Growth form

Summary

Author

This article is modified from Perez Harguindeguy et al. (2013). The “New handbook for standardised measurement of plant functional traits worldwide” is a product of and is hosted by Nucleo Diversus (with additional Spanish translation). For more on this trait and on its context as part of the entire trait handbook visit its primary site Nucleo DiverSus at http://www.nucleodiversus.org/?lang=en

Contributing Author

Peter Ray

Summary

Growth form is mainly determined by the direction and extent of growth, and any branching of the main-shoot axis or axes. These affect canopy structure, including its height, and both the vertical and horizontal distribution of leaves. Growth form may be associated with ecophysiological adaptation in many ways, including maximising photosynthetic production, sheltering from severe climatic conditions, or optimising the height and positioning of the foliage to avoid or resist grazing by particular herbivores, with rosettes and prostrate growth forms being associated with high grazing pressure by mammals.

How to record

Growth form is a hierarchical trait assessed through field observation or descriptions or figures or photographs in the literature. Because we are classifying types along a continuum, intermediate forms, between the categories recognised here, may be encountered, as well as occasional unique forms lying outside any of these categories.

  1. Terrestrial, mechanically and nutritionally self-supporting plants
    1. Herbaceous plants have either no or at most modest secondary growth, with stem and root tissues that are rather soft compared with typical wood.
      1. Rosette plant. Leaves concentrated on a short, condensed section of stem or rhizome (see Category C under Clonality bud banks and below-ground storage organs), at or very close to the soil surface; with an inflorescence (or single-flower peduncle) bearing either no or reduced leaves (bracts) produced from the rosette axis, above ground level. Graminoids whose principal photosynthetic leaves are attached to the base of their aerial stems (e.g. -bunch grasses’) fall in this category.
      2. Elongated, leaf-bearing rhizomatous plant The permanent axis is an elongated rhizome that directly bears photosynthetic leaves that extend individually up into the light. The rhizome can be located either at or below ground level (e.g. Pteridium aquilinum (bracken fern), Viola spp., Iris spp.), or (epiphytes) on an above-ground support such as a tree branch. Aerial inflorescences (or single-flower peduncles) with either reduced leaves (bracts), or none, may grow out from the rhizome.
      3. Cushion plant (pulvinate form) Tightly packed foliage held close to soil surface, with relatively even and rounded canopy form (many alpine plants have this form).
      4. Extensive-stemmed herb develops elongated aerial stem(s) whose nodes bear photosynthetic leaves that are distributed nearly throughout the canopy of the plant, except when shed from its more basal parts during later growth, and lacking in distally developed inflorescences. Graminoids (rhizomatous or not) with leafy aerial stems fall here.
      5. Tussock Many individual shoots of a dense colony or clone grow upward, leaving behind a tough, mostly dead supporting column topped by living shoots with active leaves (e.g. the Arctic cotton grass, Eriophorum vaginatum).
    2. Semi-woody plants. Stem without secondary growth but often toughened by sclerification (or, alternatively, with relatively feeble, soft or -anomalous’ secondary growth).
      1. Palmoid Bears a rosette-like canopy of typically large, often compound leaves atop a usually thick (-pachycaulous’), columnar, unbranched or little-branched stem (e.g. palms (Pandanus), tree ferns). Certain tropical or alpine Asteraceae such as Espeletia spp., cycads, Dracaena, arborescent Yucca spp. and some Bombacaceae can be regarded as having this growth form, although their stems undergo more extensive secondary growth (see also -Corner model’ within the references below).
      2. Bambusoid An excurrently branched (cf. Point A.3.d.i below) trunk lacking or having only weak secondary growth is stiffened by sclerification to support a vertically extensive, sometimes tree-sized canopy (bamboos; various tall, herbaceous dicots such as Chenopodium, Amaranthus and Helianthus).
      3. Stem succulent A usually leafless photosynthetic stem with extensive, soft, water-storage tissue and only limited secondary growth (cacti, and cactoid plants of other families; most leaf succulents fall instead into one of the subclasses of Points A.1 or A.3).
    3. Woody plants develop extensive, usually tough, secondary xylem and phloem from vascular cambium, and corky outer bark from cork cambium (woody vines are covered in Point B.3 below).
      1. Prostrate subshrub Long-lived woody stem growing horizontally at ground level (examples include many Arctic willows and ericoids).
      2. Dwarf shrub, or subshrub, with usually multiple, ascending, woody stems less than 0.5 m tall.
      3. Shrub Woody plant between 0.5 m and ~5 m tall, with canopy typically carried by several trunks that are usually thinner and younger than typical mature tree trunks.
      4. Tree Woody plant usually >5 m tall, with main canopy elevated on a long-lived, substantial, usually single (but upwardly branching), trunk.
        1. Excurrent. Single main axis (trunk) extends up to, or almost to, the top, with shorter, ascending or horizontal branches giving a conical or (in mature trees) columnar form to the crown.
        2. Deliquescent Trunk divides, somewhere above its base, into two to several, more or less equal branches that continue branching upward to produce a wider, more flat-topped crown.
      5. Dwarf tree Morphology as in one of Types (i) or (ii) as for trees above, but substantially <5 m tall. Many forest understorey trees, but also in various climatically or nutritionally challenging, unshaded habitats, such as -pine barrens’, semi-deserts, certain tropical cloud forests, bogs and near-timberline vegetation.
  2. Plants structurally or nutritionally supported by other plants or by special physical features
    1. Epiphyte Plant that grows attached to the trunk or branch of a shrub or tree (or to anthropogenic supports) by aerial roots, normally without contact with the ground (e.g. many tropical orchids and Bromeliaceae).
    2. Lithophyte Plant that grows in or on rocks (e.g. many species of ferns, species of Nepenthes, Utricularia forestii, Cymbalaria muralis).
    3. Climber Plant that roots in the soil but relies on external support for its upward growth and leaf positioning, but sometimes only during early or late life phases.
      1. Herbaceous vine Usually attaches to its support either by twining or by means of tendrils.
      2. Woody vine, including liana Often attaches to a support by means of aerial roots.
      3. Scrambler Grows up through a sufficiently dense canopy of other plants, without any means of attachment (e.g. Galium spp.).
      4. Strangler May start epiphytically (but become soil-rooted) or by climbing from ground level. However, by secondary growth, it later becomes self-supporting, and may eventually envelope the initially supporting stem (e.g. certain tropical Ficus spp.).
    4. Submersed or floating hydrophyte Herbaceous, aquatic plant that relies on surrounding water for physical support. (Emergent hydrophytes (-helophytes’) mostly fall into one of the subgroups of Point A.1 above).
    5. Parasite or saprophyte obtains important nutritional needs directly or indirectly from other vascular plants (parasite) or from dead organic matter in the soil (saprophyte).

