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 Having in the first place formed the opinion that the achenes of the early Hawaiian Compositæ are
suited for dispersal by birds, and then shown that sea-birds were probably the principal agents, we
are met with the curious difficulty that in the case of the early Hawaiian genera of Compositæ the
complete suspension for ages of the means of dispersal is involved in the circumstances that these
genera are confined to the Hawaiian group. We can attribute to the agency of existing sea-birds the
occurrence of the genus Lagenophora in the uplands of Hawaii, on the mountain-tops of Fiji, and in
Australia and New Zealand; but the agency of birds as at present in operation does not assist us
except indirectly in the case of the genera restricted to Hawaii or to Tahiti. Is it possible, we may
inquire, to penetrate this mystery? Why, we may ask with Mr. Hemsley, has the agency ceased acting,
and why have its operations been confined to the conveyance of seeds to the islands and not from
the islands as well (Intr. Bot. Chall. Exped., p. 66)? I need scarcely add that the same question
presents itself with all the other peculiar genera of these islands, and in fact with endemic genera all
over the world. What can be stranger, it may be remarked, than the limited distribution of the
Pandanaceous genus Sararanga in the Western Pacific, although suited for dispersal by frugivorous
birds. This is not, indeed, a special difficulty connected with oceanic islands; it applies to the whole
plant-world; yet it is possible that, as it is exhibited by the Compositæ in these islands, we may be in
a better position to grapple with the problem. But before doing so it will be requisite to look a little
closer at these early Hawaiian genera of the Compositæ.
The distribution within the archipelago of the genera and species of the early Compositæ of Hawaii
is worthy of notice from the light it throws, not only on the relative antiquity of the genera, but also
on the subsequent conditions of isolation. Of the nine genera here referred to five are distributed over
most of the islands of the group. These include all the genera possessing a number of species,
namely, Tetramolopium with seven species, Lipochæta with eleven, Campylotheca with twelve,
Dubautia with six, and Raillardia with twelve species. Of the four genera remaining all have only two
species, and are restricted to two or three islands, Remya and Wilkesia being in both cases found in
Kauai and Maui, whilst Argyroxiphium is confined to the adjacent islands of Maui and Hawaii, and
Hesperomannia to those of Oahu, Lanai, and Maui. These four genera that are restricted to only two
or three islands are the same before referred to as regarded by Hillebrand as the oldest, partly on
account of their isolated generic position, and partly because in each case they only possess two
species.
Although the early Hawaiian Compositæ were evidently originally transported to most of the islands
of the group, it is noteworthy that their subsequent isolation from the rest of the world has in the
later ages been repeated within the limits of the archipelago. Of the 56 species, all of which are now
endemic, 28, or just half, as shown in the table on the following page, are confined to a single island.
Of the remainder, almost all are restricted to two or three adjacent islands. Hillebrand gives only a
solitary species, Lipochæta connata, as occurring in all the islands. This suspension, to a great extent,
of the means of dispersal between the islands is also strikingly illustrated by the Lobeliaceæ.
We have only to mention the flora of Fiji and those of the adjacent groups of Samoa and Tonga to
exclude them from any share in the early era of the Compositæ in the Pacific. The prevailing
adventitious character of the Fijian Compositæ is indicated in the fact that the species of the majority
of the genera are included by Seemann in his list of Fijian weeds. There are only one or two Fijian
Compositæ, such as the mountain species of Lagenophora and the littoral species of Wedelia, that
merit the special attention of the student of dispersal. So also with Samoa, Reinecke enumerates eight
species, of which six are weeds either of aboriginal or of European introduction, the others being the
littoral Wedelia above alluded to, and a species of Blumea found also in Fiji.

Distribution of the Endemic Genera of Compositæ in the Hawaiian Islands.

Distribution of the Species.
Genus. One Two Three Four Total.
General.
island. islands. islands. islands.
 Remya 2 — — — — 2

Tetramolopium 1 4 2 — — 7

Lipochæta 3 4 3 — 1 11

Campylotheca 5 4 3 — — 12

Argyroxiphium 1 1 — — — 2

Wilkesia 2 — — — — 2

Dubautia 4 — 2 — — 6

Raillardia 9 1 — 2 — 12

Hesperomannia 1 1 — — — 2

28 15 10 2 1 56
We have now, I venture to think, gone far to establish the existence of an early “Composite” flora
with mainly American affinities in the Pacific islands, an ancient flora of which only the remnants now
occur in the uplands of Hawaii, Tahiti, and Rarotonga. That the achenes were originally transported in
birds’ plumage is, as we have seen, probable; but we are still quite in the dark as to the causes of the
subsequent suspension of the means of dispersal and of the resulting period of isolation, during which
the original immigrant plants acquired their endemic characters. In our uncertainty, therefore, we will
look to Fiji in the hope that in the absence of the early Compositæ from that group we may find a
clue that will enable us to divest this problem of some of its difficulties.
It might be at first considered that since these peculiar genera of Compositæ occur in the higher
levels of Hawaii and Tahiti their absence from Fiji might be connected with the relatively low altitude
of those islands, a character that is concerned with the exclusion from the Fijian flora of many
Hawaiian and Tahitian mountain plants (see Chapters XXIII. and XXIV.). But this view is at once
negatived by the fact that Fitchia thrives in Rarotonga, an island which does not far exceed 2,000 feet
in elevation. It is negatived also by the extensive development of shrubby and arborescent
Compositæ in the Galapagos Islands, on the equator, in St. Helena in 16° South latitude, and in other
tropical islands, which are less than, or do not exceed, the Fijian Islands in their altitude.
During the age of the Compositæ it is reasonable to suppose that the dispersal was general over
the Pacific. The absence of genera indicating this era from the islands of the Fijian region, that is,
from Fiji, Tonga, and Samoa, would become intelligible if these groups were submerged during this
age of the general dispersal of the order over this ocean. In my volume on the geology of Vanua Levu
in Fiji, I have shown that these island-groups of the Western Pacific emerged from the sea towards
the close of the Tertiary period, a conclusion that would enable us to assign the age of the general
dispersal of the Compositæ over the tropical Pacific to an earlier portion of the same period.
In order, however, to make further progress in the discussion of this difficult problem we are obliged
to approach it from the outside. We must in fact regard these genera from the standpoint of their
position as members of the vast and ancient order of the Compositæ. It is now more than thirty years
since Mr. Bentham completed his remarkable memoir on the classification, history, and geographical
distribution of the Compositæ (Journal Linnean Society, Botany, London, Vol. 13, 1873). Like De
Candolle, when dealing with the facts of distribution, he handled thousands of species, and as a result
he drew certain inferences which are of prime importance to students of plant-dispersal. In his time
the order included nearly 10,000 known species, and although this number has since no doubt been
considerably increased, it is not likely that his main conclusions, in so far as they are free from purely
hypothetical considerations, will be materially affected by the later discoveries.
 Accepting the antiquity of the order, and regarding it as probably dating far back in geological time,
he observes that the evidence points to a very wide dispersion of its original stock at an early period.
Africa, West America, and possibly Australia, possessed the order at the earliest recognisable stage.
There must have existed, he contends, at this early period some means of reciprocal interchange of
races between these regions. Then followed a stoppage of communication, or a suspension of the
means of dispersal, between the tropical regions of the Old and New Worlds; but long after
communication was broken off in the warmer regions, it still existed, as he holds, between the alpine
heights in those regions and also between the high northern latitudes of both hemispheres. Referring
particularly to the Hawaiian Group, he considers that the large endemic element among the
Compositæ indicates that the ancient connection, whether with America or with Australasia, has been
so long severed as not to have left a single unmodified common form. Fitchia, the Tahitian genus, as
we have already remarked, is regarded as the only remnant of an ancient Composite flora in the
tropical islands of the South Pacific.
In the light of these reflections it will be interesting to glance at the general distribution of the
shrubby and arborescent or woody Compositæ. Mr. Hemsley, having generally discussed the subject,
arrived at the conclusion that, “although they form so large a proportion of the floras of St. Helena,
Juan Fernandez, the Sandwich Islands, and some other islands, they are not specially insular.” There
are scores of them, he goes on to say, in South America, Africa, Madagascar, India, Australia, and
New Zealand from twenty to forty feet high, and more truly arboreous than the insular ones; whilst
nearly every sub-order has its arboreous representatives. He was, however, unable to form any
definite opinion of the method of distribution of the woody Compositæ. Taking those of St. Helena
and Juan Fernandez, he observes that they are not more closely allied to the Compositæ of the
nearest continents than they are to those of more distant regions. The occurrence of arboreous
Compositæ, belonging in each case to different tribes, in so many remote oceanic islands, coupled
with the distribution of the genera to which they bear the greatest affinity, seems, he observes, to
indicate that they are the remains of very ancient types (Introd. Bot. Chall. Exped., pp. 19-24, 66, 68;
also Parts ii. p. 61, and iii. p. 23).
The further discussion of this subject would lead us into a wide field of inquiry, quite beyond the
scope of this work. There is, however, an inference that I think we may legitimately draw from
geological evidence in this region. With respect to the antiquity of the woody Compositæ of the Pacific
as illustrated by the endemic genera, both Mr. Bentham and Mr. Hemsley view them as belonging to
ancient types. Mr. Wallace, in his Island Life, a book that becomes more and more indispensable for
the student of dispersal as years progress, dwells on the importance of these ancient Compositæ in
the floral history of the Pacific islands. We may look upon the Hawaiian Compositæ, he remarks, as
representing the most ancient portion of the existing flora, carrying us back to a very remote period
when the facilities for communication with America were greater than they are now. The date of this
period of oceanic dispersal of the Compositæ we can now approximately determine, since these
plants are absent from the Fijian region, an area of submergence during the Tertiary era. Before the
island-groups of the Fijian region had emerged towards the close of the Tertiary period the achenes of
the early Compositæ had been dispersed far and wide over the tropical Pacific.
But this is not all that we can infer from the convergence of these independent lines of botanical
and geological investigation. Mr. Bentham observes that the tribes of the Compositæ had acquired the
essential characters now employed in classification before the dispersion of the order over the Pacific.
Since this general dispersion took place, as we hold, during the Tertiary submergence of the island-
groups of West Polynesia (Fiji, Tonga, Samoa), it follows that the birth of the tribes of the Compositæ
antedates that period. If this interesting order could supply us with a “datum-mark” in the history of
the Pacific floras, it would be stated in terms of the development of specific and generic characters,
but not of those of a tribe.

Summary of Chapter.
 (1) The Hawaiian Islands present the same contrast with the Fijian and Tahitian groups as regards
the development of new species in the case of the flowering plants that they offer in the case of the
vascular cryptogams (ferns and lycopods). But the contrast is intensified, and it is further emphasised
as respecting the flowering plants by the evolution of a large number of endemic genera.
(2) This great preponderance of peculiar species and genera in Hawaii is not to be connected with
the relative antiquity of the group but with its degree of isolation.
(3) The earliest stage of the flowering plants of the islands of Hawaii and of Eastern Polynesia (the
Tahitian region) is indicated by the endemic genera, particularly those of the Compositæ and
Lobeliaceæ. Such genera are numerous in Hawaii, and occur also in the Tahitian region, as in Tahiti
and Rarotonga; but do not exist in the groups of the Fijian region (Fiji, Tonga, and Samoa).
(4) The endemic genera of the Hawaiian Compositæ are mainly American in their affinities. The
relationship of the solitary Tahitian genus (Fitchia) is still a subject of discussion.
(5) In the Hawaiian Islands, as well as in Tahiti and Rarotonga, the plants of the endemic genera of
Compositæ are, as a rule, arborescent or shrubby; and in the first two localities they are mainly
restricted to the higher levels.
(6) In discussing the mode of dispersal of the achenes of the original genera we have also to
explain why the process of dispersal has been in the main suspended.
(7) It is shown that the achenes of these early Compositæ were in all probability suited for dispersal
in birds’ plumage.
(8) Yet the isolating influence that cut off these genera from the outside world has, in later ages,
been active within the limits of the Hawaiian archipelago, with the result that half the species are not
found in more than a single island. Inter-island dispersal has, therefore, been also largely suspended.
(9) The absence of endemic genera of Compositæ from Fiji, Tonga, and Samoa cannot be
attributed to unsuitable climatic conditions connected with the relatively low elevation of those islands
as contrasted with those of Hawaii, since a species of Fitchia abounds in Rarotonga, which is not far
over 2,000 feet in elevation. Shrubby and arborescent Compositæ of peculiar types also occur in the
Galapagos and other tropical islands not more elevated than the Fijis.
(10) These endemic genera are the remains of an ancient Composite flora in the islands of the
tropical Pacific, and ages have elapsed since the severance of their connections with regions outside.
(11) According to Mr. Bentham the Compositæ were distributed over Africa, West America, and
possibly Australia, at an early period, but subsequent to the differentiation of the tribes of the order.
Some means of reciprocal interchange of races between these regions then existed. Then followed a
suspension of the means of dispersal between the tropical regions of the Old and New Worlds except
between the alpine heights of those latitudes.
(12) It is inferred by the author of this volume that the general dispersion of the early Compositæ
over the Pacific took place during the Tertiary submergence of the island-groups of the Fijian region
(Fiji, Tonga, and Samoa), and that their absence from that region may be thus explained. At the time
of this general dispersion, as above pointed out, the tribes of the Compositæ had been already
differentiated.
 CHAPTER XXII

THE ERA OF THE ENDEMIC GENERA (continued)

The Compositæ and Lobeliaceæ (continued)

The Age of the Tree-Lobelias

The distribution of the arborescent Lobeliaceæ.—On the upper flanks of Ruwenzori.—The Lobeliaceæ of
the Hawaiian Islands.—The Lobeliaceæ of the Tahitian or East Polynesian region.—The capacities for
dispersal.—The explanation of the absence of the early Lobeliaceæ from West Polynesia.—The other
Hawaiian endemic genera.—The Fijian endemic genera.—Summary.

