樹木學及實習 Dendrology and Practice 植物型態 Plant Morphology

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樹木學及實習 Dendrology and Practice

植物型態 Plant Morphology

國立臺灣大學森林環境暨資源學系鍾國芳 (Kuo-Fang Chung) School of Forestry and Resource Conservation, National Taiwan University

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How to classify plants?

• How do we put different plants in groups?• What is species? How to recognition and delimitate species? ( 物種的描述 )• Classification ( 分門別類 )

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What is a plant species? • Since the ancient Greek, a species, such as particular species of oak, was

considered the manifestation of a unique, unchangeable essence ( 本質 ). Any deviations from this essence were regarded as unimportant.

• Darwin’s “Origin of Species”

Steady state of natural populations

Superfecundity

Limited resource

Variation

Struggle for existence

Heritability of variation

Differential survival (natural selection)

Over many generations, evolution & speciation

Darwin’s observations and inferences leading to the Evolution theories


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Modern corn—a miracle of artificial selection ( 人擇 )

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What should we do with the species problems?

• Different groups of organisms differ widely in morphology, genes, ecology, geographic distribution, pollination ……

• The lack of consensus on species definition can partly attribute to idiosyncratic nature of the biodiversity.

• Each lineage is unique.• Morphology is the most accessible source of data about evolutionary relationship

and is the only basis for the easy recognition of most species today.• Taxa are hypotheses, open to repeat testing as new data or methods of analysis

become available (e.g., DNA barcoding).

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Darwin placed Linneaus scheme of classification within an evolutionary framework—Descent with modification

“… species are nested within genera, genera within families, and so forth because of a history of descent with modification within evolutionary lineages—“…the view that an arrangement (hierarchical taxonomic system) is only so far natural as it is genealogical (evolutionary)”

http://en.wikipedia.org/wiki/Public_domain

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The importance of (organism) classification

• Organizing knowledge of the natural world into a system aids communication and helps us to understand better the organisms that surround us and upon which we depend. ……. The basic function of words is that of naming (Lyons 1977), and in the human mind nothing really exists or can be communicated without a name.

Christenhusz et al. (2011) Preface to “Linear sequence, classification, synonymy, and bibliography of vascular plants: Lycophytes, ferns, gymnosperms and angiosperms” Phytotaxa 19: 4-6.


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Classification: naming ( 命名 ), grouping ( 歸類 ), and naming the groups

• Names: • Purposes of classification:

• Easy to use• Stable• An aid to memory• Predictive• concise

• The logic of classification:• Hierarchical, groups nested within groups

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The hierarchical ( 階層 ) natural of classification—groups nested within groups


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Classification: naming and grouping

• Effective and efficient communication• Think about the amount of information a name carries!• The degree of confidence on a classification system.• Reducing the heterogeneity of a group of objects• Increase our understanding and ability to predict the characteristics of any one of the

objects.• Scale, hierarchical• Purpose-driven (no single best classification)• Not a static, once-and-for-all activity

• Anything can be classified, but modern biological classification is different because it involves evolutionary relationships.

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植物分類學 Plant Classification/Taxonomy/Systematics

• 植物分類學• Classification ( 分類學 ): classifying• Taxonomy ( 命名學 ): Naming (taxon/taxa: 分類群 )• Systematics ( 系統分類學 ): classification + taxonomy + phylogenetics ( 親緣關係、譜系關係 )

• 植物辨識 (plant identification)• 野外工作、標本館、形態學

• 植物命名 (nomenclature, taxonomy)• 植物命名法規 (International Code of Botanical Nomenclature)

• 分類文獻 (priority)

• 植物親緣關係的重建 (phylogenetics, systematics)• 穩定且能反映物種演化的分類架構 (Classification)

• Angiosperm Phylogeny Group Classification (APG III)

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How to classify plants?

• Hierarchical taxonomic system is so far natural only as it is genealogical (evolutionary)

• How do we put different plants in groups?

