Folia Geobotanica

, Volume 52, Issue 3–4, pp 433–442 | Cite as

Accounting for clonality in comparative plant demography – growth or reproduction?

  • Zdeněk JanovskýEmail author
  • Tomáš Herben
  • Jitka Klimešová


Clonal growth occurs in a high proportion of herbaceous plant species, but it is difficult to deal with in demographic transition matrix models. It is primarily a growth process, but in many cases gives rise to new individuals from the viewpoint of plant demography. In the present contribution, we review how clonality is treated in existing demographic studies of clonal species from the Central European flora. We based our analyses on publicly accessible data in the COMPADRE and CLO-PLA databases and compiling from these information on plant matrix population models (hereafter MPMs) and clonal traits. Out of the 55 species × study combinations of species with known clonality, ca 30% neglect clonality. Another 30% treat clonal growth as growth of mother ramet and only the remaining 40% address clonality as a separate reproductive pathway. However, but only about half of these studies (12; 22% of total) reported a separable submatrix of clonal transitions necessary for comparative analyses. Here we show that failure to report clonal transitions separately leads to a serious bias in estimation of many demographic characteristics derived from MPMs, such as generation time. Shoot ontogeny (namely shoot lifespan and its branching architecture) largely determined whether clonality was neglected, treated as growth or treated as reproduction. Additionally, species with long runners were under-represented in demographic studies relative to their proportion in the Central European clonal flora. To conclude, we discuss how to construct MPMs for species with varying degree of ramet integration. We recommend that demographic analyses should focus on ramets – the smallest unit capable of living through all life-cycle stages of the species – as a sampling and study unit of plant populations.


CLO-PLA COMPADRE Cyclicity Population projection matrix Ramet Shoot lifespan Shoot ontogeny 



We thank Kay Gross for her valuable comments on the previous versions of manuscript. The research reported here was supported by grants from the Czech Science Foundation (16-19245S, Centre of Excellence 14-36079G PLADIAS).

Supplementary material

12224_2017_9301_MOESM1_ESM.doc (37 kb)
ESM 1 (DOC 37 kb)


