be separated on the basis of leaf anthocyanin content, but not when a number of photosynthetic and related traits were used (G. Lanigan, B. A. Osborne & G. Clabby, unpublished data). Native in Britain but doubtfully native in Ireland. Generally a plant of shaded places including woodlands, wood margins, scrub, walls and rock outcrops. In Ireland reaches its greatest abundance on open limestone pavement in the Burren. It is present throughout most of temperate continental Europe, with a distribution centred on central and southern regions (Fig. 2; Fl. Eur. 4), and may be classified phytogeographically as a European temperate species (Preston & Hill 1997). It is more sparsely distributed in Scandinavia, parts of the Russian Federation and western parts of the Iberian peninsula (Fl. Eur. 4; Atl. Fl. Eur.; Fitter 1978; Ellenberg 1988). Its northernmost European location is in Norway at c. 68.5 °N (Clapham et al. 1987). It is absent from the Azores, the Balearic Islands, Crete, the Faroe Islands, Iceland, Portugal, the Russian Federation approximately north of latitude 60 °N and the islands of Spitsbergen, Björnnöya and Jan Mayen (Fl. Eur. 4). The Northern hemisphere distribution of Mycelis muralis. Hatched areas indicate regions where the species is fairly common to common. Dots indicate isolated occurrences, and areas bounded by a line indicate regions for which precise data are not available. Amended from Hultén (1986). In the Swiss canton of Wallis (Valais) it has been documented at an altitude of 1800 m (Hegi Fl. ed. 1, 1) and in Turkey up to 1600 m (Davis 1975). In the Polish Carpathian Mountains it reaches an altitude of 940 m and is most abundant above 600 m (Zarzycki 1976). Mycelis muralis also occurs in North Africa, including Algeria (Quezel & Santa 1963) and Morocco where it is found up to 1400 m (Jahandiez & Maire 1934). It also occurs in the Caucasus where it reaches an altitude of 2300 m (Fl. URSS 29). Mycelis muralis has been introduced in North America (Gleason & Cronquist 1963) and now occurs in the Atlantic, Cordilleran and Pacific phytogeographic regions (Klinka et al. 1989). It is also present in New Zealand on Stewart Island, from Coromandel southwards on the North Island and throughout the South Island (Webb et al. 1988). In continental Europe M. muralis is a woodland species most commonly found in open woods, wood margins and woodland clearings (Ellenberg 1988), occurring to a lesser extent in scrub, on walls, rock outcrops and montane habitats (Hegi Fl. ed. 1, 1; Fl. URSS 29; Fl. Eur. 4). In Britain and Ireland M. muralis is also found in open woods, wood margins and scrub but is also locally common on natural rock outcrops and on walls. The temperate distribution of M. muralis suggests intolerance of climatic extremes. Its northern limit in Europe follows approximately the – 7 °C mean January isotherm, and its southern limit in Europe and adjacent North Africa follows approximately the 25 °C mean July isotherm (Wallén 1970). In addition, in Europe, North Africa and the Caucasus, M. muralis occurs in areas with an average annual rainfall of 500 mm or above (Wallén 1970) and this may be a factor in setting southern and eastern distribution limits. In western Europe it does not seem possible to attribute the sparse occurrence of M. muralis in Ireland, Scotland and the Iberian peninsula to any simple climatic variables. The recent and ongoing increase of M. muralis in Ireland and Scotland suggests that it has not yet completed its north-westward migration after the last glaciation, and this may also explain its absence from Iceland and western parts of the Iberian peninsula. Alternatively this could reflect climatic and land-use changes in recent decades. Typically, M. muralis is a plant of shaded or semi-shaded locations, such as open woodland, woodland clearings and margins, throughout its range (Fl. URSS 29; Jahandiez & Maire 1934; Quezel & Santa 1963; Ellenberg 1988; Grime et al. 1988; Webb et al. 1988; Klinka et al. 1989; Clabby & Osborne 1994). These habitats may often be associated with