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LORENZO CAMERANO'S CONTRIBUTION

TO EARLY FOOD WEB THEORY


JOEL E. COHEN
Rockefeller University,
1230 York Avenue, New York, NY 10021-6399, U.S.A.

INTRODUCTION
When you first learn science, you want to know the facts. You want to know
Mendel's laws, though you don't really care who Mendel was. Later, if you go on
to do research, it's quite important to know the history of your field. ... Research
students who know the past have a decided advantage over those who don't.
James D. Watson, in Campbell (1986, p. 730)

In 1880, Lorenzo Camerano, then a 24-year-old assistant in the laboratory of the Royal Zoological

Museum of Torino, Italy, published a paper "On the equilibrium of living beings by means of
reciprocal destruction" in the Acts of the Royal Academy of Sciences of Torino. This pioneering
paper contains an early, perhaps the first, graphical representation of a food web as a network of
groups of species linked by feeding relations. It also contains a vivid dynamic model of the
consequences of trophic links for popUlation dynamics. The model is based on the propagation of
sound waves in an organ pipe.
introductory essay.

An English translation of Camerano's paper follows this

The purpose of this introduction is to summarize briefly Camerano's life and work, to place his
1880 article in the context of his other works, and to relate his paper to earlier and later works of
others.

A BRIEF LIFE OF CAMERANO


The following summary of his life relies largely on the memoir of Rosa (1919). Camerano was born
April 9, 1856, in Biella, at the foot of the Alps in northern Italy, and died in Torino November 22,
1917. His father was a functionary of the Piedmont prefectorial government. With his father's job,
the family moved to Torino, where Camerano studied at Liceo Gioberti and enrolled in the School
of Painting at Accademia Albertina. Among his school friends were Camillo and Mario, sons of
Michele Lessona, a Member of the Royal Academy of Sciences of Torino since December 1, 1867.
On February 14, 1874, Camillo and Mario asked Camerano if he would make some drawings their
father needed for zoological demonstrations. After school, Camerano went with them to meet their
father at the Museum of Zoology, then located at the Academy of Sciences. The elder Lessona was

352
an "ingenious proponent of the theory of evolution, at a time when this excited passionate conflict"
(Rosa 1919, p. 688). Camerano fell under Lessona's spell and, in many ways, followed in his
footsteps. Camerano eventually married Lessona's daughter Luigia.
On June 24, 1877, Camerano's first publication was approved for publication in the Acts of the
Royal Academy of Sciences of Torino. Written when Camerano was a student in natural sciences at
the University, it dealt with the polymorphism of females of the species Hydrophylus piceus. In
1878, Camerano received his "laurea" degree and was named an assistant in the Zoological Museum
of the Royal University of Torino. Two years later, Camerano received the degree of "Dottore
aggregato" in the Faculty of Sciences of the University. That same year, Lessona introduced
Camerano's paper "On the eqUilibrium of living beings" for publication in the Acts of the Royal
Academy.
Camerano's last publication, dated May 21, 1917, appeared in the Bulletin of the Museum of
Zoology and Comparative Anatomy of the Royal University of Torino and dealt with researches on
the subspecies of Capra sibirica Mayer. Camerano was then president of the Royal Academy of
Sciences of Torino (following Lessona, who had been president from 1889 to 1895) and a Senator
of the Kingdom. He died a few months later. His bibliography contains 341 items (Rosa 1919,
pp. 714-736).

CAMERANO'S 1880 PAPER IN RELATION TO ms OTHER WORK


Camerano's early papers were largely descriptive. They were often well-illustrated but showed little
inclination towards abstraction or generalization. A year before his 1880 paper, Camerano (1879)
published his first major book (344 pages) on entomology. Two years later, he (1882a) published
another book (252 pages) on the anatomy of insects. Few of his later papers had titles as titillating
as his 1884 essay on "Anomalous loves of the amphibia" (Camerano 1884), published in Archives
of Psychiatry. Some descriptions are accompanied by attempts to construct phylogenetic trees. He
published in Italian, Latin, French, German, and English in journals originated allover Europe.
Camerano's 1880 essay, which follows, appears to be unique in his oeuvre with regard to the
abstraction and generality of its ideas and its illustrations. There are two central ideas. First, the
abundance of each plant, phytophagous animal, carnivore and parasite has a natural eqUilibrium
level, and this eqUilibrium is maintained through feeding relations or, in today's language, trophic
links. Second, when the abundance of any component of a natural community is perturbed from its
eqUilibrium level, the perturbation propagates along the food chains much as the perturbations from
equilibrium of a sound wave propagate along the length of an organ pipe. These two ideas are

353

illustrated in differing degrees of detail, and with characteristic redundancy, in plates I and n, and in
plates ill and N, respectively.
It appears that the 1880 paper was a one-time flight of fancy for Camerano. Though I have not
examined every o,ne of his publications, I saw figures like these nowhere else in his work. In two
later papers on themes closely related to that of his 1880 paper (Camerano 1882b, 1885), he cites
this 1880 paper and repeats the argument of an equilibrium in which perturbations from equilibrium
are linked through feeding, but does not repeat any theoretical pictures. No similar pictures appear
in a bound volume that contains a complete collection of his works published in the Bulletin of the
Museum of Zoology and Comparative Anatomy of the Royal University of Torino (Camerano
1886-1917).
The central ideas of Camerano's 1880 paper appear to be closely related to contemporary ideas and
graphic devices of Darwin and Helmholtz. Among Italian biologists, as among biologists
everywhere, Darwinism was ascendant. Michele Lessona, Camerano's mentor, was a leading
proponent of Darwinism in Italy and had translated Origin of Species into Italian. According to
Camerano's (1902) review of biology in the nineteenth century, the Royal Academy of Sciences of
Torino in its session of December 28, 1879, awarded Charles Darwin the first of the grand prizes of
the Bressa Foundation. Though Darwin is not cited in the 1880 essay, Camerano refers to the
"struggle for life", a phrase that suggests Darwin's "struggle for existence". The first edition of
Darwin's Origin of Species (Darwin 1859) contains a single illustration, a plate inserted between
pages 116 and 117, which illustrates the principle of divergence in phylogenetic trees. The visual
image (Figure 1) of divergent treelike branching is remarkably similar to that of Camerano's first
plate. There is one important difference. Darwin's phylogenetic tree is crossed by horizontal lines
that suggest geological strata, time progressing from bottom to top. By contrast, Camerano's first
plate is marked by concentric rings expanding outward from vegetation. The rings suggest the
propagation of perturbations along a food web, by analogy to the propagation of sound waves.