The Lobeliaceæ rank with the Compositæ in the prominence of their position in the early Pacific floras.
Though absent, as far as is known, from Fiji, they are represented in Hawaii by 58 species, all endemic
and belonging to six genera, of which five are not found elsewhere. All possess, as Hillebrand remarks,
a woody stem, by far the greater number being either tall shrubs, 5 or 6 feet high, or small trees, 10 to
20 feet or more in height. In the East Polynesian or Tahitian region, the order is represented by two
genera containing in all five known species and restricted to those islands. One genus is common to the
islands of Tahiti and Rarotonga, and the other is confined to Raiatea. The species may be shrubby or
arborescent.
It was for some time considered that the oceanic archipelagoes of the Pacific were the exclusive
centres of these singular arborescent Lobeliaceæ (I am here quoting Baillon in his Natural History of
Plants). And indeed this idea would receive some support from the circumstance that Dr. Hillebrand, in
his work on Hawaii, says little or nothing about the affinities or general relations of plants which he
enthusiastically termed “the pride of our flora.” His death in 1886 deprived his work of its crowning
piece, a discussion of “the interesting questions of the origin and development of the Hawaiian flora”
(see the Editor’s Introduction, p. ix.). In no group of plants is this want more keenly felt than with the
Lobeliaceæ. Yet in his time the explorations had yet to be made that could set the student of plant-
distribution on the road to investigate this problem.
It was true, no doubt, that types analogous to those of the Hawaiian Lobeliaceæ were known from
the American and African continents. Thus Oliver in his Flora of Tropical Africa, published in 1877, gives
an account of the species of Lobelia then known from the mountains of this region. The genus was,
however, not entirely confined to mountainous districts, but it would almost seem that most of the high
mountains of Equatorial Africa had their peculiar species, some of them being tree-like and others
shrubby. Two mountain species were recorded from Abyssinia, one of them from an elevation of 11,000
to 13,000 feet and growing to a height of 12 to 15 feet, the other from an altitude of about 8,000 feet;
another, Lobelia Deckenii, attaining a height of 4 feet, was recorded from the uplands of Kilimanjaro,
12,000 to 13,000 feet above the sea, and yet another from the mountains of Fernando Po, at an
altitude of 9,000 feet. So again, in the case of the American continent, Hemsley, writing in 1885 (Intr.
Bot. Chall. Exped., p. 32), speaks of arborescent species of the American genera Centropogon,
Siphocampylus, &c.; and Baillon in his Natural History of Plants (Engl. edit. viii. 350) refers to the similar
Tupas and Haynaldias from South America. But what the student of plant-distribution looked for was not
merely the occurrence of “tree-lobelias” in other parts of the world, but also the reproduction of these
wonderful plants under the same conditions and on the same scale as those familiar to him on the
Hawaiian mountains. He has accordingly had to wait for the results of the more recent explorations of
the mountains of Central Africa in order to obtain his wish.
On the upper flanks of Ruwenzori, Kilimanjaro, and Kenya, at elevations of 9,000 to 13,000 feet and
reaching to the snow-line, there flourish in boggy portions of the forest arborescent Lobeliaceæ that
attain a height of 15 or 20 feet. They have the habit sometimes of a Dracæna and sometimes of an
Aloe, and do not exhibit the branching trunks so characteristic of the Hawaiian genus of Clermontia.
They all belong, however, to the genus Lobelia, and thus do not display the extensive differentiation of
the endemic genera of Hawaii. Nor, apparently, has there been the same degree of formative energy in
the development of species, since only about half a dozen species are hitherto known. We find,
however, produced on these lofty mountains of Equatorial Africa the same climatic conditions under
which the arborescent Lobeliaceæ flourish in Hawaii, namely, the very humid atmosphere, the heavy
rainfall, and the mild temperature; and if there are important contrasts in their character and in the
amount of differentiation which they have undergone in the two regions, the one a continental and the
other an insular region, it will be from such contrasts that some of the most interesting results of this
comparison of a mountain of Central Africa with an island of the open Pacific will be ultimately derived
(see Sir H. Johnston’s Uganda Protectorate, 1902, and Kilimanjaro Expedition, 1886; also Trans. Linn.
Soc. Bot., ser. 2, vol. 2, p. 341.)

THE LOBELIACEÆ OF THE HAWAIIAN AND OF THE EAST POLYNESIAN OR TAHITIAN ISLANDS.[1]

Hawaiian Islands.
No. of Distribution Distribution in Height of Nature of Station.
Genus.
species. of genus. the group. plant. Elevation. Station.
Islands not Steep palis
Brighamia 1 Endemic. Molokai, Niihau. 5 to 12 feet. exceeding 3,500 or mountain
feet. gaps.

Bridges,
2,000 to 6,000
Lobelia 5 Non-endemic. General. 4 to 6 feet. gulches and
feet.
woods.

Usually 10 to 2,000 to 6,000

Clermontia 11 Endemic. General. Open woods.
20 feet.[2] feet.

Usually 4 to 6
Higher parts of
feet, one
Rollandia 6 Endemic. Oahu. Oahu, which is Woods.
species 10 to
4,000 feet high.
15 feet.

1,000 to 5,000 Woods and

Delissea 7 Endemic. General. 5 to 10 feet.
feet. gulches.

Woods,
Usually 6 to 15 1,000 to 5,000
Cyanea 28 Endemic. General. ravines,
feet.[3] feet.
gulches.

East Polynesian or Tahitian Islands.

Humid
Endemic in E. Tahiti, 1,500 to 3,000
Sclerotheca 4 6 to 25 feet. wooded
Polynesia. Rarotonga. feet.
slopes.
 In the mountains.
Apetahia. 1 Endemic. Raiatea. 3 to 6 feet. Elevation of island
3,400 feet.

1. The materials are nearly all derived from the works of Hillebrand and Drake del Castillo. Some of
those relating to the elevations in Hawaii are supplemented from my notes. All the genera are endemic
except Lobelia, of which all the species are apparently endemic, excepting perhaps one, which,
according to Hillebrand, resembles greatly a species from the Liukiu Islands.

2. The range of the heights of different species of Clermontia is from 5 or 6 feet for shrubs to 25 feet
for trees.

3. The heights attained by different species of Cyanea range from 3 or 4 feet to between 30 and 40
feet, thus:—
In 8 species 3 to 6 feet.
In 9 species 6 to 10 feet.
In 7 species 10 to 15 feet.
In 3 species 15 to 25 feet.
In 1 species 30 to 40 feet.

The Lobeliaceæ of the Hawaiian Islands.

Having thus prepared the way, I will proceed to the discussion of the Hawaiian Lobeliaceæ, dealing
first with their “station.” Their vertical distribution is well illustrated in the large and lofty island of
Hawaii. Whilst the woody Compositæ, as before described, are most at home on the open-wooded and
often scantily-forested slopes between 5,000 and 9,000 feet, the Lobeliaceæ are most characteristic of
the middle or true forest zone that extends from 2,000 or 3,000 feet to between 5,000 and 6,000 feet
above the sea. This lies within the region of clouds and mists, and it is here that the rain-belt or area of
greatest rainfall is situated, the annual amount averaging probably 150 to 200 inches. It is in such
humid conditions that, as Hillebrand observes, trees and jungle are developed in greatest luxuriance;
and it is here that “the Lobeliaceæ exhibit their most striking forms.” The traveller, as he ascends the
mountains, finds the Tree-Lobelias in the region of mist and rain-cloud; and he is lucky if he escapes the
usual downpour and encounters only a fine drizzling rain.
The mild climate of this region is indicated by a mean annual temperature ranging probably with
elevation from 65° to 55° F. It is secure from the frosts of the upper slopes of the mountain; whilst at
the same time it is above the regions of tropical heat. There is, however, no doubt that when the forests
extended to the coasts, as they occasionally do now on the north side of Hawaii, the Lobeliaceæ
occurred much lower down than they do at present, though still only attaining their greatest
development in size and number in the higher levels. Thus, at rare intervals, I noticed in the forests of
Hamakua and Kohala, where they descended to the coasts, species of Clermontia at an elevation of only
500 or 600 feet above the sea.
Probably in no part of the Hawaiian Islands are the conditions under which the “Tree-Lobelias” thrive
better illustrated than on the higher slopes of Mount Eeka, a bulky mountain mass about 6,000 feet in
height, forming the western portion of Maui. Its flat top, as Hillebrand observes, is wrapped in a cloud
of mist nearly the whole year. On the boggy surface of the summit, where Acæna exigua gives a
tussocky appearance, and Sphagnum or bog-moss abounds, flourish Cyperaceæ, Lycopods, and
Selaginellæ; and here Drosera longifolia and a peculiar species of marsh violet (Viola mauiensis) find a
home. The upper slopes, down to 4,000 feet, present similar moist conditions, and here in an open-
wooded district, associated with Cyrtandræ, Marattias, and true Tree-Ferns, the ground being covered
with Lycopods, the “Tree-Lobelias” abound. I noted four kinds within two hundred yards. Of the
humidity of the upper slopes of Mount Eeka I have a very vivid recollection, and my experience of
passing a night on that mountain is described in Chapter XIX.
The Lobeliaceæ, as Hillebrand remarks, occur invariably as isolated individuals. I was often struck,
however, with the preference the genera showed for particular localities. Thus, Clermontia is well
represented on the western slopes of Mount Eeka, Delissea on the northern slopes of Hualalai (3,800 to
4,500 feet), Cyanea on the Hamakua slopes of Mauna Kea (2,300 to 4,100), and Lobelia on the
southern slopes of Mauna Loa behind Punaluu (2,000 to 3,500 feet).
To the student of geographical distribution the Hawaiian Lobeliaceæ are of especial interest. Mr.
Hemsley observes that they have their greatest affinities in America (Intr. Bot. Chall. Exped., p. 68). M.
Drake del Castillo, in his “Mémoire couronné par l’Académie des Sciences” (Paris, 1890), remarks that
these plants connect Hawaii with America just as the Goodeniaceæ link the same group with Australia.
This is what we might have expected since the centre of the order is in America, principally in the
Mexican and Andine regions (Drake del Castillo, Flore Polyn. Franc., xi.).
Though five out of the six genera are endemic, the sixth, that of Lobelia, has a world-wide
distribution. Here then, we have a genus that belongs strictly to the next or second stage of the plant-
stocking of the Hawaiian Group, namely, when the non-endemic genera now containing endemic species
were introduced. As with the Composite genera, Campylotheca and Lipochæta, Lobelia marks the
beginning of the new or the close of the old era. It is, however, necessary to point out that many of the
conditions favouring luxuriant and rank vegetable growth are pre-eminently represented in the zone of
the Lobeliaceæ. In these soft-stemmed plants with their copious milky sap and large fleshy flowers,
sometimes two or three inches long, the very redundancy of growth would tend both to exaggerate and
to disguise the generic distinctions. To the ordinary observer these “Tree-Lobelias” call up vague notions
of a flora of a bygone age, and by their bizarre appearance he might with some excuse be led to give
play to his imagination when describing them; but the systematic botanist, seeing through their
disguise, frames rather more prosaic notions of their antiquity and degree of differentiation. According
to my view, the first Hawaiian Lobeliaceæ occupied open, exposed localities such as are held by the
decadent genus Brighamia now, and acquired their monstrous form in the humid forests of a later age.
(See Perkins in Note 80.)
In his monograph on the Campanulaceæ (Engler’s Nat. Pflanz. Fam., teil 4, abth. 5, 1894), S.
Schönland, speaking of the sub-family Lobelioideæ, places the seven endemic Hawaiian and Tahitian
genera in a group by themselves. Though, as he observes, the Hawaiian tree-forms appear at first sight
to constitute a natural group, they cannot be sharply distinguished from other forms, and even in habit
come near some Indian and Abyssinian types of Lobelia. In their treatment, he says, they should all go
together, and he does not approve of the endeavours of some botanists to isolate one of them
(Brighamia) from the rest and to connect it with the Australian genus Isotoma.
It is also to be noted that whilst four of the Hawaiian genera are more or less dispersed over the
group, one (Brighamia) with only one species is confined to the islands of Molokai and Niihau, the
double habitat being suggestive of its approaching extinction. Another (Rollandia) with six species is
restricted to the island Oahu. Cyanea, which possesses twenty-eight out of the total of fifty-eight
species, may, from the point of view of its formative energy, be regarded as in its prime. It is thus
apparent that, as with the Compositæ, the early Lobeliaceous immigrants were not all contemporaneous
arrivals.

DISTRIBUTION OF THE LOBELIACEÆ IN THE HAWAIIAN ISLANDS.[4]

Hawaiian
Brighamia. Lobelia. Clermontia. Rollandia. Delissea. Cyane
Lobeliaceæ.
Species confined
— — 6 6 4 22
to one island

Species confined
1 2 2 — 2 5
to two islands
Species confined
— 1 2 — 1 1
to three islands

Species generally
distributed, but — 2 1 — — —
still endemic

1 5 11 6 7 28

4. All the species are endemic.

Another interesting fact of distribution, brought out by an analysis of Hillebrand’s materials and
illustrated in the subjoined table, is that out of the fifty-eight Hawaiian species, all of which are
endemic, thirty-eight, or 66 per cent., are recorded from only one island. In most of the other cases
they are recorded from two or three islands, usually adjacent, like Maui and Molokai; and except in the
instance of two species of Lobelia and one species of Clermontia they never range over the length of
the group.
These facts speak eloquently of the suspension to a great extent of the agencies of dispersal in recent
times within the group. Some corrections of the figures will be rendered necessary by future
investigations, but the main conclusion will not be materially affected. Such facts are paralleled in the
distribution of the Hawaiian insects, mollusca, &c.; but these matters need only be mentioned here. We
might, indeed, have expected, apart from other considerations, that the isolation of the Hawaiian
Lobeliaceæ from their kindred in other parts of the world would not have been reproduced within the
group itself. This, however, is not the case; and we now see that not only have they been deprived for
ages of their means of distribution over the Pacific, but that even within the archipelago their
transportal from island to island has been largely suspended. We have before arrived at similar
conclusions with regard to the early Compositæ, when we saw that about half the species were not
found in more than one island. It is therefore evident that the same great principle regulating the
operations of the distributing agencies has influenced to a similar extent both the Compositæ and the
Lobeliaceæ of the Hawaiian Group.

