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Basic and Applied Ecology Basic Appl. Ecol. 3, 85–94 (2002) © Urban & Fischer Verlag Basic and Applied Ecology http://www.urbanfischer.de/journals/baecol Does plant competition intensity rather depend on biomass or on species identity? Alexandra Weigelt*, Tom Steinlein, Wolfram Beyschlag Universität Bielefeld, Lehrstuhl für Experimentelle Ökologie und Ökosystembiologie, Bielefeld, Germany, Received January 16, 2001 · Accepted July 23, 2001 Abstract In two experiments, we tested whether species specific traits or mainly biomass determines the com- petitive strength of plant individuals in resource-poor habitats. As measure of competition intensity, we calculated the log Response Ratio (lnRR) based on total biomass for three key species of early successional stages on inland dunes. Using seedlings of Corynephorus canescens and Hieracium pilosella in a pot experiment, competition intensity was significantly and positively correlated with the biomass of the respective competitors. In contrast, such a correlation was not detected in a con- trolled field experiment with adult plants of the two species and of Carex arenaria. However, in both experiments the strength of competitive interactions (measured as lnRR) significantly depend- ed on the identity of the competing species. We conclude, that a biomass advantage over the com- petitors (which can for instance be achieved by earlier germination) seems to play a crucial role only for successful seedling establishment, while competitive interactions of neighbouring plants depend on species-specific biomass allocation strategies at both developmental stages. Um zu testen, ob artspezifische Eigenschaften oder hauptsächlich die Biomasse selbst die Konkur- renzstärke zwischen Pflanzen auf ressourcenarmen Standorten bestimmen, wurde der Log Response Ratio (lnRR) der Gesamtbiomasse für drei dominante Arten früher Sukzessionsstadien auf Sand bestimmt. In einem Topfexperiment, in dem Keimlinge der Arten Corynephorus canescens und Hie- racium pilosella verwendet wurden, nahm die Konkurrenz mit zunehmender Biomasse umgebender Keimlinge signifikant zu. In einem zweiten Experiment unter kontrollierten Freilandbedingungen, dagegen, in dem ausgewachsene Pflanzen der gleichen Arten zusammen mit Carex arenaria gemessen wurden, war die Konkurrenzintensität nicht signifikant von der Biomasse benachbarter Pflanzen abhängig. Artspezifische Eigenschaften hatten jedoch einen signifikanten Einfluss auf die Konkurrenzstärke sowohl von Keimlingen als auch von älteren Pflanzen. Wir folgern daraus, dass ein zeitlicher Vorsprung bei der Besiedlung offener Flächen, der mit höherer Biomasse einher geht, für einen Keimling einen entscheidenden Konkurrenzvorteil bietet. Mit zunehmendem Alter jedoch nimmt der Einfluss der Biomasse auf das Konkurrenzgeschehen ab und die artspezifische Biomasse- Allokation gewinnt mehr und mehr an Bedeutung. Key words: Corynephorus canescens – Hieracium pilosella – Carex arenaria – log Response Ratio – sandy soil *Corresponding author: Alexandra Weigelt, Lehrstuhl für Experimentelle Ökologie und Ökosystembiologie, Universität Bielefeld W4-107, Universitätsstr. 25, D-33615 Bielefeld, Germany, Phone ++49-521-106 5573; Fax: ++49-521-106 6038, E-mail: [email protected] 1439-1791/02/03/01-085 $ 15.00/0 86 A. Weigelt et al. Introduction biomass on competition intensity, and (2) whether there are different elevations of regression lines be- So far many studies considered changing resource tween species indicating that species-specific traits do gradients in different habitats, while within one plant affect competitive strength. community or at the single plant level, the correlation of competition and biomass is much less debated. However, the change of target plant biomass with Materials and methods density of neighbours has been modelled (Shinozaki & Kira 1956, Mead 1979, Firbank & Watkinson Species description 1985, Connolly 1986, Firbank & Watkinson 1990, Humphrey & Pyke 1998) and the few mathematical All three study species are common plants of European approaches for a conceptual integration of neighbour coastal and inland sand dunes where they are predom- biomass into competition indices (Goldberg 1987, inantly found in the early successional stages. Freckleton & Watkinson 2000) reflect that density Corynephorus canescens (L.) P. beauv. is a tufted win- and biomass are obviously correlated. In both cases, tergreen perennial grass that occurs on open sand. The the function between target plant growth and neigh- species has a finely divided, fibrous root system with- bour plant density (biomass) is generally expected to out rhizome. The bulk of its roots is about 25 cm long, be either inverse linear or hyperbolic (see Fig. 5, and the maximum length is 35–40 cm. Its roots