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Methods in
Molecular Biology 2222
Pascale Besse Editor
Molecular
Plant Taxonomy
Methods and Protocols
Second Edition
METHODS IN MOLECULAR BIOLOGY
Series Editor
John M. Walker
School of Life and Medical Sciences
University of Hertfordshire
Hatfield, Hertfordshire, UK
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For over 35 years, biological scientists have come to rely on the research protocols and
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Molecular Plant Taxonomy
Methods and Protocols
Second Edition
Edited by
Pascale Besse
UMR PVBMT, Universite de la Reunion, St Pierre, Réunion, France
Editor
Pascale Besse
UMR PVBMT
Universite de la Reunion
St Pierre, Réunion, France
ISSN 1064-3745 ISSN 1940-6029 (electronic)

Methods in Molecular Biology
ISBN 978-1-0716-0996-5 ISBN 978-1-0716-0997-2 (eBook)
https://doi.org/10.1007/978-1-0716-0997-2
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Preface
Plant taxonomy is an ancient discipline facing new challenges with the availability of a vast
array of modern molecular technologies. The literature reviews and protocols that appear as
chapters in this book were selected to provide conceptual as well as technical guidelines to
plant taxonomists and geneticists. This second edition of Molecular Plant Taxonomy
appeared necessary to take into account the increasing use of next-generation sequencing
(NGS) technologies for many applications in plant taxonomy.
The introductive Chapter 1 allows the reader to travel through the historical aspects of
plant taxonomy with a focus on the strengths, limitations, and the future of molecular
techniques. Chapter 2 then proposes guidelines to choose the best sequence and molecular
technique to be used according to the taxonomic question addressed. A temporal landscape
of the most commonly used techniques is also provided. Both chapters are prerequisite
readings to understand the concepts underlying the “plant taxonomy” discipline and to fully
appreciate the strengths and limits of each molecular technique presented in this book.
DNA extraction protocols specifically focused on recalcitrant plant species (Chapter 3)
and herbarium specimens (Chapter 4) are proposed. The latter will ensure the development
of an “integrative” taxonomic approach by allowing the use of ancient DNA references from
herbarium specimens together with present-date accessions in DNA analyses.
Next-generation sequencing technologies have opened a new era for molecular plant
taxonomy. This revised edition provides literature review and wet-lab protocols and/or
decision flowcharts covering whole chloroplast (Chapter 5) and mitochondrial (Chapter 6)
genome sequencing, now more and more replacing the Sanger sequencing of specific
regions described in the earlier version of this book. We also chose to present an updated
protocol for microsatellite markers isolation based on Illumina sequencing (Chapter 11) to
complement classical enriched library construction described in the first version. This
NGS-based method is powerful enough to reveal numerous microsatellite loci, which are
markers of choice for molecular plant taxonomy. New methods to discover single nucleotide
polymorphism (SNP) markers from sequenced pangenomes (Chapter 9) are also described,
together with the simple and powerful genotyping-by-sequencing (GBS) method to
develop SNP markers without any need for whole genome sequencing and assembly,
perfectly suited for many plant species (Chapter 10).
This book still provides detailed literature reviews and detailed wet-lab protocols for
many multilocus PCR-based profiling methods that have been shown to be very efficient in
resolving many molecular plant taxonomy issues: amplified fragment length polymorphism
(AFLP, Chapter 12), random amplified polymorphic DNA (RAPD, Chapter 13) and their
multiple derived techniques, inter-simple sequence repeats (ISSR, Chapter 14), and the use
of a range of methods tagging retrotransposable elements (Chapter 15). It also provides a
protocol for Sanger sequencing and data analysis of the widely used internal transcribed
spacer (ITS) nuclear region in plants (Chapter 7), and the usefulness and power of this ITS
region together with that of various chloroplast regions as a “DNA barcoding” tool is
reviewed and assessed (Chapter 8): it is now clear that using these simple “barcode tools” as
defined by the CBOL (consortium for the barcoding of life) for resolving plant taxonomy
will not be sufficient, particularly in some plant groups. We rather highly recommend that
molecular approaches are used within an “integrative taxonomy” framework, combining a
v
vi Preface
range of nucleic acid and cytogenetic data together with other crucial information (taxon-
omy, morphology, anatomy, ecology, reproductive biology, biogeography, paleobotany,
etc.), which will help not only to best circumvent species delimitation but also to resolve
the evolutionary processes in play. In this respect, Chapters 17, 18, and 19, covering
cytogenetic techniques such as flow cytometry, chromosome banding, fluorescent in situ
hybridization (FISH), and genomic in situ hybridization (GISH), are essential to provide
tools allowing the assessment of plant genome size, ploı̈dy, aneuploidy, reproductive mode,
species relationships, and interspecific hybrids. Moreover, the generation of large sets of
SNP markers through NGS technologies now allows detailed population genomics studies
(Chapter 16) that can help to resolve the evolutionary processes in play in natural popula-
tions through the analysis of population structure, the inference of population splits and
exchanges, and the detection of footprints of natural or artificial selection. Although the
primary focus of plant taxonomy is on the delimitation of species, molecular approaches now
provide a better understanding of evolutionary processes, at species and population level, a
particularly important issue for some taxonomic complex groups and a prerequisite to
resolve speciation processes. This is essential when one wants to apply plant taxonomy to
conservation issues.
St Pierre, Réunion, France Pascale Besse

Contents
Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v
Contributors. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ix
List of Abbreviations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xi
1 Plant Taxonomy: A Historical Perspective, Current Challenges,

and Perspectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Germinal Rouhan and Myriam Gaudeul
2 Guidelines for the Choice of Sequences for Molecular Plant Taxonomy. . . . . . . . 39
Pascale Besse
3 Isolation and Purification of DNA from Complicated Biological Samples . . . . . . 57
Ruslan Kalendar, Svetlana Boronnikova,
and Mervi Sepp€ a nen
4 Herbarium Specimens: A Treasure for DNA Extraction, an Update . . . . . . . . . . . 69
Lenka Záveská Drábková
5 Sequencing of Complete Chloroplast Genomes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
Berthold Heinze
6 Utility of the Mitochondrial Genome in Plant Taxonomic Studies . . . . . . . . . . . . 107
Jérôme Duminil and Guillaume Besnard
7 Nuclear Ribosomal RNA Genes: ITS Region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119
Pascale Besse
8 Plant DNA Barcoding Principles and Limits: A Case Study
in the Genus Vanilla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131
Pascale Besse, Denis Da Silva, and Michel Grisoni
9 High-Throughput Genotyping Technologies in Plant Taxonomy . . . . . . . . . . . . . 149
Monica F. Danilevicz, Cassandria G. Tay Fernandez,
Jacob I. Marsh, Philipp E. Bayer, and David Edwards
10 Genotyping-by-Sequencing Technology in Plant Taxonomy
and Phylogeny. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 167
Félicien Favre, Cyril Jourda, Pascale Besse,
and Carine Charron
11 Development of Microsatellite Markers Using Next-Generation
Sequencing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 179
Hélène Vignes and Ronan Rivallan
12 Amplified Fragment Length Polymorphism: Applications
and Recent Developments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 187
Thotten Elampilay Sheeja, Illathidath Payatatti Vijesh Kumar,
Ananduchandra Giridhari, Divakaran Minoo,
Muliyar Krishna Rajesh, and Kantipudi Nirmal Babu
vii
viii Contents
13 Random Amplified Polymorphic DNA (RAPD)

