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

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)↑

1. Plant life span (annuals vs. perennials)
2. Carbon allocation (photosynthesis vs. non-photosynthesis)
3. Leaf life span (evergreen vs. 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

uniform dryness
periodic dryness
Ex. mangrove

• Life-form [code] position of dormancy bud (simplified category), example
• Phanerophytes [P] (trees, including epiphyte and sclerophyte)
  • Macrophanerophytes [MM] > 8 m in height (tall tree), Larix kaempferi, Populus maximowiczii, Fagus crenata
  • Mesophanerophytes [M] = 2-8 m (short tree or shrub), Camellia japonica, Aucuba japonica
• Nanophanerophytes [N] = 30 cm - 2 m (shrub), Salix reinii, Ledum palustre
• Chamaephytes [Ch] < 30 cm, Ardisia japonica, Trifolium repens
• Hemicryptophytes [H] ground surface (perennial herb), Polygonum sachalinense, Taraxacum officinale
• Geophytes [G] underground (perennial herb), Petasites japonicus var. giganteus, Phyllostachys heterocycla
• Helophytes [HH] underwater (perennial herb), Typha latifolia, Nymphaea tetragona, Ceratophyllum demersum, Hyderilla verticillata
• Therophytes [Th] producing seeds (annual herb), Polygonum longisetum

Raunkiaer's life form spectrum

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

Bioindicator

Biomonitoring

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

Hypochaeris radicata

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

Plantago lanceorata

Environmental factors

Site potential

Lamium purpureum

Historical condition

Life form and biotic indicators

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

(Grime 1973)

• Code, scientific name, common name and general indicator value of important species in Washington

Quiz

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

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

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

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

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

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

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