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Lifeform (生活型)

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

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Definition
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

Descriptive:
1. Basic (Theopratos 300 BC)
2. Complex (von Humbolt 1986, etc.)
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)

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
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

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

Some factors favoring evergreen over deciduous

high UV
high atmospheric drought (low humidity, high wind)
Low nutrients
Periodic early-season frost
Short growing season
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 or 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)

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

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
Lifeform

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
Normal spectrum (world standard)
Rainforest, Queensland (2)
Subtropical evergreen, Matheran, India
Desert, Transcaspian lowlands (3)
Mediterranean olive forest, Creta (4)
Steppe, Akron, Colorado (5)
Temperate deciduous broad-leaf
    forest, Connecticut (6)
Tundra, Spisbergen (1)

  n
1000
  141
  361
  768
1571
    79
1453

  110

Ph
46
96
65
11
9

15

1

Ch
  9
  2
17
  7
15
19
  2

22

  H
26

    2
  27
  27
  58
  49

  60

G/HH
    6
    2
    5
  14
  10
    8
  22

  15

Th
13

10
41
38
15
12

2

Raunkiaer C. 1934
Cromer DAN & Pryor LD. 1942. A contribution to rainforest ecology. Proc. Linn. Soc. New So. Wales 67: 249-268
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
Turrill WB. 1929. The plant life of the Balkan Peninsula. A phytographic study. 1-490
Paulsen O. 1915. Some remarks on the desert vegetation of America. Plant World 18: 155-161
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 dormancy	Type of dormant bud	Raunkiaer's lifeform	Phenology of shoot
		Oct.	Dec.
1.	Erigeron annuus	3	4	I	A	Hpr	WG-SG
2	Oenothera biennis	3	6	I	A	Hpr	WG-SG
3	Geum aleppicum	6	7	I	A	Hpr	EG
4	Rumex acetosella	3	4	I	A	Hpr	SG
5	Viola sp.	6	7	I	A	Hr	SG
6	Trifolium repens	5	5	I	A	H	SG
7	Plantago lanceolata	6	6	I	A	Hr	EG
8	Plantago asiatica	23	4	II	B	Hr	SG
9	Leibnitzia anandria	54	11	II	B	Hr	Sg
10	Solidago virga-aurea	39	12	II	B	protH	SG
11	Aster sp.	19	19	II	B^b	protH	SG
12	Solidago altissima	45	18	II	B^b	protH	SG
13	Artemisia japonica	37	8	II	B	protH	SG
14	Artemisia montana	54	10	II	B	protH	SG
15	Anaphalis margaritacea var. angustior	39	17	II	B	protH	SG
16	Sanguisorba tenuifolia var. tenuifolia f. alba	52	15	II	B	protH	SG
17	Tiarella polyphylla	19	10	II	C	Ch	EG
18	Pyrola incarnata	59	16	II	C^c	Ch	EG
19	Pachysandra terminalis	25	11	II	C^c	Ch	EG
20	Agrimonia japonica	23	7	II	B^b	Grh	SG
21	Geranium thunbergii	17	13	II	C	Grh	SG
22	Filipendula camtschatica	(300)	11	III	B	Grh	SG
23	Helianthus tuberosus	300	33	III	B	Gst	SG
24	Aster glehnii	300	240	III	B	Grh	SG
25	Adenocaulon himalaicum	300	60	III	B	Grh	SG
26	Cacalia hastata var. orientalis	300	24	III	B	Grh	SG
27	Polygonum filiforme	300	31	III	B	Grh	SG
28	Phrma leptostachya var. asiatica	300	23	III	B-C	Grh	SG
29	Cacalia delphiniifolia	300	19	III	B-C	Grh	SG
30	Cacalia auriculata var. kamtschatica	300	16	III	B-C	Grh	SG
31	Diphylleia grayi	300	43	III	C	Grh	SG
32	Petasites japonicus var.giganteus	(300)	8	III	C	Grh	SG
33	Lilium cordatum var. glehnii	300	53	III	C	Gbulb	SG
34	Trillium smallii	300	46	III	C^c	Grh	SG
35	Trillium tschonskii	300	50	III	C^c	Grh	SG
36	Trillium kamtschaticum	300	41	III	C^c	Grh	SG
37	Paris tetraphylla	300	25	III	C^c	Grh	SG
38	Chloranthus japonicus	300	33	III	C^c	Grh	SG
39	Erythronium japonicum	300	60	III	C^c	Gbulb	SPG
40	Gagea lutea	300	24	III	C^c	Gbulb	SPG
41	Anemone raddeana var. integra	300	31	III	C	Grh	SPG
42	Panax japonicus	300	28	III	C	Grh	SG
43	Maianthemum dilatatum	(300)	10	III	C^c	Grh	SG

*: 300 and 240 - No bud burst within 300 days and 240 days, respectively.
*: (300) - A few buds burst within 300 days.
B^b: Some buds had swollen or burst slightly in late October.
C^c: 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.

Bioindicator

≈ biomonitor
Plant, animal or chemicals used to monitor an environment and/or ecosystem

Biomonitoring

the measurement of the body burden of toxic chemical compounds, elements, or their metabolites, in biological substances

Methods and procedures on determining indicator species

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

sample the vegetation
omit indifferent species (usually common species)
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
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

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

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.
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.