and Derived Techniques . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219
Kantipudi Nirmal Babu, Thotten Elampilay Sheeja,
Divakaran Minoo, Muliyar Krishna Rajesh,
Kukkamgai Samsudeen, Erinjery Jose Suraby,
and Illathidath Payatatti Vijesh Kumar
14 Inter-Simple Sequence Repeats (ISSR), Microsatellite-Primed
Genomic Profiling Using Universal Primers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 249
Chrissen E. C. Gemmill and Ella R. P. Grierson
15 Retrotransposable Elements: DNA Fingerprinting and the Assessment
of Genetic Diversity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 263
Ruslan Kalendar, Alexander Muterko,
and Svetlana Boronnikova
16 Introduction to Population Genomics Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . 287
Thibault Leroy and Quentin Rougemont
17 The Application of Flow Cytometry for Estimating Genome Size,
Ploidy Level Endopolyploidy, and Reproductive Modes in Plants . . . . . . . . . . . . . 325
Jaume Pellicer, Robyn F. Powell, and Ilia J. Leitch
18 Molecular Cytogenetics (Fluorescence In Situ Hybridization - FISH
and Fluorochrome Banding): Resolving Species Relationships
and Genome Organization . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 363
Sonja Siljak-Yakovlev, Fatima Pustahija,
Vedrana Vičić-Bočkor, and Odile Robin
19 GISH: Resolving Interspecific and Intergeneric Hybrids . . . . . . . . . . . . . . . . . . . . . 381
Nathalie Piperidis
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 395
Contributors
KANTIPUDI NIRMAL BABU • Indian Institute of Spices Research, Kozhikode, Kerala, India
PHILIPP E. BAYER • School of Biological Sciences, University of Western Australia, Perth,
Australia
GUILLAUME BESNARD • CNRS-UPS-IRD, UMR5174, EDB, Université Paul Sabatier,
Toulouse, France
PASCALE BESSE • UMR PVBMT, Universite de la Reunion, St Pierre, Réunion, France
SVETLANA BORONNIKOVA • Department of Botany and Genetics of Plants, Faculty of Biology,
Perm State University, Perm, Russia
CARINE CHARRON • CIRAD, UMR PVBMT, St Pierre, La Réunion, France
MONICA F. DANILEVICZ • School of Biological Sciences, University of Western Australia,
Perth, Australia
DENIS DA SILVA • Université de La Réunion, UMR PVBMT, St Pierre, La Réunion, France
JÉRÔME DUMINIL • DIADE, University of Montpellier, IRD, Montpellier, France
DAVID EDWARDS • School of Biological Sciences, University of Western Australia, Perth,
Australia
FÉLICIEN FAVRE • Université de La Réunion, UMR PVBMT, St Pierre, La Réunion, France
CASSANDRIA G. TAY FERNANDEZ • School of Biological Sciences, University of Western
Australia, Perth, Australia
MYRIAM GAUDEUL • Institut de Systématique, Evolution, Biodiversité (ISYEB), Muséum
national d’Histoire naturelle, Sorbonne Université, Ecole Pratique des Hautes Etudes,
Université des Antilles, CNRS, Paris, France
CHRISSEN E. C. GEMMILL • School of Science, University of Waikato, Hamilton, New Zealand
ANANDUCHANDRA GIRIDHARI • Indian Institute of Spices Research, Kozhikode, Kerala, India
ELLA R. P. GRIERSON • Plant & Food Research, Palmerston North, New Zealand
MICHEL GRISONI • CIRAD, UMR PVBMT, St Pierre, La Réunion, France
BERTHOLD HEINZE • Department of Genetics, Austrian Federal Research Centre for Forests
(BFW), Vienna, Austria
CYRIL JOURDA • CIRAD, UMR PVBMT, St Pierre, La Réunion, France
RUSLAN KALENDAR • Department of Agricultural Sciences, Viikki Plant Science Centre and
Helsinki Sustainability Centre, University of Helsinki, Helsinki, Finland; National
Laboratory Astana, Nazarbayev University, Nur-Sultan, Kazakhstan
ILLATHIDATH PAYATATTI VIJESH KUMAR • Indian Institute of Spices Research, Kozhikode,
Kerala, India
ILIA J. LEITCH • Department of Comparative Plant and Fungal Biology, Royal Botanic
Gardens, Kew, Richmond, Surrey, UK
THIBAULT LEROY • Montpellier Institute of Evolutionary Sciences (ISEM), Université de
Montpellier, Montpellier, France; Department of Botany and Biodiversity Research,
University of Vienna, Vienna, Austria
JACOB I. MARSH • School of Biological Sciences, University of Western Australia, Perth,
Australia
DIVAKARAN MINOO • Providence Women’s College, Kozhikode, Kerala, India
ALEXANDER MUTERKO • The Federal Research Center Institute of Cytology and Genetics,
Novosibirsk, Russian Federation
ix
x Contributors
JAUME PELLICER • Department of Comparative Plant and Fungal Biology, Royal Botanic
Gardens, Kew, Richmond, Surrey, UK; Department of Biodiversity, Institut Botànic de
Barcelona (IBB, CSIC-Ajuntament de Barcelona), Barcelona, Spain
NATHALIE PIPERIDIS • SRA, Sugar Research Australia, Te Kowai, QLD, Australia
ROBYN F. POWELL • Department of Comparative Plant and Fungal Biology, Royal Botanic
Gardens, Kew, Richmond, Surrey, UK
FATIMA PUSTAHIJA • Faculty of Forestry, University of Sarajevo, Sarajevo, Bosnia and
Herzegovina
MULIYAR KRISHNA RAJESH • Central Plantation Crops Research Institute, Kasaragod,
Kerala, India
RONAN RIVALLAN • CIRAD, UMR AGAP, Montpellier, France; AGAP, University of
Montpellier, CIRAD, INRAe, Montpellier SupAgro, Montpellier, France
ODILE ROBIN • University Paris-Saclay, CNRS, AgroParisTech, Ecologie Systématique
Evolution, Orsay, France
QUENTIN ROUGEMONT • Département de Biologie, Institut de Biologie Intégrative et des
Systèmes (IBIS), Université Laval, Quebec, QC, Canada
GERMINAL ROUHAN • Institut de Systématique, Evolution, Biodiversité (ISYEB), Muséum
national d’Histoire naturelle, Sorbonne Université, Ecole Pratique des Hautes Etudes,
Université des Antilles, CNRS, Paris, France
KUKKAMGAI SAMSUDEEN • Central Plantation Crops Research Institute, Kasaragod, Kerala,
India
€
MERVI SEPPANEN • Department of Agricultural Sciences, Viikki Plant Science Centre and
Helsinki Sustainability Centre, University of Helsinki, Helsinki, Finland
THOTTEN ELAMPILAY SHEEJA • Indian Institute of Spices Research, Kozhikode, Kerala, India;
Division of Crop Improvement and Biotechnology, ICAR-Indian Institute of Spices
Research, Kozhikode, Kerala, India
SONJA SILJAK-YAKOVLEV • University Paris-Saclay, CNRS, AgroParisTech, Ecologie
Systématique Evolution, Orsay, France
ERINJERY JOSE SURABY • Indian Institute of Spices Research, Kozhikode, Kerala, India
VEDRANA VIČIĆ-BOČKOR • Faculty of Science, Department of Molecular Biology, University of
Zagreb, Zagreb, Croatia
HÉLÈNE VIGNES • CIRAD, UMR AGAP, Montpellier, France; AGAP, University of
Montpellier, CIRAD, INRAe, Montpellier SupAgro, Montpellier, France
LENKA ZÁVESKÁ DRÁBKOVÁ • Laboratory of Pollen Biology, Institute of Experimental Botany
of the Czech Academy of Sciences, Prague, Czech Republic
List of Abbreviations
ADBC Advancing digitization of biological collections

AFLP Amplified fragment length polymorphism
AFLP-RGA Resistance gene analog anchored AFLP
AIMS Amplification of insertion mutagenized sites
AMP-PCR Anchored microsatellite-primed PCR
APG Angiosperm phylogeny group
AP-PCR Arbitrary primed polymerase chain reaction
CAPS Cleaved amplified polymorphic sequences
CBC Compensatory base change
CBD Convention on biological diversity
CBOL Consortium for barcoding of life
cDNA-AFLP Complementary DNA AFLP
CETAF Consortium of European taxonomic facilities
CID Cultivar identification diagram
CNV Copy number variation
cpDNA Chloroplast DNA
CTAB Cetyltrimethyl ammonium bromide
DAF DNA amplification fingerprinting
DArT Diversity array technology
DD Differential display
DNA Deoxyribonucleic acid
DOI Digital object identifiers
EDGF Enzymatic digestion and glass-fiber filtration
EDIT European Distributed Institute of Taxonomy
ERV Endogenous retrovirus
FCM Flow cytometry
FCSS Flow cytometric seed screening
FISH Fluorescence in situ hybridization
GBS Genotyping by sequencing
GEA Genotype-environment associations
GISH Genomic in situ hybridization
GPA Genotype-phenotype associations
GSPC Global strategy for plant conservation
GTI Global taxonomic initiative
HTS High-throughput sequencing technologies
IAM Infinite allele model
ICN International Code of Nomenclature for algae, fungi, and plants
iDigBio Integrated Digitized Biocollections
Indel Insertion/deletion
iPBS Inter-PBS amplification
IPNI International Plant Names Index
IRAP Inter-retrotransposons amplified polymorphism
ISSR Inter-simple sequence repeats
ISTR Inverse sequence tagged repeat
ITS Internal transcribed spacer
xi
xii List of Abbreviations
LARDs Large retrotransposon derivatives

LCNG Low-copy nuclear genes
LD Linkage disequilibrium
LGT Lateral gene transfer
LINE Long interspersed repetitive element
LTR Long terminal repeat
M-AFLP Microsatellite-amplified fragment length polymorphism
MGE Mobile genetic element
MITE Miniature inverted-repeat transposable element
MOTU Molecular operational taxonomic units
MSAP Methylation-sensitive amplified polymorphism
mtDNA Mitochondrial DNA
mtpt Plastid-derived region
nDNA Nuclear DNA
NGS Next-generation sequencing
NOR Nucleolar organizing region
nrDNA Nuclear ribosomal RNA genes
NSF National Science Foundation
ORF Open reading frame
PAV Presence and absence variation
PBS Primer binding site
PCA Principal component analysis
PCR Polymerase chain reaction
PEET Partnerships for Enhancing Expertise in Taxonomy
pGWAS Population genome-wide association study
PLOP Pre-labeled oligonucleotide probes
QTL Quantitative trait loci
RAD Restriction-site reduced complexity approach
RAHM Random amplified hybridization microsatellites
RAMPO Random amplified microsatellite polymorphism
RAPD Randomly amplified polymorphic DNA
rDNA Ribosomal DNA
RE Restriction Enzyme
REMAP Retrotransposon-microsatellite amplification polymorphism
RFLP Restriction fragments length polymorphism
RL Restriction ligation
RNA Ribonucleic acid
RTE Retrotransposable elements
SAMPL Selective amplification of microsatellite polymorphic loci
SCAR Sequence characterized amplified region
SDAFLP Secondary digest AFLP
SINE Short interspersed nuclear element
SMM Stepwise mutation model
SNP Single nucleotide polymorphism
SRAP Sequence-related amplified polymorphism
SSAP Sequence-specific amplified polymorphism
SSCP Single strand conformation polymorphism
SSR Simple sequence repeats
STR Simple tandem repeats
List of Abbreviations xiii
TCG Taxonomic complex group

TDF Transcript-derived fragment
TE Transposable element
TE-AFLP Three endonucleases AFLP
TRIM Terminal-repeat retrotransposons in miniature
TSDs Target site duplications
VNTR Variable number of tandem repeats
Chapter 1
Plant Taxonomy: A Historical Perspective, Current

