(Upload on February 25 2015) [ 日本語 | English ]

Lifeform (生活型)

Mount Usu / Sarobetsu post-mined peatland
From left: Crater basin in 1986 and 2006. Cottongrass / Daylily


Spelling: life form, life-form or lifeform

Raunkiaer's life form
Raunkiaer's life form spectrum in the world
Raunkiaer, Christen Christensen
Bud dormancy and lifeform
Indicator species

Mode of life and species attributes
  1. Descriptive:
    1. Basic (Theopratos 300 BC)
    2. Complex (von Humbolt 1986, etc.)
  2. Ecological:
    1. Surviving the bad times (Raunkiaer 1905, 1907)
    2. Growth form (du Rietz 1931, etc.)
      1. Migule form
      2. Radicoid form (Numata & Asano 1969)

Lifeform (生活型)

  1. Growth form of the species, e.g., coniferous tree, broad-leaved tree, evergreen shrub, deciduous shrub, fern, graminoid, forb, parasite or saprophyte, dwarf woody plant
  2. Avoid stressful periods (cool/hot temperature, or drought) by dormant organs

Evolution of lifeform through nutrient aquisition

Maximizing the good times (良好タイミング最大化) (Boysen-Jensen 1937, Monsi 1960, Walter 1973, Schulze 1982)

The performance of a specific life span is measured by the marginal return of carbon per unit resource invested.
(carbon gain)/(carbon use)↑
(carbon use)/(water use)↑
(carbon gain)/(nutrient use)↑

Plant form adaptations to adjust this ratio
  1. Plant life span (annuals vs. perennials)
  2. Carbon allocation (photosynthesis vs. non-photosynthesis)
  3. Leaf life span (evergreen vs. deciduous)
Some factors favoring evergreen over deciduous
  1. high UV
  2. high atmospheric drought (low humidity, high wind)
  3. Low nutrients
  4. Periodic early-season frost
  5. Short growing season
  6. Dry summer - wet summer

Wetland plants

hydrophyte (水生植物) = aquatic plants
hygrophyte (湿生植物/水生植物) = between hydorphytes and mesophytes, not developing special organs for water conditions
halophyte (塩生植物): tolerant to salts
mesophytes (中生植物): most plants
xerophyte (乾生植物): adapted to

uniform dryness
periodic dryness
Ex. mangrove

Raunkiaer's life form (dormancy form) (ラウンケアの生活型)

Species classified into a few life forms based on the place of the dormant organs during adverse seasons, i.e., cold and/or dry seasons
Table. Raunkiaer's life form or dormancy form spectra (Raunkiaer 1934) Epiphyte (着生植物): a plant growing non-parasitically on another plant (such as a tree) → tropical regions
Sclerophyte (硬葉植物): a evergreen and xerophytic plant with thick, hard leaves
Fig. Raunkiaer's life form. Brown squares indicate the positions of dormant buds. (a) phanerophytes, (b) nanophanerophytes - chamaephytes, (c) hemicryptophytes, (d) geophytes, (e) therophytes, (f) epiphytes

Raunkiaer's life form spectrum

Table 4. Selected Raunkiaerian life-form spectra for certain major climate. n: number of species examined. (Cain 1950)
  Vegetation type  climate and locality       Life form
                                            n Ph Ch  H G/HH Th

  Normal spectrum (world standard)       1000 46  9 26   6  13
  Rainforest, Queensland (2)              141 96  2      2
  Subtropical evergreen, Matheran, India  361 65 17  2   5  10
  Desert, Transcaspian lowlands (3)       768 11  7 27  14  41
  Mediterranean olive forest, Creta (4)  1571  9 15 27  10  38
  Steppe, Akron, Colorado (5)              79    19 58   8  15
  Temperate deciduous broad-leaf         1453 15  2 49  22  12
      forest, Connecticut (6)
  Tundra, Spisbergen (1)                  110  1 22 60  15   2

