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 Another Random Document on
Scribd Without Any Related Topics
 When dealing with the general character and composition of the
strand-plants of the tropical Pacific, it is shown that in Fiji the beach-
plants often assert their primary xerophilous habit or fitness for
occupying any dry station by extending into the inland plains on the
dry sides of the islands. The Fijian shore-plants are divided into
three formations, those of the beach, those of the mangrove-
swamp, and those of intermediate stations on the borders of the
swamps. The great majority of the Fijian shore-plants are dispersed
by the currents. The Tahitian Islands, which are representative of
Eastern Polynesia, lack the mangroves and most of the plants that
grow at the margin of a mangrove-swamp; and their strand-flora is
mainly composed of plants of the beach, such as are dispersed by
the currents far and wide in tropical regions. The Hawaiian strand-
flora is very meagre in its character, lacking not only the plants of
the mangrove and intermediate formations, but almost all the large-
fruited beach-trees of the South Pacific. Since Hawaii possesses but
few current-dispersed shore-plants that are not found in the New
World, reasons are given for the inference that such shore-plants
were originally brought by the currents from America, and not from
the South Pacific.
We are led on various grounds to the conclusion that tropical
shore-plants distributed by currents belong to two great regions, the
American including the west coast of Africa, and the Asiatic, or Old
World Region, which includes the African east coast. It is held that
America is so placed with regard to the currents, that it is a
distributor, and not a recipient of tropical shore-plants dispersed by
that agency. From this it follows that all cosmopolitan tropical beach-
plants that are dispersed by the currents have their homes in
America.
The results of observation and experiment are given to show that
there is no direct relation between the specific weight of seeds and
fruits and the density of sea-water. Yet, although the floating or
sinking of a seed or fruit is but an accidental attribute, it has had
indirectly a far-reaching influence not only on plant-distribution, but
on plant-development. In accordance with this want of relation
between the specific weight of seeds and fruits and the density of
sea-water, the great variety of structures concerned with buoyancy
are regarded in the main, after a detailed examination of their
character, as not arising from adaptation. Rather, it is urged, is
buoyancy connected with structures that now serve a purpose for
which they were not originally developed. Nature, it is held, has
never concerned herself directly with providing means of dispersal of
any sort.
In the discussion of the relation between the littoral and inland
Pacific floras, it is shown, as a result of the examination of those
genera possessing both shore and inland species, that they have
been on the whole developed on independent lines. Two special
difficulties in explaining the modes of dispersal of plants of the
Pacific islands here come into prominence. There is the Hawaiian
difficulty, where with genera containing both shore and inland
species only the last are found in Hawaii; and although the shore-
plants are known to be dispersed by the current, the inland plants
display little or no capacity for this or any other mode of dispersal.
Here belong the Leguminous genera Canavalia, Erythrina,
Mezoneuron, and Sophora, and the Apocynaceous genus Ochrosia;
and it is assumed that the inland Hawaiian species are derived from
a current-dispersed shore-plant that has since disappeared from the
group. The Fijian difficulty is displayed in those genera where both
coast and inland species occur in the islands, but no known existing
means of dispersal across an ocean can be postulated for the inland
plants, though the shore species are distributed by the currents. Of
such genera Pandanus is the best example, and it is pointed out that
this genus presents the same difficulty in the Mascarene Islands, in
which case the agency of the extinct Columbæ is invoked.
As illustrating the methods of observation and experiment
employed by the author, the Leguminous shore-plants Afzelia bijuga,
Cæsalpinia bonducella, and Entada scandens are discussed at
length; and in the chapters on the enigmas of the Leguminosæ in
the Pacific it is pointed out that the behaviour of the plants of this
order is a source of much perplexity, and that they conform to no
single rule of dispersal.
Coming to the inland plants of this region, the Fijian, Tahitian, and
Hawaiian groups are taken as the chief centres of distribution in the
Pacific. After discussing the relative sizes, the altitudes, and the
climates of these three archipelagoes, it is shown that Hawaii, on
account of the far greater altitude of the islands, is characterised by
a special mountain flora, and that it is comparable with Fiji, and to a
great extent also with Tahiti, only as regarding the plants of the
levels below 4,000 or 5,000 feet.
The first era of the plant-stocking is designated the Age of Ferns,
and it is observed that, whilst in Hawaii nearly half of the ferns and
lycopods are peculiar to that group, very few new species have been
developed in the Fijian and Tahitian regions.
The next era in the floral history of these islands is represented in
the first era of the flowering plants. This is indicated by the endemic
genera, which are particularly numerous in Hawaii, relatively scanty
in Fiji, and very few in Tahiti. On account of their preponderance, the
era is designated the Age of Compositæ and Lobeliaceæ. The genera
of these two orders, though mainly characteristic of Hawaii, are also
to be found in the Tahitian region, but they are absent from the
Fijian area. Chiefly American in their affinities, their dispersion over
the Pacific took place during the Tertiary submergence of the
archipelagoes of the Western Pacific, in which are included the
groups of the Fijian area (Fiji, Samoa, Tonga). These early forms of
Compositæ and Lobeliaceæ are often arborescent in habit; and it is
observed that Tree-Lobelias also occur high up the slopes of lofty
mountains in tropical regions, as in Equatorial Africa, under
