Topic 04 History of Plant Systematics & Classification

Home , Carolus (plant), Genera Plantarum, Jardin des Plantes, Species Plantarum

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I. Primer on Classification A. Definition ‐process of organizing thoughts and ideas about the world around us.

I. Primer on Classification A. Definition ‐process of organizing thoughts and ideas about the world around us. B. Primary Operations 1. Grouping & Ranking

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Grouping results in a horizontal or coordinate arrangement of objects or organisms.

AB B D D B A D A C C C D objects or organisms. B E A B E C D E

Grouping results in a horizontal or coordinate arrangement of objects or organisms.

AB B D D B A D A C C C D objects or B E organisms. A B E C D E

Group A Group B Group C Group D Group E Groups of objects or organisms AB C D D E E B B D E A A A BB C C C DD E

Grouping results in a horizontal or coordinate arrangement of objects or organisms.

Group A Group B Group C Group D Group E

AB C D D E E B B D E A A A B B C C C DD E

groups have equal ranks or importance (e.g. species or genera, etc.)

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Ranking results in a vertical or hierarchical arrangement of those groups.

Group A Group B Group C Group D Group E

AB C D D E E B B D E A A A B B C C C DD E Genus 1 Genus 2 Genus 3

Family 1 Family 2

Order

I. Primer on Classification A. Definition ‐process of organizing thoughts and ideas about the world around us. B. Primary Operations 1. Grouping & Ranking 2. Universally found ‐from folk taxonomies to scientific classifications

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I. Primer on Classification A. Definition ‐process of organizing thoughts and ideas about the world around us. B. Primary Operations 1. Grouping & Ranking 2. Universally Found C. Artificial vs. Natural Systems 1. AS have little basis in reality a. a priori decisions about which chars are used at 1st, 2nd, 3rd – order ranks, etc.

e.g. the number of legs

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e.g. the number of legs

Bipods

Tripods

Tetrapods

I. Primer on Classification A. Definition ‐process of organizing thoughts and ideas about the world around us. B. Primary Operations 1. Grouping & Ranking 2. Universally Found C. Artificial vs. Natural Systems 1. AS have little basis in reality 2. NS: meant to recognize “real” groups a. a posteriori reasoning

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Group A

Group C

Group B

Group D

I. Primer on Classification

. 2. NS: meant to recognize “real” groups a. a posteriori reasoning b. Groups may be harder to recognize b/c no one character used to delimit each b. But have greater predictive power

II. The Early Days A. Theophrastus (ca. 371‐287 BC)

Given name: Tyrtamus

Nicknames:

‘Theophrastus’ by Aristotle (theos, god, phrasis, declare or tell)

‘Father of Botany’ by Linnaeus

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II. The Early Days A. Theophrastus (ca. 371‐287 BC) Important Works: • De Historia Plantarum • De Causis Plantarum (only Latin translations remain)

Characteristics: • Empirical approach to biology of plants • Primary subdivision based on habit • Subsidiary grps based on other chars • Fundamentally an AS.

Significance: • First systematic treatment of plants

II. The Early Days A. Theophrastus B. Herbalists 1. Herbals Characteristics: • Utilitarian compendiums of medicinal or useful plants where they grew, what they look like, and how to use them • Artificial systems (primary subdivisions based on habit or use)

Significance: • Partial floras • Elements of modern floras: names of plants, where they grow, what they look like (incl. illustrations)

II. The Early Days A. Theophrastus B. Herbalists 1. Herbals 2. Dioscorides (40‐90 AD)

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II. The Early Days A. Theophrastus B. Herbalists 1. Herbals 2. Dioscorides (40‐90 AD)

Important Work: • De Materia Medica

Characteristics: • Medicinal plants (ca. 600 spp) of Roman Empire • Used Theophrastus’s system

Significance: • Partial Mediterranean flora • Promoted importance of knowing plants and their names • Elements of modern floras: names of plants, where they grow, what they look like (incl. illustrations) • Used ca. 1500 yrs

De Materia Medica – Arabic translation from Spain (12-13th Century) Photo courtesy of PHGCOM.