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How will you classify? Which plants should be classified as groups?


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Plant habit is important? (woody vs. herbaceous plants)


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•Plant habit is important? (woody vs. herbaceous plants)•Leaf shape is more important? (elliptic vs. linear)


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•Plant habit is important? (woody vs. herbaceous plants)•Leaf shape is more important? (elliptic vs. linear)•Number of petals is more important (5 vs. 4)


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•Plant habit is important? (woody vs. herbaceous plants)•Leaf shape is more important? (elliptic vs. linear)•Number of petals is more important (5 vs. 4)•Color of petals is more important (yellow vs. red)


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•Plant habit is important? (woody vs. herbaceous plants)•Leaf shape is more important? (elliptic vs. linear)•Number of petals is more important (5 vs. 4)•Color of petals is more important (yellow vs. red)•Number of stamen is more important (5 vs. 4 vs. 2)


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•Plant habit is important? (woody vs. herbaceous plants)•Leaf shape is more important? (elliptic vs. linear)•Number of petals is more important (5 vs. 4)•Color of petals is more important (yellow vs. red)•Number of stamen is more important (5 vs. 4 vs. 2)•Pollen surface is more important (smooth vs. spiny)


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• Why should I believe you?• A deeper understanding of morphological characters• developmental, ontongenetic studies


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The tradition of taxonomy……

“Young taxonomists are trained like performing monkeys, almost wholly by imitation, and that only in the rarest cases are they given any instruction in taxonomy theory” —Cain (1959)

“…… because of the subjective nature of the problem, it is difficult to lay down any hard and fast procedure for attaining satisfactory results……” —Mayr, Linsley, and Usinger (1953)

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Giants in plant taxonomy and their classification systems

• Carolus Linnaeus (L.) (1707-1778)‒ Species plantarum (1753), starting point of plant taxonomy

• de Jussieu‒ Genera plantarum (1789)—the idea of family

• Augustin Pyramus de Candolle (DC.)‒ Prodromus systematis naturalis regni vegetabilis (1824-1873)

• George Bentham and Joseph D. Hooker‒ Genera plantarum (1862-1883)

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Giants in plant taxonomy and their classification systems (cont.)

• Engler and Prantl—Die Naturlichen Pflanzenfamilien (1889-1907)• C. E. Bessey—The Essentials of Botany (1884)• J. Hutchinson—The Families of Flowering Plants (1973)• A. Takhtajan—Outline of the Classification of Flowering Plants (1980)• A. Croquist—An Integrated System of Classification of Flowering Plants (1981)• Angiosperm Phylogeny Group (1998- )

‒ Hopefully, from now on, this is the only system we have to remember and follow!

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分類學原理 (Principles of Classification)

• How to classify plants? How do we put different plants in groups?‒ By comparing characters possessed by different plants and then placing them

in groups, guided by trained intuition, almost by instinct, taxonomy is largely still practicing this way ( ～ 1960’s)

‒ By calculating the overall similarity of all (as many as possible) characters that you can observed—phenetics ( 數值分類學 / 相似性分類學 ) (1960’s～ )

‒ By tracking the transformation of character state series and find the shortest possible trajectory (among the numerous possibilities) of all character transformations—cladistics [ 支序 ( 分類 ) 學 ] (1960’s ～ )

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Numerical Phenetics (numerical taxonomy)數值分類、表形學派• P.H.A. Sneath (bacteriologists) and R.R. Sokal (biostatistist, anthropologist)• Searching for objective classification based on purely numerical (phenetic),

overall similarity among taxa.• Hope to recover history but did not intend to reconstruct phylogenetic

relationship.