  1. Raunkiaer C. 1934
  2. Cromer DAN & Pryor LD. 1942. A contribution to rainforest ecology. Proc. Linn. Soc. New So. Wales 67: 249-268
  3. Paulsen O. 1912. The second Danish Pamir expedition. Studies on the vegetation of the Transcaspian lowland. Arbejder fra den bot. Have i Københaven 90: 1-279
  4. Turrill WB. 1929. The plant life of the Balkan Peninsula. A phytographic study. 1-490
  5. Paulsen O. 1915. Some remarks on the desert vegetation of America. Plant World 18: 155-161
  6. Ennis B. 1928. The life forms of Connecticut plants and their significance in relation to climate. Conn. State Geol. Nat. Hist. Surv., Bull. 43: 1-100

Bud dormancy and lifeform

Table. Depth of bud dormancy, structures of dormant bud and life form of herbs in the temperate region. (After Yoshie's seminar)
Species Depth of
bud dormancy
Type of
Type of
dormant bud
of shoot
1.Erigeron annuus 34IAHprWG-SG
2Oenothera biennis 36IAHprWG-SG
3Geum aleppicum 67IAHprEG
4Rumex acetosella 34IAHprSG
5Viola sp. 67IAHrSG
6Trifolium repens 55IAHSG
7Plantago lanceolata 66IAHrEG
8Plantago asiatica 234IIBHrSG
9Leibnitzia anandria 5411IIBHrSg
10Solidago virga-aurea 3912IIBprotHSG
11Aster sp. 1919IIBbprotHSG
12Solidago altissima 4518IIBbprotHSG
13Artemisia japonica 378IIBprotHSG
14Artemisia montana 5410IIBprotHSG
15Anaphalis margaritacea var. angustior 3917IIBprotHSG
16Sanguisorba tenuifolia var. tenuifolia f. alba 5215IIBprotHSG
17Tiarella polyphylla 1910IICChEG
18Pyrola incarnata 5916IICcChEG
19Pachysandra terminalis 2511IICcChEG
20Agrimonia japonica 237IIBbGrhSG
21Geranium thunbergii1713IICGrhSG
22Filipendula camtschatica(300)11IIIBGrhSG
23Helianthus tuberosus30033IIIBGstSG
24Aster glehnii300240IIIBGrhSG
25Adenocaulon himalaicum30060IIIBGrhSG
26Cacalia hastata var. orientalis30024IIIBGrhSG
27Polygonum filiforme30031IIIBGrhSG
28Phrma leptostachya var. asiatica30023IIIB-CGrhSG
29Cacalia delphiniifolia30019IIIB-CGrhSG
30Cacalia auriculata var. kamtschatica30016IIIB-CGrhSG
31Diphylleia grayi30043IIICGrhSG
32Petasites japonicus var.giganteus(300)8IIICGrhSG
33Lilium cordatum var. glehnii30053IIICGbulbSG
34Trillium smallii30046IIICcGrhSG
35Trillium tschonskii30050IIICcGrhSG
36Trillium kamtschaticum30041IIICcGrhSG
37Paris tetraphylla30025IIICcGrhSG
38Chloranthus japonicus30033IIICcGrhSG
39Erythronium japonicum30060IIICcGbulbSPG
40Gagea lutea30024IIICcGbulbSPG
41Anemone raddeana var. integra30031IIICGrhSPG
42Panax japonicus30028IIICGrhSG
43Maianthemum dilatatum(300)10IIICcGrhSG

*: 300 and 240 - No bud burst within 300 days and 240 days, respectively.
*: (300) - A few buds burst within 300 days.
Bb: Some buds had swollen or burst slightly in late October.
Cc: Shoot which develop in the following vegetative period are performed in the dormant bud.
*: Hpr - Partial rosette plant of hemicryptophytes, Hr - Rosette plant of hemicryptophytes, protoH - protohemicryptophyte, Ch - Chamaephyte, Grh - Rhizome geophyte, Gst - Stem-tuber geophyte, and Gbulb-Bulb geophyte.
++: WC - Wintergreen, SC - Summergreen, EG-Evergreen, SPC-Springgreen.

