Journal of Biogeography (J. Biogeogr.

) (2006) 33, 761–769

GUEST The first biogeographical map

EDITORIAL
Malte C. Ebach1 and Daniel F. Goujet2

1
Laboratoire Informatique et Systématique
(LIS), Université Pierre et Marie Curie UMR
5143 Paléobiodiversité et Paléoenvironnements
Équipe Systématique, Recherche Informatique
et Structuration des Cladogrammes, 12 rue
Cuvier, 75005 Paris, France. E-mail:
ebach@ccr.jussieu.fr
2
Muséum National d’Histoire Naturelle,
Département Histoire de la Terre USM 203-
UMR 5143 Paléobiodiversité et
Paléoenvironnements Équipe Systématique,
Recherche Informatique et Structuration des
Cladogrammes, 8 rue Buffon, 75005 Paris,
France
ABSTRACT
Unbeknownst to many historians of biology, the first biogeographical map was
published in the third edition of the Flore française by Lamarck and Candolle in
1805, the same year in which Humboldt’s famous Essai sur la Geographie
appeared. Lamarck and Candolle’s map marks the beginning of a descriptive or
classificatory biogeography focusing on the study of biota rather than on the
distributional pathways of taxa. The map is relevant because it heralds the
beginning of the creation of biogeographical maps popularized by zoogeogra-
phers in the mid- to late nineteenth century together with the study of biogeo-
graphical regions.
Keywords
Biogeographical maps, biogeography, Candolle, Flore française, history of
biogeography, Lamarck.

theory of descent (Deszendenzstheorie) was the modus oper-

INTRODUCTION
The history of biogeography is poorly known and hitherto has
aroused little interest. Only a handful of detailed historical
accounts of biogeography exist that outline its general aims
(e.g. Nelson, 1978), its theory (e.g. Kinch, 1980; Richardson,
1981; Browne, 1983) and its early history (Hofsten, 1916;
Papavero et al., 1997, 2003; Heads, 2005). Nils Gustaf Erland
von Hofsten (1874–1956) presented the first in-depth histor-
ical account of distributional studies from Aristotle to Daniele
Rosa (Hofsten, 1916). The next seminal work by Nelson
(1978), details the roles of ecological and historical biogeog-
raphy and their history and influence on late twentieth century
comparative biology. The most recent work to rival von
Hofsten (1916) is that of Papavero et al. (1997) (Spanish
translation see Papavero et al., 2003), representing the most
comprehensive history of ‘pre-evolutionary’ biogeography to
date.
All of these works differ in their interpretation of biogeog-
raphy. Hofsten in his Zur älteren Geschichte des Disk-
ontinuitätsproblems in der Biogeographie considered the study
of distributional pathways on earth as the primary aim of
biogeography, in order to solve the problem of disjunct
distributions (Hofsten, 1916, pp. 197–198, a similar sentiment
echoed later by Platnick & Nelson, 1978, p. 1). Like other
evolutionists in the late nineteenth and early twentieth
centuries (e.g. Engler, 1879), Hofsten believed that Darwin’s
andi of distributional patterns. Hofsten went as far as declaring
that alternative evolutionary theories such as Rosa’s (1909)
Hologenesis were ‘‘opposed to the biogeographical study of
origins’’ (Hofsten, 1916, p. 338, our translation). More
significantly, however, Hofsten declared that ‘‘the biogeo-
graphical study of origins has, since the theory of descent, two
separate but related aims…’’ (Hofsten, 1916, p. 332, our
translation), namely to uncover Earth history and phylogenetic
change. Despite Hofsten’s aims, he disagreed with those who
believed in different theories of evolution, such as hologenesis.
In particular, he disagreed with the Austrian zoologist Ludwig
Karl Schmarda (1819–1908) and the English ornithologist
Philip Lutley Sclater (1829–1913), who rejected evolutionary
models in favour of multiple origins.