354
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FIGURE I,

Three years before Camerano published his essay. Helmholtz (1877) published the fourth and last
German edition of his great book On the Sensations of Tone as a Physiological Basisfor the Theory
of Music , Helmholtz described the vibrations of sound by means of a graphical device (Figure 2).

with a horizontal line indicating equilibrium, and small vertical lines above or below the horizontal
indicating deviations from equilibrium, that is remarkably close to Camerano's plates ill and IV.
It is tempting to speculate that the graphics of Darwin and Helmholtz provided the inspiration for

Camerano's graphic representations of the connections and mutual perturbations of groups of


species in a natural community. But the graphics of Darwin and Helmholtz do not explain why
Camerano adapted them to ecological purposes, I speculate that Camerano's childhood artistic
inclinations and training led him to seek a visual representation of the population-dynamical

355

phenomena he observed in his entomological field work, and that he adapted the graphical ideas
available to him at that time.

FIGURE 2.
Another recurrent motif of the 1880 essay is that the equilibrium among different kinds of
organisms is maintained, and re-established following disturbance, by the organisms and the habitat
"without any action of man". Camerano may have had practical or scientific motives in emphasizing
this theme. Given the contemporary influence of agricultural entomology, he may have been trying
to show that many interventions in nature by applied zoologists were not really necessary. Or, as a
question of pure science, he may have sought to remove man's throne from ecology just as Darwin
removed it from evolution.

THE HISTORICAL SETTING OF CAMERANO'S 1880 ESSAY


Camerano was not the first to try to describe the food of animals, either in a general way or exactly.
He cites papers by agricultural entomologists, published in the 1870s, which described the feeding
habits of insects and birds in agricultural areas. But nearly half a century earlier, in February, 1832,
Darwin (1839, p. 10) described the food web of St. Paul's Rocks near the equator in the middle of
the Atlantic Ocean, and remarked with surprise on the apparent absence of plants. The notion that
the abundance of each species has a natural equilibrium level maintained through feeding relations
apparently also did not originate with Camerano. According to D' Ancona (1954, p. 20), "Moebius,
1877 [unfortunately, the citation is missing from the bibliography] ... recognized the importance of
interspecific nutritive relationships while he was studying the organisms living on the oysterbeds of
Schleswig-Holstein. To Moebius is due also the credit for noting that the effect of these
interspecific relationships is to establish a state of eqUilibrium. "

356
In America, detailed quantitative studies of the diet of birds, fishes, and insects were already well

under way by 1880. Forbes ([1878-1883] 1977), in papers published from 1878 onwards, cites
several works on animals of the Great Lakes region published in the decade or so prior to his own
works, and gives numerous quantitative tables of food taken by different consumers, in a format
now called a predation matrix or feeding matrix. Some ideas of Forbes and Camerano show
remarkable convergence. Forbes (1880a, p. 3) states the theme of a theoretical essay thus: "The
serious modification of any group [of organisms], either in numbers, habits, or distribution, must
modify, considerably, various other groups; and each of these must transmit the change in tum, or
initiate some other form of change, the disturbance thus propagating itself in a far extending circle. "
Introducing his empirical work on the food of birds, Forbes (1880b, p. 84) states:
Now a species is stable because the rate of its reproduction is uniform, because the
checks upon its increase are substantially unvarying, and because these two forces
balance each other. To set up any vibration in anyone of these checks, will necessarily
cause a corresponding vibration in the numbers of the species limited by it. More
explicitly, to set up an oscillation in a predaceous or parasitic species must produce a
reverse oscillation in the species parasitized or preyed upon. As the former increases,
the latter must diminish, and vice versa. But either a marked decrease or a marked
increase of a species will cause it to oscillate, unless made with extreme slowness, a
slowness so extreme as to allow progressive adjustments of all kinds to keep pace with
it.
The 1880 papers of Forbes and Camerano clearly share the ideas of a natural equilibrium population
level maintained through checks by food supplies and consumers, and the propagation of
perturbations along food webs. The last sentence quoted shows Forbes's awareness of the
importance of the rate of change of population abundance. Forbes (1880a, p. 11) also remarks on a
relation between the connectivity of food webs and the stability of communities:
It is a general truth, that those animals and plants are least likely to oscillate widely
which are preyed upon by the greatest number of species, of the most varied habit.
Then the occasional diminution of a single enemy will not greatly affect them, as any
consequent excess of their own numbers will be largely cut down by their other
enemies, and especially as, in most cases, the backward oscillations of one set of
enemies will be neutralized by the forward oscillations of another set. But by the
operations of natural selection, most animals are compelled to maintain a varied food
habit,-- so that if one element fails, others may be available.

Nevertheless, none of the reprinted papers of Forbes ([1878-1883] 1977) contains a food web
graph. Camerano's introduction of the food web graph appears to be a genuine innovation.
That innovation was evidently not noticed for some time. The earliest citation in the bibliography of
D' Ancona (1954; first German edition 1939) is to Camerano (1880), but the reference in the text
(D' Ancona 1954, p. 26) makes no mention of the food web graph. The development of food web
data, concepts and representations that included Camerano, Forbes and D' Ancona seems to have

357
been generally ignored recently. For example, Worster's (1977) history of ecological ideas cites
none of these three writers in its index and traces food webs and food chains no further back than
Charles Elton (see Worster 1977, pp. 295-298). D' Ancona and Forbes, but not Carnerano, are
cited by Kingsland (1985). Elton (1927) refers to neither Camerano nor Forbes in what is hardly
the only instance when an English writer has overlooked the work of continental and American
scientists. It seems likely that food web graphs had to be invented several times before their general
use became established. Food web graphs are not unique in this respect (Merton 1961, 1965).

