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On the concept of chorotype
Simone Fattorini
APPENDIX S1 Excerpts from works by Hofmann, Hooker & Thomson, Forbes,
Areschoug, Christ and Watson.
Text S1 Excerpt illustrating Hofmann’s division of the European butterflies into groups
with similar ranges (Hofmann, 1873, p. 14)
Dieselben vertheilen sich in folgende geographische Gruppen:
1. Sibirisch-europäische Arten ......................................................................................... 173
2. a) Europäisch-asiatisch Arten .........................................................................................39
b) Europäisch-asiatisch-afrikanische Arten..............................................................12
3. Europäisch-afrikanische Arten......................................................................................... 8
4. Europäische Arten allein ...................................................................................................21
5. Alpine Arten ...........................................................................................................................23
6. Hochnordische Arten............................................................................................................ 5
Summa 281 Arten
Translation
The same species can be divided into the following geographical groups:
1. Siberian–European species ........................................................................................... 173
2. a) European–Asian species ..............................................................................................39
b) European–Asian–African species.............................................................................12
3. European–African species .................................................................................................. 8
4. Strictly European species .................................................................................................21
5. Alpine species ........................................................................................................................23
6. High Nordic species ............................................................................................................... 5
Total 281 species
Text S2 Excerpts illustrating the analysis by Hooker & Thomson (1855, pp. 103–113) of
the Indian flora according to its similarity with the flora of other regions
The following geographical alliances or affinities (if we may use the terms) of the Indian
Flora, with more or less remote countries, we consider well established; they are capable
of much illustration, even in the present state of our knowledge, but it is obviously
impossible to dilate upon them here.
1. The Australian type.—The Flora of Australia is well known to contain far more
endemic species and families than any other country does, and of these a few representatives extend into India. […]
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2. The Malayan Archipelago type.—This forms the bulk of the Flora of the perennially
humid regions of India; as of the whole Malayan peninsula, the upper Assam valley, the
Khasia mountains, the forests of the base of the Himalaya from the Bramaputra to Nipal, of
the Malabar coast, and of Ceylon. It is of course impossible to specify the genera or even
families of so predominant an element; to do so would be to enumerate a very large
proportion of the Indian genera, and to except only the north temperate and the comparatively few African types. The extent, however, to which this element predominates is not
yet appreciated, nor do we ourselves know its total amount; for constantly, during our
examination of the temperate as well as tropical plants of the Nilghiri, Khasia, Ceylon, and
the Himalaya, we find them identical in species with Javanese mountain plants. […]
3. The China and Japan type.—In the Indian flora we meet with many temperate
genera and species, which are also common to North America west of the Rocky Mountains, and which are foreign to Europe, to America east of that range, and to Western
Siberia; besides many tropical species that are also Malayan and West Polynesian. The
Chinese type is abundant in the temperate regions of the Himalaya, extending westward to
Garhwal and Kumaon, but is most fully developed in Sikkim, Bhotan, and the Khasia. […]
4. The Siberian type.—This is characteristic of the colder temperate parts of Asia, and
is very fully represented in the upper temperate and alpine regions of the Himalaya,
descending in the north-western and drier parts of the chain to very low levels. It
approaches, in many respects, to the South European vegetation, but is characterized by
the predominance of Fumariaceæ, Potentillæ, Leguminosæ, especially Hedysarum and
Astragaleæ, of Umbelliferæ, Lonicera, Artemisia, Pedicularis, and Boragineæ; and by the
rarity or total absence of certain groups or genera which are especially abundant in
Europe, such as Cistaceæ, Rosa, Rubus, Trifolium, Erica, Ferns, and other cryptogams. […]
5. The European type.—The extent to which European plants abound in India has
never hitherto been even approximately appreciated. Dr. Royle was the first to indicate
this affinity between the vegetation of the eastern and western continents of the old
world; and throughout his writings we find constant evidence of his never having lost
sight of this being a marked feature. […]
6. The Egyptian type.—Egypt, Southern Arabia, and the warmer parts of Persia,
possess a remarkable similarity of climate to Beluchistan, Sind, and the Panjab, and at the
same. time a nearly complete identity of vegetation. Many North African or Arabian forms,
such as Peganum Harmala, Fagonia Cretica, Balanites Ægyptiaca, Acacia Arabica, Alhagi,
Grangea, Calotropis, Salvadora Persica, extend throughout all the drier parts of India.
Others have a less extensive range, being only found in Northern and Western India […].
