The Lichenologist 42(1): 83–96 (2010) doi:10.1017/S0024282909008627 © British Lichen Society, 2009 The elevation gradient of lichen species richness in Nepal Chitra Bahadur BANIYA, Torstein SOLHØY, Yngvar GAUSLAA and Michael W. PALMER Abstract: This study of elevation gradients of lichen species richness in Nepal aimed to compare distribution patterns of different life-forms, substratum affinities, photobiont types, and Nepalese endemism. Distribution patterns of lichens were compared with elevational patterns shown by a wide range of taxonomic groups of plants along the Nepalese Himalayan elevational gradient between 200–7400m. We used published data on the elevation records of 525 Nepalese lichen species to interpolate presence between the maximum and minimum recorded elevations, thereby giving estimates of lichen species richness at each 100-m elevational band. The observed patterns were compared with previously published patterns for other taxonomic groups. The total number of lichens as well as the number of endemic species (55 spp.) showed humped relationships with elevation. Their highest richness was observed between 3100–3400 and 4000–4100m, respectively. Almost 33% of the total lichens and 53% of the endemic species occurred above the treeline (>4300m). Non-endemic richness had the same response as the total richness. All growth forms showed a unimodal relationship of richness with elevation, with crustose lichens having a peak at higher elevations (4100–4200m) than fruticose and foliose lichens. Algal and cyanobacterial lichen richness, as well as corticolous lichen richness, all exhibited unimodal patterns, whereas saxicolous and terricolous lichen richness exhibited slightly bimodal relationships with elevation. The highest lichen richness at mid altitudes concurred with the highest diversity of ecological niches in terms of spatial heterogeneity in rainfall, temperature, cloud formation, as well as high phorophyte abundance and diversity implying large variation in bark roughness, moisture retention capacity, and pH. The slightly bimodal distributions of saxicolous and terricolous lichens were depressed at the elevational maximum of corticolous lichens. Key words: altitude, endemism, Himalaya Introduction The Himalayas are the highest mountain range in the world and show a rich diversity of eco-climatic zones. In such a landscape, elevational gradients are of particular interest in applied as well as in theoretical ecology (Wolf 1993). The pattern of changes in species-richness with elevation characterizes C. B. Baniya (corresponding author) and T. Solhøy: Department of Biology, University of Bergen, Allégaten 41, P.O. Box 7803, N-5020 Bergen, Norway. Email: chitra.baniya@bio.uib.no Current address: Central Department of Botany, Tribhuvan University Kirtipur, Kathmandu, Nepal. Y. Gauslaa: Department of Ecology and Natural Resource Management, Norwegian University of Life Sciences, P.O. Box 5003, NO-1432 Ås, Norway. M. W. Palmer: Department of Botany, Oklahoma State University, 104 LSE Stillwater OK 74078 USA 405-744-7717. the vegetation in a simple but powerful way. For example, Yoda (1967) found gradually decreasing tree species richness with increasing altitude. Later, Hunter & Yonzon (1993) reported a similar pattern for Nepalese birds and mammals. A number of studies have produced macro-scale elevational gradients for the whole of Nepal, using interpolation methods on data from secondary sources. Several groups of organisms have been studied and exhibit maximum richness at group-specific intermediate elevations, referred to as a unimodal, or humped, elevation pattern. For example, Himalayan vascular plants showed their highest total richness at altitudes between 1500 to 2500m and their highest endemic richness at 4000m (Grytnes & Vetaas 2002; Vetaas & Grytnes 2002), while fern richness peaked at 2000m (Bhattarai et al. 2004). However, orchids 84 THE LICHENOLOGIST showed their maximum richness at 1600m whereas the highest orchid endemic richness was bimodal with two separate peaks at 1200 and 3300m (Acharya 2008). Liverworts and mosses had their maximum richness at 2800 and 2500m, respectively, and the peak for endemic liverwort richness occurred at 3300m (Grau et al. 2007). Relatively few studies on elevational richness patterns for lichens have been undertaken (Kessler 2000; Bruun et al. 2006; Grytnes et al. 2006) and among terrestrial photosynthetic organisms they are the only major group that has not yet been investigated in this respect in Nepal. They are among the most successful organisms in extreme environments such as cold arctic and alpine environments where few other plants can grow (e.g. Schroeter et al. 1994; Kappen et al. 1996). Lichens also show a high diversity as eipiphytes and thus may benefit from high diversity of trees and shrubs with species-specific bark chemistry, texture and stability (Kessler 2000; Bruun et al. 2006; Grytnes et al. 2006). This paper describes and interprets the Himalayan elevational distribution of lichen species richness in Nepal. Material and Methods Study area Nepal is a mountainous country in