CAMERANO'S IDEAS TODAY


Generalized food web graphs in which large numbers of functionally similar species are lumped
together in broad categories such as plants, herbivores, and carnivores have become standard
illustrations in introductory ecology texts. Though they are pedagogically useful, such pictures are
no longer acceptable as data for analyses of food web structure (Pimm, Lawton and Cohen 1991).
Camerano's and Forbes' descriptions of the propagation of disturbances from equilibrium along
food chains resemble remarkably the cascade model of population dynamics in lakes developed
recently by Carpenter and colleagues (e.g., Carpenter, Kitchell and Hodgson 1985; Carpenter
1988). "[W]e consider a lake food web that includes limiting nutrients and four trophic levels:
piscivores such as bass, pike, or salmon, zooplanktivores, herbivorous zooplankton, and
phytoplankton... Simply put, a rise in piscivore biomass brings decreased planktivore biomass,
increased herbivore biomass, and decreased phytoplankton biomass" (Carpenter et al. 1985, p.
634).
Camerano did not develop further his 1880 symbolic formalism for the propagation of disturbances
from equilibrium along a food chain, and others seem to have overlooked his suggestion. The
development of a coherent formalism for the interactions Camerano described had to await the work
ofLotka (1880-1949) and Volterra (1860-1940).
Oksanen (1991) pointed out that some insights of another early ecologist, the Finn A. K. Cajander,
have also recently been overlooked, then independently re-invented. As Oksanen suggested, an
acquaintance with earlier work could add efficiency to progress in ecology today.

ACKNOWLEDGMENTS
I thank Stephen Jay Gould, Claudia M. Jacobi, Richard D. Keynes, Robert K. Merton, and Mario
Rasetti for very helpful comments on a previous draft. James Drake obtained a copy of Camerano's

358

1880 article and shared it with Stuart L. Pimm, who shared it with me. Mario Rasetti, Member of
the Academy of Sciences of Torino, made it possible for me to use the magnificent library of the
Academy of Sciences of Torino. Guido Donini, Chancellor of the Academy of Sciences of Torino,
assisted me in locating biographical and bibliographical material on Camerano. Richard D. Keynes
pointed out Darwin's 1832 description of a food web. This work was supported in part by National
Science Foundation grant BSR87-05047 and by the hospitality of Mr. and Mrs. William T.
Golden.

REFERENCES
Many of Camerano's papers were published both in journals and separately as booklets. This dual
publication accounts for the discrepancy in the numbering of the plates of his 1880 paper and for the
incompleteness of the journal references to some of his other papers. The numbers given in square
brackets at the end of some references are the call numbers in the library of the Academy of Sciences
of Torino.
Camerano, L. (1879). Gli insetti: introduzione aZZo studio deZZ'entomologia. Torino e Roma:
Ermanno Loescher. 80 viii + 344 pp. [M2V94].
Camerano, L. (1880). Dell'equilibrio dei viventi merc la reciproca distruzione. Atti della Reale
Accademia delle Scienze di Torino. 15: 393-414. 80 24 pp. 4 tav. [Misc. B 296(21)].
Camerano, L. (1882a). Anatomia degli insetti. Torino: Ermanno Loescher 80 viii + 252 pp. 57
fig. [JK.VI.9].
Camerano, L. (1882b). De alcuni mezzi atti a preservare Ie piante dagli insetti nocivi. Torino. 80
18 pp. Annali d. R. Accademia d'agricoltura 24 [Misc. B783(16)].
Camerano, L. (1884). Amori anomali degli anfibi. Torino. 80 14 pp. Archivio di psichiatria Vol.
V [Misc B783(25)].
Camerano, L. (1885). n congresso ornitologico di Vienna e la questione degli uccelli e degli insetti
in rapporto coll'agricoltura (7 luglio 1884) [Misc. B296(42)] in Annali d. R. Accademia
d'agricoltura 27 :97 -113. Torino.
Camerano, L. (1886-1917). Collegione completa dei lavori inscriti nel Bollettino dei musei di
zoologia ed anatomia comparata della R. Universita di Torino. Torino. 80 1 vol. [130.527].
Camerano, L. (1903). La biologia nel secolo XIX. Milano: Francesco Vallardi. 40 50 pp. [Misc.
C97(25)].
Campbell, N. A. (1986). Discoverer of the double helix. BioScience, 36(11):728-731.
Carpenter, s. R. (1988). Transmission of variance through lake food webs. In: S. R. Carpenter,
ed., Complex Interactions in Lake Communities. Springer-Verlag: New York, pp. 119-135.

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Carpenter, S. R., Kitchell, J. F., and Hodgson, J. F. (1985). Cascading trophic interactions and
lake productivity. BioScience. 35(10): 634-639.
D' Ancona, U. (1954). The Struggle for Existence, Anne Charles and R. F. J. Withers, trans.,
Leiden (The Netherlands): E. J. Brill.
Darwin, C. (1839). Journal of Researches into the Geology and Natural History of the Various
Countries Visited by H. M. S. Beagle. Under the Command of Captain Fitzroy. R.N. from
1832 to 1836. London: Henry Colburn. Reprinted: Culture et Civilisation, Brussels, 1969.
Darwin, C. (1859). On the Origin of Species. Facsimile of 1st ed. Intro. by Ernst Mayr.
Cambridge, MA: Harvard University Press, 1966.
Elton, C. (1927). Animal Ecology. [New impression with additional notes 1935.] New York:
Macmillan.
Forbes, S. A. (1880a). On some interactions of organisms. Illinois Laboratory of Natural History
Bulletin 1(3): 3-17.
Forbes, S. A. (1880b). The food of birds. Illinois Laboratory of Natural History Bulletin 1(3):
80-148.
Forbes, S. A. (1977). Ecological Investigations of Stephen Alfred Forbes. New York: Arno
Press.
Helmholtz, H. L. F. (1954). On the Sensations of Tone. New York: Dover Publications.
Kingsland, S. E. (1985). Modeling Nature: Episodes in the History of Population Ecology.
Chicago: University of Chicago Press.
Merton, R. K. (1961). Singletons and multiples in scientific discovery: a chapter in the sociology
of science. Proceedings of the American Philosophical Society 105: 470-486.
Merton, R. K. (1965). On the Shoulders of Giants. New York: Free Press; also Harcourt Brace
Jovanovich 1985.
Oksanen, L. (1991). A century of community ecology: how much progress? Trends in Ecology
and Evolution 6(9): 294-296.
Pimm, S. L., Lawton, J. H. and Cohen, J. E. (1991). Food web patterns and their consequences.
Nature 350: 669-674, 25 April.
Rosa, D. 1919 L'opera scientific a di Lorenzo Camerano. Atti della Reale Accademia delle Scienze
di Torino (1918-1919) 54: 686-737.
Worster, D. (1977). Nature's Economy: A History of Ecological Ideas. Cambridge: Cambridge
University Press.