Sclater, however, had different aims in mind:
‘‘Organic beings are not scattered broadcast over the
earth’s surface without regularity or arrangement, as the
casual observer might suppose, nor are they distributed
according to the variations of climate or of any other
physical external agent, although the latter have, unques-
tionably, much influence in modifying their forms. But
each species (or assemblage of similar individuals),
whether of the animal or vegetable kingdom, is found
to occupy a certain definite and continuous geographical
area on the earth…It thus happens that the various parts

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Journal compilation ª 2006 Blackwell Publishing Ltd doi:10.1111/j.1365-2699.2006.01477.x
Guest Editorial
of the world are characterized by possessing special
groups of animals and vegetables, and that, as a general
rule, such tracts of land as are most nearly contiguous
have their Faunæ and Floræ most nearly resembling one
another; while, vice versâ, those that are farthest asunder
are inhabited by most different forms of animal and
vegetable life. When any similar forms, or two regions far
apart exhibit similar forms, it is the task of the student of
geographical distribution to give some reason why this
has come about, and so to make the ‘exception prove the
rule’ ’’ (Sclater, 1864, p. 213).
Sclater, too, saw that Earth’s history played an important
role in defining the geographical regions and modifying the
organisms that inhabited them. For Sclater, organisms origin-
ated where they are currently found and any discontinuity, that
is, disjunct distributions, were a result of earth processes.
Contrary to the ideas of evolutionists at the time, Sclater
believed that organisms did not disperse to favourable areas,
rather they changed over time (modifying their forms) in the
same area. The implication of disjunct distributions was,
therefore, that earth was changing too.
‘‘To conclude, therefore, granted the hypothesis of the
derivative origin of species, the anomalies of the
Mammal-fauna of Madagascar can best be explained
by supposing that, anterior to the existence of Africa in
its present shape, a large continent occupied parts of the
Atlantic and Indian Oceans stretching out towards (what
is now) America on the west, and to India and its islands
on the east; that this continent was broken up into
islands, of which some have become amalgamated with
the present continent of Africa, and some possibly with
what is now Asia – and that [in] Madagascar and the
Mascarene Islands we have existing relics of this great
continent, for which as the original focus of the ‘Strips
Lemurum’ I should propose the name Lemuria!’’
(Sclater, 1864, p. 219).
Surprisingly, exactly 100 years before Croizat (1964), Sclater
had presumed that life and earth evolved together. Whether this
makes Sclater a vicariance biogeographer, or in the case of his
super-continent Lemuria, a supporter of continental drift or an
expanding earth is not a matter for debate here. Although
Sclater (1864, p. 219) coined the term Lemuria he neither
defined it nor suggested that it sank into the Indian Ocean, as
Haeckel believed. Sclater disassociated the discontinuity prob-
lem from evolutionary or migration models. For him, distri-
bution was a result of changes in the area that the organisms
inhabited and not due to migration or dispersal.
The striking difference between Hofsten (explanatory bioge-
ography) and Sclater (classificatory biogeography) illustrates the
different aims of biogeography although the latter never
discusses or mentions Haeckel’s chorology, namely the ‘‘the
science of the geographic spread of organisms’’ (Haeckel, 1866,
p. 287, translation from Williams, in press), which is clearly what
Hofsten’s biogeography was all about. Nor was Haeckel’s
762
ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd
chorology mentioned by the eminent American zoogeographer
and mammalogist, Clinton Hart Merriam (1855–1942), who
considered the study of geographical differences, namely the
general change ‘‘that takes place in the fauna and flora in passing
from one region to another, or from low valleys or plains to high
mountains – geographic differences’’ (Merriam, 1892, p. 3), to
be the main focus of biogeography. Merriam understood
biogeography to be the study of life zones, areas that encompass
the distribution of organisms (Merriam, 1898). Life zones are
equivalent to Lamarck & Candolle’s (1805) floristic provinces
(see below), which are analogous to Candolle’s stations, namely
relating ‘‘…essentially to climate, to the terrain of a given
place…’’ (Candolle, 1820 in Nelson, 1978, p. 280) that is the
smaller division within habitations or regions:
‘‘In accordance with these facts naturalists long ago began to
divide the surface of the globe into zoological and botanical
regions irrespective of the long recognized geographic
and political divisions’’ (Merriam, 1892, pp. 3–4).