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特徵 (character) 與特徵狀態 (character state)

•莖 : 木本 (woody) vs. 草本 (herbaceous)•葉形 : 橢圓形 (elliptic) vs. 線形 (linear)•花瓣數目 : 5 vs. 4•花瓣顏色 : 黃 (yellow) vs. 紅 (red)•雄蕊數目 : 5 vs. 4. vs. 2•花粉表面 : 平滑 (smooth) vs. 有刺 (spiny)


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葉形莖花瓣數目花瓣顏色雄蕊數目花粉表面X. alba E W 5 R 4 Sp

X. lutea E H 5 R 4 Sm

X. nigra L W 4 Y 2 Sm

X. purpurea L W 4 Y 2 Sp

X. rubens L W 4 Y 4 Sm

X. elliptica E W 5 Y 5 Sm

特徵資料矩陣Character data matrix•莖 : 木本 (woody) vs. 草本 (herbaceous)•葉形 : 橢圓形 (elliptic) vs. 線形 (linear)•花瓣數目 : 5 vs. 4•花瓣顏色 : 黃 (yellow) vs. 紅 (red)•雄蕊數目 : 5 vs. 4. vs. 2•花粉表面 : 平滑 (smooth) vs. 有刺 (spiny)



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From data matrix to distance table

• Calculate the differences between two plants

‒ X. alba: E W 5 R 4 Sp‒ X. lutea:E H 5 R 4 Sm‒ difference = 2‒ X. alba: E W 5 R 4 Sp‒ X: nigra:L W 4 Y 2 Sm‒ difference = 5

• Fill the matrix

A B C D E FA 0 2 5 4 4 3B 2 0 5 6 4 3C 5 5 0 1 1 3D 4 6 1 0 2 4E 4 4 1 2 0 3F 3 3 3 4 3 0

葉形莖花瓣數目花瓣顏色雄蕊數目花粉表面A- X. alba E W 5 R 4 Sp

B- X. lutea E H 5 R 4 Sm

C- X. nigra L W 4 Y 2 Sm

D- X. purpurea L W 4 Y 2 Sp

E- X. rubens L W 4 Y 4 Sm

F- X. elliptica E W 5 Y 5 Sm



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Unweighted Pair Group Method with Arithmatic Mean (UPGMA) ( 無權重群組算數平均法 )

• clustering the pair of plants (operational taxonomic units/OTUs) with the smallest distance (and divided by 2)

• re-calculating the distance matrix‒ Dist (A,CD) = [(DistAC + DistAD)]/2 =[(5+4)/2] = 4.5‒ Dist (B,CD) = [(DistBC + DistBD)]/2 =[(5+6)/2] = 5.5‒ Dist (CD,E) = [(DistCE + DistDE)]/2 =[(1+2)/2] = 1.5‒ Dist (CD,F) = [(DistCF + DistDF)/2] = =[(3+4)/2] = 3.5

CD

0.5

0.5

A B C D E FA 0 2 5 4 4 3B 2 0 5 6 4 3C 5 5 0 1 1 3D 4 6 1 0 2 4E 4 4 1 2 0 3F 3 3 3 4 3 0

A B CD E FA 0 2 4.5 4 3B 2 0 5.5 4 3CD

4.5 5.5 0 1.5 3.5

E 4 4 1.5 0 3F 3 3 3.5 3 0



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Unweighted Pair Group Method with Arithmatic Mean (UPGMA)

• re-clustering the pair of OTUs with the smallest distance

• re-calculating the distance matrix‒ Dist(A,CDE)=[(DistA,CD)+(DistAE)]/2

=[(4.5+4)/2]=4.25

‒ Dist (B,CDE) =[(DistB.CD)+(DistBE)]/2 =[(5.5+4)/2]=4.75

‒ Dist (CDE,F) =[(DistCD.F)+(DistEF)]/2 =[(3.5+3)/2]=3.25

A B CDE FA 0 2 4.25 3B 2 0 4.75 3

CDE 4.25 4.75 0 3.25F 3 3 3.25 0

A B CD E FA 0 2 4.5 4 3B 2 0 5.5 4 3

CD 4.5 5.5 0 1.5 3.5E 4 4 1.5 0 3F 3 3 3.5 3 0

CDE

0.75

0.751.5/2 = 0.75



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• re-calculating the distance matrix‒ Dist(AB,CDE)=[(DistA,CDE)+(DistB.CDE)]/2