conditions similar to those prevailing on the slopes of the Hawaiian
mountains, where the Tree-Lobelias, termed by Dr. Hillebrand “the
pride of our flora,” abound.
The other Hawaiian endemic genera, marking the first chapter in
the history of the flowering plants, arrange themselves in two
groups, one chiefly American in general affinities, and containing
highly differentiated Caryophyllaceæ, Labiatæ, &c.; the other largely
Malayan, and indicating the close of the first era of the flowering
plants, when the main source of the plants was shifted from America
to the Old World. The Fijian endemic genera, which are few in
number, miscellaneous in appearance, and disconnected in character,
are regarded as having probably acquired their endemic reputation
through their failure at their sources in the regions to the west.
The second era of the flowering plants is indicated by the non-
endemic genera. Here we are concerned on the one hand with a
mountainous flora mainly Hawaiian, in which genera from the New
Zealand and Antarctic floras take a conspicuous part, and on the
other with a low-level flora chiefly derived from Indo-Malaya, and
including the plants of the lower slopes of Hawaii below 4,000 and
5,000 feet, and the floras in mass of Fiji and Tahiti.
On account of their lower altitude, the extensive mountain flora of
Hawaii is but scantily developed in Tahiti, and is represented by a
mere remnant in Fiji and Samoa. Two-thirds of the Hawaiian non-
endemic mountain genera contain only species restricted to the
group, and, although amongst these disconnected genera, Acæna,
Gunnera, Coprosma, Lagenophora, &c., of the New Zealand and
Antarctic floras take a prominent part, a large proportion of the
genera like Ranunculus, Rubus, Artemisia, Vaccinium, and Plantago
represent generally the flora of the north temperate zone on the
summits of tropical mountains. The Tahitian mountain flora, scanty
as it is when judged by the non-endemic genera, displays much
kinship with the Hawaiian mountain flora; but this kinship is mainly
confined to genera from high southern latitudes, such as Coprosma,
Cyathodes, Astelia, &c. In the possession on its mountain slopes of
the three genera of the Coniferæ, Dammara, Podocarpus, and
Dacrydium, the Fijian region is distinguished from that of Tahiti and
Hawaii; and it is assumed that they mark the site of a continental
area in the Mesozoic period, when the Tahitian and Hawaiian groups
did not exist.
The era of the non-endemic genera, in so far as it is concerned
with the low-level flora of Hawaii and the floras in mass of the areas
of Fiji, Samoa and East Polynesia, is termed Malayan, because many
of the genera are thence derived. Here we are dealing with all the
oceanic groups of the tropical Pacific, and not with a portion of
them, as in the case of the Age of Coniferæ, in the Secondary
period, that was limited to the Western Pacific, or in the case of the
Age of Compositæ and Lobeliaceæ that was restricted during the
Tertiary epoch to the Hawaiian and Tahitian regions. The first part of
this era, as is indicated by the endemic species, is an age of
complete isolation in Hawaii, and of partial isolation in the groups of
the southern region. Amongst the genera typical of this period are
Pittosporum, Gardenia, Psychotria, Cyrtandra, and Freycinetia. A
later period in this era of the general dispersal of Malayan plants
over the Pacific is one where the extremely variable or polymorphous
species plays a conspicuous part, as represented in such genera as
Alphitonia, Dodonæa, Metrosideros, Pisonia, and Wikstrœmia, the
general principle being that each genus is at first represented by a
widely ranging very variable species, which ultimately ceases to
wander and settles down, and becomes the parent of different sets
of species in the several groups.
The facts of distribution in this age of general dispersion are just
such as we might look for in the case of a general dispersal over the
oceanic groups of the Pacific, with the altitudes of the islands playing
a determining part. But it should be remarked that the greater
number of the genera that have entered the Pacific from the Old
World have not advanced eastward of the Fijian region, half of the
Fijian genera not occurring in the Hawaiian and Tahitian regions. The
explanation of this is to be found, not in any lack of capacities for
dispersal, but in a want of opportunities. The story of plant-
distribution in the Pacific is bound up with the successive stages of
decreasing activity in the dispersing agencies. The area of active
dispersion, as illustrated by the non-endemic genera, at first
comprised the whole of the tropical Pacific. It was afterwards
restricted to the South Pacific, and finally to the Western Pacific only.
The birds that carried seeds all over this ocean became more and
more restricted in their ranges, probably on account of increasing
diversity of climatic conditions. The plants of necessity responded to
the ever narrowing conditions of bird-life in this ocean, and the
differentiation of the plant and the bird have taken place together.
During the stages of decreasing activity in the dispersing agencies,
the widely-ranging highly variable species continued to be an
important factor in the development of new species in the different
groups. The rôle of the polymorphous species has always been a
conspicuous one in the Pacific.
Yet, as in the case of the Cyrtandras, it is shown that the display
of great formative power within a genus is not a peculiarity of an
insular flora; that the isolation of an oceanic archipelago does not
exclusively induce “endemism,” but only intensifies it; that the
development of new species may be nearly as active on a mountain
in a continent as on an island in mid-ocean; and that this is equally
true of a land genus, like Embelia, exposed to an infinite variety of
conditions, and of an aquatic genus, like Naias, where the conditions
of existence are relatively uniform all the world over.
In framing a scheme by which the eras of the floral history of the
Pacific are brought into correlation with those of geological time, the
age of the Coniferæ is placed in the Secondary period, that of the
Compositæ and Lobeliaceæ in the Tertiary period, whilst the era of
Malayan immigration is regarded as mainly post-glacial. The age of
the Coniferæ is concerned only with the Western Pacific, since the
