Transcriber’s Note
When italics were used in the original book, the corresponding text has
been surrounded by _underscores_; bold or larger characters (such as the
large numbers in the index) have been surrounded by =equal signs=.
Superscripted characters are preceded by ^. Mixed fractions have been
displayed with a hyphen between whole number and fraction for clarity.
When a fractional number has been used in a range (i.e. one and a half
to two and a half), the dash separating the two numbers has been spaced
(i.e. 1-1/2 - 2-1/2).
Some corrections have been made to the printed text. These are listed in
a second transcriber’s note at the end of the text.
OBSERVATIONS OF A NATURALIST IN THE
PACIFIC BETWEEN 1896 AND 1899
[Illustration]
[_Frontispiece._
[Illustration: Selala (Vanua Levu, Fiji) (1/3).]
[Illustration: Fruits with seedlings nearly ready to fall from the tree
of Rhizophora mangle (shortest) and Rhizophora mucronata (longest). From
Vanua Levu, Fiji. (1/4 of the true length.)]
[Illustration: Rhizophora mangle (Vanua Levu, Fiji) (1/4).]
[Illustration: Rhizophora mucronata (Vanua Levu, Fiji) (1/5).]
OBSERVATIONS OF
A NATURALIST IN
THE PACIFIC BETWEEN
1896 AND 1899
BY
H. B. GUPPY, M.B., F.R.S.E.
VOLUME II
_PLANT-DISPERSAL_
London
MACMILLAN AND CO., LIMITED
NEW YORK: THE MACMILLAN COMPANY
1906
_All rights reserved_
RICHARD CLAY AND SONS, LIMITED,
BREAD STREET HILL, E.C., AND
BUNGAY, SUFFOLK.
Dedication
TO THOSE NUMEROUS PERSONS TO WHOM I WAS INDEBTED FOR
GREAT KINDNESS AND ASSISTANCE DURING MY
SOJOURN IN HAWAII AND FIJI
PREFACE
ALTHOUGH this volume contains a great amount of original material, I am
largely indebted to the labours of my predecessors for its present form;
and a scheme that at first was limited only to my own observations in
the Pacific has gradually extended itself to the general subject of
plant-dispersal. The farther I proceeded in my work the more I realised
that the floras of the Pacific islands are of most interest in their
connections, and that the problems affecting them are problems
concerning the whole plant-world. Deprived of the writings of Seemann,
Hillebrand, Drake del Castillo, and other botanists, several of whom
have lived and died in the midst of their studies of these floras, and
without the aid of the works of Hemsley and Schimper, generalisers who
have mainly cleared the way for the systematic study of
plant-distribution and plant-dispersal, it would not have been possible
for me to accomplish such an undertaking.
My interest in plant-dispersal dates back to 1884, when, whilst surgeon
of H.M.S. _Lark_, in the Solomon Islands, I made some observations on
the stocking of a coral island with its plants, which were published in
the _Report on the Botany of the “Challenger” Expedition_. In 1888 I
followed up the same line of investigation during a sojourn of three
months on Keeling Atoll, and during a journey along the coasts of West
Java. But realising that as yet I had barely touched the fringe of a
great subject, and that several years of study would be required before
one could venture even to appreciate the nature of the problems involved
and much less to weigh results, I took advantage of the circumstances of
my life to make, between the years 1890 and 1896, a prolonged
investigation of the plants of the British flora, mainly from the
standpoint of dispersal by water. This involved the study of the
seed-drift of ponds and rivers and of the plants supplying it, a study
which brought me into close relation with aquatic and sub-aquatic
plants. This line of investigation led me into contact with many other
aspects of plant-life; and as time went on my field of interest extended
to the plants of dry stations and to the bird as an agent in
plant-dispersal. Only a few of these results have been published, as in
the journals of the Linnean Society and of the Royal Physical Society of
Edinburgh as well as in the pages of _Science Gossip_. They lie for the
most part still within my note-books, and fitly so, since I regarded
such studies chiefly as a preparation for the investigation of the
general question of plant-dispersal.
