Evolutionary Biology: A Plant

Perspective Mitchell B Cruzan

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Evolutionary Biology:
A Plant Perspective
Evolutionary Biology:
A Plant Perspective

MITCHELL B. CRUZAN

1
1
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ISBN 978–​0–​19–​088267–​9

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Printed by Sheridan Books, Inc., United States of America

Cover photo: Scalesia pedunculata forest (Galapagos daisy tree),

Los Gemelos, Isla Santa Cruz, Ecuador. Photo by the author.
 For Carol

Thank you for your tremendous support

CONTENTS

Preface   • ix
Introduction   • xi

Pa rt I | Or ig ins of Di ver si t y   • 1

1. Origin of Plant Life   • 3
2. Diversification of Land Plants   • 22

Pa rt II | Me ch a n i sms of Di ver sification   • 57

3. Origins of Variation   • 59
4. Distribution of Genetic Variation   • 99
5. Selection   • 155

Pa rt III | Pat t er ns of Di ver si t y   • 205

6. Phenotypic Variation   • 207
7. Quantifying Phenotypes   • 254
8. Intraspecific Variation   • 288

Pa rt I V | P ro ce ss of Di ver sification    • 323

9. Reproduction   • 325
10. Reproductive Isolation and Hybridization   • 385
11. Speciation and Diversification   • 427

Literature Cited   • 485

Index   • 529

vii
PREFACE

This book was initiated as a result of my experience developing a course in plant evolutionary biology. The
first day of class, I challenged the students to list unique characteristics of plants and to think about how these
features affected the patterns and modes of plant evolution. This became the theme for the course and eventu-
ally for this text. As we worked through each topic, I continued to find examples of aspects of the biology and
ecology of plants that had implications for evolutionary processes. I was motivated to begin writing reviews of
topics that were not covered in the text we were using in class. As the term progressed, I continued writing to fill
in gaps, and very soon, I had most of the outline for this book.
Writing this book has been one of the most rewarding experiences of my career. This project has given me
an excuse to dive into many topics with which I previously had limited experience. For each topic, I have been
pleased and amazed not only with the quantity of information, but also with the quality and creativity of the
work. It was perhaps most enjoyable to explore areas of the literature with which I had the least familiarity.
I have learned many new things along the way, and some of these have significantly changed my understanding
of evolutionary processes in plants. Some of the ideas that were new to me include the pathways for the origins
of eukaryotes and the first land plants, the processes affecting mutation accumulation in plant populations, and
models for the evolution of sexual reproduction.
In writing this book, I have striven to provide discussions and clear explanations of difficult concepts, as well
as new perspectives on topics that have already been thoroughly reviewed. It would not be worth writing such
a volume if I did not have something new to say. I am sure that there will be other opinions, and in fact, I hope
these ideas will promote more discussion, because otherwise we will learn nothing new. Science cannot prog-
ress if we just repeat what has already been said, and it is our duty to advance new ideas with the understanding
that some will wane while others may flourish. If I can inspire just a few to engage in new avenues of investiga-
tion, whether they agree with me or not, then the book will have been a success.
This book is not solely focused on plants. Much of the discussion of evolutionary processes is applicable to a
wide range of organisms. Moreover, the “plant perspective” only makes sense if we have something to compare
it to—​namely, animals. For each principle, I start out with a general discussion and in later sections, explore the
ways that plant biology and ecology may modify our understanding of these principles with regard to plants.
I was motivated to take this approach because too often I have noticed that concepts that have been developed
from our understanding of animals have been blindly applied to plants without considering the implications of
their biology. Virtually everything about plants is different—​they are truly the most alien group of organisms
we may ever have the privilege of encountering—​and these differences should define the core of our under-
standing of their past, present, and future evolution.

ix
x Preface

It is probably not possible, and certainly not desirable, to put everything about plant evolution into a single
book. I have attempted to be comprehensive without subjecting the reader to oppressive levels of detail. Of
course, the topics selected necessarily represent my perspective, and I am sure there are many who will point
out important topics that I overlooked. Similarly, I was limited on the number of excellent papers I could cite.
My goal was to provide relevant examples from the literature, both historical and contemporary, so that anyone
interested would have a window into the topic for further exploration. I purposefully tried to avoid over-​citation
of the work coming out of my own lab, but sometimes these papers provided the best examples. Out of the many
thousands of papers I reviewed during the preparation of this work, I was only able to include a small fraction,
and I am sure that many readers could provide a list of very important papers that could have been cited.
Completion of this book would not have been possible were it not for the contributions of many people, and
most of all, the students with whom I have had the privilege of working. The best learning comes about through
discussion, and I have benefited from discussions with all of my colleagues, and especially my mentors, fellow
graduate students, postdocs, and every student who has ever spent time in my lab. Many others have had a di-
rect impact on the content of this text. In particular, I deeply appreciate the contributions of Michelle Nicole
Williamson, who generously provided many fine drawings and beautiful illustrations of paleo plants. Those who
have provided edits, comments, and participated in discussions that have improved the presentation include
Tina Arredondo, Tanya Cheeke, Carol Cruzan, Nic Diaz, Monica Grasty, Lizzi Hendrickson, Brendan Kohrn,
Caitlin Lee, Caitlin Maraist, Gina Marchini, Trieste Musial, Jessica Persinger, Avery Pheil, Alisa Ramakrishnan,
Jaime Schwoch, Pam Thompson, Michelle Williamson, and Rachael Workman, as well as many students from
my courses on evolutionary biology and plant reproduction.
INTRODUCTION