Hawaiian and Tahitian islands had not then been formed. The age of
the Compositæ and Lobeliaceæ is concerned only with Hawaii and
Tahiti, since the islands of the Western Pacific were then more or
less submerged. That of the Malayan plants affects the whole Pacific
as at present displayed to us.
In the chapter on the viviparous mangroves of Fiji it is shown that
both the Asiatic and the American species of Rhizophora (R.
mucronata and R. mangle) exist in that group, and that there is in
addition a seedless form, the Selala, which, although intermediate in
character between the two other species, comes nearest to the
Asiatic plant. Reasons are given for the belief that the Selala is
derived from the Asiatic species (R. mucronata), not as the result of
a cross but as connected with its dimorphism; and in support of this
it is pointed out that on the Ecuador coast of South America, where
only the American species exists, a dimorphism is also displayed,
one of the forms approaching in several of its characters the Fijian
Selala, though fruiting abundantly and bearing the impress of a
closer connection with the typical American species than with the
Asiatic plant. The view that Rhizophora mangle reached the Western
Pacific from America is rejected, and it is considered that this species
was originally as widely diffused in the Old World as in America, and
that it now survives only in a few places in the tropics of the Old
World. The results of detailed observations on the modes of
dispersal and on the germinating process both with Rhizophora and
Bruguiera are given; and the absence, as a general rule, of any
period of rest between the fecundation of the ovule and the
germination of the seed is established.
A special chapter is devoted to the significance of vivipary, and it
is considered that a record of the history of vivipary on the globe is
afforded in the scale of germinative capacity that begins with the
seedling hanging from a mangrove and ends with the seed that is
detached in an immature condition from an inland plant. It is
suggested that with the drying up of the planet in the course of ages
the viviparous habit, which was once nearly universal, has been for
the most part lost except in the mangrove swamp, which to some
extent represents an age when the earth was enveloped in cloud
and mist and the atmosphere was saturated with aqueous vapour.
The lost habit is at times revived in the abnormal vivipary of some
inland plants, and traces of it are seen in the abnormal structure of
the seeds of some genera of the Myrtaceæ, like Barringtonia, and in
the seeds of genera of other orders. With the desiccation of the
planet and the emergence of the continents there has been
continual differentiation of climate resulting in seasonal variation and
in the development of the rest-period of the seed.
With the secular drying of the globe and the consequent
differentiation of climate is to be connected the suspension to a
great extent of the agency of birds as plant-dispersers in later ages,
not only in the Pacific Islands but over all the tropics. The changes
of climate, bird, and plant have gone on together, the range of the
bird being controlled by the climate, and the distribution of the plant
being largely dependent on the bird.
The history of climate, the history of the continents and of the
oceans, the history of life itself, but only in the sense below defined,
all belong to that of a desiccating world, or rather of a planet once
sunless and enveloped in mist and cloud, that through the ages has
been drying up. Life’s types were few and the sea prevailed, and one
climate reigned over the globe. With the diminution of the aqueous
envelopes the continents began to emerge, climates began to
individualise, and organisms commenced to differentiate, and thus
the process has run on through the past, ever from the general to
the special both in the organic and in the inorganic world.
The same story of a world drying up is told by the marine remains
left stranded far up some mountain slope, or by the bird akin to no
other of its kind that Time has stranded on some island in mid-
Pacific. The bird generalised in type that once ranged the globe is
now represented over its original range by a hundred different
groups of descendants, confined each to its own locality. Climate,
once so uniform, now so diversified, has by restricting the range of
the bird favoured the process of differentiation, and the plant
dependent on the bird for its distribution has in its turn responded to
these changes.
The rôle of the polymorphous species belongs alike to the plant
and to the bird. A species that covers the range of a genus varies at
first in every region and ultimately gives birth to new species in
some parts of its range. Then the wide-ranging species disappears
and the original area is divided up into a number of smaller areas
each with its own group of species. Each smaller area breaks up
again, and forms, yet more specialised, are produced; and thus the
process of subdivision of range and of differentiation of form goes
along until each island in an archipelago owns its bird and each hill
and valley has its separate plants. This is not the path that Evolution
takes, since beyond lies extinction whether of plant or of bird. Such
is the upshot of the process of differentiation exhibited in the
development of species and genera in the Pacific Islands, or, indeed,
in any oceanic groups. It can never do more than produce a Dodo or
a Kiwi, or amongst the plants a Tree-Lobelia.
Evolution here and elsewhere is a thing apart from species and
genera, which are but eddies on the surface of its stream. It is a
scheme of life introduced into a much conditioned world, and
adaptation in endless forms is the price it has had to pay. The whole
story of life on this earth is a story of a sacrifice, of an end to be
won, but of a price to be paid. Immortality is in the scheme, but
death is the price of adaptation. The same theme runs through our
conceptions of the spiritual life. There is the same duality, evolution
adapting its scheme to the exigencies of the physical world, the
good principle ever in conflict with the evil, and at times compelled
to adapt itself to attain its ends. There is the tale of adaptation in
the one case and of sacrifice in the other, and success is reached in
both.
 APPENDIX