The Lobeliaceæ of the Tahitian or East Polynesian Region.

The order is represented in this region by two endemic genera, Sclerotheca of Tahiti and Rarotonga,
and Apetahia of Raiatea. These islands are, however, not sufficiently large for the extensive
development of the arborescent Lobeliaceæ, such as we find in Hawaii. The species in both genera are
either arborescent or shrubby; but I do not gather that they give any character to the floras of these
islands. According to the data given by Drake del Castillo for one of the two peculiar species of
Sclerotheca occurring in Tahiti, these plants grow on the humid wooded slopes of the mountains at
elevations of 2,000 to 3,000 feet. Whilst in one species the plants attain a height of 10 to 25 feet, in the
other they do not exceed 10 feet. Rarotonga possesses a peculiar species of Sclerotheca, 4 to 6 feet
high, which was discovered by Cheeseman growing plentifully on the upper slopes of the highest
mountain of the island at altitudes of 1,500 to 2,200 feet. The same botanist also came upon a second
species of the genus on another mountain in Rarotonga at elevations of 1,000 to 1,500 feet, but it was
rare and has not yet been described. The other genus, Apetahia, has only been recorded from Raiatea,
where it is represented by a solitary species (6 feet high) growing, according to Nadeaud, in the
mountains of that island.
It is apparent that the dispersal of these genera of the Lobeliaceæ amongst the groups of Eastern
Polynesia ceased long ago. From the circumstance that Sclerotheca exists in Tahiti and in Rarotonga,
which are about 650 miles apart, it may be inferred either that the genus was introduced into this
region from outside, or else, which is perhaps more probable, that it was developed in Tahiti whence it
was transported to Rarotonga. Hemsley speaks of this Tahitian genus as seemingly marking a former
wide extension of the Hawaiian arborescent type of the Lobeliaceæ (Introd. Bot. Chall. Exped., p. 68).
This is the view that will be adopted in this chapter, and it is precisely the view advocated by Bentham
and followed here, in the case of the early Compositæ of the Pacific.
With regard to the absence of these arborescent Lobeliaceæ from the island-groups of the Western
Pacific, and notably from Fiji and Samoa, where no members of the order seem to occur, it is probable
that, as in the case of the similar distribution of the early Compositæ described in the preceding
chapter, this is to be attributed to the fact that the Western Pacific archipelagoes were more or less
submerged during the general dispersion of the Compositæ and Lobeliaceæ over the Pacific in the
earliest age of the floral history of these islands. The occurrence of the early Compositæ and
Lobeliaceæ in Rarotonga, which is almost half-way between Tahiti and Tonga on the outskirts of the
Fijian region, sufficiently indicates that they are not lacking in that region from inability to reach there in
the past. During the age of general dispersal of these two orders over the Pacific, probably only a few
rocky islets, tenanted perhaps by Conifers, marked the situation in the Tertiary period of the present
archipelagoes of Fiji and Samoa.
One may note in passing the general absence of these arborescent types of the Lobeliaceæ from
Malaya, since they do not seem to have been recorded either from the Owen Stanley Range in New
Guinea or from Kinabalu in North Borneo, the highest mountain in the Malayan Islands, or from the
mountains of Java.
The consideration of the occurrence of these plants in other tropical or subtropical oceanic islands
need not detain us long, since, with the exception of the solitary Lobelia scævolifolia of St. Helena, they
seem rarely to be found. This species, which is endemic, is a shrub growing on the upper slopes and
summit of the island at elevations of 2,000 to 2,700 feet (Introd. Bot. Chall. Exped., p. 40, and Part ii.
pp. 54, 76).
There are two herbaceous species of Lobelia in Juan Fernandez, of which one only, according to
Hemsley, could be regarded as indigenous. This is a showy Chilian and Peruvian species (Lobelia tupa)
noticed by Bertero as very common in 1829 (Bot. Chall. Exped., Part iii.). Since, however, it would
belong to the present age of plant-dispersal in the Pacific, it does not require further mention here; and
indeed it would almost appear, when we bear in mind the geographical position and the history of this
island since its discovery in 1563, that even as a truly indigenous plant it is not above suspicion.
Lobelias of this type are now amongst the commonest plants of the coast regions of northern Chile,
where I noticed some as much as 9 or 10 feet high.
On the Capacities of Dispersal of the Lobeliaceæ of the Pacific.—Of actual observations, with the
exception of the instance of birds pecking at the capsules of our garden Lobelias, I have come upon few
that bear directly on this point. When writing of the flora of the Kermadec Group, many years ago, Sir
Joseph Hooker referred (Journ. Linn. Soc. Bot., vol. i.) to the minute seeds of Lobelia as not adapted for
transport unless their minuteness and number fit them for it; but since he associates in this connection
the tiny seeds of Metrosideros, which is now represented by a species found all over the Pacific, it would
seem that the difficulty in the case of Lobelia is not connected so much with the nature as with the
suspension of these means of distribution during the later stages of the plant-stocking of the oceanic
islands of the tropical Pacific. It will be gathered from the following remarks that the descendants of the
early Pacific Lobeliaceæ are probably as well fitted for dispersal as their ancestors, and that the break in
the communication is the ultimate subject for inquiry.
The fruits of the Hawaiian endemic genera are in four out of five cases baccate, with usually fleshy or
pulpy contents. Such berries, which are generally yellow, but sometimes bluish in colour, vary in size
from about half an inch in Rollandia and Delissea to an inch in Cyanea, and not infrequently to more
than an inch in Clermontia. The fruits of Lobelia and Brighamia are capsular and dehiscent. With regard
to the two genera of the Society Islands and Rarotonga, the fruits of Sclerotheca are hard-walled
capsules, opening by two pores; whilst those of Apetahia are seemingly dry and indehiscent. I do not
imagine, therefore, that the character of the fruit has determined to any important degree the
distribution of these plants.
Nor is there reason to suppose that the fruits have acquired their baccate character in Hawaii, and
that they were originally dry and capsular. Both types of fruit are found among the arborescent
Lobeliaceæ of America, with which the Hawaiian genera have their affinities. Centropogon, for instance,
which occurs in Central America and in the warm parts of North and South America, has, according to
Baillon, a somewhat fleshy berry. It is noteworthy that a similar question is raised with respect to
Cyrtandra as to the relation between fleshy fruits in the Pacific islands and dry or capsular fruits in the
continental home of the genus (see Chapter XXV.).
The berries of the Tree-Lobelias would attract birds. We learn from Mr. Perkins that one of the
Hawaiian Drepanids, the Ou, is very partial to the berries of some of the Tree-Lobelias and especially
those of Clermontia, the seeds passing unharmed in the droppings. The mode of dispersal of the seeds
of the dry-capsular fruits is not so apparent; but the fruits could scarcely be less inviting to birds than
the dry capsules of Metrosideros, the small seeds of which have in some way or other been carried to
almost every island-group of the Pacific. I have beside me the dark brown, smooth crustaceous seeds of
a species of Clermontia. They measure 1⁄42 of an inch or 0·6 of a millimetre, and about 500 go to a
grain. Mr. Wallace, in his book on Darwinism, advocates the paramount influence of winds over birds for
carrying small seeds, like those of Orchis and Sagina, over tracts of ocean a thousand miles across. I
am, however, not inclined to think that, except as regards the spores of cryptogams, winds have done
very much for Hawaii. For small seeds we can appeal not only to the agency of birds and bats but also
to insects (see Chapter XXXIII.).
Observations of this kind, however, merely indicate that these early Lobeliaceæ possessed the same
capacities for dispersal that in the succeeding stages of the plant-stocking of the Pacific islands have
belonged to Metrosideros, Cyrtandra, Ophiorrhiza, Freycinetia, and many other small-seeded genera.
They go no way to explain why the same agencies which transported the minute seeds in a later age
could not have been available for continuing the dispersal of the early Lobeliaceæ. To find an
explanation we are compelled to go behind the mere capacities for dispersal and to appeal to the
general laws of distribution in so far as our facts enable us to interpret them.
We have seen that the two principal components of the early Pacific flora, the Compositæ and the
Lobeliaceæ, have American affinities. The plants of the later ages are mainly Old World in their
connections. Though containing often endemic species in the various groups, the genera occur also
outside each group. The stream of migration that came from America during the early age of the
Compositæ and the Lobeliaceæ, when the islands of the Western Pacific were more or less submerged,
was during the later ages (after these islands had re-emerged) suspended or diverted, giving place to a
stream that brought plants in numbers from tropical Asia, Malaya, and Australia. The general dispersion
of the Compositæ and Lobeliaceæ took place during the Tertiary submergence of the islands of the
Western Pacific, including the island-groups of Fiji, Samoa, and Tonga. The migration from the west,
mainly Indo-Malayan in character, occurred after the re-emergence of those archipelagoes. Thus we get
to understand how genera like those of the early Lobeliaceæ and Cyrtandra, which possess, as regards
the minute size of their seeds, closely similar capacities for dispersal, have such different distributions,
the first confined to Hawaii and Tahiti and American in their affinities, the second widely spread over the
Pacific with its home in Malaya.
We have yet to inquire whether this suspension of the means of transport in the later ages of the
Pacific Lobeliaceæ is confined to the tropics or whether it extends to the colder latitudes in the southern
hemisphere. The indications of the Lobeliaceæ of the “antarctic flora” go to establish that the dispersal
of the order is still, or was very recently, in operation in these high latitudes. It is well illustrated, among
other plants, by Lobelia anceps, which is found in extra-tropical South America, Australia and New
Zealand, and South Africa. This, indeed, recalls Bentham’s view concerning the Compositæ, that whilst
communication was broken off in the tropics, it was kept up in higher latitudes.
Here ends, therefore, our consideration of the Tree-Lobelias of the Pacific islands; but as it is not
quite complete without a discussion of the remaining endemic genera of other orders than the
Compositæ and Lobeliaceæ which also belong to the same early age of the Pacific floras, I will proceed
at once to their consideration.

The Hawaiian Endemic Genera excepting those of the Compositæ and Lobeliaceæ.
 It will not be possible for me to do more than point out a few general indications that can legitimately
be drawn from these genera. The subject bristles with difficulties for the systematist; but on one point
there can be but little danger of going astray, namely, in imputing to them a high antiquity in the floral
history of Hawaii. This can be said of all of them, whether or not the generic distinction adopted in Dr.
Hillebrand’s work is always adopted by botanists. It is therefore in this general sense that they may be
regarded as belonging to the early age of the Hawaiian flora.
Although the genera of Compositæ and Lobeliaceæ are prominent amongst the representatives of the
original flora of the Hawaiian Islands, forming about two-fifths of the whole, the genera of other orders
are by no means inconspicuous, and their variety is shown in the fact that though twenty-three in
number they belong to twelve orders. It is possible to divide these genera into two groups—one the
older and perhaps more or less contemporaneous with the Lobeliaceæ and Compositæ, the affinities
when apparent being American; the other the more recent and marking the close of the first era of the
plant-stocking of these islands, the affinities being all with the Old World, and especially with Malaysia.
This grouping is indicated in the list subjoined; and it may be here remarked that whilst shrubs,
undershrubs, and perennial herbs of the Caryophyllaceæ, Labiatæ, and Urticaceæ form the features of
the earlier group, trees of the Rubiaceæ and Araliaceæ are the most conspicuous members of the later
group. At the close of the earliest era known to us of the floral history of the Hawaiian Islands we
observe the commencement of those forests that now throughout Polynesia as well as in Hawaii betray
their Asiatic origin.
In making this distinction I am proceeding on the assumption that the stream of migration, at first
chiefly American in its source, came ultimately in the main from the Asiatic side of the Pacific. The
change commenced, as I hold, in the latter portion of the first era of plant-stocking, an era
characterised by the arrival of those early plants that are now represented by the endemic genera of the
archipelago. The genera of this early period that belong neither to the Compositæ nor to the
Lobeliaceæ are, as above observed, arranged by me in two groups, one regarded as contemporaneous
with, the other as of later origin than, the genera of these two orders. To the first belong the shrubby,
highly differentiated genera of the Caryophyllaceæ, Schiedea and Alsinidendron, and the Labiate
genera, similarly differentiated, of Phyllostegia and Stenogyne. To the second belong the Rubiaceous
genera Kadua, Gouldia, Bobea, and Straussia, the Araliads Cheirodendron, Pterotropia, and Triplasandra,
and the Loganiaceous Labordea.
In the earlier group the fruits are dry in half the genera, and in such cases granivorous birds probably
were usually the transporting agents. Only in one case (Nothocestrum) is the fruit a berry, and in the
other cases we have fruits like the fleshy nucules of Phyllostegia and Stenogyne which would probably
attract birds. In the later group two-thirds or three-fourths of the genera have moist fruits such as
would be eaten by frugivorous birds. Of these most are drupes, possessing not a single stone, but two
or more pyrenes. This is the first appearance of the drupe in the plant-history of the archipelago. The
Rubiaceous type of drupe inclosing two or more pyrenes plays a very conspicuous part in the
distribution of plants over the Pacific in the succeeding eras.
I would here lay stress on an important characteristic of all the fruits of the endemic genera of the
Hawaiian Islands. There are no “impossible” fruits of this era in Hawaii, such as we occasionally find in
the succeeding eras. I mean by this term, fruits that defy the efforts of the student of distribution to
explain their transport in their present condition. The discovery of a new inland genus possessing dry
indehiscent fruits three or four inches long, or even of a single species of the coniferous Dammara,
would play havoc with all our views respecting the stocking of these islands with their plants. The
finding here of a large marsupial would scarcely produce more astonishment. The fruits indeed of this
early era are very modest in their size, the dry indehiscent fruits and the stone-fruits rarely exceeding
half an inch (12 mm.) in size.
There is another interesting point which is connected with the deterioration of some of the fruits in
their capacity for dispersal. Some of the species of Phyllostegia, and a few also of the Araliads, as well
as those of Nototrichium, are ill fitted for dispersal by birds now, the coverings of the seeds being not
sufficiently hard to protect them from injury in a bird’s stomach. At the same time there are in some
cases other species of the same genera that are better suited for this mode of transport. The effect of
dispersal by frugivorous birds is that only the hard-coated seeds propagate the plant in a new locality.
When, however, as has occurred in the Hawaiian Islands, bird-agency largely ceases to act, this
selective influence is removed (see Note 68).

