Hostname: page-component-78c5997874-mlc7c Total loading time: 0 Render date: 2024-11-13T11:57:34.113Z Has data issue: false hasContentIssue false
  • English
  • Français

A hypervolume approach to niche specialism, tested for the old-growth indicator status of calicioids

Published online by Cambridge University Press:  13 December 2022

Christopher J. Ellis Open the ORCID record for Christopher J. Ellis [Opens in a new window]
Christopher J. Ellis*
Affiliation:
Royal Botanic Garden Edinburgh, 20A Inverleith Row, Edinburgh EH3 5LR, UK
*
Author for correspondence: Christopher J. Ellis. E-mail: c.ellis@rbge.org.uk
Get access Rights & Permissions [Opens in a new window]

Abstract

Certain lichen epiphytes are restricted to old-growth forest stands with long ‘ecological continuity’, explained by i) niche specialism and their dependence on microhabitats associated with old stands including veteran or senescent trees, and/or ii) dispersal limitation with probabilities of colonization being relaxed over extended time periods. ‘Calicioid’ species are among the most important old-growth indicators, yet they reproduce sexually via small spores that appear widely dispersed at ecological scales. This suggests that they should have a high level of niche specialism compared to lichen epiphytes in general, explaining their role as old-growth indicators. However, comparisons of niche specialism are challenging, and this study uses epiphytic, corticolous calicioid species as an appropriate test case. Having measured 20 variables that constrain the lichen epiphyte niche, these were collapsed into a ‘hypervolume’ representing the sampled environmental space available for occupancy by lichens in Scotland as a study system. It was then possible to examine the occupancy of this hypervolume by individual lichens (niche breadth), with the proportion/percent occupied used to estimate a niche specialism score. Consequently, epiphyte calicioid species are confirmed to have a high degree of niche specialism compared to lichen epiphytes in general, and compared to other old-growth indicators, with their niche position directed towards drier climates including locally sheltered microhabitats associated with old-growth forest structure.

Keywords

niche positionold-growth structurepin-head lichensrealized niche
Type
Standard Paper
Information
The Lichenologist , Volume 54 , Issue 6 , November 2022 , pp. 379 - 387
Check for updates
Copyright
Copyright © The Author(s), 2022. Published by Cambridge University Press on behalf of the British Lichen Society

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

Alam, MA, Gauslaa, Y and Solhaug, KA (2015) Soluble carbohydrates and relative growth rates in chloro-, cyano- and cephalolichens: effects of temperature and nocturnal hydration. New Phytologist 208, 750–762.CrossRefGoogle ScholarPubMed
Angelini, A, Corona, P, Chianucci, F and Portoghesi, L (2015) Structural attributes of stand overstory and light under the canopy. Annals of Silvicultural Research 39, 23–31.Google Scholar
Antoine, ME and McCune, B (2004) Contrasting fundamental and realized ecological niches with epiphytic lichen transplants in an old-growth Pseudotsuga forest. Bryologist 107, 163–173.CrossRefGoogle Scholar
Austin, MP and Smith, TM (1989) A new model for the continuum concept. Vegetatio 83, 35–47.CrossRefGoogle Scholar
Bates, JW (1992) Influence of chemical and site factors on Quercus and Fraxinus epiphytes at Loch Sunart, western Scotland: a multivariate analysis. Journal of Ecology 80, 163–179.CrossRefGoogle Scholar
