Abstract
The effect of competition on species coexistence is usually strongly modified by other factors especially in non-equilibrium systems of sessile organisms with limited availability of propagules. As a consequence, competition-based assembly rules (even if their existence seems to be unambiguously detected) would result in incomplete understanding of the coexistence of species in plant communities. J. Bastow Wilson suggested measuring variance deficit in the number of co-occurring species as a means to detect niche limitation in a community. The method provides a relatively simple and quick “snap-shot” analysis of a community. However, it has been questioned whether niche limitation is the only factor which might account for variance deficit.
The paper presents a spatially explicit simulation experiment in which artificial communities are produced by pre-defined rules for competitive interactions. Then we examine whether these rules can be detected by a proposed method for pattern analysis. Two limiting cases are simulated: (A) all the species share the same niche, and (B) all the species have different niches. The difference between these cases in the variance of species numbers is examined. Using the simulation results, some basic spatial constraints upon species assembly are emphasized.
It is argued that the assumptions of Wilson’s approach confine its applicability to species-saturated equilibrium communities. The study of assembly rules in dynamically changing, spatially structured communities requires the consideration of a set of coenological characteristics and the use of careful spatio-temporal scaling to detect their patterns. The use of spatially explicit individual-based models to study the mechanisms and constraints limiting species coexistence at different scales is suggested.
This is a preview of subscription content, log in via an institution to check access.
Similar content being viewed by others
References
Bartha S. (1992a): Gyomnövényközösségek szünmorfogenezise külszíni szénbányák meddőhányóin. (Vegetation development on dumps from strip-coal mining).—Ph.D. thesis, Vácrátót.
Bartha S. (1992b): Preliminary scaling for multi-species coalitions in primary succession.—Abstr. Bot. (Budapest) 16: 31–41.
Bartha S. &Horváth F. (1987): Application of long transects and information theoretical functions to pattern detection I. Transects versus isodiametric sampling units.—Abstr. Bot. (Budapest) 11: 9–26.
Bycroft C.M., Nicolaou N., Smith B. &Wilson J.B. (1993). Community structure (niche limitation and guild proportionality) in relation to the effect of spatial scale, in aNothofagus forest sampled with a circular transect.—New Zealand J. Ecol. 17: 95–101.
Cale W.G., Henebry G.M. &Yeakley J.A. (1989): Inferring process from pattern in natural communities.—BioScience 39: 600–605.
Czárán T. (1993): PATPRO: A Monte-Carlo simulation program for multispecies neighborhood competition. —Abstr. Bot. (Budapest) 17: 275–281.
Czárán T. &Bartha S. (1992): Spatiotemporal dynamic models of plant populations and communities.— Trends Ecol. Evol. 7: 38–42.
Diamond J.M. (1975): Assembly of species communities.—In:Cody M.L. &Diamond J.M. [eds.]: Ecology and evolution of communities, Harvard University Press, Cambridge, pp. 342–444.
Drake J.A. (1990): Communities as assembled structures: do rules govern pattern?—Trends Ecol. Evol. 5: 159–164.
Greig-Smith P. (1979): Pattern in vegetation.—J. Ecol. 67: 755–779.
Greig-Smith P. (1983): Quantitative plant ecology. Ed. 3.—University of California Press, Berkeley.
Harvey P.H., Colwell R.K., Silwertown J.W. &May R.M. (1983): Null models in ecology. —Annual Rev. Ecol. Syst. 14: 189–211.
Herben T., Krahulec F., Hadincová F. &Kovářová M. (1993): Small-scale spatial dynamics of plant species in a grassland community over six years.—J. Veg. Sci. 4: 171–178.
Hogeweg P., Hesper B., van Schaik C.P. &Beeftink W.G. (1985): Patterns in vegetation succession, an ecomorphological study.—In:White J. [ed.]: The population structure of vegetation, Dr. W. Junk Publ., Dordrecht, pp. 637–666.
Huston M., DeAngelis D. &Post W. (1988): New computer models unify ecological theory.— BioScience 38: 682–691.
Journel A.G. &Huijbregts C. (1978): Mining geostatistics.—Academic Press, London.