Early in the history of life on Earth, the rise of bacteria capable of photosynthesis established a novel mode of
energy capture. The new photosynthetic lineages embarked on an evolutionary trajectory that differed from
the dominating consumers—​one that was governed by the unique opportunities afforded by an autotrophic
mode of resource acquisition. This simple difference had far-​ranging implications for the future evolution of
photosynthetic microorganisms and eventually, for land plants. Many characteristics of plants stem directly
from the ability to obtain energy directly from the sun. The goal of this book is to improve our understanding
of the origin and evolutionary consequences of the unique traits that characterize the vast and diverse group of
organisms generally referred to as plants.
Developing an understanding of the consequences of evolutionary processes for proliferation and diver-
sification of plants requires an appreciation of their unique biological features. The processes of mutation,
selection, genetic drift, gene flow, and diversification remain the same for plants and animals, but specific char-
acteristics of plants impact the way evolution is implemented. Unique traits of plants affect everything from the
elevated potential for the accumulation of mutations during vegetative growth, to the distribution of genetic
variation within and among populations, and ultimately to rates and patterns of diversification. My primary goal
is to develop a deeper understanding of the origins of these features, and an appreciation of their implications
for evolutionary processes.
Evolutionary biology is an exceedingly complicated and conceptually rich science. This field of study has
tremendous depth, and I believe that all evolutionary biologists who are working in this field—​no matter how
experienced or knowledgeable—​are constantly learning more of the nuances and developing a deeper appre-
ciation of the complexity of plant evolutionary processes. This point has become even more relevant in recent
decades as improved technologies allow us to explore genomes with increasingly greater resolution and as we
strive to make appropriate evolutionary interpretations of the patterns we find.
A thorough appreciation of concepts in evolutionary biology requires familiarity with a range of fields.
Students will find the discussions in this book most rewarding if they have some fluency in ecology, population
genetics, physiology, cellular and molecular biology, and basic principles of evolutionary biology. This book is
specifically designed for advanced undergraduates and graduate students and as a reference for researchers in
plant evolutionary biology. My goal is to provide a balanced and thorough coverage of traditional and contem-
porary topics in evolutionary biology with reference to aspects of plant biology that are relevant to evolutionary
processes. Evolutionary principles are discussed using classical and contemporary examples from the primary
literature.

xi
xii Introduction

This book is organized by major topics in plant evolutionary biology, starting with the origin and diver-
sity of plants, origins of genetic variation, principles of population and quantitative genetics, and processes of
evolutionary diversification. To the degree that the information enhances our understanding of evolutionary
processes, I have included discussion of more phytocentric topics, such as plant morphology and development
and plant reproductive biology. Each chapter begins with a general introduction to increase accessibility to a
broader audience. Basic principles are presented first, followed by expanded discussions that integrate more
complicated aspects of plant biology. Several of the chapters include sections on the history and development
of evolutionary thought to provide a contextual backdrop for discussion of contemporary issues in plant evolu-
tionary biology. Many of the examples used to illustrate concepts are taken from applied biology (agricultural
and conservation biology), and are informed by other fields such as plant developmental biology, anatomy,
physiology, ecology, and molecular genetics and genomics.
I have provided readers with a list of questions for contemplation and discussion at the end of each chapter
that should be particularly useful for students and instructors. The goal of these questions is to provide students
with a tool for reviewing the information presented in each chapter and to provide topics for class discussion.
Understanding of the topics addressed can be further developed by reading additional papers from the primary
literature. Instructors using this text may consider incorporating regular discussions that utilize papers from the
literature to expand coverage of each topic and provide examples of more recent advances in the field. The text
is designed to provide an introduction to each topic, and it is up to the reader to determine which topics they
wish to explore further.
Fig. 1.1.