natural or anthropogenic disturbance such as storms and fires, clear-felling, etc. Mycelis muralis is also found in open habitats such as limestone pavement in the Burren, Co. Clare, Ireland or, for example, as part of the weedy flora of small, non-forested islands in Barkley Sound, Vancouver Island, British Columbia, Canada. In Ireland and Britain it also occurs in fissures in limestone pavement (grikes) and is most common in shallow grikes or above a depth of 100 cm in deeper grikes (Silvertown 1982, 1983; Clabby 1992). In Britain M. muralis appears to be confined to grikes in limestone pavement habitats (Silvertown 1982, 1983). Microclimatically, grikes may resemble woodland edge environments where plants are subject to comparable changes in light level, temperature and humidity and this could account, in part, for this distribution. In the Sheffield area of Britain, M. muralis is most frequent on shaded north-facing slopes (Grime et al. 1988). In Ireland it is most frequent on north-facing walls (Clabby & Osborne 1994), although this does not seem to be the case in south-eastern Britain (R. Payne, personal communication). Mycelis muralis occurs predominantly on north- or south-facing slopes in north-western parts of the Carpathian mountains (Zarzycki 1976). Mycelis muralis does not appear to be adversely affected by exposure to wind as it is widespread on the windswept limestone pavements of the Burren, and the Aran Islands, western Ireland. Mycelis muralis grows on a range of base-rich substrata in Ireland, Britain and continental Europe, including mineral soils and a number of skeletal substrates such as limestone pavements and walls. In woodland M. muralis is typically found over brown earth soils and rendzinas, which are often base-rich and calcareous throughout (Ellenberg 1988; Rodwell 1991; Clabby & Osborne 1994). These soils are normally rich in invertebrates and lumbricid worms, with rapid incorporation of leaf litter which produces a mull humus (Rodwell 1991). In some places M. muralis occurs on more acidic brown earth soils but only where base-rich conditions are maintained (Ellenberg 1988; Rodwell 1991). On limestone pavement and walls, plants are usually rooted in a variable base-rich substrate derived from the parent material, wind-blown debris and organic matter (Clabby & Osborne 1994). Mycelis muralis occurs mainly on free-draining soils that may be seasonally flooded (Rodwell 1991; Clabby 1992). Distribution data and Al sensitivity (Rode & Runge 1991) suggest that M. muralis is a calcicole. Despite this, M. muralis occurs on soils over a wide range of pH values in Ireland and Britain (Table 1; Grime et al. 1988) and in Poland on woodland soils with a pH ranging from 3.5 to 7.0 (Czarnecka 1986). This may indicate the presence of locally adapted edaphic populations. Soils from woodland sites, limestone pavement and walls in Ireland and Britain had a variable range of mineral nutrient contents, although they were generally base-rich (Table 1). Czarnecka (1986) reports M. muralis from woodland soils in Poland with 0–14% humus or organic matter and with variable mineral nutrient contents. Mycelis muralis occurs mainly in beech or related woodland types (class Querco–Fagetea) and in woodland clearings (class Epilobietea angustifolii) and margins (class Artemisetea vulgaris) in Europe (Oberdorfer 1978, 1983; Ellenberg 1988). It also occurs less frequently on basic rocks and on walls (class Asplenietea rupestria) (Oberdorfer 1977; Ellenberg 1988; Clabby & Osborne 1994; A.J.C. Malloch, personal communication). The following account details occurrences in woodland and related vegetation types and then addresses other communities in which M. muralis is found. In Ireland M. muralis occurs in woodland communities in the Corylo–Fraxinetum (Clabby & Osborne 1994) in the alliance Alno–Ulmion and the order Fagetalia (sensuEllenberg 1988). The Corylo–Fraxinetum corresponds broadly to the Dryopterido–Fraxinetum phyllitidetosum of Klötzli (1970), which corresponds in part with the Geranium robertianum subcommunity of