Skip to main content
Log in

The intimacy between sexual traits and Grime’s CSR strategies for orchids coexisting in semi-natural calcareous grassland at the Olive Lawn

  • Published:
Plant Ecology Aims and scope Submit manuscript

Abstract

The diversity of orchid species in semi-natural calcareous grassland is thought to depend on floral variability. However, differences in primary life-history strategy, or the overall suite of functional traits, could also affect coexistence. Both primary strategy (affecting day-to-day survival) and reproductive traits (representing periodic events) have been hypothesized to contribute to a general mechanism regulating the local persistence and relative abundance of species. We recorded species identity at 8,000 points at the centimetre scale along sixteen 5-m long transects at the Olive Lawn, a xeric sand calcareous grassland near Lecco, Italy. Transects allowed the relative abundance and spatial aggregation of Grime’s competitor, stress-tolerator, ruderal (CSR) strategies to be quantified. Orchid flowering and fruiting phenology was recorded alongside the phenology of graminoid growth. Seven orchid species were present and exhibited a spectrum of strategies spanning two extremes, (1) small early-flowering ruderals (R/CR-selected), and (2) larger-leaved competitors (C/CR) flowering later as dominant graminoids attained maximum height. These orchids, and other subordinate species in the community, exhibited random spatial distributions amongst dominant graminoids that exhibited a high degree of spatial aggregation and similar, but non-identical, stress-tolerator (S) strategies. The reproductive phenology of meadow orchids is an intimate component of their CSR strategies, and appears to promote coexistence by allowing segregation between temporal niches and the exploitation of opportunities between clumps of dominant species. Only for closely related species with identical CSR strategies, such as Ophrys benacensis and Ophrys sphegodes, could specific floral differences alone explain coexistence.

This is a preview of subscription content, log in via an institution to check access.

Access this article

Price excludes VAT (USA)
Tax calculation will be finalised during checkout.

Instant access to the full article PDF.

Fig. 1
Fig. 2
Fig. 3
Fig. 4

Similar content being viewed by others

References

  • Arditti J (1992) Fundamentals of orchid biology. Wiley, New York

    Google Scholar 

  • Armbruster WS, Edwards ME, Debevec EM (1994) Floral character displacement generates assemblage structure of Western Australian triggerplants (Stylidium). Ecology 75:315–329

    Article  Google Scholar 

  • Benzing DH, Atwood JT (1984) Orchidaceae: ancestral habitats and current status in forest canopies. Syst Bot 9:155–165

    Article  Google Scholar 

  • Cerabolini BEL, Brusa G, Ceriani RM, De Andreis R, Luzzaro A, Pierce S (2010) Can CSR classification be generally applied outside Britain? Plant Ecol 210:53–261

    Article  Google Scholar 

  • Cornelissen JHC, Lavorel S, Garnier E, Diaz S, Buchmann N et al (2003) A handbook of protocols for standardised and easy measurement of plant functional traits worldwide. Aust J Bot 51:335–380

    Article  Google Scholar 

  • Cozzolino S, Widmer A (2005) Orchid diversity: an evolutionary consequence of deception? Trends Ecol Evol 20(9):487–494

    Article  PubMed  Google Scholar 

  • Craine JM (2009) Resource strategies of wild plants. Princeton University Press, Princeton

    Google Scholar 

  • Díaz S, Hodgson JG, Thompson K, Cabido M, Cornelissen JHC et al (2004) The plant traits that drive ecosystems: evidence from three continents. J Veg Sci 15:295–304

    Article  Google Scholar 

  • Dorland E, Willems JH (2006) High light availability alleviates the costs of reproduction in Ophrys insectifera (Orchidaceae). J Europäischer Orchideen 38(2):501–518

    Google Scholar 

  • Dressler RL (1981) The orchids. Natural history and classification. Harvard University Press, Cambridge

    Google Scholar 

  • Elliott JM (1993) Some methods for the statistical analysis of samples of benthic invertebrates. Scientific publication 25. Freshwater Biological Association, Ambleside

    Google Scholar 

  • Fargione J, Tilman T (2006) Plant species traits and capacity for re source reduction predict yield and abundance under competition in nitrogen-limited grassland. Funct Ecol 20:533–540