Belinchón, R, Martínez, I, Otálora, MAG, Aragón, G, Dimas, J and Escudero, A (2009) Fragment quality and matrix affect epiphytic performance in a Mediterranean forest landscape. American Journal of Botany 96, 1974–1982.10.3732/ajb.0900040CrossRefGoogle Scholar
Belinchón, R, Yahr, R and Ellis, CJ (2015) Interactions among species with contrasting dispersal modes explain distributions for epiphytic lichens. Ecography 38, 762–768.CrossRefGoogle Scholar
Belinchón, R, Harrison, PJ, Mair, L, Várkonyi, G and Snäll, T (2017) Local epiphyte establishment and future metapopulation dynamics in landscapes with different spatiotemporal properties. Ecology 98, 741–750.CrossRefGoogle ScholarPubMed
Bidussi, M, Gauslaa, Y and Solhaug, KA (2013) Prolonging hydration and active metabolism from light periods into nights substantially enhances lichen growth. Planta 237, 1359–1366.CrossRefGoogle ScholarPubMed
Blonder, B (2018) Hypervolume concepts in niche- and trait-based ecology. Ecography 41, 1441–1455.CrossRefGoogle Scholar
Boucher, VL and Nash, TH, III (1990) Growth patterns in Ramalina menziesii in California: coastal vs. inland populations. Bryologist 93, 295–302.Google Scholar
Boulangeat, I, Lavergne, S, Van, Es J, Gaurraud, L and Thuiller, W (2012) Niche breadth, rarity and ecological characteristics within a regional flora spanning large environmental gradients. Journal of Biogeography 39, 204–214.CrossRefGoogle Scholar
Braidwood, DW and Ellis, CJ (2012) Bioclimatic equilibrium for lichen distributions on disjunct continental landmasses. Botany 90, 1316–1325.CrossRefGoogle Scholar
Carnes, BA and Slade, NA (1982) Some comments on niche analysis in canonical space. Ecology 63, 888–893.CrossRefGoogle Scholar
Colesie, C, Scheu, S, Green, TGA, Weber, B, Wirth, R and Büdel, B (2012) The advantage of growing on moss: facilitative effects on photosynthetic performance and growth in the cyanobacterial lichen Peltigera rufescens. Oecologia 169, 599–607.CrossRefGoogle ScholarPubMed
Coppins, AM and Coppins, BJ (2002) Indices of Ecological Continuity for Woodland Epiphytic Lichen Habitats in the British Isles. London: British Lichen Society.Google Scholar
Coppins, BJ (1976) Distribution patterns shown by epiphytic lichens in the British Isles. In Brown, DH, Hawksworth, DL and Bailey, RH (eds), Lichenology: Progress and Problems. London: Academic Press, pp. 249–278.Google Scholar
Demmig-Adams, B, Máguas, C, Adams, WW, Meyer, A, Kilian, E and Lange, OL (1990) Effect of high light on the efficiency of photochemical energy conversion in a variety of lichen species with green and blue-green phycobionts. Planta 180, 400–409.CrossRefGoogle Scholar
Dettki, H, Klintberg, P and Esseen, P-A (2000) Are epiphytic lichens in young forests limited by local dispersal? Écoscience 7, 317–325.CrossRefGoogle Scholar
Devictor, V, Clavel, J, Julliard, R, Lavergne, S, Mouillot, D, Thuiller, W, Venail, P, Villéger, S and Mouquet, N (2010) Defining and measuring ecological specialization. Journal of Applied Ecology 47, 15–25.CrossRefGoogle Scholar
Doering, M and Coxson, D (2010) Riparian alder ecosystems as epiphytic lichen refugia in sub-boreal spruce forests of British Columbia. Botany 88, 144–157.CrossRefGoogle Scholar
Dymytrova, L, Brändli, U-B, Ginzler, C and Scheidegger, C (2018) Forest history and epiphytic lichens: testing indicators for assessing forest autochthony in Switzerland. Ecological Indicators 84, 847–857.CrossRefGoogle Scholar
Ellis, CJ (2012) Lichen epiphyte diversity: a species, community and trait-based review. Perspectives in Plant Ecology, Evolution and Systematics 14, 131–152.CrossRefGoogle Scholar