[393] Member Michele Lessona, Esq., submits the following work on behalf of the author, Dr. L. Camerano,
Assistant at the Zoological Museum of the Royal University of Torino 1

ON THE EQUILmRIUM OF LIVING BEINGS


BY MEANS OF RECIPROCAL DESTRUCTION
LORENZO CAMERANO*
[Translated by Claudia M. Jacobi. Edited by Joel E. Cohen.]
The building of the world is held together by the forces of hunger and love.
Schiller

The recent history of Zoology, starting some time before the middle of this century and still
continuing, is principally characterized by many researches on both the utility and the damages to
man due to animals. Applied zoology, in a word, was born in this period of development of the
zoological sciences.
Much has been written about animals that are useful and harmful, in various ways, to mankind.
Practical results have been, up to now, very scarce.
The reasons for this are many, starting with the tremendous difficulty presented by this type of
studies but, above all, because of an inclination to hasten to applications while disregarding data
from pure science.
This not only does not produce the expected results but also - more seriously - frequently gives
rise to erroneous theories about phenomena related to living animals and plants, theories which
hinder the development of science.
One of the most debated issues these days is that of animals which are useful and those harmful
to crops. [394]
This issue is one of the most complex regarding animals and about which there are no very clear
and conclusive ideas. This probably comes mostly from separately studying the relations
between animals and plants, that is, without taking into account the many and very important
relations among the different groups of animals which, as I will now explain, influence very
importantly the relations between animals and plants.
1 [Page numbers in the original appear here in brackets 1
* [CAMERANO, LORENZO. 1880 Dell'equilibrio dei viventi merce la reciproca distruzione. Atti della Reale
Accademia delle Scienze di Torino 15:393-414. 80 24 pp. 4 plates.]

361

To give an idea of the main current theories on the abovementioned issues I will briefly discuss
the case of insectivorous birds and insects harmful to crops, and the relations between these
groups of living beings.
Naturalists are divided on this topic, as is well known, in two categories.
One category admits the usefulness of birds since these destroy insects which damage crops, and
believes that by promoting increase of bird numbers the number of insects and the extent of the
damage they do could be reduced.
The other category, on the other hand, believes that the effect of birds is of little importance
concerning the destruction of insects harmful to crops, and that the development of birds would
not prevent the development of insects.
Naturalists belonging to the first category reason in this way: the number of insects which cause
damage to crops increases; that of birds, on the other hand, decreases. Now, birds feed to a great
extent on insects; so if we increase the number of birds, the number of insects will decrease.
The second category of naturalists think differently: the number of birds is high particularly in
those places where insects are very abundant. When the number of insects decreases, so does
the number of birds. Regions with low insect abundance also have few birds. The amount of
insects in a region depends essentially on the amount of plant food found in it: the [395] total
number of insects in a given locality is quite constant; what changes is the number of species and
of individuals belonging to each species (1) [see end-notes]. Hence, they conclude: birds play
only a small effect in destroying insects which damage crops.
Well-known naturalists have argued in favor of either one of the theories mentioned.
However, the number of naturalists who support the first theory is decreasing every day, while
those in favor of the second one increase. What has been said for the special case of insects and
insectivorous birds may be applied more generally to many other groups of animals.
To have an exact and clear idea, I repeat, of the relations between, for example, insectivorous
birds and phytophagous insects, and between these and plants, these groups cannot be studied
[396] separately. Rather it is necessary to study each in relation to all other animals to see the
general laws governing the equilibrium of animal and plant species.

362

I will now leave aside the special cases and will consider comprehensively all animals.
Before continuing, nevertheless, it is useful that I explain the meaning of some expressions
which I will frequently employ hereafter.
I say that a species is in equilibrium when the number of individuals is not significantly above or
below what is usually observed in a given locality.
I say that a species is increasing when the number of individuals is higher than what is usually
observed in a particular place.
I further say that a species is decreasing when the number of individuals in a given area is
reduced beyond what is usually observed in that place.
These observations apply to both animal and plant species.
Considering all animals, we can divide them in two broad groups:
1) phytophagous animals;
2) carnivorous animals.
Animals of the first group take their nourishment directly at the expense of the plant kingdom.
Animals of the second group feed on phytophagous animals.
It is a fact accepted by all that animals and plants develop in direct proportion to the available

food.
From this it follows that no species, be it carnivore or herbivore, can develop beyond a certain
limit which, if surpassed, would destroy the source of its own nourishment. Equilibrium, broken
by the excessive growth of either kind of animal, would again be reestablished.
To make clear the law I have just stated, let us look at a practical example. Let us consider a
portion of unploughed land, in which a great number of herbs and wild plants grow. [397] If we
visit this site, we

~ill

find a great number of insects, some phytophagous, which feed on the

various kinds of plants that grow there, some carnivorous, feeding on the phytophagous species.
Let us suppose that, for some reason, the number of carnivores increases extraordinarily, thus
soon consuming the herbivores almost completely. This in turn will benefit the plants upon

363
which this last group was feeding. But this effect would be temporary, for, once the herbivores
are destroyed, a great proportion of carnivores will have either to migrate or to die for lack of
food. Moreover, it would be almost impossible that not a single individual of the phytophagous
species would survive the slaughter. Now, these few individuals, thanks to the extraordinary
fecundity of insects, will soon repopulate the place with herbivores. Carnivores being very
scarce for the moment, herbivores will quickly develop and soon the plants which they feed upon
will suffer greatly and also die. The herbivores will thus be deprived of food and so these too
will have to die for the most part. Furthermore, the great number of herbivores will have already
fed the carnivores, which will have again started to increase in number, and in tum will resume
the destruction of herbivores, and so on.
The equilibrium of species, which is disturbed every now and then due to the abnormal growth
of some component, is reestablished naturally without the action of man.
From what we have said we can conclude that if carnivores are absent from a certain area,
herbivores will not multiply indefinitely, and that carnivores are not indispensable to prevent the
development of phytophagous species.
From what I have said above, it follows that:
1) the number of phytophagous animals depends directly on the vegetation;
2) the number of carnivorous animals depends directly on the number of phytophagous
animals and indirectly on the vegetation. [398]
It can be seen that the immense quantity and variety of animals are strictly related to the

vegetation, and that the different animals are related among themselves through feeding relations.
Carnivorous animals can be divided, in a very general way, essentially in two categories which
are:

1) carnivores which feed on phytophagous animals;


2) carnivores which feed on other carnivores.