    Article  Google Scholar 

  • Fitter AH (1982) Influence of soil heterogeneity on the coexistence of grassland species. J Ecol 70:139–148

    Article  Google Scholar 

  • Frenette-Dussault C, Shipley B, Léger J-F, Meziane D, Hingrat Y (2012) Functional structure of an arid steppe plant community reveals similarities with Grime’s C-S-R theory. J Veg Sci 23:208–222

    Article  Google Scholar 

  • Freschet GT, Aerts R, Cornelissen HC (2012) A plant economics spectrum of litter decomposability. Funct Ecol 26:56–65

    Article  Google Scholar 

  • Fridley JD, Grime JP, Askew AP, Moser B, Stevens CJ (2011) Soil heterogeneity buffers community response to climate change in species-rich grassland. Glob Change Biol 17:2002–2011

    Article  Google Scholar 

  • GIROS (Italian Group for Research on Spontaneous Orchids) (2009) Orchidee d’Italia (Orchids of Italy). Il Castello, Cornaredo (in Italian)

    Google Scholar 

  • Grime JP (1998) Benefits of plant diversity to ecosystems: immediate, filter and founder effects. J Ecol 86:901–910

    Article  Google Scholar 

  • Grime JP (2001) Plant strategies, vegetation processes and ecosystem properties, 2nd edn. Wiley, Chichester, p 417

    Google Scholar 

  • Grime JP, Pierce S (2012) The evolutionary strategies that shape ecosystems. Wiley-Blackwell, Chichester

    Book  Google Scholar 

  • Guidi V (2011) Study of the biodiversity of native orchids in the province of Lecco. Unpublished BSc thesis, University of Milan Bicocca, Milan, p 81

  • Hutchings MJ (1987) The population biology of the early spider orchid, Ophrys sphegodes Mill. I. Demographic study from 1975 to 1984. J Ecol 75(3):711–727

    Google Scholar 

  • Hutchings MJ, Mendoza A, Havers W (1998) Demographic properties of an outlier population of Orchis militaris L. (Orchidaceae) in England. Bot J Linn Soc 126:95–107

    Google Scholar 

  • Jacquemyn H, Brys R, Honnay O, Hermy M (2008) Effects of coppicing on demographic structure, fruit and seed set in Orchis mascula. Basic Appl Ecol 9:392–400

    Article  Google Scholar 

  • Kullenberg B (1950) Investigations on the pollination of Ophrys species. Oikos 2(1):1–19

    Article  Google Scholar 

  • Martin EM (1996) Fitness costs of resource overlap among coexisting bird species. Nature 380:338–340

    Article  CAS  Google Scholar 

  • Otero JP, Ackerman JD, Bayman P (2002) Diversity and host specificity of endophytic Rhizoctonia-like fungi from tropical orchids. Am J Bot 89(11):1852–1858

    Article  CAS  Google Scholar 

  • Pärtel M, Kalamees R, Zobel M, Rosén E (1998) Restoration of species-rich limestone grassland communities from overgrown land: the importance of propagule availability. Ecol Eng 10:275–286

    Article  Google Scholar 

  • Pedersen HÆ, Faurholdt M (2007) Ophrys: the bee orchids of Europe. Kew Publishing, London

    Google Scholar 

  • Pierce S, Ceriani RM, Villa M, Cerabolini B (2006) Quantifying relative extinction risks and targeting intervention for the orchid flora of a natural park in the European pre-alps. Conserv Biol 20(6):1804–1810

    Article  PubMed  Google Scholar 

  • Pierce S, Luzzaro A, Caccianiga M, Ceriani RM, Cerabolini B (2007) Disturbance is the principal α-scale filter determining niche differentiation, coexistence and biodiversity in an alpine community. J Ecol 95:698–706

    Google Scholar 

  • Pierce S, Brusa G, Vagge I, Cerabolini BEL (2013) Allocating CSR plant functional types: the use of leaf economics and size traits to classify woody and herbaceous vascular plants. Funct Ecol 27(4):1002–1010

    Article  Google Scholar 

  • Pignatti S (1982) Flora d’Italia. Edagricole, Bologna

    Google Scholar 

  • Rasmussen HN (2002) Recent developments in the study of orchid mycorrhiza. Plant Soil 244:149–163

