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Epizoochorous Seed Dispersal by Large Herbivores

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Epizoochorous Seed Dispersal by Large Herbivores ouvreur C ISPERSAL D Martine EED S ERBIVORES BY H ARGE L PIZOOCHOROUS E 2005 EPIZOOCHOROUS SEED DISPERSAL BY LARGE HERBIVORES MARTINE COUVREUR ISBN 90-5989-052-3 ISBN 90-5989-052-3 UNIVERSITEIT GENT FACULTEIT BIO-INGENIEURSWETENSCHAPPEN Vakgroep Bos- en Waterbeheer _________________________ Academiejaar 2004 - 2005 21 maart 2005 EPIZOOCHOROUS SEED DISPERSAL BY LARGE HERBIVORES EPIZOÖCHORE ZAADVERBREIDING DOOR GROTE HERBIVOREN door Martine Couvreur Thesis submitted in fulfillment Proefschrift voorgedragen tot het of the requirements for behalen van de graad van Doctor the degree of Doctor (Ph.D.) in de Toegepaste Biologische in Applied Biological Sciences Wetenschappen (Land- en Bosbeheer) Op gezag van: Rector Prof. Dr. A. De Leenheer Promotor : Prof. Dr. ir. K. Verheyen, Universiteit Gent Copromotor: Prof. Dr. M. Hermy, Katholieke Universiteit Leuven Decaan: Prof. Dr. ir. H. Van Langenhove, Universiteit Gent Leden van de Examencommissie: Prof. Dr. J. Bakker, Rijksuniversiteit Groningen, Nederland Prof. Dr. H. Van Dyck, Université Catholique de Louvain Prof. Dr. M. Hoffmann, Universiteit Gent Prof. Dr. P. Goetghebeur, Universiteit Gent Prof. Dr. ir. R. Lemeur, Universiteit Gent Prof. Dr. ir. G. Hofman, Universiteit Gent (voorzitter) Wijze van citeren: Couvreur M. 2005. Epizoochorous seed dispersal by large herbivores. Ph.D. thesis, Ghent University, Ghent. Nederlandse vertaling van de titel: Epizoöchore zaadverbreiding door grote herbivoren. Illustraties: Bert Delanoeije, Eric Cosyns, Dries Adriaens ISBN 90-5989-052-3 The author and the promotor give the authorization to consult and to copy parts of this work for personal use only. Every other use is subject to the copyright laws. Permission to reproduce any material contained in this work should be obtained from the author. Table of contents Samenvatting............................................................................................................................iii Summary..................................................................................................................................vii CHAPTER 1......................................................................................................................... p. 1 Introduction CHAPTER 2 ...................................................................................................……........... p. 17 Large herbivores as mobile links within and between isolated nature reserves through adhesive seed dispersal. Adapted from: Couvreur, M., Christiaen, B., Verheyen, K. & Hermy, M. (2004). Applied Vegetation Science 7(2): 229-236. CHAPTER 3 .........................................................................................................…......... p. 39 Complementarity of epi- and endozoochory by free ranging donkeys. Adapted from: Couvreur, M., Cosyns, E., Hermy, M. & Hoffmann, M. (2005). Ecography 28(1): 37-48. CHAPTER 4 ...................................................................................................................... p. 57 Experimental assessment of seed adhesivity on animal furs. Adapted from: Couvreur, M., Vandenberghe, B., Verheyen, K. & Hermy, M. (2004). Seed Science Research 14(2): 147-159. CHAPTER 5 ...................................................................................................................... p. 81 Experimental assessment of plant seed retention times in fur of cattle and horse. Adapted from: Couvreur, M., Verheyen, K. & Hermy, M. (2005). Flora 200(2): 126-137. CHAPTER 6 ...................................................................................................................... p. 97 Epizoochory by large herbivores: merging data with models. Adapted from: Couvreur, M., Verheyen, K., Vellend, M., Lamoot, I., Cosyns E., Hoffmann, M. & Hermy, M. Ecography (submitted). CHAPTER 7 .................................................................................................................... p. 121 Conclusions References ........................................................................................……......…….......... p. 135 Curriculum and Publications ......................................................…….........