I must mention that many species of carnivores feed indifferently on these both categories.
Carnivorous animals may also be subdivided in two groups, taking into account their life history
and their relations with the animals upon which they feed:
1) carnivorous predators;
2) carnivorous parasites.

364
Carnivorous parasites may in tum be divided into:
1) carnivorous parasites of herbivores;
2) carnivorous parasites of predatory carnivores;
3) carnivorous parasites of other parasites of herbivores or carnivorous predators.
I believe it would be useful for greater clarity to put together in a table the divisions of animals
mentioned, taking as a point of departure their feeding habits.

Phytophagous

predators

Carnivorous

A
L

parasites
(endoparasites)
(ectoparasites)

of phytophagous animals
{ of carnivorous animals
of phytophagous animals and
carnivorous animals

of phytophagous animals

lOf carnivorous animals

{ predators
parasites

S
[399]

I do not intend to elaborate on the various kinds of parasitism or of predation of animals. It is


enough to say that the phenomenon of parasitism is one of the most widespread in the animal
kingdom and that virtually no animal is free of parasites. The number of cases of parasitism
grows every day as animals continue to be studied.
There are parasites of herbivores or carnivores which are themselves parasitized by other
animals, just as carnivorous animals preying on herbivores or carnivores are themselves preyed
on by other carnivores, which in turn can be preyed on by other carnivores.
Most animals are at the same time the prey of other animal and subject to varied sorts of
parasites.
A great number of animals can be at the same time prey of other animals and subjected to various
kinds of parasites, and also predators or parasites of other animals which can be predators or
parasites of others, etc.
I have graphically displayed in Plate I the relations which exist among the various categories of
animals from the point of view of their nourishment. [Plates/ollow the text.]

365

Vegetation serves as food to herbivores upon which carnivores feed.


Carnivores that feed on herbivores are divided in two groups, carnivorous predators and
carnivorous parasites. Each of these groups is in tum devoured by other predators and other
parasites, etc. up to those groups which are not preyed upon by other predators but are
essentially subject to endoparasites, which are in charge of preventing their growth.
I have considered a special case in Plate II. I have pictured the enemies of phytophagous
Coleoptera and the enemies of those enemies, etc. This plate is constructed analogously to the
preceding.
Phytophagous Coleoptera eat vegetation and are decimated by other predators and parasites.
[400] These in tum are decimated by other animal predators and animal parasites and so on.
For example, phytophagous Coleoptera are in part eaten by certain mammals. These mammals
are prey of various reptiles, birds and larger mammals, and are subjected to all manner of
parasites. These birds, reptiles and mammals are themselves prey of other mammals, birds and
reptiles, and are also variously parasitized.
Phytophagous Coleoptera, besides being eaten by mammals, are parasitized by several
Hymenoptera. These Hymenoptera can be devoured by amphibians, reptiles, birds, mammals,
Orthoptera, other Hymenoptera, arachnids, etc., and these are themselves subject to other
parasites, etc.
Plate II shows the complexity of the causes affecting the enemies of phytophagous Cpleoptera,
that is, Hymenoptera, Orthoptera, carnivorous Coleoptera, arachnids, amphibians, reptiles,
birds, mammals, Diptera and parasitic spiders.
Now that we have seen the feeding relations between animals and plants, and among animals, let
us see how the law stated above applies to an increase in animal numbers: that animals develop
in direct proportion to their available food.
Essentially two categories of causes act to increase or diminish animal numbers. One could be
named extrinsic and the other one intrinsic. The extrinsic causes come from the outside world
and act directly or indirectly on animals. Included in this category are climatic, physical, and
geological conditions from the locality in which the animals live.

366

Suppose, for example, that the forest of a certain area is cut, and this is done at very large scales
in our days. It is evident that the climatic conditions in that region will change, more or less
quickly and [401] more or less profoundly. Cutting down a forest frequently modifies the nature
of the soil, favoring river overflow and flooding of the neighboring plains, on which they
deposit different sorts of material which can directly affect the animals that live in those regions,
or have an indirect effect by preventing the development of many kinds of plants.
Cutting the forest also reduces the number of animals: first because it directly affects the animals
that lived on the felled plants; second because plants which grew in the shadow of the trees cut
and that served as food to many groups of animals can no longer develop; third because it has a
direct effect on animals by modifying climatic and physical conditions in the area.
An example of what I have said just now can be seen in the surroundings of Turin.

Insects that were abundant some thirty years ago along the forested banks of the rivers Po, Dora
and Stura have nowadays become quite scarce, and many species have completely disappeared or
are rapidly disappearing.
A very important extrinsic cause is also that which comes from climatic variations.
Continuous and abundant rains, for example, long-lasting droughts, and also intense and
prolonged cold weather, can cause decreases in animal numbers.
In a word, it can be said that animal numbers tend to vary according to any cause (climatic,
physical or geological) that alters normal conditions in a given area.
These changes have almost always an indirect effect on animals, while they affect vegetation
more directly.
Damages caused directly to animals by these factors are relatively small, and are certainly not
enough effectively to prevent their development in great numbers.
The intrinsic causes directly affect [402] the animals and originate from their feeding relations.
For example, the growth in number of a species A, predator of another species B, is an intrinsic
cause directly affecting the latter by reducing its number of individuals.

367
On the other hand the development of a large number of individuals of a species C, predator of

species A, is a direct cause of decrease of individuals of species A, while indirectly preventing


the destruction of species B by species A.
More generally, we can say that considering a species A which affects another species B, that is,
feeds on it either by preying upon it or parasitizing it, we have (2) [see end-notes]
A~B=-B

If a third species C acts on species A we have: C

A=-A

resulting in: - A and + B.


Let us suppose that a fourth species D affects species C, then: D

=- C

resulting in: - C and + A


so that we will again have + A
and hence - B.
Suppose a fifth species E affects species D,
then: E

D=-D

and so + C and - D
[403] and again + C and - A
and consequently + B.

