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Environmental Niche Partitioning Among Riparian Sedges {Carex, Cyperaceae) in the St. Lawrence Valley, Quebec

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Environmental Niche Partitioning Among Riparian Sedges {Carex, Cyperaceae) in the St. Lawrence Valley, Quebec Environmental niche partitioning among riparian sedges {Carex, Cyperaceae) in the St. Lawrence Valley, Quebec Laura Plourde Master of Science Department of Biology McGill University Montreal, Quebec, Canada A thesis submitted to the Faculty of Graduate Studies and Research in partial fulfillment of requirements of the degree of Master of Science August 31st, 2007 ©Copyright Laura Plourde 2007. All rights reserved. Library and Bibliotheque et 1*1 Archives Canada Archives Canada Published Heritage Direction du Branch Patrimoine de I'edition 395 Wellington Street 395, rue Wellington Ottawa ON K1A0N4 Ottawa ON K1A0N4 Canada Canada Your file Votre reference ISBN: 978-0-494-51322-4 Our file Notre reference ISBN: 978-0-494-51322-4 NOTICE: AVIS: The author has granted a non­ L'auteur a accorde une licence non exclusive exclusive license allowing Library permettant a la Bibliotheque et Archives and Archives Canada to reproduce, Canada de reproduire, publier, archiver, publish, archive, preserve, conserve, sauvegarder, conserver, transmettre au public communicate to the public by par telecommunication ou par Plntemet, prefer, telecommunication or on the Internet, distribuer et vendre des theses partout dans loan, distribute and sell theses le monde, a des fins commerciales ou autres, worldwide, for commercial or non­ sur support microforme, papier, electronique commercial purposes, in microform, et/ou autres formats. paper, electronic and/or any other formats. The author retains copyright L'auteur conserve la propriete du droit d'auteur ownership and moral rights in et des droits moraux qui protege cette these. this thesis. Neither the thesis Ni la these ni des extraits substantiels de nor substantial extracts from it celle-ci ne doivent etre imprimes ou autrement may be printed or otherwise reproduits sans son autorisation. reproduced without the author's permission. In compliance with the Canadian Conformement a la loi canadienne Privacy Act some supporting sur la protection de la vie privee, forms may have been removed quelques formulaires secondaires from this thesis. ont ete enleves de cette these. While these forms may be included Bien que ces formulaires in the document page count, aient inclus dans la pagination, their removal does not represent il n'y aura aucun contenu manquant. any loss of content from the thesis. Canada Abstract To understand maintenance of the within-habitat diversity of closely related species, I investigated 11 Carex species growing along rivers in the south-western St. Lawrence Valley of Quebec. Microenvironments within a half meter of focal plants characterized for Carex comosa, C. crinita, C. grayi, C. intumescens, C. lacustris, C. lupulina, C. pseudocyperus, C. retrorsa, C. tuckermanii, C. typhina, and C. vesicaria revealed significant differences among the species in their environmental affinities. Species appear to fall into groups based on their tolerance of flooding and are secondarily differentiated on other environmental gradients such as insolation, soil pH and soil organic matter. Several traits were related to the environments that species inhabit: diaspore weight, diaspore floating duration, and root aerenchyma. The absence of any phylogenetic trend in niche differences for pairs of species supports the idea that evolutionary differentiation of the alpha-niche is the basis for coexistence of congeners. Resume Pour comprendre la preservation, a I'interieur d'un meme habitat, d'une diversite d'especes rapprochees phylogenetiquement, j'ai examine 11 especes de Carex vivant le long des rivieres dans le sud-ouest de la vallee du St-Laurent au Quebec. La characterisation de microenvironnements a I'interieur d'un demi metre de plantes focales characterises pour Carex comosa, C. crinita, C. grayi, C. intumescens, C. lacustris, C. lupulina, C. pseudocyperus, C. retrorsa, C. tuckermanii, C. typhina, and C. vesicaria a revele des differences significatives parmi les especes en ce qui concerne leurs affinites environnementales. Les especes sembleat s'agencer en groupes bases sur leurs tolerances face aux inondations et divises secondairement selon d'autres facteurs environnementaux tel I'insolation, le pH du sol, et le taux de matiere organique present dans le sol. Plusieurs traits ont ete relies a I'environnement que I'espece occupe: poids du diaspore, duree de flottation du diaspore et racine aerenchyme. L'absence de tendance phylogenetique en ce qui a trait aux differences entre les niches occupees par les differentes pairs d'especes supporte I'idee que la differentiation evolutive de I'alpha-niche est a la base de la coexistence d'especes appartenant a un meme genre. ii Table of Contents Abstract i Resume ii Table of Contents iii List of Tables iv List of Figures ' v Acknowledgements vi Chapter 1: Introduction 1 The neutral theory: a null hypothesis 2 Niche conservatism, differentiation and phylogenetic hierarchy 2 Riparian plant communities and their environmental gradients 4 Study genus 6 Study species 7 Objectives. 