the British Fraxinus excelsior–Acer campestre–Mercurialis perennis woodlands (NVC code W8) (Rodwell 1991). In Britain there are two woodland types on base-rich calcareous soils in which M. muralis occurs frequently: the Fraxinus–Acer–Mercurialis woodlands (NVC W8) and the Fagus sylvatica–Mercurialis perennis woodlands (NVC W12) (Rodwell 1991). In W8 woodlands M. muralis occurs most commonly with low frequency and with low cover values in the Hedera helix subcommunity (W8d), the Geranium robertianum subcommunity (W8e) and the Teucrium scorodonia subcommunity (W8g). Constant species in W8 woodlands include Acer campestre, Corylus avellana, Eurhynchium praelongum, Fraxinus excelsior, Mercurialis perennis and Rubus fruticosus agg. Other species common in the field layer include Circaea lutetiana, Dryopteris filix-mas, Geum urbanum and Hyacinthoides non-scripta, and in the ground layer Brachythecium rutabulum and Plagiomnium undulatum. In the Yorkshire dales the Geranium robertianum subcommunity occurs in grikes in the limestone pavement with a greater range of ferns, including Dryopteris submontana (D. villarii) and Polystichum lonchitis (Rodwell 1991). In W12 woodlands, M. muralis occurs in the Mercurialis perennis subcommunity (W12a), the Sanicula europaea subcommunity (W12b) and the Taxus baccata subcommunity (W12c), all of which occur in southern England. The constant species in W12 woodlands are Fagus sylvatica and Mercurialis perennis. The field layer is qualitatively similar to the field layer of the W8 woodlands and frequently includes species such as Circaea lutetiana, Geum urbanum and Sanicula europaea. Bryophytes are infrequent, with the most common being Brachythecium rutabulum and Eurhynchium praelongum. In phytosociological terms these woodland communities could be included in the alliances Carpinion betuli (W8), Tilio–Acerion (W8) or Fagion sylvaticae (W12), in the order Fagetalia. Mycelis muralis also occurs infrequently in the Hedera helix–Urtica dioica and the Viburnum lantana subcommunities of Crataegus monogyna–Hedera helix scrub (NVC W21a and W21d, respectively), a community of neutral to base-rich soils common throughout lowland Britain. It also occurs occasionally in communities that develop over base-poor and often moist brown soils, such as the typical subcommunity of Fraxinus excelsior–Sorbus aucuparia–Mercurialis perennis woodland (NVC W9a), the Hedera helix subcommunity of Quercus robur–Pteridium aquilinum–Rubus fruticosus woodland (NVC W10c), and the Blechnum spicant subcommunity of Quercus petraea–Betula pubescens–Oxalis acetosella woodland (NVC W11b). In continental Europe M. muralis occurs most often in communities in the order Fagetalia and the alliance Fagion sylvaticae, although it also occurs in the alliances Tilio–Acerion and Carpinion betuli (Ellenberg 1988). In the Fagion, Ellenberg (1988) distinguishes four suballiances, all of which contain associations in which M. muralis occurs. These include montane to submontane pure beech woods in the Luzulo–Fagion (association Luzulo–Fagetum), which are found from Switzerland to southern Sweden and from the Ardennes to the Carpathians. In the Galio odorati–Fagion M. muralis occurs in several associations including the Galio odorati–Fagetum (or montane limestone fir beech woods) typical of the Vosges, the Sudeten mountains, the Bavarian forest, the eastern Black forest, the north-west Swabian Jura, the Alps and the mountains of Croatia and Slovenia, the Dentario glandulosae–Fagetum in the Polish Carpathians (Zarzycki 1976; Czarnecka 1986), and the Melico–Fagetum in central Europe (Ellenberg 1988). In the Polish Carpathians M. muralis also occurs in the Carici–Fagetum (Zarzycki 1976) or dry-slope beech woods in the suballiance Cephalanthero–Fagion. Mycelis muralis is also found in silver-fir (Abies alba) woods, which occur extensively on the Balkan peninsula and the Swiss Alps and are assigned to the suballiance Galio rotundifolii–Abietion (Ellenberg 1988). Typical associations include the Querco–Abietetum in the outer Alps and the Abietetum