are Mead 1979, Firbank & Watkinson 1985, 1990, Con- mainly directed downwards with only few or no hori- nolly 1986, Austin et al. 1988, Humphrey & Pyke zontal ones (Marshall 1967). Hieracium pilosella L. 1998), i.e. the curve shows a considerable descend at (the mouse-ear hawkweed) is a stoloniferous perenni- low and a rather shallow decrease at high neighbour al, with extended clonal growth in often denser vegeta- densities. tion cover. H. pilosella forms a fine, spiders’ web like Accordingly, competition between small plants (e.g. rather superficial root system with intensive lateral seedlings), or at low neighbour density, should follow growth and only some thicker deep growing roots of a pattern, where a small increase in neighbour biomass 30 to 40 cm length (personal observation). The third results in a considerable decrease of target biomass. In species, Carex arenaria L. (the sand sedge) is a sympo- contrast, one should expect only minimal changes of dial plant forming an extensive perennial rhizome sys- target biomass in competitive interaction between tem. C. arenaria is generally more abundant on dry adult plants (and/or at higher neighbour density) even grassland stages and sometimes open pine forests on if neighbour biomass intensively changes. In the field, sand. A distinctive morphological feature of the plant densities are often rather in the stages with high species is the root dimorphism. At the base of each neighbouring biomass, which is why we hypothesise, shoot one or two large sinker roots emerge, together that neighbouring species biomass might play a subor- with several finer roots. Fine roots may reach about dinate role in determining competition patterns in nat- 50 cm in length and the large sinker roots may grow 2 ural vegetation. m in one summer, while the maximum reported root- We choose three dominant and often co-occurring ing depth for the plant was 3–4 m in deep sand (Noble species of early successional stages on inland dunes to 1982). study this question with both, a pot and a field-like The three species are the only dominant perennials competition experiment. To analyse competitive on these rather species poor stages that could be se- strength in the target-neighbour design we used the lected for their differences in growth of both above- “log Response Ratio” (lnRR) rather than the “relative and belowground structures, which we assumed competition intensity” (RCI) for mainly statistical rea- might influence competitive patterns. Rather than sons (see methods). We performed the analysis with forming one successional stage together, where the both RCI and lnRR and found no qualitative differ- three are all key species, they are dominant in differ- ences between the two indices. Therefore only the ent but subsequent stages of succession on inland lnRR will be presented. dunes. However, one can often find the species co-oc- The present study was set up to investigate, whether curring on intermediate vegetation patches and as in a situation of existing competition the biomass of single plants within each others dominant stages, neighbouring species or rather species-specific at- which is comparable to the density and ground cover tributes determine the competitive strength of plants in the field-like experiment with adult plants (see coexisting in the same habitat. Therefore we analysed below). Seeds of H. pilosella and C. canescens both linear regressions of target plant lnRR on total neigh- germinate during late summer under favourable cli- bour biomass to ask, (1) whether there are significant matic conditions often together in small disturbed regression slopes which indicate an effect of plant patches. Basic Appl. Ecol. 3, 1 (2002) Biomass or species specific competition intensity 87 Seedling competition experiment In a pot experiment with H. pilosella and C. canescens we first tested the influence of different biomass of seedlings on competitive interactions. H. pilosella plants were grown from seeds collected from approximately 20 maternal plants from an area of 100 m × 20 m with patchy vegetation of all early successional stages in an inland dune area called “Senne” near Bielefeld (co-or- dinates: 08° 40′ E 51° 57′ N). For C. canescens seed material from seven german botanical gardens was mixed. Subsequently, seedlings of three different age classes (10, 32 and 54 days old, respectively) were planted either alone (control) or with 4 individuals per pot (pot diameter 9 cm; pot height 7.5 cm) filled with pure sand. The competitive design was set up as two Fig. 1. Planting design of the adult-plant competition experiment. 120 pairs of two equally aged plants, and included all com- hexagonal plots (diameter d = 55 cm, distance target-neighbour plants binations of species and age.
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