the central Himalayas between 26°22# to 30°27# N and 80°04# to 88°12# E with an area of 147 181 km2. The elevation ranges from 60m above sea level to 8848m (Mt. Everest). The country is surrounded by the Indo-Gangetic Plain in the east, west and south, and the Tibetan Plateau towards the north. Parallel mountain ranges run southeast and north-west dissecting the country into northsouth running river valleys that are associated with fertile plains, inner valleys, lakes, and the outer flood plains. The Indian monsoon beginning in May and ending in September is the main source of precipitation (more than 80%). Monsoon rain forms after condensation of water vapour arising from the Bay of Bengal, and consequently there is a spatial gradient of monsoon rainfall decreasing from east to west. Precipitation also occurs in winter formed from central Asian water vapour and diminishes eastwards. Physiographically, Nepal has been divided into five zones (Hagen 1969; Upreti 1999). 1) A 30–40km wide southernmost plain is called Terai between 60 and 300m. It is the most fertile, densely populated zone with a tropical climate (between 22–27°C during winter and above Vol. 42 35°C during the warmest summer months); it is mainly cultivated but has still some tropical forests. 2) The Siwalik located from 700 to 1500m in the foothills of the Mahabharat Range. 3) The Mahabharat Range lies between 1500 and 3000m; it belongs to the Midlands or lesser Himalaya and includes the principal inner valleys such as Kathmandu and Pokhara. The climate is mild throughout the year and receives the highest annual rainfall (above 4000mm) south of Annapurna, central Nepal. It includes the lower cloud zone at 2000 to 2500m (Miehe 1989, 1990). 4) The Greater-Himalaya extends above 2700 m. Here the climate varies from dry summers to cold winters with snow, and subzero temperatures on the mountain tops. This zone includes the uppermost limit of cloud-zone forest in Nepal at 4000m (Miehe 1989, 1990). 5) Trans-Himalaya lies towards the northern side of the Greater-Himalaya with elevations above 3000m. It is in the rain-shadow zone with an average annual rainfall between 200 and 400mm. Vegetation is one of the prime determinants of lichen diversity and distribution (Awasthi 2007). In Nepal, vegetation has been studied and classified into 8 elevational forest zones (Yoda 1967; Stainton 1972, 2001; Dobremez & Jest 1969; Dobremez 1976; Miehe 1982, 1989). 1) Tropical forest between 60 and 1000 m (Stainton 1972) dominated by Shorea robusta Gaertn., Dalbergia sissoo Roxb., Adina cordifolia (Willd. ex Roxb.) Benth. & Hook. f. ex Brandis, Terminalia spp., Lagerstroemia spp., Michelia champaca L., and Bombax ceiba L. 2) Subtropical forest occurring between 1000 to 2000m with Pinus roxburghii Sarg. as the dominant species on south-facing slopes in the Midlands. 3) Lower temperate broad-leaved forest at 1700–2400m in the east and 2000–2700m in the west. Dominant trees include Alnus nitida (Spach.) Endel., Castanopsis tribuloides A. DC., Castanopsis hystrix A. DC., Lithocarpus pachyphylla (Kurz.) Rehder., and Quercus spp. Moister slopes between 1700–2200m are dominated by Cinnamomum spp. and represent the lower temperate mixed broadleaved forest. Upper temperate broad-leaved forests are represented by various types. Drier southern slopes at 2200–3000m are dominated by Schima wallichii (DC.) Korth., Castanopsis indica (Roxb.) Miq., and Quercus semecarpifolia J. E. Smith. Moist, north and west slopes between 2500–3500m from central to east Nepal are dominated by Acer spp., Rhododendron spp., Aesculus spp. and Juglans spp. 4)Temperate coniferous forests between 2000 and 3700m dominated by Pinus wallichiana A. B. Jack. Other conifers include Cedrus deodara (Roxb.) G. Don, Picea smithiana (Wall.) Boiss., Juniperus indica Bertol., Larix himalaica W. C. Cheng & L. K. Fu, Larix griffithiana Carriere, Cupressus torulosa D. Don, and Tsuga dumosa (D. Don) Eichler. 5) Sub-alpine forests occurring between 3000 and 4100m (Stainton 1972) dominated by Betula utilis D. Don, Abies spectabilis (D. Don) Mirb., and Rhododendron spp. This forest includes the climatic treeline at 4100–4300m. 6) Open, low-alpine shrub communities including Caragana spp., Lonicera spp., Rosa spp., Sophora spp., Rhododendron anthopogon D. Don, R. lepidotum Wall. ex G. Don, Ephedra gerardiana Wall. ex C. A. Meyer, and Hippophae tibetana Schldl. 7) Mid-alpine zone with dwarf-shrubs 2010 Elevation gradients of lichens in Nepal—Baniya et al. but largely dominated by herb communities. 