Biogeography, in effect has two histories, not an ecological
and historical history (sensu Nelson, 1978), but rather a
chorological history that uses evolutionary models such as
Buffon’s Law (Buffon, 1761) to trace distribution pathways,
and a systematic biogeographical history that describes, com-
pares, classifies biota and asks ‘‘…whether the continental
areas – the Neartic, the Neotropics, etc. – are real units, or
whether they are geological and biological conglomerates of
smaller and more local units with diverse histories’’ (Nelson,
1983, p. 490; see also Ebach & Morrone, 2005). The former is a
concept that dates as far back as Aristotle (see Aristotle Historia
animalium, VIII, pp. 156–165; also see Hofsten, 1916, p. 201,
footnote 1) and was later revised by Buffon (1761) as an
explanatory law of distribution (see Nelson, 1978, pp. 275–
276). The latter has its beginnings in taxonomy and classifi-
cation of French flora and floristic regions (Lamarck, 1778;
Lamarck & Candolle, 1805), although the first study was until
now considered by many to have been made by von Humboldt
(1805). Lamarck & Candolle (1805) also mark the initial stages
of the science of phytogeography, rather than the later work by
Swiss botanist Alphonse Louis Pierre Pyramus de Candolle
(1806–1893) (Candolle, 1855). Chorology and systematic
biogeography are equally historical and ecological. The divi-
sion between ecological and historical biogeography is artificial
and redundant. The beginning nineteenth century marks a
systematic way of approaching biogeography (phytogeography)
that was led by the Swiss botanist Augustin-Pyramus de
Candolle (1779–1841), father of Alphonse.
Candolle teamed up with fellow naturalist Jean Baptiste
Pierre Antoine de Monet, Chevalier de Lamarck (1744–1829)
to write the third edition of Flore française, already an
important work of late eighteenth century botany. Lamarck’s
dedication to and knowledge of Linnean taxonomy had
heralded his first two editions of the Flore française, the
seminal work on botanical taxonomy and classification. Next,
Lamarck turned to the invertebrates, a term he coined, and
Journal of Biogeography 33, 761–769

greatly advanced animal taxonomy by separating the crusta-
ceans, arachnids and annelids from the poorly described
‘Insecta’. He also provided the most detailed classification of
the molluscs. Unfortunately for Lamarck, his theory of
evolution received more attention (through controversy) than
his botanical and zoological classification and his development
of this, the first biogeographical map denoting floristic areas of
France.
Lamarck & Candolle (1805) applied a systematic method
through the use of biotic maps, rather than the usual
transverse maps that showed the ecological gradients along
the slopes of mountains (see below). The biotic map ‘‘…is
designed to highlight two very different things: 1. the
knowledge of vegetation in different parts of France that
are known by Botanists and; 2. the general plant distribution
on French soil…The second object of this map, namely, to
show the general distribution of plants in France, precedes
the first in its importance, or at least given our present state
of knowledge of French flora. The map should be considered
more of an attempt to apply a specific methodology rather
than an attempt to show the complete plant geography of
France’’ (our italics).
The use of a systematic method, namely to map the
distributions of biota based on their composition, marks a
move away from explanatory distributional models (e.g.
Buffon’s Law) toward a formal biotic classification.
The same systematic approach is not seen in zoogeography
until 50 years later in the works of English physician James
Cowles Prichard (1786–1848), Ludwig Karl Schmarda, Scottish
naturalist Andrew Murray (1812–1878), Philip Lutley Sclater
and his son, ornithologist William Lutley Sclater (1863–1944)
(Prichard, 1836; Schmarda, 1853; Murray, 1866, Sclater, 1858;
Sclater & Sclater, 1899). The application of a systematic
method in zoogeography led to a plethora of biotic maps, that
is, biogeographical maps (two terms that will be used
interchangeably).
BIOGEOGRAPHICAL MAPS
The development of biogeographical maps also reveals a
divergent history. Maps either showed the direction of
distributional pathways (e.g. Engler, 1879, first map; Haeckel,
1866) or they showed the biota (life zones) or regions (e.g.
Engler, 1879, second map; Murray, 1866; Merriam, 1892), the
latter being the more common in the late 19th and early 20th
centuries.