Ellis, CJ (2020) Microclimatic refugia in riparian woodland: a climate change adaptation strategy. Forest Ecology and Management 462, 118006.CrossRefGoogle Scholar
Ellis, CJ and Ellis, SC (2013) Signatures of autogenic succession for an aspen chronosequence. Journal of Vegetation Science 24, 688–701.CrossRefGoogle Scholar
Ellis, CJ, Coppins, BJ, Dawson, TP and Seaward, MRD (2007) Response of British lichens to climate change scenarios: trends and uncertainties in the projected impact for contrasting biogeographic groups. Biological Conservation 140, 217–235.CrossRefGoogle Scholar
Ellis, CJ, Eaton, S, Theodoropoulos, M and Elliott, K (2015) Epiphyte Communities and Indicator Species. An Ecological Guide for Scotland's Woodlands. Edinburgh: Royal Botanic Garden Edinburgh.Google Scholar
Ellis, CJ, Geddes, H, McCheyne, N and Stansfield, A (2017) Lichen epiphyte response to non-analogue monthly climates: a critique of bioclimatic models. Perspectives in Plant Ecology, Evolution and Systematics 25, 45–58.CrossRefGoogle Scholar
Englund, SR, O'Brien, JJ and Clark, DB (2000) Evaluation of digital and film hemispherical photography and spherical densiometry for measuring forest light environments. Canadian Journal of Forest Research 30, 1999–2005.CrossRefGoogle Scholar
Etienne, RS and Alonso, D (2007) Neutral community theory: how stochasticity and dispersal-limitation can explain species coexistence. Journal of Statistical Physics 128, 485–510.10.1007/s10955-006-9163-2CrossRefGoogle Scholar
Fedrowitz, K, Kaasalainen, U and Rikkinen, J (2011) Genotype variability of Nostoc symbionts associated with three epiphytic Nephroma species in a boreal forest landscape. Bryologist 114, 220–230.Google Scholar
Fridley, JD, Vandermast, DB, Kuppinger, DM, Manthey, M and Peet, RK (2007) Co-occurrence based assessment of habitat generalists and specialists: a new approach for the measurement of niche width. Journal of Ecology 95, 707–722.CrossRefGoogle Scholar
Fritz, O and Heilmann-Clausen, J (2010) Rot holes create key microhabitats for epiphytic lichens and bryophytes on beech (Fagus sylvatica). Biological Conservation 143, 1008–1016.CrossRefGoogle Scholar
Gauslaa, Y (1985) The ecology of Lobarion pulmonariae and Parmelion caperatae in Quercus dominated forests in south-west Norway. Lichenologist 17, 117–140.CrossRefGoogle Scholar
Gauslaa, Y and Solhaug, KA (1996) Differences in the susceptibility to light stress between epiphytic lichens of ancient and young boreal forest stands. Functional Ecology 10, 344–354.CrossRefGoogle Scholar
Gauslaa, Y, Palmqvist, K, Solhaug, KA, Holien, H, Hilmo, O, Nybakken, L, Myhre, LC and Ohlson, M (2007) Growth of epiphytic old forest lichens across climatic and successional gradients. Canadian Journal of Forest Research 37, 1832–1845.CrossRefGoogle Scholar
Goward, T (1994) Notes on old growth-dependent epiphytic macrolichens in inland British Columbia, Canada. Acta Botanica Fennica 150, 31–38.Google Scholar
Goward, T and Arsenault, A (2018) Calicioid diversity in humid inland British Columbia may increase into the 5th century after stand initiation. Lichenologist 50, 555–569.CrossRefGoogle Scholar
Gu, W-D, Kuusinen, M, Konttinen, T and Hanski, I (2001) Spatial pattern in the occurrence of the lichen Lobaria pulmonaria in managed and virgin boreal forests. Ecography 24, 139–150.CrossRefGoogle Scholar
Hannah, L, Lohse, D, Hutchinson, C, Carr, JL and Lankerani, A (1994) A preliminary inventory of human disturbance of world ecosystems. Ambio 23, 246–250.Google Scholar
Hannah, L, Carr, JL and Lankerani, A (1995) Human disturbance and natural habitat: a biome level analysis of a global data set. Biodiversity and Conservation 4, 128–155.CrossRefGoogle Scholar