Challenges, and Perspectives
Germinal Rouhan and Myriam Gaudeul
Abstract
Taxonomy is the science that explores, describes, names, and classifies all organisms. In this introductory
chapter, we highlight the major historical steps in the elaboration of this science, which provides baseline
data for all fields of biology and plays a vital role for society but is also an independent, complex, and sound
hypothesis-driven scientific discipline.
In a first part, we underline that plant taxonomy is one of the earliest scientific disciplines that emerged
thousands of years ago, even before the important contributions of the Greeks and Romans (e.g., Theo-
phrastus, Pliny the Elder, and Dioscorides). In the fifteenth–sixteenth centuries, plant taxonomy benefited
from the Great Navigations, the invention of the printing press, the creation of botanic gardens, and the use
of the drying technique to preserve plant specimens. In parallel with the growing body of morpho-
anatomical data, subsequent major steps in the history of plant taxonomy include the emergence of the
concept of natural classification, the adoption of the binomial naming system (with the major role of
Linnaeus) and other universal rules for the naming of plants, the formulation of the principle of subordina-
tion of characters, and the advent of the evolutionary thought. More recently, the cladistic theory (initiated
by Hennig) and the rapid advances in DNA technologies allowed to infer phylogenies and to propose true
natural, genealogy-based classifications.
In a second part, we put the emphasis on the challenges that plant taxonomy faces nowadays. The still
very incomplete taxonomic knowledge of the worldwide flora (the so-called taxonomic impediment) is
seriously hampering conservation efforts that are especially crucial as biodiversity has entered its sixth
extinction crisis. It appears mainly due to insufficient funding, lack of taxonomic expertise, and lack of
communication and coordination. We then review recent initiatives to overcome these limitations and to
anticipate how taxonomy should and could evolve. In particular, the use of molecular data has been
era-splitting for taxonomy and may allow an accelerated pace of species discovery. We examine both
strengths and limitations of such techniques in comparison to morphology-based investigations, we give
broad recommendations on the use of molecular tools for plant taxonomy, and we highlight the need for an
integrative taxonomy based on evidence from multiple sources.
Key words Classification, Floras, DNA, History, Molecular taxonomy, Molecular techniques, Mor-
pho-anatomical investigations, Plant taxonomy, Species, Taxonomic impediment
Pascale Besse (ed.), Molecular Plant Taxonomy: Methods and Protocols, Methods in Molecular Biology, vol. 2222,
https://doi.org/10.1007/978-1-0716-0997-2_1, © Springer Science+Business Media, LLC, part of Springer Nature 2021
1
2 Germinal Rouhan and Myriam Gaudeul
1 Introduction
Adapting the famous aphorism of Theodosius Dobzhansky [1],

could we dare to say that nothing in biology makes sense except
in the light of taxonomy? Maybe yes, considering that most of
biology relies on identified—and so described—species that are
end products of taxonomy. Taxonomic information is obviously
crucial for studies that analyze the distribution of organisms on
Earth, since they need taxonomic names for inventories and
surveys. But names are also needed to report empirical results
from any other biological study dealing with, e.g., biochemistry,
cytology, ecology, genetics, or physiology: even if working an entire
life on a single species, e.g., Arabidopsis thaliana (L.) Heynh., a
molecular biologist will focus all his/her research on numerous
plants that all represent this species as delimited by taxonomy.
Thus, taxonomy provides names, but it is not only a ‘biodiversity-
naming’ service: it is also a scientific discipline requiring theoretical,
empirical, and epistemological rigor [2]. Names represent scientific
hypotheses on species boundaries, and to put forward such hypoth-
eses involves gathering information from characters of the organ-
isms and adopting a species concept (see Note 1 for an overview of
the main species concepts). Morphology, anatomy, and genetics are
the main sources of characters used in today’s plant taxonomy. Not
without noting that these types of characters all bring potentially
valuable evidence, the focus of this book is on the use of nucleic
acids—and genome size and chromosomes—for a reliable and
efficient taxonomy.
Before discussing how to choose genomic regions to be studied
in order to best deal with particular taxonomic issues (Chapter 2),
this chapter aims to summarize the history of taxonomy and to
highlight that plant molecular taxonomy emerged from an ancient
discipline that has been, and is still, central to other scientific
disciplines and plays a vital role for society. We will also give a
brief overview of the general background into which plant taxon-
omy is performed today and propose some general considerations
about molecular taxonomy.
2 Taxonomy and Taxon: Terminology and Fluctuating Meanings
It is not before 1813 that the Swiss botanist Augustin Pyramus De

Candolle (1778–1841) invented the neologism ‘taxonomy’ from
the Greek ταξις (order) and νoμoς (law, rule) and published it for
the first time in his book Théorie élémentaire de la Botanique (‘Ele-
mentary Theory of Botany,’ [3]). He defined this scientific disci-
pline as the ‘theory of the classifications applied to the vegetal
kingdom,’ which he considered as one of the three components
Plant Taxonomy History and Prospects 3
of botany along with glossology—‘the knowledge of the terms used

to name plant organs,’ and phytography—‘the description of plants
in the most useful way for the progress of science.’
Much later, the Global Biodiversity Assessment of the United
Nations Environment Programme (UNEP; [4]) defined taxonomy
as ‘the theory and practice of classifying organisms,’ including the
classification itself but also the delimitation and description of taxa,
their naming, and the rules that govern the scientific nomenclature.
Today, depending on the authors, taxonomy is viewed either as a
synonym for the ‘systematics’ science—also called biosystematics
[5, 6]—including the task of classifying species, or only as a com-
ponent of systematics restricted to the delimitation, description,
and identification of species. This latter meaning of taxonomy
emerged lately, with the advent of phylogenetics as another com-
ponent of systematics that allows classifications based on the evolu-
tionary relationships among taxa [7].
Thus, it is ironical that taxonomy and systematics, which deal in
particular with classifications and relationships between organisms,
often themselves require clarifications on their relative circumscrip-
tions and meanings before being used [8]. This book will consider
plant taxonomy in the broadest sense, from, e.g., species delimita-
tion based on different molecular techniques—to focuses on popu-
lation genomics methods, or studies resolving interspecific and
intergeneric hybrids.
Incidentally, it is interesting to note that the word ‘taxon’—
plural: taxa—was invented much later (Lam, in [9]) than ‘taxon-
omy’: a taxon is a theoretical entity intended to replace terms such
as ‘taxonomic group’ or ‘biodiversity unit’ [10], and ‘taxon’ refers
to a group of any rank in the hierarchical classification, e.g., species,
genus, or family.
3 A Historical Perspective to Plant Taxonomy
3.1 One Delimiting, describing, naming, and classifying organisms are activ-
of the Earliest ities whose origins are obviously much older than the word ‘taxon-
Scientific Disciplines omy’—which dates back to the nineteenth century; see above. The
use of oral classification systems likely even predated the invention
of the written language ca. 5600 years ago. Then, as for all vernac-
ular classifications, the precision of the words used to name plants
was notably higher for plants that were used by humans. There was
no try to link names and organisms in hierarchical classifications
since the known plants were all named following their use: some
were for food, others for medicines, poisons, or materials. As early
as that time, several hundreds of plant organisms of various kinds
were identified, while relatively few animals were known and
named—basically those that were hunted or feared [11].
These early classifications, that were exclusively utilitarian, per-

sisted until the fifteenth–sixteenth centuries although some major
advances were achieved, mainly by ancient Greeks and Romans. It
was perceptible that the Greeks early considered plants not just as
useful, but also as beautiful, taking a look at paintings in Knossos
(1900 BC) that indeed show useful plants like barley, fig, and olive,
but also narcissus, roses, and lilies. The Greek Theophrastus
(372–287 BC), famous as the successor of Aristotle at the head of
the Lyceum, is especially well known as the first botanist and the
author of the first written works on plants. Interested in naming
plants and finding an order in the diversity of plants, he could have
been inspired by Aristotle who started his Metaphysics book with the
sentence: ‘All men by nature desire to know.’ Theophrastus is
indeed the first one to provide us with a philosophical overview of
plants, pointing out important fundamental questions for the
development of what will be later called taxonomy, such as ‘what
have we got?’ or ‘how do we differentiate between these things?’
He was moreover the first one to discuss relationships among
plants, and to suggest ways to group them not just based on their
usefulness or uses. Thus, in his book Enquiry into Plants, he
described ca. 500 plants—probably representing all known plants
at that time—that he classified as trees, shrubs, undershrubs, and
herbs. He also established a distinction between flowering and
nonflowering plants, between deciduous and evergreen trees, and
between plants that grew in water and those that did not. Even if
80% of the plants included in his works were cultivated, he had
realized that ‘most of the wild kinds have no names, and few know
about them,’ highlighting the need to recognize, describe, and
name plants growing in the wild [12]. Observing and describing
the known plants, he identified many characters that were valuable
for later classifications. For instance, based on his observations of
plants sharing similar inflorescences—later named ‘umbels’—he
understood that, generally, floral morphology could help to cluster
plants into natural groups and, several centuries later, most of these
plants showing umbels were indeed grouped in the family Umbel-
liferae—nowadays Apiaceae.
Theophrastus was way ahead of his time, to such a point that his
botanical ideas and concepts became lost during many centuries in
Europe. But his works survived in Persia and Arabia, before being
translated back into Greek and Latin and rediscovered in Europe in
the fifteenth century. During this long Dark Age for botany—like
for all other natural sciences—in Europe, the Roman Pliny the
Elder (23–79 AD) and the Greek Dioscorides (~40–90 AD), in
first century AD, have however been two important figures.
Although they did not improve the existing knowledge and meth-
ods about the description, naming, or classifications of plants, they
compiled the available knowledge and their written works were
renowned and widely used. The Naturalis Historia of Pliny
(77 AD) was indeed a rich encyclopedia of the natural world,

gathering 20,000 facts and observations reported by other authors,
mostly from Greeks like Theophrastus. At the same time in Greece,
plants were almost only considered and classified in terms of their
medical properties. The major work of Dioscorides De Materia
Medica (ca. 77 AD) was long the sole source of botanical informa-
tion (but at that time, botany was only considered in terms of
pharmacology) and was repeatedly copied until the fifteenth cen-
tury in Europe. Juliana’s book—Juliana Anicia Codex, sixth cen-
tury; Fig. 1—is the most famous of these copies, well known
because it innovated by adding beautiful and colorful plants illus-
trations to the written work of Dioscorides. If some paintings could
be seen as good visual aids to identification—which should be
considered as an advance for taxonomy—others, however, were
fanciful [12]. All those plant books, called ‘herbals’ and used by
herbalists—who had some knowledge about remedies extracted
from plants—throughout the Middle Ages, did not bring any
other substantial progress.
3.2 Toward With the Renaissance, the fifteenth and sixteenth centuries saw the
a Scientific beginning of the Great Navigations—e.g., C. Columbus discovered
Classification of Plants the New World from 1492; Vasco da Gama sailed all around Africa
to India from 1497; F. Magellan completed the first circumnavi-
gation of Earth in 1522—allowing to start intensive and large-scale
naturalist explorations around the world: most of the major terri-
tories, except Australia and New Zealand, were discovered as soon
as the middle of the sixteenth century, greatly increasing the num-
ber of plants that were brought back in Europe either by sailors
themselves or naturalists on board. At that time, herbalists still
played a major role in naming and describing plants, in association
with illustrators who were producing realistic illustrations. But
naming and classifying so numerous exotic and unknown plants
from the entire world would not have been possible without three
major inventions. Firstly, the invention of the Gutenberg’s printing
press with moveable type system (1450–1455) made written works
on plants largely available in Europe—the first Latin translation of
Theophrastus’ books came out in 1483. Secondly, the first botanic
gardens were created in Italy in the 1540s, showing the increasing
interest of the population for plants and allowing teaching botany.
Thirdly, in the botanic garden of Pisa, the Italian Luca Ghini
(1490–1556) invented a revolutionary method for preserving—
and so studying—plants, consisting in drying and pressing plants
to permanently store them in books as ‘hortus siccus’ (dried gar-
den), today known as ‘herbaria’—or ‘herbarium specimens.’ These
perennial collections of dried plants were—and are still—a keystone
element for plant taxonomy and its development: from that time,
any observation and experimental result could be linked to specific
plant specimens available for further identification, study of
Fig. 1 Painting of a Cyclamen plant, taken from the Juliana’s book, showing the flowering stems rising from
the upper surface of the rounded corm. According to Dioscorides, those plants were used as purgatives,
antitoxins, skin cleansers, labor inducers, and aphrodisiacs
morphology, geographic distribution, ecology, or any other fea-

