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FIGURE 15. A in Disentangling the diversity and taxonomy of Hymenophyllaceae (Hymenophyllales, Polypodiidae) in the Mascarene archipelago, with ecological implications
FIGURE 15. A. Hymenophyllum capillare Desv., typical form, as epiphyte (La Réunion, 'Forêt de Bon Accueil, Les Makes'). B. H. capillare, typical form, detail of fronds. C. H. capillare f. nanum Hennequin & Dubuisson, f. nov., as epiphyte (La Réunion, 'Pitons Mare à Boue'). D. H. capillare f. nanum, habit, type individual, scale = 0.5 cm (La Réunion, 'Forêt de Bon Accueil, Les Makes', J.-Y. Dubuisson & L. Bauret HR 2016-10, P). E. H. digitatum (Sw.) Forsberg, as epiphyte (La Réunion, 'Forêt de Bon Accueil, Les Makes'). F. H. digitatum, detail of laminar margin showing minute dark trichomes. G. H. digitatum, detail of fronds and sori with tubular bases (s) (photographs. A–B: L. Bauret; C–G: J.-Y. Dubuisson).
FIGURE 9. A in Disentangling the diversity and taxonomy of Hymenophyllaceae (Hymenophyllales, Polypodiidae) in the Mascarene archipelago, with ecological implications
FIGURE 9. A. Crepidomanes fallax (Christ) Ebihara & Dubuisson, as lithophyte on wet rock (La Réunion, 'Rivière des Marsouins'). B. C. fallax, habit (from Madagascar, F. Rakotondrainibe 6467, P00248564). C. C. frappieri (Cordem.) J.P.Roux, covering a rock in a wet ravine (La Réunion, 'Ravine Rivière Ste Anne'). D. C. frappieri, habit, type specimen (Bédier s.n., P00477825). E. C. frappieri, detail of long branched rhizome trichomes. F. C. frappieri, detail of sori (photographs. A, C & F: J.-M. Tamon).
FIGURE 18. A in Disentangling the diversity and taxonomy of Hymenophyllaceae (Hymenophyllales, Polypodiidae) in the Mascarene archipelago, with ecological implications
FIGURE 18. A. Polyphlebium diaphanum (Kunth) Ebihara & Dubuisson, as epiphyte (La Réunion, 'Forêt de Bon Accueil, Les Makes'). B. P. diaphanum, detail of a frond and sori. C. Vandenboschia gigantea (Bory ex Willd.) Pic.Serm., climbing on a tree trunk (La Réunion, 'Mare Longue'). D. V. gigantea, terrestrial parts (La Réunion, 'Grand Etang'). E. V. gigantea, detail of a long-creeping climbing rhizome (photographs. A: L. Bauret; B–E: J.-Y. Dubuisson).
FIGURE 19 in Disentangling the diversity and taxonomy of Hymenophyllaceae (Hymenophyllales, Polypodiidae) in the Mascarene archipelago, with ecological implications
FIGURE 19. Importance of gametophyte ecology and biology for the sporophyte settlement on distant islands according to Dassler & Farrar's (2001) hypothesis. The long-lived and colonial epiphytic gametophytes (above) would have the possibilities to realize fertilization many seasons/years (T) after the spore germination, whereas short-lived and non-colonial terrestrial gametophytes (below) would fail in realizing the fertilization if they are not close enough, and die at the end of the season (red crosses). Hence, the epiphytic taxa with longlived and colonial gametophytes would have a higher probability of establishment than terrestrial taxa. Hymenophyllaceae and the genus Ctenitis (Dryopteridaceae) illustrate such contrasts in the Mascarene archipelago, explaining the difference in the level of endemicity (see text for an in-depth discussion).
FIGURE 7 in Disentangling the diversity and taxonomy of Hymenophyllaceae (Hymenophyllales, Polypodiidae) in the Mascarene archipelago, with ecological implications
FIGURE 7. Abrodictyum tamarisciforme (Jacq.) Ebihara & Dubuisson. A. A dense population of many individuals covering a trunk (La Réunion, 'Grand Etang'). B. Single individual as epiphytic on a tree-fern trunk, which is also covered here by Didymoglossum cuspidatum (La Réunion, 'Mare Longue'). C. Lithophytic individual, the little individual at the bottom right is A. parviflorum (Poir.) Bauret & Dubuisson (La Réunion, 'Grand Etang'). D. Detail of segments with typical distinct marginal row of cells (mc) (photographs. A, C: J.-Y. Dubuisson; B: C. Chaussidon; D: L. Bauret).