Dill (anti-gas, indigestion soother) Cumin (parasites)

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II. The Early Days A. Theophrastus B. Herbalists 1. Herbals 2. Dioscorides (40‐90 AD) 3. Others (15th‐17th century) Important Works: • Various

Characteristics: • Utilitarian, partial floras. • Printed (printing press invented in 1440).

Significance: • Renaissance in botany ushered in by Age of Exploration • Widely and more economically distributed. • Herbals were repository for this information.

II. The Early Days A. Theophrastus B. Herbalists 1. Herbals 2. Dioscorides (40‐90 AD) 3. Others (15th‐17th century) 4. De La Cruz ( ? ‐ 1552 ‐ ?)

Martin de la Cruz • Aztec physician/botanist in Mexico (Spain)

Important Works: • Badianus Manuscript (translated from Nahuatl to Latin by Badiano)

Characteristics: • 250 medicinal spp of Aztec (Nahua) • Rediscovered in Vatican library in 1929, returned by Pope John Paul II in 1990.

Significance: • Earliest systematic record of Mexican (New World) flora, including Nahuatl names and uses.

(Calliandra anomala; stalky cornsilk flower; Tlacoxiloxochitl)

For persistent cough.

-Drink juice of the root. -Mix some of juice with honey and smear on throat.

-Fls of this plus water were said to improve eyesight and heal ulcers.

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(Mixture including Pinus sp.)

For lightening strike.

-Drink made from lvs of a pine and other species.

(thistle, pepper)

For “Black Blood” (depression).

-Grind, cook in water. Add pearl, wolf’s liver and wine. Drink. Dance.

(Urtica chichicaztli; water-nettle)

For nose bleeds (Atzitzicaztli)

-Grind juice w/ salt in urine, milk. -Pour into nose to stop flow of blood.

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III. Pre-Linnaean Taxonomists A. Caesalpino (1519‐1603) • Tuscany, Italy • ‘First Taxonomist’

Important Works: • De Plantis Libri • Herbaria (both personal and commissioned)

Characteristics: • 1500 spp of Italy and region • Principal grps based on habit • Subsidiary groupings by fr, seed, & fl chars

Significance: • Broke free of 1500 yrs of herbalism format • Natural subsidiary groups based on fr, seed, and fl chars • Created some of the earliest herbaria

III. Pre-Linnaean Taxonomists B. Ray (1628‐1705) • British

Important Works: • Synopsis Methodica Stirpium Brittannicarum • Historia Plantarum

Characteristics: • First British flora • Classification of 18,000: Herbae Imperfectae (Free‐sporing “cryptogams”) Perfectae (Seed Plants) Monocotyledons Dicotyledons Arborae Monocotyledons Dicotyledons Significance: • Complex mix of artificial and natural groupings • First recognition of cryptogams vs. seed plants, monocots vs. dicots • First to define species (variation within interbreeding populations not worthy of species distinction)

III. Pre-Linnaean Taxonomists C. Bauhin & Tournefort

• French‐Swiss • French • (1560‐1624) • (1656‐1708)

• Both contributed to establishing the genus concept

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III. Pre-Linnaean Taxonomists C. Bauhin & Tournefort

• Pinax Theatri Botanici • Shortened polynomials, flirted with binomials for species • Listed synonymy

III. Pre-Linnaean Taxonomists C. Bauhin & Tournefort

• Institutiones Rei Herbariae • Very explicit about genus concept and descriptions

IV. Linnaeus A. Carl von Linné (1707‐1778) • Sweden • aka Carolus Linnaeus • ‘Founder of Modern Taxonomy’

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IV. Linnaeus A. Carl von Linné (1707‐1778) • Sweden • aka Carolus Linnaeus • ‘Founder of Modern Taxonomy’

Historical Context: • Much old disjunct info, • New info (Age of Exploration, Microscopy, Herbaria) • Potential to synthesize & disseminate (herbaria and printing) • Need for easy, rapid way to ID, classify, and communicate

IV. Linnaeus A. Carl von Linné (1707‐1778) Important Works: Important Works: • Species Plantarum (May 1, 1753) • Systema Naturae (10th ed., 1758)

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IV. Linnaeus A. Carl von Linné (1707‐1778) Important Works: Characteristics • Species Plantarum (May 1, 1753) • Starting pt for modern bot nom • 5,490 names (he est. 10K spp) • Binomial

Before: Plantago foliis ovato‐lanceolatis pubescentibus, spica cylindrica, scapo tereti

After: Plantago media

IV. Linnaeus

Polynomial

sp. epithet (for ease of reference)

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IV. Linnaeus

Synonyms and source.

