Species

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Species (種)

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

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☛ plant taxonomy 植物分類学|Engler's syllabus エングラー体系] [alga 藻類|mosses 蘚苔類|fungi 菌類]

Species richness (種数)

The divergence of modern taxonomical groups (mostly divisions), based on the assumption of the origin of common ancestor.

Angiospermae    2-300,000 spp
   (12,000 gen)
Gymnospermae            800
Ferns                        11,000
Mosses                     24,000
Lichens                     20,000
Fungi                        36,000
Bacteria                      2,000

Variety: region - size

Asia

Table 3. Flora and fauna in East Asia. spp = number of species. end = percentages of endemic species. rpr = number of reproductive species. (World Resouces 2000-2001, WRI, 2001)
				mammal		aves		reptile		amphibia		vascular	plant
region	area × 10⁴ km²）	islands or continent	forest (%)	spp	end	rpr	end	spp	end	spp	end	spp	end
Japan	37	islands	68%	188	22%	250	8%	87	38%	61	74%	5,565	36%
Indonesisa	182	islands	60%	457	49%	1,530	27%	514	59%	285	40%	29,375	60%
Malaysia	33	islands	45%	300	12%	508	4%	350	20%	189	37%	15,500	23%
Philippines	30	islands	23%	158	65%	196	95%	190	84%	92	79%	8,931	39%
Singapore	0.06	islands	7%	85	1%	118	0%	140	0%	24	0%	2,282	0%
Sri Lanka	6	islands	33%	88	17%	250	10%	144	53%	39	51%	3,314	27%
Bangladesh	13	continent	8%	109	0%	295	0%	119	2%	19	0%	5,000	-
Bhutan	5	continent	60%	99	0%	448	0%	19	11%	24	0%	5,446	1%
Cambodia	18	continent	56%	123	0%	307	0%	82	1%	28	0%	-	-
China	933	continent	14%	400	21%	1,103	6%	340	24%	290	54%	32,200	56%
India	297	continent	22%	316	14%	926	6%	390	48%	209	58%	16,000	31%
North Korea	12	continent	50%	-	-	115	1%	19	11%	14	14%	2,898	4%
South Korea	10	continent	80%	49	0%	112	0%	25	4%	14	0%	2,898	8%
Laos	23	continent	52%	172	0%	487	0%	66	2%	37	5%	-	-
Mongolia	157	continent	6%	133	0%	426	0%	22	0%	6	0%	2,823	8%
Myanmar	66	continent	41%	251	2%	867	0%	203	18%	75	13%	7,000	15%
Nepal	14	continent	36%	181	1%	611	0%	100	1%	43	26%	6,973	5%
Pakistan	77	continent	3%	151	3%	375	0%	172	13%	17	24%	4,950	8%
Thailand	51	continent	24%	265	3%	616	0%	298	12%	112	19%	11,625	-
Vietnam	33	continent	27%	213	4%	535	2%	187	25%	80	34%	10,500	12%
Russian Federation	1,689	continent	45%	269	8%	628	2%	58	0%	41	0%	-	-
Europe (reference)
England	24	islands	8%	50	0%	230	0%	8	0%	7	0%	1,623	1%
Finland	30	continent	67%	60	0%	248	0%	5	0%	5	0%	1,102	-
France	55	continent	27%	93	0%	269	0%	32	3%	32	9%	4,630	3%
Germany	35	continent	31%	76	0%	239	0%	12	0%	20	0%	2,632	0%
Italy	29	continent	22%	90	3%	234	0%	40	3%	41	29%	5,599	13%
Spain	50	continent/ islands	16%	82	5%	278	2%	53	21%	28	14%	5,050	19%

Species (種)

Biological and taxonomic species

Table. Relationship between classification units and applicable sphere (Cain 1954, Simpson 1961)
	Units being classified	Sphere in which applicable
Taxonomic species (分類学的種)	Taxa; groups of morphologically similar individuals	Groups with morphological differences that are useful in formal classification
Biological species (生物学的種)	Sexually reproducing population systems	Sexual organisms on a single time level
Successional species (遷移種)	Phyletic lineages	Uniparental organisms, including asexual and parasexual organisms
Biosystematic species (系統学的種)	Fertility groups	Sexual plant groups which have been artificially hybridized in the experimental garden

Good species (良い種)

One taxonomic species = one biological species
Several taxonomic species = one biological species
Several taxonomic species = syngameon (sunsu Grant 1981)