8) Nival zone begins above the mid-alpine zone with more lichens and fewer mosses and angiosperms. Within the nival zone, there are vascular plants of open wind exposed areas as well as those tolerant of snow-lie. Data sources The main source of data for this study is the Lichens of Nepal (Sharma 1995). However, many species reported in Sharma (1995) lack elevation data and since then many new species have been described. Other relevant and easily accessible literature as well as the recent keys for both microlichens and macrolichens of Nepal, India and Sri Lanka by Awasthi (1991; 2007) have also been checked. In addition, the following literature on Nepalese lichens (Lamb 1966; Poelt 1966a, b, 1974; Bystrek 1969; Poelt & Reddi 1969; Abbayes 1974; Jahns & Seelen 1974; Kurokawa 1974; Mitchell 1974; Schmidt 1974; Vězda & Poelt 1975; Hellmich & Poelt 1977; Awasthi 1986; Vitikainen 1986; Awasthi & Mathur 1987; Miehe 1990; Awasthi 1991, 2007; Esslinger & Poelt 1991; Poelt & Hinteregger 1993; Sharma 1995; Baniya 1996; Jørgensen 2001) have provided information on the elevation ranges of Nepalese lichens. These sources have provided elevational distribution data for 525 species, including 55 Nepalese endemics (Appendix 1). Among the 525 species, 172, including 29 endemics, are reported from above the treeline (> 4300m). In addition, data about life-forms (crustose, foliose and fruticose), photobiont types (cyanobacteria and green algae) and five substratum types (tree bark, wood, mosses, rocks and soil) were obtained. Tripartite lichens were treated as cyanobacterial. There were 50 species of lichens for which elevational data or other important information are either incomplete or lacking; these species were excluded from the analysis. Lignicolous and muscicolous substrata had been reported for a few species (5 and 17, respectively) and were thus also excluded during the analyses involving substrata. Lichen nomenclature follows Awasthi (1991; 2007). The altitudinal range of lichens in Nepal, 200– 7400m, was divided into 73 bands of 100m each and a complete data matrix for all species was assembled. Presence of a species indicates that the species occurs in, or has been collected in the past, from that elevation band and absence means either that the species does not occur or has previously not been collected from that elevation. A species is assumed to be present in all possible 100m bands between its upper and lower elevation limits. For example, a lichen that has elevational occurrences between 210 to 451m in the literature falls between the 200- and 500-m bands. A list of the lichens included in this study together with their elevation ranges is provided in Appendix 1. Species richness is an estimate of the total number of lichen species occurring in each 100m elevation band. This is a macro-scale study (gamma diversity, sensu Whittaker 1972) that covers the entire elevational range of Nepal. Fifty-six taxa, varieties, forms or subspecies, were treated as species (Appendix 1). 85 Data analysis Patterns related to total lichen species richness and their sub-sets (life forms, algal partners, and substrata) as responses and their elevations as a predictor variable were extracted by using a cubic smooth spline (s) within the framework of Generalized Additive Models (GAM, Hastie & Tibshirani 1990; Heegaard 2004) with a default of c. 4 degrees of freedom. Response variables are counts; thus, the variance changes with the mean and negative predictions are meaningless. We found overdispersion in our data. We thus applied a Quasi-poisson family error distribution with a logarithmic link function (Crawley 2006). We confirmed our assumption of normal distribution of error after the Q-Q diagnostic plots plotted against residuals. The change in deviance follows the F-distribution. We used R 2.7.0 (R Development Core Team 2008) to analyse our data and smoothers were fitted with library GAM (Hastie & Tibshirani 1990). GAM was used because it is a non-parametric approach that does not make a priori assumptions about the species-elevational relationship. One of the main purposes of this study is to contrast observed lichen richness patterns with other studies. Biogeographic data have biases and constraints, such as a focus on specific taxa and/or a restriction to easily accessible landscapes. However, these problems are shared among data sets with various taxonomic groups (Grytnes & Vetaas 2002; Vetaas & Grytnes 2002; Bhattarai et al. 2004; Grau et al. 2007). Thus, a comparison of empirical patterns among available studies is possible. Results The 525 lichen species recorded (Appendix 1) represented 40 families and 121 genera. Among these lichens, 35·4% were crustose, 46·3% were foliose and 18·3% were fruticose. A total of 12% had a cyanobacterial photobiont and 88% had a green algal photobiont. Lichens endemic to Nepal (n=55) represented 10% of the total Nepalese lichen flora. There were 172 species reported from above the treeline (R 4300m) and 29 of these were endemics. Carbonea vorticosa in Nepalese Himalaya at 7400m was the world’s highest reported lichen. Heterodermia pseudospeciosa represented one of the lichens occurring at the lowest elevation ranges in Nepal (150–2100m). The total lichen species richness showed a unimodal relationship with elevation. The maximum modelled total richness occurred at 3100–3400m (Fig. 1A), whereas the observed maximum richness (144 species) 86 THE LICHENOLOGIST Vol. 42 F. 1. Relationship between elevation and lichen species richness in Nepal. A, total lichen species richness; B, total endemic lichen species richness. The fitted regression lines represent statistically significant (P % 0·001) smooth spline (s) after