IV. Linnaeus

Nativity

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IV. Linnaeus A. Carl von Linné (1707‐1778) Important Works: Characteristics • Species Plantarum (May 1, 1753) • Starting pt for modern bot nom • 5,490 names (he est. 10K spp) • Binomial • Synonymy • Nativity • Ca. 5,500 spp, ca. 1000 gen, into ca. 100 ord, 24 classes, 1 kingdom

IV. Linnaeus

Classes based on Orders based on number (and number (and arrangement) of arrangement) of stamens pistils

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IV. Linnaeus A. Carl von Linné (1707‐1778) Anthropomorphicizing Linnaeus’s Sexual System • The flowers' leaves. . . serve as bridal beds which the Creator has so gloriously arranged, adorned with such noble bed curtains, and perfumed with so many soft scents that the bridegroom with his bride might there celebrate their nuptials with so much the greater solemnity. . . (L., 1729) • Class Pentandria, Order Monogynia was described as “5 husbands in the same marriage” • Johann Siegesbeck (1686‐1785; St. Petersburg Bot Gard): o Such “loathsome harlotry as several males with one female would not be permitted in the vegetable kingdom by the Creator!”

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In response, Linnaeus named a foul- smelling weed Siegesbeckia.

V. The French Push for Natural Systems A. Jussieu (1748‐1836) • Antoine de J. • Uncle Bernard de J. (Jardin des Plantes)

Important Works: • Genera Plantarum

Significance: • Natural classification • Invented the family

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V. The French Push for Natural Systems B. Adanson (1727‐1806) • Extensive fieldwork in Africa • Baobobs (Adansonia L.)

Important Works: • Familles des Plantes

V. The French Push for Natural Systems B. Adanson (1727‐1806) • Extensive fieldwork in Africa • Baobobs (Adansonia L.)

Important Works: • Familles des Plantes

Significance: • Natural classification • Compared families quantitatively, in table format with 65 characters

V. The French Push for Natural Systems C. Lamarck (1744‐1829) • Early advocate for evolution • Theory of evolution by inheritance of acquired traits

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V. The French Push for Natural Systems C. Lamarck (1744‐1829) • Early advocate for evolution • Theory of evolution by inheritance of acquired traits

Important Works: • Flora Francoise

Significance: • Early French flora • Dichotomous‐like key

VI. Evolutionary Taxonomy

A. Darwin Darwin (1809-1882)

Origin of Species 1859

VI. Evolutionary Taxonomy

B. Engler (1844-1930): Die Natürlichen Pflanzenfamilien

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C. Bessey (1845-1915)

“Bessey’s Dicta”

D. Cronquist (1919-1992)

An Integrated System of Classification of Flowering Plants (1981, 1988)

Evolutionary Taxonomists: Armen Tahktajan & Arthur Cronquist

NYBG ca. 1985

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Cronquist (1919-1992)

An Integrated System of Classification of Flowering Plants (1981, 1988)

VII. Phenetics (Phenetic Approaches to Classification) Recognize taxa based on similarity (overall similarity)

Characteristics of:

1. Many characters used (morphological, genetic, etc.) 2. Computational (computers & similarity metric) 3. Objective 4. Repeatable

VII. Phenetics (Phenetic Approaches to Classification) Recognize taxa based on similarity (overall similarity)

A. Adanson

B. Sneath & Sokal

Sneath & Sokal. 1973. Numerical taxonomy: The Principles and Practice of Numerical Classification.

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Example: Let’s classify the following three primate genera into two separate families.