Merriam’s interest in distributional maps led to the first
biogeographic map that encompassed all life zones within a
region, in this case the life zones of North America. In order
to represent the biogeographical regions of North America,
mammals were ‘‘chiefly used as illustrations because they
answer the purpose better than any other single group…’’
(Merriam, 1892, p. 64). In outlining his ‘principles’ Merriam
coined the term biogeography with the heading ‘Principles on
which biogeographic regions should be established’. The
following colour map of North America also carried the title,
Journal of Biogeography 33, 761–769
ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd
Guest Editorial
‘Second Provisional Bio-geographic map of North America
showing the principal Life Areas’. Merriam rarely used the
term biogeography again, however, many have since used the
term to describe the study of chorology (Williams, in press).
Merriam never defined biogeography, although the term was
at first exclusively associated with the study of life zones and
biotic maps. Considering Merriam’s usage of the term,
biogeography would have originally been defined as the
classification of the life zones (biota) and endemic areas
within biogeographical regions, the result of which would be
a biogeographical map rather than a distributional pathway,
as proposed by Haeckel (1870). If we accept the above
definition, then who produced the first biogeographical
map?
Before Merriam’s biogeographical maps, floristic and fau-
nistic maps were strictly designated as phytogeographical or
zoogeographical, respectively. Merriam (1892) compiled a list
of all biogeographical studies that either did or did not include
maps, starting with Latreille (1817). The first biogeographical
maps describing the world regions were the most popular, as
they delimited vast areas defined by climate and larger
geographical features such as oceans or mountain ranges (see
Murray, 1866). Smaller scale maps that outlined the faunistic
or floristic areas within a region were rare. Most maps dealt
with the distributions of single taxa.
Mennema (1985) proposed that the Danish botanist
Joachim Frederik Schouw (1789–1852) was the first to draw
a modern distributional map in his Grundzüge einer
Allgemeinen Pflanzengeographie (Schouw, 1823). According
to Mennema (1985), Schouw’s map provides a ‘birds-eye’
view rather than the earlier transverse views provided by
Wahlenberg (1812) and Humboldt & Bonpland (1807).
Given the Dane’s ingenuity, Mennema proclaimed that
Schouw ‘‘should be considered the father of plant geography’’
(Mennema, 1985, p. 117). The reason given by Mennema was
that Schouw’s map appeared earlier than Candolle’s (1855)
map, which according to Stearn (1951) was the oldest known
biogeographical map. Surprisingly, however, the French
Abbott Jean-Louis Giraud-Soulavie (1752–1813) drew the
first botanical map with a transverse perspective in 1784,
which is 23 years before the map illustrated by von
Humboldt & Bonpland (1807). The map entitled Coupe
verticale des montagnes vivaroises. Limites respectives des
climats des plantes et mesures barométriques de leur hauteur
sur le niveau de la Méditerranée (Giraud-Soulavie, 1770–1784)
shows the succession of flora delimited by latitudinal climatic
gradients. The maps pioneered by Giraud-Soulavie and later
drawn by Humboldt & Bonpland and Wahlenberg are
ecological transverse sketches that detail a particular region
or biome in detail, rather than showing the range and variety
of biotic regions as in later biogeographical maps (e.g.
Merriam, 1892).
In the third edition of Flore française by Lamarck & Candolle
(1805), there is a biogeographical map outlining the floristic
provinces in France (Fig. 1; a translation of the introduction to
the biogeographical map within the second volume of the third
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Figure 1 Carte Botanique de France, pour la 3ème Edition de la Flore française par A.G. Dezauche fils Ingénieur Hydrogéologue de la Marine an
13 (1805) ‘Botanical map of France for the 3rd Edition of Flore française by A.G. Dezauche the son, Marine Hydrological Engineer on the
13th year of the Revolution (1805)’ (our translation). Water colour on machine print. Note that Corsica is coloured crimson, which is
characteristic of the 1815 hand painted version.

edition of Flore française is given in Appendix 1). Although the
existence of such a map has been noted (Nelson, 1978, p. 269,
footnote 1; Drouin, 1997, p. 249; see Appendix 2) its relevance
to phytogeography was understated as it is rarely ever
mentioned in the botanical or biogeographical literature, with
a few exceptions like Drouin (1997) who notes that the map
appeared in the same year that Humboldt presented his Essai
sur la geographie des plantes (Drouin, 1997, p. 249). Perhaps
Lamarck and Candolle’s map was over-shadowed by Hum-
boldt’s monumental work (Humboldt, 1805)?