List of Notes

Note 1. On the number of known species of Fijian flowering plants.

Note 2. The littoral plants of Fiji.
Note 3. Results of long flotation experiments on the seeds or
seedvessels of tropical littoral plants.
Note 4. Table illustrating the degree of buoyancy of the seeds and fruits
of inland Fijian plants.
Note 5. The inland Fijian plants possessing buoyant seeds or fruits.
Note 6. Table showing the degree of buoyancy of the seeds and fruits of
some inland Hawaiian plants.
Note 7. Some inland Hawaiian plants possessing buoyant seeds or fruits.
Note 8. The pyrenes of Morinda.
Note 9. The buoyancy of the fruits of Calophyllum.
Note 10. The buoyancy experiments on British plants.
Note 11. The effect of sea-water immersion on the germinating capacity
of seeds and seed-vessels.
Note 12. The buoyancy of the fruits of Galium aparine.
Note 13. The buoyancy of the seeds of Convolvulus sepium.
Note 14. Other long flotation experiments.
Note 15. The occurrence inland of Silene maritima.
Note 16. The buoyancy of the seeds or fruits of the British beach-plants
that also occur inland.
Note 17. The buoyancy of the seeds or fruits of the British littoral plants
that frequent salt-marshes and muddy shores.
Note 18. The buoyancy of the seeds or fruits of the British littoral plants
that are confined to the beach.
Note 19. On germination in sea-water.
Note 20. On the maximum heights reached by some shore plants in
their extension inland in Vanua Levu, Fiji.
Note 21. On the dwarfing of shore plants when extending inland in the
“talasinga” plains in Vanua Levu.
Note 22. The “talasinga” plains of Vanua Levu.
Note 23. Schimper’s grouping of the Indo-Malayan strand flora.
Note 24. Grouping of some of the characteristic plants of the strand
flora of Fiji.
Note 25. The strand flora of the Tahitian region.
Note 26. The Fijian shore plants not found in Tahiti.
Note 27. The intruders into the beach flora from the inland plants of
Tahiti.
Note 28. The littoral plants of the Hawaiian islands.
Note 29. Botanical notes on the coast plants of the Hawaiian islands.
Note 30. The beach drift of the Hawaiian islands.
Note 31. The inland extension of the shore plants of the Hawaiian
islands.
Note 32. The Fijian species of Premna.
Note 33. De Candolle’s list of plants dispersed exclusively by currents.
Note 34. The littoral plants of the eastern-most Polynesian islands.
Note 35. Distribution of the littoral plants with buoyant seeds or fruits
that occur in the Fijian, Tongan, Samoan, Tahitian, and Hawaiian
Groups.
Note 36. Hawaiian plants with buoyant seeds or fruits known to be
dispersed by the currents either exclusively or with the assistance of
frugivorous birds.
Note 37. On vivipary in the fruits of Barringtonia racemosa and Carapa
obovata.
Note 38. On the temperature and density of the surface water of the
estuaries of the Rewa River in Fiji and of the Guayaquil River in
Ecuador.
Note 39. On the Pacific species of Strongylodon.
Note 40. Precautions in testing seed-buoyancy.
Note 41. The buoyancy of the seeds of Convolvulus soldanella in fresh-
water and sea-water compared.
Note 42. On secular changes in sea-density.
Note 43. On the mucosity of small seeds and seed-like fruits when wet.
Note 44. Upon the effects of inland extension on the buoyancy of the
seeds or fruits of littoral plants.
Note 45. Tabulated results of the classification, according to Schimper’s
application of the Natural Selection Theory, of the buoyant seeds and
fruits of tropical littoral plants.
Note 46. On the modes of dispersal of the genus Brackenridgea.
Note 47. On the transport of gourds by currents.
Note 48. On the useless dispersal by currents of the fruits of the Oak
and of other species of Quercus, as well as of the Hazel (Corylus).
Note 49. On the distribution of Ipomœa pes capræ, Convolvulus
soldanella, and Convolvulus sepium.
Note 50. On the structure of the seeds and fruits of Barringtonia.
Note 51. On a common inland species of Scævola in Vanua Levu, Fiji.
Note 52. On the capacity for dispersal by currents of Colubrina
oppositifolia.
Note 53. On the genus Erythrina.
Note 54. On the genus Canavalia.
Note 55. The inland extension of Scævola kœnigii.
Note 56. On the capacity for dispersal by currents of Sophora
tomentosa, S. chrysophylla, and S. tetraptera.
Note 57. On the species of Ochrosia.
Note 58. On Pandanus.
Note 59. Seeds in petrels.
Note 60. Schimper on the halophilous character of littoral Leguminosæ
and of shore plants generally.
Note 61. Meteorological observations on the summit of Mauna Loa.
Note 62. On the relative proportion of vascular cryptogams in Fiji.
Note 63. On the table of vascular cryptogams of Tahiti, Hawaii, and Fiji.
Note 64. On the distribution of the Tahitian ferns and lycopods.
Note 65. Distribution of some of the mountain ferns of Hawaii that are
not found either in Fiji or in Tahiti.
Note 66. Endemic genera of ferns in Hawaii.
Note 67. On the dispersal of Compositæ by birds.
Note 68. On some of the Hawaiian endemic genera excluding those of
the Compositæ and Lobeliaceæ.
Note 69. On the germination of Cuscuta.
Note 70. On beach-temperature.
Note 71. On the buoyancy of the seeds or seed-vessels of some Chilian
shore plants.
Note 72. On the southern limit of the mangrove formation in Ecuador.
Note 73. Additional note on the temperature of the dry coast of
Ecuador, between the island of Puna and the equator.
Note 74. Observations on the temperature of the Humboldt current from
Antofagasta northward between January and March, 1904.
Note 75. On the stranded massive corals of the genus Porites (?) found
on the coast of North Chile and Peru at Arica, Callao, and Ancon.
Note 76. Stranded pumice on English and Scandinavian beaches.
Note 77. On the mode of dispersal of Kleinhovia hospita.
Note 78. On the “Sea”, an unidentified wild fruit tree in Fiji.
Note 79. On willow-leaved river-side plants.
Note 80. Mr. Perkins on the Hawaiian Lobeliaceæ.
Note 81. On the vertical range of some of the most typical and most
conspicuous of the plants in the forests on the Hamakua slopes of
Mauna Kea, Hawaii.
Note 82. Aboriginal weeds.
Note 90. On the buoyancy of the seeds of Euphorbia amygdaloides and
E. segetalis.
Note 91. Mr. E. Kay Robinson on Aster tripolium.
 NOTE 1 (page 13)

On the Number of Known Species of Fijian Flowering Plants

Rather over 600 species of flowering plants are included in Seemann’s
Flora Vitiensis, excluding the weeds and the plants introduced by man.
Horne’s collections would probably add another 300 species; and many
more remain to be discovered.

NOTE 2 (page 13)