ENDEMIC HAWAIIAN GENERA, EXCLUDING THOSE OF THE COMPOSITÆ AND LOBELIACEÆ, AS GIVEN
IN HILLEBRAND’S “FLORA OF THE HAWAIIAN ISLANDS.”
[Those preceded by * are not usually regarded now by botanists as endemic, though they nearly
take that rank.]
The Earlier Group.
Number
Genus. Order. of Character. Fruit. Affinities.
species.
Isodendrion Violaceæ. 3 Shrubs. Capsule. American (H).

Undershrubs,
Schiedea Caryophyllaceæ. 17 Capsule.
&c. Near Colobanthus of the
Antarctic islands, temperate
Capsule, South America, and Australia
Alsinidendron Caryophyllaceæ. 1 Undershrubs. with fleshy (C).
calyx.

Small trees or
Platydesma Rutaceæ. 4 Capsule. —
shrubs.

Hillebrandia Begoniaceæ. 1 Herbs. Capsule. —

Nothocestrum Solanaceæ. 4 Small trees. Berry. South American (H).

Regarded by Gray as a
*Haplostachys Labiatæ. 3 Herbs. Dry nucules.
section of Phyllostegia.

Fleshy Belong to the tribe Prasiæ,

*Phyllostegia Labiatæ. 16 Undershrubs.
nucules. which is mostly Asiatic. Two
other species of Phyllostegia
Trailers or Fleshy recorded from Tahiti and
Stenogyne Labiatæ. 17
climbers. nucules. Paumotu Islands.

Charpentiera Amarantaceæ 2 Trees. Utricle. American (H).

Achene with
Touchardia Urticaceæ. 1 Shrubs. fleshy —
perigone.

Achene with
Allied to Bœhmeria, a genus
Neraudia Urticaceæ. 2 Shrubs. fleshy
of Old and New Worlds.
perigone.

The Later Group.

Belongs to Melicope, an Old

*Pelea Rutaceæ. 20 Trees. Capsular.
World genus (IK).

Broussaisia Saxifragaceæ. 2 Small trees. Berry. Malayan (H).

 Referred to Panax, an Old
*Cheirodendron Araliaceæ. 2 Trees. Drupe.
World genus (IK).

*Pterotropia Araliaceæ. 3 Trees. Drupe. Malayan (H). Pterotropia

Trees or referred to Heptapleurum of
Triplasandra Araliaceæ. 4 Drupe. Old World (IK).
shrubs.

Approaches both Asiatic and

Kadua Rubiaceæ. 16 Shrubs, &c. Capsular
American types (C).

Small trees or Drupaceous

Gouldia Rubiaceæ. 5 American (C).
shrubs. berry.

Malayan (H). Genus also in

*Bobea Rubiaceæ. 5 Small trees. Drupe.
Malaya (IK).

Near Psychotria, a genus of

Straussia Rubiaceæ. 5 Trees. Drupe.
Asia and America (H).

Small trees or Capsule with

Labordea Loganiaceæ. 9 Malayan (H).
shrubs. pulp.

Trees or Referred to the Australian

*Nototrichium Amarantaceæ. 3 Utricle.
shrubs. Ptilotus (IK).

(H) = Hillebrand’s Flora of the Hawaiian Islands.

(C) = Drake del Castillo’s Remarques sur la Flore de la Polynésie.
(IK) = Index Kewensis.
Note.—Probably Schumann’s genus, Pteralyxia, should be placed in the later group (see p. 154).
Another feature of interest is to be found in the distribution within the archipelago of the species of
the peculiar genera. As in the case of the Compositæ and Lobeliaceæ, but few of the species are
generally distributed, most being restricted to one island or to two or three adjacent islands. The
suspension of the dispersal among the islands is, however, not so marked as with the species of the two
orders just named.
Note.—Some further remarks on some of these genera are given in Note 68.

The Endemic Genera of the Fijian Islands.

The interest that is associated with the endemic genera of Hawaii fails to attach itself to those of Fiji.
For this there are several reasons. In the first place, our acquaintance with the Fijian flora is much less
complete. In the next place, the group holds a much less isolated position, and the history of an
endemic genus may have a significance quite different from that connected with it in Hawaii. Fiji also
lacks, on account of its submergence in the Tertiary period, those highly interesting genera of the
Compositæ and Lobeliaceæ that form the chief feature in the early history of the flowering plants of
Hawaii. Then, again, on account of our imperfect knowledge of the floras of the neighbouring groups of
continental islands to the westward, the New Hebrides, Santa Cruz, and Solomon Groups, we can never
feel quite confident that any particular genus is really peculiar to the Fijian archipelago. This is well
brought out in the later history of the genera designated by Dr. Seemann in his Flora Vitiensis as
peculiar to Fiji.
Of the sixteen genera enumerated by Dr. Seemann, and given in the table below, only about half now
retain their character of being restricted to Fiji. Nor does it seem likely that future investigations will
increase this number, since, judging from a remark made by Mr. Hemsley in his paper on the botany of
the Tongan Group, explorations subsequent to those of Dr. Seemann, more especially those of Mr.
Horne, have not apparently added a single new endemic genus to the Fijian flora. It will be seen from
the list that at least four of the sixteen genera have since been found in the Malayan region, and in one
case (Smythea pacifica) the same species occurs in both regions; whilst a fifth genus (Haplopetalon)
has been recorded from New Caledonia.
There are, however, some peculiarities about the Fijian endemic genera that will attract our attention
from the standpoint of dispersal. One remarkable feature is the paucity of species. Almost all the genera
are monotypic, that is to say, they are only known by a single species. Amongst the twenty-eight
Hawaiian genera that are strictly endemic, only four or five are monotypic, and they are mostly
regarded by Hillebrand as worn-out, decadent types found in only one or two islands. In Hawaii there
are on the average six species to each endemic genus; and it is thus apparent that in the display of
formative energy Nature has worked on very different lines in these two groups. Since the nine Fijian
endemic genera belong to nearly as many different orders, the composition of this endemic generic flora
is by no means homogeneous. It is, I venture to think, such a motley collection as one might expect in
a region that has been exposed to wave after wave of migration from the west, with no lofty mountains,
as in Hawaii, to afford a refuge against extinction. It by no means follows that all these endemic genera
have been produced in Fiji. Some of them may represent genera that have become extinct in the large
continental groups to the westward.

SEEMANN’S SIXTEEN FIJIAN ENDEMIC GENERA.

Number of Affinities or other
Genus. Order. Character. Fruit.
species. localities.
Richella Anonaceæ. 1 Tree. Baccate(?). Indian in type (C).

Trimenia Ternstrœmiaceæ. 1 Tree. Unknown.

Small spinose Related to Australian genera

Pimia Sterculiaceæ. 1 Tree.
capsule. (S).

Near Trichospermum, a
Græffea Tiliaceæ. 1 Tree. Unknown. Fijian and Malayan genus
(S).

Order mainly South

Thacombauia Humiriaceæ. 1 Shrub. Drupe.
American.

Near Soulamea, a Malayan

Amarouria Simarubeæ. 1 Tree. Dry drupe.
genus (S).

Straggling Also in Burma, New Guinea,

*Smythea Rhamneæ. 1 Capsule.
shrub. and Malaya (IK), (Sc).

*Oncocarpus Anacardiaceæ. 2(H) Tree. Drupe. Also in New Guinea (IK).

*Haplopetalon Rhizophoreæ. 2 Shrub. Unknown. Also in New Caledonia (IK).

*Nesopanax Plerandreæ. 1 Tree. Drupe. =Plerandra (IK).

Bakeria Plerandreæ. 1 Tree. Drupe.

Pelagodendron Rubiaceæ. 1 Shrub. Berry.

 =Agapetes, a Malayan
*Paphia Ericaceæ. 1 Shrub. Berry.
genus (IK).

*Carruthersia Apocyneæ. 2(H) Climber. Berry. Also in Philippines (IK).

Also in Kaiser Wilhelmsland,

*Couthovia Loganiaceæ. 2 Tree. Drupe.
New Guinea (So).

Canthiopsis Loganiaceæ. 1 Shrub. Drupe.

Those genera marked * have since been found outside the group.
The authorities are thus indicated: (C)=Drake del Castillo; (H)=Horne; (IK)=Index Kewensis
(S)=Seemann; (Sc)=Schimper; (So)=Solereder in Engler’s Nat. Pflanz. Fam.
The fact that several of them are fitted for dispersal by frugivorous birds is very suggestive of the lack
of means of transport in later times. In the instance of Couthovia corynocarpa the drupes are known to
be the food of fruit-pigeons at the present time (Seemann), whilst this is also true of Oncocarpus
vitiensis, though this genus has since been found in New Guinea. Since, as will be pointed out in a later
chapter, birds must still be fairly active in carrying seeds to Fiji from regions westward, it would seem
that genera only become peculiar to Fiji when they fail at their source, and it is indeed doubtful whether
any of the Fijian peculiar genera are home productions. One may instance in this connection the genus
Pimia, the fruits of which are especially well suited for attachment to a bird’s plumage, yet it is only
known from Fiji.
It should be here observed that no peculiar generic types have been recorded from the adjacent
Tongan Group, and scarcely any from Samoa. Except perhaps with the Palmaceæ, no peculiar genera
seem to be mentioned in Dr. Reinecke’s memoir on Samoa.

Summary.
(1) The Lobeliaceæ, like the Compositæ, take a prominent place in the early Pacific flora, being
represented, more particularly in Hawaii but also in the East Polynesian or Tahitian region, by endemic
genera of tall shrubby and tree-like species.
(2) Tree-Lobelias occur in other parts of the world, as in South America and tropical Africa; but it is
especially on the higher slopes of the mountains of Equatorial Africa that they attain a development
comparable with that of Hawaii.
(3) In Hawaii the Tree-Lobelias are most characteristic of the middle forest-zone (3,000-6,000 feet),
where the temperature is mild, the rainfall heavy, and the atmosphere laden with humidity.
(4) The affinities of these endemic genera of the Lobeliaceæ are mainly American; but their generic
distinctions have been both exaggerated and disguised by redundant growth.
(5) From the distribution of the genera and species within the Hawaiian Group it is evident that, as
with the early Compositæ, the original Lobeliaceous immigrants were not all contemporaneous arrivals.
Some of the genera are on the point of extinction, whilst others are in their prime.
(6) The absence of the Lobeliaceæ from the groups of the Fijian area (Fiji, Tonga, Samoa) is probably
to be connected, as in the case of the absence of the early Compositæ, with the circumstance that the
general distribution of these two orders over the tropical Pacific occurred during the Tertiary
submergence of these archipelagoes.
(7) These endemic genera of the Lobeliaceæ possess the same facilities for dispersal that are owned
by other genera with minute seeds, such as Cyrtandra, &c., that are dispersed over the Pacific; but in
the case of the Lobeliaceæ the agencies of dispersal have been for ages suspended.
(8) This suspension is to be associated with the diverting of the main stream of migration from its
source in America, during the early age of the Lobeliaceæ and Compositæ, to a source on the Asiatic
side of the Pacific.
 (9) The Hawaiian endemic genera other than those of the Compositæ and Lobeliaceæ arrange
themselves in two groups—an earlier group containing highly differentiated Caryophyllaceæ and
Labiatæ, and belonging to the age of the Compositæ and Lobeliaceæ; and a later group, characterised
by Rubiaceæ and Araliaceæ, which marks the close of the first era, as well as the change in the main
source of the plants from America to the Old World, the beginning of the Hawaiian forests, the
appearance of the Rubiaceous drupe, and the first active intervention of frugivorous birds.
(10) Though there are no “difficult” or “impossible” fruits (fruits, the dispersal of which is not easy to
explain) amongst the forty and odd endemic genera of Hawaii and Tahiti, it is noteworthy that in some
cases the fruits are seemingly little fitted for dispersal now, and that this deterioration in capacity for
dispersal is to be frequently associated with more or less failure of the inter-island dispersal in the case
of Hawaii.
(11) The interest associated with the Hawaiian endemic genera fails to attach itself to those of Fiji,
where genera only seem to have become peculiar because they have failed at their sources in the
regions to the west. The endemic genera of the Compositæ and Lobeliaceæ are here lacking, and this is
true also of the neighbouring Samoan and Tongan Groups, it being held that the age of the general
dispersion of these orders over the Pacific corresponded with the Tertiary submergence of the
archipelagoes of the Western Pacific. Those of Fiji, which do not amount to ten in number, belong to
nearly as many orders and present a motley collection such as one might look for in a group much less
isolated than Hawaii and exposed to wave after wave of migration from the west.
 CHAPTER XXIII

THE ERA OF THE NON-ENDEMIC GENERA OF FLOWERING PLANTS

The Mountain-Floras of the Pacific Islands as illustrated by the Non-endemic Genera

The mountain-flora of Hawaii.—A third of it derived from high southern latitudes.—An American
element.—Compared with Tahiti and Fiji.—Capacities for dispersal of the genera possessing only
endemic species.—Acæna, Lagenophora, Plantago, Artemisia, Silene, Vaccinium, &c.—Capacities for
dispersal of the genera possessing non-endemic species.—Cyathodes, Santalum, Carex,
Rhynchospora.—Fragaria chilensis, Drosera longifolia, Nertera depressa, Luzula campestris.—
Summary.