=[(4.25+4.75)/2]=4.5

‒ Dist (AB,F) =[(DistAF)+(DistBF)]/2 =[(3+3)/2]=3

AB

1

12/2 = 1

AB CDE FAB 0 4.5 3

CDE 4.5 0 3.25F 3 3.25 0

A B CDE FA 0 2 4.25 3B 2 0 4.75 3

CDE 4.25 4.75 0 3.25F 3 3 3.25 0



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• re-calculating the distance matrix‒ Dist(ABF,CDE)=[(DistAB,CDE)+(DistF.CDE)]/2

=[(4.5+3.25)/2]=3.875

ABF

1

13/2 = 1.5

ABF CDEABF 0 3.875CDE 3.875 0

AB CDE FAB 0 4.5 3

CDE 4.5 0 3.25F 3 3.25 0

C- X. nigra D- X. purpurea E- X rubens F- X. elliptica A- X. alba B- X. lutea

0.00.51.01.52.0

3.875/2 = 1.9375




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C- X. nigra

D- X. purpurea

E- X rubens

F- X. elliptica

A- X. alba

B- X. lutea

0.00.51.01.52.0



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Advantages and pitfalls of phenetic approaches

Easy to operateComputationally fast Most alike ≠ most closely related (parallel and convergent evolution;

e.g. Euphorbia vs. cacti) Aiming for objective, but difficult to achieve

‒ Selection of characters• Phenetic approaches are now widely used in ecology

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Phylogenetic Systematics/Cladistics ( 支序學 )• Willi Hennig (German entomologist)• Hennig recognized the logical consequences of evolutionary constraint

—new structure and functions resulting from modification of existing ones.

‒ Evolution is recognized as changes from a preexisting ancestral (primitive, plesiomorphic) condition to a new, derived (advanced, apomorphic) condition.

• Reconstructing history of lineage diversification (cladogensis) by tracing the character transformation

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The concept of Characters and Homology( 特徵與同源性 )

• Homology ( 同源性 ): character shared by a group of organisms or taxa due to inheritance from a common ancestor.

• Homoplasy ( 同塑性、非同源相似性 ): character that appears similar among a group of organism or taxa, but the similarity is due to parallel or convergent evolution rather than inheritance from a common ancestor (wings of bats, birds, flying squirrel, butterflies, etc. .

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Homoplasy

• Convergent evolution ( 趨同演化 ): evolution of similar features independent in different evolutionary lineages, usually from different antecedent features or by different development pathways. (e.g., 仙人掌的刺 vs. 小檗的刺；鳥、蝙蝠、翼手龍的翅膀 )

• Parallel evolution ( 平行演化 ): the evolution of similar or identical features independently in related lineage, though usually to be based on similar modification of the same developmental pathway. (e.g., 寄生或腐生植物在不同分類群獨立演化 )

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Cladogram (phylogenetic tree)


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X nigra

X purpurea

X rubens

X alb

a

X lutea

Outgroup

0.5

X nigra X purpurea X rubens X alba X lutea Outgroup


cladogram (支序圖 ): depicting relationships only

diagonal linesperpendicular lines

X nigra X purpurea

X rubens X alba X lutea

Outgroup

0.5

phylogram (譜系圖 ): relationships with branch lengths

Unrooted tree (無根樹 )


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Anatomy of Cladistics1. Monophyletic (Holo-) group (單系 /源群 )

• Grouping (clade 支序群 ) includes all descendant taxa and their common ancestor

• All taxonomic groups (taxa) should be monophyletic.

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Paraphyletic group ( 並 / 側系群 )

• Grouping includes an ancestor and some but not all of its descendents


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Paraphyletic group ( 並 / 側系群 )



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Monophyly and paraphyly


The problem with ranked taxonomy!