Fig. 1.5.
 Flowers
Dicots
Seeds Monocots
-Vasculature Basal Angiosperms

Vascular Plants
-Sporophyte is Gymnosperms
Dominant
Ferns
-Embryo
-Vertical Growth Lycophytes
-Gametophyte is
Dominant Hornworts

Bryophytes
Apical Growth Mosses

Liverworts

Charophyceae

Chlorophyll a + b
Green Algae

Chlorophyceae

Red Algae

Other Protista

Origin of Land Plants

Fig. 2.1.
Fig. 2.11.

Sporophyte Meiosis
(a) Sperm Megaspores
Microspores 2N
Meiosis
2N 1N
Microgametophyte
1N
Megagametophyte

(b)

Rhizoids

(c)

Archegonia

(d) Nucellus
Sporophyte
Pollen Embryo
Pollen Tube
Necellus
Megagametophyte Seed

Fig. 2.15.
 Petal

Pistil

Stamen

Sepal

Peduncle

Evolutionary Origin of the Flower

Fig. 2.18.

Fig. 2.22.

Fig. 2.23.
 Selective Sweeps Background Selection
Hard Sweep Soft Sweep High Density Low Density

Selection
Before
Selection
After
Polymorphism

Fig. 3.4.

Recombination
High Low
Selection
Before
Selection
After
Polymorphism

Fig. 3.5.

(a) (b)

Central zone L1 Cell layer

Peripheral zone L2 Cell layer
Rib zone L3 Cell layers

Fig. 3.13.
Fig. 3.15.

Fig. 3.16.
 Neutral Neutral
Beneficial Deleterious Beneficial Deleterious

Benefi
s
tion

Not A
c ial M
uta

ccumu
Elim rious M

u t at io
ed

lated
in at

ns
ete
Del
Mutation
Generations

Accumulation Mutation
with Intra- Accumulation
without Intra-
Generation
Initial Initial Generation
Selection
Population Population Selection

0.0 0.0
Proportion of Mutations Proportion of Mutations

Fig. 3.20.

Fig. 4.5.
Fig. 4.11.

Inbreeding (f )

1 0.0

2 0.0
Generations

3 0.125

4 0.125

5 0.141

Fig. 4.14.
Fig. 5.3.

Selection

Strong

Migration Weak Drift

Larg
Low e
e Pop
Hig
h
al Rat ulat
ion Sma
pers Size ll
Dis
The Allele
Frequency Fulcrum

Fig. 5.17.

0.9 0.6

0.8 0.5

0.4
Multiplicative fitness

0.7
Gene diversity (Hs)

0.3
0.6
0.2
0.5
0.1

0.4 0.0

0.3 –0.1
E4 M1 S2 C6 C1 C10
Population

Fig. 5.19.
 1.0

0.9

0.8

0.7

0.6

Un-
colonized

Fig. 5.21.

Differentiation Selection and Drift

within and
between
Habitats

FST HIGH

FST MID
Genetic Drift

FST FST FST FST

LOW LOW LOW LOW

FST MID

FST HIGH

Fig. 5.27.
 Adaptation along an Environmental Gradient
FST LOW FST LOW FST LOW FST LOW

FST MID FST MID FST MID FST MID

Adaptation with Isolation by Distance

FST LOW Dispersal

FST LOW IBDist

Allopatric
Di

al
ers
spe

isp
rsa

D
l

FSTLOW FSTMID FSTLOW

FST LOW FST HIGH FST LOW

Parapatric

Fig. 5.28.

Seconds Minutes Hours Days Years Centuries Millennia

Genetic Lineages

Epigenetic Populations
Selection
Generations Responses
Gene Expression
Module Developmental

Acclimation Phenotypic
Plasticity
Physiological Adjustments

0 1 2 3 4 5 6 7 8 9 10 11 12
E)
Time Scale (Seconds 10

Fig. 6.1.
Fig. 6.3.

Fig. 6.4.
 Ground Vascular
Tissue Bundle Phloem Xylem Epidermis

Fig. 6.5.

cuticle
uppder epidermis
chloroplast
vacuole
nucleus
palisade cell wall
cytoplasm
mesophyll

spongy

lower epidermis
cuticle

guard cell
stoma

Fig. 6.6.

Endodermis Parenchyma

Pericycle Phloem

Vessel
Element

Tracheids
Amyloplasts

Fig. 6.7.
 Secondary Phloem

Secondary Phloem

Ray
Pith

Vascular Cambium

Cork Cambium

Fig. 6.8.

Fig. 6.9.
Phenotypic Trait Two

Phenotypic Trait One

Fig. 6.12.
Fig. 6.13.
Reproductive Value (log scale)

Age (log scale)

Annual Perennial Large Tree

Fig. 6.14.
Fig. 6.18.