The Age of the Endemic Genera of Flowering Plants.

We are now entering an era distinguished from the preceding age of the endemic genera, the age
chiefly of the Compositæ and Lobeliaceæ, by the fact that the extreme isolation that followed that era
no longer prevails. In a sense these island-floras are in touch again with the world around, though the
main stream of plant-migration now comes from the south and from the west. Yet in a large number of
cases, the amount varying greatly in the different groups, it is evident that this stream has not flowed
continuously to the present day. The agencies of dispersal are often no longer active; but the period of
inactivity has not been sufficiently prolonged to produce generic distinction, and the differentiating
energy has been restricted to the development of new species.
Yet within these limits the development of new forms, as indicated in Table B on p. 233, has often
been very great. Thus, nearly half the Hawaiian genera that are non-endemic are composed entirely of
species not found outside the group; and in this sense they may be regarded as cut off from the regions
around. In Fiji and Tahiti only about a fourth are in this manner isolated, the agencies of dispersal being
still effective with the majority of the genera. It is apparent, therefore, that the same question
concerning the cause of the failure of the means of dispersal presents itself in this era as in the last, and
most markedly in the instance of Hawaii.
The simplest and quickest plan for bringing into relief the prominent features of this age is first to
regard the genera from the standpoint of the elevation of their stations. We have before remarked that
in the occurrence of extensive regions of great altitude the Hawaiian Islands differ conspicuously from
the groups of Tahiti and Fiji (and I may add Samoa); and that they present conditions for the
development of a temperate mountain-flora that are not found at all in Fiji and are barely represented in
Tahiti. That the Hawaiian flora responds to this contrast between the elevations of the three groups is
well established; and I will now proceed to refer more in detail to the subject.

The Mountain-Floras of the Pacific Islands.

In the Hawaiian Islands there are at least 37 or 38 genera, making up about 19 or 20 per cent. of
those belonging to this era, that may be designated mountain genera, nearly all of them being
characterised as appertaining exclusively or in the main to temperate regions, or as frequenting
mountain-tops in tropical latitudes. In Tahiti there are only 7 or 8 of such genera, about 4 per cent. of
the total for the era. In Fiji, excluding the Conifers, there are only 4 or 5, or not 2 per cent. of the
whole. In Samoa, which may be included in the Fijian area, there are 3, or about 2 per cent. of the
total. These are results which we might have expected from the varying altitudes of these groups, as
described in Chapter XIX.
Few things give more pleasure to the botanist than his recognition in some remote locality of plants
long familiar to him in other regions. This will often be his lot on the mountain summits of Hawaii. If he
has been a mountain-climber in many countries, he will there notice again the genera Artemisia,
Geranium, Plantago, Ranunculus, Rubus, Sanicula, Vaccinium, and others that he has met perhaps
either in the Rocky Mountains or in the Andes or in Equatorial Africa or in the Himalayas. If fresh from
Chile he will find on these heights the familiar Gunnera and the Chilian Strawberry (Fragaria chilensis).
If he has been in New Zealand and in the islands of the Southern Ocean he will find old friends in the
genera Acæna and Coprosma. He may handle once again plants like Nertera depressa, that he gathered
on Tristan da Cunha; and on the boggy summits of some of the mountains he will find the ubiquitous
Sun-dew (Drosera longifolia).
Within the limited area occupied by the peaks of Tahiti he will find genera like Astelia and Coprosma
that are at home in New Zealand or in Antarctic America, and may even find, as in the cases of Coriaria
ruscifolia and Nertera depressa, the identical species that are at home in those distant regions. Even on
the summit of Rarotonga he will gather a species of Vaccinium. In Fiji, here and there on some isolated
mountain-top he may come upon a remnant of this Antarctic flora, such as a solitary species of
Coprosma or Lagenophora, that will carry him back for a moment to high southern latitudes; and in the
highlands of Savaii, in the neighbouring Samoan Group, he will find again Nertera depressa and a
species of Vaccinium. But that which will interest him most in Fiji will be the tall conifers of the genera
Dammara, Podocarpus, and Dacrydium, which will bring to him memories perhaps of New Zealand and
southern Chile, of South Africa, and of the mountain-woods of Java and of Southern Japan.
Yet the influence of isolation has been at work amongst the mountain-plants of all these groups. The
agencies that have dispersed over the tropical Pacific plants from the cold latitudes of the southern
hemisphere, and those that have borne the seeds of Plantago, Sanicula, and Vaccinium from mountain-
top to mountain-top, even though it be to a peak in mid-ocean, are to a great extent inactive now.

The Mountain-Flora of Hawaii as illustrated by the Non-endemic Genera.

Let us look in the first place at Hawaii, where the breaking off of communication with the outside
world is especially pronounced. Here, all the species of two-thirds or more of the mountain-genera are
confined to that group. Only in a relatively small number of cases are the species in touch with the
regions outside. The mystery of disconnection that is so evident in the instance of the peculiar or
endemic mountain-genera of the Compositæ and Lobeliaceæ and other orders is here again presented
to us, and once more in the upland regions 4,000 to 10,000 feet above the sea. We will now endeavour
to discover from an examination of the present distribution of the isolated mountain-genera (those non-
endemic genera possessing only peculiar species) along what tracks they arrived at the Hawaiian
uplands, tracks, as indicated by the local distribution of the species, that have been more or less
abandoned since.
The Mountain Genera with only Endemic Species.—By referring to the Table on the following page it
will be observed that nearly a third of these mountain genera have now their principal homes in the
high latitudes of the southern hemisphere. They are components of what Forster and Hooker have
termed the “Antarctic” flora, a collection of plants that range round the globe in high southern latitudes,
namely, over Fuegia, New Zealand, southern Australia, South Africa, and the islands of the Southern
Ocean, the “Antarctic” islands, as they have been termed. These genera are Acæna, Gunnera,
Coprosma, Lagenophora, Astelia, Oreobolus, and Uncinia. (It is necessary to observe that I am entirely
indebted to the Introduction to the Botany of the “Challenger” Expedition for my information on the
“Antarctic” flora.)
 We are thus led to expect that some of the other mountain genera may have been similarly derived
from cool southern latitudes, even though they may be scarcely included in the “Antarctic” flora. This is
very probably true of Myoporum and Exocarpus, two genera that are chiefly centred in Australia. A
species of Sophora (S. tetraptera) is now one of the most widely dispersed of the plants of high
southern latitudes, a circumstance which at all events explains the capacity for transport that the
ancestor of the Hawaiian “Mamani” (S. chrysophylla) must have originally possessed (see Chapter XV.).
Kinship between the Hawaiian species and southern forms has been found in the case of a few of the
widely ranging genera here represented. Thus Decaisne placed Plantago princeps next to P.
fernandeziana of Juan Fernandez; whilst according to Hillebrand, Plantago pachyphylla resembles P.
aucklandica from the Auckland Islands. These resemblances are consistently associated with the
respective range in altitude of the Hawaiian plants, since Plantago princeps occurs usually between
2,000 and 4,000 feet, and P. pachyphylla between 6,000 and 8,000 feet, the species of greatest
elevation being related with the species of highest latitude. It is thus seen that these endemic mountain
genera with peculiar species have very evident affinities with the plants of extra-tropical southern
latitudes, and especially with the “Antarctic” flora. This affinity will also be found, as will subsequently
be noticed, in the case of genera like Cyathodes and Nertera, where there is still a specific connection
with the outside world.

THE MOUNTAIN-FLORA OF HAWAII, AS REPRESENTED BY THE NON-ENDEMIC

GENERA (Compiled from Hillebrand’s Flora).
Usual Distribution in Hawaii, Fiji,
Distribution outside Polynesia.
altitude and Tahiti.
Genus of Australia All
Both Old New Antarctic Hawaii Hawaii, Hawaii,
station and New three
Worlds. World. World. flora. only. Fiji. Tahiti.
in feet. Zealand groups.
With all Species Endemic.

Ranunculus 6,000-
+ ... ... ... ... + ... ... ... A
(2) 7,000

2,000-
Viola (5) + ... ... ... ... + ... ... ... C
6,000

2,000-
Silene (4) + ... ... ... ... + ... ... ... C
9,000

5,000-
Geranium (6) + ... ... ... ... + ... ... ... C
10,000

7,000-
Vicia (1) + ... ... ... ... + ... ... ... P
8,000

5,000-
Sophora (1) + ... ... ... ... ... ... ... + P
10,000

4,000-
Rubus (3) + ... ... ... ... ... + ... ... B
7,000

5,000- S
Acæna (1) ... ... ... + ... + ... ... ...
6,000 a

3,000-
Gunnera (1) ... ... ... + ... + ... ... ... D
6,000
 6,000- P
Sanicula (1) ... ... + ... ... + ... ... ...
8,000 c

3,000-
Coprosma (9) ... ... ... ... + ... ... ... + D
9,000

Lagenophora V
6,000- ... ... ... + ... ... + ... ...
(1) a

4,000-
Artemisia (2) + ... ... ... ... + ... ... ... A
8,000

2,000-
Lobelia (5) + ... ... ... ... + ... ... ... C
6,000

3,000- +
Vaccinium (2) + ... ... ... ... ... ... ... B
8,000 Samoa

Myoporum Coast to
... ... ... ... + + ... ... ... D
(1) 10,000

2,000-
Plantago (2) + ... ... ... ... + ... ... ... C
8,000

3,000- F
Exocarpus (2) ... ... ... ... + + ... ... ...
6,000 n

Sisyrinchium 4,000-
... ... + ... ... + ... ... ... C
(1) 7,000

2,000-
Astelia (2) ... ... ... + ... ... ... ... + B
6,000

T
Oreobolus (1) 6,000 ... ... ... + ... + ... ... ...
n

3,000- A
Uncinia (1) ... ... ... + ... + ... ... ...
5,000 n

4,000- A
Agrostis (3) + ... ... ... ... + ... ... ...
6,000 g

Deschampsia 3,000- A
+ ... ... ... ... + ... ... ...
(3) 6,000 g

3,000- A
Trisetum (1) + ... ... ... ... + ... ... ...
5,000 g

Poa (2) + ... ... ... ... + ... ... ... G

With Endemic and Non-endemic Species.

2,000-
Cyathodes (2) ... ... ... ... + ... ... + ... D
10,000
Lysimachia Coast to
+ ... ... ... ... + ... ... ... C
(6) 6,000

Chenopodium Up to S
+ ... ... ... ... + ... ... ...
(2) 7,000 li

Coast to
Santalum (3) ... + ... ... ... ... ... ... + D
10,000

2,000-
Carex (5) + ... ... ... ... ... + ... ... N
7,000

Rhynchospora Up to
+ ... ... ... ... ... ... ... + N
(4) 10,000

Coast to
Panicum (14) + ... ... ... ... ... ... ... + G
6,000

Up to A
Deyeuxia (3) + ... ... ... ... + ... ... ...
10,000 g

With no Endemic Species.

Fragaria 4,000-
... ... + ... ... + ... ... ... F
chilensis 6,000

Drosera
4,000 + ... ... ... ... + ... ... ... C
longifolia

Nertera 2,500-
... ... ... + ... ... ... ... + D
depressa 5,000

Luzula 3,000-
+ ... ... ... ... ... ... + ... C
campestris 10,000
It is evident that in one or two cases the connection between the representatives of the “Antarctic”
genera on the Hawaiian uplands and those of high southern latitudes has only been recently broken off.
Thus with reference to the Hawaiian species of the Cyperaceous genus, Uncinia, it may be observed
that although Hillebrand regards it as a distinct species, Hemsley (Intr. Bot. Chall. Exped., p. 31)
remarks that it is very near if not the same as a New Zealand species, an affinity very significant of the
source of the mountain plants of this group that are derived from these southern latitudes.
The next component to be recognised in these Hawaiian mountain genera with peculiar species is a
small special American element; and in this connection Sanicula and Sisyrinchium may be especially
mentioned. The first is mainly North American, and particularly Californian; but there are two solitary
species found on the continents and in oceanic islands such as the Azores. The continental species,
Sanicula europæa, occurs not only in Europe and Central Asia, but in South Africa, and at high
elevations on the mountains of Equatorial Africa and of Madagascar. It is not, however, with this widely
ranging species that Sanicula sandwicensis is related, but with S. menziesii, a species from California
and Oregon (Hillebrand). Sisyrinchium is confined to temperate and tropical America; but a singular and
suggestive outlier of the genus (S. bermudiana) is found in Bermuda.
The mountain genera that are distributed on both sides of the Pacific constitute about three-fifths of
the total. So far as my scanty data show, they seem to have reached Hawaii from the four quarters of
the compass. The probable southerly origin of Plantago has been already indicated. Hillebrand notes the
great resemblance between Lobelia gaudichaudii and an undescribed species from the Liukiu Islands,
lying on the west side of the Pacific. It is likely, also, that the genus Ranunculus reached Hawaii from
the west, since one of the species, R. mauiensis, resembles R. repens of the Old World (Hillebrand);
whilst the other, R. hawaiiensis, comes near R. sericeus of Mauritius (Drake del Castillo). On the other
hand, the genus Rubus may hail from an American source, since, in the opinion of Gray, Rubus
hawaiiensis, one of the mountain raspberries, finds its nearest relative in R. spectabilis from the north-
west coast of America; and there are reasons for believing, as will subsequently be shown, that the
genus Artemisia has an American source. It is also probable that some of these genera have reached
Hawaii from the north, since it is likely, as pointed out in a later page, that the Carices of the Hawaiian
uplands came originally from north-eastern Asia.
In the previous paragraphs the mountain genera have been considered with especial reference to
their distribution and source beyond the confines of the Pacific. If we now briefly discuss them from the
standpoint of their distribution within the Pacific, or rather as concerning their presence or absence in
the Fijian and Tahitian groups, we shall see that to a large extent Hawaii has received its mountain
genera of this era independently of the other Pacific groups.