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Angiosperm phylogeny


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Polyphyletic group ( 多源群 )

• Grouping includes taxa that trace back through two or more separate ancestors before reaching a common ancestor.


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Chirita 唇柱苣苔屬 , a polyphyletic taxon


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Cladogram ( 支序樹 ), ingroup ( 內群 ), sister group ( 姊妹群 ), outgroup ( 外群 )

• Ingroup: focal set of taxa within a monophyletic group.• Sister group: outgroup that most close to the ingroup• Outgroup is use to root the tree and provide an ancestor-descendent

orientation of characters.Additionaloutgroups

Sisteroutgroups

Ingroups

X Y Z


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葉形莖花瓣數目花瓣顏色雄蕊數目花粉表面X. alba E W 5 R 4 Sp

X. Lutea E H 5 R 4 Sm

X. Nigra L W 4 Y 2 Sm

X. Purpurea L W 4 Y 2 Sp

X. Rubens L W 4 Y 4 Sm

X. ellipticaOutgroup

E W 5 Y 5 Sm

E↔L W ↔ H 5 ↔ 4 Y ↔ R 5 → 4 → 2 Sp ↔ Sm


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1 2 3 4 5 6

X. alba 0 0 0 1 1 1

X. lutea 0 1 0 1 1 0

X. nigra 1 0 1 0 2 0

X. purpurea 1 0 1 0 2 1

X. rubens 1 0 1 0 1 0

Outgroup 0 0 0 0 0 0




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1 2 3 4 5 6

X. alba 0 0 0 1 1 1

X. lutea 0 1 0 1 1 0

X. nigra 1 0 1 0 2 0

X. purpurea 1 0 1 0 2 1

X. rubens 1 0 1 0 1 0

Outgroup 0 0 0 0 0 0

homoplasious character




52Tree length (樹長 ) = 8


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Tree length (樹長 ) = 8

Tree length (樹長 ) = 11



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Parsimony methods

• The goal is to find the most parsimonious tree (MP)• Parsimony: 檢約、吝嗇• The criteria are to calculate the changes of character states, i.e. the

evolutionary steps• First, we have to know the way to evaluate a given tree

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Molecular (DNA) data

• Molecular data are particular suitable for cladistic analyses• The number of characters is theoretically unlimited• Everyone interprets the data in the same way• Easier to assess homology• Less subject to selection; independent of the morphological data and

thus can be used to test previous taxonomic hypotheses.• Better in keeping the signature of the evolutionary history of

organisms

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Character optimization ( 特徵最佳化 )Sp_1: TCAGACGATTGTCAGACCATTGSp_2: TCAGTCGACTGTCAAACCATTGSp_3: TCGGTCAATTGTCAAACGATTGSp_4: TCGGTCAATTGTCAAACGATTG

Sp_1 is outgroup. Three different tree topologies.


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Cha

ract

er o

ptim

izat

ion:

cha

ract

er 3

0000000000111111111122

1234567890123456789012Sp_1TCAGACGATTGTCAGACCATTG

Sp_2TCAGTCGACTGTCAAACCATTG

Sp_3TCGGTCAATTGTCAAACGATTG


Step=2 Step=2 Step=1

Step=2

A→G

G→A

1 2 3

1

A A G G A A G G A G G A

A A G G


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0000000000111111111122





A→G

G →A

A

G

A

G

A

G

A

G

A

G

A

G

A

A

G

GStep=1Step=2Step=2

Step=2

A

G

A

GStep=2

A

G

A

A

G

G

A→GA

G A←G

Step=1

Unrooted trees


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Cha

ract

er o

ptim

izat

ion:

cha

ract

er 7

0000000000111111111122






Step=2

G→A

A→G

1 2 3

1

G G A A G G A A G A A G

G G A A


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1 2 3

Most parsimonious tree


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How many trees out there are we dealing with?

1 tree

3 trees

3 OTUs Root


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Four taxa

Unrooted trees

3 trees

Rooted trees 15 trees


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How many trees we need to search?