Table. Biological species and stage
of divergence. Local race, population
level or variety

Geographical race
Allopatric semispecies
Sympatric semispecies
of hybrid origin
Allopatric biological sp.
Sympatric biological sp.
Species gorup
Syngameon

ssp. or var.
taxonomic sp.
taxonomic sp.
taxonomic sp.
taxonomic sp.
taxonomic sp.
series
series or section

Character (形質)

phylogeny (= natural classification)

genetics – micro-evolution
taxonomy – macro-evolution

(Maekawa 1950)

Characters in multifaceted nature (多面性の形質)

= characters that reflect the phylesis
1. structure of flaggella (鞭毛構造)

Ex. Phycomycetes (藻菌類)

Basidiomycetes (担子菌): basidiospore, dicaryon
Ascomycetes (子嚢菌): ascospore, monocaryon
Phycomycetes: coenocyte (多核体)

Chytridiales (ツボカビ類): uniflagellate flaggella (単毛)
Oomycetes (卵菌類): two comb-shaped flaggella
Zygomycetes (接合菌): conjugation (接合)

→ the common character in Phycomycetes is only noncellular structrue

2. chemical structural components in cell wall
3. assimilatory pigment → assimilates
4. formation mode and behavior of reproductive cells

Ex. Rhizophydium (Zygomycetes)________♀: transfer of cell contents

______________________________♂: active reproductive behavior
Ex. Pyronema (Ascomycetes)

ascogonium (造嚢器) / antheridium (造卵器) → trichogyne (受精毛) → ascogenous hypha (造嚢糸), forming 2n → forming ascus (担子器)
→ oomycetes is phylogenetically different from the other phycomycetes → establishing Dichotomycetes (二毛菌) (Maekawa 1947)

5. biosynthetic pathway

Ex. biosynthesis of lysin – two pathways (Vögel 1960)

AAA-pathway: α-KG acid + CH3COO-Coenzyme A → L-α-aminoadaphic δ-semialdehyde → lysin

bacteria, cyanophyta, green algae, dichonomycetes

DAP-pathway: L-asparatic-α-semialdehyde + pyruvic acid → L-α-ε-diamino pimelic acid → lysin

Phycomycetes, ascomycetes, basidiomycetes

→ dichonomycetes is essentially different from the other fungal taxa

Characters in tendency (偏向性の形質)

1. number of flagella (鞭毛数)
2. life phase (生活相)

Ex. life phase (Maekawa 1947) – considered as interfacial film

1. virus phase
2. amoeboid phase
3. monadoid phase (鞭毛相) - primitive
4. cystoid phase (包膜相) – cell wall present = high environmental adaptation
5. gymnoid phase (裸出相) – specific for animals

phase

Speciation (種分化)

Character displacement (形質置換)

Fig. Character displacement inferred from character states in sympatry (同所) and allopatry (異所) (Grant 1972).

Ex. Drosophilla pavlistrum complex

allopatry ___________sympatry
speciation
_____________________premating isolation
_____________________(the mechanisms unknown)

Ex. Litoria ewingii vs L. verreauxi

differences in sounds (or calls)

Ex. Phlox

____________allopatry sympatry
Phlox pilosa___pink______white
P. globerrina___pink______pink

Varation (変異)

1942 Mayer E: Geographycal race (≈ ecotype)

relationships of distribution between relative species: sympatric vs allopatric

1949, 1963 Anderson E: Introgressive hybridization

speciation
Introgression = immigration or penetration

introgressive genes to different race(s) in allopatric region(s), due to geological factors, etc.

1963 Anderson 1963: Introgression = hybridization of habitats
Summary
individuals [genes] → local population (forma) [genes, chromosome] → ecotype (variety) [genes, chromosome] → ecospecies (species) → coenospecies (species) → comparium (species or genus)
ramet (ラメート): all shoots have the same genotype

Ex. populations propagated by division = one clone

Evolutional entities (or units)

individuals
├─────┐
↓              ↓
ramets  biotype: many heterogenous genotypes
             (genotype) - the same genotypes
clones      ↓
             species
                ↓ closely remoted, related

Race (レース)

Geographical races

continuous geographical race – inappropriate because race is discontinuous
disjunctive gegographical race

Ex. Gilia latifolia (continuous distribution), G. leptantha (disjunctive) – sympatric = can breed between the races

ecological (geograpical) race

Ex. savanna, slope of alpine zone
Ex. Gilia achilleaefolia

Sun races: large flower – pollinator = insect
Shade races: small flower – self-crossing