A
13

12 High exsertion
selection lines
11

10

Low exsertion
9 selection lines
Long filament length (mm)

8
7 8 9 10 11

B
13

12

11

10

8
7 8 9 10 11
Corolla Tube Length (mm)

Rep1 High Rep1 Control Rep1 Low

Rep2 High Rep2 Control Rep2 Low

Fig. 6.21.
 Fig. 6.23.

Fig. 6.24.
Trait Trait Trait
Environment Environment Environment
Fitness Fitness Fitness Fitness Trait
1

Time
Plasticity
Environment
2
Fig. 6.25.

Fig. 6.26.

6.0

Dry c a
4.5
Length/Width

3.0 Wet
b

1.5

1.0
Leaf Length

Fig. 6.28.
 Glabrous Moderate High

sity
Den
me dien
t
Tr icho Gra
on
ressi

Phenotypic Trait
Gene Expression
p
e Ex
Gen

Gene A Gene B Gene C

Environmental Gradient

Fig. 6.29.

6.0

AA′
1
Locus
4.0
Frequency

2
Loci
AA A′A′
1 2 4 2 1
2.0 Locus Loci Loci Loci Locus
4 4
Loci Loci

4 4
Loci Loci
0.0
Phenotypic Trait Value

Fig. 7.2.

2
Pollen load residuals (W)

–2

–4
H V C
–6

–8
0 0.5 1 1.5 2 2.5 3
Floral area difference (cm2)

Fig. 7.11.
 QTL Genetic Map

AA X aa

F1 Hybrids Aa X Aa

AA AA Aa Aa Aa Aa aa aa
F2 Hybrids

AA Aa aa
Red Purple Blue

Fig. 7.14.

A. m. pseudomajus A. m. striatum
A

WT

rostr

sulf inc

B 3 2 1

Fig. 7.16.
Fig. 7.17.
 a10 f 10 m10

a9 f9 m9

a8 f8 k8 I8 m8

a7 f7 k7 I7 m7

a6 f6 k6 m6

a5 d5 k5 m5

a4 d4 t4 m4

a3 t3 m3
Time

a2 s2 m2

a1 m1

Fig. 8.1.

Laurentide Ice Sheet

Mixed Hardwoods
Shoreline 20,000 bp

Fig. 8.2.
 (a)
+

Abundance/Growth Rate
0

–
(b) +
Abundance

Gene Flow

(c)
+
Abundance

–
(d)
+
Limited Gene Flow

Gene Flow
Abundance

Gene Flow

0
Geographic Distance

Fig. 8.4.

Pollen Dispersal Pollen Dispersal

cpDNA Nuclear cpDNA Nuclear cpDNA

Genetic Marker Migration

Fig. 8.5.
 Contemporary Haplotypes
HAGE F DA BC E

Dispersal
Time

Dispersal

Fig. 8.11.

Geographic Distance
Time

Gene
Flow
Gene
Flow

Fig. 8.12.
 Anther
Stigma
Filament Stamen
Pistil
Style
Petal
Stamen
Ovary Sepal

Cosexual Monoecious Pistil

Dioecious Gynodioecious Androdioecious

Fig. 9.1.

Fig. 9.2.

Asexual/Hermaphrodite Dioecious

Fig. 9.4.
Fig. 9.8.

Archesporium Megagametophyte Ovule

Pollen
Primordial Synergids Micropyle Tube
Integuments
Nucellus Integuments
Egg
Cell Funiculus

Meiosis Fertilization Sperm

Nuclei
Polar
Nucellus
Nuclei Embryo
Megasporocyte Antipodals Sac
Chalaza

Fig. 9.9.

Sporophytic Apomixis Cotyledon

Archesporium
Mitosis Endosperm
5
Unreduced Nucellus
Somatic Embryo Seed Coat
2 Apospory
Hypocotyl
Mitosis Autonomous
Apomixis Mitosis
Mitosis
Generative 3
Apospory 1
Pseudogamy

Gametophytic 4
Apomixis
Unreduced Megagametophyte

Fig. 9.10.
Fig. 9.11.