    Article  CAS  Google Scholar 

  • Rathcke BJ (1983) Competition and facilitation among plants for pollination. In: Real L (ed) Pollination Biology. Academic Press, New York, pp 305–329

    Google Scholar 

  • Reynolds HL, Hungate BA, Chapin FS, D’Antonio CM (1997) Soil heterogeneity and plant competition in an annual grassland. Ecology 78:2076–2090

    Google Scholar 

  • Schlüter PM, Ruas PM, Kohl G, Ruas CF, Stuessy TF, Paulus HF (2009) Genetic patterns and pollination in Ophrys iricolor and O. mesaritica (Orchidaceae): sympatric evolution by pollinator shift. Bot J Linn Soc 159:583–598

    Article  Google Scholar 

  • Scopece G, Musacchio A, Widmer A, Cozzolino S (2007) Patterns of reproductive isolation in Mediterranean deceptive orchids. Evolution 61(11):2623–2642

    Article  PubMed  Google Scholar 

  • Silvera K, Santiago LS, Winter K (2005) Distribution of crassulacean acid metabolism in orchids of Panama: evidence of selection for weak and strong modes. Funct Plant Biol 32:397–407

    Article  CAS  Google Scholar 

  • Soliva M, Widmer A (2003) Gene flow across species boundaries in sympatric, sexually deceptive Ophrys (Orchidaceae) species. Evolution 57(10):2252–2261

    PubMed  Google Scholar 

  • Tilman D (1988) Plant Strategies and the Dynamics and Structure of Plant Communities. Princeton University Press, Princeton

    Google Scholar 

  • Vereecken NJ, Cozzolino S, Schiestl FP (2010) Hybrid floral scent novelty drives pollinator shift in sexually deceptive orchids. BMC Evol Biol 10:103

    Article  PubMed Central  PubMed  Google Scholar 

  • Waser NM (1983) Competition for pollination and floral characters differences among sympatric species: a review of evidence. In: Jones DE, Little RJ (eds) Handbook of experimental pollination biology. Van Nostrand Reinhold, New York, pp 277–293

    Google Scholar 

  • Waterman RT, Bidartondo MI, Stofberg J, Combs JK, Gebauer G, Savolainen V, Barraclough TG, Pauw A (2011) The effects of above- and below-ground mutualisms on orchid speciation and coexistence. Am Nat 177(2):E54–E68

    Article  PubMed  Google Scholar 

  • Wells TCE, Rothery P, Cox R, Bamford S (1998) Flowering dynamics of Orchis morio L. and Herminium monorchis (L.) R.Br. at two sites in eastern England. Bot J Linn Soc 126:39–48

    Google Scholar 

  • Wright IJ, Reich PB, Westoby M et al (2004) The worldwide leaf economics spectrum. Nature 428:821–827

    Article  CAS  PubMed  Google Scholar 

Download references

Acknowledgments

We thank Salvatore Valsecchi (the owner of the fieldsite) the Native Flora Centre of the Lombardy Region (CFA) and Parco Monte Barro (acting on behalf of the Lombardy Region) for permission to work at the site and collect plant material, and for access to the equipment and laboratory at Monte Barro where leaf area and weight were determined. Pierfranco Arrigoni originally made the authors aware of the presence of the orchids at the site.

Author information

Authors and Affiliations

Authors

Corresponding author

Correspondence to Simon Pierce.

Additional information

Communicated by Satoki Sakai.

Electronic supplementary material

Below is the link to the electronic supplementary material.

Supplementary material 1 (DOC 7451 kb)

Supplementary material 2 (XLS 580 kb)

Rights and permissions

Reprints and permissions

About this article

Cite this article

Pierce, S., Vagge, I., Brusa, G. et al. The intimacy between sexual traits and Grime’s CSR strategies for orchids coexisting in semi-natural calcareous grassland at the Olive Lawn. Plant Ecol 215, 495–505 (2014). https://doi.org/10.1007/s11258-014-0318-y

Download citation

  • Received:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI: https://doi.org/10.1007/s11258-014-0318-y

Keywords

Navigation