  1. Aarssen LW (2008) Death without sex – the 'problem of the small' and selection for reproductive economy in flowering plants. Evol Ecol 22:279–298CrossRefGoogle Scholar
  2. Adler PB, Salguero-Gómez R, Compagnoni A, Hsu JS, Ray-Mukherjee J, Mbeau-Ache C, Franco M (2014) Functional traits explain variation in plant life history strategies. Proc Natl Acad Sci USA 111:740–745CrossRefPubMedGoogle Scholar
  3. Barkham JP (1980) Population dynamics of the Wild Daffodil (Narcissus pseudonarcissus). I. Clonal growth, seed reproduction, mortality and the effects of density. J Ecol 68:607–633Google Scholar
  4. Berg H (2002) Population dynamics in Oxalis acetosella: the significance of sexual reproduction in a clonal, cleistogamous forest herb. Ecography 25:233–243Google Scholar
  5. Brys R, Jacquemyn H, Endels P, De Blust G, Hermy M (2005) Effect of habitat deterioration on population dynamics and extinction risks in a previously common perennial. Conservation Biol 19:1633–1643CrossRefGoogle Scholar
  6. Burns JH, Pardini EA, Schutzenhofer MR, Chung YA, Seidler KJ, Knight TM (2013) Greater sexual reproduction contributes to differences in demography of invasive plants and their noninvasive relatives. Ecology 94:995–1004Google Scholar
  7. Carlsson BÅ, Callaghan TV (1990) Programmed tiller differentiation, intraclonal density regulation and nutrient dynamics in Carex bigelowii. Oikos 58:219–230CrossRefGoogle Scholar
  8. Carlsson BA, Callaghan TV (1991) Simulation of fluctuating populations of Carex bigelowii tillers classified by type, age and size. Oikos 60:231–240CrossRefGoogle Scholar
  9. Caswell H (2001) Matrix population model: construction, analysis, and interpretation, 2nd edition ed. Sinauer Associates, Inc. Publishers, Sunderland, MA, USAGoogle Scholar
  10. Černá L, Münzbergová Z (2013) Comparative population dynamics of two closely related species differing in ploidy level. PLOS ONE 8Google Scholar
  11. Cochran ME, Ellner S (1992) Simple methods for calculating age-based life history parameters for stage-structured populations. Ecol Monogr 62:345–364CrossRefGoogle Scholar
  12. Colling G, Matthies D (2004) The effects of plant population size on the interactions between the endangered plant Scorzonera humilis, a specialised herbivore, and a phytopathogenic fungus. Oikos 105:71–78CrossRefGoogle Scholar
  13. Curle CM, Stabbetorp OE, Nordal I (2004) Eryngium maritimum, biology of a plant at its northernmost localities. Nordic J Bot 24:617–628Google Scholar
  14. Dinnétz P, Nilsson T (2002) Population viability analysis of Saxifraga cotyledon, a perennial plant with semelparous rosettes. Pl Ecol 159:61–71Google Scholar
  15. Ehrlén J (1995) Demography of the perannial herb Lathyrus vernus – 2. Herbivory and population dynamics. J Ecol 83:297–308Google Scholar
  16. Ehrlén J, Lehtilä K (2002) How perennial are perennial plants? Oikos 98:308–322CrossRefGoogle Scholar
  17. Ehrlén J, Syrjänen K, Leimu R, García MB, Lehtilä K (2005) Land use and population growth of Primula veris: an experimental demographic approach. J Appl Ecol, 42, 317–326Google Scholar
  18. Endels P, Jacquemyn H, Brys R, Hermy M (2005) Rapid response to habitat restoration by the perennial Primula veris as revealed by demographic monitoring. Pl Ecol 176:143–156Google Scholar
  19. Endels P, Jacquemyn H, Brys R, Hermy M (2007) Genetic erosion explains deviation from demographic response to disturbance and year variation in relic populations of the perennial Primula vulgaris. J Ecol 95:960–972Google Scholar
  20. Eriksson O (1988) Ramet behaviour and population growth in the clonal herb Potentilla anserina. J Ecol 76:522–536Google Scholar
  21. Forbes J C (1977) Population flux and mrotality in a ragwort (Senecio jacobaea L.) infestation. Weed Res 17:387–391Google Scholar
  22. Fröborg H, Eriksson O (2003) Predispersal seed predation and population dynamics in the perennial understorey herb Actaea spicata. Canad J Bot 81:1058–1069Google Scholar
  23. Gabbard B L (2003) The population dynamics and distribution of the exotic grass, Bothriochloa ischaemum. University of Texas at Austin, AustinGoogle Scholar
  24. García MB, Picó FX, Ehrlén J (2008) Life span correlates with population dynamics in perennial herbaceous plants. Amer J Bot 95:258–262Google Scholar
  25. García MB, Goñi D, Guzmán D (2010) Living at the edge: local versus positional factors in the long-term population dynamics of an endangered orchid. Conservation Biol 24:1219–1229Google Scholar
  26. Gustafsson C, Ehrlén J (2003) Effects of intraspecific and interspecific density on the demography of a perennial herb, Sanicula europaea. Oikos 100:317–324Google Scholar
  27. Harper JL (1977) Population biology of plants. Academic Press, New YorkGoogle Scholar
  28. Herben T, Šerá B, Klimešová J (2015) Clonal growth and sexual reproduction: tradeoffs and environmental constraints. Oikos 124:469–476CrossRefGoogle Scholar