If we now suppose that a sixth species F affects species E, we will have the same pattern:
F ~ E=-E
which will result in: - E and + D
and hence + D and - C
resulting in - C and + A and
again - B
and so on.
It is seen from the preceding that a cause acting directly on a species is in most cases an indirect
cause for the reduction of another species, and that a direct cause of increase in a species can be
an indirect cause of increase of another species. Or even, a direct cause of decrease in a species
can be indirectly a cause of either increase or decrease of other species.
The result of this series of intricate relations between living beings with regards to their
nourishment is the struggle for life. This comprises essentially two series of actions, that is, a

368

series of actions that the animals perform to obtain their food, and another series that animals
perform to avoid being themselves food for other animals.
The struggle for life, understood in its broadest sense, is itself a very important intrinsic cause
and exercises a major influence on the equilibrium of beings.
For greater clarity, I will now put together in a table the main causes that modify the equilibrium
of living beings. [404]

C
A
U
S
E
S

extrinsic [dUm
indirect

intrinsic

[dUm
indirect

{ climatic. physical, and other conditions that act directly on the animal species

climatic. physical (geological, etc.) conditions that directly affect the plants which are food for
phytophagous animaJs and thus indirectly affect the animaJs

{predators}

struggle for life

{ parasites

Now that the main causes for change have been pointed out, let us see what effects these produce
on the eqUilibrium of living beings and in which way eqUilibrium can be explained in spite of the
action of these forces, which tend to alter the reciprocal relations between the various groups of
animals.
Examining a given locality, we see that the various animal species have for a certain time an
almost constant amount of individuals, and the relations between phytophagous species and
vegetation and between phytophagous species and carnivores are also substantially constant.
When this occurs we will say that in that locality the animals are in eqUilibrium.
It might happen that in the same area a certain species increases suddenly and extensively, or that

a given species decreases suddenly, or even that a species is increasing or decreasing gradually
but significantly.
When any of these things happens we say that the equilibrium of the animals of that place is
broken, and we will be able to distinguish a period of growth or reduction according to the
species under consideration.

369
If we continue observing the animals of the same place, it will be frequently observed that those

species which have grown extraordinarily, as well as those which have undergone gradual
reduction will eventually return sooner or later to constant levels. [405]
Equilibrium is thus reestablished without the intervention of man.
Examples of these phenomena are many and can be taken from any group of animals. Insects,
however, are the animals which provide us with the most outstanding and easily studied ones.
It is frequent, for example, that in a particular area an insect species develops in such a way as to

cause damage to some particular plant species for one, two, three years or more. It can later
disappear almost completely from the place, only to reappear some time later. The grapevines
from several places in Astigiana were damaged some years ago by a great number of Anomala

vilis, a Coleopteran which although not new in the region, usually occurred in low abundance.
Nowadays the insect is again found in small numbers.
Also in this case human interference was very small, even if in a few places, not very extensive,
some action was taken to destroy the species named.
This localized development of some species can be observed in very restricted areas. A species
may be very abundant in a garden or a field while this is not seen in other gardens or fields which
are apparently subjected to the same conditions.
An example of this nature can be easily seen in the surroundings, or rather, within the city of
Turin itself. There are years in which the poplars from the old main square are entirely
devastated by thousands and thousands of Liparis salicis caterpillars, while the poplars in
Queen's A venue, not far from there, remain almost immune.
Also in this case we see that periods of great abundance of individuals are followed by periods of
less but constant activity and by periods of scarcity of the same individuals.
Also in this case the part played by man is negligible.
How can these facts be explained? How can the practically always constant quantities of many
categories of animals be explained, etc.? [406]
There is no doubt that these phenomena are a product of the various causes mentioned above.

370

To explain the very complex facts I believe that it is possible to resort to a reasoning analogous to
that which serves to explain the propagation of sound waves in a generic cylinder, or better still,
in a generic medium. This may seem rather daring but the concept will become clearer after I
have exposed some general considerations.
Although the ideas can be applied to the majority of cases, I will use here for simplicity and
clarity the example of the relations between the reciprocal actions of vegetation and herbivores,
carnivorous animals which prey on herbivores, carnivores which parasitize predators, those
which prey on parasites and finally endoparasites of parasite predators.
Suppose that for some reason the vegetation in a given locality happens to grow abnormally.
This increase in vegetation will necessarily result in an increase in phytophagous animals,
following the rule already noted which states that animals grow in direct proportion to the
available food. This increase in phytophagous animals will, for the same reason, promote an
increase in carnivores who prey upon herbivores. This in turn will promote the development of
animals which parasitize those predators. The increase of parasites will cause an increase of
animals which prey on these parasites. Finally, this will increase the endoparasites of these
predators.
Now, an increase of endoparasites will tend to reduce the number of animals in which they live,
that is predators, resulting in the growth of their prey.
In our case, this will increase the number of parasites, and this will again result in a decline of
other predators. Consequently, the number of phytophagous animals will increase, which in
turn will diminish the quantity of vegetation. [407]

Assuming that other causes have not changed, this decrease of the vegetation will result in a
decrease of herbivores; this will reduce the predators; this, the parasites; this in turn the other
predators and then their endoparasites.
Now the decrease of endoparasites will cause an increase of their prey and hence the number of
predators will increase. This will result in a decrease of parasites, promoting growth of other
predators, which will reduce the number of herbivores. The decrease of phytophagous animals
is one of the causes of increase in the quantity of vegetation.

371

If the same causes continue to exist, increase in vegetation will cause the increase of herbivores
which in tum will increase the number of predatory carnivorous animals, etc. The same cycle
will be repeated again.
Figure 1 in Plate ill represents graphically what is stated above.
The lines that rise from the horizontal line indicate an increase in growth of a given category of
animals; the lines that go down from the same horizontal line indicate decrease in development of
a given group of animals.
The signs + and - indicate these facts.
Equilibrium is represented by the horizontal line 0 already mentioned.
The arrows show the directions in which the causes for disturbance are acting.
This notation serves for all figures in Plates ill and IV in this work.
The case just mentioned is the simplest, and assumes that a complex of causes affects the
vegetation and that the results of this disequilibrium is slowly propagated successively to the
different categories of animals. When equilibrium is broken by the increase in [408] vegetation,
it starts a reaction which propagates in a given direction and brings about certain determinate
changes in the equilibrium of other categories of animals. While this first impulse proceeds, a
second one is produced as a direct consequence of it which will modify the action produced by
the first one, as I will explain better soon.
Let us first consider what occurs between vegetation and herbivores, and between predatory
animals and endoparasitic animals.
The increase in vegetation increases the number of herbivores. Now, while on the one hand this
will increase the number of their predators, it is also on the other hand cause for vegetation
reduction. Thus the great increase in vegetation is overwhelmed as a direct consequence of this
increase, by the great development of herbivores. This decrease in vegetation produces a
decrease in the number of herbivores, so that in this case too the increase in herbivore numbers is
overwhelmed by its consequences. This decrease of herbivore numbers will again make
vegetation thrive, and this in tum will make the number of herbivores grow, and so on.