When again, in October, 1896, I found myself once more in the Pacific,
the subject was taken up again with zeal; but my larger experience had
only increased my diffidence, and the unknown looked so overwhelming
that I settled down for the next three years content with merely making
experiments and recording observations. Here again the main problem was
attacked through the study of seed-buoyancy, and gradually it led me to
the systematic study of the mangroves and of the beach-plants, whilst my
inland excursions brought me into familiarity with the plants of the
interior. My geological exploration of the island of Vanua Levu, in
Fiji, greatly assisted me by giving a method to my botanical examination
of the island.
Whilst working out my geological collections in England, in the years
1900-1902, I devoted an hour or two daily to the elaboration of my
botanical notes and to a consideration of the problems concerned. During
a winter in Sicily I took up again the subject of the beach-plants; and
after the publication of the volume on the geology of Vanua Levu I was
able to accomplish a plan, for years in my dreams, of visiting the
eastern shores of the Pacific. During a period of three months from
December, 1903, to March, 1904, I examined the littoral flora of the
west side of South America at various localities between Southern Chile
and Ecuador; and finally completed this investigation by comparing the
shore-plants on the Pacific and Atlantic coasts of the isthmus of
Panama. Returning to England with a fresh collection of data, I passed
many months in elaborating and arranging all my notes, waiting vainly
for a clue to guide me in framing a scheme by which I could bring the
results of many years of work into some connected form. At last I
decided once again to take the floating seed as my clue, and without any
prearranged plan I allowed the work to evolve itself. Now that it is
finished, I can see some obvious defects; but if any other plan had been
adopted I scarcely think that I should have been more successful in
piecing together in a single argument materials resulting from so many
years of research and relating to so many aspects of plant-life.
Yet the final object of a naturalist would be but a sorry one, if his
aim were only to write a treatise and append his name to it. His
personal faith lies behind all his work; and no one can pursue a long
line of study of the world around him without rising from his task with
some convictions gained and some convictions lost.
As far as the observation of Nature’s processes at present in operation
can guide us, the world presents itself to us only as a differentiating
world. We can perceive, it is true, a progressive arrangement of types
of organisms from the lowest to the highest, and we can perceive a
development of varieties of the several types; but the only process
evident to our observation is that concerned with the production of
varieties of the type. Nature does not enlighten us as to the mode of
development of the type itself. We can, for instance, detect in actual
operation the process by which the different kinds of bats or the
different kinds of men have been developed; but there is no principle in
Nature evident to our senses that is concerned with type-creation.
Though we can supply it by hypothesis, we cannot discover it in fact. On
the other hand, the evidence of differentiation is abundant on all sides
of us, both in the organic and in the inorganic worlds. The history of
the globe has ever proceeded from the uniform to the complex; and in the
closing chapter of this book an endeavour is made to connect the
differentiation of plant and bird with the differentiation of the
conditions of existence on the earth. But this leaves no room for the
development of new types of organisms; and so far as observation of the
processes of Nature at present working around us can guide us, each type
might well be regarded as eternal. We can never hope to arrive at an
explanation of the progressive development of types by studying the
differentiating process; and since the last is alone cognisable for us,
evolution, as it is usually termed, becomes an article of our faith, and
of faith only.
In illustration of this argument, let me take the case of the races of
men. We see mankind in our own day illustrating the law of
differentiation all over the globe, as far as physical characters are
concerned. Just as the ornithologist would postulate a generalised type
in tracing the origin of various allied groups of birds, so the
anthropologist, guided by his observation of the changes now offered by
man in different regions, would postulate a generalised original type as
the parent-stock of mankind. Observation of the processes of change now
in operation by no means leads us to infer that such a generalised type
was an anthropoid ape, or even simian in character. In so doing we
should be forming a conclusion not warranted by the observation of
existing agencies of change, and we should be confusing the two distinct
processes of evolution and differentiation, or rather of progressive and
divergent evolution, of which the last alone comes within our field of
cognition. The study of variation can do no more than enable us to
ascertain the mode of development of different kinds, we will say, of
birds or of men. The origin of the type lies outside our observation.
“Given the type, to explain its origin”: this is the problem we can
never solve, and Nature aids us nothing by the study of her ways. On the
other hand, there is the subsidiary problem.... “Given a type, to
explain its varieties” ...; and here Nature’s processes are apparent to
us in a thousand different shapes.