Not only were Lamarck & Candolle (1805) the first to draw
a biogeographical map depicting floristic provinces, they also
formulated a systematic method to classify biota (namely, to
map floristic provinces), emphasizing the classificatory nature
of biogeography rather than the study of distributional
pathways. Whether this makes Lamarck and Candolle fathers
or founders of the field of phytogeography or biogeography is
a matter of speculation. This thinking in terms of floristic
regions and endemic areas may, however, mark the beginnings
of studying biotic regions and biogeographical classification,
central concepts that Candolle was to develop 15 years later
(Candolle, 1820).
ACKNOWLEDGEMENTS
We are grateful to M. Didier Geffard-Kuriyama for scanning
Lamarck and Candolle’s map, to Michael Heads and Juan J.
Morrone for their comments and suggestions, and to
Lynne R. Parenti for providing access to the Smithsonian
(Washington DC) copy of the Flore française. We also
thank René Zaragüeta i Bagils, David M. Williams, Gareth
Nelson, Dennis McCarthy, Antoine Chalubert, Guillaume
Dubus and Caitlin E. Hulcup for reading through earlier
drafts.

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BIOSKETCHES
Malte C. Ebach is a researcher and author in the history and
philosophy of comparative biology, Goethe’s way of science,
trilobite taxonomy and systematics at the Laboratoire Informa-
tique et Systématique, Université Pierre et Marie Curie, Paris.
Daniel F. Goujet is professor of palaeontology at the
Muséum National d’Histoire Naturelle, Paris. His research
interests include Palaeozoic early vertebrates, mainly Placo-
derm anatomy and phylogeny, the history and philosophy of
science and cladistics in all its aspects, and including
palaeobiogeography.
Editor: John Lambshead
APPENDIX 1 ‘EXPLANATION OF THE
BOTANICAL MAP OF FRANCE’ FROM FLORE
F R A N Ç A I S E ( 3 R D E D N , 2 N D V O L . ) B Y L A M A R C K
& CANDOLLE (1805), PP. I–XII). A
TRANSLATION OF THE INTRODUCTION
The botanical map of France, which we thought would be a
useful addition to this book, is designed to highlight two very
different things: (1) the knowledge of vegetation in different
parts of France that are known by Botanists; and (2) the
general plant distribution on French soil.
The main objectives of the map are to help practising field
botanists easily orient their research on those points which
have been insufficiently visited, that is, on parts of the flora
that have not been described in printed works. We will
therefore know which parts of French flora have been studied
in detail.
To give an idea of the different provinces of this vast territory
which have been explored by Naturalists, the map will show only
the names of towns and villages where plants have been collected.
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On the map the unknown Western provinces, for example,
appear nearly entirely void, thereby calling attention to the need
for further observation and study. Areas on the map crowded
with names, however, like the surrounding parts of Paris,
Montpellier and Turin [Italy], clearly show that nearly all the
plant content of these areas has been observed.
In order to highlight our current knowledge of the French
flora, we have noted in capital letters the towns where several
distinguished botanists have lived and collected (Paris, Mont-
pellier, Turin). Towns labelled in lower case letters indicate the
towns and surrounding areas that have a well-known Flora, or
several fragmentary ones (e.g. Grenoble, Barrèges and Geneva
[Geneva at the time being part of France]). Towns labelled with
roman letters indicate areas known from incomplete catalogues
(e.g. Rouen and Soissons). Finally, towns labelled in italics
indicate floras of neighbouring countries or areas containing
some seedless plants. Given these guidelines, one can, with a
simple map, investigate and determine with precision whether
floras of one or another country are sufficiently known. These
guidelines, quite different from those of ordinary maps, explain
why the names of some villages are written in capitals and those
of towns are in smaller fonts, as well as why there is a large
discrepancy between the numbers of villages mentioned in
different provinces. Due to a lack of space on the map and from a
botanical point of view, it has been necessary to exclude several
well-known countries and areas.
The second object of this map, namely to show the general
distribution of plants in France, precedes the first in its
importance, or at least given our present state of knowledge of
French flora. The map should be considered more of an
attempt to apply a specific methodology rather than an
attempt to show the complete plant geography of France.