Hanski, I (1999) Metapopulation Ecology. Oxford: Oxford University Press.Google Scholar
Hanski, I (2002) Extinction debt at extinction threshold. Conservation Biology 16, 666–673.CrossRefGoogle Scholar
Holien, H (1996) Influence of site and stand factors on the distribution of crustose lichens of the Caliciales in a suboceanic spruce forest area in central Norway. Lichenologist 28, 315–330.CrossRefGoogle Scholar
Hollis, D, McCarthy, M, Kendon, M, Legg, T and Simpson, I (2019) HadUK-Grid – a new UK dataset of gridded climate observations. Geoscience Data Journal 6, 151–159.Google Scholar
Hubbell, SP (2001) The Unified Neutral Theory of Biodiversity and Biogeography. Princeton: Princeton University Press.Google Scholar
Hutchinson, GE (1957) Concluding remarks. Cold Spring Harbour Symposia on Quantitative Biology 22, 415–427.CrossRefGoogle Scholar
Johansson, NR, Kaasalainen, U and Rikkinen, J (2021) Woodpeckers can act as dispersal vectors for fungi, plants and microorganisms. Ecology and Evolution 11, 7154–7163.CrossRefGoogle ScholarPubMed
Jüriado, I, Liira, J, Paal, J and Suija, A (2009) Tree and stand level variables influencing diversity of lichens on temperate broad-leaved trees in boreo-nemoral floodplain forests. Biodiversity and Conservation 18, 105–125.10.1007/s10531-008-9460-yCrossRefGoogle Scholar
Jüriado, I, Liira, J, Csencsics, D, Widmer, I, Adolf, C, Kohv, K and Scheidegger, C (2011) Dispersal ecology of the endangered woodland lichen Lobaria pulmonaria in managed hemiboreal forest landscape. Biodiversity and Conservation 20, 1803–1819.CrossRefGoogle Scholar
Kenkel, NC and Bradfield, GE (1986) Epiphytic vegetation on Acer macrophyllum: a multivariate study of species-habitat relationships. Vegetatio 68, 43–53.CrossRefGoogle Scholar
Kruys, N and Jonsson, BG (1997) Insular patterns of calicioid lichens in a boreal old-growth forest-wetland mosaic. Ecography 20, 605–613.CrossRefGoogle Scholar
Kubiak, D and Osyczka, P (2020) Non-forested vs forest environments: the effect of habitat conditions on host tree parameters and the occurrence of associated epiphytic lichens. Fungal Ecology 47, 100957.CrossRefGoogle Scholar
Kuusinen, M (1996) Epiphyte flora and diversity on basal trunks of six old-growth forest tree species in southern and middle boreal Finland. Lichenologist 28, 443–463.CrossRefGoogle Scholar
Kuusinen, M and Siitonen, J (1998) Epiphytic lichen diversity in old-growth and managed Picea abies stands in southern Finland. Journal of Vegetation Science 9, 283–292.CrossRefGoogle Scholar
Lemmon, PE (1956) A spherical densiometer for estimating forest overstory density. Forest Science 2, 314–320.Google Scholar
Lewis, JEJ and Ellis, CJ (2010) Taxon- compared with trait-based analysis of epiphytes, and the role of tree species and tree age in community composition. Plant Ecology and Diversity 3, 203–210.CrossRefGoogle Scholar
Lieffers, VJ, Messier, C, Stadt, KJ, Gendron, F and Comeau, PG (1999) Predicting and managing light in the understory of boreal forests. Canadian Journal of Forest Research 29, 796–811.CrossRefGoogle Scholar
Lõhmus, A and Lõhmus, P (2011) Old-forest species: the importance of specific substrata vs. stand continuity in the case of calicioid fungi. Silva Fennica 45, 1015–1039.Google Scholar
Loppi, S and Frati, L (2004) Influence of tree substrate on the diversity of epiphytic lichens: comparison between Tilia platyphyllos and Quecus ilex (central Italy). Bryologist 107, 340–344.CrossRefGoogle Scholar