The Littoral Plants of Fiji

In the following table are incorporated the results of an extensive
series of observations and experiments on the buoyancy of the seeds
and fruits of the shore plants made by the author during his sojourn of
two years in Fiji, and based not only on prolonged buoyancy-tests, but
also on systematic examination of the stranded and floating seed-drift,
both of sea and river. The details would occupy many chapters: and it is
only possible here to give the bare results. Since Professor Schimper
went over much the same ground in the Malayan region, one enjoys in
many cases the great advantage of his authority; but a fair proportion
of the results are new; and, besides, there are a number of plants
included, the buoyancy of whose seeds or fruits has long been well
established. In all cases the seed or fruit is taken as it presents itself for
dispersal by the currents. Many of the plants are discussed with some
detail in various parts of this book, as indicated in the reference column
of the table.
Since the Gramineæ and the Cyperaceæ contain very few species
suited for direct transport by the currents over wide areas of sea, this
list may be regarded as containing nearly all the littoral flowering plants
possessing seeds or seed-vessels with any buoyancy of importance.
Nearly all the Tahitian strictly littoral plants are represented in Fiji, and
the few that have not been found there yet, such as Sesbania
grandiflora, Heliotropium anomalum, &c., may exist, as in the first-
named species, in the neighbouring Tongan group, and may probably
even exist in Fiji. Two other Tahitian littoral plants, that are widely
spread in the Pacific, namely, Suriana maritima and Sesuvium
Portulacastrum, are found in Tonga, and are included in my list of Fijian
shore plants, though not yet recorded from that group, where, however,
they will, without a doubt, be found by some future observer.
Table showing the Buoyancy of the Seeds or Fruits of the Littoral Plants of
Fiji, excluding the Grasses and, with one exception, the Sedges
The letters placed before the plant name indicate that the species is
also found in Hawaii (H), in Tahiti (T), and in the Marquesas (M). The
Marquesan locality is only given where the plant is not in Tahiti.
The abbreviations in the reference column are as follows:
S=Schimper; G=Guppy; P=Earlier authorities and particularly the list
given by Hemsley in the Introduction to the Botany of the Challenger
Expedition.
Buoyancy of
seeds or
fruits.
Sink Pages of
at further
Species. Family. once Authorities. reference.
Float
or in See also
for
a Index.
months.
week
or
two.
HT
Calophyllum Guttiferæ. + ... S.G.P. 18
inophyllum

HT Hibiscus
Malvaceæ. + ... S.G.P. 21
tiliaceus

Hibiscus Malvaceæ. + ... G. 21

diversifolius
(Jacq.)

HT Thespesia
Malvaceæ. + ... S.G.P. Note 3
populnea

H Gossypium
tomentosum Malvaceæ. ... + G.
(Nutt.)

Heritiera
Sterculiaceæ. + ... S.G.P. 45, 48
littoralis

T Kleinhovia
Sterculiaceæ. + ... G. 21
hospita

T Triumfetta
Tiliaceæ. ... +?
rhomboidea

T Triumfetta
Tiliaceæ. ... + G. 45
procumbens

T Suriana
Simarubeæ. + ... S.G.
maritima

Carapa
Meliaceæ. + ... S.G.P. 45
moluccensis

Carapa
Meliaceæ. + ... S.G.P. 45
obovata

T Ximenia
Olacineæ. + ... S.G. 113
americana

Smythea
pacifica Rhamneæ. + ... G.P. 106
(Seem.)

HT Colubrina
Rhamneæ. + ... G. 137
asiatica
HT Dodonæa
Sapindaceæ. + ... S.G.
viscosa

HT Tephrosia
Papilionaceæ. ... + G. 45
piscatoria

M Desmodium
Papilionaceæ. ... + G.
umbellatum

HT Dioclea
Papilionaceæ. + ... G.P. 82
violacea

T Canavalia
Papilionaceæ. + ... S.G.P. Note 54
obtusifolia

T Canavalia
Papilionaceæ. + ... G. Note 54
sericea

T Canavalia
ensiformis, var. Papilionaceæ. + ... S.G.P.? Note 54
turgida.

HT Mucuna
Papilionaceæ. + ... S.G.P. 81
gigantea

T Erythrina
Papilionaceæ. + ... S.G.P.
indica

HT
Strongylodon Papilionaceæ. + ... G. 82
lucidum

HT Vigna lutea Papilionaceæ. + ... S.G. 139

Dalbergia
Papilionaceæ. + ... S.G. 106
monosperma

Derris Papilionaceæ. + ... S.G.P. 111

uliginosa

Pongamia
Papilionaceæ. + ... S.G.P.
glabra

T Sophora
Papilionaceæ. + ... S.G. Note 56
tomentosa

T Inocarpus
Papilionaceæ. +? ... G.P.
edulis

HT Cæsalpinia
Cæsalpinieæ. + ... S.G.P. 193
Bonducella

T Cæsalpinia
Cæsalpinieæ. + ... G.P. 193
Bonduc

Afzelia bijuga Cæsalpinieæ. + ... G. 173

Cynometra sp. Cæsalpinieæ. +? ... S.G.

Entada
Mimoseæ. + ... G.P. 181
scandens

Acacia
Mimoseæ. ... + G. 164
laurifolia

T Leucæna
Mimoseæ. ... + G.
Forsteri

T Serianthes
Mimoseæ. ... + G. 424
myriadenia

Parinarium
Rosaceæ. + ... G.P.
laurinum

Eugenia Richii Myrtaceæ. ... + G.

T Barringtonia Myrtaceæ. + ... S.G.P.

speciosa

Barringtonia
Myrtaceæ. + ... G.
racemosa

Rhizophora
Rhizophoreæ. + ... S.G.P.
mucronata

Rhizophora
Rhizophoreæ. + ... S.G.P.
mangle

Bruguiera
Rhizophoreæ. + ... G.P. &nbsp
Rheedii

HT Terminalia
Combretaceæ. + ... S.G.P.
Katappa

M Terminalia
Combretaceæ. + ... S.G.P.
littoralis

Lumnitzera
Combretaceæ. + ... S.G.P.
coccinea

T Gyrocarpus
Combretaceæ. + ... G. 423
Jacquini

T Pemphis
Lythraceæ. + ... S.G.
acidula

T Luffa
insularum Cucurbitaceæ. + ... G. 426
(Gray)

HT Sesuvium
Ficoideæ. ... + G.
Portulacastrum

HT Morinda
Rubiaceæ. + ... S.G.P.
citrifolia
T Guettarda
Rubiaceæ. + ... S.G.P.
speciosa

T Wedelia
Compositæ. + ... G.
biflora

HT Scævola
Goodeniaceæ. + ... S.G.P.
Koenigii

Table showing the Buoyancy of the Seeds or Fruits of the

Littoral Plants of Fiji, excluding the Grasses, and with
one exception, the Sedges (continued)

Buoyancy of
seeds or
fruits.
Sink Pages of
at further
Species. Family once Authorities. reference.
Float
or in See also
for
a Index.
months.
week
or
two.
T Cerbera
Apocynaceæ. + ... S.G.P.
Odollam

T Ochrosia
Apocynaceæ. + ... G.P.
parviflora

HT Cordia
Boraginaceæ. + ... S.G.P.
subcordata

T Tournefortia
Boraginaceæ. + ... S.G.P.
argentea

HT Ipomœa Convolvulaceæ. + ... S.G.P.

pes capræ

H Ipomœa
glaberrima Convolvulaceæ. + ... G.
(Boj.)