7. The Tropical African type.—Though tropical Asia and Africa are separated by a
vast expanse of ocean, there is a striking similarity in their vegetation. This is shown not
only by the identity of the annual vegetation which springs up during the rainy season, but
by a great similarity in the families and genera of the trees and shrubs […].
Too little is known of the African Flora to enable any definite conclusions to be
drawn as to the numerical value of this type in India, but it is evidently an important one.
Text S3 Excerpts illustrating Forbes’ classification of the British flora into five groups of
species on the basis of their distribution and immigration time into the British Isles
(Forbes, 1846, pp. 338–347)
The vegetation of the British Isles presents a union of five well-marked floras, four of
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which are restricted to definite provinces, whilst the fifth, besides exclusively claiming a
great part of the area, overspreads and commingles with all the others.
I. Commencing with the smallest, we find the mountainous districts of the west and
south-west of Ireland characterized by botanical peculiarities which depend on the presence of a few prolific species belonging to the families Saxifrageæ, Ericaceæ, Lentibulariæ,
and Cruciferæ. The nearest points of Europe where these plants are native is the north of
Spain. […]
II. In the south-west of England and south-east of Ireland we find a flora which
includes a number of species not elsewhere seen in the British Isles, and which is intimately related to that of the Channel Isles, and the neighbouring part of France. [...] This is
the Atlantic type in Mr. Watson’s arrangement of British types of vegetation. […]
III. In the south-east of England, where the rocks of the cretaceous system are chiefly
developed, we find the vegetation distinguished by the presence of a number of species
common to this district and the opposite coast of France. Here are the localities of the
well-known chalk plants, much sought after by botanists from the north. They form part of
Mr. Watson’s second, or Germanic, and of his third, or English, type of British vegetation.
IV. The summits of our British Alps have always yielded to the botanist a rich
harvest of plants which he could not meet with elsewhere among these islands. The
species of these mountain plants are most numerous on the Scotch mountains […]
V. The fifth and general flora of the British Islands—everywhere present, alone or in
company with the others—is identical as to species with the Flora of Central and Western
Europe—that which may be properly styled Germanic. […]
During the post-pliocene epoch, over the elevated bed of the glacial sea, the great
mass of the flora and fauna of the British Isles migrated from the Germanic regions of the
continent. The whole of the flora I have numbered V., including the great mass of British
plants, is Germanic. […]
We have seen that the great Germanic and central British plains themselves were
portions of the elevated bed of a pre-existing sea, which sea […] was the sea of the Glacial
period […].
Now it was during this epoch (the epoch of my IVth flora), that Scotland and Wales,
and part of Ireland, then groups of islands in this ice-bound sea, received their alpine flora
and a small portion of their fauna. […]
As a general rule, we may regard the most southern floras to be oldest, especially
when, as in these cases, they are more and more fragmentary, and their character is more
and more southern.
That which I have numbered III. is the most extensive, and from the number of
species which are exclusively or chiefly found in chalk districts in this country, I have
called it the Kentish flora. […] Still more ancient appears to have been the flora numbered
II., the peculiarities of which are seen more especially in Cornwall and Devon, and in the
south-east of Ireland. This flora—a relic of a larger—is undoubtedly a part of that which
we find in the Channel Isles, and in the adjacent provinces of France. It is still more
southern in character than No. III., exhibiting the features of the transition between the
great flora of central Europe and that of the southern or Mediterranean region. […]
Whatever doubts may be entertained respecting the antiquity of the Kentish and
Devon floras, there can be none (if my premises be granted) respecting that which I have
numbered I., and from which the peculiar botanical character of the south-west and west
of Ireland is derived.
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Text S4 Excerpts illustrating Areschoug’s groups of Scandinavian plants based on their
assumed immigration routes. Note the use of the word ‘element’ (Areschoug, 1867, p.
88).
Såsom resultat af mina här gjorda undersökningar tror jag mig kunna antaga, att i Skandinaviens närvarande vegetation ingå åtminstone trenne, till tid och ursprung skilda
elementer, nämligen:
1) En arktisk vegetation, som under istidens sednare period invandrat österifrån och
som […]. Detta element skulle man kunna benämna den Nordsibiriska Floran.
2) En nordöstlig och östlig vegetation, som efter istiden och före bokens invandring
inkommit till Europa från Sibirien. […] Denna vegetation blir sålunda i södra Europa mer
eller mindre alpin. Man skulle kunna benämna detta element Altai-Floran.