It might seem that the presumptive evidence connecting man in his origin
with the monkeys is so strong that, supposing his simian descent were
regarded as a crime, a jury would without hesitation pronounce his
guilt; but until some observer of the processes followed by Nature can
bridge over the gap that divides man from the ape, until indeed he can
offer a legitimate illustration of how it is accomplished in similar
cases in our own day, the gap remains. Those who have read the recent
work of Prof. Metchnikoff on the Nature of Man will properly regard his
chapter on the simian origin of man as a brilliant argument advanced by
a most competent authority. Yet he fails to complete his case by
bridging over this gap, and can only appeal to the results of the now
famous researches of De Vries concerning the mutations of the evening
primrose (Œnothera). It is probable, he says, that man owes his origin
to a similar phenomenon (English edition, p. 57). Several objections
could be raised against this illustration from the plant-world, the most
important of them lying in the circumstance that these mutations could
only be urged as instances of the sudden development of new species of
the evening primrose type. They merely illustrate the process of
differentiation from a given type, and by no means represent the process
of progressive evolution from a simian to a man.
However, look where we may—and this is the great lesson I have learned
from my researches in the Pacific islands—Nature does not present to our
observation any process in operation by which a new type of organism is
produced. The processes involved lie hidden from our view. The channels
by which impressions from the outside world reach us are comparatively
few; and although it seems likely that the future development of man
will be mainly concerned with the acquirement of additional
sense-channels, no newly acquired sense will enable him to be at once an
actor in and a spectator of the great drama presented in the organic
world. That a creature should be able to get at the back of its own
existence, or, in other words, to penetrate the secret of its own
creation, is unthinkable. Outside the limited field of observation that
immediately surrounds us extends the region where reason alone can guide
us, and beyond lies the realm where reason fails and faith begins.
H. B. GUPPY.
_November 8th, 1905._
LIST OF SOME OF THE PRINCIPAL AUTHORITIES QUOTED IN THIS VOLUME,
WITH AN ENUMERATION OF THE AUTHOR’S BOTANICAL PAPERS
BURKILL, I. H., “The Flora of Vavau, one of the Tonga Islands,” Journal
of the Linnean Society, vol. xxxv., Botany, 1901.
CHEESEMAN, T. F., “The Flora of Rarotonga,” Transactions of the Linnean
Society, 2nd Ser., Botany, vol. vi., part 6, 1903.
DRAKE DEL CASTILLO, E., “Flore de la Polynésie Française,” Paris, 1893.
“Remarques sur la Flore de la Polynésie” (Mémoire couronné par
l’Académie des Sciences), Paris, 1890.
EGGERS, BARON H. VON, “Die Manglares in Ecuador,” Botanisches
Centralblatt, No. 41, 1892.
“Das Küstengebiet von Ecuador,” Deutsche Geographische Blätter, heft
4, band 17, Bremen, 1894.
EKSTAM, O., “Einige blütenbiologische Beobachtungen auf Novaja Semlja,”
Tromso Museums Aarshefter, 18, 1895.
“Einige blütenbiologische Beobachtungen auf Spitzbergen,” Tromso
Museums Aarshefter, 20, 1897.
GUPPY, H. B., “The Dispersal of Plants as illustrated by the Flora of
Keeling Atoll,” Journal of the Victoria Institute, London, 1889.
“The Polynesians and their Plant-Names,” Journal of the Victoria
Institute, London, 1896.
“The River Thames as an Agent in Plant-Dispersal,” Journal of the
Linnean Society, Botany, vol. xxix., 1891-93.
“River-Temperature,” part iii., Proceedings of the Royal Physical
Society of Edinburgh, 1896. (The first two parts deal principally with
the temperature of ponds and rivers, whilst in the last part the
thermal conditions are discussed especially in connection with the
life of aquatic plants.)
“On the Postponement of the Germination of the Seeds of Aquatic
Plants,” Proceedings of the Royal Physical Society of Edinburgh, 1897.
“On the Temperature of Springs as especially illustrated by the
Wandle and other Tributaries of the Thames.” (This paper, of which I
have no copy, was published in the Journal of the Royal Meteorological
Society, about 1895. It throws light on the thermal conditions of
plants in springs.)
“Water-Plants and their Ways,” _Science Gossip_, Sept., Oct., Nov.,
1894. (The various modes of dispersal of land as well as water plants
are here dealt with, their thermal conditions are discussed, and in
the November number are given the results of four years’ observations
on the life-history and life-conditions of Ceratophyllum demersum.)