(Huxley 1938)

Cline (クライン)

Ex. Chenopodium rubrum (Cumming 1963)

34.3°N      50.2°N            60.8°N          62.8°N
California  Suskatchwan  Yukon           Yukon
Short-day                         Quantitatic    Mostly ventral
short-day  →→→→→→→→→→→→  long-day

also observed on cultivated plants
likely reversed pattern shown by plants of southern elements

Selection factors on race formation

1922 Turesson: race is classified into two types:

1. adaptation to major environmental factors in a given habitat
2. replacement between species

Ex. Geum, Potentilla, Campanula, etc. → edaphic factor / biotic factor

Isolation mechanisms (隔離機構)

Table 2.1. classification of isolation types
1 Geographical isolation (地理的隔離)
2 Ecological isolation (生態的隔離)
3 Reproductive isolation (生殖的隔離)

a) Seasonal isolation
b) Diurnal isolation
c) Floral isolation
d) Incompatibility (不和合性)
e) Hybrid inviability or weakness
f) Hybrid sterility
g) Self fertilization

1-3c: External isolation or premating isolation
3d-g: Internal isolation or postmating isolation

*: mechanical isolation, ethological isolation

Semi-species (半種)

1. progressive branching (漸進的分岐) Ex. Catalpa bignoides (USA), C. ovata (China)
Ex. Pleatnus racemosa (California), P. wrightii (Arizona)

→ convenient when treated as taxonomic species
= not good biological species

Overlapping rings (重複環)

rings Fig. Overlapping rings
representative species groups that show overlapping rings: Gilia capitata group, Ipomopsis aggregata group, Aquilegia carysantha group, Mimulus longiflorus group, Pinus ponderosa group

2. natural hybridization (自然交雑)
a. hybridized group

semi-sepcies drived by semi-species: Ex. Geum rivale × G. urbanum, Aquilegia formosa × A. pubescens

Fig. 18.4. Laeves of Quercus alba and two of its hybrids (Hardin 1975)

b. permeable cross population (浸透性交雑個体群)

Iris hexagona var. giganticaerulea × I. fulva
→ conducted permeable cross after ecological isolation occurred (need revisit)

Ecological species (生態種)

Ecotype (エコタイプ) ≈ ecospecies (生態種)

Same species with different characters along environmental gradient
ecotype

Product arising as a result of the response of the genotype of an organism to the particular habitat in which it lives. For example, Plantago maritima has a height of about 17.5 cm in waterlogged mud and 56 cm in fertile meadow. Cf. phenotpye plasticity

Ecocline (エコクライン)

consisting of ecotypes that exhibit gradual phenotypic and/or genetic differences along environmental gradients

Isozyme (アイソザイム)

= enzyme gene or isoenzyme
allozyme = allelic isozyme
multiple gene = non-allelic isozyme

Hybrid (雑種)

Def. the offspring resulting from combining the qualities of two organisms of different taxa through sexual reproduction
Ancient Greek Aristoteles: consider the presence of hybrid

discussed in "De Generatione Animalium (Generation of animals)"

1761 Kölreuter, Joseph Gottlieb (1733-1806)

plant hybridization experiments - confirmed the hybrids
discovered paternal inheritance

⇒ species immutability → evolution

Intraspecific hybrid (種内雑種)

progeny between the same species but different infraspecific taxa

Interspecific, interspecies or species hybrid (種間雑種)

progeny from the mating of individuals from different species

Ex. mule (ラバ) = ♂donky × ♀horse
Ex. leopon (レオポン) = ♂panther × ♀lion
Ex. liger (ライガー) = ♂lion × ♀tiger

☛ breed improvement (品種改良): breeding, embryology
1780 Spallanzani: intraspecific hybridization conducted using dogs
1776 Hunter, John (1728-1793, England), anatomist

firstly reported successful artificial insemination

1907 Ivanov, Ll'ya Ivanovich (1834-1901): applyed to livestock science

horse hybridization → cow, sheep, etc.

Animal and plant

Mutatiion
X-ray

1949 γ-field (Brookhaven National Laboratory, USA)

☛ bioethics

Breed improvement (品種改良)

Animal

Artificial insemination
1952 Polge & Rowson: established cryopreservation of bovine semen

Plants

Artificial pollination
Grafting (接木)
Tissue culture (組織培養) ☛ Daucus carota (ニンジン)