Malíček, J, Palice, Z, Vondrák, J, Kostovčík, M, Lenzová, V and Hofmeister, J (2019) Lichens in old-growth and managed mountain spruce forests in the Czech Republic: assessment of biodiversity, functional traits and bioindicators. Biodiversity and Conservation 28, 3497–3528.CrossRefGoogle Scholar
Manrique, E, Balaguer, L, Barnes, J and Davison, AW (1993) Photoinhibition studies in lichens using chlorophyll fluorescence analysis. Bryologist 96, 443–449.CrossRefGoogle Scholar
McCune, B (2007) Improved estimates of incident radiation and heat load using non-parametric regression against topographic variables. Journal of Vegetation Science 18, 751–754.CrossRefGoogle Scholar
McCune, B and Grace, JB (2002) Analysis of Ecological Communities. Gleneden Beach, Oregon: MjM Software.Google Scholar
McCune, B and Keon, D (2002) Equations for potential annual direct incident radiation and heat load. Journal of Vegetation Science 13, 603–606.CrossRefGoogle Scholar
McCune, B and Mefford, MJ (2011) PC-ORD. Multivariate Analysis of Ecological Data. Version 6.22. Gleneden Beach, Oregon: MjM Software.Google Scholar
McCune, B, Rosentreter, R, Ponzetti, JM and Shaw, DC (2000) Epiphyte habitats in an old conifer forest in western Washington, U.S.A. Bryologist 103, 417–427.CrossRefGoogle Scholar
Merinero, S, Martínez, I, Rubio-Salcedo, M and Gauslaa, Y (2015) Epiphytic lichen growth in Mediterranean forests: effects of proximity to the ground and reproductive stage. Basic and Applied Ecology 16, 220–230.CrossRefGoogle Scholar
Mežaka, A, Brūmelis, G and Piterāns, A (2012) Tree and stand-scale factors affecting richness and composition of epiphytic bryophytes and lichens in deciduous woodland key habitats. Biodiversity and Conservation 21, 3221–3241.CrossRefGoogle Scholar
Miller, JED, Villella, J, Stone, D and Hardman, A (2020) Using lichen communities as indicators of forest stand age and indicator value. Forest Ecology and Management 475, 118436.CrossRefGoogle Scholar
Mistry, J and Beradi, A (2005) Effects of phorophyte determinants on lichen abundance in the cerrado of central Brazil. Plant Ecology 178, 61–76.CrossRefGoogle Scholar
Nascimbene, J, Marini, L, Motta, R and Nimis, PL (2009) Influence of tree age, tree size and crown structure on lichen communities in mature Alpine spruce forests. Biodiversity and Conservation 18, 1509–1522.CrossRefGoogle Scholar
Nascimbene, J, Marini, L and Nimis, PL (2010) Epiphytic lichen diversity in old-growth and managed Picea abies stands in Alpine spruce forests. Forest Ecology and Management 260, 603–609.CrossRefGoogle Scholar
Nitare, J (2000) Signalarter. Jönköping: Skogsstyrelsens Förlag.Google Scholar
Öckinger, E, Niklasson, M and Nilsson, SG (2005) Is local distribution of the epiphytic lichen Lobaria pulmonaria limited by dispersal capacity or habitat quality? Biodiversity and Conservation 14, 759–773.CrossRefGoogle Scholar
Paletto, A and Tosi, V (2009) Forest canopy cover and canopy closure: comparison of assessment techniques. European Journal of Forest Research 128, 265–272.CrossRefGoogle Scholar
Palmqvist, K and Sundberg, B (2000) Light use efficiency of dry matter gain in five macro-lichens: relative impact of microclimate conditions and species-specific traits. Plant, Cell and Environment 23, 1–14.CrossRefGoogle Scholar
Palmqvist, K, Dahlman, L, Jonsson, A and Nash, TH, III (2010) The carbon economy of lichens. In Nash, TH, III (ed.), Lichen Biology. Cambridge: Cambridge University Press, pp. 184–217.Google Scholar
Prieto, M, Baloch, E, Tehler, A and Wedin, M (2013) Mazaedium evolution in the Ascomycota (Fungi) and the classification of mazaediate groups of formerly unclear relationship. Cladistics 29, 296–308.CrossRefGoogle ScholarPubMed