  29. Honnay O, Jacquemyn H, Roldán-Ruiz I, Hermy M (2006) Consequences of prolonged clonal growth on local and regional genetic structure and fruiting success of the forest perennial Maianthemum bifolium. Oikos 112:21–30CrossRefGoogle Scholar
  30. Iriondo JM, Albert MJ, Giménez Benavides L, Domínguez Lozano F, Escudero A (2009) Poblaciones en peligro: viabilidad demográfica de la flora vascular amenazada de España. Dirección General de Medio Natural (MMARM) Google Scholar
  31. Jacquemyn H, Brys R (2008) Effects of stand age on the demography of a temperate forest herb in post-agricultural forests. Ecology 89:3480–3489Google Scholar
  32. Jacquemyn H, Brys R, Neubert M G (2005) Fire increases invasive spread of Molinia caerulea mainly through changes in demographic parameters. Ecol Applic 15:2097–2108CrossRefGoogle Scholar
  33. Jacquemyn H, Brys R, Honnay O, Hermy M, Roldán-Ruiz I (2005) Local forest environment largely affects below-ground growth, clonal diversity and fine-scale spatial genetic structure in the temperate deciduous forest herb Paris quadrifolia. Molec Ecol 14:4479–4488CrossRefGoogle Scholar
  34. Jacquemyn H, Brys R, Jongejans E (2010) Size-dependent flowering and costs of reproduction affect population dynamics in a tuberous perennial woodland orchid. J Ecol 98:1204–1215Google Scholar
  35. Jacquemyn H, Brys R, Davison R, Tuljapurkar S, Jongejans E (2012) Stochastic LTRE analysis of the effects of herbivory on the population dynamics of a perennial grassland herb. Oikos 121:211–218Google Scholar
  36. Jäkäläniemi A, Crone EE, Närhi P, Tuomi J (2011) Orchids do not pay costs at emergence for prolonged dormancy. Ecology 92:1538–1543CrossRefPubMedGoogle Scholar
  37. Jongejans E, de Kroon H (2005) Space versus time variation in the population dynamics of three co-occurring perennial herbs. J Ecol 93:681–692CrossRefGoogle Scholar
  38. Jongejans E, de Vere N, de Kroon H (2008) Demographic vulnerability of the clonal and endangered meadow thistle. Pl Ecol 198:225–240CrossRefGoogle Scholar
  39. Kiviniemi K (2002) Population dynamics of Agrimonia eupatoria and Geum rivale two perennial grassland species. Pl Ecol 159:153–169CrossRefGoogle Scholar
  40. Klimeš L, Klimešová J, Hendriks R, van Groenendael J (1997) Clonal plant architecture: a comparative analysis of form and function. In de Kroon H, van Groenendael J (eds) The ecology and evolution of clonal plants. Backhuys Publishers, Leiden, The Netherlands, pp 1–29Google Scholar
  41. Klimešová J, Doležal J, Sammul M (2011) Evolutionary and organismic constraints on the relationship between spacer length and environmental conditions in clonal plants. Oikos 120:1110–1120CrossRefGoogle Scholar
  42. Klimešová J, Nobis MP, Herben T (2015) Senescence, ageing and death of the whole plant: morphological prerequisites and constraints of plant immortality. New Phytol 206:14–18CrossRefPubMedGoogle Scholar
  43. Klimešová J, Nobis MP, Herben T (2016) Links between shoot and plant longevity and plant economics spectrum: Environmental and demographic implications. Perspect Pl Ecol Evol Syst 22:55–62CrossRefGoogle Scholar
  44. Klimešová J, Danihelka J, Chrtek J, de Bello F, Herben T (2017) CLO-PLA: a database of clonal and bud-bank traits of the Central European flora. Ecology 98:1179CrossRefPubMedGoogle Scholar
  45. Lehtilä K Syrjänen K Leimu R García M B, Ehrlén J (2006) Habitat change and demography of Primula veris: identification of management targets. Conservation Biol 20:833–843Google Scholar
  46. Lemke T, Salguero-Gómez R (2016) Land use heterogeneity causes variation in demographic viability of a bioindicator of species-richness in protected fen grasslands. Populat Ecol 58:165–178CrossRefGoogle Scholar
  47. Lindborg R, Ehrlén J (2002) Evaluating the extinction risk of a perennial herb: demographic data versus historical records. Conservation Biol 16:683–690CrossRefGoogle Scholar
  48. Magda D, Duru M, Theau J-P (2004) Defining management rules for grasslands using weed demographic characteristics. Weed Sci 52:339–345CrossRefGoogle Scholar
  49. Marcante S, Winkler E, Erschbamer B (2009) Population dynamics along a primary succession gradient: Do alpine species fit into demographic succession theory? Ann Bot (Oxford) 103:1129–1143CrossRefGoogle Scholar
  50. Mildén M, Münzbergová Z, Herben T, Ehrlén J (2006) Metapopulation dynamics of a perennial plant, Succisa pratensis, in an agricultural landscape. Ecol Modelling 199:464–475CrossRefGoogle Scholar
  51. Münzbergová Z (2005) Determinants of species rarity: population growth rates of species sharing the same habitat. Amer J Bot 92:1987–1994CrossRefGoogle Scholar
  52. Münzbergová Z, Křivánek M, Bucharová A, Juklíčková V, Herben T (2005) Ramet performance in two tussock plants – Do the tussock-level parameters matter? Flora 200:275–284CrossRefGoogle Scholar
  53. Münzbergová Z (2007) Population dynamics of diploid and hexaploid populations of a perennial herb. Ann Bot (Oxford) 100:1259–1270CrossRefGoogle Scholar