FIGURE 6 in Disentangling the diversity and taxonomy of Hymenophyllaceae (Hymenophyllales, Polypodiidae) in the Mascarene archipelago, with ecological implications
FIGURE 6. Abrodictyum parviflorum (Poir.) Bauret & Dubuisson. A. typical habit, here as terrestrial (La Réunion, 'Mare Longue'). B. Brush-like 'meifolium' form (La Réunion, 'Grand Etang'). C. Plane 'parviflorum' form (La Réunion, 'Grand Etang'). D. Detail of subcapillary ultimate segments and sori (photographs. A: C. Chaussidon; B, D: L. Bauret; C: J.-Y. Dubuisson).
FIGURE 5 in Disentangling the diversity and taxonomy of Hymenophyllaceae (Hymenophyllales, Polypodiidae) in the Mascarene archipelago, with ecological implications
FIGURE 5. Distribution of Hymenophyllaceae species richness in relation to elevations, here on the windward side of La Réunion and on Mauritius (data are reported in Appendices 1 & 2). The range of each taxon is assumed to correspond to the range between the minimum and the maximum elevation at which they have been observed, even if the taxon has not been observed in all intermediate localities between such limits.
FIGURE 14. A in Disentangling the diversity and taxonomy of Hymenophyllaceae (Hymenophyllales, Polypodiidae) in the Mascarene archipelago, with ecological implications
FIGURE 14. A. Hymenophyllum fumarioides Bory ex. Willd, as epiphyte (La Réunion, 'Mare Longue'). B. H. fumarioides, detail of curled/crisped segments and sori with toothed margins. C. H. peltatum (Poir.) Desv., as epiphyte (La Réunion, 'Pitons Mare à Boue'). D. H. peltatum, detail of toothed/serrulate laminar margins. E. H. sibthorpioides (Bory ex Willd.) Mett. ex Kuhn, as epiphytic (La Réunion, 'Vallée Heureuse'). F. H. sibthorpioides, fertile frond and sori with toothed margins (photographs. A, E: C. Chaussidon; B, F: L. Bauret; C: J.-Y. Dubuisson; D: J.-M. Tamon).
FIGURE 4. Mauritian ecosystems. A in Disentangling the diversity and taxonomy of Hymenophyllaceae (Hymenophyllales, Polypodiidae) in the Mascarene archipelago, with ecological implications
FIGURE 4. Mauritian ecosystems. A. Lowland Sapotaceae rainforest, here invaded by the exotic Malagasy traveller's trees Ravenala madagascariensis (Strelitziaceae) ('Vallée d'Osterlog' forest, ~260 m). B. 'Pétrin' locality showing particular habitat with thickets of the endemic Erica brachyphylla (Ericaceae) (~670 m) (photographs by J.-Y. Dubuisson).
FIGURE 11 in Disentangling the diversity and taxonomy of Hymenophyllaceae (Hymenophyllales, Polypodiidae) in the Mascarene archipelago, with ecological implications
FIGURE 11. Crepidomanes trinerve (Baker) Dubuisson & Ebihara. A. As lithophyte in a wet ravine of the western side of La Réunion ('Les Canots'; photograph by J.-M. Tamon). B. Habit (from Comores, F. Rakotondrainibe 6651, P00310478; modified from Dubuisson et al. 2013).
FIGURE 3. Particular Mascarenan ecosystems. A in Disentangling the diversity and taxonomy of Hymenophyllaceae (Hymenophyllales, Polypodiidae) in the Mascarene archipelago, with ecological implications
FIGURE 3. Particular Mascarenan ecosystems. A. Hyperhumid montane 'Avoune' vegetation dominated by the endemic Erica reunionensis (Ericaceae), with most trunks and branches fully covered by epiphytes (liverworts, mosses, peat mosses, lycopods and ferns, especially here Hymenophyllum capillare and H. inaequale, and orchids) (La Réunion, 'Pitons Mare à Boue', ~1,600 m). B. Montane 'Tamarinaie' dominated by endemic Acacia heterophylla (Fabaceae), showing as for 'Avoune' tree-trunks and branches fully covered by epiphytes, including here H. inaequale (La Réunion, 'Bélouve', ~1,520 m). C. Remnant of leeward lowland semi-dry forest (La Réunion, 'Ilet Solitude', ~550 m). D. Typical wet ravines that host lithophytic Hymenophyllaceae on the wet and shaded rocks close to streams and waterfalls. (photographs. A: J.-Y. Dubuisson; B–C: J.-M. Tamon; D: A. Mercier).