3) En sydöstlig och sydlig vegetation, hvars invandring varit samtidig med bokens
och som kommit dels från de Kaukasiska och de kring Kaspiska samt Svarta hafvet belägna
länderna, dels äfven från Medelhafsregionen. […] Denna vegetation kan kallas den
Kaukasiska och Medelhafs Floran.
Translation
As a result of my studies reported here, I think I can assume that the current vegetation in
Scandinavia includes at least three elements that differ in time and origin, namely:
1) An Arctic vegetation, which during the latter part of the ice age immigrated from
the east […]. This element may be termed the North Siberian flora.
2) A north-eastern and eastern vegetation that came into Europe from Siberia after
the ice age, but before the immigration of the beech. […] This vegetation is thus in southern Europe more or less alpine. One could call this element the Altai flora.
3) A south-eastern and southern vegetation, which came simultaneously with the
beech, partly from the Caucasus and the countries around the Caspian and Black seas,
partly from the countries of the Mediterranean. […] This vegetation may be called the
Caucasian and Mediterranean flora.
Text S5 Excerpts illustrating Christ’s analysis of the Alpine plants on the basis of their
overall distribution (Christ, 1867, pp. 10–26).
Unsre Tabelle ergiebt 693 Species, die sich nach ihrem Verbreitungsbezirk in zwei Hauptgruppen trennen: in solche, welche die mitteleuropäischen Gebirgsmassen und deren Ausläufer: von den Pyrenäen bis zum Caucasus ausschliesslich bewohnen, und in solche, die
neben diesem, im weitesten Sinn alpinen Gebiet, einen zweiten, nordischen: Scandinavien,
Nordasien und Nordamerika umfassenden Verbreitungsbezirk haben.
A. Nordischer Arten zählen wir, wenn wir sämmtliche 10 Colonnen von Island bis
zum Ural mitrechnen, 271, also fast ⅖ unserer Gesammtzahl (697). […]
Bleiben wir jedoch – bei der Möglichkeit abweichender Ansichten über die Heimat
dieser Arten – bei den 422 im Norden absolut fehlenden Species stehen.
Diese Arten bilden die alpine Gruppe; sie haben ihre Heimat, ihren Entstehungsheerd unzweifelhaft in den Gebirgen, welche im Norden des Mittelmeers sich hinziehen,
vor Allem in der grossen Alpenaxe.
[…]
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B. Mediterrane Arten. Neben diesen Alpen im weitern Sinne kann man nun aber
nicht umhin, ein anderes Gebiet als Heimat einer gewissen Quote unserer Liste zu
betrachten. Es ist dies die Mittelmeerregion. […]
Ziehen wir diese Arten – deren genaue Zahl nicht zu ermitteln ist – ab von der
Gesammtzahl der 422 Species, so bleibt die echte Alpenflora übrig, deren Ursprungsort die
grosse Alpenaxe mit ihren Zweigen ist.
C. Alpine Arten. Wie innerhalb der nordischen Gruppe, so zeigt sich auch hier – auf
beschränkterm Feld allerdings – die allergrösste Mannigfaltigkeit und Eigenartigkeit: es
sind kaum 2 Arten, deren Verbreitungsbezirk, deren Centrum, genau gleich ist. Jede
einzelne Art hat also ihre Specialgeschichte und ihre Eigenthümlichkeit gegenüber den
physischen Einflüssen.
Translation
Our table includes 693 species, which can be separated into two main groups according to
their area of distribution: those that occur only in the mountain ranges of Central Europe
and their spurs, from the Pyrenees to the Caucasus; and those which, in addition to this
Alpine distribution in a broad sense, also occur in a second, nordic, distribution district,
which includes Scandinavia, northern Asia and North America.
A. The Nordic species are 271, i.e. about ⅖ of the total number (697), if we count the
10 columns of our table from Iceland to the Urals.
[…]
Regardless of the different points of view about the area of origin of these species,
there is a set of about 422 species that are completely absent from the Nordic group; these
form the Alpine group and have their area of origin undoubtedly in the mountain ranges
that stretch to the north of the Mediterranean, especially on the Alpine arc.
[…]
B. Mediterranean species. In addition to the species of the Alps in a wider sense, we
cannot forget another area as the place of origin of a certain proportion of the species in
our list. This is the Mediterranean region. […]
If, from the 422 species [belonging to the Nordic group], we omit these species,
whose number is not known with certainty, we obtain the true Alpine flora, which has its
area of origin in the great Alpine arc and its branches.