using GAM with approximately 4 degrees of freedom. was at approximately 4000m. Endemic lichens also had a unimodal relationship with elevation, but their maximum modelled richness occurred at 3900–4400 m (Fig. 1B) approximately 800m higher than the peak for total species. Non-endemic richness matched closely the elevational patterns of the total species richness (data not shown). Species richness of all three growth forms of lichens had unimodal responses to elevation (Fig. 2A–C). Among the three morphological types, crustose lichens peaked at the highest altitude (4100–4200m) while foliose lichens had their maximum richness at consistently lower altitudes (2400–2500m). Fruticose lichens peaked at an intermediate elevation (3200m). With respect to photobiont type, cyanolichens exhibited maximum richness at lower altitudes (2900–3000m; Fig. 3A) than green algal lichens (3300– 3500m; Fig. 3B). Among the five substratum categories studied, corticolous lichens had a clear unimodal relationship to elevation, with the highest species richness occurring between 2500 and 2700m (Fig. 4A) in the lower part of the temperate forests. Saxicolous and terricolous lichens had slightly bimodal patterns F. 2. Relationship between elevation and lichen species life-form richness in Nepal. A, foliose lichen species richness; B, fruticose lichen species richness; C, crustose lichen species richness. The fitted regression lines represent statistically significant (P % 0·001) smooth spline (s) after using GAM with approximately 4 degrees of freedom. with a prominent hump at 3900–4200m (Fig. 4B & C). The regression analysis results showing the best selected model for each response variable are recorded in Appendix 2. Discussion Total lichen species richness in Nepal varies strongly with elevation (Figs 1A & B) in line with previous findings for vascular plants (Grytnes & Vetaas 2002; Vetaas & Grytnes 2002; Bhattarai & Vetaas 2003; Bhattarai et al. 2004) including orchids (Acharya 2008), and for bryophytes (Grau et al. 2007). In all these studies, as well as studies dealing with lichens from other countries (Wolseley & Aguirre-Hudson 1997; Negi 2000, 2003; Wolf & Alejandro 2003; Pinokiyo et al. 2010 Elevation gradients of lichens in Nepal—Baniya et al. 87 F. 3. The relationship between elevation and lichen richness for lichens with different photobionts. A, cyanolichen species richness; B, green algal lichen species richness. The fitted regression lines represent statistically significant (P % 0·001) smooth splines (s) after using GAM with approximately 4 degrees of freedom. 2008), species richness tends to peak at intermediate altitudes. The various major taxonomic groups exhibit maximum richness at different altitudes: vascular plants (1500 to 2500m; orchids 1600m, ferns 2000m), liverworts 1800m, mosses 2500m and as high as 3100 to 3400m for lichens. Thus, total richness for lichens occurs at higher elevations than for any of the other groups studied. The total lichen species richness in the Himalayas peaked in the upper cloud-forest zone (following the vegetation classification of Hagen 1969; as modified by Miehe 1982; 1989). This is the zone of the highest rainfall on southernmost slopes (> 4000mm), and the lowest rainfall (< 300mm) on northernmost slopes where the Himalayan föhn causes local drying (Miehe 1989). The zone with maximum lichen richness represents the temperate zone with extremely large local variations in water availability and the accompanying gradients in vegetation cover. According to Bhattarai et al. (2004), the high annual rainfall (> 4000mm) and cool summer temperatures (14–17°C) in this zone are likely to favour forests.The large number of temperate broad-leaved and coniferous trees F. 4. The relationship between elevation and lichens preferring specific substrata. A, corticolous lichen species richness; B, saxicolous lichen species richness; C, terricolous lichen species richness. The fitted regression lines represent statistically significant (P % 0·001) smooth splines (s) after using GAM with approximately 4 degrees of freedom. with bark differing in roughness, moisture retention capacity and pH present a wide variety of habitats for lichens. The slightly bimodal peaks for saxicolous (Fig. 4B) and terricolous lichens (Fig. 4C) tend to show the lower peak in species richness occurring at the same altitude at which corticolous species reach their maximum richness (Fig. 4A). These patterns may be determined by habitat availability and specificity. Corticolous lichens apparently peak at elevations with a high abundance of forests, implying a reduced occurrence of natural and well-lit terricolous and saxicolous habitats. Fruticose lichens have their maximum richness at a substantially higher altitude than the foliose lichens (Figs 2A & B). A similar relationship, but on a very different scale, can be seen within forest canopies 88 THE LICHENOLOGIST where the biomass of fruticose lichens increases and that of foliose lichens decreases with height above the ground (Goward 1998; Campbell & Coxson 2001; Gauslaa et al. 2008). In well-lit and open high altitude