Aniseia uniflora Convolvulaceæ. + ... G.

T Premna
Verbenaceæ. + ... G. Note 32
tahitensis

Clerodendron
Verbenaceæ. + ... S.G.
inerme

HM Vitex
Verbenaceæ. + ... G.
trifolia

HT Cassytha
Lauraceæ. + ... G.
filiformis

T Hernandia
Lauraceæ. + ... S.G.
peltata

HT
Wikstrœmia Thymelæaceæ. ... + G.
fœtida

Drymispermum
Thymelæaceæ. ... + G.
Burnettianum

T Euphorbia
Euphorbiaceæ. ... + S.P.G.
Atoto

Excæcaria
Euphorbiaceæ. + ... S.G.
Agallocha

T Casuarina
Casuarineæ. ... + G.
equisetifolia
HT Tacca
Taccaceæ. + ... G. 19
pinnatifida

HT Cocos
Palmeæ. + ... P.
nucifera

HT Pandanus
Pandaneæ. + ... S.G.P.
odoratissimus

Crinum
Amaryllideæ. ... + G.P.
asiaticum

Scirpodendron
Cyperaceæ. + ... G. 407
costatum

Cycas circinalis Cycadeæ. + ... G.P.

NOTE 3 (page 13)

Results of Long Flotation Experiments on the Seeds or Seed-

vessels of Tropical Littoral Plants

At various times during the past twenty years I have made

lengthened experiments in England on the buoyancy in sea-water of the
seeds or seed-vessels of beach plants collected by me in the Solomon
Islands, the Fijis, Hawaii, Keeling Atoll, &c. In all the species
enumerated below, the floating powers were retained after twelve
months’ immersion, the seed-contents being to all appearance
unharmed. In six species I succeeded in getting the seeds to germinate
after the experiment; and there can be no doubt that the number of
successful results would have been largely increased, if I had not been
obliged to resort to very primitive methods in conducting the
experiments. Some of the results are referred to in a note to my paper
on the flora of Keeling Atoll, dated about 1889; and if I remember
aright, Mr. Hemsley mentioned those relating to Thespesia populnea
and Ipomœa grandiflora in the Annals of Botany, not long after. The
others have not been previously published. In one instance (Cæsalpinia
bonducella) the flotation experiment was prolonged to two and a half
years, the seeds floating buoyantly and being apparently quite sound at
the end of the experiment.
As demonstrating that tropical seeds can be transported unharmed by
currents through cold latitudes, it should be noted that all these
experiments were conducted in England. In the cases of the Keeling
Atoll seeds the experiment was carried on through a very severe winter,
the vessel of sea-water being exposed to a degree of cold that kept
fresh-water frozen for three weeks on the same table. This did not
prevent the subsequent germination of the seeds of Thespesia populnea
and Ipomœa grandiflora. The same thing was established in a more
natural way by Lindman, who planted seeds of Entada scandens and
Mucuna urens, that had been stranded on the Norwegian coast, and
found that they retained their germinating capacity (see Sernander, p.
7).
The following are the seeds or seed-vessels that remained afloat after
a year’s flotation in sea-water, those that subsequently germinated
being preceded by G. In the other cases the germinating capacity was
not tested; but they were always sound in appearance when cut across
at the close of the experiment.
G Thespesia populnea (Malvaceæ)
Dioclea (violacea?) (Papilionaceæ)
G Mucuna gigantea, D C (Papilionaceæ)
G Mucuna urens, D C (Papilionaceæ)
Mucuna, sp. (Papilionaceæ)
Mucuna, sp. (Papilionaceæ)
G Strongylodon lucidum, Seem. (Papilionaceæ)
Sophora tomentosa, (Papilionaceæ)
G Cæsalpinia bonducella (Cæsalpinieæ)
Entada scandens (Mimoseæ)
Morinda citrifolia (Rubiaceæ)
Scævola Koenigii (Goodeniaceæ)
Cordia subcordata (Boragineæ)
Tournefortia argentea (Boragineæ)
G Ipomœa grandiflora, Lam. (Convolvulaceæ)
 Tacca pinnatifida (Taccaceæ)

NOTE 4 (page 13)