Fig. 9.12.
 Trimeniaceae Austrobaileyales
Luzuriagaceae
Liliaceae
Iridaceae
Amaryllidaceae
Agavaceae

Monocots
Tecophilaeaceae
Orchidaceae
Poaceae
Bromeliaceae
Heliconiaceae
Zingiberaceae
Pontederiaceae
Commelinaceae
Cyclanthaceae
Araceae
Annonaceae
Winteraceae Magnoliids
Saururaceae
Papaveraceae
Fumariaceae Ranunculales
Berberidaceae
Ranunculaceae
Proteaceae Proteales
Rosaceae
Phamnaceae
Fagaceae
Betulaceae
Fabaceae
Celastraceae
Oxalidaceae
Connaraceae
Erythroxylaceae
Passifloraceae
Turneraceae
Chrysobalanaceae
Linaceae
Clusiaceae
Malpighiaceae
Euphorbiaceae
Cochlospermaceae

Rosids
Malvaceae
Dipterocarpaceae
Cistaceae
Rutaceae
Meliaceae
Sapindaceae
Anacardiaceae
Brassicaceae
Resedaceae
Tropaceolaceae
Myrtaceae
Melastomataceae
Onagraceae
Lythraceae
Geraniaceae
Staphyleaceae
Hamamelidaceae
Saxifragaceae
Crassulaceae
Paeoniaceae
Caryophyllaceae
Amaranthaceae
Nyctaginaceae
Cactaceae
Plumbaginaceae
Polygonaceae
Droseraceae
Olacaceae
Santalaceae
Cornaceae
Ericaceae
Theaceae
Lecythidaceae
Sapotaceae
Myrsinaceae
Primulaceae
Polemoniaceae
Fouquieriaceae
Asterids

Solanaceae
Convolvulaceae
Boraginaceae
Genseriaceae
Plantaginaceae
Scrophulariaceae
Lentibulariaceae
Bignoniaceae
Acanthaceae
Orobanchaceae
Oleaceae
Rubiaceae
Sl unknown Apocynaceae
Gametophytic Sl Loganiaceae
Gentianaceae
Sporophytic Sl Campanulaceae
Menyanthaceae
Heteromorphic Sl Goodeniaceae
Asteraceae
Argophyllaceae
Caprifoliaceae

Fig. 9.14.
 Selection to
produce only
male or
Monoecious female flowers

Loss of female
function in
cosexual
flowers

Gynodioecious

Selection for
Cosexual Dioecious
increased
specialization
towards male
and female
function
Distyly

Fig. 9.27.

Fig. 9.28.
Fig. 9.29.

Speciation
Reproductive

Fusion
Isolation

Diversification
Time

Geographic
Variation

Fig. 10.1.

Mechanisms of
Reproductive
Isolation
Phenological
Ethological
Morphological
Post-Zygotic
Pre-Mating
Fertility:
Pollen
Ovule Post-Mating/
Post-Zygotic
Viability:
Survival Pollen
Seeding Germination
Germination Pollen Tube
Attrition
Seed Pollen
Abortion Competition

Fig. 10.4.
Fig. 10.6.

Fig. 10.7.
 aabb

aaBB AAbb

Fitness
F1

AaBb
aa Recombinants bb

aA bB

AA BB

Fig. 10.10.

Antioch

San Lucas Idria

Tehachapi

Bear
Valley

Cholame
Oceana

Casmalia

Fig. 10.12.

Phlox
cuspidata

Texas

Phlox
drummondii

Phlox
drummondii

Fig. 10.13.
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 Table of the Twelve Houses.
Astrologers draw their table of the TWELVE HOUSES into a triple
quadrangle prepared for the purpose, of which there are four
principal angles, two of them falling equally upon the horizon, and
the other two upon the meridian, which angles are sudivided into 12
triangles for the 12 houses, in which they place the 12 signs of the
Zodiac, to each of which is attributed a particular quality,—viz.