C. Alpine species. As in the case of the Nordic group, we can see in this group a great
variety and oddity: we can hardly find two species whose area of distribution, their centre
of origin, is exactly the same. Each species has its special history and its peculiar
characteristics in relation to the physical influences.
Text S6 Excerpts illustrating Watson’s groups of British plants based on their
distribution within the British Isles (Watson, 1847, pp. 43–55).
In addition to their distribution by provinces and climatic zones, there is a third mode of
indicating the geographical relations of plants, which may also require some explanation.
It has been before observed that certain species are spread over the whole island, while
others are limited to one, two, three or more of the provinces. The same holds true in the
zones; some species occurring in all of them, others in one or more. Perhaps no two
species have exactly the same distribution or relative frequency; and yet certain general
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similarities may be traced, by which the species may be grouped together under a few
leading Types of distribution. In the small volume before alluded to, by the title of
‘Remarks,’ certain “geographic types” were indicated (pp. 86–89); and in the ‘Tabular
Appendix’ to the same volume (115–184) the species were severally assigned to their
peculiar types. This was simply an attempt to express, by a single term, the leading
character of their distribution, with reference to geographical position and climate. Six
types of distribution were particularly mentioned; under one or other of which, it was
thought, nearly all the species of plants indigenous in Britain might respectively be
arranged. No attempt, however, was made to define the precise limits of the types geographically. Nor, indeed, could any exact boundary lines be traced on a map, without
abruptly cutting asunder the fine gradations of Nature; for the types pass into each other
without any hard or abrupt lines of distinction. In slightly describing the several types, in
the former volume, a different order of succession was adopted, and consequently the nos.
affixed to them were different also; but in other respects they were essentially the same as
the following:–
1. The British Type. – In this group will be included those species which are found in
all, or nearly all, of the eighteen provinces before explained; and which, moreover, are not
so exclusively prevalent or predominant in any particular portion of the island, as to bring
them clearly within one or other of the following types. Some of the species may be
regarded as of universal occurrence in this country, growing in all the eighteen provinces,
probably in every county, and even in all the six ascending zones of vegetation or climate
also. Few species, however, even of this most general type, are so very general in their
distribution. […]
2. The English Type. – The plants of this geographic type are distinguished from
those of the British type by having their chief prevalence in England, and particularly in its
more southern provinces; whence they gradually become rare in a northern direction, and
finally (with few peculiar exceptions) find an earlier northern limit or cessation than those
of the preceding type. […]
3. The Scottish Type. – This may be deemed the opposite of the English type; the
distribution of the species referred hereto being characterized by a northern tendency,
either by absolute limitation to Scotland or the north of England, or otherwise by a chief
prevalence there and increased rarity southward. […]
4. Highland Type. – This may be considered the boreal flora in a more intense degree,
as respects climate, than that of the Scottish type. The species referred hereto are distinguished from those of the Scottish type by being more especially limited to the mountains
or their immediate vicinity. […]
5. The Germanic Type. – The distribution of several species which might otherwise
be associated with those of the English type, is peculiarly characterized by a tendency to
the eastern side of the island. […]
6. The Atlantic Type. – Contrary to the peculiarity of distribution which constitutes
the Germanic type, there is in that of other species a marked tendency towards the western and south-western coasts or counties. […]
7. A Local or doubtful Type. – Interspersed about the island, there are some species
whose distribution is restricted to single or few counties. […]
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REFERENCES
Areschoug, F.W.C. (1867) Bidrag till den skandinaviska vegetationens historia. Berlingska Boktryckeriet,
Lund.
Christ, H. (1867) Ueber die Verbreitung der Pflanzen der alpinen Region der europäischen Alpenkette.
Neue Denkschriften der Allgemeinen Schweizerischen Gesellschaft für die Gesammten Naturwissenschaften, 22, 1–85.
Forbes, E. (1846) On the connexion between the distribution of the existing fauna and flora of the British
Isles, and the geological changes which have affected their area, especially during the epoch of the
Northern Drift. Memoirs of the Geological Survey of Great Britain, 1, 336–432.
Hofmann, E. (1873) Isoporien der europäischen Tagfalter. E. Schweizerbart’sche Buchdruckerei (E. Koch),
Stuttgart.
Hooker, J.D. & Thomson, T. (1855) Flora Indica: being a systematic account of the plants of British India,
together with observations on the structure and affinities of their natural orders and genera, Vol. 1,
Ranunculaceæ to Fumariaceæ. W. Pamplin, London.
Watson, H.C. (1847) Cybele Britannica: or British plants, and their geographical relations. Longman,
London.
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