Leontopithecus (tamarin) Hylobates (gibbon) Homo (human, man)

Leontopithecus (golden lion tamarin) Pongo Homo

Hylobates (lar gibbon) Homo

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Homo (human, man) Homo erectus

Example: Let’s classify the following three primate genera into two separate families.

1. Makes observations, collect data. Hairiness Habit Tail Pedalism Leontopithecus (tamarin) high (0) arboreal (0) yes (0) 4‐ped (0) Hylobates (gibbon) high (0) arboreal (0) no (1) 4‐ped (0) Homo (human, man) low (1) terrestrial (1) no (1) 2‐ped (1)

Example: Let’s classify the following three primate genera into two separate families.

2. Calculate similarities (states shared/Max possible) Hairiness Habit Tail Pedalism Leontopithecus (tamarin) 0 000 Hylobates (gibbon) 00 10 Homo (human, man) 11 11

Record using a pairwise similarity matrix

tamarin gibbon human tamarin 1.00 0.75 0.00 gibbon ‐‐‐‐‐ 1.00 0.25 human ‐‐‐‐‐ ‐‐‐‐‐ 1.00

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Example: Let’s classify the following three primate genera into two separate families.

3. Construct phenogram (type of dendrogram)

tamarin gibbon human tamarin 1.00 0.75 0.00 gibbon ‐‐‐‐‐ 1.00 0.25 human ‐‐‐‐‐ ‐‐‐‐‐ 1.00

1. Group most similar taxa first

Example: Let’s classify the following three primate genera into two separate families.

3. Construct phenogram (type of dendrogram)

tamarin gibbon human tamarin 1.00 0.75 0.00 gibbon ‐‐‐‐‐ 1.00 0.25 human ‐‐‐‐‐ ‐‐‐‐‐ 1.00

Human – tamarin: 0.00 Human – gibbon: 0.25 Average similarity 0.13 of human to gibbon/tamarin group

1. Group most 2. Add next most similar taxon similar taxa first (average similarity may have to calculated)

Example: Let’s classify the following three primate genera into two separate families.

3. Construct phenogram (type of dendrogram)

tamarin gibbon human tamarin 1.00 0.75 0.00 gibbon ‐‐‐‐‐ 1.00 0.25 human ‐‐‐‐‐ ‐‐‐‐‐ 1.00

1. Group most 2. Add next most similar taxon similar taxa first (average similarity may have to calculated)

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Example: Let’s classify the following three primate genera into two separate families.

4. Classify

Hylobatidae: Hylobates (gibbon) + Leontopithecus (tamarin)

Hominidae: Homo (human)

VII. Phenetic Approaches to Classification

D. Considerations

1.) Why did phenetics florish after 1950’s?

VII. Phenetic Approaches to Classification

D. Considerations

2.) Pros?

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VII. Phenetic Approaches to Classification

D. Considerations

3.) Cons?

VIII. Cladistic Approaches to Classification

A. Willi Hennig (German; 1913‐1976)

Grundzüge einer Theorie der Phylogenetischen Systematik (Hennig, 1950).

Phylogenetic Systematics (Hennig, 1966)

VIII. Cladistic Approaches to Classification

B. Hennigian Principles

1. Classification consistent w/ phylogeny i.e., named taxa above sp‐level are clades or are monophyletic

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VIII. Cladistic Approaches to Classification

B. Hennigian Principles

1. Classification consistent w/ phylogeny i.e., named taxa above sp‐level are clades or are monophyletic

a. Clade/monophyletic group = grp. incl. a common ancestor and all of its descendants.

Cladogram depicting phylogeny of 3 taxa

VIII. Cladistic Approaches to Classification

B. Hennigian Principles

1. Classification consistent w/ phylogeny i.e., named taxa above sp‐level are clades or are monophyletic

a. Clade (monophyletic group) = grp. incl. a common ancestor and all of its descendants. Good: Genus 1: Sp B and C. Genus 2: Sp A.

VIII. Cladistic Approaches to Classification

B. Hennigian Principles

1. Classification consistent w/ phylogeny i.e., named taxa above sp‐level are clades or are monophyletic

a. Clade (monophyletic group) = grp. incl. a common ancestor and all of its descendants. Good: Genus 1: Sp B and C. Genus 2: Sp A.