forests and the upper canopies of trees, fruticose lichens may have the advantage in being able to utilise light from all directions, whereas many foliose and flat lichens maximize the harvest of more or less unidirectional light in shady positions in dense forests and on low canopy branches (Gauslaa et al. 2009). The foliose lichen richness peaks in the same zone as the highest richness for mosses (Grau et al. 2007), taxa that are also relatively shade tolerant. Furthermore, finely dissected fruticose lichens on twigs lack thick boundary layers. Thus, they are more closely coupled to ambient air than flat foliose lichens and absorb water vapour more readily from the air as discussed by Jonsson et al. (2008). By contrast, flat lichens and bryophyte carpets are more dependent on rainwater. The elevation range between 4000 to 4300m is the sub-alpine zone of Nepal (Stainton 1972). This humid upper-cloud zone (Miehe 1989) is densely colonized by epiphytic mosses reflecting the high humidity (Grau et al. 2007). This humidity may also facilitate high lichen richness associated with open canopies with more light facilitating lichen growth (Gauslaa et al. 2007). A high degree of isolation creates endemism (Cox & Moore 2000), and isolation might increase with elevation. High mountain peaks with an exceptionally cold and harsh climate act as islands (Grau et al. 2007), and phytogeographical isolation can lead to localized speciation events. The occurrence of 29 endemic species (17%) out of the 172 total species above the treeline (R 4300 m) emphasises the link between elevation and endemism. The Nepalese lichen endemism rate is higher than that for mosses (3 out of 480 spp. i.e., < 1%) and liverworts (33 out of 368 spp. i.e., 9%), but slightly less than that for vascular plants (303 out of 1957 spp. i.e., 16%). Above 4500m both total and endemic lichen diversity (Figs 1A & B) decline until 6100m, which is the highest reported elevation for lichens en- Vol. 42 demic to Nepal. Cold winds leading to rapid freezing and drying may reduce competition from surviving vascular plant species (Grime 1977). The harsh climate might lead to various specializations and thus facilitate formation of endemic species (Aptroot & Bungartz 2007). Land area might impose a unimodal richness pattern. In general, the larger the area sampled, the greater will be the number of species encountered (Rosenzweig 1995; Qian et al. 2007). A decreasing trend in land area in each 500m band occurs with increasing elevation (Vetaas & Grytnes 2002) as estimated from the Digitized Map by the International Centre for Integrated Mountain Development (ICIMOD, Kathmandu). This is not always a common pattern (Körner 2007). If we consider species-area relationships, more species should be expected towards lower elevations than we observed. However, lichen diversity can be greatly limited by lack of long ecological continuity in forests (Rose 1976) and lichens may be less prevalent at low altitudes because of dense forest canopies and intensive land-use. Furthermore, as lichen species richness also tends to be universally greater in cool or cold climates (Mattick 1953), we believe that the strong reduction in lichen richness at low elevations is real and not a sampling artefact. In conclusion, lichens and various subgroups of lichens exhibit unimodal patterns similar to those found in other major taxonomic groups, but the highest total lichen richness peaks at higher altitudes than in any other group. The maximum lichen richness occurred at the altitude with the highest diversity of ecological niches in terms of spatial heterogeneity, rainfall, temperature, cloud formation, as well as high phorophyte abundance and diversity implying large variation in bark roughness, moisture retention capacity, and pH. We are grateful to the Norwegian State Education Loan Fund (Lånekassen) for providing funding and the Central Department of Botany, Tribhuvan University, Nepal, for providing study leave for CBB. Thanks are due to John Birks for his valuable comments, Walter Obermayer and Per Magnus Jørgensen for provision of literature on lichens and to Govind Ghimire and Bijaya 2010 Elevation gradients of lichens in Nepal—Baniya et al. Kattel for their feedback. Thanks are also due to Louise Olley, Teuvo Ahti and Tor Tönsberg for their help in revising the species list and to the Senior Editor, Peter Crittenden and two anonymous referees for their useful comments and suggestions. 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Annals of the Missouri Botanical Garden 80: 928–960. Wolf, J. H. D. & Alejandro, F.-S. (2003) Patterns in species richness and distribution of vascular epiphytes in Chiapas, Mexico. Journal of Biogeography 30: 1689–1707. 