On this map, France is divided into five regions distin-
guished by different colours; one should note that these areas
are not etched into nature exactly as they appear here. It would
be difficult to represent their exact delimitation therefore these
regions have to be considered only as very generalized
indicators.
The area coloured green, marking the coasts from Ostende
[Belgium] to Oneille, indicates the realm of maritime [aquatic]
plants. Other areas marked in green within France, such as the
saline lakes of Dieuze, Château-Salins, Salins, Durkheim and
Frankenstatt, indicate that the same aquatic plants occur in
places with a sufficient amount of marine salt.
All aquatic plants from northern France are also found on
southern coasts, but the reverse does not occur, and most
maritime plants of the Mediterranean realm only grow in small
quantities on the ocean border around Gascogne [Bay of
Biscay] and develop northwards only around the Loire estuary,
or, at the very most until the south of Brittany. Despite this
difference, I [A.P. de Candolle, see below] did not think it
necessary to split into two classes the maritime plants, because
of the extreme similarity we observe in their bearing and their
vegetation.
The blue colour represents areas in France that are occupied
by mountain plants. These demarcation lines are less pro-
Journal of Biogeography 33, 761–769

nounced than in the preceding region. Valleys exposed to the
sun share the same vegetation of southern provinces, whereas
cooler valleys contain plants that are shared with the vast
northern and central plains of France. These regions, however,
offer a very high number of specific plants, the majority of
which are present on all mountains. Despite any botanical
differences that may exist between the mountain chains of
Vosges, Jura, Alps, Auvergne, Cévennes, Pyrénées, one cannot
dismiss the fact that aspects of their vegetation offer high traits
of similarity and that most mountain plants are found in these
different chains.
The area coloured crimson red which tints the island of
Corsica and the southern parts of France is meant to represent
the space occupied by the class of plants that I will term
Mediterranean plants, since they are found in nearly all
countries that encircle the Mediterranean sea. One may note
that this area occupies the southern side of our great mountain
chains and the space between the sea and the base of
mountains, spreading a little to the north around Montélimart
and in the Rhône valley because in the high terrain in this part
of France there is a higher temperature than in other cities
located at the same latitude.
The vast area in yellow, which covers more than half of
France and notably all the plains situated north of the
mountain chains, indicates the uniformity in vegetation of
these large plains. This region contains plants that are similar
and present in most of the other regions, but lack endemics.
The area in vermilion provides us with knowledge of French
provinces where the vegetation is intermediate between those
of northern plains and those of southern provinces. By a
simple inspection of this map one may see that meridional
[southern] plants appear further to the north on the western
side than on the eastern side. So, if the floras from Le Mans or
Nantes1 are studied, it shows they differ only slightly from
those of Dax and Agen, which are located three of four degrees
south of them. On the eastern side, however, floras from Dijon
and Strasbourg differ totally from those of Aix and Turin
although they are located at quite similar distances.
This fact would appear singular if it were not compared to
another observed by Mr Arthur Young. This esteemed
traveller, who has attentively studied cultivated plants,
remarked that if lines are traced between the northern points
where olive trees, grapes and maize are grown, three nearly
parallel lines are obtained that all tend to meet to the north on
the eastern side. This is exactly the reverse to what we observe
for wild plants. We have copied the three lines of Mr Arthur
Young, to serve as points of comparison with our divisions.
Why there is such an apparent contradiction may be
explained by the comparison of the physical nature of Eastern
1 northern of coastal provinces also belong to this class, including
For the plants of Nantes, I refer to the publication of Mr Bonamy. I
learn at the time of publication of this note, that this botanist, without
informing others, has naturalized several exotic plants around Nantes:
thus this part of the French flora will have to perhaps undergo some
revision.
Journal of Biogeography 33, 761–769
ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd
Guest Editorial
and Western France and in the choice of cultivated plants
relative to wild plants.