12 Chapter 2: Methods 13 Study area 13 Study species and focal sample selection 13 Environmental data........ 16 Trait data 18 Statistical analyses 19 Chapter 3: Results ...22 Univariate analysis of environmental variables 22 Multivariate analysis of the environment 23 Trait analysis 23 Phylogenetic analysis of the niche 24 Chapter 4: Discussion 40 Filtering by flood tolerance and segregation along multiple environmental gradients 40 Upper swamp 41 Lower swamp 42 Marsh 43 Role of traits 44 Absence of phylogenetic signal 46 Summary and conclusion 48 References 49 Appendix I: Canopy Analysis 57 Photograph Collection: 57 WinsCANOPY Data: 58 Appendix II: Estimation of percent time flooded 60 Appendix III: Soil collection and preparation 62 Soil collection: 62 Considerations on sampling and soil preparation: 62 Soil crushing: 63 Quartering the crushed and sifted soil sample: 63 Appendix IV: Aerenchyma Estimation 64 Killing and preservation agent: 64 Field root collection, killing, and preserving: 64 Photography: 68 Appendix V: Electronic archive of thesis material 69 List of Tables Table 1: Matrix of patristic distance between pairs of species based on branch lengths (number of changes or mutations along each branch) in a parsimony analysis using two nuclear and four chloroplast non-coding DNA regions (tree shown in Figurel) 10 Table 2: Habitat occurrence of the 11 study species from comments in published floras and journals -. 11 Table 3: Site descriptions 14 Table 4: Environmental data collected 20 Table 5: Univariate statistics for environmental variables at each focal plot (n=229). Species grouped by elevation on floodplain 25 Table 6: Total-sample standardized canonical coefficients of environmental variables, P values (P>0) and percent variance explained for each axis 32 Table 7: Niche distances. Squared Mahalanobis distance among species are given above the diagonal, with associated p-values below the diagonal. Bold numbers indicate the few species pairings that are oot significantly different (p>0.01) 33 Table 8: Univariate statistics for trait data 35 iv List of Figures Figure 1: Phylogram of riparian sedge species (Waterway unpublished). Bootstrap values based on 100 replicates are shown on branches with >50% support. The dashed line collapses in a strict consensus tree 9 Figure 2: Site Locations: CSJ: Parc-Nature du Cap-St-Jacques, DIL: Dowker Island, HUL: Hull, HUM: Huntingdon Marsh, MAS: Masson, OKA: Pare National d'Oka, PLS: Pare National de Plaisance, RIG: Pointe-Riviere-a-la-Raquette, VAU: Vaudreuil-Sur-Le-Lac; Gauging Stations: 1) Hull Dam, 2) Carillon Dam, 3) Summerstown, 4) Ste-Anne-De-Bellevue, 5) Pointe-Calumet, 6) Pointe-Claire 15 Figure 3: Boxplots of species elevation from the average river level (in cm) displaying habitat groupings used in the discussion of ecological differences among the study species. Dots are outliers, whiskers are the 10th and 90th percentiles. The box is defined by the 25th and 75th percentiles. The bar within the box is the median. In this compilation and comparison of foal plant data sampled across sites the average water level from 2003-2005 is standardized at zero elevation for each site 30 Figure 4: Biplots of species distribution on the first two axes of a Canonical Variates Analysis based on 9 environmental variables for 11 riparian Carex species. The first and second canonical axes explain 63 and 17% of the variation in this data set, respectively. Circles represent individual focal plots. Outer ellipses are the 95% confidence interval of the species distribution; the 95% confidence interval of the mean is the inner ellipse. Sets of biplots grouped within bold lines contain species of the same riparian habitat type; from top to bottom of the figure these are upper swamp, lower swamp, and marsh. Smaller boxes within each grouping denote pairs of sister species ....31 Figure 5: Biplots of species by habitat groups in canonical space. Canonical axes in each biplot are those which best visually separate the species for that particular group. Crosses (+) are species means, ellipses are 95% bivariate confidence intervals of focal plots. IN: Carex intumescens, GR: C. grayi, TY: C. typhina, CR:C. crinita, RE: C. retrorsa, LU: C. lupulina, TU: C. tuckermanii, VE: C. vesicaria, CO: C. comosa, LA: C. lacustris, PS: C. pseudocyperus 34 Figure 6: Scatterplot of species average location on flooding gradient in relation to floating duration. Species acronyms are as in Figure 5. Triangles indicate marsh species, squares: lower swamp, and diamonds:
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