Indicator species (指標種)

Species that indicates the characteristics of the environmental conditions, such as serpentine.


≈ biomonitor
Plant, animal or chemicals used to monitor an environment and/or ecosystem
the measurement of the body burden of toxic chemical compounds, elements, or their metabolites, in biological substances

Methods and procedures on determining indicator species

    functional approach: easily-recognizable functional groups that represent adaptive strategies

    literature, asking
    direct ordination inverse analysis
    binary discriminant analysis: basically χ²-test
    Grime-strategy analysis: basically life form
    life or growth form
    Klinka life form type → weighted average
    it may be useless what environment(s) is(are) indicated is decided before making indicator species

  1. sample the vegetation
  2. omit indifferent species (usually common species)
  3. spectrum approach
    Frequency species (j) in attribute category (k) (e.g., moisture 1 → 6)
    F = [sum of species (i)j, k]/[total of all cover(ΣiΣjCi, j, k)] × 100, sum of species (i)j, k = Cijk
  4. indicator index II ... weighting factor analysis
    weight = Zj ... various weighting methods
    IIk = Σ(each species weight)/Σ(all species weight) × 10

Indicator plant

Hypochaeris radicata

Plant species used as indicator species.
Plants can not move greatly except when they are seeds (and pollens). Therefore, each plant establishes in a respective soil type, topography and climate, and generally indicates the environments better than animals.

Phragmites australis - indicates wet habitat
Drosera rotundifolia - indicates nutrient-poor habitat
exotic species - indicates urbanization

Plantago lanceorata

Environmental factors
pH, nutrients, moisture, light, temperature, salinity, etc.
+ disturbance (grazing, fire frequency and intensity, flood)
→ assumption: the past environment s could be estimated based on the established plants
+ successional status
Site potential

Lamium purpureum

Historical condition

Life form and biotic indicators

individual species
morphological patterns

life form spectrum
Ex. WW (western Washington): a few annuals ↔ EW: many → related to precipitation, wildfire and openness

(Grime 1973)

  1. The indicator species concept can be extended to higher taxa. In general, what might each of these taxa indicate in our region: Orchidaceae; Asteraceae; Poaceae; Carex; Vaccinium, shrub Rosaceae; Ericaceae. For each group, name a species that is an exception to the general rule.
  2. For each of conditions listed, list several excellent indicators for low, medium and high levels of the condition: soil moisture, soil pH, salinity, grazing, trampling, nutrients, light, fire, and bioclimatic zone. List other conditions that could be inferred from species present on a site.

(Bartelheimer & Poschlod 2016)

Ellenberg indicator value
can be used as a numerical system to classify species habitat niches and their peak occurrence along gradients
The picture emerging is that multiple determinants can be identified for:
N (nitrogen/nutrients)

1 = only growing on the poorest soils → 9 = only growing on excessively nitrogen (nutrient)-rich soils

R (soil reaction numbers)

1 = strong acidity (never moderately acidic or alkaline) → 9 = alkaline and calcareous conditions, only calcareous soils

M (soil moisture numbers): 1-12

1 = species only on dry soils → 9 = only on wet or hypoxic soils
10–12 = aquatic species

L (light numbers)

1 = deep shadow with 1–30% of full light available → 9 = only growing in full light to minimum 50% of full light

C (continentality)

the distance to the sea in Central Europe, where 'eucontinental' (C = 9) represents a species peak distribution at a far distance to the sea and 'euoceanic' (C = 1) indicates close proximity to the sea

T (tmperature numbers)

predominantly relate to species occurrence at respective elevations above sea level (lower T) correspond to higher elevations) but additionally include planar/especially warm habitats in Europe

Proably this indicator value can not be applied to Japan.