Literature references

References on theory, significance and large datasets:

Barkman J (1988) New systems of plant growth forms and phenological plant types. In -Plant form and vegetation structure’. (Eds MJA Werger, PJM Van der Aart, HJ During, JTA Verhoeven) pp. 9-44. SPB Academic Publishers: The Hague, The Netherlands. (and references therein)

Box EO (1996) Plant functional types and climate at the global scale. Journal of Vegetation Science 7, 309-320. doi:10.2307/3236274

Cain SA (1950) Life-forms and phytoclimate. Botanical Review 16, 1-32. doi:10.1007/BF02879783

Cramer W (1997) Using plant functional types in a global vegetation model. In -Plant functional types’. (Eds TM Smith, HH. Shugart, FI Woodward) pp. 271-288. Cambridge University Press: Cambridge, UK

Ellenberg H, Müller-Dombois D (1967) A key to Raunkiaer plant life forms with revised subdivisions. Berichte des geobotanischen Institutes der ETH. Stiftung Rübel 37, 56-73.

Ewel JJ, Bigelow SW (1996) Plant life forms and ecosystem functioning. In -Biodiversity and ecosystem processes in tropical forests’. (Eds GH Orians, R Dirzo, JH Cushman) pp. 101-126. Springer: Berlin.

Lüttge (1997) -Physiolgical ecology of tropical plants.’ Springer-Verlag: Berlin.

McIntyre S, Lavorel S (2001) Livestock grazing in subtropical pastures: steps in the analysis of attribute response and plant functional types. Journal of Ecology 89, 209-226. doi:10.1046/j.1365-2745.2001.00535.x

Medina E (1999) Tropical forests: diversity and function of dominant lifeforms. In -Handbook of plant functional ecology’. (Eds FI Pugnaire, F Valladares) pp. 407-448. Marcel Dekker: New York.

Richter M (1992) Methods of interpreting climatological conditions based on phytomorphological characteristics in the cordilleras of the Neotropics. Plant Research and Development 36, 89-114.

Rundel PW (1991) Shrub life-forms. In -Response of plants to multiple stresses’. (Eds HA Mooney, WE Winner, EJ Pell) pp. 345-370. Academic Press: San Diego, CA.

Whittaker RH (1975) -Communities and ecosystems.’ 2nd edn. Macmillan Publishing: New York.

More on methods:

Barkman J (1988) New systems of plant growth forms and phenological plant types. In -Plant form and vegetation structure’. (Eds MJA Werger, PJM Van der Aart, HJ During, JTA Verhoeven) pp. 9-44. SPB Academic Publishers: The Hague, The Netherlands. (and references therein)

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