tures. In short, Ghini provided with herbaria the basis of reproduc-
ibility that is an essential part of the scientific method [13].
A student of Ghini, Andrea Cesalpino (1519–1603), was the
first one since the Ancient Greeks to take over the work of Theo-
phrastus, and to discuss it. He highlighted that plants should be
classified in a more natural and rational way than the solely utilitar-
ian thinking. Convinced that all plants have to reproduce, he
provided a new classification system primarily based on seeds and
fruits: in De Plantis libri XVI (1583), he described 1500 plants that
he organized into 32 groups such as the Umbelliferae and Compo-
sitae—currently Apiaceae and Asteraceae, respectively. Cesalpino
also made a contribution to the naming of plant names, sometimes
adding adjectives to nouns designing a plant, e.g., he distinguished
Edera spinosa (spiny ivy) from Edera terrestris (creeping ivy). This
could be seen as a prefiguration of the binomial naming system that
was established in the eighteenth century and is still used in taxon-
omy. But the science of scientific naming was only starting and
plants—like other living beings—were usually characterized by
several words forming polynomial Latin names: for instance,
tomato was designed as Solanum caule inermi herbaceo, foliis pin-
natis incisis, which means ‘Solanum with a smooth herbaceous
stem and incised pinnate leaves’ [14] (Fig. 2).
Cesalpino contributed to the emergence of the concept of
natural classification, i.e., a classification reflecting the ‘order of
Nature.’ This latter expression involved different interpretations
and classifications through the history of taxonomy, but a natural
classification was always intended to reflect the relationships among
plants. Because the Evolutionary thought was not developed yet, it
basically resulted in clustering plants with similar morphological
features. So, it must be noted that the distinction between artificial
and natural classifications—respectively named ‘systems’ and
‘methods’ at the end of the eighteenth century—is a modern
interpretation of the past classifications. Taking advantage of both
technical progresses like microscopy—in the seventeenth century—
and scientific methods inspired by Descartes (1596–1650), several
attempts were made to reach such a natural classification. For
example, Bachmann—also known as Rivin or Rivinus
(1652–1723)—based his classification on the corolla shape in
Introductio ad rem herbariam in 1690. Altogether, the major inter-
est of these classifications is that they triggered investigations on
many morpho-anatomical characters that could be used by later
taxonomists to describe and circumscribe plant species. The British
John Ray (1627–1705) innovated by not relying anymore on a
single characteristic to constitute groups of plants: he suggested
natural groupings ‘from the likeliness and agreement of the princi-
pal parts’ of the plants, based on many characters—mostly relative
to leaves, flowers, and fruits. He documented more than 17,000
worldwide species in Historia Plantarum (1686–1704) and distin-
guished flowering vs. nonflowering plants, and plants with one
cotyledon, which he named ‘monocotyledons,’ vs. plants with
two cotyledons, ‘dicotyledons.’ Ray also played a major role in
the development of plant taxonomy—and more generally of plant
science—by creating the first text-based dichotomous keys that he
used as a means to classify plants [15].
Fig. 2 Herbarium specimen from the Tournefort’s Herbarium (housed at the Paris
national Herbarium, Muséum national d’Histoire naturelle, MNHN) displaying a
label with the hand-written polynomial name ‘Aconitum caeruleum, glabrum,
floribus consolid(ae) regalis’
In contrast to Ray and his method intended to be natural, his

French contemporary Joseph Pitton de Tournefort (1656–1708)
explored, in his Elements de Botanique (1694), the possibility of
classifying plants based on only a few characteristics related to the
corolla of flowers, creating an artificial system. The success of
Tournefort’s system resulted from the ease to identify groups of
plants based on the number and relative symmetry of the petals of a
flower. Within his system, Tournefort precisely defined 698 enti-

ties—‘Institutiones rei herbariae,’ 1700—each being called a genus,
plural: genera. The genus concept was new and contributed to a
better structuration of the classification.
3.3 Naming Plant In spite of the numerous new ideas and systems produced from the
Names: Major 16th to the middle of the eighteenth century, names of plants still
Advances by Linnaeus consisted in polynomial Latin names, i.e., a succession of descrip-
tors following the generic name. This led to a rather long, compli-
cated, and inoperative means to designate plants and became
problematic in the context of the Great Explorations, which
allowed the discovery of more and more plants from all over the
world (major explorations with naturalists on board included, e.g.,
the circumnavigation of La Boudeuse under Bougainville from
1766 to 1769, and the travels to the Pacific of J. Cook between
1768 and 1779). To overcome this impediment involving the
naming of plants, the Swedish Carolus Linnaeus (1707–1778)
took a critical step forward for the development of taxonomy.
He suggested dissociating the descriptors of the plant from the
name itself, because according to him, the name should only serve
to designate the plant. Therefore, he assigned a ‘trivial name’ to
each plant (more than 6000 plants in Species Plantarum, 1753)
[16] and this name was binomial, only consisting of two words: the
‘genus’ followed by the ‘species,’ e.g., Adiantum capillus-veneris is
a binomen created by Linnaeus that is still known and used as such
to designate the Venus-hair fern. Although there had been some
attempts of binomials as early as Theophrastus (followed by Cesal-
pino and a few others), Linnaeus succeeded in popularizing his
system as new, universal—applied for all plants and, later on, even
for animals in Systema Naturae [17], and long-lasting. Truly, Species
Plantarum [16] has been a starting point for setting rules in plant
taxonomy. Used since Linnaeus until today, the binomial system
along with other principles for the naming of plants were devel-
oped, standardized, synthesized, and formally accepted by taxono-
mists into a code of nomenclature—initially called ‘Laws of
botanical nomenclature’ [18] and nowadays called the Interna-
tional Code of Nomenclature for algae, fungi, and plants (ICN).
The current code is slightly evolving every 6 years, after revisions
are adopted at an international botanical congress.
Linnaeus also proposed his own artificial classification. With the
goal to describe and classify all plants—and other living beings—
that were ‘put on Earth by the Creator,’ he grouped them based on
the number and arrangement of stamens and pistils within flow-
ers—contrary to Tournefort, who only focused on petals. He called
this classification a ‘sexual system,’ referring to the fundamental
role of flowers in sexual reproduction (Fig. 3). This system included
five hierarchical categories: varieties, species, genera, orders—
equivalent to current families, and classes.
Fig. 3 Linnaeus’s sexual system as drawn by G. D. Ehret for the Hortus Cliffortianus (1735–1748); this
illustration shows the 24 classes of plants that were defined by Linnaeus according to the number and
arrangements of stamens
3.4 The Advent The end of the eighteenth century was conducive to revolutionary
of the Theory ideas in France, including new principles to reach the natural classi-
of Evolution and Its fication. Studying how to arrange plants in space for creating the
Decisive Impact new royal garden of the Trianon in the Palace of Versailles, Bernard
on Taxonomy de Jussieu (1699–1777) applied the key principle of subordination
of characters, which will be published in 1789 by his nephew
Antoine Laurent de Jussieu (1748–1836) in Genera Plantarum
[19]. Bernard and A. L. de Jussieu stated that a species, genus, or
any other taxon of the hierarchical classification should group
plants showing character constancy within the given taxon, as
opposed to the character variability observed among taxa. Since
not all characters are useful at the same level of the classification, the
principle of subordination led to a character hierarchy: characters
displaying higher variability should be given less weight than more
conserved ones in plant classifications. As a result, B. and A. L. de
Jussieu subordinated the characters of flowers—judged more vari-
able and therefore less suitable at higher levels—to the more con-
served characters of seeds and embryos. It was the first application
of this principle in taxonomy, and it could be interpreted today as a
way to limit homoplasy, though the concept of homoplasy had not
been elaborated yet [20].
Whereas botanical taxonomy had long been preponderant and
faster in its development than its zoological counterpart, the trend
was reversed at the beginning of the nineteenth century, especially
with the application of the principle of subordination of characters
to animals by the French biologists Jean-Baptiste de Lamarck
(1744–1829) and Georges Cuvier (1769–1832). New questions
then arose in the mind of taxonomists, who were not only inter-
ested in naming, describing, and classifying organisms anymore,
but also in elucidating how the observed diversity had been gener-
ated. Early explanatory theories included the theory of the trans-
mutation of species, proposed by Jean-Baptiste de Lamarck in 1809
in his Philosophie zoologique [21]. This was the first theory to
suggest the evolution of species, although it involved several mis-
leading assumptions such as the notion of spontaneous genera-
tions. Charles Darwin (1809–1882) published his famous theory
of evolution in On the Origin of Species (1859) [22], and intro-
duced the central concept of descent with modification that later
received extensive support and is still accepted today. This implied
that useful characters in taxonomy, the so-called homologous char-
acters, are those inherited from a common ancestor. Darwin indeed
predicted that ‘our classifications will come to be, as far as they can
be so made, genealogies’ (Darwin 1859, p. 486) [22]. In other
words, since the history of life is unique, only one natural classifica-
tion is possible that reflects the phylogeny. This latter word was
however not coined by Darwin himself, but in 1866 in his Generelle
Morphologie der Organismen [23] by Ernst Haeckel (1834–1919),
who is commonly known for the first illustration of a phylogeny,
although Dayrat [24] evidenced that all Haeckel’s illustrations