372
The same occurs with predatory animals and endoparasites.

A great development of

endoparasites will result in a decrease in predators and this will produce a reduction of
endoparasites. This will increase the number of predators. The increase in predators will cause
an increase of endoparasitic animals. This will again reduce the number of predators and so on.
This is depicted graphically in Figures 2 and 3, Plate ill.
It follows from what has been said thus far that there are two centers of disturbance which
vibrate continuously, which are vegetation on the one hand and endoparasites on the other.
These are not, however, the only sources of disturbance that can be observed in the animal series
mentioned above. [409]
It may happen, for example, that for some of the reasons already stated the number of predators
or parasites either increases or decreases abnormally. What will then happen to the other
categories?
Suppose that the predatory animals increase (Plate ill, fig 4). Then, on one side the number of
parasites of these predators will grow, and hence the number of other predators, and lastly of
endoparasites; on the other side we would see a decrease in herbivores, whence an increase in
vegetation.
If we now suppose that predators decrease (Plate ill, fig. 5), parasites will decrease. This will

reduce on the one hand the number of predators and of their endoparasites, but on the other hand
will increase herbivores which will result in less vegetation.
What I have conjectured for carnivorous predators can be applied to carnivorous parasites etc.
It is now clear that besides these two disturbance centers at the extremes of the animal series
considered, every other group within the series can become in its turn a center of disturbance.
Examining one of the ends of the series of living beings, for example vegetation, we see that an
increase in vegetation will promote an increase in phytophagous animals, that is, a pulse or cause
of disequilibrium that moves forward and is felt by the other categories of living beings.
Increasing numbers of phytophagous animals will immediately reduce the amount of vegetation,
and this will be a second cause of disequilibrium which continues in the direction of the first one,
canceling the effects of the first one.

373
In this way, the balance broken at any point of the series of living beings is reestablished by the

relations between them and are based on food, in other words, on reciprocal destruction (Plate
III, fig. 6).
The same reasoning may be repeated starting from any of the disturbance centers just mentioned.
[410]
Figures 7 and 8 in Plate ill show the effects of considering other centers of disturbance.
What has been said for two causes of disequilibrium that are generated in a given point can be
also said for any number of causes of disequilibrium that rise successively from the same point.
Let us suppose (Plate ill, Figure 9) that the amount of vegetation increases. This will make the
number of phytophages increase and will be a first cause of disequilibrium that will move
forward along the series. The increase in phytophages will produce a second cause of
disequilibrium - less vegetation and less phytophagous individuals - which will proceed in the
same direction as the first. But the decrease of phytophages will again favor an increase in
vegetation and this again will promote an increase of herbivores, creating a third cause of
disequilibrium. The increase in herbivores will produce a fourth cause of disequilibrium, as it
decreases the amount of vegetation, and this again reduces the number of herbivores. In this
particular case we then have four causes of disequilibrium which rise from the same points of
propagation, vegetation and phytophages, that tend to suppress each other alternately. What has
been said for four causes of disequilibrium can be repeated naturally for any number of causes
and for any starting point.
Let us consider now, for simplicity, what happens when a single cause of disequilibrium
originates in the vegetation and arrives at the endoparasites (Plate IV, Fig. 1). This category
reacts, so to speak, causing another disequilibrium which will return to affect the vegetation.
It can be thus said that the cause of disturbance produced by the vegetation which has reached the
endoparasites is reflected and modifies its own effects, either by increasing it or suppressing it.
In the case of Figure 1, both the first cause of [411] disequilibrium and the second one - which is

but the consequence of the reflection of the first one - would produce equilibrium among
predators, increasing instead the parasites which would again produce the equilibrium of
predators, increase phytophages and finally produce the equilibrium of the vegetation.

374

This case - I repeat - is one of the simplest: things are much more complicated in nature.
Figures 2, 3, 4 and 5 in Plate IV show the consequences when the causes of disturbance rise
from any of the points of propagation and are later reflected.
Figure 6 in Plate IV shows the consequences of four causes of disequilibrium that are reflected in
pairs.
In nature, the various causes of disequilibrium do not act in isolation, as I have supposed here

for clarity. Instead they superpose, intertwine, at times canceling one another, at times
increasing, at times decreasing.
In Figure 7, Plate IV, I have considered the case in which eleven causes of disequilibrium rise

from all the points of propagation already mentioned and go in various directions, sometimes
reflecting on themselves and sometimes not. The directions of the arrows and their color [color

is omitted in the reproductions of the plates] clearly show the development of the causes of
disequilibrium so that there is no need to repeat here the explanations given for the various
special cases of disequilibrium. In Figure 7, I have indicated the final result produced by the
action of the causes of disequilibrium considered. To obtain this curve it is enough naturally to
make the algebraic sum of the causes above (+) and below (-) the line of equilibrium (0) for each
category of living beings.
The case I have considered is one among many that are observed in nature. It will suffice to
change the number of causes of disequilibrium and the numbers of propagation points to obtain
many others.
It is now clear that:

1) the categories that constitute the animal series have an indefinite number of disequilibrium
[412] centers, and each group of animals is a potential center of disequilibrium for the
other groups of animals;
2) all the causes of disequilibrium produced by single groups of animals ultimately reflect at
one end on the vegetation and at the other end on the endoparasitic animals.
These two categories of living beings reflect, though modified, the causes of disequilibrium
themselves;

375
3) all these causes of disequilibrium tend either to suppress each other, or to increase and to
decrease one another alternately.
From what was said it is clear that the propagation of causes of disequilibrium can be very well
compared to the propagation of sound waves from any center of disturbance along a radius. In
the same way as there are sound interferences when waves corning from two or more centers of
disturbance meet, it is also frequent to see cases of interference among the causes which govern
various groups of living beings and tend to modify their equilibrium.