…….......... p. 151 i ii Samenvatting De verbreiding van plantenzaden via de vacht van dieren (epizoöchorie) is één van de belangrijke mechanismen die de lange-afstandsverbreiding van planten kunnen mediëren. In vele ecosystemen is zaadverbreiding een ‘bottleneck’ voor kolonisatie en migratie van planten. In het snel veranderende landschap in Vlaanderen en west-Europa zijn vele (half)natuurlijke habitats bovendien verloren gegaan, en zijn de resterende gebieden sterk versnipperd. De doorgedreven modernisering van de landbouw heeft ook heel wat traditionele gebruiken doen verdwijnen, hetgeen in het algemeen tot een verminderd zaadaanbod en een verminderde connectiviteit van het landschap heeft geleid. Epizoöchorie door in het wild levende zoogdieren en loslopend vee heeft bijgevolg sterk aan belang ingeboet. Anderzijds is begrazing met grote herbivoren momenteel één van de belangrijkste natuurbeheerpraktijken geworden, waarbij regelmatig transport van vee komt kijken. Via deze grazers kan dus potentieel nog een belangrijke uitwisseling van zaden tussen soortenrijke gebieden gebeuren. Lange-afstandszaadverbreiding is echter moeilijk te traceren, zeker wanneer er complex diergedrag bij komt kijken. Ondanks het toenemend aantal onderzoeken naar zaadverbreiding is epizoöchorie dan ook nog weinig gekend. In deze studie worden verschillende aspecten van epizoöchorie belicht, via beschrijvend, experimenteel en modelleringsonderzoek. De voornaamste doelstellingen waren (1) inzicht verwerven in de hoeveelheid, identiteit en eigenschappen van zaden die via epizoöchorie verbreid worden, (2) belangrijke aspecten van epizoöchorie experimenteel kwantificeren, en (3) velddata rond epizoöchorie integreren in een model. Hiertoe bestudeerden we vooral grote grazers die ingezet worden voor de begrazing in Vlaamse natuurgebieden. Deze kunnen fungeren als model voor wilde zoogdieren in verleden, heden en toekomst, zijn relatief gemakkelijk te onderzoeken, en spelen een cruciale rol in het hedendaags natuurbeheer in onze streken. In een eerste deel van deze studie werden de vachten van een 200-tal grote herbivoren (Galloway-runderen, paarden en ezels) uit Vlaamse natuurgebieden geïnspecteerd op zaden. We identificeerden 6385 kiemkrachtige zaden van 75 plantensoorten, met een aanzienlijke variatie aan planteigenschappen en zaadmorfologieën. In tegenstelling tot wat oorspronkelijk werd verondersteld, worden dus niet alleen zaden met morfologische aanpassingen voor epizoöchorie (bv. haakjes) verbreid in de vacht van dieren, maar ook ongespecialiseerde zaden en zaden aangepast aan andere verbreidingsmechanismen (bv. wind, water). Onze resultaten geven aan dat de rol van grote herbivoren in zaadverbreiding aanzienlijk is, en dat iii zij als ‘mobiele links’ binnen en tussen habitats fungeren -bij diertransporten in het kader van seizoensbegrazing zelfs tussen natuurgebieden op grote afstanden. Dieren verbreiden niet enkel zaden via epizoöchorie, maar ook via endozoöchorie (na doorgang van de zaden door het spijsverteringsstelsel). Beide mechanismen opereren gelijktijdig, maar werden nog maar zelden vergeleken als gevolg van hun mechanistische verschillen. Bij onderzoek naar de relatie tussen epi- en endozoöchorie vonden we dat in een kustduinengebied 20% van de lokale plantensoorten via de grote grazers werd verbreid, met seizoenale verschillen in soortensamenstelling. Een complementaire set aan plantensoorten werd verbreid via epi- respectievelijk endozoöchorie (slechts één vierde van de 66 zoöchore soorten werd verbreid door beide mechanismen). Deze complementariteit werd bevestigd op niveau van verbreidingsgerelateerde plantkenmerken, die gebruikt werden om de soorten in het studiegebied op te delen in functionele planttypes. Een vergelijking van het aandeel epi- en endozoöchoor verbreide soorten in deze functionele groepen gaf aan dat epizoöchorie specifieker was dan endozoöchorie (geassocieerd met een kleiner aantal functionele plantgroepen). De resultaten tonen aan dat epi- en endozoöchorie beide belangrijk zijn, als additieve en complementaire verbreidingsmechanismen. Een sleutelaspect van epizoöchorie is de adhesieve interactie tussen zaden en vachten. In een tweede, experimenteel luik, kwantificeerden we deze interactie met behulp van gelooide vachten van gedomesticeerde en wilde dieren, via een eenvoudige gestandardiseerde methode. Op die manier bekwamen we een maat voor de epizoöchore verbreidings- mogelijkheden van uiteenlopende zaadsoorten in relatie tot verschillende vachten. Diepe vachten met lange, golvende haren en een verticale haarinplant (Galloway-rund, schaap, wild zwijn) waren beter geschikt voor de aanhechting van zaden dan ondiepe vachten met korte, gladde haren die horizontaal ingeplant zijn (ree, Holstein rund, konijn, paard). In het algemeen werd de zaadaanhechting gefaciliteerd door gespecialiseerde zaadaanpassingen (haken, borstels, kafnaalden, haren, …) en bemoeilijkt door niet-gespecialiseerde aanpassingen zoals steeltjes of vleugels. Nochtans was de functionaliteit van bepaalde zaadmorfologieën sterk afhankelijk van het type vacht (interactie-effect). Zaadmorfologie bleek een goede predictor te zijn voor zaadaanhechting, alhoewel ook minder adhesieve zaden relatief goed scoorden. Dit suggereert dat zo goed als alle soorten tot op zekere hoogte intrinsiek in staat zijn tot epizoöchore zaadverbreiding.
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