Of all the factors that influence the habitat [l’habita-
tion ¼ ‘physical environment’ or ‘regions’, see Nelson, 1978,
pp. 280–281] of plants, temperature is without doubt the most
essential. The latitude and the height above sea level may
determine the average temperature of one site, independent of
local conditions. One estimates that in general, areas 200 m
above sea level have, on average, temperatures that are more or
less equivalent to one degree of latitude to the North. The
reader may compare this ratio for the different parts of France
using the lines that we have traced on the map, which indicate
the general altitude of the different provinces relative to sea
level. Mr Dupaintriel’s ingenious idea of applying such a
method to continents, which indicates continental height, is
the same process that is used on marine charts to indicate
depths. We copied the heights of French plains and mountains
from Mr Dupaintriel’s map of France and from the recorded
observations of geologists. The heights of the Alps are extracted
from the Voyages de De Saussure (Saussure, 1779–1796),
Mr. Ramond communicated those for the Pyrenees and
Mr. Léopold de Buch observed those for the Jura.
If we compare the western and eastern provinces, we see that
the first are not very high above sea level because, at greater
distances from the coast, the altitude is still only 100 m. On the
contrary, the eastern provinces that encircle the mountain
chains are generally at four to 500 m above sea level. It is true
that this height diminishes at the Belgian side but then
temperature changes based on a second factor, namely the
distance from equator. So there is nothing more than
the conformity to laws of physics to explain why plants of the
south appear more northerly on the west than on the east sides.
Even if the average temperature is the same, the distribution
of plants between these two parts of France should be different
because of the different way the same temperature is distri-
buted among the seasons of the year. It is a generally accepted
fact that, on the same latitude, islands and coastal areas have
more equal temperatures than countries further from the sea.
In other words, they have fewer warmer summers and colder
winters. The uniform temperature of coastal lands is due to
mild winds and to the proximity to large bodies that have little
variations in temperature. Therefore, the western provinces of
France, which all are coastal, benefit by this sort of uniformity
that does not affect the eastern provinces that are far away
from the seas and closer to the mountains.
Plants are divided into two classes: those that dislike the harsh
winter cold but do not need the summer heat and those that can
endure cold winters but require hot summers. Included with the
first class are trees (including those that are not conifers) that
retain their leaves and consequently their sap during wintertime.
Most indigenous or acclimatized southern trees that occur in the
the holm oak, the cork oak, the kermes oak, the arbutus, the bay
tree, the fig tree, the philaria and the large flower periwinkle. On
the other hand, the second class consists of plants that resist the
harsh winter cold by stopping sap flow and shedding their leaves
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Guest Editorial
(e.g. like the vine, etc.), and those that are able to escape from
such cold conditions (e.g. maize, etc.). Plants of this second class
easily acclimatize to the eastern rather than the western parts of
France.
As for cultivated plants, one should add a last observation
that is to say that those cultivated to bear fruit are restricted to
regions with hot summers. Vines for example, are grown for
profit on the southern slopes of the Alps and in places where
the average temperature is colder than in Brittany or
Normandy, but where summers are very hot and where the
grapes will surely ripen. This same shrub is not grown in
northern France, not because it will die, but because its fruit
will not ripen since summers are not warm enough. On the
contrary, plants that are grown for their fruit, even indigenous
varieties from the most southern countries, are easily cultiva-
ted all over France – as are the artichoke, lavender, the nettle
tree, etc. I need not develop these observations further as they
appear sufficient to explain why, in France, plants from the
south get nearer to the north on the west side than on the east
and why several cultivated plants do the reverse.
I place great importance on altitude as a major factor
influencing temperature. I also believe that temperature and not
air density greatly influences vegetation, a factor that well known
scientists support. How can you reconcile the influence of air
density with well known facts? In mountains where the soil
allows a vegetation to grow, plants are found at all attitudes up to
the permanent snow caps. Plants found in the high Alps are
found in northern Europe and in places where the air is denser.
But where the temperature is equal to that of these mountains,
these alpine plants can, with certain precautions, be cultivated on
the lowest plains. Even some of those that grow in the high Alps
are found on the coast, and in the same mountains the same
plants grow higher up on the southern slopes than on the
northern ones. In temperate zones where the altitude does not
alone determine the temperature, many anomalies are observed
relative to the altitude to which these plants are found, although
very few are noticed in countries close to the equator where the
altitude alone determines the temperature. Given these facts, I
believe that the temperature level, which varies according to
mountain height, influences vegetation.