Table illustrating the Degree of Buoyancy of the Seeds and

Fruits of Inland Fijian Plants

(Unless otherwise indicated, the seeds or fruits sink at once or in a day

or two)
Abrus precatorius.
Acacia Richii.
Ageratum conyzoides.
Alphitonia excelsa.
Alpinia sp.
Alyxia (scandens?).
Artocarpus incisa.
Artocarpus integrifolia.
Barringtonia edulis (1 month)
Barringtonia sp.
Bauhinia sp.
Bischoffia javanica.
Cæsalpinia sp.
Calophyllum spectabile (2-4 weeks).
Calophyllum Burmanni (4-10 days).
Cananga odorata.
Canarium sp.
Canarium sp.
Canna indica.
Citrus aurantium (3-4 weeks).
Citrus decumana (1 month).
Citrus limonum (5 weeks).
Citrus vulgaris, R. (6-7 weeks).
Coix lachryma (2-7 days).
Commersonia platyphylla.
Cordyline sepiaria.
Couthovia corynocarpa (a few days).
Cucumis acidus (a few days).
Cucurbita sp. (several months).
Cupania sp.
Dammara vitiensis (7-10 days).
Dioscorea sativa (a few days).
Dioscorea sp.
Dracontomelon sylvestre.
Dracontomelon sp.
Elæocarpus sp.
Elæocarpus sp. (a few days).
Eranthemum sp.
Eugenia malaccensis (2-4 weeks).
Eugenia effusa? (4-7 days).
Eugenia confertiflora? (10-12 days).
Eugenia rariflora (a few days).
Eugenia corynocarpa (a few days).
Eugenia rivularis (a week).
Fagræa Berteriana (a few days).
Ficus Harveyi (7-10 days).
Ficus scabra (7-10 days).
Ficus sp. (7-10 days).
Gardenia vitiensis (4-5 weeks).
Geissois ternata.
Geophila reniformis.
Gnetum gnemon.
Grewia sp.
Guettarda sp. (a few weeks).
Hibiscus Abelmoschus (months).
Hibiscus seculentus.
Hydrocotyle asiatica (months).
Ipomœa batatas.
Ipomœa insularis (nil or months).
Ipomœa peltata (weeks or months).
Ipomœa turpethum (nil or weeks or months).
Ipomœa sp. (7-10 days).
Lindenia vitiensis (weeks or months).
Maba sp. (7-10 days).
Macaranga sp. (1-2 weeks).
Melastoma denticulatum.
Micromelum minutum.
Momordica Charantia (a few days).
Morinda Forsteri.
Mussænda frondosa.
Myristica sp. (3-7 days)
Myristica sp. (3-7 days)
Myrmecodia sp.
Nelitris vitiensis (a few days).
Nephelium pinnatum (a few days).
Ophiorrhiza leptantha.
Phyllanthus sp.
Phyllanthus sp.
Piper Macgillivrayi.
Pittosporum sp.
Pleiosmilax vitiensis.
Portulaca (lutea?).
Portulaca quadrifida.
Premna serratifolia.
Pritchardia pacifica.
Psychotria sp.
Psychotria sp.
Psychotria sp.
Psychotria sp.
Psychotria sp.
Ptychosperma sp.
Rhaphidophora vitiensis.
Sapota sp. (a few days)
Sapota sp. (a few days)
Scævola floribunda.
Spondias dulcis (a month).
Sterculia sp. (seeds nil, fruits months).
Stylocoryne sambucina (2 or 3 days).
Tabernæmontana (orientalis?) (a few days).
Tacca maculata (nil or a few days).
Trichospermum Richii (a few days).
Urena lobata.
Veitchia Joannis.
Veitchia sp.

NOTE 5 (page 14).

The Inland Fijian Plants possessing Buoyant Seeds or Fruits

They come under the following heads:
(a) Plants of the stream-border or the pond-side or of the inland
swamp, e.g., Lindenia vitiensis and Hydrocotyle asiatica. The extension
of the principle by which plants with buoyant seeds or fruits are located,
not only at the sea-side but at the water-side generally, is here involved,
as explained in Chapter III.
(b) Plants following the rule deduced by Schimper for Terminalia, that
when a genus comprises several species possessing buoyant fruits, only
those having fruits with the greatest floating power are found at the
coast, the least buoyant plants occurring inland; examples, Calophyllum
and Guettarda.
(c) Plants that like Ipomœa behave irregularly in respect to seed-
buoyancy, a difference in behaviour often associated with varying
stations both at the coast and inland.
(d) Plants with dehiscent buoyant capsular fruits, like Sterculia, where
dehiscence takes place on the tree and the seeds have no buoyancy.
Although the unopened fruit may float a long time, it does not in that
condition come under the influence of the currents.
(e) Plants like Citrus Decumana, Gardenia, sp., &c., that, although
apparently exceptions to the principle, do not offer much opposition to
it, since the first is most at home at the river-side and the second often
displays a decided inclination for a station at the coast.
(f) Genuine exceptions to the principle, such as Hibiscus Abelmoschus
(see page 21).
 NOTE 6 (page 15)

Table showing the Degree of Buoyancy of the Seeds and

Fruits of some Inland Hawaiian Plants

(Unless otherwise stated, the seeds or fruits sink at once or in a day or

two)
Acacia Koa.
Aleurites moluccana (1-2 weeks).
Alyxia olivæformis.
Argemone mexicana.
Argyreia tiliæfolia (nil or months).
Bidens pilosa.
Campylotheca sp.
Canavalia galeata.
Capparis sandwicensis.
Cassia Gaudichaudii.
Cassia occidentalis.
Cheirodendron Gaudichaudii.
Colubrina oppositifolia (weeks).
Commelina nudiflora.
Coprosma ernodeoides.
Coprosma sp.
Coprosma sp.
Cyathodes Tameiameiæ (a few days).
Cyrtandra sp. (a few days).
Cyrtandra sp. (a few days).
Cyrtandra sp. (a few days).
Dianella odorata (a few days).
Dracæna aurea.
Eclipta alba (months).
Erythrina monosperma.
Gossypium tomentosum (a week).
Gossypium barbadense (a few days).
Gossypium sp. cultiv. (a few days).
Hibiscus Youngianus (weeks).
Hydrocotyle verticillata (weeks).
Ipomœa bona nox (nil or months).
Ipomœa insularis.
Ipomœa pentaphylla.
Ipomœa reptans.
Ipomœa tuberculata.
Jacquemontia sandwicensis.
Jussiæa villosa (a few days).
Lobeliaceæ (Clermontia).
Maba sandwicensis.
Metrosideros polymorpha.
Mezoneuron kauaiense (pod, a week).
Mucuna urens (months).
Myoporum sandwicense.
Olea sandwicensis, see page 364.
Phyllostegia grandiflora.
Phyllostegia mollis.
Plectronia odorata.
Pritchardia Gaudichaudii (5 or 6 weeks).
Ricinus communis (7-10 days).
Rubus Macraei.
Scævola Chamissoniana.
Scævola Gaudichaudii.
Sida fallax.
Sisyrinchium acre.
Solanum aculeatissimum.
Sophora chrysophylla (pod, 1-2 weeks).
Viola Chamissoniana.
Waltheria americana.