Pulliam, HR (2000) On the relationship between niche and distribution. Ecology Letters 3, 349–361.CrossRefGoogle Scholar
Quine, CP and White, IMS (1994) Using the relationship between rate of tatter and topographic variables to predict site windiness in upland Britain. Forestry 67, 345–356.CrossRefGoogle Scholar
R Development Core Team (2020) R: a Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna, Austria. [WWW resource] URL https://www.R-project.org.Google Scholar
Rambo, TR (2010) Habitat preferences of an arboreal forage lichen in a Sierra Nevada old-growth mixed-conifer forest. Canadian Journal of Forest Research 40, 1034–1041.CrossRefGoogle Scholar
Rikkinen, J (2003) Calicioid lichens and fungi in the forests and woodlands of western Oregon. Acta Botanica Fennica 175, 1–41.Google Scholar
Rikkinen, J, Oksanen, J and Lohtander, K (2002) Lichen guilds share related cyanobacterial symbionts. Science 297, 357.CrossRefGoogle ScholarPubMed
Roberts, AJ, Russel, C, Walker, GJ and Kirby, KJ (1992) Regional variation in the origin, extent and composition of Scottish woodland. Botanical Journal of Scotland 46, 167–189.CrossRefGoogle Scholar
Rose, F (1974) The epiphytes of oak. In Morris, MG and Perrring, FH (eds), The British Oak. London: E.W. Classey, pp. 250–273.Google Scholar
Rose, F (1976) Lichenological indicators of age and environmental continuity in woodlands. In Brown, DH, Hawksworth, DL and Bailey, RH (eds), Lichenology: Progress and Problems. London and New York: Academic Press, pp. 279–307.Google Scholar
Rubio-Salcedo, M, Merinero, S and Martínez, I (2015) Tree species and microhabitat influence the population structure of the epiphytic lichen Lobaria pulmonaria. Fungal Ecology 18, 1–9.CrossRefGoogle Scholar
Sætersdal, M, Gjerde, I and Blom, H (2005) Indicator species and the problem of spatial inconsistency in nestedness patterns. Biological Conservation 122, 305–316.CrossRefGoogle Scholar
Selva, SB (1994) Lichen diversity and stand continuity in the northern hardwoods and spruce-fir forests of northern New England and western New Brunswick. Bryologist 97, 424–429.CrossRefGoogle Scholar
Selva, SB (2003) Using calicioid lichens and fungi to assess ecological continuity in the Acadian Forest Ecoregion of the Canadian Maritimes. Forestry Chronicle 79, 550–558.CrossRefGoogle Scholar
Sexton, JP, Montiel, J, Shay, JE, Stephens, MR and Slatyer, RA (2017) Evolution of ecological niche breadth. Annual Review of Ecology, Evolution and Systematics 48, 183–206.CrossRefGoogle Scholar
Sillett, SC and McCune, B (1998) Survival and growth of cyanolichen transplants in Douglas-fir forest canopies. Bryologist 101, 20–31.CrossRefGoogle Scholar
Sillett, SC, McCune, B, Peck, JE, Rambo, TR and Ruchty, A (2000) Dispersal limitations of epiphytic lichens result in species dependent on old-growth forests. Ecological Applications 10, 789–799.CrossRefGoogle Scholar
Slatyer, RA, Hirst, M and Sexton, JP (2013) Niche breadth predicts geographical range size: a general ecological pattern. Ecology Letters 16, 1104–1114.CrossRefGoogle Scholar
Smart, SM, Scott, WA, Whitaker, J, Hill, MO, Roy, DB, Critchley, CN, Marini, L, Evans, C, Emmett, BA, Rowe, EC, et al. (2010) Empirical realised niche models for British higher and lower plants – development and preliminary testing. Journal of Vegetation Science 21, 643–656.Google Scholar
Solhaug, KA, Chowdhury, DP and Gauslaa, Y (2018) Short- and long-term freezing effects in a coastal (Lobaria virens) versus a widespread lichen (L. pulmonaria). Cryobiology 82, 124–129.CrossRefGoogle Scholar