Mountain genera possessing only peculiar species, in Hawaii only 20

Mountain genera possessing only peculiar species, in Hawaii and Fiji 2

Mountain genera possessing only peculiar species, in Hawaii and Tahiti 0

Mountain genera possessing only peculiar species, in all three groups 4

26

It is here shown that three-fourths of the genera of the Hawaiian mountains in this era are not found
either in Fiji or Tahiti. This, as before pointed out, is mainly to be attributed to the greater elevation of
the Hawaiian Islands. Had there been an island 13,000 to 14,000 feet in height in Fiji, we cannot think
that any such contrast in the floras would have existed. The temperate genera of the Hawaiian uplands
would have been largely represented in the Fijian flora. Yet although we do not find such genera as
Ranunculus, Geranium, Sanicula, Uncinia, &c., in Fiji and Tahiti, a small number of the Hawaiian
mountain genera have obtained a scanty footing. This is what we might have expected. Thus,
Lagenophora has been found on the mountains of Vanua Levu, and Vaccinium in Tahiti and Rarotonga;
whilst Coprosma and Astelia occur on the tops of some of the mountains in both regions. In Fiji their
distribution seems sporadic, as shown not in Lagenophora alone, but also by Astelia, which has been
found only on the summit of Kandavu.
The Capacities for Dispersal of the Hawaiian Non-endemic Mountain Genera possessing only Peculiar
Species.—As shown in the Table, seven, or 27 per cent., of these genera have fleshy fruits that would
attract frugivorous birds. In three cases (Gunnera, Coprosma, Myoporum) they are drupes, in three
others (Rubus, Vaccinium, Astelia) they are berries, and in one (Exocarpus) there is a nut with a fleshy
perigone. It is particularly interesting to notice that frugivorous birds, and I include here granivorous
birds that are known to be frugivorous at times, could have transported seeds of the “Antarctic” flora to
this group. We can observe the process in operation in our own time within the limits of the group. It
has been long known, and we find it referred to in the pages of Hillebrand’s work, that the wild
mountain-goose (Bernicla sandwicensis) feeds upon the fruits of Coprosma ernodeoides, and of
Vaccinium reticulatum, the famous “ohelo.” The fruits of the first are known to the natives as “kukai
neenee” (droppings of geese), and the hard stones or pyrenes are very well suited for withstanding the
risks of the digestive process. I found a number of these pyrenes in the stomach of a mountain-goose
shot by my companion, Dr. Krämer, high up the slopes of Mauna Loa.
According to Mr. Perkins, Chloridops kona, a big Hawaiian finch, feeds on the fruits of the bastard
sandal-tree (Myoporum sandwicense). There are no “impossible fruits” among the mountain genera of
Hawaii, that is to say, fruits so large that bird agency must be excluded. All of them are practicable in
point of size. Thus amongst the largest, the “stones” of Gunnera would not exceed 1⁄5 of an inch (5
mm.), and those of Myoporum scarcely 1⁄4 of an inch (6 mm.); whilst the nuts of Exocarpus range in the
Hawaiian species from 3⁄10 to 6⁄10 of an inch (7-15 mm.), and the beans of Sophora chrysophylla do not
at the most exceed 1⁄4 of an inch (6 mm.).
The principal feature, however, which these mountain genera exhibit from the point of view of their
dispersal is the number of plants possessing seeds or fruits capable of adhering to plumage. Half of
these genera are thus characterised. Of these Sanicula and Acæna represent the ordinary hooked fruits;
whilst the fruits of the Grasses and Sedges, Agrostis, Deschampsia, Trisetum, Poa, Oreobolus, and
Uncinia, are enabled by means of their awns or of their serrated beaks to attach themselves to
plumage, and the same may be said of the carpels of Geranium. The fruits of Lagenophora and the
seeds of Plantago display the capacity of adhesiveness by means of a gummy secretion.
One or two of these genera need further mention. I will first take Acæna, which is spread all over the
south temperate zone both on the continents and on the islands. The Hawaiian species (A. exigua)
forms tussocky growths on the swampy summits of Mount Eeka in Maui, and in Kauai, at an elevation of
6,000 feet above the sea. Numerous observers refer to the probable mode of dispersal of the genus in
the “Antarctic” and neighbouring islands. Captain Carmichael, in the instance of Acæna sanguisorbæ on
Tristan da Cunha, observes that it overruns the low ground. Its burr-like fruit, as he describes, “fixes
itself on the slightest touch into one’s clothes, and falling into a hundred pieces covers one all over with
an unseemly crust of prickly seeds not to be got rid of without infinite labour” (Trans. Linn. Soc., xii.
483, 1818). Both Mr. Moseley (Wallace’s Island Life, p. 250) and Dr. Kidder (Bull. U.S. Nat. Mus., 2) refer
to the burrowing habits of the Petrels, Puffins, and other sea-birds amongst the vegetation covering the
ground in Tristan da Cunha, Marion Island, Kerguelen, &c., in places where Acæna, amongst other
plants, thrives. Mr. Moseley remarks that the fruits of this genus stick like burrs to feathers, and he
looks to sea-birds for the dispersal of this and similar plants over the ocean. He especially notes that the
Petrels and other seafowl burrow and breed high up the mountain-slopes of tropical islands as in Tahiti,
Viti Levu, Hawaii, and Jamaica.... It should be noted in the case of the Hawaiian endemic species that it
has been found only on two mountain tops; and that however active may be the dispersal of the genus
in south temperate latitudes now, the Hawaiian Islands lie outside the present area of dispersal.
The next mountain genus I will specially refer to is Lagenophora, one of the Compositæ. The solitary
Hawaiian endemic species, L. mauiensis, is restricted to the summit of Mount Eeka, in Maui. In the
mountains of Vanua Levu, Fiji, another peculiar species, L. pickeringii, has been found; and there is a
species, L. petiolata, in the Kermadec Islands (Hooker, in Journ. Linn. Soc., i. 127); but the genus is
chiefly characteristic of Australia, New Zealand, and temperate South America, one species occurring
both in Fuegia and Tristan da Cunha. The genus has no pappus; but Hooker in the case of the
Kermadec species considered that the “viscid fruit” favoured its dispersal; and this may probably be true
of the genus.
With regard to the capacity for dispersal of the seeds of Plantago, it may be pointed out that the
seeds of Plantago major, P. lanceolata, &c., become coated with a mucilaginous material when wetted.
In 1892, when experimenting on these plants, I found that the wetted seeds adhered firmly to a
feather, so that it could be blown about without their becoming detached. Species of Plantago are so
characteristic of the “alpine” floras of the summits of lofty mountains in the tropics, as in Java and many
other regions, that the mode of dispersal has always been a subject of curiosity. I cannot myself doubt
that this is the explanation of the occurrence of the representatives of the genus that now thrive as
endemic species on the higher slopes of the Hawaiian mountains. This method of dispersal for Plantago
is recognised by recent writers on the subject of seed-dispersal. (In a paper in Science Gossip for
September, 1894, I dealt with the “mucous adhesiveness” of such seeds as a factor in dispersal. The
subject had previously been discussed by Kerner in one of the earlier volumes of his Pflanzenleben; and
I have summed up some of the results in Note 43 of the present volume.) My readers can readily
ascertain by a simple experiment that a bird pecking the fruit-spikes in wet weather would often carry
away some of the sticky seeds in its plumage. Several years ago, when I was endeavouring to examine
the condition of these seeds in the droppings of a canary, my efforts were defeated by the bird itself,
since, in spite of all my care, some seeds and capsules were always carried by the bird on its feathers
into the clean cage reserved for the experiment.
 The plants of these mountain genera possessing dry seeds or fruits neither very large nor very minute
and suitable for bird-food are Ranunculus, Viola, Vicia, Sophora, Artemisia, Sisyrinchium, six in all, or 24
per cent. of the total. On the probable method of transport of the ancestors of these endemic species
the following remarks may be made. With regard to Ranunculus, some authors like C. M. Weed (Seed-
Travellers, p. 48, Boston, 1899) perceive in the curved or hooked beaks of the achenes a means of
attaching the fruit to plumage. This no doubt applies to some species, and it is advocated by Ekstam for
some of the plants of the Nova Zembla flora. There are others to which this explanation would not be
applicable, and the achenes of the Hawaiian species do not appear to be specially fitted for this mode of
transport. I have found the achenes of Ranunculus frequently in the stomachs of birds in England, in
partridges frequently, and in wild ducks at times. Those of certain species that possess buoyancy are
common in the floating seed-drift of rivers, as of the Thames (Journ. Linn. Soc. Bot., xxix. 333), and
they would probably in this way be often swallowed by waterfowl.
I have but few data directly relating to the dispersal of seeds of Viola by birds. From the frequent
occurrence of species in alpine floras, as in the Caucasus, the Great Atlas, in the mountains of
Equatorial Africa, in Madagascar, &c., it may be inferred that birds transport the seeds between the
higher levels of many continental ranges in tropical regions and to the mountain-slopes of neighbouring
large islands. Viola abyssinica, for instance, which occurs in Madagascar, is spread over the elevated
mountain ranges of tropical Africa. With regard to the five Hawaiian species, it may be remarked that
three of them are bog species and two occur in dry situations. The first are most characteristic of the
mountains, one species occurring on the summit of Mount Eeka, 6,000 feet above the sea. Judging from
the stations alone, at least two species were originally introduced into the Hawaiian Group.
Viola seeds, as indicated by my experiments on the different British species, including Viola palustris,
are not buoyant, and there is no possibility of the seeds being picked up by birds in floating drift. There
is, however, a possible means of dispersal in birds’ plumage by means of the mucosity of the seeds of
some species. Thus, although this is not exhibited, as shown by my experiments, by Viola canina and V.
palustris, it is well displayed by the Field-Pansy (V. tricolor). I found that the seeds of this species, after
lying a little time in water, were thickly covered with mucus, and that they adhered to a feather, on
drying, as firmly as if gummed. This did not, however, come under my notice in the case of the seeds of
one of the Hawaiian species, V. chamissoniana, examined by me. One sometimes observes Viola canina
in England growing in places, as in the crevices and on the tops of old walls, where its seeds could have
only been carried by birds. In some cases the propellent force of the seed ejected by the contracting
valves of the capsule would explain queer stations. In its power of seed-expulsion, Viola chamissoniana,
the common Hawaiian species, is just as active as our British species.
With regard to the Leguminous genus Vicia we have the observation of Focke on the dispersal of its
seeds by pigeons, as described before on page 150.
Sophora chrysophylla, the “Mamani” of the natives and one of the most familiar of the trees of the
Hawaiian mountains, is discussed at length in Chapter XV., where the difficulty of supposing that its
seeds could be transported unharmed in a bird’s stomach half-way across the Pacific is pointed out; and
it is suggested that it was more probably derived from a littoral species brought by the currents.
However, the point is a debatable one, and the seeds of the “Mamani” can scarcely be regarded as
“impossible” from the standpoint of dispersal.
With reference to the possibilities of dispersal of the achenes of Artemisia, some very suggestive
indications are to be obtained from a paper by Mr. D. Douglas on the North American Tetraonidæ
published in the Transactions of the Linnæan Society for 1833. The “Cock of the Plains” (Tetrao
urophasianus), as we here learn, makes its nest on the ground under the shade of Artemisia bushes,
and lives on the foliage and fruits of these and other plants. This bird is plentiful in Columbia and North
California, and another allied species is mentioned which lives on the same sort of food. Later authors
refer to these and other birds of the same family as living chiefly on the Sage-brush (Artemisia
tridentata), a plant prevailing over great regions of the plains as well as on the slopes of the Sierra
Nevada and of the Rocky Mountains. According to Dr. Sernander (page 228), birds when feeding on the
fruits of Artemisia vulgaris in the district of Upsala scatter them about and thus aid in its dispersal.
Artemisia achenes, since they have neither pappus nor other appendages, nor any special adhesiveness
when wetted, depend largely on their small size and light weight to aid them in dispersal. (Those of A.
absinthium measure a millimetre in length, or 1⁄25 of an inch, whilst those of A. vulgaris measure 1·8
mm., or 1⁄14 of an inch.) Driven as we are to look to bird-dispersal for the means of transport of
Artemisia achenes, it is interesting to find a possible source of the Hawaiian endemic species on the
nearest American mainland, even though it is some 2,000 miles away. It is assumed that they would be
ordinarily carried in adherent soil or entangled in the feathers, and on rare occasions in the bird’s
stomach.
The small seeds of Sisyrinchium possess no means of adherence to plumage. They are crustaceous,
and in cases where the stomach and intestines of a bird are well filled with other food they are quite
capable of resisting injury. The solitary Hawaiian species has, according to Hillebrand, a range in altitude
from 3,500 to 7,000 feet. I found this pretty herb most abundant on the “cattle-plains” of Hawaii
between 5,000 and 6,000 feet, where it is evidently in part dispersed by the cattle and other animals.
The seeds are very small, being about a millimetre in size, and when dried nearly 100 go to a grain
(0·65 decigramme). They might thus also be transported in mud on birds’ feet.
For the mode of dispersal of the minute seeds of Lobelia, the last of the mountain genera to be
specially noticed, I must refer the reader to the remarks on this subject in Chapter XXII. They would
probably be carried in soil adhering to the legs or feet of a bird.
There are one or two interesting points relating to the temperate genus Silene, which is represented
on these mountains. The four Hawaiian species show a great range in altitude. Thus, whilst S.
struthioloides finds its home in Hawaii and Maui at elevations of 5,000 to 9,000 feet, another species (S.
lanceolata) thrives equally at elevations of 5,000 or 6,000 feet on the central plateau of Hawaii and at
heights only of 300 to 500 feet above the sea. Although I have not yet come upon any direct reference
to the mode of dispersal of the small seeds of this genus, there is little doubt that their rough
tuberculated surfaces would favour their attachment to plumage. A very significant observation,
however, is made by Jens Holmboe in a paper on littoral plants in the interior of Norway. He refers to
the occurrence in no small quantity of Silene maritima on the top of “Linnekleppen,” 331 metres high,
one of the highest peaks of Smaalenene, and distant about 29 kilometres from the nearest coast
(Strandplanter i det indre af Norge, “Naturen,” Bergen, 1899). Sernander (p. 405), commenting on this
observation, remarks that since bare hill-tops are frequented by birds, such an agency in this instance is
not impossible.
I will conclude these remarks on the non-endemic Hawaiian mountain genera possessing only peculiar
species, with a few observations on the genus Vaccinium in the Pacific. This genus is known to be
distributed over the northern hemisphere and to occur on the uplands of tropical mountains, as, for
instance, on the summits of the Java mountains and on the high levels of the Equatorial Andes at
altitudes even of 15,000 to 16,000 feet. There are apparently only some four or five species known from
the Pacific islands, from Hawaii, the Marquesas, Tahiti, Rarotonga, Samoa, and the New Hebrides, and it
would almost seem that these can be reduced to one or two species. Although not yet recorded from
Fiji, the probability of the genus being represented on some of the mountains is pointed out by