91 Wolseley, P. A. & Aguirre-Hudson, B. (1997) The ecology and distribution of lichens in tropical deciduous and evergreen forests of northern Thailand. Journal of Biogeography 24: 327–343. Yoda, K. (1967) A preliminary survey of the forest vegetation of eastern Nepal II. General description, structure and floristic composition of sample plots chosen from different vegetation zones. Journal of the College of Arts and Sciences, Chiba University National Science Series 5: 99–140. Accepted for publication 28 March 2009 Appendix 1. Elevation ranges of lichen species of Nepal. Name of lichen species Altitudinal range (m) Alectoria ochroleuca Allocetraria flavonigrescens* A. globulans A. oakesiana A. sinensis A. stracheyi Anthracothecium himalayense A. leucostomum Arctocetraria nigricascens Arctoparmelia subcentrifuga Aspicilia cinerea Awasthia melanotricha* Bacidia millegrana B. nigrofusca B. personata B. rubella B. spadicea Baeomyces pachypus B. roseus Bryonora castanea var. castanea B. castanea var. euryspora B. curvescens B. pulvinar var. microspora* B. pulvinar var. pulvinar* B. pulvinar* B. reducta B. rhypariza var. cyanotropha B. rhypariza var. lamaina B. rhypariza var. rhypariza B. selenospora* B. stipitata* B. yeti Bryoria bicolor B. confusa 4600–5100 4600–4800 3700–4700 3900–4200 4830 3000–5000 1650 2400–2600 3800 4600 4300 4350–4510 1800–3200 3100 300–1800 300 2600 2000–4200 2000–2500 3750 4420 3500–5000 4720–5070 4830–5100 4720–5390 3200 3750–3900 5100–5200 4000–4100 3800–4400 3000–5080 4190–5200 3000–5050 3450 Appendix 1. Continued Name of lichen species Altitudinal range (m) B. furcellata B. himalayana B. implexa B. lactinea B. lanestris B. levis B. nadvornikiana B. nepalensis* B. nitidula B. perspinosa B. poeltii B. smithii B. variabilis Buellia geophila B. granularis B. inornata B. papillata Bulbothrix isidiza B. meizospora B. setschwanensis Calicium abietinum C. lenticulare C. viride Caloplaca arnoldii C. aureosora C. borealis var. borealis C. borealis var. oligosperma C. castellana C. cerina var. cerina C. cerina var. chloroleuca C. cerinopsis* C. cirrochroa C. cirrochroopsis* C. citrina C. cupreobrunnea* 3000 3900–4000 3700–4000 3300–3450 3800–3900 3450 3800–3900 3900–4000 3900 3800–4500 4400 3000–4511 3800–4000 4200 2100 700–900 4650 900–2180 2100–2300 1260–2300 3000–3300 2250 3000–3300 3500–4000 2500–3000 3650 4000 4000–5540 3200 4950–5050 1800–3200 3900–4500 1480 3750 2700 92 THE LICHENOLOGIST Appendix 1. Continued Vol. 42 Appendix 1. Continued Name of lichen species Altitudinal range (m) Name of lichen species C. cupulata* C. epiphyta C. epithallina C. exsecuta var. aphanes* C. farinosa* C. grimmiae C. holocarpa C. holochracea C. insularis C. isabellina* C. leptocheila C. lithophila C. lobulascens* C. lypera* C. maura* C. obliterans C. ochroplaca C. phoenicopta* C. praeruptorum* C. procerispora* C. rinodinopsis C. sancta* C. saxicola var. chamaeleon C. saxicola var. saxicola C. saxifragarum C. tetraspora C. ulcerata* C. variabilis Calvitimela aglaea C. armeniaca Candelaria crawfordii C. sphaerobola Candelariella aurella C. coralliza C. grimmiae C. himalayana C. nepalensis C. sorediosa C. vitellina var. glacialis C. vitellina var. vitellina Canomaculina subsumpta C. subtinctoria Canoparmelia aptata C. ecaperata C. eruptens Carbonea vorticosa Catapyrenium cinereum C. daedaleum Catillaria leptocheiloides Catolechia wahlenbergii Cetraria ambigua C. nepalensis* Cetrelia braunsiana 3200–3800 4400–4500 4350–4450 2060 3000–3500 3500 3700–5000 1700–2300 4600–5540 4100–5200 5000 3200–4850 1500–1800 2000–3000 4950–5000 3200–3700 1800–3000 3200 3200–3650 4100–5000 2000–3000 4000–4340 3200–4000 2900–5000 4950–5000 4400–5000 3500–3750 2000–3500 5000 5000 1300–3000 3000–3900 1600–3500 3800–5000 4250–5000 3700–5100 5000–5400 3700–5200 4900–5400 1600–5540 1200–1500 1500 1500–2000 1300–2350 5000 5000–7400 4300–5080 3900–5080 2100 4500 3400–5390 4500 3150 C. cetrarioides C. olivetorum Cetreliopsis rhytidocarpa ssp. langtangi Chaenotheca brunneola C. chrysocephala C. furfuracea C. hispidula C. phaeocephala C. stemonea Chrysothrix chlorina Cladia aggregata Cladonia amaurocraea C. awasthiana C. calyciformis C. cariosa C. carneola C. cartilaginea C. ceratophyllina C. chlorophaea C. ciliata C. coccifera C. corniculata C. corymbescens C. delavayi C. fenestralis C. fimbriata C. fruticulosa C. furcata C. humilis C. laii C. luteoalba C. macilenta C. macroptera C. mongolica C. nitida C. ochrochlora C. pocillum C. pyxidata C. ramulosa C. rangiferina C. scabriuscula C. singhii C. squamosa C. stellaris C. stricta C. subconistea C. subulata C. yunnana Coccocarpia erythroxyli C. palmicola Collema nepalense* C. poeltii* C. pulcellum Altitudinal range (m) 2000–3000 2850 2880 3000–3556† 2700–3879† 3000 †2947–3250 3000–3869† 3000 3100–3200 3900–4000 4530–5230 950–5100 1500–2700 3100 3900–4100 1100–2000 3000–4600 2000–5350 4480 2000–4000 1650–2300 1500–5100 2200–5230 3900–5100 3900–4250 1500–5100 1700–3900 1200–4250 2700–4600 3200–3660 1500–3800 2700–3900 1600–4500 3900 1500–3500 2000–3600 4000–4100 2000–3366 3400–4500 2845–3000 750–2700 1800–4000 4400–4500 3300–4000 2500–3500 2200–2700 3300–4000 3150 1800 3900–4000 3900–4000 1350–1500 2010 Elevation gradients of lichens in Nepal—Baniya et al. Appendix 1. Continued Name of lichen species C. rugosum C. subconveniens C. substipitatum Dermatocarpon miniatum var. miniatum D. vellereum Dibaeis sorediata Dimelaena oreina Dimerella lutea Diploicia canescens Diploschistes muscorum D. muscorum subsp. bartlettii