Stehn, SE, Nelson, PR, Roland, CA and Jones, JR (2013) Patterns in the occupancy and abundance of the globally rare lichen Erioderma pedicellatum in Denali National Park and Preserve, Alaska. Bryologist 116, 2–14.CrossRefGoogle Scholar
Suárez, J, Gardiner, B and Quine, CP (1999) A comparison of three methods for predicting wind speeds in complex forested terrain. Meteorological Applications 6, 329–342.CrossRefGoogle Scholar
Svensson, M, Caruso, A, Yahr, R, Ellis, C, Thor, G and Snäll, T (2016) Combined observational and experimental data provide limited support for facilitation in lichens. Oikos 125, 278–283.CrossRefGoogle Scholar
Tibell, L (1992) Crustose lichens as indicators of forest continuity in boreal coniferous forests. Nordic Journal of Botany 12, 427–450.CrossRefGoogle Scholar
Tibell, L (1994) Distribution patterns and dispersal strategies of Caliciales. Botanical Journal of the Linnean Society 116, 159–202.CrossRefGoogle Scholar
Tibell, L (2001) Photobiont association and molecular phylogeny of the lichen genus Chaenotheca. Bryologist 104, 191–198.CrossRefGoogle Scholar
Tibell, L and Beck, A (2001) Morphological variation, photobiont association and ITS phylogeny of Chaenotheca phaeocephala and C. subroscida (Coniocybaceae, lichenized Ascomycetes). Nordic Journal of Botany 21, 651–660.10.1111/j.1756-1051.2001.tb00824.xCrossRefGoogle Scholar
Van Dort, K and Horvers, B (2021) Coniocarps: Rainshadow Specialists. Tilburg: KNNV-Afdeling.Google Scholar
Vela Díaz, DM, Blundo, C, Cayola, L, Fuentes, AF, Malizia, LR and Myers, JA (2020) Untangling the importance of niche breadth and niche position as drivers of tree species abundance and occupancy across biogeographic regions. Global Ecology and Biogeography 29, 1542–1553.CrossRefGoogle Scholar
Walser, J-C (2004) Molecular evidence for the limited dispersal of vegetative propagules in the epiphytic lichen Lobaria pulmonaria. American Journal of Botany 91, 1273–1276.CrossRefGoogle ScholarPubMed
Warren, RJ, Bahn, V and Bradford, MA (2012) The interaction between propagule pressure, habitat suitability and density-dependent reproduction in species invasion. Oikos 121, 874–881.CrossRefGoogle Scholar
Wedin, M and Tibell, L (1996) Phytogeny and evolution of Caliciaceae, Mycocaliciaceae, and Sphinctrinaceae (Ascomycota), with notes on the evolution of the prototunicate ascus. Candian Journal of Botany 75, 1236–1242.CrossRefGoogle Scholar
Whittaker, RH (1972) Evolution and measurement of species diversity. Taxon 21, 213–251.CrossRefGoogle Scholar
Whittet, R and Ellis, CJ (2013) Critical tests for lichen indicators of woodland ecological continuity. Biological Conservation 168, 19–23.CrossRefGoogle Scholar
Whittet, R, Hope, J and Ellis, CJ (2015) Open structured woodland and the ecological interpretation of Scotland's Ancient Woodland Inventory. Scottish Geographical Journal 131, 67–77.CrossRefGoogle Scholar
Wiersma, YF and McMullin, RT (2022) Are calicioids useful indicators of boreal forest continuity or condition? Biodiversity and Conservation 31, 1647–1664.CrossRefGoogle Scholar
Wild, M and Gagnon, D (2005) Does lack of available suitable habitat explain the patchy distributions of rare calcicole fern species? Ecography 28, 191–196.CrossRefGoogle Scholar
Williams, L and Ellis, CJ (2018) Ecological constraints to ‘old-growth’ lichen indicators: niche specialism or dispersal limitation? Fungal Ecology 34, 20–27.CrossRefGoogle Scholar
Yahr, R, Vilgalys, R and DePriest, PT (2004) Strong fungal specificity and selectivity for algal symbionts in Florida scrub Cladonia lichens. Molecular Ecology 13, 3367–3378.CrossRefGoogle ScholarPubMed