albae, typical of the transition zone between the northern central Alps and the dry central valleys (Ellenberg 1988). A recent account of Austrian plant communities (Mucina et al. 1993) also lists M. muralis among the dominant and constant companion species in the ‘diagnostic species combination’ for many associations in the alliance Fagion sylvaticae. In the alliance Tilio–Acerion, M. muralis is found in the Phyllitido–Aceretum in Poland (Zarzycki 1976) and in the Scolopendrio–Fraxinetum, the Acer–Carpinetum and the Asperulo taurii–Tilietum in Austria (Mucina et al. 1993), and in the association Tilio– Carpinetum in the alliance Carpinion betuli (Czarnecka 1986). In Austria, Mucina et al. (1993) distinguish another alliance, the Aremonio–Fagion, and list M. muralis in the Lamio orvalae–Fagetum, the Anemono trifoliae–Fagetum and the Polysticho lonchitis–Fagetum. In the montane spruce woods of the central Swiss Alps, on more acidic soils, M. muralis occurs in the Galio–Abietetum, which may be assigned to the order Fagetalia and in the Melico–Piceetum, an association of dry continental valleys in the order Vaccinio–Piceetalia (class Vaccinio–Piceetea) (Ellenberg 1988). Soldatenkova (1982) described M. muralis from a spruce grove (Piceetum–Hylocomiosum) near Moscow and M. muralis also occurs in the Vaccinio–Piceetea in the Polish Carpathians in associations such as the Querco roboris–Pinetum and Abietetum polonicum (Zarzycki 1976; Czarnecka 1986). Mycelis muralis is common in many woodland clearing communities in the class Epilobietea angustifolii, in the alliances Epilobion angustifolii, Atropion and the Sambuco–Salicion capreae (Oberdorfer 1978). Mycelis muralis is also frequent in the closely related woodland margin communities in the alliance Alliarion (of which it is a character species) in the class Artemisietea vulgaris, and is particularly common in the association Epilobio–Geranietum robertiani that occurs at woodland edges, around rocks, walls and at the entrances to caves (Oberdorfer 1983; Mucina et al. 1993). In coastal British Columbia, Canada, M. muralis often occurs in similar communities in open forest, forest edges and on cut-over or burnt sites with species such as Achlys triphylla (Sm.) DC., Chamerion angustifolium, Kindbergia oregana (Sullivant) Ochyra, Polystichum munitum and Tiarella trifoliata L. (Klinka et al. 1989). Similarly, in the Barkley Sound region of Vancouver Island M. muralis occurs in open habitats on small non-forested islands and around the periphery of larger forested islands (Cody et al. 1996). Mycelis muralis also occurs in rock crevice and wall communities in the class Asplenietea rupestria. Oberdorfer (1977) lists M. muralis in three rock crevice communities, the Drabo–Hieracietum humilis in the Schwäbische Alb, the Cardaminopsietum petraea in the Franconian Jura and the Gymnocarpietum robertiani, a pioneer community of moist calcareous screes in the Jura. This community also occurs locally in Britain (NVC OV38, Gymnocarpium robertianum–Arrhenatherum elatius community) in patchworks of grassland, scrub and woodland on screes and rocky limestone slopes (A.J.C. Malloch, personal communication). These communities are often placed in the order Potentilletalia caulescentis and the alliance Potentillion caulescentis, which contain vegetation of limestone rocks and walls (Oberdorfer 1977; Ellenberg 1988). A recent treatment places the Gymnocarpietum robertiani in the alliance Stipion calamagrostis, which includes fern assemblages of base-rich screes in the submontane and montane zones of central Europe and Scandinavia (Jarolímek et al. 1997). In Britain, in situations where there is no grazing, OV38 is often quickly replaced by scrub or woodland, although M. muralis and other species such as Geranium robertianum, Mercurialis perennis and Viola riviniana can persist in abundance under the trees (A.J.C. Malloch, personal communication). The community is more frequent on rocky limestone slopes and outcrops where woodland development is patchy and it may sometimes be found in the Teucrium