• The number of unrooted trees:

• The number of rooted trees:

)!2(2)!32(

2

nnN nrooted

)!3(2)!52(

3

nnN nunrooted

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Taxon number All possible unrooted tree number3 14 35 156 1057 9458 10,3959 135,13510 2,027,02511 34,459,42512 654,729,07513 13,749,310,57514 316,234,143,22515 7,905,853,580,62516 213,458,046,676,87517 6,190,283,353,629,37518 191,898,783,962,510,62519 6,332,659,870,762,850,62520 221,643,095,476,699,771,87521 8,200,794,532,637,891,559,37522 319,830,986,772,877,770,815,62523 13,113,070,457,687,988,603,440,62524 563,862,029,680,583,509,947,946,87525 25,373,791,335,626,257,947,657,609,375 >1029

- etc.


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In real world, how many trees can we search?

• Say a computer can evaluate 106 trees per second.• If we want to evaluate all of the trees for 25 taxa, we will need

x = 1029/106/60/60/24/365 = 3.17x1015

>三千兆年• Many ways to get-by this problems

‒ faster computer, software……..

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Phenetics vs. Cladistics

• Phentics: Algorithm—by defining a specific sequence of steps that leads to the determination of tree (UPGMA, NJ)—fast and you get one tree

• Cladistics: Optimality criteria—by defining a (optimal) criterion for comparing alternative phylogenies to one another and deciding which is better (parsimony, maximum likelihood, Bayesian)—slow and you can get many trees

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Which method is better in reflecting true evolutionary relationship?

Phenogram-UPGMA Maximum Parsimony Tree


X alba X lutea Outgroup X purpurea X nigra X rubens

0.00.51.01.52.0



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Angiosperm families with nodular nitrogen-fixing symbioses and the frequency of this association in each familyProkaryote Angiosperm family [Genera having root nodules/Approx. total

genera] Bold, family/genus in Taiwan

Rhizobiaceae Fabaceae 豆科 [530/730]Cannabaceae 大麻科 (Trema) [1/11]

Frankia (Actinorhizal plants, 放線根瘤植物 )

Betulaceae (Alnus) 樺木科 [1/6]Cassuarinacedae 木麻黃科 (Allocasuarina, Casuarina,

Gymnostoma) [4/4]Elaeagnaceae 胡頹子科 (Elaeagnus, Hippophae, Shepherdia)

[3/3]Myricaceae 楊梅科 (Morella, Myrica, Comptonia) [2/3]Rhamnaceae 鼠李科 (Ceanothus, Colletia, Discaria,

Kentrothamnus, Retanilla, Trevoa) [7/55]Rosaceae 薔薇科 (Cercocarpus, Chamaebatia , Purshia, Dryas)

[5/100]Datiscaceae 疣柱花科 (Datisca) [1/1]Coriariacaee 馬桑科 (Coriaria) [1/1]

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fabids 豆類

維管束植物約 400 個科中，僅有分屬於 10 個科的少數植物在根瘤與固氮細菌共生具固氮的能力，這 10 個科在 APG 的分類中，分屬於瓜目、殼斗目、薔薇目及豆目。These orders share a genetic predisposition for nitrogen fixation via root nodules, and this condition represents a possible synapomorphy for this group of four orders.

Cucurbitales 瓜目 Fagales 殼斗目 Rosales 薔薇目 Fabales 豆目 Celastrales 衛矛目 Oxalidales 酢醬草目 Malpighiales 黃褥花目 Zygophyllales 蒺藜目

Nitrogen-fixing clade


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版權聲明頁碼作品版權標示作者 /來源

3製表：國立臺灣大學鍾國芳參考來源： Charles Darwin, On the Origin of Species by Natural Selection, 1859.