should not be interpreted as real evolutionary-based phylogenetic
trees [23] (Fig. 4). However, Darwin did not provide any new
techniques or approaches to reconstruct the phylogeny or assist
practicing taxonomists in their work [25] and, in spite of his major
contributions, plant taxonomists therefore kept applying the
method of classification described by B. and A. L. de Jussieu even
after the onset of the evolutionary thought.
3.5 New Methods In the 1960s, facing the subjectivity of the existing methods to
and New Sources reconstruct phylogenies, the new concept of numerical taxonomy
of Characters proposed an entirely new way of examining relationships among
for a Modern taxa. Robert Sokal (1926–2012) and Peter Sneath (1923–2011)
Taxonomy started developing this concept in 1963 [26], and elaborated it as
an objective method of classification. The method consisted in a
quantitative analysis of overall similarities between taxa, based on a
characters-by-taxa data matrix—with characters divided into char-
acter states—and resulting in pairwise distances among taxa. But
this method was not based on any evolutionary theory and the
resulting diagrams could therefore not be reasonably interpreted
in an evolutionary context, or as an evolutionary classification.
Nevertheless, this theory flourished for a while, greatly benefiting
from rapid advances in informatics.
A crucial change in the way botanists practice taxonomy
occurred with the development of the cladistic theory and recon-
struction of phylogenies—using diagrams called cladograms—to
infer the evolutionary history of taxa. Willi Hennig (1913–1976)
initiated this revolution with his book Grundzüge einer Theorie der
Phylogenetischen Systematik, published in 1950 [27], but his ideas
were much more widely diffused in 1966 with the English transla-
tion entitled Phylogenetic Systematics [28]. The primary principle of
cladism, or cladistics, is not to use the overall similarity among taxa
to reconstruct the phylogeny, since similarity does not necessarily
reflect an actual close evolutionary relationship. Instead, Hennig
only based the phylogenetic classification on derived characters,
i.e., the characters that are only inherited from the last common
ancestor to two taxa—as opposed to the primitive characters. Every
taxonomic decision, from a species definition to a system of higher
classification, was to be treated as a provisional hypothesis, poten-
tially falsifiable by new data [29]. This new method benefited from
an increasing diversity of sources of characters to be considered,
thanks to the important technological advances accomplished in
the 1940s and 1950s in cytology, ecology, and especially in
genetics.
The discovery of the double helical structure of the DNA
molecule in 1953, by James Watson and Francis Crick, followed
by the possibility to target specific fragments of the genome for
selectively amplifying DNA—the Polymerase Chain Reaction
Fig. 4 Illustration from ‘Monophyletischer Stammbaum der Organismen’ (Haeckel 1866): plants form one of
the three main branches of the monophyletic genealogical tree of organisms
(PCR) was invented by Karry Mullis in 1986 [30]—have dramati-

cally changed biology. In particular, the introduction of DNA
sequence data has been era-splitting for plant taxonomy, offering
access to numerous characters and statistical approaches. Thus, at
the turn of the twenty-first century, the use of molecular data and
new tree-building algorithms—with probabilistic approaches—led
the Angiosperm Phylogeny Group (APG) to better circumscribe all
orders and families of flowering plants [31–34]. Similarly, the Pte-
ridophyte Phylogeny Group reached a consensual classification for
free-sporing vascular plants (ferns and lycophytes) to the genus
level [35]. Such collaborative initiatives have improved to a great
extent our understanding of the plant classification based on evo-
lutionary relationships. Many long-standing views of deep-level
relationships were drastically modified at the ordinal level, and to
a lesser extent at the familial level in flowering plants. One of the
most striking changes is the abandonment of the long-recognized
monocot-dicot split, since monocots—class Liliopsida—were
found to be derived from within a basal grade of families that
were traditionally considered as dicots—class Magnoliopsida.
Another outstanding finding resulting from analyses of molecular
data has been that horsetails and ferns together are the closest
relatives to seed plants, necessitating the abandonment of the pre-
vailing view that ferns and horsetails represent paraphyletic succes-
sive grades of increasing complexity in early vascular plant
evolution, which eventually led to the more complex seed plants,
and ultimately to angiosperms [36]. Thus, the more or less intuitive
classifications proposed since the beginning of the twentieth cen-
tury [37–41] have progressively been less used, as a consequence of
the modifications brought to the classification by molecular
results [42].
Taxonomy took advantage of molecular data not only for
improving plant classification or species delineation, but also for
species-level identification with the development of the DNA bar-
coding initiative since the early 2000s. DNA barcoding is based on
the premise that a short standardized DNA sequence can allow
distinguishing individuals of different species because genetic vari-
ation between species is expected to exceed that within species. It
was first promoted by Paul Hebert for animals [43] and later
supported by international alliances of research organizations like
the Consortium for Barcoding of Life (CBOL; http://barcoding.si.
edu), which includes a Plant working group, or the China Plant
Barcoding of Life Group.
The long history of plant taxonomic research and its numerous
contributors, both for theoretical concepts and the practical accu-
mulation of knowledge, allowed the development of an indepen-
dent, complex, and sound hypothesis-driven scientific discipline
that explores, describes, documents the distribution of, and classi-
fies taxa. It is clearly not restricted to, e.g., identifying specimens
and establishing species lists, but it nevertheless also provides basic

knowledge that is required to address a wide range of research
questions and serve stakeholders in government agencies and inter-
national biodiversity organizations (for management of agriculture
pests, development of new pharmaceutical compounds, control of
trade in endangered species, management of natural resources, etc.;
[29, 44–48]). However, taxonomy is faced with the enormous
existing plant diversity, and one still unanswered question resides
in the extent of plant diversity: how many species are there on
Earth?
4 Plant Taxonomy Today: Current Challenges, Methods, and Perspectives
4.1 How Many Plant Linnaeus’ Species Plantarum, published in 1753, was one of the
Species Are There? first key attempts to document the diversity of plants on a global
scale [16]. In this work, Linnaeus recognized more than 6000
species but erroneously concluded that ‘the number of plants in
the whole world is much less than commonly believed, I ascertained
by fairly safe calculation [. . .] it hardly reaches 10,000’ [16]. Later
on, in 1824, the Swiss A.P. de Candolle, in his Prodromus Systematis
Naturalis Regni Vegetabilis [49], aimed to produce a flora of the
world: he included 58,000 species in seven volumes. Today, we
know that the magnitude of plant diversity is much larger, although
we are uncertain of the exact number of plant species.
There are two questions in estimating the total number of plant
species: the first one is how many species have already been described;
the second one is how many more species are presently unknown to
science.
Our uncertainty about the number of described species is
mostly due to the fact that taxonomists sometimes gave different
names to the same species inadvertently, especially in the past due to
poor communication means between distant scientists. This led to
the existence of multiple names for a single biological entity, a
phenomenon called synonymy. As a consequence, we know that
more than 1,064,908 vascular plant names were published, as
evidenced by the International Plant Names Index (IPNI)
[50, 51], but they would actually represent only 223,000 to
422,000 accepted species—depending on the method of calcula-
tion ([46, 52] and references therein, [53, 54]), with the most
recent estimates of 383,671 [51] and 351,176 according to
The Leipzig Catalogue of Vascular Plants (LCVP) v.1.0.2 by
Freiberg et al. (unpublished). In addition, the disagreement on a
single species concept (see Note 1) among plant taxonomists means
that species counts can easily differ by an order of magnitude or
more when the same data are examined by different botanists
[55]. This leads to a taxonomic inflation, i.e., an increased number
of species in a given group that is not due to an actual discovery of
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CHAPTER LII
LIFE IN SOUTH AMERICA
While the variety of conditions in South America makes any

treatment of this subject necessarily superficial, a few words beyond
those already said may not be out of place, since it is evident that for
the successful conduct of our trade many persons from the United
States must spend some years or reside permanently in the several
countries. From the descriptions given one should have a fair idea as
to climatic conditions in these and make an intelligent choice of
locality if any is offered. Some persons will be happy in warm Rio or
even in more tropical Pará. Others will prefer Andean heights, from
7000 to 14,000 feet altitude, the higher for persons with sound hearts
only. Most of the cities where one is likely to be stationed have a
fairly temperate climate, and good health conditions, except as
previously indicated.
In respect to social advantages there is considerable variety. In
general the smaller the city the greater the hospitality and the more
will one’s society be cultivated, as is true in the United States also.
On the other hand in the important commercial cities, the English-
speaking folk are numerous enough to make an agreeable society
for themselves, and some South Americans have made the criticism
that the English and Americans hold aloof, apparently preferring their
own company: a mistake from a business point of view and also
nationally. One must, however, have the right qualifications for
cordial recognition anywhere. It has been stated of Buenos Aires that
the fact of membership in the diplomatic corps did not entitle the
gentleman and his family to more than official courtesies; to be
received socially he must be persona grata. This is true to some
extent everywhere. At the same time one who is at the head of a
large commercial establishment is more likely to have social
opportunities than members of the office staff, one of whom, a young
man of unusually good manners and attractive personality
complained to me in Lima some years ago, that he had no social
opportunities. It is different now. On the other hand a young dentist in
a city of Argentina where Americans are few associated with the best
people and married into one of the first families.
At the mining camps of the Americans provision is now made for
the social life of the employes and for exercise and recreation, also
by other large corporations. In general I believe that men enjoy the
life in South America better than their wives. Some of the latter
decline to go or to remain after being there a while: a great mistake if
they have any regard for their husband’s welfare, unless the care of
children or other serious matter compels their return. Many women
are perfectly contented, this depending in some degree on their
location, but chiefly upon their disposition. In the town of Sorata,
Bolivia, I chanced to meet one who seemed perfectly happy, though
she was the only English speaking woman in the place, or within 100
miles.
It is desirable for banks and business houses to give their young
men sufficient salaries to permit them to marry and take their wives
along. It will be better for both parties in the long run. Perhaps there
are no more temptations than in our own large cities, but in most
places there are fewer forms of wholesome recreation. Too many
men in cities and in mining camps have gone to pieces as they say.
Some men prefer life abroad for the reason that they feel less
restraint than in their native home or even in our metropolis, rather
than the responsibility which a real patriot should recognize of
presenting the highest American ideals of manners, conduct, and
business practices. If one cannot be contented without going
somewhere every night, except in Rio or Buenos Aires he might be
unhappy or worse. It would be well if persons everywhere had
sufficient intelligence to enjoy themselves at home with a good book,
a quiet game of cards, even cribbage; but especially books that are
worth while, valuable as literature or as containing information about
the world in general or on matters connected with business. “Movies”
are found almost everywhere; in the larger cities, theatres and a long
or short season of opera; clubs with opportunities for golf, tennis,
and other sports; often horse races. In smaller towns horseback
riding is a common, sometimes the chief diversion; but in such
places one sooner enters the social life of the community. Some
Americans say they would not take a wife to such a place, but if she
is wise she will go.
Punctiliousness in dress as well as in manners is more highly
regarded in South America than in the United States. Evening dress
is more general in large cities than in most of ours, and correct
afternoon dress for men is a more rigid requirement. Some persons
on important missions have astonished the Latins by their
negligence in this regard. Of course a gentleman is a gentleman the
world over and such an one will have no trouble. It is unnecessary to
imitate certain mannerisms of many South Americans, yet a little
more effusiveness is easily acquired and may be an improvement on
the coldness of the Anglo Saxon. It will be noticed that men regularly
lift their hats to each other, that they shake hands much oftener,
when you come and when you go, make more inquiries after your
health, etc. That they pat each other on the back, give mild hugs, or
at times kiss one another (not you), will perhaps not seem so terrible
as formerly, now that so much has been written about brave
marshals and generals kissing soldiers on both cheeks when
conferring decorations.
Courtesy must not be considered hypocrisy because phrases like
“The house is yours” mean no more than our remarks “I am glad to
see you” or “How are you?” though we may wish the caller in Africa
and have no real interest in his health. Not everywhere is the same
courtesy evident. On my first visit to La Paz in 1903 I noticed that
men frequently stepped from the narrow sidewalk into the gutter to
allow a lady to pass. More recently in a much larger city, still with
some narrow sidewalks, I frequently stepped into the street myself to
avoid crowding past a gentleman (?) who made no move to give
way.
The cost of living is an item of practical interest. Some remarks on
this subject have seemed to me exaggerated. Great diversity exists
in this respect in the different cities: the larger the more expensive,
as in the world generally. In most of the capital cities and chief ports
rents are high; in some places many articles of food are dear with
others cheap; similarly with dry goods and other articles, some
higher than in New York, others lower. Recent unusual conditions
have made sudden changes which may be repeated. Living
expenses were increased by the War, and on account of the influx of
foreigners for business houses. In 1916 rents in Buenos Aires were
lowered in the business centre; but they have now advanced to their
former price or higher. Years ago one of our diplomats there paid
more for his house rent than his entire salary on which others
perhaps have managed to live.
In remote sections, for instance in the Huailas Valley, Peru, in
1906, a sheep cost $1, a lamb 40 cents, a chicken 15 cents,
beefsteak, 9 cents a pound. The cook received $1.50 a month and
board. Fresh fruit and vegetables were almost given away. In Lima
then as now it was very different, some kinds of food were
expensive, others cheap. Coal and kerosene oil are dear everywhere
except in the Huailas Valley where coal is abundant with no market.
In Buenos Aires a few apartment houses and hotels have hot water
heating, but in many places in winter one freezes, or uses an oil
stove or an electric heater, the former the cheaper and more
effective.
Persons of adaptable disposition may spend a few years in South
America with pleasure and profit, returning with broader minds, and
with the ability to command higher salaries than if they had remained
at home.
APPENDIX I
POSTAL REGULATIONS
Much repetition is avoided and probably greater convenience