In a word, we could compare the different groups of living beings to a set of diapasons which
vibrate together generating a series of waves which interfere with one another, with well-known
effects.
From what has been said it can be concluded that:
1) the equilibrium between vegetation and animals, and among the various groups of animals
is maintained by the animals themselves;
2) the causes of disequilibrium produced by man in any animal group, such as deforestation,
agriculture, etc., have many times insignificant effects because they are counterbalanced
by other causes corning from the various groups of animals themselves;
3) the complete extinction of any animal group may produce either light effects or very
pronounced ones. In any case, even when the disequilibrium caused by disappearance
of an animal group is severe, this is eventually compensated by other causes generated
by the animals themselves. [413]
4) the part played by man is very small regarding changes in the equilibrium of the various
groups of living beings, and hence in the destruction of certain species, or the abnormal
development of others. In any case the momentary effects produced by these factors
tend to stop by the effects of other factors coming from the various groups of living
beings themselves.
Anybody could, nevertheless, object concerning this last conclusion that man can very easily
destroy completely many species of living beings. This is true and there are many examples of
species now extinct by human action. Man destroys especially big animals, but has not been
successful in destroying directly those small species of animals which are so frequently harmful
to humans.
The destruction of a big mammal species, for example in a given area, by man, is certainly not a
small cause of disequilibrium, but this disequilibrium is momentary, and balance is reestablished
without the action of man. It is useful that I mention on this topic what happened in the region of

376
Piedmont. Professor Michele Lessona in his account on the Snakes of the Piedmont (3) [see
end-notes] speaks about the gathering of snakes that took place in the Lanzo valleys between the
end of the past century and the beginning of ours. He refers to a passage in the book by Count
Luigi Francesetti di Mezzenile, entitled Lettres sur les vallees ck Lanzo, in which the gathering of
snakes is discussed. From this passage it is evident that in the valleys of the Lanzo alone many
thousands of snakes were caught for pharmaceutical uses.
Snake gathering ceased entirely in the Lanzo valleys and ~so in other places some twenty years
ago. It could be deduced, from the original number of snakes, that their actual number would be
enormous. But this is not so. Stopping the snake hunt must have broken the equilibrium of
many [414] categories of living beings in the Lanzo valleys. The numerous snakes must have
greatly decreased the number of frogs and small mammals of these Valleys. Lower numbers of
frogs and small mammals must in tum have been a cause of lower numbers of snakes. The
reasoning from the general cases can be applied here.
What is important to notice is that the equilibrium which comes from the relations among the
animals in the Lanzo area disappeared in a relatively brief time, due to the animals themselves
without any action of man.

ENDNOTES
(1) Also consult on this topic, among other works, Camerano, The Insects, Turin, E. Loescher,

1879, p. 26, etc. I have treated the main questions concerning the relations between insects and
agriculture in a passage of this book: "Let us consider a portion of unploughed land, in which a
large number of herbs and wild plants grow. If we visited this place, we would find a large
popUlation of insects; some phytophagous, which feed on the various kinds of plants available,
some carnivorous, feeding on the phytophagous species ... Suppose that the area is now tilled.
The farmer, by plucking out the plants that filled up the place and turning over the soil with the
plough, destroys a large number of insect species which either lived on the uprooted plants or
metamorphosed there."
"Following the seeding, only one or two species of plants will take the place of the many more that
the farmer took away. Very few insect species will remain, and these will precisely be .those which
feed on the cultivated plants. Since these plants are in great number, the number of insects will soon
grow to such an extent as to cause great damage to the plants."
"This does not mean, as is the general belief, that the total number of insects has increased. Before
the tillage the area was populated with many types of insects. After the tillage the number of species
is reduced, and instead the number of individuals from the remaining species has increased."
(2) The direction of the arrows shows the direction in which one species acts on the other one. I
use the symbol =to indicate the result of the action of one species on the other one, the symbols when this action reduces the number of individuals of a species, and + when the action of a
species does not reduce the number of individuals of a second one.

(3) Atti della Reale Accademia delle Scienze di Torino, vol. XII, 1877.

377

Plate I
vegetazione = vegetation
predatori = predators
parassiti = parasites
endoparassiti = endoparasites
carnivori =carnivores
1ill"lJj,., .IX
J.illltlj>ttrtr.uili

~.

--------"',

378
Plate n
coleotteri fitofagi =phytophagous Coleoptera
imenotteri = Hymenoptera
ortotteri =Orthoptera
acaridi = mites
anfibi = amphibians
rettili = reptiles
uccelli =birds
mammiferi = mammals
ditteri = Diptera
vermi endoparassiti = endoparasitic worms
parasiti vegetali = parasitic plants
rincoti =true bugs (Hemiptera)
aracnidi =arachnids
pesci predatori =carnivorous fish
crostacei parassiti =parasitic crustaceans
7'lwoln X

I __ ~",,..'I

ru

./""(''''11';/;''

,'rl,,.,'~/v

fn

379

Plate ill
aumento = increase
equilibrio = eqUilibrium
diminuzione = decrease
[In the originals of plates III and IV, a grid of Cartesian coordinates is printed in red. Various
arrows are printed in red, green or black, but no explanation is given for the different colors.]

~1:1' ~Tl' 1'T' ~T ~r

";A"

"

'Y; ' i"

~l"" ii, -J".v \'J-"'::

:..- .... ...... . . . ,.-......' "'-1"'' -,""1'..... ""'1"-'' 1'-l~li. "~-I': -;-:~,
' .::-. "-'f- ----l'~'"
,: : I ;II ,: :

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........

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'I
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....

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_,

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iiW"

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~I

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~

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. ': ' ;~ 1 1 I
+" ',---,-I
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I "",.I""'r- Ir
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'+ ;

., :

r.,..--",""

"-I .

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hM.

itr,",", 1'IrwI. -t.

380

Plate IV
[no new words]

TtiI'offl XII.

Pill'
'~IiJ.

1f1..L

1trtI.

:e: I I

"'it~

h .....

1"tWI.

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." i :'

I I 1
,

1.,"",,*,,,
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:t= I

1 I.: 1

'0

Etf";';,,.itl .
/hit"#IW',w,

:t: I \ \

.....,Ii'A.

I"~/.

/:-I"",/~

\ I
f;~.S

';,,-f.,ft-'-

~-

I",-wl, I'MI .

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FI

,..,.,.

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til"';

fh..I.

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~ 11' 1\ ' I i ~
'I';: j

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