Juhász-Nagy P. (1984): Spatial dependence of plant populations. Part 2. A family of new models.—Acta Bot. Acad. Sci. Hung. 30: 363–402.
Juhász-Nagy P. &Podani J. (1983): Information theory methods for the study of spatial processes and succession.—Vegetatio 51: 129–140.
Keddy P.A. (1992): Assembly and response rules: two goals for predictive community ecology.—J. Veg. Sci. 3: 157–164.
Kolasa J. &Pickett S.T.A. [eds.] (1991): Ecological heterogeneity.—Springer-Verlag, New York.
Lepš J. (1990): Can underlying mechanisms be deduced from observed patterns?—In:Krahulec F., Willems J., Agnew A.D.Q. &Agnew S. [eds.]: Spatial processes in plant communities, Academia, Praha, pp. 1–13.
Lepš J. (1995): Variance deficit is not reliable evidence for niche limitation.—Folia Geobot. Phytotax. 30: 455–459.
van der Maarel E. &Sykes M.T. (1993): Small-scale plant species turnover in a limestone grassland: the carousel model and some comments on the niche concept.—J. Veg. Sci. 4: 179–188.
Palmer M.W. (1987): Variability in species richness within Minnesota oldfields: a use of the variance test.— Vegetatio 70: 61–64.
Palmer M.W. &White P.S. (1994): Scale dependence and the species-area relationship.—Amer. Naturalist 144: 717–740.
Palmer M.W. &van der Maarel E. (1995): Variance in species richness, species association, and niche limitation.—Oikos 73: 203–213.
Pickett S.T.A. (1980): Non-equilibrium coexistence of plants.—Bull. Torrey Bot. Club 107: 238–248.
Pickett S.T.A. &White P.S. [eds.] (1985): The ecology of natural disturbance and patch dynamics.— Academic Press, New York.
Podani J., Czárán T. &Bartha S. (1993): Pattern, area and diversity: the importance of spatial scale in species assemblages.—Abstr. Bot. (Budapest) 17: 37–51.
Rossi R.E., Mulla D.J., Journel A.G. &Franz E.H. (1992): Geostatistical tools for modeling and interpreting ecological spatial dependence.—Ecol. Monogr. 62: 277–314.
Schluter D. (1984): A variance test for detecting species associations, with some example applications.— Ecology 65: 998–1005.
Shmida A. &Ellner S. (1984): Coexistence of plant species with similar niches.—Vegetatio 58: 29–55.
Tilman D. (1994): Competition and biodiversity in spatially structured habitats.—Ecology 75: 2–16.
Tilman D. &Pacala S. (1993): The maintenance of species richness in plant communities.—In:Ricklefs R.E. &Schluter D. [eds.]: Species diversity in ecological communities, University of Chicago Press, Chicago, pp. 13–25.
Tóthmérész B. (1994): Statistical analysis of spatial pattern in plant communities.—Coenoses (Trieste) 9: 33–41.
Tóthmérész B. &Erdei Zs. (1992): The effect of species dominance on information theory characteristics of plant communities.—Abstr. Bot. (Budapest) 16: 43–47.
Watkins A.J. &Wilson J.B. (1992): Fine-scale community structure of lawns.—J. Ecol. 80: 15–24.
Wilson J.B. (1989): A null model of guild proportionality, applied to stratification of a New Zealand temperate rain forest.—Oecologia (Berlin) 80: 263–267.
Wilson J.B. (1991): Does vegetation science exist?—J. Veg. Sci. 2: 289–290.
Wilson J.B. (1994): Who makes the assembly rules?—J. Veg. Sci. 5: 275–278.
Wilson J.B., Gitay H. &Agnew A.D.Q. (1987): Does niche limitation exist?—Funct. Ecol. 1: 391–397.
Zobel K., Zobel M. &Peet R.K. (1993): Changes in pattern diversity during secondary forest succession in Estonian forests.—J. Veg. Sci. 4: 489–498.
Author information
Authors and Affiliations
Rights and permissions
About this article
Cite this article
Bartha, S., Czárán, T. & Oborny, B. Spatial constraints masking community assembly rules: A simulation study. Folia Geobot 30, 471–482 (1995). https://doi.org/10.1007/BF02803977
Issue Date:
DOI: https://doi.org/10.1007/BF02803977