secured by presenting a summary of the Postal Regulations. All of
the South American Republics are members of the Postal Union. In
November, 1920, a Pan American Postal Federation was formed.
According to the convention adopted, domestic rates will apply to
letters, postal cards, and printed matter, among the various countries
of Latin America and the United States, as soon as they have ratified
the agreement. At present, October, 1921, this has been done by the
United States and by the South American Republics, Argentina,
Bolivia, Brazil, Colombia, and Peru. To these the letter rate is two
cents, postal cards, one cent, return cards two cents; printed matter,
newspapers and periodicals, one cent for four ounces. The old rates
now effective in the other countries will doubtless soon be reduced,
and should therefore be investigated.
Parcel post service has been extended so that parcels weighing
up to 22 pounds may be sent to Argentina, Brazil, Colombia,
Paraguay, and Peru. To Ecuador 20 pounds is the limit; to Bolivia,
Chile, British, Dutch, and French Guiana, Uruguay, and Venezuela,
11 pounds is the maximum. The rate to all is 12 cents a pound or a
fraction thereof; except that to Paraguay, on account of transit
through Argentina, 30 cents additional must be paid for a parcel
weighing 11 pounds or less, and 60 cents for one above that to 22
pounds. In Brazil, this service is limited to Bahia, Bello Horizonte,
Curityba, Manaos, Pará, Pelotas, Pernambuco, Porto Alegre, Rio de
Janeiro (including Petropolis), Rio Grande do Sul, and São Paulo.
Parcels are subject to customs duties, and these with other details
should be investigated. Parcels may be registered for Bolivia, Brazil,
British Guiana, Chile, Ecuador, Peru, Venezuela, but not for the other
countries.
Money orders may be sent to Peru, Bolivia, and Uruguay.
Changes resulting from the Pan American Postal Congress at
Buenos Aires in 1921 will be inaugurated January, 1923 or earlier.
Cable Facilities
On the North Coast, Cartagena has direct cable connection with

Colon and so with New York. To Puerto Colombia a cable has been
laid, which, however, December, 1921, has not yet been opened. A
French company has a line from Salinas near Pará to Cayenne,
Paramaribo, and Martinique, another from La Guaira, to Curaçao,
and Santo Domingo. The Venezuelan Government has its own cable
along the coast from Maracaibo, to La Guaira, Barcelona, and other
points. A British line connects Georgetown, Guiana, with the Port of
Spain, Trinidad.
The West Coast is connected with North America by three lines of
the All America system: one from Nicaragua and two from Panamá
to Santa Elena, Ecuador, one of the latter by way of Buenaventura
and Esmeraldas. The three lines continue south to Callao, one
touching at Paita. Two go on to Iquique and Valparaiso, one touching
at Antofagasta, while a branch comes north from Iquique to Arica to
make connection with La Paz. A cable of another company from
Callao touches at Mollendo, Arica, Antofagasta, La Serena,
Valparaiso, and Concepción.
The East Coast is connected with the cables of the West Coast by
three private wires of the All America Cables over the Andes from
Valparaiso to Buenos Aires, so that they can handle messages to the
Argentine metropolis, 7452 miles from New York, by automatic
methods in 15 minutes. Another cable company has a land line from
Valparaiso to La Plata, where connection is made with its Trans-
Atlantic cable to Africa and Europe. Both companies have short lines
to Montevideo, the focus of the East Coast lines. From here the All
America has a cable to Santos and one to Rio de Janeiro. The other,
the Western Telegraph, has one to Chuy, Uruguay, thence to Rio
Grande do Sul, Santa Catharina, Santos, Rio de Janeiro, Bahia,
Pernambuco, Fortaleza, Maranhão, and Pará, Brazil; and one from
Chuy direct to Rio de Janeiro and Pernambuco. Four cables from the
latter port connect with Africa and Europe. The Western Telegraph
was to lay a cable from Maranhão to Barbados, there to connect with
the Western Union line to Florida. The All America expects to lay a
cable from Cuba south to Rio de Janeiro. The Amazon Company
has a cable up that river from Pará.
Metric System
The Metric System of weights and measures is legal and official in

all the Republics and obligatory in most, in Argentina, Brazil, Chile,
Colombia, Peru, and Venezuela. In the other countries and in some
of these, the old Spanish measures (Portuguese in Brazil) are more
or less used, but these differ in the various countries and are
nowhere like ours. Always to employ the metric system is highly
important and in the above mentioned countries necessary, though
for shipping to some, the weight in pounds must also be given. In
Chile the use of other weights and measures is prohibited; also in
Uruguay, where their importation is forbidden.
APPENDIX II
LEADING BANKS OF SOUTH AMERICA
Including the branches and affiliations of American banks and

banking houses, British banks, and the most important local banks of
each country.
United States Banks
The National City Bank, 55 Wall St., New York City, which led the
way, has branches in six of the South American Republics,
The Mercantile Bank of the Americas, 44 Pine St., New York,
The American Foreign Banking Corporation, 53 Broadway, New
York,
W. R. Grace and Company’s Bank, 7 Hanover Square, New York,
The First National Bank, 70 Federal St., Boston,
The American Express Company, 65 Broadway, New York, with
offices in Buenos Aires, Argentina; Montevideo, Uruguay; and
Valparaiso, Chile; and with correspondents in other cities, performs
some banking service.
British Banks
Important banks with New York offices and with many branches in
South America are:
The Anglo South American Bank, 49 Broadway, New York,
affiliated with
The British Bank of South America, and with
The Commercial Bank of Spanish America, 49 Broadway, New
York;
The London and River Plate Bank, 51 Wall St., New York,
The London and Brazilian Bank, 56 Wall St., New York,
The Royal Bank of Canada, 68 William St., New York.
Branches and Affiliations
National City Bank, Branches: Argentina, Buenos Aires, Rosario;