1.— Aries, denoted by the sign ♈︎, is, in their extravagant opinion,
a masculine, diurnal, cardinal, equinoctial, easterly sign, hot
and dry,—the day house of Mars.
2.— Taurus, ♉︎, is a feminine, nocturnal, melancholy, bestial,
furious sign—cold and dry.
3.— Gemini, ♊︎, is a masculine sign, hot and moist, diurnal, aërial,
human, double-bodied, &c.
4.— Cancer, ♋︎, is a feminine, nocturnal, phlegmatic sign, by
nature cold and moist, the only house of Luna.
5.— Leo, ♌︎, is a sign, masculine, diurnal, bestial, choleric and
barren; a commanding, kingly sign—hot and dry, the only
house of the sun.
6.— Virgo, ♍︎, is a feminine, nocturnal, melancholy, and barren
sign.
7.— Libra, ♎︎, is a sign masculine, cardinal, equinoctial, diurnal,
sanguine and human, hot and moist.
8.— Scorpio, ♏︎, is a feminine, nocturnal, cold and phlegmatic
northern sign.
9.— Sagittarius, ♐︎, is a sign masculine, choleric, and diurnal, by
nature hot and dry.
 10. Capricorn, ♑︎, is a feminine, nocturnal, melancholy, solstitial,
— moveable, cardinal, and southern sign.
11.— Aquarius, ♒︎, is a masculine, diurnal, fixed, sanguine, and
human sign.
12. Pisces, ♓︎, is a feminine, nocturnal, phlegmatic, northerly
— double-bodied sign, the last of the twelve.
Having thus housed their signs and directed them in their
operations, they afterwards come to enquire of their tenants, what
planet and fixed stars they have for LODGERS, at the moment of the
nativity of such person; from whence they draw conclusions with
regard to the future incident of that person’s life. For if at the time of
that person’s nativity they find Mercury in 27° 52 min. of Aquarius,
and in the sextile aspect of the horoscope, they pretend to foretel that
that infant will be a person of great sagacity, genius, and
understanding; and therefore capable of learning the most sublime
sciences.
Astrologers have also imagined, for the same ridiculous purpose,
to be in the same houses different positions of the signs and planets,
and from their different aspects, opposition and conjunction, and
according to the rules and axioms they have prescribed to themselves
and invented, have the sacrilegious presumption to judge, in dernier
resort, of the fate of mankind, though their pretended art or science
is quite barren either of proofs or demonstrations.
 Signs to the Houses of the Planets.
The planets have allowed themselves each, except Sol and Luna,
two signs for their houses; to Saturn, Capricorn and Aquarius; to
Jupiter, Sagittarius and Pisces; to Mars, Aries and Scorpio; to Sol,
Leo; to Venus, Taurus and Libra; to Mercury, Gemini and Virgo;
and to Luna, Cancer.
 Angles or Aspects of the Planets.
By their continual mutations among the twelve signs, the planets
make several angles or aspects; the most remarkable of which are the
five following, viz.—
☌ Conjunction.—Δ Trine.—☐ Quadrate.—⚹ Sextile.—☍
Opposition.
A Conjunction is when two planets are in one and the same
degree and minute of a sign; and this, according to Astrological cant,
either good or bad, as the planets are either friends or enemies.
A Trine is when two planets are four signs, or 120 degrees distant,
as Mars in twelve degrees of Aries, and Sol in twelve degrees of
Leo. Here Sol and Mars are said to be in Trine Aspect. And this is
an aspect of perfect love and friendship.
A Quadrate Aspect is when two planets are three signs, or 90
degrees distant, as Mars in 10 degrees, and Venus in 10 degrees of
Leo. This particular aspect is of imperfect enmity, and Astrologers
say, that persons thereby signified, may have jars at sometime, but of
such a nature as may be perfectly reconciled.
A Sextile Aspect, is when two planets are two signs, or 60 degrees
distant, as Jupiter in 15 degrees of Aries; and Saturn in 15 degrees
of Gemini; here Jupiter is in a sextile aspect to Saturn. This is an
aspect of friendship.
An Opposition is, when two planets are diametrically opposite,
which happens when they are 6 signs, or 180 degrees (which is one
half of the circle) asunder; and this is an aspect of perfect hatred.
A Partile Aspect, is when two planets are in a perfect aspect to
the very same degree and minute.
Dexter Aspects, are those which are contrary to the succession of
signs; as a planet, for instance, in Aries, casts its sextile dexter to
Aquarius.
Sinister Aspect, is with the succession of signs, as a planet in
Aries, for example, casts its sextile sinister in Gemini.
In addition to these, Astrologers play a number of other diverting
tricks; hence we read of the Application—Prohibition—
Translation—Refrenation—Combustion—Exception—
Retrogradation, &c. of planets.
 The Application of Planets.
Application of the planets is performed by Astrologers in three
different ways.
1. When a light planet, direct and swift in its motion, applies to a
planet more ponderous and slow in motion; as Mercury in 8° of
Aries, and Jupiter in 12° of Gemini, and both direct; here Mercury
applies to a sextile of Jupiter, by direct application.
2. When they are both retrograde, as Mercury in 20° of Aries, and
Jupiter in 15° of Gemini; here Mercury, the lighter planet, applies to
the sextile aspect of Jupiter; and this is by retrogradation.
3. When one of the planets is direct, and the other retrograde; for
example, if Mercury were retrograde in 18° of Aries, and Jupiter
direct in 14° of Gemini; in this case Mercury applies to a sextile of
Jupiter, by a retrograde motion.
 Prohibition,
is when two planets are applying either by body or aspect; and
before they come to their partile aspect, another planet meets with
the aspect of the former and prohibits it.
 Separation,
is when two planets have been lately in conjunction, or aspect, and
are separated from it.
 Translation of Light and Virtue,
is when a lighter planet separates from the body or aspect of a
heavier one, and immediately applies to another superior planet, and
so translates the light and virtue of the first planet to that which it
applies to.
 Refrenation,
is when a planet is applied to the body or aspect of another; and,
before it comes to it, falls retrograde, and so refrains by its
retrograde motion.
 Combustion.
A planet is said to be combust of Sol, when it is within 8° 30″ of