• B and C comprise a clade. • B & C share a more recent common ancestor than either does w/ A.

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VIII. Cladistic Approaches to Classification

B. Hennigian Principles

1. Classification consistent w/ phylogeny i.e., named taxa above sp‐level are clades or are monophyletic

a. Clade (monophyletic group) = grp. incl. a common ancestor and all of its descendants. Good: Genus 1: Sp B and C. Genus 2: Sp A. Family: Genus 1 & 2

VIII. Cladistic Approaches to Classification

B. Hennigian Principles

1. Classification consistent w/ phylogeny i.e., named taxa above sp‐level are clades or are monophyletic

a. Clade (monophyletic group) = grp. incl. a common ancestor and all of its descendants. Bad: Genus 1: Sp B and A. Genus 2: Sp C.

VIII. Cladistic Approaches to Classification

B. Hennigian Principles

1. Classification consistent w/ phylogeny i.e., named taxa above sp‐level are clades or are monophyletic

a. Clade (monophyletic group) = grp. incl. a common ancestor and all of its descendants. b. Not all clades have to be named. Ranks applied are arbitrary.

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How many clades are represented here?

D E

F G

D E

F G

4 genera 1 family I

D E

F G

2 families H

4 genera 1 order I

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VIII. Cladistic Approaches to Classification

B. Hennigian Principles

1. Classification consistent w/ phylogeny i.e., named taxa above sp‐level are clades or are monophyletic

a. Clade (monophyletic group) = grp. incl. a common ancestor and all of its descendants. b. Not all clades have to be named. Ranks applied are arbitrary. c. Phylogenetic inference usu not appropriate below species.

B & C related by phylogeny

Pops & orgs within B and C are related by tokogeny

VIII. Cladistic Approaches to Classification

B. Hennigian Principles

2. Clades marked by synapomorphy(s). Shared, derived, homologous character state(s)

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VIII. Cladistic Approaches to Classification

B. Hennigian Principles

2. Clades marked by synapomorphy(s). Shared, derived, homologous character state(s)

a. Homology vs. Non‐homology & Analogy

Homologous character states: Forelimbs & bones within

Non-homologous states: Wings

Homologous character states: Presence of stems, leaves, flowers, etc.

Non-homologous States: Arborescent growth.

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Vasculature Seeds Flowers Mosses 0 00 Ferns 100 Conifers 110 Angios 111

VIII. Cladistic Approaches to Classification

B. Hennigian Principles

2. Clades marked by synapomorphy(s). Shared, derived, homologous character state(s)

a. Homology vs. Analogy or Convergence

b. Types of Homology: Apomorphy Synapomorphy Autapomorphy Plesiomorphy Symplesiomorphy

Apomorphy Plesiomorphy

Hairiness Habit Tail Pedalism tamarin high (0) arboreal (0) yes (0) 4‐ped (0) gibbon high (0) arboreal (0) no (1) 4‐ped (0) human low (1) terrestrial (1) no (1) 2‐ped (1)

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Hairiness Habit Tail Pedalism tamarin high (0) arboreal (0) yes (0) 4‐ped (0) gibbon high (0) arboreal (0) no (1) 4‐ped (0) human low (1) terrestrial (1) no (1) 2‐ped (1)

Phenogram was based on overall similarity (plesiomorphies weighted equally to apomorphies)

Hairiness Habit Tail Pedalism tamarin high (0) arboreal (0) yes (0) 4‐ped (0) gibbon high (0) arboreal (0) no (1) 4‐ped (0) human low (1) terrestrial (1) no (1) 2‐ped (1)

Plesiomorphies are uninformative.

No evidence for this clade

Hairiness Habit Tail Pedalism tamarin high (0) arboreal (0) yes (0) 4‐ped (0) gibbon high (0) arboreal (0) no (1) 4‐ped (0) human low (1) terrestrial (1) no (1) 2‐ped (1)

Plesiomorphies are uninformative.