Brazil, Rio de Janeiro, Santos, São Paulo, Pernambuco; Chile,
Santiago, Valparaiso; Peru, Lima; Uruguay, Montevideo; Venezuela,
Caracas.
Mercantile Bank of the Americas: Affiliated Banks: Colombia,
Banco Mercantil Americano de Colombia, Bogotá Barranquilla,
Cartagena, Medellín, Cali, Girardot, Manizales; Peru, Banco
Mercantil Americano de Peru, Lima, Arequipa, Chiclayo, Callao,
Piura, Trujillo; Venezuela, Banco Mercantil Americano de Caracas,
Caracas, La Guaira, Maracaibo, Puerto Cabello, Valencia; Agency in
Ecuador.
The American Foreign Banking Corporation: Argentina, Buenos
Aires; Brazil, Rio de Janeiro.
W. R. Grace and Company’s Bank: Argentina, Buenos Aires, W. R.
Grace y Cia.; Bolivia, La Paz, W. R. Grace and Company; Brazil, Rio
de Janeiro, Grace and Company; Chile: Santiago, Grace y Cia.,
Valparaiso, W. R. Grace and Company, Iquique, Nitrate Agencies,
Ltd.; Ecuador, Guayaquil, Guayaquil Agencies Company; Peru,
Lima, W. R. Grace and Company; Venezuela, Caracas, Venezuela
Commercial Company.
The First National Bank, Boston: Argentina, Buenos Aires.
The Anglo South American Bank: Chile, Antofagasta, Chillán,
Concepción, Copiapó, Coquimbo, Iquique, Punta Arenas, Santiago,
Talcahuano, Valparaiso; Argentina, Buenos Aires, Bahia Blanca,
Comodoro Rivadavia, Mendoza, Puerto Deseado, Rio Gallegos,
Rosario de Santa Fé, San Julian, San Rafael, Santa Cruz, Trelew;
Peru, Lima; Uruguay, Montevideo.
The British Bank of South America: Argentina, Buenos Aires,
Rosario de Santa Fé; Brazil, Rio de Janeiro, Bahia, Pernambuco,
Porto Alegre, Rio Grande do Sul, São Paulo; Uruguay, Montevideo.
The Commercial Bank of Spanish America: Colombia, Bogotá,
Barranquilla, Medellín; Ecuador, Guayaquil, Manta; Peru, Iquitos;
Venezuela, Caracas, Puerto Cabello.
The London and River Plate Bank: Argentina, Buenos Aires,
Rosario de Santa Fé, Mendoza, Bahia Blanca, Concordia, Córdoba,
Paraná, Tucumán; Brazil, Rio de Janeiro, Pará, Maceió,
Pernambuco, Bahia, Santos, São Paulo, Curityba, Pelotas, Porto
Alegre, Rio Grande do Sul; Chile, Santiago, Valparaiso, Antofagasta;
Colombia, Bogotá; Paraguay, Asunción; Uruguay, Montevideo, Salto,
Paysandú.
The London and Brazilian Bank: Argentina, Buenos Aires, Rosario;
Brazil, Rio de Janeiro, Manaos, Pará, Maranhão, Ceará,
Pernambuco, Bahia, Santos, São Paulo, Curityba, Rio Grande do
Sul, Pelotas, Porto Alegre; Uruguay, Montevideo.
The Royal Bank of Canada: Argentina, Buenos Aires; Brazil, Rio
de Janeiro, Santos, São Paulo; British Guiana, Georgetown, Rose
Hall (Corentyn); Colombia, Barranquilla; Uruguay, Montevideo;
Venezuela, Caracas, Ciudad Bolívar, Maracaibo, Puerto Cabello.
Most if not all of the banks mentioned have correspondents or
agents in the chief cities of the countries where they have no
branches and some have connections in the smaller cities.
The Irving National Bank, Woolworth Building, New York, has
correspondents in the principal cities of South America.
The Guaranty Trust Company, 140 Broadway, New York, is
affiliated with the Mercantile Bank of the Americas and has other
correspondents.
Other Important Banks in South America

Argentina: Buenos Aires, Banco de la Nación Argentina, with 18
branches in as many Argentine cities, Ernesto Tornquist and
Company, Banco de la Provincia de Buenos Aires, American Bank of
the River Plate; La Plata, the Central Bank of the Provincia de
Buenos Aires, which has branches in many cities of the Province.
Bolivia: La Paz, Banco de la Nación Boliviana, branches in
Cochabamba, Oruro, Potosí, Tarija, Uyuni; Banco Francisco
Argandoña, also in Cochabamba and Oruro; Banco Mercantil, also in
Cochabamba, Oruro, Potosí, Tarija, Tupiza, Uyuni; Banco Nacional
de Bolivia, branches in Cochabamba, Oruro, Potosí, Tupiza, Uyuni.
Brazil: Rio de Janeiro, Banco do Brasil, with branches in most of
the Brazilian cities, Banco Nacional Brasileiro; São Paulo, Banco
Commercial do Estado de São Paulo; Bahia, Banco de Bahia; Pará,
Banco de Pará; Pernambuco, Banco do Recife; Bello Horizonte,
Banco Hypothecario e Agricola de Estado de Minas Geraes; etc.
Chile: Santiago, Banco de Chile, branches in many cities; Banco
Español de Chile with branches; Banco de A. Edwards y Cia.;
Valparaiso, Banco de Chile y Argentina, branches in Punta Arenas,
and also in San Julian and Santa Cruz, Argentina.
Colombia: Bogotá, Banco de Bogotá, Banco de Colombia. These
banks have fewer branches, if any, than the Bancos de la Nación
Argentina, de Brasil, or de Chile, Medellín has the Banco de la
Mutualidad, Banco Dugand, and Banco Lopez, found also in
Bucaramanga, and in other cities.
Ecuador: Guayaquil, Banco Comercial y Agricola, Banco del
Ecuador, Mercantile Overseas Corporation, Juan Marcas y Cia.,
correspondent of the Guaranty Trust Company.
Guiana: British, Georgetown, Colonial Bank of London (22 William
St., New York), branches in Henrietta and New Amsterdam; Dutch,
Paramaribo, De Surinaamsche Bank; French, Cayenne, Banque de
la Guyane.
Peru: Lima, Banco del Peru y Londres, branches in most of the
Peruvian cities, Credito Hipotecario del Peru.
Paraguay: Asunción, Banco Mercantil del Paraguay, branches in
Concepción, Encarnación, Pilar, Villa Rica; Banco de la Republica,
branch in Encarnación.
Uruguay: Montevideo, Banco de la Republica Oriental del
Uruguay, with branches in many cities of the country.
Venezuela: Caracas, Banco de Venezuela, many branches; Banco
de Caracas, some branches.
Other American Banks
with facilities for South American Trade are:

New York, American Exchange National Bank, 128 Broadway,
Bank of New York, 48 Wall St., Battery Park National Bank of New
York, 2 Broadway, Canadian Bank of Commerce, 16 Exchange
Place, Lincoln Trust Company, 7 Wall St.
Boston, The Merchants National Bank, 28 State St.
Chicago, Central Trust Company of Illinois, 125 West Monroe St.,
Great Lakes Trust Company.
Cincinnati, The Fifty-Third National Bank.
Detroit, The Peoples State Bank, Fort & Shelby Sts.
Philadelphia, The Philadelphia National Bank, 421 Chestnut St.
Pittsburgh, Mellon National Bank, 514 Smithfield St.
San Francisco, The Crocker National Bank.
Additional banking information may be found in the Exporters’
Encyclopaedia, annual edition; in Commercial Travelers’ Guide to
Latin America, containing lists of banks for each city; and in the
Bankers’ Almanac and Year Book, London, annual, with complete
lists of banks in the cities of all countries.
APPENDIX III
STEAMSHIP LINES TO SOUTH AMERICA
The North Coast
Colombia: Passenger and Freight Lines
New York to Puerto Colombia and Cartagena, Caribbean

Steamship Company, 10 Bridge St., weekly, Five Continent
Steamship Company, 2 Stone St., weekly, United Fruit
Steamship Company Service, 17 Battery Place, weekly, also
to Santa Marta.
Boston to Cartagena, Puerto Colombia, Santa Marta,
United Fruit Company Steamship Service, Long Wharf.
New Orleans to Puerto Colombia, Pacific-Caribbean-Gulf
Line, 630 Common St., fortnightly; Caribbean Steamship
Company, Lykes Bros., monthly.
Grace Line, to Colombian ports, monthly.
Colombia: Freight Only
New York to Cartagena and Puerto Colombia, Tropical

Steamship Corporation, 44 Whitehall St.
Seattle to Cartagena and Puerto Colombia, Tropical
Steamship Pacific-Caribbean-Gulf Line, A. M. Gillespie, Inc.,
Arctic Building, monthly.
Venezuela: Passengers and Freight

New York to La Guaira, Puerto Cabello, Maracaibo, Red
“D” Line, 82 Wall St., weekly to La Guaira, fortnightly to the
other ports.
To Ciudad Bolívar, Trinidad Line, 29 Broadway, fortnightly to
Port of Spain, transshipment.
To Curaçao, Puerto Cabello, La Guaira, Cumaná,
Carupano, and Port of Spain, Trinidad, Royal Netherlands
West India Mail, Funch, Edye, and Company, 25 Broadway,
fortnightly.
New Orleans to La Guaira, Puerto Cabello, Maracaibo,
New Orleans and South American Steamship Company,
Queen and Crescent Bldg., semi-monthly.
Grace Line to Venezuelan ports, monthly.
Venezuela: Freight Only
New York to La Guaira, Puerto Cabello, Maracaibo,

Caribbean Steamship Company, 10 Bridge St., fortnightly.
New Orleans to La Guaira, Puerto Cabello, Maracaibo,
Caribbean Steamship Company, Lykes Bros., monthly.
Guiana: British, Dutch, and French
British Guiana Passengers and Freight
New York to Georgetown, Quebec Steamship Company,

34 Whitehall St., every 10-14 days; Trinidad Line, 22 Pearl
St., fortnightly; Royal Netherlands West India Mail, monthly,
25 Broadway.
New York to Georgetown, Paramaribo, Cayenne, Clyde
Steamship Company, leave Pier 44 North River; fortnightly,
freight only.
Mobile to Georgetown, Windward Island Line, Passengers
and freight, every three weeks.
Dutch and French Guiana: Passengers and Freight
New York to Paramaribo, Royal Netherlands West India

Mail Line, 25 Broadway, monthly.
New York to Cayenne, Trinidad Line, 22 Pearl St.,
transshipment at Port of Spain.
The West Coast
Through Lines to Chile by Panama Canal, and from Pacific

Ports.
Passengers and Freight
New York: Grace Line, 10 Hanover Square, fortnightly to

Callao and Mollendo, Peru; Arica, Iquique, Antofagasta,
Valparaiso, Talcahuano, Chile; 20 days to Valparaiso.
Pacific Steam Navigation Company, Sanderson and Son,
26 Broadway, monthly to Callao, Mollendo, Peru; Arica,
Iquique, Antofagasta, Valparaiso, Chile; 20 days; a line from
Liverpool to same ports, also a line every three weeks from
Arica to Iquique, Antofagasta, Valparaiso, Talcahuano,
Coronel, Corral, Puerto Montt, Punta Arenas.
Compañia sud Americana de Vapores, Wessel, Duval, and
Company, 25 Broad St., monthly to Guayaquil, Ecuador,
Salaverry, Callao, Mollendo, Peru; Arica, Iquique,
Antofagasta, Valparaiso, Chile.
Seattle and San Francisco: Grace Line, Hoge Building,
Seattle, monthly to Talara, Paita, Salaverry, Callao, Pisco,
Mollendo, Peru; Arica, Iquique, Antofagasta, Valparaiso,
Chile; also to Ecuador.
Portland and San Francisco: South American Line, to
Guayaquil, Ecuador; Talara, Callao, Mollendo, Peru;
Antofagasta, Chile.
Freight Only
New York to Peru and Chile, New York and Isthmian