subcommunity of the Fraxinus–Acer–Mercurialis woodland (NVC W8g). On such rocky limestone slopes OV38 may often be a secondary community following woodland clearance, and the Asplenium trichomanes–A. ruta-muraria community (NVC OV39) may also occur in the vegetation mosaic. Mycelis muralis is also found infrequently in this community, which is characteristic of sunny crevices in lime-rich bedrocks and walls at low to moderate altitudes, occurring particularly in western Britain (A.J.C. Malloch, personal communication). OV39 corresponds to the association Asplenietum trichomano–rutae–murariae and in Ireland M. muralis occurs commonly in this community on walls with species such as Asplenium ruta-muraria, A. trichomanes, Cymbalaria muralis and Homalothecium sericeum (Clabby & Osborne 1994). On fully exposed limestone pavement in the Burren, Co. Clare, Ireland, M. muralis occurs with species such as A. ruta-muraria, Ceterach officinarum, Geranium robertianum, Hedera helix, Prunus spinosa, Senecio jacobaea, Sesleria caerulea, Teucrium scorodonia and Thymus polytrichus. Ivimey-Cook & Proctor (1966) refer to this community as the Ceterach officinarum–Asplenium trichomanes association, which probably corresponds to the association Asplenietum trichomano–rutae–murariae (White & Doyle 1982). The Asplenietum has generally been placed in the alliance Potentillion caulescentis, although some authors follow Segal (1969) and place it in the Cymbalario–Asplenion (White & Doyle 1982). In the Burren, M. muralis also occurs in grikes with species such as Asplenium ruta-muraria, Geranium robertianum and Phyllitis scolopendrium (Osborne & Clabby 1991) and in Yorkshire with a range of species including Allium ursinum, Asplenium trichomanes, Epilobium montanum, Geranium robertianum, Mercurialis perennis, Oxalis acetosella, Phyllitis scolopendrium, Stachys sylvatica and Urtica dioica (Silvertown 1983). In shaded crevices and on ledges of lime-rich bedrocks in western and northern Britain, M. muralis occurs in the Asplenium viride–Cystopteris fragilis community (NVC OV40) (A.J.C. Malloch, personal communication). This community corresponds to the Asplenio–Cystopteridetum, which has generally been placed in the alliance Cystopteridion fragilis (Ellenberg 1988). Segal (1969) placed similar vegetation from walls in the Austrian and French mountains in a new association, the Polysticho lonchitis–Asplenietum viridis in the alliance Cymbalario–Asplenion, which has been followed by White & Doyle (1982) although M. muralis has not been recorded from this vegetation type in Ireland. Mycelis muralis is also found on walls dominated by Hedera helix, Sambucus nigra and Stachys sylvatica in Ireland and at wall bases, with species such as Acer pseudoplatanus, Circaea lutetiana, Fraxinus excelsior, Geranium robertianum, Hedera helix, Heracleum sphondylium, Poa annua, Sambucus nigra and Urtica dioica (Clabby & Osborne 1994). In Britain, M. muralis occurs infrequently in a range of other communities, most notably the Arrhenatherum elatius grasslands (NVC MG1) (A.J.C. Malloch, personal communication), which may reflect an ability to colonize disturbed, open sites on base-rich substrates. The exclusion of cattle from a woodland in Durrow, Co. Laois, Ireland, led to an increase in the population size of M. muralis, which suggests that plants may be susceptible to grazing. There was little or no indication of an effect of grazing by large mammals on the depth distribution of M. muralis in grikes in limestone pavement in Yorkshire (Silvertown 1982, 1983). In the Burren, Co. Clare, Ireland, there is no evidence of cattle or goat grazing of M. muralis. In high-light habitats large concentrations of anthocyanins in leaves and stems may possibly discourage grazing animals. In wall habitats grazing by large mammals is effectively avoided owing to location. There is no information on the extent of insect herbivory on M. muralis. There is no experimental evidence on the competitive ability of M. muralis. In high-light habitats M. muralis almost always occurs as a