  54. Nicolé F, Brzosko E, Till-Bottraud I (2005) Population viability analysis of Cypripedium calceolus in a protected area: longevity, stability and persistence. J Ecol 93:716–726CrossRefGoogle Scholar
  55. Ott JP, Hartnett DC (2015) Vegetative reproduction and bud bank dynamics of the perennial grass Andropogon gerardii in mixedgrass and tallgrass prairie. Amer Midl Naturalist 174:14–32CrossRefGoogle Scholar
  56. Pfeifer M, Wiegand K, Heinrich W, Jetschke G (2006) Long-term demographic fluctuations in an orchid species driven by weather: implications for conservation planning. J Appl Ecol 43:313–324CrossRefGoogle Scholar
  57. Ramula S, Toivonen E, Mutikainen P (2007) Demographic consequences of pollen limitation and inbreeding depression in a gynodioecious herb. Int J Pl Sci 168:443–453CrossRefGoogle Scholar
  58. Salguero-Gómez R, Jones OR, Archer CR, Buckley YM, Che-Castaldo J, Caswell H, Hodgson D, Scheuerlein A, Conde DA, Brinks E, de Buhr H, Farack C, Gottschalk F, Hartmann A, Henning A, Hoppe G, Roemer G, Runge J, Ruoff T, Wille J, Zeh S, Davison R, Vieregg D, Baudisch A, Altwegg R, Colchero F, Dong M, de Kroon H, Lebreton JD, Metcalf CJE, Neel MM, Parker IM, Takada T, Valverde T, Vélez-Espino LA, Wardle GM, Franco M, Vaupel JW (2015) The COMPADRE Plant matrix database: an open online repository for plant demography. J Ecol 103:202–218CrossRefGoogle Scholar
  59. Salguero-Gómez R, Jones OR, Jongejans E, Blomberg SP, Hodgson DJ, Mbeau-Ache C, Zuidema PA, de Kroon H, Buckley YM (2016) Fast-slow continuum and reproductive strategies structure plant life-history variation worldwide. Proc Natl Acad Sci USA 113:230–235CrossRefPubMedGoogle Scholar
  60. Sarukhán J, Harper, JL (1973) Studies on plant demography – Ranunculus repens L. R. bulbosus L. and R. acris L. I. Population flux and survivorship. J Ecol 61:675–716Google Scholar
  61. Serebriakova TI (1977) On major architectural models of herbaceous perennials and patterns of their reconfiguration [in Russian]. Byull Moskovsk Obshch Isp Prir Otd Biol 82:112–128Google Scholar
  62. Silvertown J, Franco M, Pisanty I, Mendoza A (1993) Comparative plant demography – Relative importance of life-cycle components to the finite rate of increase in woody and herbaceous perennials. J Ecol 81:465–476CrossRefGoogle Scholar
  63. Söyrinki N (1938) Studien über die generative und vegetative Vermehrung der Samen-pflanzen in der alpinen Vegetation Petsamo Lappland. Ann Bot Soc Zool-Bot Fenn “Vanamo” 11:1–311Google Scholar
  64. Suzuki JI, Herben T, Krahulec F, Hara T (1999) Size and spatial pattern of Festuca rubra genets in a mountain grassland: its relevance to genet establishment and dynamics. J Ecol 87:942–954CrossRefGoogle Scholar
  65. Svensson BM, Carlsson BÅ, Karlsson PS, Nordell KO (1993) Comparative long-term demography of three species of Pinguicula. J Ecol 81:635–645CrossRefGoogle Scholar
  66. ter Braak CJF, Šmilauer P (2012) CANOCO reference manual and user's guide: software for ordination (version 5.0). Microcomputer Power, Ithaca, USAGoogle Scholar
  67. Valverde, T, Silvertown J. (1998) Variation in the demography of a woodland understorey herb (Primula vulgaris) along the forest regeneration cycle: projection matrix analysis. J Ecol 86:545–562CrossRefGoogle Scholar
  68. Wallin L, Svensson BM (2012) Reinforced traditional management is needed to save a declining meadow species. A demographic analysis. Folia Geobot 47:231–247CrossRefGoogle Scholar
  69. Weppler T, Stoll P, Stöcklin J (2006) The relative importance of sexual and clonal reproduction for population growth in the long-lived alpine plant Geum reptans. J Ecol 94:869–879CrossRefGoogle Scholar
  70. Widén B, Cronberg N, Widén M (1994) Genotypic diversity, molecular markers and spatial distribution of genets in clonal plants, a literature survey. Folia Geobot Phytotax 29:245–263CrossRefGoogle Scholar
  71. Wikberg S, Svensson BM (2006) Ramet dynamics in a centrifugally expanding clonal sedge: a matrix analysis. Pl Ecol 183:55–63CrossRefGoogle Scholar
  72. Winter S, Jung LS, Eckstein RL, Otte A, Donath TW, Kriechbaum M (2014) Control of the toxic plant Colchicum autumnale in semi- natural grasslands: effects of cutting treatments on demography and diversity. J Appl Ecol 51:524–533CrossRefGoogle Scholar
  73. Zhang LQ, Hytteborn H (1985) Effect of groundwater regime on development and distribution of Fritillaria meleagris. Holarc Ecol 8:237–244Google Scholar

Copyright information

© Institute of Botany, Academy of Sciences of the Czech Republic 2017

Authors and Affiliations

  1. 1.Department of Botany, Faculty of ScienceCharles UniversityPraha 2Czech Republic
  2. 2.Institute of BotanyAcademy of Science of the Czech RepublicPrůhoniceCzech Republic
  3. 3.Institute of BotanyAcademy of Science of the Czech RepublicTřeboňCzech Republic

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