Steamship Lines, J. W. Ryan, 39 Cortland St., monthly.
West Coast Line, Wessel, Duval, and Company, 25 Broad
St., monthly or oftener to Paita, Etén, Salaverry, Callao,
Pisco, Mollendo, Peru; Arica, Iquique, Antofagasta, Taltal,
Chañaral, Coquimbo, Valparaiso, Talcahuano, Chile.
Grace Line, Paita, Etén, Salaverry, Callao, Coquimbo,
Valparaiso, Talcahuano, monthly.
Also from Baltimore, Clarence Cottman Company,
according to demand.
Baltimore to Peru and Chile, Pacific Steam Navigation
Company, Furness, Withy, and Company, 19 South St.,
monthly.
New Orleans to Ecuador, Peru, and Chile, New Orleans
and South American Steamship Line Company, Queen and
Crescent Bldg., monthly to Guayaquil, Ecuador, Callao,
Mollendo, Peru; Arica, Iquique, Antofagasta, Valparaiso,
Chile.
Grace Line, monthly to Ecuador, Peru, and Chile.
Seattle, Portland, San Francisco, and San Pedro to
Colombia, Ecuador, Peru, and Chile, General Steamship
Corporation, Colman Bldg., Seattle, every 20 days to
Buenaventura, Colombia; Guayaquil, Ecuador; Paita, Callao,
Mollendo, Peru; Arica, Antofagasta, Valparaiso, Chile.
Seattle to Colombia, Ecuador, Chile, Rolph Steamship
Company, Hind, Rolph and Company, Henry Building,
monthly to Buenaventura, Colombia; Bahia, Manta,
Guayaquil, Ecuador; Arica, Iquique, Antofagasta, Valparaiso,
Chile.
Portland, Oregon, and San Francisco to Peru and
Chile, Toyo Kisen Kaisha Oregon Pacific Company, Wilcox
Bldg., monthly to Callao, Mollendo, Peru; Arica, Iquique,
Valparaiso, Chile.
Colombia, Ecuador, and Peru
New York to Cartagena, Buenaventura, Guayaquil, Paita,

Etén, Pimentel, Pacasmayo, and Salaverry, every three
weeks; freight only, Grace Line, 10 Hanover Square.
Colombia and Ecuador
New York: Pacific Line every three weeks to

Buenaventura, Colombia; Esmeraldas, Bahia, Manta,
Guayaquil, Ecuador; freight.
Colombia
New York to Buenaventura and Tumaco, Caribbean

Steamship Company, 10 Bridge St., passengers and freight,
monthly.
Other Lines with Transshipment at Colon
New York to Colon, Panama Railroad Steamship Line, 24

State St., weekly, passengers and freight; United Fruit
Company Steamship Service, twice a week to Colon,
passengers and freight; other service to Colon from Boston
and New Orleans.
West Coast Lines from Colon and Panama

Pacific Steam Navigation, 26 Broadway, New York,
fortnightly, to Paita, Pimentel, Etén, Pacasmayo, Salaverry,
Callao, Cerro Azul, Tambo de Mora, Pisco, Lomas, Chala,
Mollendo, Peru; Arica, Iquique, Antofagasta, Coquimbo,
Valparaiso, Talcahuano, Penco, Tomé, Coronel, Chile;
another line fortnightly to Buenaventura, Tumaco, Colombia;
Esmeraldas, Bahia de Caraquez, Manta, Cayo, Machalilla,
Manglar Alto, Ballenita, P. Bolívar, Guayaquil, Ecuador.
Compañia Peruana de Vapores (Peruvian Line), 32
Broadway, New York, every ten days to Guayaquil, Ecuador;
Paita, Pimentel, Etén, Pacasmayo, Salaverry, Chimbote,
Samanco, Casma, Callao, Cerro Azul, Tambo de Mora, Pisco,
Lomas, Chala, Mollendo, Ilo, Peru.
Compañia Sud Americana de Vapores, 25 Broad St.,
New York, fortnightly to Guayaquil, Ecuador, and primary
ports of Peru and Chile; and by transfer to caletero boats
serving Paita, Pimentel, Etén, Pacasmayo, Salaverry,
Chimbote, Samanco, Casma, Huarmey, Supe, Huacho,
Callao, Cerro Azul, Tambo de Mora, Pisco, Lomas, Chala,
Mollendo, Ilo, Peru; Arica, Pisagua, Caleta Buena, Iquique,
Tocopilla, Gatico, Antofagasta, Taltal, Chañaral, Caldera,
Huasco, Coquimbo, Valparaiso, Talcahuno, Penco, Tomé,
Coronel, Lota, Chile.
The Colombian Maritime Company serves Buenaventura
and Tumaco, Colombia.
The East Coast
Lines to Brazil, Uruguay, Argentina
From New York, Passenger and Freight
Lamport and Holt Line, 42 Broadway, fortnightly to

Pernambuco, Bahia, Rio de Janeiro, Santos, Rio Grande do
Sul, Brazil; Montevideo, Uruguay; Buenos Aires, Argentina.
Munson Steamship Line, 67 Wall St., fortnightly to Rio de
Janeiro, Brazil; Montevideo, Buenos Aires.
Lloyd Brasileiro, 44 Whitehall St., fortnightly to
Pernambuco, Bahia, Rio de Janeiro, Santos, Brazil.
Booth Steamship Company, 17 Battery Place, monthly or
oftener to Pará, Manaos (transshipment for Iquitos, Peru),
Maranhão, Ceará, Parnahyba, Maceió, Pernambuco,
Cabedello, Natal; also semi-monthly service to Rio de
Janeiro, Santos, and Rio Grande do Sul, with calls when
required at Bahia, Victoria, Paranaguá, Florianopolis, and São
Francisco.
Norton Line, Norton, Lilly, and Company, 26 Beaver St.,
passenger and freight service expected bi-monthly to
Montevideo and Buenos Aires; sometimes to Rosario. Freight
service semi-monthly to Montevideo, Buenos Aires, Rosario,
occasionally to Santa Fé.
The Royal Mail Steam Packet Company has services from
Liverpool and from Southampton to Brazil, Uruguay, and
Argentina; also a Line around South America by the Straits of
Magellan and through the Panama Canal, and vice versa,
calling at the principal East and West Coast ports.
From New York, Freight Only
Munson Line, 67 Wall St., fortnightly to Rio de Janeiro,

Santos, Montevideo, Buenos Aires.
Donald Line, Oriental Navigation Company, 39 Broadway,
to Rio de Janeiro, Santos, Montevideo, La Plata, Buenos
Aires, Rosario.
Ward Line, New York and Cuba Mail Steamship Company,
foot of Wall St., fortnightly to Pará, Maranhão, Ceará, Natal,
Cabedello, Pernambuco, Maceió, Bahia, Montevideo, La
Plata, Buenos Aires, Rosario.
Prince Line, Furness, Withy and Company, 34 Whitehall
St., fortnightly to Rio de Janeiro, Santos, Rio Grande do Sul,
Porto Alegre, Pelotas, La Plata, Buenos Aires, Rosario.
Commercial South American Line, Moore and McCormack,
Inc., 5 Broadway, monthly to Pernambuco, Bahia, Rio de
Janeiro, Santos, Paranaguá, Rio Grande do Sul, Montevideo,
La Plata, Buenos Aires, Rosario.
National Line, National Steamship Lines, 11 Broadway,
monthly to Pernambuco, Bahia, Rio de Janeiro, Santos, La
Plata, Buenos Aires, Rosario.
New York and Argentine Steamship Company, 50
Broadway, fortnightly to Rio de Janeiro, Santos, Buenos
Aires.
North and South Line, P. Kleppe and Company, 11
Broadway, monthly to Rio de Janeiro, Santos, Buenos Aires.
To Brazil Only
United States and Brazil Steamship Line, Arthur Lewis, 39

Cortlandt St., fortnightly to Bahia, Rio de Janeiro, Santos.
Prince Line, 34 Whitehall St., monthly to Pará,
Pernambuco, Bahia.
Lamport and Holt Line, 42 Broadway, monthly to Pará,
Maranhão, Ceará, Natal, Cabedello, Pernambuco, Maceió,
Bahia.
Ward Line, foot of Wall St., monthly to Rio de Janeiro and
Santos.
Argentina and Uruguay
Barber Steamship Line, 17 Battery Place, fortnightly to

Montevideo, La Plata, Buenos Aires, Rosario.

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© Springer Science+Business Media, LLC, part of Springer Nature 2014, 2021

St Pierre, Réunion, France Pascale Besse

1 Plant Taxonomy: A Historical Perspective, Current Challenges,

13 Random Amplified Polymorphic DNA (RAPD)

ADBC Advancing digitization of biological collections

LARDs Large retrotransposon derivatives

TCG Taxonomic complex group

Plant Taxonomy: A Historical Perspective, Current

Adapting the famous aphorism of Theodosius Dobzhansky [1],

2 Taxonomy and Taxon: Terminology and Fluctuating Meanings

It is not before 1813 that the Swiss botanist Augustin Pyramus De

of botany along with glossology—‘the knowledge of the terms used

3 A Historical Perspective to Plant Taxonomy

These early classifications, that were exclusively utilitarian, per-

(77 AD) was indeed a rich encyclopedia of the natural world,

morphology, geographic distribution, ecology, or any other fea-

In contrast to Ray and his method intended to be natural, his

flower. Within his system, Tournefort precisely defined 698 enti-

although Dayrat [24] evidenced that all Haeckel’s illustrations

(PCR) was invented by Karry Mullis in 1986 [30]—have dramati-

and establishing species lists, but it nevertheless also provides basic

4 Plant Taxonomy Today: Current Challenges, Methods, and Perspectives

While the variety of conditions in South America makes any

Much repetition is avoided and probably greater convenience

On the North Coast, Cartagena has direct cable connection with

The Metric System of weights and measures is legal and official in

Including the branches and affiliations of American banks and

United States Banks

Branches and Affiliations

National City Bank, Branches: Argentina, Buenos Aires, Rosario;

Other Important Banks in South America

Other American Banks

with facilities for South American Trade are:

The North Coast

Colombia: Passenger and Freight Lines

New York to Puerto Colombia and Cartagena, Caribbean

Colombia: Freight Only

New York to Cartagena and Puerto Colombia, Tropical

Venezuela: Passengers and Freight

Venezuela: Freight Only

New York to La Guaira, Puerto Cabello, Maracaibo,

Guiana: British, Dutch, and French

British Guiana Passengers and Freight

New York to Georgetown, Quebec Steamship Company,