FIGURE 8. A in Disentangling the diversity and taxonomy of Hymenophyllaceae (Hymenophyllales, Polypodiidae) in the Mascarene archipelago, with ecological implications
FIGURE 8. A. Crepidomanes bipunctatum (Poir.) Copel., as epiphyte on a tree-trunk (Mauritius, 'Bar le Duc'). B. C. bipunctatum, detail of segments with false veins parallel to the margins (fv). C. C. bipunctatum, sori (s) with triangular lips. D. C. bonapartei (C.Chr.) J.P.Roux, as lithophyte on wet rock (La Réunion, 'Grand Etang'). E. C. bonapartei, detail of subsessile or short-stipitate fronds and one sorus (photographs. A: J.-Y. Dubuisson; B–C: L. Bauret; D–E: E. Grangaud).
FIGURE 2. Wet Mascarenan ecosystems. A in Disentangling the diversity and taxonomy of Hymenophyllaceae (Hymenophyllales, Polypodiidae) in the Mascarene archipelago, with ecological implications
FIGURE 2. Wet Mascarenan ecosystems. A. Lowland windward rainforest dominated by Sapotaceae trees, showing the understory with numerous low epiphytic and terrestrial free-sporing plants (liverworts, mosses, ferns, lycopods and spikemosses) and a few herbaceous angiosperms (especially orchids) (La Réunion, 'Mare Longue' forest, ~450 m). B. Montane rainforest dominated by Dombeyoideae (Malvaceae) trees and Alsophila spp. tree-ferns (Cyatheaceae) as emergents from the canopy (La Réunion, 'Cassé de Takamaka, Bébour', 1,100–1,200 m). C. Montane ericoid thickets (La Réunion, 'Enclos du Piton de la Fournaise', ~2,200 m). (Photographs. A, C: C. Chaussidon; B: J.-M. Tamon).
FIGURE 10. A in Disentangling the diversity and taxonomy of Hymenophyllaceae (Hymenophyllales, Polypodiidae) in the Mascarene archipelago, with ecological implications
FIGURE 10. A. Crepidomanes inopinatum (Pic.Serm.) J.P.Roux var. inopinatum, as lithophyte (La Réunion, 'Petite Plaine'). B. C. inopinatum var. tamonii Y.Robert & Dubuisson, habit (Y. Robert 913, P02433924). C. C. minutum (Blume) K.Iwats. var. minutum, as epiphyte (La Réunion, 'Bébour'). D. C. minutum var. minutum, frond with long terete wingless stipe. E. C. minutum var. minutum, showing stipe proliferation, scale = 1 cm (from Madagascar, F. Rakotondrainibe 3698, P00085267). F. C. minutum var. mascarenense Pynee & Dubuisson, as epiphyte (La Réunion, 'Grand Etang'). G. C. minutum var. mascarenense, detail of fronds with short stipes. (photographs. A: J.-Y. Dubuisson; C–D: C. Chaussidon; F–G: E. Grangaud).
FIGURE 13 in Disentangling the diversity and taxonomy of Hymenophyllaceae (Hymenophyllales, Polypodiidae) in the Mascarene archipelago, with ecological implications
FIGURE 13. Didymoglossum cuspidatum (Willd.) Ebihara & Dubuisson. A. Typical sub-circular, and lanceolate to deltoid sterile forms as usually observed (La Réunion, 'Mare Longue'). B. Linear sterile forms (La Réunion, 'Grand Etang'). C. Ovate to lanceolate fertile fronds with apical sori (La Réunion, 'Forêt de Bon Accueil, Les Makes'). D. Narrowly lanceolate fertile frond with apical sori (La Réunion, 'Grand Etang'). E. Small sub-circular/ovate fertile frond with a single sorus, showing sub-marginal false vein (mfv) and false veins parallel to true veins (fv) (photographs. A–D: J.-Y. Dubuisson; E: L. Bauret).