4

Wikimedia Commons / Author: John Doebleyhttp://commons.wikimedia.org/wiki/File:Maize-teosinte.jpg，http://teosinte.wisc.edu/images.html，瀏覽日期： 2014/10/13 。本作品採創用CC 「姓名標示」 3.0 未本地化授權釋出。

6Wikimedia Commons / illustration by Charles Darwin, SVG file by Inductiveloadhttp://commons.wikimedia.org/wiki/File:Origin_of_Species.svg，瀏覽日期：2014/10/13 。本作品屬公共領域之著作。

6“…the view that an arrangement is only so far natural as it is genealogical”

Charles Darwin, On the Origin of Species by Natural Selection, 1859. 本作品屬公共領域之著作。

7Organizing knowledge …be communicated without a name.

Christenhusz et al. (2011) Preface to “Linear sequence, classification, synonymy, and bibliography of vascular plants: Lycophytes, ferns, gymnosperms and angiosperms” Phytotaxa 19: 4-6.瀏覽日期： 2014/08/23 。本作品依據著作權法第 46 、 52 、 65 條合理使用。

9圖表繪製：國立臺灣大學鍾國芳

http://commons.wikimedia.org/wiki/File:Maize-teosinte.jpg

http://teosinte.wisc.edu/images.html

http://commons.wikimedia.org/wiki/File:Origin_of_Species.svg

http://creativecommons.org/licenses/by/3.0/deed.en






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版權聲明頁碼作品版權標示作者 /來源13

繪製：國立臺灣大學鍾國芳

14

15, 16

17

18

19

20


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“Young taxonomists…in taxonomy theory”

Cain, A.J. (1959) The post-Linnaean development of taxonomy. Proceedings of the Linnean Society, London, 170, 234–244. 本作品依據著作權法第 46 、 52 、 65 條合理使用。

21“because of the subjective nature …results…”

Mayr, Ernst, Linsley, E. Gorton and Usinger, Robert L. 1953. Methods and Principles of Systematic Zoology. p. 168; also pp. 176–177. McGraw-Hill. 本作品依據著作權法第 46 、 52 、 65 條合理使用。

25、 26、 27、49、 50、 67


27繪製：國立臺灣大學鍾國芳







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32, 33繪製：國立臺灣大學鍾國芳








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圖表繪製：國立臺灣大學鍾國芳

39Wikimedia Commons / Author: Petter Bøckmanhttp://commons.wikimedia.org/wiki/File:Cladogram_vertebrata.jpg，瀏覽日期：2014/10/13 。本作品屬公共領域之著作。


41、 42、 46

Wikimedia Commons / Author: Petter Bøckmanhttp://commons.wikimedia.org/wiki/File:Monophyly-paraphyly-polyphyly.jpg，瀏覽日期： 2014/10/13 。本作品採創用 CC 「姓名標示 - 相同方式分享」 3.0 未本地化授權釋出。

43

Wikimedia Commons / Author: Petter Bøckmanhttp://commons.wikimedia.org/wiki/File:Traditional_Reptilia.pngOriginal Source: Stanislav Traykov http://commons.wikimedia.org/wiki/File:Paraphyletic.png ，瀏覽日期：2014/10/13 。本作品依據著作權法第 46 、 52 、 65 條合理使用。

44

Wikimedia Commons / Author: Petter Bøckmanhttp://commons.wikimedia.org/wiki/File:Traditional_Reptilia.jpg (Based on the file File:Paraphyletic.svg) ，瀏覽日期： 2014/10/13 。本作品採創用 CC0 1.0 通用 (CC0 1.0) 公眾領域授權釋出。

http://commons.wikimedia.org/wiki/File:Cladogram_vertebrata.jpg

http://commons.wikimedia.org/wiki/File:Monophyly-paraphyly-polyphyly.jpg

http://commons.wikimedia.org/wiki/File:Traditional_Reptilia.jpg

http://commons.wikimedia.org/wiki/File:Paraphyletic.svg




http://creativecommons.org/publicdomain/zero/1.0/deed.en



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76






56、 57、 59、

60







77












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