Seemann. Of these Pacific forms a single species, V. cereum, is spread over the East Polynesian region
including the Marquesas, Tahiti, and Rarotonga; and, according to Hillebrand, V. reticulatum, one of the
two endemic Hawaiian species, is nearly related to it. Even the New Hebrides species (V. macgillivrayi)
resembles it, according to Seemann, in general appearance. That there has been a single Pacific
polymorphous species is, as shown below, not impossible; but Reinecke, in describing in 1898 the
Samoan species, V. antipodum, was under the impression that it was the only species known from the
southern hemisphere, and says nothing of its affinity to other Pacific plants.
A few words on the station and habit of Vaccinium in the Pacific islands may be here of interest. In
Hawaii there are, according to Hillebrand, two species, a high-level form, V. reticulatum, occurring at
elevations of 4,000 to 8,000 feet, and a low-level form, V. penduliflorum, ranging between 1,000 and
4,000 feet. I may, however, remark that the last species occasionally came under my notice at
elevations of 6,000 to 7,000 feet. This species exhibits much variation, and Gray, Wawra, and other
botanists have evidently not been always able to distinguish between the two species in their varying
forms. It is not only distinguished from the high-level species by its lower station, but also by its
epiphytic habit, a circumstance that, as pointed out below, may explain some of the differences, since
such a habit is bound up with the difference in station. It seems, therefore, safer to regard them as
station forms of one species which is closely allied to V. cereum, the species of the South Pacific, an
inference which, if well founded, would make highly probable the view that there has been a single
polymorphous Pacific species.... In Tahiti, as we learn from Nadeaud, V. cereum occurs on the
mountain-tops at altitudes exceeding 800 metres (2,600 feet). In Rarotonga, according to Cheeseman,
it is found on the summits of most of the higher hills extending almost to the summit of the island,
2,250 feet above the sea. The Samoan species, V. antipodum of Reinecke, which that botanist considers
as probably one with V. whitmei, a Polynesian (Samoa?) species originally described by Baron F. von
Müller, grows in the central mountains of Savaii at an elevation of 1,500 metres (4,920 feet).
These Pacific species of Vaccinium, as on tropical mountains of the continents, occasionally assume
an epiphytic habit, and it is here, as above observed, that lies one of the distinctions between the
Hawaiian species. V. penduliflorum, the low-level form, occurs typically in the forests, where, according
to Hillebrand, it grows on the trunks of old trees. The trees, however, may be quite in their prime, and I
have observed it growing in the fork of the trunk of an Olapa tree (Cheirodendron gaudichaudii). It is in
this connection of significance to notice that a variety found in open glades and on grassy slopes is
described by Hillebrand as terrestrial in habit. The other high-level form, V. reticulatum, grows
gregariously on open ground, and is typically terrestrial in its habit. The Samoan species, as we learn
from Reinecke, grows on trees, as on the branches of Gardenia. The epiphytic habit of species of
Vaccinium is especially discussed by Schimper in the case of plants growing on the Java mountains. He
there shows (Plant-Geography, i. 14) that species which are epiphytes in the virgin forest become
terrestrial plants in the treeless alpine region. This interchange of station, which is exhibited by several
other plants, including orchids and ferns, is connected with their xerophilous characteristics, and is
given by Schimper as an example of the interchange of physiologically dry habitats.
Of the mode of dispersal of Vaccinium by frugivorous birds, much has been written and much will be
familiar to my readers. The berries of V. reticulatum are known to be the principal food of the Hawaiian
mountain-goose. But probably birds of the grouse family have been the chief agents in distributing the
genus over the continents. I have frequently found the fruits in the stomachs of the Black Cock (Tetrao
tetrix), the Scotch Grouse (Lagopus scoticus), and the Capercailzie (Tetrao urogallus); but the same
story comes from all over the northern hemisphere. The Willow Grouse (Lagopus albus), which travels
round the globe, is known to feed on them. Hesselman in Sweden and Ekstam in Nova Zembla have
especially investigated the dispersal of Vaccinium by Tetrao tetrix and Lagopus (see Sernander, pp. 6,
226); and according to Mr. Douglas and others the different species of Tetrao that frequent the
subalpine regions of the Rocky Mountains and the uplands of Columbia and North California subsist on
Vaccinium fruits. This family is not now represented in the Hawaiian avifauna; but it is noteworthy, as
indicated by the differentiation of the Pacific species of Vaccinium, that dispersal of the genus is there
almost suspended except within the region of Eastern Polynesia. It is probable that numerous other
birds, except the Hawaiian goose, aided the original dispersal.
The Mountain Genera with both Endemic and Non-endemic Species.—I pass on now to consider those
Hawaiian mountain genera that possess species some of which are confined to the group, whilst others
occur in regions outside the islands. They are not many, as may be seen from the table before given,
and but few of them are entirely restricted to the high levels, a range in altitude that may be frequently
associated with great lateral extension of the genus over different latitudes. Here the agents of dispersal
have through some species in each genus preserved a connection with the outer world, though it may
be restricted to the limits of the Pacific islands.
Cyathodes tameiameiæ, an Epacridaceous species found also in the uplands of Tahiti, occurs,
according to Hillebrand, on all the Hawaiian Islands, from 1,800 feet up to the limit of vegetation 10,000
feet and over above the sea. I found it, however, at even lower levels. On the Puna coast of Hawaii,
associated with Metrosideros polymorpha, Osteomeles anthyllidifolia, and other inland plants, it
descends on the surface of ancient lava-flows to the coast wherever the bolder spurs reach the sea-
border. The other species, C. imbricata, is more exclusively confined to the greater altitudes. It is
endemic, and may possibly be a station form of the other species.
The six species of Lysimachia are found at different elevations, one near the sea-shore, others at
altitudes of 2,000 to 3,000 feet, and others again at elevations of 6,000 feet. Chenopodium
sandwicheum occurs at all elevations from near the coast to the high inland plains of Hawaii and to the
upper slopes of Mauna Kea, that is to say, up to altitudes of 6,000 or 7,000 feet. Hillebrand observes
that it is a low decumbent plant at the coast, and may become arborescent with a height of 12 to 15
feet in the upper forests of Mauna Kea.
The species of Santalum (sandal-wood trees) also display great vertical range in these islands.
Though S. freycinetianum, which is also a Tahitian species, is most at home in the forests 2,000 to
4,000 feet above the sea, it has, as Hillebrand informs us, a dwarfed form that extends far up the
mountain slopes of Mauna Loa and Hualalai to elevations of 7,000 or 8,000 feet, and another dwarfed
shrubby variety that grows only near the sea-shore. Another species, S. haleakalæ, occurs as a tall
shrub on Haleakala at elevations of 8,000 to 10,000 feet. Among the sedges, most of those of the
genera Carex and Rhynchospora are found at altitudes of between 3,000 and 7,000 feet, and two
grasses of the genus Deyeuxia occur at elevations of 6,000 to 8,000 feet.
Amongst these Hawaiian mountain genera with both endemic and non-endemic species there are no
plants possessing fruits which from their size could be with difficulty regarded as dispersed by birds.
The mode of dispersal of these plants is in some cases indeed not far to seek. Thus in the stomach of
an Hawaiian goose (Bernicla sandwicensis), shot by my companion Dr. Krämer on the slopes of Mauna
Loa, I found a number of the “stones” of Cyathodes tameiameiæ, the plant being abundant in fruit in
the immediate vicinity. It is highly probable that the seeds of Santalum have been carried over the
Pacific by frugivorous birds. We learn from Dr. Brandis that Santalum album in India is mainly spread
through the agency of birds (Bot. Chall. Exped., iii. 13). The drupes of the Pacific species, S.
freycinetianum, that occurs alike in Hawaii, the Marquesas, and Tahiti (Drake del Castillo), measure
about half an inch. There can be little doubt that with this tree, as with the species of Cyathodes above
mentioned, which also links together Tahiti and Hawaii, there has been up to recent times an
interchange by means of frugivorous birds between these two regions, some 2,000 miles apart.
The small seeds of the capsular fruits of Lysimachia could be transported in birds’ plumage or in dried
soil attached to their feet or feathers. The seed-like fruits of Chenopodium were probably dispersed by
some granivorous bird, much as nowadays our partridges carry about in their stomachs the similar fruits
of Atriplex. The long-awned fruits of Deyeuxia were, it is likely, transported in birds’ plumage, and
doubtless also those of Panicum; whilst the nutlets of Carex and Rhynchospora might have been carried
about in a similar fashion.
The distribution of the non-endemic species of these Hawaiian mountain genera may perhaps aid us
in determining the original source of the genus as well as in confirming the conclusions formed
concerning the other mountain genera that only possess species restricted to the group. Lysimachia,
Chenopodium, Carex, Rhynchospora, Deyeuxia, and Panicum are found in both the Old and New
Worlds. Since Hillebrand remarks that one of the six species of Lysimachia (L. spathulata) occurs in
Japan and in the Liukiu, Bonin, and Marianne groups, we have here a valuable indication of the route
followed by a genus that has not been recorded from the oceanic groups of the South Pacific.
The capricious distribution of the genus Carex in the Pacific is remarkable, and it is noticed by
Hemsley in the Introduction to the Botany of the “Challenger” Expedition. No species have been
recorded from Tahiti, the Marquesas, and Rarotonga, but three Fijian species are mentioned by
Hemsley, and there is another in Samoa. Of the five Hawaiian species given by Hillebrand, two are
endemic. Of the rest, C. wahuensis (oahuensis), Meyer, occurs also in Korea and Japan, whilst C.
brunnea, Thunb., is found in Japan and Australia, and the third, C. propinqua, Nees., occurs all round
the border of the Pacific Ocean, from Kamschatka through Alaska south to the Straits of Magellan.
These three species all possess a home in common in north-east Asia, and probably there lies the
source of the Hawaiian species of Carex—a conclusion which would help to explain the irregular
distribution of the genus amongst the South Pacific groups.
The genus Rhynchospora occurs alike in the Hawaiian, Tahitian, and Fijian islands; but the groups in
the North and South Pacific seem to have been independently supplied with the original species, since
R. aurea, a widely spread tropical species, ranging the South Pacific from New Caledonia to Tahiti, has
not been recorded from Hawaii. A connection between Hawaii and the Australian region seems to be
indicated by a species of Deyeuxia (D. forsteri) that is found also in Easter Island, Australia, and New
Zealand, and by the presence of the Australian and New Zealand genus Cyathodes in Hawaii, though
the existence of a species common to both Tahiti and Hawaii goes to show that the route followed by
the genus lay through Eastern Polynesia. It is also not unlikely that the genus Santalum reached Hawaii
through Eastern Polynesia, since two forms found in Hawaii and Tahiti are closely allied, and are, in fact,
regarded by Drake del Castillo as the same species. The genus occurs in tropical Asia, Australia, and
New Zealand.
Looking at the indications above given, I should be inclined to think that the genera Lysimachia and
Carex reached the Hawaiian mountains from temperate Asia or the islands off its Pacific coast, and that
Cyathodes, Santalum, and Deyeuxia hail from the Australian or New Zealand region by way of Eastern
Polynesia.
The Mountain Genera possessing no Endemic Species.—The few remaining mountain plants of Hawaii
to be considered are solitary, widely ranging species of genera that here possess no peculiar species.
Such may be regarded as belonging to the latest age of the indigenous plants. They still keep up, or
kept up until recently, the connection with the world outside Hawaii, and among them one may name
here Fragaria chilensis, Drosera longifolia, Nertera depressa, and Luzula campestris.
Fragaria chilensis, the Chilian strawberry, flourishes at elevations of between 4,000 and 6,000 feet on
the Hawaiian mountains. Its fruits, according to Hillebrand and other authors, are much appreciated by
the wild goose of the islands. This plant ranges in America from Chile north to Alaska; and Drake del
Castillo is doubtless on safe ground when he assumes that a congener of this bird originally brought the
species from the nearest part of the American continent, namely from California (Remarques, &c., p. 8).
In this connection it should be remembered that one of the endemic mountain-raspberries of Hawaii
(Rubus hawaiiensis) finds its nearest relative, according to Gray, in Rubus spectabilis, a species from the
north-west coast of America.
The species of Sun-dew, Drosera longifolia, hitherto found only on the marshy tableland of Kauai at
an elevation of 4,000 feet above the sea, occurs both in Asia and North America. Its minute fusiform
seeds are very light in weight, and might readily become entangled in a bird’s plumage, or they could
be carried in adherent dried mud.
Luzula campestris, which grows on the high mountains of the Hawaiian group from 3,000 feet
upward, is also found in Tahiti. It is widely distributed in cool latitudes, and there is no special indication
of its source. Its seeds are especially well suited for adhering to birds’ feathers. When experimenting on
these seeds in 1893 I ascertained that whether freshly gathered or kept for more than a year they
became on wetting coated with mucus, and adhered firmly to a feather on drying. There are many ways
in which the “sticky” seeds in wet weather might fasten themselves to a bird’s plumage. The plant-
materials might be used, for instance, for making nests. The Sea Eagle (Aquila albicilla), as we learn
from Mr. Napier (Lakes and Rivers), uses materials derived from Luzula sylvatica in the construction of
its nest.
Nertera depressa, a creeping Rubiaceous plant, with red, fleshy drupes containing two coriaceous
pyrenes, is found in all the Hawaiian Islands at elevations of 2,500 to 5,000 feet, and it grows on the
mountains of Tahiti at altitudes over 3,000 feet. The genus is widely diffused over the southern
hemisphere. This particular species is characteristic of the Antarctic flora, being found all round the
south temperate zone (excepting South Africa) in New Zealand, Fuegia, the Falkland Islands, and Tristan
da Cunha, and extending up the Andes to Mexico, occurring also on the summits of Malayan mountains
at elevations of 9,000 to 10,500 feet above the sea, as on Pangerango in West Java (Schimper), and on
Kinabalu in North Borneo (Stapf). Captain Carmichael, who resided on Tristan da Cunha in the early part
of last century, states (Trans. Linn. Soc., xii. 483) that its drupes are eaten by a species of thrush and
by a bunting. Professor Moseley, who visited the island in the Challenger many years after, remarks that
its fruits are “the favourite food of the remarkable endemic thrush, Nesocichla eremita,” the bunting
being Emberiza brasiliensis (Bot. Chall. Exped., ii. 141). It would seem most likely that the Hawaiian
Islands received this representative of the Antarctic flora through the Tahitian Islands, as in the case of
the species of Cyathodes common to both these groups.
Looking at the indications of these four widely ranging plants, the Chilian strawberry (Fragaria
chilensis), the Sun-dew (Drosera longifolia), Nertera depressa, and Luzula campestris, it may be inferred
that with the exception of Nertera they all reached Hawaii from either the Asiatic or American sides of
the North Pacific, the last route being evident in the case of the strawberry. Nertera depressa was
probably derived from southern latitudes.