his body, either before or after his conjunction: but Astrologers
complain, that a planet is more afflicted when it is applying to the
body of Sol, than when it is separating from combustion.
 Reception,
is when two planets are in each other’s dignities, and it may either
be by house, exultation, triplicity, or term.
 Retrogradation,
is when a planet moves backward from 20° to 9°, 8°, 7°, and so out
of Taurus into Aries.
 Frustration,
is when a swift planet applies to the body or aspect of a superior
planet; and before it comes to it, the superior planet meets with the
body or aspect of some other planet.
 The Dragon’s Head and Tail.
To the seven planets, viz. Saturn, Jupiter, Mars, Sol, Venus,
Mercury, and Luna; Astrologers add, two certain nodes or points,
called the Dragon’s head, distinguished by this sign ☋, and the
Dragon’s tail by ☊. In those two extremities of the beast, our students
in Astrology place such virtues, that they can draw from thence
wealth, honour, preferments, &c. enough to flatter the avarice,
ambition, vanity, &c. of the fools who follow them. Sensible,
however, that the admirers of this art support their principles and
defend their doctrines by examples founded on their own experience
and on the authority of history; there is no necessity for us here to
expose the weakness and futility of their arguments. Tully’s proof
will suffice; who, amidst the darkest clouds of superstition and
ignorance, and in the very heyday of paganism and idolatry, and
whilst religion itself seemed to countenance Astrology, inveighs
severely against it in Lib. 2, de devinat. “Quam multa ego Pompeis,
quam multa Crasso, quam multa huic ipsi Cæsari a Chaldæis dicta
memini, neminem eorum nisi senectute, nisi domi, nisi cum clantate
esse moriturum? ut mihi per Mirum videatur quem quam extare,
qui etiam nunc credastis, quorum predicta quotidie videat re et
eventis refelli[9].”
 Climacteric.
Astrologers have used their best artifices, and employed all the
rules of their art, to render those years of our age, which they call
climacterics, dangerous and formidable.
Climacterick from the Greek, κλιμακτης, which means by a scale or
ladder, is a critical year, or a period in a man’s age, wherein,
according to Astrological juggling, there is some notable alteration to
arise in the body; and a person stands in great danger of death. The
first climacterick, say they, is the seventh year of a man’s life; the rest
are multiples of the first, as 21, 49, 56, 63, and 84; which two last are
called the grand climactericks, and the danger more certain.
Marc Ficinus accounts for the foundation of this opinion: he tells
us there is a year assigned for each planet to rule over the body of a
man, each in his turn; now Saturn being the most maleficent
(malignant) planet of all, every seventh year, which falls to its lot,
becomes very dangerous; especially those of 63 and 84, when the
person is already advanced in years. According to this doctrine, some
hold every seventh year an established climacteric; but others only
allow the title to those produced by the multiplication of the
climacterical space by an odd number, 3, 5, 7, 9, &c. Others observe
every ninth year as a climacterick.
There is a work extant, though rather scarce, by Hevelius, under
the title of Annus Climactericus, wherein he describes the loss he
sustained by his observatory, &c. being burnt; which, it would
appear, happened in his grand climacterick. Suetonius says, that
Augustus congratulated his nephew upon his having passed his first
grand climacterick, of which he was very apprehensive.
Some pretend that the climacterick years are fatal to political
bodies, which perhaps may be granted, when they are proved to be so
to natural ones; for it must be obvious that the reason of such danger
can by no means be discovered, nor what relation it can have with
any of the numbers above-mentioned. Though this opinion has a
great deal of antiquity on its side; Aulus Gellius says, it was borrowed
from the Chaldeans, who, possibly, might receive it from Pythagoras,
whose philosophy turned much on numbers, and who imagined an
extraordinary virtue in the number 7.
The principal authors on the subject of climactericks, are Plato,
Cicero, Macrobius, Aulus Gellius, among the ancients; Argol,
Magirus, and Salmatius, among the moderns. St. Augustine, St.
Ambrose, Beda, and Bœtius, all countenance the opinion.
 Lucky and Unlucky Days.
Astrologers have also brought under their inspection and controul
the days of the year, which they have presumed to divide into lucky
and unlucky days; calling even the sacred scriptures, and the
common belief of Christians, in former ages, to their assistance for
this purpose. They pretend that the 14th day of the first month was a
blessed day among the Israelites, authorised therein, as they pretend,
by the several following passages out of Exodus, c. xii. v. 18, 40, 41,
42, 51. Leviticus, c. xxiii. v. 5. Numbers, c. xxviii. v. 16. “Four
hundred and thirty years being expired of their dwelling in Egypt,
even in the self same day departed they thence.”
With regard to evil days and times, Astrologers refer to Amos, c. 5,
v. 13, and c. vi. v. 3. Ecclesiasticus, c. ix. v. 12. Psalm, xxxvii. v. 19.
Obadiah, c. xii. Jeremiah, c. xlvi. v. 21, and to Job cursing his birth
day, chap. iii. v. 1 to 11. In confirmation of which they also quote a
calendar, extracted out of several ancient Roman catholic prayer
books, written on vellum, before printing was invented, in which
were inserted the unfortunate days of each month, as in the following
verses;—
January.—Prima dies mensis, et septima truncat ensis.
February.—Quarta subit mortem, prosternit tertia fortem.
March.—Primus mandentem, disrumpit quarta bibentem.
April.—Denus et undenus est mortis vulnere plenus.
May.—Tertius occidit, et Septimus ora relidit.
June.—Denus Pallescit, quindenus fædera nescit.
July.—Ter denus mactat, Julii denus labefactat.
August.—Prima necat fortem, perditque secunda cohortem.
September.—Tertia Septembris, et denus fert mala membris.
October.—Tertius et denus, est sicut mors alienus.
November.—Scorpius est quintus, et tertius est vita tinctus.
December.—Septimus exanguis, virosus denus ut Anguis.