High hairiness plesiomorphic in mammals

No evidence for this clade

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Hairiness Habit Tail Pedalism tamarin high (0) arboreal (0) yes (0) 4‐ped (0) gibbon high (0) arboreal (0) no (1) 4‐ped (0) human low (1) terrestrial (1) no (1) 2‐ped (1)

Plesiomorphies are uninformative.

High hairiness plesiomorphic in mammals Arboreal is plesiom. in primates No evidence for this clade

Hairiness Habit Tail Pedalism tamarin high (0) arboreal (0) yes (0) 4‐ped (0) gibbon high (0) arboreal (0) no (1) 4‐ped (0) human low (1) terrestrial (1) no (1) 2‐ped (1)

Plesiomorphies are uninformative.

High hairiness plesiomorphic in mammals Arboreal is plesiom. in primates No evidence 4-ped is plesiom. in vertebrates. for this clade

Hairiness Habit Tail Pedalism tamarin high (0) arboreal (0) yes (0) 4‐ped (0) gibbon high (0) arboreal (0) no (1) 4‐ped (0) human low (1) terrestrial (1) no (1) 2‐ped (1)

0 1 Autapomorphies are uninformative. 0 1 0 1

Wrong.No evidence for this clade

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Hairiness Habit Tail Pedalism tamarin high (0) arboreal (0) yes (0) 4‐ped (0) gibbon high (0) arboreal (0) no (1) 4‐ped (0) human low (1) terrestrial (1) no (1) 2‐ped (1)

Synapomorphies are informative

0 1

Hairiness Habit Tail Pedalism tamarin high (0) arboreal (0) yes (0) 4‐ped (0) gibbon high (0) arboreal (0) no (1) 4‐ped (0) human low (1) terrestrial (1) no (1) 2‐ped (1)

Autapomorphies do not change this. 0 1 0 1 0 1

0 1

Same data, different groupings (Cladistics vs. Phenetics):

1. Both based on lots of data, repeatable, objective, computational.

2. Both use dendrograms to represent how they presume taxa are related.

3. Phenetics based on overall similarity, cladistics based on special similarity (synapomorphies).

Phenetic classification Cladistic classification Family 1: gibbons & tamarins Family1: gibbons & humans Family 2: humans alone Family 2: tamarins alone

Phenogram Cladogram

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Cronquist vs. Hennig:

1. Both evolutionary.

2. Both based on lots of data, but Cronquist’s methodology was subjective (authoritarian) and not as objective or repeatable as Hennig’s.

3. Both used tree-like diagrams to represent how they presume taxa are related. (Though Cronquist’s Besseygrams are less precise than a cladogram)

4. Cronquist willing to recognize nonmonophyletic groups. Hennig not willing.

Hamamelidae

Angiosperms are monophyletic: therefore, we could recognize as a Sp (e.g., Magnoliophyta or Magnoliopsida)

nymphaeids (water-lilies & friends) magnoliids monocots Ranunculids & other primitive eudicots rosids caryophyllids asterids

But what about classic subdivision into “monocots” &“dicots?” e.g., Cronquist (1981) Which of Cronquist’s classes is not monophyletic?

Magnoliophyta (angiosperms) Magnoliopsida (dicots) Liliopsida (monocots) nymphaeids (water-lilies & friends) magnoliids monocots Ranunculids & other primitive eudicots rosids caryophyllids asterids

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But what about classic subdivision into “monocots” &“dicots?” e.g., Cronquist (1981) Which of Cronquist’s classes is not monophyletic?

Magnoliophyta (angiosperms) Magnoliopsida (dicots) Liliopsida (monocots) nymphaeids (water-lilies & friends) magnoliids monocots Ranunculids & other primitive eudicots rosids caryophyllids asterids

More: Cronquist’s Magnoliidae included water-lilies, Magnoliids, and some primitive eudicots. Emphasis on floral Symplesiomorphies.

nymphaeids (water-lilies & friends) magnoliids monocots Ranunculids & other primitive eudicots rosids caryophyllids asterids

Dr. Hardy’s classifications of angiosperms:

Magnoliopsida (angiosperms) Various monophyletic subclasses.

Nymphaeidae

Magnoliidae

Liliidae

Ranunculidae

Proteidae

Rosidae

Caryophyllidae

Asteridae