Summary.
(1) The second era of the flowering plants of the Pacific islands is indicated by the non-endemic
genera. Here also the isolating influences have been generally active, and the work of dispersal is in
some regions largely suspended. Thus in Hawaii nearly half the non-endemic genera possess only
species that are restricted to the group, whilst in Fiji and Tahiti about a fourth are thus isolated.
(2) The contrast in the elevations of the islands of the Hawaiian, Tahitian, and Fijian regions is
reflected in the development of an extensive mountain-flora in Hawaii, in its scanty development in
Tahiti, and, excluding the Fijian conifers, in a mere remnant in Fiji and Samoa.
(3) The influence of isolation has been very active in the Hawaiian mountains, since about two-thirds
of the genera contain only species confined to the group, and are thus disconnected from the world
outside.
(4) Amongst these disconnected Hawaiian mountain genera, Antarctic or New Zealand genera, like
Acæna, Gunnera, Coprosma, and Lagenophora, constitute nearly a third. The American element,
represented, for instance, by Sanicula and Sisyrinchium, is small; whilst the genera found on both sides
of the Pacific form more than one-half of the total, and include genera like Ranunculus, Viola, Rubus,
Artemisia, Vaccinium, and Plantago, that often represent the flora of the temperate zone on the
summits of tropical mountains. Three-fourths of these genera are not found either in Fiji or in Tahiti.
(5) The proportion of the disconnected Hawaiian mountain genera possessing seeds or seedvessels
suited for dispersal in a bird’s plumage is very large, quite half belonging to this category; whilst only
about a fourth have fruits that would be dispersed by frugivorous birds.
(6) The Hawaiian mountain genera that still remain in touch with the external world through species
found outside the islands whilst other species are confined to the group, present a later stage in the
plant-stocking. Their widely ranging species, which would be dispersed either by frugivorous birds, as
with Santalum and Cyathodes, or in birds’ plumage, as with Lysimachia, Carex, and Deyeuxia, seem to
indicate that the main lines of migration for these genera have been from temperate Asia and from the
Australian and New Zealand region, the last by way of Eastern Polynesia.
(7) The latest stage of the Hawaiian mountain-flora is exemplified by those genera that are only
represented in the group by a solitary widely-ranging species, such as Fragaria chilensis, Nertera
depressa, Drosera longifolia, and Luzula campestris. It is our own age; and birds are shown to be actual
agents in the dispersal of the two first-named species and to be probable agents with the two other
species. The two last-named species probably reached Hawaii from one or other side of the North
Pacific; whilst Fragaria chilensis doubtless hails from the adjacent part of the American continent, and
Nertera depressa from high southern latitudes by way of Tahiti.
 CHAPTER XXIV

THE MOUNTAIN-FLORAS OF THE TAHITIAN AND

FIJIAN REGIONS

The mountain-flora of the Tahitian region, as illustrated by the non-

endemic genera.—Derived chiefly from high southern latitudes.—
Weinmannia, Coprosma, Vaccinium, Astelia, Coriaria, Cyathodes,
Nertera depressa, Luzula campestris.—The mountain flora of
Rarotonga.—The mountain-flora of the Fijian region, as illustrated
by the non-endemic genera.—Weinmannia, Lagenophora,
Coprosma, Astelia, Vaccinium, Nertera depressa.—The Fijian
Coniferæ.—Dammara, Podocarpus, Dacrydium.—Not belonging to
the present era of dispersal.—The age of dispersal of the Coniferæ
in the Pacific.—Earlier than the age of Compositæ and Lobeliaceæ.
—The first in the Mesozoic period.—The last in the Tertiary period.
—Summary.

The Mountain-Flora of theTahitian Region as Illustrated

by the Non-Endemic Genera
This floral region of the Pacific corresponds with the limits of Eastern
Polynesia, and includes not only the Tahitian group proper, but also
the Cook, Austral, Paumotuan, and Marquesan groups. It is only,
however, in Tahiti, the peaks of which rise to over 7,000 feet above
the sea, that we should expect to find such a mountain-flora, since
the islands of the other groups are much lower, the highest of them
in the Marquesan group barely exceeding 4,000 feet. Yet even in
Tahiti it is not possible to speak of a mountain-flora in the sense that
we attach to it in Hawaii. The elevated area of its interior is, as
described in Chapter XIX., relatively very small; whilst, as Drake del
Castillo points out, the conditions presented by the steep mountain-
slopes rarely afford a hold for trees of any size, ferns often
predominating in the higher levels. Still, we can observe the traces
of such a flora, and it is in this sense only that the term “mountain-
genera” is used in relation with this group.

Mountain-Genera of the Tahitian or East Polynesian Region.

all
Weinmannia, Saxifragaceæ, from New Zealand.
species
Coprosma, Rubiaceæ, from New Zealand.
endemic.
Vaccinium, Vacciniaceæ, from the northern hemisphere.
Astelia, Liliaceæ, from New Zealand.

Coriaria, Coriariaceæ, from New Zealand some species

Cyathodes, Epacridaceæ, from New Zealand endemic

Nertera depressa, Rubiaceæ, a species of the Antarctic flora.

Luzula campestris, Juncaceæ, from the northern hemisphere.
The Tahitian non-endemic mountain-genera, though scanty in
number, are of considerable interest to the student of plant-
dispersal. Among those possessing only species that are confined to
Eastern Polynesia, genera that would be regarded as belonging to a
past era of dispersal, Weinmannia, Coprosma, Vaccinium, and Astelia
may be mentioned.
Weinmannia, a Saxifragaceous genus of trees and shrubs, not
represented in Hawaii, but recorded from almost all the elevated
oceanic groups of the tropical South Pacific, as well as from the New
Hebrides and New Caledonia, has its home in South America, more
particularly in the Andes, and also occurs in New Zealand, Tasmania,
and the Mascarene Islands. One can scarcely doubt that, as in the
case of Coprosma, the Pacific Islands derived their species originally
from high southern latitudes, as from New Zealand, the absence of
the genus from Hawaii negativing an American origin. Of the two
Tahitian species, one is peculiar to Tahiti, whilst the other, W.
parviflora, which is conspicuous on the mountain-crests at elevations
of 3,000 feet and over, occurs also in the Marquesas. Another
species grows in abundance in the interior of Rarotonga. Samoa
possesses two species, one of which, W. affinis, occurs in Fiji, and
the other, W. samoensis, which frequents the mountains at
elevations of 1,500 to 3,300 feet, is seemingly endemic. Fiji
possesses four or five species of Weinmannia occurring at all
altitudes up to 2,000 feet, of which some are evidently peculiar. The
capsular fruits of this genus contain hairy seeds that would probably
become entangled in a bird’s plumage. Dispersal by birds is
distinctedly indicated in the curious observation of Dr. Reinecke in
the case of the Samoan peculiar species. The seeds, he says, appear
to germinate by preference on the bark of other trees, young plants
growing epiphytically being of frequent occurrence.
There is some evidence that the species of Weinmannia, about ten
in all, found in the tropical islands of the open Pacific are derived
from one or two polymorphous species. As we learn from Mr.
Cheeseman, the Rarotongan species, W. rarotongensis, has
considerable affinity to several closely allied Polynesian species, and
its nearest allies are a Fijian and Samoan species, W. vitiensis and W.
samoensis. Possibly, he remarks, fuller materials may lead to the
union of several of these forms under one species.
The interesting New Zealand genus Coprosma, which we have
noticed in Hawaii, occurs also in the Tahitian region and Fiji; and it
will be further discussed under the last-named locality. The genus
Vaccinium has been previously dealt with in Chapter XXIII.
The Liliaceous genus Astelia may be considered as representing,
like Coprosma, the Antarctic or New Zealand flora in the higher
levels (usually) of the islands of the tropical Pacific, where it grows
both on trees and on the ground. The genus, according to Hemsley,
is chiefly at home in New Zealand, but is also found in Fuegia and in
South-east Australia. It is represented in Hawaii, Tahiti, Samoa, and
Fiji. In Hawaii there are two peculiar species ranging between 2,000
to 6,000 feet in elevation. The solitary Tahitian species, A. nadeaudi,
is found in the central mountains of Tahiti, reaching to the crests of
Mount Aorai, which attains a height of 6,700 feet. Fiji and Samoa
possess a species in common, A. montana, which is only recorded by
Seemann, from the summit of Kandavu, 2,750 feet above the sea;
whilst in Samoa it frequents, according to Reinecke, moist coast
districts. The fruits of Astelia are berries with crustaceous seeds that
would be dispersed by frugivorous birds.
Amongst the Tahitian mountain-genera that possess species
ranging far beyond this region as well as species confined to the
group may be mentioned Coriaria and Cyathodes. It is to their non-
endemic species that we look for further clues as to the general lines
of migration by which the mountain-genera that only possess
peculiar species reached this group. The evidence afforded by
Coriaria is of some importance. The genus has not been recorded
from Hawaii, and, so far as the collections of Seemann and Home
show, not from Fiji. It is found in the Mediterranean region, the
Himalayas, Japan, New Zealand, and Antarctic America, including
Chile; and there are two particular species, C. ruscifolia and C.
thymifolia, that occur in both cases in New Zealand and the adjacent
islands and in South America (Introd. Chall. Bot. p. 53). The first of
these, which is very common in Chile, exists also in Tahiti on the
crest of Aorai, 6,700 feet above the sea. Drake del Castillo also
describes a peculiar Tahitian species, C. vescoi, of which the altitude
is not given. Here one is in doubt whether Tahiti derived its wide-
ranging species from New Zealand or from Chile; but in the New
Zealand home of Coprosma, another Tahitian mountain-genus, we
are afforded the clue. The fruits of Coriaria possess fleshy cocci that
attract birds, though it would seem that the seeds of plants of this
genus are poisonous for man. Among the numerous fruits that form
the diet of the New Zealand fruit-pigeon (Carpophaga novæ
zealandiæ) are included, as we learn from Sir W. Buller in his Birds
of New Zealand, those of the “tupakihi” or “tutu” shrub, which Kirk
identifies with C. ruscifolia, the species that also occurs on the
summit of Tahiti.
The Australian and New Zealand genus Cyathodes (Epacridaceæ)
has been already noticed in the case of Hawaii (page 282). The two
Tahitian species occur on the elevated mountain-ridges forming the
summits of Tahiti, one of them, C. tameiameiæ, occurring also in
Hawaii, and the other, C. pomaræ, being restricted to the group. I
have shown that the fruits are dispersed by frugivorous birds, and I
can only include the genus as another example of the representation
of the New Zealand flora in Tahiti.... There remain of these so-called
Tahitian mountain-genera the Antarctic Nertera and the north-
temperate Luzula, each represented by a solitary widely ranging
species, N. depressa and L. campestris, which I have fully discussed
under Hawaii (Chapter XXIII), in which group they also occur.
When we look at the evidence of origin supplied by the four
Tahitian mountain-genera possessing species that are found outside
the group, namely Coriaria, Cyathodes, Nertera, and Luzula, we find
that the first three hail from high southern latitudes, and more
especially from New Zealand; and when with this clue in our hands
we take up the four genera Weinmannia, Coprosma, Vaccinium, and
Astelia, possessing only species restricted to the Tahitian region, we
find that all but the third-named genus hail also from the south. It
would thus appear that the element of the Antarctic flora is much
more evident in the Tahitian mountain-genera than with those of
Hawaii. In the Hawaiian mountain-flora, excluding, of course, the
endemic genera, it includes about a fourth of the mountain-genera,
which number about thirty-eight or forty in all; whilst in the Tahitian
mountain-flora it comprises six out of the eight genera. It may,
indeed, be said that the resemblance between the mountain-genera
of Hawaii and Tahiti is mainly restricted to genera that are found in
high southern latitudes, namely, Nertera, Coprosma, Cyathodes, and
Astelia, the only other genera linking the mountain-floras of both
groups together being Vaccinium and Luzula, which probably hail
from high northern latitudes. The agency of the frugivorous bird is
plainly marked in the case of five out of the six genera that connect
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