In some texts, importance is placed on the chemical nature
of soil in which plants grow and, perhaps on further
consideration, I should have included it on the botanical
map. I will however, point out that all the general facts prove
otherwise. I do not deny that the nature of the compost, and
even sometimes the nature of earth, influences the strength and
properties of plants. What I think can be affirmed, is that this
influence is too insignificant to determine the general region
[habitation] of plants, so that plants which flourish on certain
soils can just as well propagate on a different ones, especially
those that occur nearby. As an example, I will take two of the
most characteristic terrains on which the diversity of plants
have been most clearly delineated – granitic and calcareous
terrains – and, like in the preceding example, I will use
evidence rather than details. In France we have two important
mountain chains that break this assumption: the Vosges are
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granitic, the Jura is calcareous and we hardly find plants that
are not common to both chains. The Jura contains a high
number of plants that also grow in the granitic Alps. The Alps,
when compared to the high Pyrenees summits, again show that
both terrains share numerous plants. Furthermore, if we
exclude the very rare plants, we would not find a single
example that is present only in calcareous or granitic terrains.
From the preceding considerations, I believe that in a given
country, such as France, the causes that determine the plant
region [habitation] could be reduced to three:
1. Temperature, as determined by distance from the equator,
height above sea level and southern or northerly exposure.
2. The mode of watering, which is more or less the quantity of
water that reaches the plant. The manner by which water is filtered
through the soil and the matter that is dissolved in the water
which may or may not be harmful to the growth of the plant.
3. The degree of soil tenacity or mobility.
APPENDIX 2 THE HISTORY OF FLORE
F R A N Ç A I S E
The ‘Flore française, ou descriptions succinctes de toutes les
plantes qui croissent naturellement en France, disposées selon
une nouvelle méthode d’analyse, et précédées par un exposé des
principes élémentaires de la botanique, troisième édition’ was
the only volume in the series of three editions spanning over half
a century that contained a colour map, ‘ouvrage accompagné
d’une grande carte botanique coloriée…’. The Flore française
survived the French revolution. The first edition published by
the Royal Printers (de l’Imprimerie Royale) in 1778 as three
volumes with Lamarck as the sole author. The second edition,
printed by H. Agasse in 1795 (l’an 3e. de la République), was ‘‘an
almost word by word reprint’’ of the first edition (Stafleu &
Cowan, 1976–1988, p. 732). The third edition grew by two
volumes and was ‘‘almost entirely rewritten by Candolle’’ in
1805 (Stafleu & Cowan, 1976–1988, p. 442).
The original third edition of Flore française of 1805 was
reissued as five volumes in 1815 with a new title page
‘‘…troisième edition, augmentée du tome V, ou sixième
volume, contenant 13000 espèces non décrites dans les cinq
premiers volumes’’ and again in 1829, ‘‘but printed by
Huzard-Courcier ‘imprimeur depuis 1820’ ’’ (Stafleu & Cow-
an, 1976–1988, p. 442). The coloured plate, appearing in the
second volume and designed for A.G. Dezauche, varies with
each reissue of the 3rd edition. The map consists of an original
print that is hand painted with water colours, usually in blue,
red, crimson, orange, green and aquamarine. In the 1805 print
the colours are bright and bold, however in the 1815 prints the
colours are washed out pastels. The colours of the south-west
region of France differ in both prints, being crimson in the
1805 edition (as noted by Lamarck and Candolle, above) and a
light pink in the 1815 version. One major difference between
them is that Corsica is coloured (pink) in the later 1815 reissue
and not coloured at all in the original 1805 print, although
designated as so in the introductory text. Another difference
between both reissues is that the 1815 edition makes note that
Journal of Biogeography 33, 761–769
 Guest Editorial

the 3rd edition contains a map and that the map is located in
the front of the volume. The maps were produced independ-
ently of the text and were inserted afterwards by the binder –
also a limited number of maps were produced, resulting in the
absence of the map in many existing copies of the original 1805
edition. The second reprinting of 1815 has the map placed in
different parts of the book, either before the introduction, after
the introduction or in the back (copies viewed: 1805 editions
Muséum National d’Histoire Naturelle, Paris; 1815 editions
Muséum National d’Histoire Naturelle, Paris; Smithsonian
Institution Washington DC; Private Collection of D. Goujet;
Private Collection of Anon. No 1829 editions were sighted).

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