This poetry is a specimen of the rusticity and ignorance at least of

the times; and is a convincing proof that Christianity had yet a very
strong tincture of the Pagan superstitions attached to it, and which
all the purity of the gospel itself, to this very day, has not been able
entirely to obliterate.
That the notion of lucky and unlucky days owes its origin to
paganism, may be proved from Roman history, where it is mentioned
that that very day four years, the civil wars were begun by Pompey
the father; Cæsar made an end of them with his son, Cneius
Pompeius being then slain; and that the Romans accounted the 13th
of February an unlucky day, because on that day they were
overthrown by the Gauls at Allia; and the Fabii attacking the city of
the Recii, were all slain with the exception of one man: from the
calendar of Ovid’s “Fastorum,” Aprilis erat mensis Græcis
auspicatissimus; and from Horace, lib. 2, ode 13, cursing the tree
that had nearly fallen upon it; ille nefasto posuit die.
The number of remarkable events that happened on some
particular days have been the principal means of confirming both
Pagans and Christians in their opinion on this subject. For example,
Alexander the Great, who was born on the 6th of April, conquered
Darius and died on the same day. The Emperor Bassianus Caracalla
was born and died on a sixth day of April. Augustus was adopted on
the 19th of August, began his Consulate, conquered the Triumviri,
and died the same day.
The Christians have observed that the 24th of February was four
times fortunate to Charles the Fifth. That Wednesday was a fortunate
day to Pope Sixtus V. for on a Wednesday he was born, on that day
made a Monk, on the same day made a General of his order, on that
day created a Cardinal, on that day elected Pope, and also on that day
inaugurated. That Thursday was a fatal day to Henry VIII. King of
England, and his posterity, for he died on a Thursday; King Edward
VI. on a Thursday; Queen Mary on a Thursday; and Queen Elizabeth
on a Thursday. The French have observed that the feast of Pentecost
had been lucky to Henry III. King of France, for on that day he was
born, on that day elected king of Poland, and on that day he
succeeded his brother Charles IX. on the throne of France.
 Genethliaci.
(From γενεθλη, origin, generation, nativity.)
These, so called in Astrology, are persons who erect Horoscopes;
or pretend what shall befal a man, by means of the stars which
presided at his nativity[10]. The ancients called them Chaldæi, and by
the general name mathematici: accordingly the several civil and
canon laws, which we find made against the mathematicians, only
respect the Genethliaci, or Astrologers; who were expelled Rome by a
formal decree of the senate, and yet found so much protection from
the credulity of the people, that they remained unmolested. Hence an
ancient author speaks of them as hominum genus, quod in civitate
nostra sempe et vetabitur, et retinebitur.
Genethliacum, (Genethliac poem,)

Is a composition in verse, on the birth of some prince, or other

illustrious person; in which the poet promises him great honours,
advantages, successes, victories, &c. by a kind of prophecy or
prediction. Such, for instance, is the eclogue of Virgil to Pollio,
beginning
Sicelides Musæ, paulo majora Canamus.

There are also Genethliac speeches or orations, made to celebrate a

person’s birth day.