“Caddis-Worms and Duckweed,” _Science Gossip_, March, 1895. (A short
note.)
“Stations of Plants and Buoyancy of Seeds,” _Science Gossip_, April
and May, 1895.
“Irregularity of some Cotyledons,” _Science Gossip_, September,
1895.
“Plants of the Black Pond, Oxshott,” _Science Gossip_, October,
1895.
“On the Habits of Lemna minor, L. gibba, and L. polyrrhiza,” Journal
of the Linnean Society, Botany, vol. xxx. (This paper contains the
results of three years’ systematic observations of these plants.)
“The Distribution of Aquatic Plants and Animals,” _The Scottish
Geographical Magazine_, January, 1893.
HEMSLEY, W. B., “Report on the Botany of the _Challenger_ Expedition,”
1885.
“The Flora of the Tonga Islands,” Journal of the Linnean Society,
Botany, vol. xxx.
HILLEBRAND, W., “Flora of the Hawaiian Islands,” Heidelberg, 1888.
HORNE, J., “A Year in Fiji,” London, 1881.
KOLPIN RAVN, F., “Om Flydeevnen hos Froene af vore Vand-og Sumpplanter,”
Botanisk Tidsskrift, 19 bind., 2 hefte, Kjobenhavn, 1894 (“On the
Floating Capacity of the Seeds of Aquatic and Marsh Plants”). (A
_résumé_ in French is appended to the paper.)
MARTINS, CH., “Expériences sur la Persistance de la Vitalité des Graines
flottant à la Surface de la Mer,” Bull. Soc. Botanique de France tome
iv., 1857.
NADEAUD, J., “Enumération des Plantes indigènes de l’Ile de Tahiti,”
Paris, 1873.
PENZIG, O., “Die Fortschritte der Flora des Krakatau,” Annales du Jardin
Botanique de Buitenzorg, 2 ser., tome 3, Leide, 1902.
PERKINS, R. C. L., “Fauna Hawaiiensis,” vol. i., part iv. (Vertebrata)
Cambridge University Press, 1903.
REINECKE, F., “Die Flora der Samoa-Inseln,” Engler’s “Botanische
Jahrbücher,” band xxv., heft v., Leipzig, 1898.
SCHIMPER, A. F. W., “Die indo-malayische Strandflora,” Jena, 1891.
SEEMANN, B., “Flora Vitiensis,” London, 1865-73.
SERNANDER, R., “Den Scandinaviska Vegetationens Spridnings-biologi,”
Upsala, 1901.
THURET, G., “Expériences sur des Graines de diverses Espèces plongées
dans de l’eau de Mer,” Archives des Sciences (Phys. et Nat.) de la
Bibliothèque Universelle, tome 47, Geneva, 1873.
TREUB, M., “Notice sur la nouvelle Flore de Krakatau,” Annales du Jardin
Botanique de Buitenzorg, 1888.
_Note._—Amongst the works quoted which are not specially particularised
in the text are Scott Elliot’s “Nature Studies,” 1902, and Beal’s
“Seed Dispersal,” Boston, 1900.
CONTENTS
PREFACE _Pages_
vii—x
LIST OF SOME OF THE PRINCIPAL AUTHORITIES QUOTED, _Pages_
WITH AN ENUMERATION OF THE AUTHOR’S BOTANICAL xiii—xv
PAPERS
LIST OF ILLUSTRATIONS _Page_
xxvii
ADDITIONS AND CORRECTIONS _Page_
xxviii
CHAPTER I
INTRODUCTION
The study of insular floras.—Their investigation in this work from the
standpoint of dispersal.—The significance of plant-distribution in the
Pacific.—The problems connected with the mountain-flora of Hawaii.—The
persistence of dispersing agencies at the coast, their partial
suspension on the mountain-top, their more or less complete suspension
in the forest, and the effect on the endemic character of plants.—The
connection between the endemism of birds and plants.—The relative
antiquity of plants of the coast, forest, and mountain-top.—The
genetic relation between coast and inland species of the same
genus.—The ethics of plant-dispersal.—Evolution takes no heed of modes
of dispersal.—The seed-stage is the price of Adaptation.
_Pages_ 1-11