D. muscorum subsp. muscorum D. nepalensis D. scruposus Dirinaria aegialita D. applanata D. consimilis Erioderma meiocarpum Eumitria pectinata Evernia mesomorpha Everniastrum cirrhatum E. nepalense E. rhizodendroideum Flavocetraria cucullata F. nivalis Flavocetrariella leucostigma F. melaloma Flavoparmelia caperata Flavopunctelia flaventior Fuscopannaria poeltii* F. praetermissa Glyphis cicatricosa Graphis scripta G. subglauconigra Haematomma puniceum Heterodermia angustiloba H. awasthii H. boryi H. comosa H. dactyliza H. dactyliza f. serpens H. dendritica H. diademata H. dissecta var. dissecta H. firmula H. flabellata H. himalayensis H. incana H. isidiophora H. japonica H. obscurata H. pellucida H. propagulifera 93 Appendix 1. Continued Altitudinal range (m) Name of lichen species Altitudinal range (m) 1800 1260 3000–4000 3200 2000–3600 2500 3200–4884 3000–4000 4650 900–2160 900–2100 1500–2160 900 2000–3000 1800 600 600 3000–3050 400–2000 3962–4572 1900–3300 1410–3600 2490–3800 3600–4500 3800–5300 3500–4500 3900–4200 2250–2743 2250–2800 3500 †3805–4000 1600 1600–3366 2000–3200 1800 1800–2300 1900 1500–3000 1500–2500 2100 3800–5150 1800 410–3807† 1500–3000 1200–2200 3100 1500–2000 1800–2100 2100–2250 3000–4000 1400–3000 2500–3992 1500–3000 H. pseudospeciosa H. punctifera H. rubescens H. speciosa H. togashii H. tremulans Hyperphyscia granulata H. minor Hypogymnia delavayi H. hypotrypa H. vittata Hypotrachyna adducta H. crenata H. exsecta H. flexilis H. imbricatula H. infirma H. koyaensis H. majoris H. neodissecta H. osseoalba H. revoluta H. rhabdiformis H. scytophylla H. sinuosa H. sublaevigata Immersaria athroocarpa Ingvariella bisporus Ioplaca pindarensis Lasallia freyana L. pertusa Lecanora adolfii L. amorpha* L. chlarotera L. chondroderma L. demissa L. emodi* L. formosa L. garovaglii L. hellmichiana* L. himalayae* L. kirra* L. lesleyana* L. meridionalis L. muralis var. dubyi L. muralis var. muralis L. phaeodrophthalma L. rubina var. australis L. rugosella L. sherparum* L. somervellii L. sulphurea L. terestiuscula 150–2100 2250 1800–2800 1650–2100 3000–4000 1500 1200 1200–1400 1800–4080 3600–4050 2800–4200 2100 2500–2800 1800–2300 2160 1400–2600 1000–2100 1500–2100 1650 2250–2550 1100–2200 2000–2250 2500–2800 2250–3250 3500 2250 3900–4340 3750 2900–5200 3800–3900 3800–5050 5540 5000 200 3800–5400 2700–4000 5540 4850–5860 3000–5050 1800–3500 3900–4850 4400–4500 5029–5334 1600–3366 4930 3500–4340 4100–4500 3300–4250 2732–3200 3300–4600 3700–5500 4300–5639 4500–5200 94 THE LICHENOLOGIST Appendix 1. Continued Vol. 42 Appendix 1. Continued Name of lichen species Altitudinal range (m) Name of lichen species Altitudinal range (m) L. tschomolongmae* Lecidea advena L. auriculata L. bella* L. brachyspora L. bucculenta L. diducens L. epiiodiza L. fuscoatra var. indecora L. haerjedalica var. gyrodisca L. himalaica L. khumbuensis L. lactea L. leptoboloides L. molybdochroa L. poeltii L. secernens L. silacea L. steineri L. tessellata Lecidella carpathica L. dimelaenophila L. stigmatea Leprocaulon arbuscula Leproplaca chrysodeta Leptogium asiaticum L. askotense L. azureum L. brebissonii L. burnetiae var. burnetiae L. cochleatum L. delavayi L. delavayi f. fuliginosulum L. isidiosellum L. javanicum L. pedicellatum L. phyllocarpum L. resupinas L. saturninum L. trichophorum Lethariella cladonioides Letrouitia domingensis Lobaria discolor L. isidiosa L. kurokawae L. pindarensis L. pseudopulmonaria L. retigera L. subretigera Lobothallia alphoplaca L. praeradiosa Melanelia tominii Melanohalea poeltii* 3000–5000 5000–5540 5000–5200 4950–5000 4300–5900 5000–5540 4720 4850 4340 5150–5200 5150–5200 5540 5460 4340–5200 3900–4000 4500–5000 5000 5000 5150–5200 5080–5639 3900–4000 3800–3900 4520–5200 1900–3100 3300–4850 3900–4100 1500–1800 1410 1800–3200 1500–2250 1800 3000–4100 3900–4000 1500 1410–1800 1500–3000 2000–3100 1600 1500–2100 1450–1500 4724 240 3150 2800–3962 1800–3400 2700–4000 2550–4050 1600–3650 2800 4400 4000–4500 3200–3600 4500–4600 Menegazzia terebrata Mycobilimbia hunana Myelochroa aurulenta M. entotheiochroa M. subaurulenta Nephroma helveticum var. helveticum N. nakaoi Nephromopsis ahtii N. isidioidea N. nephromoides N. pallescens N. stracheyi Ochrolechia bryophaga O. glacialis O. margarita O. rosella f. sorediascens O. subviridis O. trochophora Pachyphiale himalayensis Parmelaria subthomsonii P. thomsonii Parmelia adaugescens P. erumpens P. latissima var. marmariza P. masonii* P. meiophora P. omphalodes P. ricasolioides P. squarrosa P. submutata P. sulcata Parmelina tiliacea Parmelinella simplicior P. wallichiana Parmelinopsis expallida P. minarum Parmotrema austrosinense P. cooperi P. hababianum P. maclayanum P. melanothrix P. mellissii P. nilgherrensis P. praesorediosum P. pseudonilgherrense P. pseudotinctorum P. rampoddense P. ravum P. reticulatum P. sancti–angelii P. stuppeum P. tinctorum P. ultralucens 1300–4000 1650 1410–2250 2000–2200 1950 2160–2800 2800 4090–4530 2900–3100 2400–3600 2400–3048 2700–3300 3900–4000 5000–5200 5000 3500 2900 3000–4000 3900–4000 1800–3200 1800–3366 3300–3600 1900–2200 2900 3000–6100 1800–3350 3500–4500 2700 3000–3800 2800 