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Termite Mediated Heterogeneity of Soil and Vegetation Patterns in a Semi‐Arid Savanna Ecosystem in Namibia

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Termite Mediated Heterogeneity of Soil and Vegetation Patterns in a Semi‐Arid Savanna Ecosystem in Namibia Termite mediated heterogeneity of soil and vegetation patterns in a semi‐arid savanna ecosystem in Namibia Dissertation zur Erlangung des naturwissenschaftlichen Doktorgrades der Julius‐Maximilians‐Universität Würzburg vorgelegt von Constanze Grohmann geboren in der Hansestadt Lübeck Würzburg, 2010 Termite mediated heterogeneity of soil and vegetation patterns in a semi‐arid savanna ecosystem in Namibia Dissertation zur Erlangung des naturwissenschaftlichen Doktorgrades der Julius‐Maximilians‐Universität Würzburg vorgelegt von Constanze Grohmann geboren in der Hansestadt Lübeck Würzburg, 2010 Eingereicht am: 11.11.2010 Mitglieder der Promotionskommission: Vorsitzender: Prof. Dr. Thomas Dandekar Gutachter: Prof. Dr. K. Eduard Linsenmair Gutachter: Prof. Dr. Jürgen Tautz Tag des Promotionskolloquiums: 14.01.2011 Doktorurkunde ausgehändigt am: _________ Um es vorweg zu nehmen: Die Namib schluckte mich mit Haut und Haaren. Dort war ich glücklich und ruhig in mir. Wer die Wüste nicht selbst erlebt hat, ihre Weite und ihren Frieden, ihre Härte und Unerbittlichkeit, wird es nicht verstehen, und ich werde es ihm nicht erklären können. Ich kannm ih nur sagen, dass ich seither ein kleines Stück Wüste in mir trage. Und es ist ein größerer Schatz für mich als manches Schulwissen. Barbara Imgrund This thesis is based on the following manuscripts Grohmann C. and Linsenmair K.E. (in prep.). Biodiversity of termite species along a rainfall gradient in Namibia. Grohmann C., Gröngröft A., Petersen A. and Linsenmair K.E. (in prep.). Macrotermes michaelseni mounds enhance the spatial heterogeneity of nitrogen distribution in the central Namibian savanna. Graiff A., Grohmann C. and Linsenmair K.E. (in prep.). The impact of a fungus growing termite species (Isoptera, Macrotermitinae) on soil fertility and plant growth. Grohmann C., Oldeland J., Stoyan D. and Linsenmair K.E. 2010. Multi‐scale pattern analysis of a mound building termite species. Insectes Sociaux 57(4): 477‐486. Table of contents Zusammenfassung .............................................................................................................. 1 Abstract .............................................................................................................................. 3 I. General introduction........................................................................................................ 5 Introduction....................................................................................................................... 5 Thesis outline .................................................................................................................. 12 The BIOTA‐project........................................................................................................... 13 Study sites ....................................................................................................................... 13 References....................................................................................................................... 15 II. Biodiversity of termites along rainfall gradients in Namibia.......................................... 19 Abstract ........................................................................................................................... 19 Introduction..................................................................................................................... 20 Methods .......................................................................................................................... 21 Results ............................................................................................................................. 25 Discussion........................................................................................................................ 31 Conclusion ....................................................................................................................... 34 References....................................................................................................................... 35 III. Macrotermes michaelseni mounds enhance the spatial heterogeneity of nitrogen distribution....................................................................................................................... 39 Abstract ........................................................................................................................... 39 Introduction..................................................................................................................... 40 Methods .......................................................................................................................... 41 Results ............................................................................................................................. 44 Discussion........................................................................................................................ 52 References....................................................................................................................... 54 Supplementary Material ................................................................................................. 57 IV. Impact of Macrotermes michaelseni on soil fertility and plant growth ........................ 61 Abstract ........................................................................................................................... 61 Introduction..................................................................................................................... 62 Methods .......................................................................................................................... 63 Results ............................................................................................................................. 66 Discussion........................................................................................................................ 70 References....................................................................................................................... 73 V. Multi‐scale pattern analysis of Macrotermes mounds .................................................. 77 Abstract ........................................................................................................................... 77 Introduction..................................................................................................................... 78 Methods .......................................................................................................................... 79 Results ............................................................................................................................. 83 Discussion........................................................................................................................ 89 References....................................................................................................................... 91 VI. General discussion....................................................................................................... 95 References....................................................................................................................... 99 Acknowledgements ............................................................................................................. i Authors’ contributions........................................................................................................iii List of publications.............................................................................................................vii Ehrenwörtliche Erklärung ...................................................................................................ix Zusammenfassung 1 Zusammenfassung Termiten sind die bedeutendsten Ökosystem‐Ingenieure in den Böden arider und semi‐ arider Gebiete. Sie beschleunigen Zersetzungsprozesse und damit auch die nachfolgende Mineralisation von Nährstoffen durch Bakterien und Pilze. Durch den Bau ihrer Galerien, Nester und Hügel fördern sie die Umwälzung des Bodens und beeinflussen die Nährstoffver‐ teilung, die Textur sowie die hydrologischen Eigenschaften der Böden. Durch diese Prozesse erhöhen sie die Heterogenität in den von ihnen bewohnten Ökosystemen. Die vorliegende Arbeit beschäftigt sich vorrangig mit dem Einfluss von Termiten auf Ökosystemfunktionen eines semi‐ariden Ökosystems. Als Grundlage dazu habe ich die Diver‐ sität von Termitentaxa und ihre Abhängigkeit von Niederschlagsmenge, Vegetationsmustern und Landnutzungssystemen auf verschiedenen Untersuchungsflächen in Namibia bestimmt. In dem darauffolgenden Teil der Arbeit habe ich mich auf die pilzzüchtende und hügel bauende Art Macrotermes michaelseni (Sjöstedt, 1914) konzentriert, eine Art, die in vielen afrikanischen Savannen in hoher Abundanz vorkommt. Ich habe mich damit beschäftigt, wie diese Art die räumliche Heterogenität von Böden und Vegetationsmustern beeinflusst. Durch wiederholtes Sammeln an 13 verschiedenen Untersuchungsgebieten in Namibia konnte ich 17 Termitentaxa aus 15 Gattungen erfassen. Während die Landnutzung einen unwesentlichen Einfluss auf die Termitenfauna hatte, konnte die Variation in der Taxa‐ diversität zu 96% durch den mittleren jährlichen Niederschlag erklärt werden und zu 81% durch den Simpson‐Diversitäts‐Index der Gefäßpflanzen. Die Anzahl der Termitentaxa nahm mit diesen beiden Variablen zu. Im Gegensatz zu vorherigen Studien
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