NOTE 7 (page 15)

Some Inland Hawaiian Plants possessing Buoyant Seeds or

Fruits
 Three of these, Eclipta alba, Hibiscus Youngianus, and Hydrocotyle
verticillata, frequent wet places, and come under the principle that
water-side plants generally have buoyant seeds or fruits. The buoyancy
of the seeds of Argyreia tiliæfolia and of Ipomœa bona nox varies with
station and may be explained as under Ipomœa in Note 5. The floating
power of the fruits of Colubrina oppositifolia may be akin to that of
inland species of Terminalia as indicated in Note 5, since another
species of the genus C. asiatica, which is a coast plant, has very
buoyant seeds. Mucuna urens was no doubt originally, as it now is in
tropical America, a littoral plant. The buoyant fruits of Pritchardia
Gaudichaudii offer a genuine exception to the principle (see page 330).

NOTE 8 (pages 18, 112)

The Pyrenes of Morinda

The pyrenes of the two Malayan inland species of Morinda (M.
umbellata and M. longiflora) examined by Professor Schimper do not
possess the bladder-like cavity to which those of M. citrifolia owe their
floating power, and it is to be inferred from his remarks (p. 183) that
they have little or no buoyancy. The pyrenes of a Fijian inland species,
near M. Grayi, had no floating power as tested by me, and they lacked
the bladder-like cavity.

NOTE 9 (page 18)

The Buoyancy of the Fruits of Calophyllum

Professor Schimper found that whilst the fruits of Calophyllum
inophyllum, the shore tree, remained afloat after 126 days, those of C.
amœnum, an inland species, sank in from three to fourteen days, both
possessing similar buoyant structures, but to a less degree in the case
of the inland species. This genus presents a parallel case to Terminalia
referred to on page 17; but the general discussion of the subject will be
found in Chapter XIII. According to the above authority C. Calaba, a
West Indian coast tree, has buoyant fruits. The same is also true of the
fruits of a large inland tree in the Solomon Islands experimented on by
me (Solomon Islands, p. 305). It would thus appear that the fruits of
the genus are as a rule buoyant, and that, as in Terminalia, the least
buoyant fruits belong to the inland species. Professor Schimper also
shows (p. 182) that the diminished floating power of the fruits of the
inland species is associated with diminution in thickness of the buoyant
seed-shell which is most developed in the buoyant fruits of the strand
species.

NOTE 10 (page 24)

The Buoyancy Experiments on British Plants

The experiments in all cases were made to test the floating power of
the seed or fruit in the condition in which it is detached from the plant.
It usually makes very little difference whether sea-water or fresh water
is employed, since in my numerous experiments there were but few
exceptions to the general rule that seeds or seed-vessels that sink in
fresh water sink also in sea-water. This subject is discussed in Chapter
X. However, it may be here observed that the chief effect of the
increased density of sea-water is merely to increase the proportion of
buoyant seeds or fruits in any particular species.
It is necessary in such experiments to imitate Nature as much as
possible. The seed or fruit, as the case may be, must be experimented
upon in the condition in which it falls from the plant, or in the condition
in which it would be ultimately found in river and pond drift. The seed
or fruit should be thoroughly wetted, and air-bubbles removed.
Prolonged drying has but a slight effect on the great majority of seeds
and seed-vessels experimented on, and this is just as true of tropical
plants. Those that sink at once in the mature and fresh condition rarely
float more than a day or two even after drying for a year. The usual
effect is to increase the floating capacity of seeds and fruits already
buoyant, and not to develop the capacity.
The results given in the table refer only to sound seeds. In fresh-
water experiments, in nearly all cases, the seeds ultimately germinate in
the water, and this is the usual cause of the close of the experiment. In
an ordinary collection of floating seed drift from a pond or river,
germination will go on for years at each successive spring, the
postponement of germination being a very striking feature with a fair
proportion of seeds in river and pond-drift. This subject is dealt with in
detail in my paper published in the Proceedings for 1897 of the Royal
Physical Society of Edinburgh.
The Table of Results of Observations and Experiments on the
Buoyancy of the Seeds or Seed-vessels of more than 300
British Flowering Plants
Explanation of Table.—The capacity of floating for months is thus
indicated, ++; of floating for 1 to 4 weeks, +; and where sinking occurs
at once or within a week there is no entry. When buoyancy continued in
my experiments after 6 and 12 months, it is indicated by Roman
numerals (VI and XII). A=an aquatic plant; M=a beach plant; R=a river-
side or pond-side plant; var.=variable in floating power.
R Thalictrum flavum

A Ranunculus aquatilis

R Ranunculus hederaceus

R Ranunculus flammula

Ranunculus ficaria

R Ranunculus sceleratus VI ++ Var.

Ranunculus acris

Ranunculus repens XII ++ Var.

R Caltha palustris +
 Berberis vulgaris

A Nymphæa alba

A Nuphar luteum

Papaver rhœas

Papaver dubium

Chelidonium majus

Rœmeria hybrida

M Glaucium luteum

Barbarea vulgaris

R Nasturtium officinale

R Nasturtium sylvestre

R Nasturtium amphibium

Arabis hirsuta

Arabis thaliana

R Cardamine pratensis

Cardamine hirsuta

Alliaria officinalis

Brassica campestris

Brassica alba

M Cochlearia officinalis
M Alyssum maritimum

Draba verna

Thlaspi arvense

Capsella bursa pastoris

M Cakile maritima +

M Crambe maritima +

M Raphanus maritimus +

Reseda luteola

Helianthemum vulgare

R Viola palustris

Viola canina

Viola tricolor

Polygala vulgaris

Silene cucubalus

M Silene maritima

Lychnis diurna

Sagina procumbens

M Arenaria peploides (Honckeneya) XII ++

Mœnchia erecta

Cerastium vulgatum
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