component of a sparse vegetation cover and rarely in a closed sward. Its occurrence ranges from weedy, disturbed, ephemeral habitats (Grime et al. 1988; Clabby & Osborne 1994; Cody & Overton 1996), to a presence in long established, stable, open habitats such as old walls or limestone pavement with many available microsites for colonization (Clabby & Osborne 1994). In many shaded locations M. muralis is often found in sparse vegetation, for example on walls or in disturbed microsites in woodland/scrub or along woodland paths and rides. Mycelis muralis also occurs in closed woodland floor vegetation where it may compete with co-occurring species (Clabby & Osborne 1991). In spruce forest in Waldviertel, Lower Austria, the addition of fertilizer and lime, combined with the removal of litter, led to the appearance of M. muralis in the ground layer with other species including Athyrium filix-femina, Polygonatum verticillatum and Urtica dioica (Leitgeb 1994). In the Hanoverian Münden, in an experiment in which beech woodland (Luzulo–Fagetum) plots were fertilized with various combinations of calcium, potassium and phosphorus, or ‘cultivated’ prior to the addition of nutrients, M. muralis appeared with a range of herbs typical of the more nutrient-rich Melico–Fagetum (Grabherr 1942 cited by Ellenberg 1988). Ellenberg (1988) also describes an experiment in which M. muralis appeared (with Galeopsis pubescens Besser and Galium odoratum) in the ground flora of a typical herb-free beech wood (Fagetum nudum in the Cephalanthero–Fagion) in response to the cutting of beech roots in the upper soil layers. As these woods are very dry the appearance of M. muralis may have been due to increased water or nutrient availability as a result of root cutting and subsequent root decomposition (Ellenberg 1988). Mycelis muralis usually occurs as scattered individuals or in small groups (generally two or three plants). It does not form patches of appreciable size. Cover values for M. muralis in many types of vegetation are low, often being less than 10% (Zarzycki 1976; Petrov 1981; Czarnecka 1986; Ellenberg 1988; Rodwell 1991; Clabby & Osborne 1994). In a range of habitats in Britain M. muralis was most frequently found in < 10% of 100-cm2 subsections of a 1-m2 quadrat, although in woodland it occurred infrequently in up to 40% of 100 cm2 subsections (Grime et al. 1988). In Ireland numbers of plants per 1-m2 quadrat ranged from 1 to 16 in woodland. Vegetative reproduction does not take place. Height of flowering individuals typically ranges from 300 to 600 mm on walls in the open, or on limestone pavement. In shaded locations plant height can be up to 1300 mm. In a woodland site at Durrow, Co. Laois, Ireland, plant height at maturity averaged 1010 mm ± 80 SD (n = 10; includes flowering panicle). In Ireland plants flower and set seed in a variety of habitats including wall tops, wall faces, wall bases, wall embankments, waste ground, scrub/ woodland, limestone pavement and grikes in limestone pavement. Non-flowering rosettes are typically less than 400 mm in height (Grime et al. 1988). An examination of seasonal performance of M. muralis growing on open limestone pavement and in woodland showed that more biomass was produced at the exposed site (Clabby & Osborne 1994). Peak biomass of flowering individuals occurred in July on the open pavement (2.20 g ± 0.61 SE) and in August at the woodland site (0.64 g ± 0.18 SE). Similar data were obtained in laboratory experiments (Clabby & Osborne 1997). Allocation of plant biomass also varied from habitat to habitat. Woodland plants typically showed greater root investment than plants from open limestone pavement or grikes (Clabby & Osborne 1994). Total flowering panicle biomass (capitula + support structures) accounted for up to 55% of total biomass in plants from open pavement but never more than 20% in woodland plants. In contrast to the large differences in plant biomass and total flowering panicle biomass between open and shaded sites, there were much smaller differences in the biomass of