FIGURE 17. A in Disentangling the diversity and taxonomy of Hymenophyllaceae (Hymenophyllales, Polypodiidae) in the Mascarene archipelago, with ecological implications
FIGURE 17. A. Hymenophyllum balfourii Baker, holotype (La Réunion, I.B. Balfour s.n., K000435603). B. H. inaequale (Poir.) Desv., as epiphyte, narrowly lanceolate form (La Réunion, 'Mare Longue'). C. H. inaequale, widely lanceolate to ovate form (La Réunion, 'Forêt de Bon Accueil, Les Makes'). D. H. inaequale, detail of glabrous segments and sori (photographs. B. L. Bauret; C–D: J.-Y. Dubuisson).
FIGURE 1 in Disentangling the diversity and taxonomy of Hymenophyllaceae (Hymenophyllales, Polypodiidae) in the Mascarene archipelago, with ecological implications
FIGURE 1. Location of the Mascarene Archipelago (or Mascarenes) in the western Indian Ocean (from Pynee et al. 2012).
FIGURE 16. A in Disentangling the diversity and taxonomy of Hymenophyllaceae (Hymenophyllales, Polypodiidae) in the Mascarene archipelago, with ecological implications
FIGURE 16. A. Hymenophyllum hirsutum (L.) Sw., as epiphyte (La Réunion, 'Grand Etang'). B. H. hirsutum, detail of a frond showing the hairy winged stipe. C. H. hirsutum, detail of bilabiate hairy sori. D. H. hygrometricum (Poir.) Desv., as epiphyte (La Réunion, 'Bébour'). E. H. hygrometricum, wet fronds become greener as shown here (photographs. A: J.-Y. Dubuisson; B–C: L. Bauret; D–E: C. Chaussidon).
FIGURE 20 in Disentangling the diversity and taxonomy of Hymenophyllaceae (Hymenophyllales, Polypodiidae) in the Mascarene archipelago, with ecological implications
FIGURE 20. Elevational distribution on each Hymenophyllaceae taxon on the windward (red plots) and leeward (blue plots) sides of La Réunion, and correspondence with thermal zones or ranges according to Cadet (1980) and provided here for the windward side, with ranges of annual average temperatures. The dotted horizontal lines correspond to the limits of leeward ecosystems. The plots indicate the average elevation. Crepidomanes trinerve is here lacking because it was only observed at a single locality on the leeward side ('Les Canots' at ~800 m). The range of each taxon, as illustrated by the vertical lines, is assumed to correspond to the range between the minimum and the maximum elevation at which they have been observed, even if the taxon has not been observed in all the intermediate localities between such limits.
Data from: Multiple facets of stream macroinvertebrate alpha diversity are driven by different ecological factors across an extensive altitudinal gradient
Environmental filtering and spatial structuring are important ecological processes for the generation and maintenance of biodiversity. However, the relative importance of these ecological drivers for multiple facets of diversity is still poorly understood in highland streams. Here, we examined the responses of three facets of stream macroinvertebrate alpha diversity to local environmental, landscape-climate and spatial factors in a near-pristine highland riverine ecosystem. Taxonomic (species richness, Shannon diversity and evenness), functional (functional richness, evenness, divergence and Rao's Quadratic entropy) and a proxy of phylogenetic alpha diversity (taxonomic distinctness and variation in taxonomic distinctness) were calculated for macroinvertebrate assemblages in 55 stream sites. Then Pearson correlation coefficient was used to explore congruence of indices within and across the three diversity facets. Finally, multiple linear regression models and variation partitioning were employed to identify the relative importance of different ecological drivers of biodiversity. We found most correlations between the diversity indices within the same facet, and between functional richness and species richness were relatively strong. The two phylogenetic diversity indices were quite independent from taxonomic diversity but correlated with functional diversity indices to some extent. Taxonomic and functional diversity were more strongly determined by environmental variables, while phylogenetic diversity was better explained by spatial factors. In terms of environmental variables, habitat-scale variables describing habitat complexity and water physical features played the primary role in determining the diversity patterns of all three facets, whereas landscape factors appeared less influential. Our findings indicated that both environmental and spatial factors are important ecological drivers for biodiversity patterns of macroinvertebrates in Tibetan streams, although their relative importance was contingent on different facets of diversity. Such findings verified the complementary roles of taxonomic, functional and phylogenetic diversity, and highlighted the importance of comprehensively considering multiple ecological drivers for different facets of diversity in biodiversity assessment.
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