3000–3366 2100 2000–2900 1800–2400 1800 1500 750–3000 1500–2100 1500–2000 1500 2400–3200 1500 1080–2800 1410–1800 1800–3530 2900–3366 900–2800 1500–2500 1500–2500 1410–2100 1800 240–2250 1500 2010 Elevation gradients of lichens in Nepal—Baniya et al. Appendix 1. Continued 95 Appendix 1. Continued Name of lichen species Altitudinal range (m) Name of lichen species Altitudinal range (m) P. yodae* Peltigera canina P. dolichorrhiza f. dolichorrhiza P. dolichospora* P. elisabethae P. malacea P. membranacea P. polydactylon P. pruinosa P. rufescens P. scabrosa Pertusaria hemisphaerica Phaeographina pyrrhochroa Phaeophyscia endococcina P. endococcina var. khumbuensis P. endococcinoides var. megalospora P. hispidula var. exornatula P. hispidula var. hispidula P. lygaea* P. primaria* P. pyrrhophora* P. sciastra Phlyctella indica Physcia aipolia P. caesia P. clementei P. dilatata P. dubia P. phaea P. stellaris ssp. intestiniformis* P. tribacia P. tribacioides Physciella nepalensis* Physconia distorta P. enteroxantha P. grisea P. muscigena Physma byrsaenum Placidium squamulosum Pleopsidium chlorophanum* Porpidia aerolotera P. crustulata P. elegantior P. hydrophila P. macrocarpa Protoparmelia badia var. badia P. effigurans* Punctelia borreri P. rudecta P. subrudecta Pyrenula cayennensis P. immersa Pyxine coccifera 2300 3150–3300 1800–2250 3000–4100 1350 2700–3300 1900–2400 1950–2920 1800 2100 3800 2400–3366 2100 3800–3900 3900 3900–4000 365 3800–3900 4500–4600 3800–3900 1420 3900–5200 1800 1800–2200 3900–5000 3500–4000 1600–5000 5000 4340–5000 4250–4340 3800–4600 1400 1400 3000–4000 2740 1400–4200 4250–5540 1350 4400–4500 3500–4000 5150–5200 2800–5000 4700 3900–4000 2950–4000 3750–4000 3750 1600–3200 1500–3200 1500–2500 3000–3200 2000 750 P. meissnerina P. philippina P. sorediata Ramalina conduplicans R. farinacea R. flabelliformis* R. hossei var. hossei R. sinensis R. subfarinacea Rhizocarpon geographicum Rhizoplaca chrysoleuca var. chrysoleuca R. melanophthalma var. obscura R. peltata Sagema potentillae Sclerophora coniophaea Solorina bispora Stereocaulon claviceps S. foliolosum var. foliolosum S. foliolosum var. botryophorum S. foliolosum var. strictum S. glareosum S. himalayense S. myriocarpum S. paradoxum S. piluliferum S. pomiferum S. sasaki var. sasaki Sticta henryana S. nylanderiana S. platyphylloides S. praetextata S. weigelii var. weigelii Sulcaria sulcata S. virens Tephromela aglaea T. armeniaca T. glacialis* T. siphulodes var. siphulodes* Thamnolia vermicularis Thelenella luridella Tremolecia atrata Tuckneraria laureri Tylophoron moderatum Umbilicaria badia U. cinereorufescens U. decussata var. decussata U. decussata var. rhizinata* U. indica var. indica U. indica var. nana* U. krascheninnikovii U. leiocarpa U. nenella U. nepalensis 300 900–1800 1410–1800 2133 2700 3200 1200–2500 2700–4267 1610 4500 2100–5150 4270 3660 4000 3000 3600–4200 4000 3300–4000 3600–4600 2400–3600 4200–4400 2500–5400 3900–5303 2000–3366 2160–5150 2500–4700 3600–4200 3657–3962 1800–3600 1900–3150 2100–3500 800–2250 3150–3300 3000–3600 5000–5200 5000–5200 4500 4830 3869–5455 1410 5000–5860 3200–4900 1800 2550 4250–5200 5100–5500 4950–5000 1800–3150 4200–4300 3700–5100 5150 5100 3600–5100 96 THE LICHENOLOGIST Vol. 42 Appendix 1. Continued Appendix 1. Continued Name of lichen species Altitudinal range (m) Name of lichen species Altitudinal range (m) U. thamnodes U. trabeculata U. vellea U. yunnana Usnea aciculifera U. baileyi U. compressa U. dendritica U. galbinifera var. subfibrillosa* U. himalayana U. longissima U. montisfuji U. nepalensis U. norkettii* U. pectinata U. pseudomontisfuji U. robusta 3600–5050 5000–5100 3600–5050 2550 1200–2250 1500–2400 2400–2700 1200–3000 2200 1800–3300 2700–3750 2200–3900 3800–4000 3000–3500 3962 3200 2500–3000 U. rubicunda U. splendeus U. thomsonii Xanthoparmelia coreana X. dentata Xanthoparmelia isidiosa X. mexicana X. nepalensis* Xanthoria borealis X. elegans X. fallax X. fulva X. sorediata X. ulophyllodes var. ulophyllodes 2700 1800–3300 1800–3150 3400–3850 4480–4572 2800–3200 3070–3700 3900 3900–4000 3750–6000 3200–3400 2700–3200 3200–5000 3200–4850† * species endemic to Nepal †Louise Olley’s unpublished personal observation Appendix 2. The elevational gradient of lichen species richness regression analysis results modelled after different species richness as response variables and their elevation as predictor variable. The Quasi-poisson family of error fitted in the GAM model after the cubic smooth spline (s) with approximately 4 degrees of freedom. (P%0·05) Response variables Null df Res. df Total species Endemic species Crustose species Foliose species Fruticose species Cyanolichens Green algal lichens Corticolous Saxicolous Terricolous 72 72 72 72 72 72 72 72 72 72 68 68 68 68 68 68 68 68 68 68 D2 0·945 0·915 0·914 0·941 0·961 0·944 0·941 0·970 0·908 0·895 Deviance 3598·1 362·3 1008 1764·9 1252·2 632·2 3047·7 2374·3 1075·7 422·3 Df = degree of freedom, Res. = residual, D2 = regression coefficient of determination F Pr (>F) 304·5 205 181·4 288·4 560·9 338·4 280·9 588 190·2 192·8 <0·001 <0·001 <0·001 <0·001 <0·001 <0·001 <0·001 <0·001 <0·001 <0·001