capitula, indicating that shading had a relatively small effect on reproductive potential (Clabby & Osborne 1994). Plants do not appear to be vulnerable to frosts in Ireland. Despite a southerly distribution in Britain the continental European distribution of M. muralis suggests that low winter temperatures do not limit its distribution. Grime et al. (1988) suggest that M. muralis is a drought-avoiding species but there is also evidence for drought resistance. In a laboratory experiment withholding water until plant pots were reduced to 70% of their initial fully watered weight did not affect plant biomass over a 12-week period (Manto 1995). Studies in the Burren, Co. Clare, Ireland, indicate that M. muralis may be more drought-resistant than some co-occurring species (Osborne & Clabby 1991). In open habitats the leaf lobes often curl inwards in periods of dry weather during the summer months. This may reduce excessive water loss and prevent high-temperature and high-light induced damage. In open habitats such as wall tops and limestone pavement, wind can result in damage to the plant when the flowering panicle has developed. Plants often respond by producing a second flowering panicle. Leaves have four to five variably dissected lobes of unequal size. The terminal lobes are generally larger, broadly triangular or hastate (Fig. 3). Lower lobes are hastate or rhombic. A flange connects individual lobes; flange development is greatest in leaves from flowering plants. Leaves developed at high light have more indented margins, with pointed tips and smaller terminal lobes. The largest lower lobes are found in flowering plants (Fig. 3b,d). Upper leaves tend to be sessile, auriculate and less divided, becoming progressively smaller upwards. Leaf shapes exist that are intermediate between the examples shown. Variability in lower leaf shape from mature plants growing in exposed (a, b) and shaded (c, d) conditions. Leaves shown are from vegetative (a, c) and flowering (b, d) plants. Exposed plants collected from the open limestone pavement of the Burren, Co. Clare, Ireland, and shaded plants from a woodland in Durrow, Co. Laois, Ireland. The number of stomata on the terminal lobes varies with irradiance. Upper leaves of plants from a woodland at Durrow, Co. Laois, Ireland, had 80.7 ± 17.1 SD and 38.8 ± 11.2 SD stomata mm–2 on their lower (abaxial) and upper (adaxial) surfaces, respectively, whereas upper leaves of plants from Mullach Mór, in the Burren, Co. Clare, Ireland had 108.4 ± 14.5 SD and 91.2 ± 22.6 SD stomata mm–2 on their abaxial and adaxial surfaces (n = 3 leaves). The leaf surface is smooth, with the major veins prominent. The epidermal cells are convex, particularly on the adaxial surface. Palisade mesophyll tissue is poorly developed and often comprises only one cell layer. There is little development of sclerenchyma (Osborne et al. 1994). Rooting depth at the Durrow site was 13.75 cm ± 1.71 SD. Roots comprise two to three major axes with a relatively low density of fine roots. Boullard & Dominik (1960) report an ectomycorrhizal infection in M. muralis collected from a woodland site at Babia Góra, Poland. There are also several records of vesicular–arbuscular mycorrhizas occurring on M. muralis in continental Europe and Britain (Harley & Harley 1987). The absence of any mycorrhizal infection has also been reported (Harley & Harley 1987). A search of the MYCOLIT database (Trappe & Castellano 1991) revealed no information on mycorrhizal associations of M. muralis. Mycelis muralis is a herbaceous perennial hemicryptophyte. The plant is wintergreen, dying completely back after flowering in the summer but then producing a small overwintering rosette(s)" @default.
W1976945654 created "2016-06-24" @default.
W1976945654 creator A5007225553 @default.
W1976945654 creator A5029057147 @default.
W1976945654 date "1999-01-01" @default.
W1976945654 modified "2023-09-24" @default.
W1976945654 title "<i>Mycelis muralis</i>(L. ) Dumort. (<i>Lactuca muralis</i>(L. ) Gaertner)" @default.
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